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Kiwifruit: The Genus
ACTINIDIA
 Supported by the National Fund for
Academic Publication in Science and Technology

Kiwifruit: The Genus

ACTINIDIA
HUANG Hongwen
Wuhan Botanical Garden, Chinese Academy of Sciences, Wuhan, China

AMSTERDAM • BOSTON • HEIDELBERG • LONDON

NEW YORK • OXFORD • PARIS • SAN DIEGO
SAN FRANCISCO • SINGAPORE • SYDNEY • TOKYO
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Contents
Contributors............................................................................................ix
Foreword...................................................................................................xi
Preface......................................................................................................xiii
Acknowledgments..............................................................................xv
Introduction................................................................................................ 1
1 Systematics and Genetic Variation of Actinidia.............................. 9
1.1 Taxonomic Revision............................................................................. 9
1.1.1 Actinidia Lindl................................................................................... 12
1.1.2 Clematoclethra (Franch.) Maxim....................................................... 16
1.1.3 Saurauia Willdenow......................................................................... 16
1.2 Treatment of Taxa in the Most Recent Revision
of Actinidia............................................................................................. 17
1.2.1 Three Varieties of Actinidia arguta (Siebold & Zuccarini)
Planchon ex Miquel........................................................................ 20
1.2.2 Actinidia melanandra Franchet, Varieties and
Synonyms....................................................................................... 21
1.2.3 Actinidia kolomikta (Ruprecht & Maximowicz)
Maximowicz................................................................................... 21
1.2.4 Actinidia tetramera Maximowicz....................................................... 21
1.2.5 Actinidia valvata Dunn...................................................................... 21
1.2.6 Actinidia callosa Lindley and A. umbelloides C.F. Liang...................... 21
1.2.7 The Actinidia chinensis Planchon Complex........................................ 21
1.2.8 Actinidia eriantha Bentham.............................................................. 22
1.2.9 Synonomy of Actinidia fortunatii Finet & Gagnepain,
A. glaucophylla F. Chun, A. henryi Dunn and A. gracilis
C.F. Liang........................................................................................ 23
1.2.10 Actinidia fulvicoma Hance, A. cinerascens C.F. Liang and
A. fulvicoma f. hirsuta (Finet & Gagnepain) C.F. Liang..................... 23
1.2.11 Actinidia hemsleyana var. kengiana (F.P. Metcalf)
C.F. Liang and A. kengiana F.P. Metcalf............................................ 23
1.2.12 Actinidia carnosifolia C.Y. Wu, A. carnosifolia var. glaucescens
C.F. Liang, and A. henryi var. polyodonta Handel-Mazzetti.............. 23
1.2.13 Actinidia flavofloris H.Z. Jiang......................................................... 24
1.2.14 Actinidia leptophylla C.Y. Wu, A. maloides H.L. Li and
A. maloides f. cordata C.F. Liang...................................................... 24
1.2.15 Actinidia jiangkouensis S.D. Shi & Z.S. Zhang and
A. laevissima var. floscula S.D. Shi................................................... 24
1.2.16 Actinidia guilinensis C.F. Liang........................................................ 24
1.2.17 Actinidia linguiensis R.G. Li & X.G. Wang and
A. wantianensis R.G. Li & L. Mo....................................................... 24
1.2.18 Actinidia longicarpa R.G. Li & M.Y. Liang and
A. rubrafilmenta R.G. Li & J.W. Li..................................................... 24
1.2.19 Actinidia petelotii Diels and A. rudis Dunn....................................... 24
1.2.20 Actinidia pentapetala R.G. Li & J.W. Li............................................. 24
1.2.21 Actinidia zhejiangensis C.F. Liang and Actinidia persicina
R.H. Huang & S.M. Wang................................................................ 24
1.2.22 Actinidia vitifolia C.Y. Wu and A. rubus H. Léveillé........................... 24
1.2.23 Actinidia rubricaulis Dunn and A. fasciculoides C.F. Liang................ 24
1.2.24 Actinidia ulmifolia C.F. Liang, A. sorbifolia C.F. Liang,
A. stellatopilosa C.Y. Chang, A. obovata Chun ex C.F. Liang,
and A. grandiflora C.F. Liang............................................................ 25
1.3 Paleopolyploid Origin, Biogeographic Patterns,
and Reticulate Evolution.................................................................. 25
1.3.1 Paleopolyploid Origin and Evolution of Actinidia.............................. 25
1.3.2 Biogeographic Patterns and Reticulate Evolution............................. 27
1.4 Actinidia Palynology.......................................................................... 30
1.4.1 Pollen Size and Morphology............................................................. 30
1.4.2 Phylogenetic Relationships in Actinidia as Revealed
by Pollen Morphology..................................................................... 31
1.4.3 Applications of Palynology............................................................... 32
1.5 Flower Morphology and Sex Variation............................................ 35
1.6 Ploidy Variation and Breeding Applications.................................. 37
1.6.1 Basic Chromosome Number............................................................. 37
1.6.2 Ploidy Races and Interploidy Hybridization...................................... 37
1.6.3 Nuclear DNA Content and Flow Cytometry........................................ 42
1.6.4 Ploidy Variation for Kiwifruit Breeding............................................. 43
1.7 Genomic Diversity.............................................................................. 43
2 Species....................................................................................................45
2.1 Actinidia arguta.................................................................................. 46
2.1a Actinidia arguta var. arguta................................................................ 46
2.1b Actinidia arguta var. giraldii............................................................... 48
vi Contents
2.2 Actinidia callosa.................................................................................. 49
2.2a Actinidia callosa var. callosa............................................................... 49
2.2b Actinidia callosa var. acuminata......................................................... 51
2.2c Actinidia callosa var. discolor.............................................................. 52
2.2d Actinidia callosa var. henryi................................................................ 54
2.2e Actinidia callosa var. strigillosa........................................................... 56
2.3 Actinidia chengkouensis..................................................................... 57
2.4 Actinidia chinensis.............................................................................. 59
2.4a Actinidia chinensis var. chinensis........................................................ 59
2.4b Actinidia chinensis var. deliciosa......................................................... 61
2.4c Actinidia chinensis var. setosa............................................................. 64
2.5 Actinidia chrysantha........................................................................... 66
2.6 Actinidia cylindrica............................................................................. 68
2.6a Actinidia cylindrica var. cylindrica....................................................... 68
2.6b Actinidia cylindrica var. reticulata....................................................... 70
2.7 Actinidia eriantha............................................................................... 71
2.8 Actinidia farinosa................................................................................ 74
2.9 Actinidia fasciculoides........................................................................ 76
2.9a Actinidia fasciculoides var. fasciculoides............................................. 76
2.9b Actinidia fasciculoides var. cuneata.................................................... 77
2.9c Actinidia fasciculoides var. orbiculata................................................. 78
2.10 Actinidia fortunatii........................................................................... 79
2.11 Actinidia fulvicoma........................................................................... 81
2.11a Actinidia fulvicoma var. fulvicoma.................................................... 81
2.11b Actinidia fulvicoma var. cinerascens.................................................. 83
2.11c Actinidia fulvicoma var. hirsuta......................................................... 85
2.11d Actinidia fulvicoma var.
pachyphylla.................................................................................... 86
2.12 Actinidia glaucocallosa.................................................................... 87
2.13 Actinidia grandiflora........................................................................ 89
2.14 Actinidia hemsleyana....................................................................... 91
2.15 Actinidia henryi................................................................................. 92
2.16 Actinidia holotricha.......................................................................... 94
2.17 Actinidia hubeiensis......................................................................... 96
2.18 Actinidia indochinensis.................................................................... 97
2.18a Actinidia indochinensis var. indochinensis......................................... 98
2.18b Actinidia indochinensis var. ovatifolia............................................... 99
2.19 Actinidia kolomikta........................................................................100
2.20 Actinidia laevissima........................................................................103
2.21 Actinidia lanceolata........................................................................104
2.22 Actinidia latifolia..........................................................................106
2.22a Actinidia latifolia var. latifolia......................................................... 106
2.22b Actinidia latifolia var. mollis........................................................... 108
2.23 Actinidia liangguangensis.............................................................109
2.24 Actinidia lijiangensis......................................................................111
2.25 Actinidia linguiensis.......................................................................113
2.26 Actinidia longicarpa.......................................................................113
2.27 Actinidia macrosperma..................................................................115
2.27a Actinidia macrosperma var. macrosperma...................................... 115
2.27b Actinidia macrosperma var. mumoides........................................... 117
2.28 Actinidia melanandra.....................................................................118
2.28a Actinidia melanandra var. melanandra........................................... 118
2.28b Actinidia melanandra var. glabrescens............................................ 121
2.29 Actinidia melliana...........................................................................122
2.30 Actinidia obovata............................................................................123
2.31 Actinidia persicina..........................................................................125
2.32 Actinidia pilosula............................................................................126
2.33 Actinidia polygama.........................................................................128
2.34 Actinidia rongshuiensis..................................................................130
2.35 Actinidia rubricaulis.......................................................................130
2.35a Actinidia rubricaulis var. rubricaulis................................................ 130
2.35b Actinidia rubricaulis var. coriacea................................................... 132
2.36 Actinidia rubus................................................................................134
2.37 Actinidia rudis.................................................................................135
2.37a Actinidia rudis var. rudis................................................................. 135
2.37b Actinidia rudis var. glabricaulis....................................................... 137
2.38 Actinidia rufa...................................................................................138
2.39 Actinidia rufotricha........................................................................140
2.39a Actinidia rufotricha var. rufotricha.................................................. 141
2.39b Actinidia rufotricha var. glomerata................................................. 142
2.40 Actinidia sabiifolia..........................................................................144
2.41 Actinidia sorbifolia.........................................................................146
2.42 Actinidia stellatopilosa..................................................................147
2.43 Actinidia styracifolia......................................................................148
2.44 Actinidia suberifolia.......................................................................150
2.45 Actinidia tetramera........................................................................151
2.46 Actinidia trichogyna.......................................................................153
2.47 Actinidia ulmifolia..........................................................................154
2.48 Actinidia umbelloides.....................................................................155
2.48a Actinidia umbelloides var. umbelloides........................................... 155
2.48b Actinidia umbelloides var. flabellifolia............................................ 157
2.49 Actinidia valvata.............................................................................157
 Contents
2.50 Actinidia venosa..............................................................................159
2.51 Actinidia vitifolia............................................................................161
2.52 Actinidia zhejiangensis..................................................................163
2.53 Actinidia hypoleuca........................................................................165
2.54 Actinidia strigosa............................................................................167
3 Natural Distribution of Genus Actinidia...................................... 169
3.1 Regional Distribution, Natural Habitats, and Wild
Production........................................................................................169
3.1.1 China.............................................................................................. 169
3.1.2 East Asia and Southeast Asia.......................................................... 183
3.2 Vertical Distribution........................................................................185
3.3 Geographic Distribution of Commercially Important
Species...............................................................................................187
3.4 Geographically Overlapping Distributions, Natural
Hybridization, and Gene Introgression of Species......................189
4 Domestication Commercialization of Actinidia......................... 191
4.1 Use of Actinidia Natural Resources in Ancient
China..................................................................................................191
4.2 Early Western Introductions of Actinidia Species........................193
4.3 Domestication and Commercialization of A. chinensis var.
deliciosa.............................................................................................194
4.3.1 First Botanical Specimens............................................................... 194
4.3.2 Cultivation of A. chinensis var. deliciosa Outside China.................... 195
4.3.3 Commercialization of A. chinensis var. deliciosa.............................. 197
4.3.4 Early Attempts to Cultivate A. chinensis var. chinensis
Outside China............................................................................... 199
4.4 Domestication and Commercialization of A. chinensis
var. chinensis.....................................................................................200
4.4.1 Initial Experimental Cultivation...................................................... 200
4.4.2 Evaluation of Wild Actinidia Germplasm in China........................... 201
4.4.3 Commercial Cultivation of A. chinensis var. chinensis in
China............................................................................................ 203
4.4.4 Commercialization of A. chinensis var. chinensis Outside
China............................................................................................ 204
4.5 Domestication and Commercialization of A. kolomikta,
A. arguta, A. polygama, and A. eriantha........................................204
4.6 The World Kiwifruit Industry..........................................................205
4.6.1 World Plantings of Kiwifruit........................................................... 205
4.6.2 World Production of Kiwifruit......................................................... 206
4.6.3 Important Kiwifruit Cultivars in International Trade....................... 207
4.6.4 Segmentation of the Kiwifruit Market............................................ 208
vii
5 Biology, Genetic Improvement, and Cultivar
Development..................................................................................... 211
5.1 Morphological Characteristics.......................................................211
5.1.1 Roots.............................................................................................. 211
5.1.2 Buds............................................................................................... 212
5.1.3 Stems............................................................................................. 213
5.1.4 Leaves............................................................................................ 213
5.1.5 Flowers........................................................................................... 213
5.1.6 Fruits.............................................................................................. 214
5.1.7 Seeds.............................................................................................. 215
5.2 Growth and Development Characteristics...................................215
5.2.1 Growth........................................................................................... 215
5.2.2 Flower Development...................................................................... 215
5.2.3 Fruiting Characteristics................................................................... 217
5.3 Variation in Important Fruit Characteristics of
Actinidia.............................................................................................217
5.3.1 Fruit Size......................................................................................... 217
5.3.2 Fruit Shape..................................................................................... 221
5.3.3 Fruit Hairs and Skin........................................................................ 221
5.3.4 Fruit Color....................................................................................... 223
5.3.5 Ripening Indicators........................................................................ 225
5.3.6 Fruit Texture................................................................................... 225
5.3.7 Fruit Flavor..................................................................................... 226
5.3.8 Health Benefits of Kiwifruit............................................................ 227
5.3.9 Nutrient Composition..................................................................... 227
5.3.10 Other Fruit Characteristics............................................................ 230
5.4 Progress in the Selection and Breeding of Actinidia..................230
5.4.1 Direct Introduction and Selection from the Wild............................. 230
5.4.2 Selection from Seed Collected in the Wild...................................... 231
5.4.3 Selection from Open Pollination of Cultivars or Named
Selections..................................................................................... 231
5.4.4 Clonal Selections............................................................................ 231
5.4.5 Budsport Selections....................................................................... 231
5.4.6 Cultivars Developed from Systematic Breeding Programs............... 231
5.4.7 Pollenizer Selections....................................................................... 233
5.4.8 Cultivars of A. arguta, A. kolomikta, and A. rufa.............................. 233
5.5 Future Cultivars of Actinidia...........................................................233
5.5.1 Further Exploration of Wild Genetic Resources............................... 234
5.5.2 Gene Introgression Selection and Breeding from Natural
Populations.................................................................................. 234
5.5.3 Gene Introgression Breeding: Theoretical and Practical
Aspects......................................................................................... 235
viii Contents
6 Main Cultivars in Commercial Production................................... 239
6.1 Main Cultivars of A. chinensis var. deliciosa..................................239
6.1.1 Hayward......................................................................................... 239
6.1.2 Qinmei........................................................................................... 241
6.1.3 Miliang No. 1.................................................................................. 241
6.1.4 Emihoutao No. 1 (Jinkui)................................................................ 242
6.1.5 Xuxiang.......................................................................................... 243
6.1.6 Bruno............................................................................................. 244
6.1.7 Guichang........................................................................................ 245
6.1.8 Cuixiang (Ximi No. 9) ..................................................................... 245
6.1.9 Yumihoutao No. 2 (Huamei No. 2) .................................................. 246
6.1.10 Jinxiang........................................................................................ 247
6.1.11 Zesh004 (Green 14) ...................................................................... 248
6.2 Main Cultivars of A. chinensis var. chinensis.................................248
6.2.1 Hort16A (Earligold, Fruit Marketed as Zespri®
Gold Kiwifruit) ............................................................................. 248
6.2.2 Jintao (Wuzhi 81-1, C6, WIB-C-6, Fruit Marketed Outside
China Under the Names Jingold™ and Kiwita) ............................. 249
6.2.3 Hongyang....................................................................................... 251
6.2.4 Huayou........................................................................................... 252
6.2.5 Jinyan............................................................................................. 252
6.2.6 Cuiyu.............................................................................................. 254
6.2.7 Ganmi No. 1 (Zaoxian).................................................................... 255
6.2.8 Ganmi No. 2 (Kuimi)....................................................................... 256
6.2.9 Ganmi No. 3 (Jinfeng)..................................................................... 256
6.2.10 Chuhong....................................................................................... 257
6.2.11 Wuzhi No. 3.................................................................................. 258
6.2.12 Zesy002 (Gold3)........................................................................... 259
6.2.13 Zesy003 (Gold9)........................................................................... 260
6.2.14 Soreli............................................................................................ 260
6.3 Pollenizer Cultivars..........................................................................261
6.3.1 Tomuri............................................................................................ 261
6.3.2 Matua............................................................................................. 261
6.3.3 Chieftain......................................................................................... 262
6.3.4 Moshan No. 4................................................................................. 262
6.4 Cultivar of A. eriantha......................................................................262
6.4.1 Huate (White)................................................................................ 262
7 Cultivation and Management........................................................ 265
7.1 Kiwifruit Plant Propagation...........................................................265
7.1.1 Seedling Propagation..................................................................... 265
7.1.2 Grafting.......................................................................................... 266
7.1.3 Vegetative Propagation.................................................................. 268
7.1.4 Tissue Culture Micropropagation.................................................... 268
7.2 Culture and Management...............................................................269
7.2.1 Orchard Establishment................................................................... 270
7.2.2 Orchard Management.................................................................... 272
7.2.3 Training and Pruning...................................................................... 275
7.2.4 Management of Flowers and Fruit.................................................. 279
7.3 Pests, Diseases, and Control...........................................................280
7.3.1 The History of Pests and Diseases on Kiwifruit................................ 280
7.3.2 Important Diseases and Their Control............................................. 281
7.3.3 Major Pests, Prevention, and Control.............................................. 292
8 Harvest and Storage........................................................................ 297
8.1 Harvest and Postharvest Handling...............................................297
8.1.1 Harvesting...................................................................................... 297
8.1.2 Postharvest Handling..................................................................... 298
8.2 Storage..............................................................................................302
8.2.1 Storage Methods............................................................................ 302
8.2.2 Main Factors in Storage and CA Storage Parameters....................... 305
8.2.3 Storage Diseases............................................................................. 306
Bibliography........................................................................................ 309
Index........................................................................................................ 329
Contributors

HUANG Hongwen

In Association with
ZHONG Caihong WANG Shengmei

LI Xinwei GONG Junjie

LI Dawei SHI Tao

JIANG Zhengwang ZHANG Lei

LIU Yifei LI Li

LI Zuozhou JIA Shuowei

YAO Xiaohong CHEN Bin

Foreword
I first started working on kiwifruit about 1975. Kiwifruit
were then a very minor crop. There were perhaps 1000 ha
of commercial kiwifruit orchards in New Zealand, most
recently planted, and as New Zealand was then the only
country with significant plantings, this meant there were
about 1000 ha of commercial kiwifruit orchards through-
out the world. Kiwifruit were still a uniquely New Zealand
crop, and the kiwifruit in world trade all came from
New Zealand.
Scientists in New Zealand knew the kiwifruit as the
“Chinese gooseberry” since the new name of “kiwifruit,”
first devised for the export trade, had not yet achieved uni-
versal acceptance. We had a collection of some of the old
kiwifruit cultivars—although by this time “Hayward” was
becoming dominant—and representatives of a few Actinidia
taxa obtained from the nursery firm of Hillier and Sons in
the United Kingdom: several genotypes of A. arguta, in-
cluding one then classified as A. arguta var. purpurea, and
single male plants of A. callosa var. henryi, A. hypoleuca,
and A. melanandra.
We knew that kiwifruit came from China—hence
the old common name Chinese gooseberry—but little
more. Reading the taxonomic treatment of Actinidia by Li
Hui-Lin made it obvious that there were many species and
varieties in China, and that the diversity in the genus was
much greater than suggested by the few taxa we had seen.
However, the literature on the genus was very scattered, of-
ten in journals that were not available to us, and frequently
in languages we could not read. Hence we had a very nar-
row view of the kiwifruit and its relatives.
The scientific research was also very narrow and lim-
ited. Almost all the research on kiwifruit as a crop plant was
carried out in New Zealand and by only a few people—five
or six of us could meet, and between a late morning tea and
an early lunch we could quickly summarize the priorities
for research and discuss what research was actually being
undertaken, almost always in New Zealand.
Since then, of course, the cultivation of kiwifruit has ex-
panded rapidly so that today the area throughout the world
of commercial kiwifruit orchards is nearly 150,000 ha and
total annual production is about 2.5 million tons. Kiwifruit
are no longer a uniquely New Zealand crop but are culti-
vated in a number of countries. In 1975, kiwifruit in China
were almost entirely collected from the wild. Today, China
has more kiwifruit orchards and produces more kiwifruit
than any other country.
Furthermore, the kiwifruit available to consumers are
no longer all brown and hairy on the outside and green
inside. Different cultivars are appearing in the marketplace
with a range of flesh colors—green, yellow, or red—and
with rather different flavors. Many of the older cultivars
were simply selections from the wild, but increasingly the
newer cultivars are the result of planned breeding programs,
sometimes even involving interspecific hybridization.
As kiwifruit cultivation and production have expanded,
so too has the research on kiwifruit and the other Actinidia
species. This year (2014), the 8th International Symposium
on kiwifruit will be held at Dujiangyan in Sichuan. This is
the second time that the symposium has been held in China
and it emphasizes the greatly increased importance and
sophistication of Chinese research on kiwifruit. A good ex-
ample is the work that has resulted in a much better under-
standing of the relationships between taxa in the Actinidia
chinensis complex and the gene flow between them.
xii Foreword
There have been several books and many detailed
reviews of different aspects of kiwifruit biology and cul-
tivation. Kiwifruit: The Genus ACTINIDIA is the most
welcome addition to the literature, as it brings together
knowledge from both China and the rest of the world and
also, most importantly, takes into account the taxonomic
and nomenclatural changes.
Kiwifruit: The Genus ACTINIDIA demonstrates just
how much we have learned over the past 40 years. From
knowing almost nothing of the wild kiwifruit, we now have
detailed and comprehensive information on the different
Actinidia taxa, their distributions, the relationships between
them, and their commercial potential. The many illustra-
tions will also be of great assistance.
Professor Huang Hongwen and his colleagues are to
be congratulated on their notable achievements. I can only
regret it was not available when I first started my work.
Kiwifruit: The Genus ACTINIDIA is essential reading for
anybody interested in kiwifruit and the other Actinidia spe-
cies. It provides a secure foundation for further research,
as there is still a great deal we have yet to learn about this
important crop.
Ross Ferguson
The New Zealand Institute for Plant & Food Research Ltd
Auckland, New Zealand
Preface
Kiwifruit is a most successful paradigm of plant domes-
tication in the twentieth century, from a wild fruit to current
worldwide commercial cultivation in only 110 years. I am
proud to say that the kiwifruit (Actinidia chinensis) and most
species of genus Actinidia are endemic plants to my home-
land. My research in Actinidia as an Actinidia germplasm
hunter and breeder began in 1979 and took me through
35 years of experience and progress. This includes partici-
pating in a national Actinidia germplasm survey, collection
and inventory, capacity building repositories, germplasm
evaluation and breeding and commercialization of the kiwi-
fruit industry in China. I am most grateful for the guidance
of my senior teachers and the support from many colleagues.
My long-term dedicated efforts in many studies from ger-
mplasm evaluation and exploration, repository manage-
ment, population genetics, and breeding to fundamental
biological questions not only progressively d
­ eepended my
understanding on genus Actinidia, but also rendered per-
sonal enjoyment and contentment. The history of kiwifruit
domestication dated back to 1904 when Actinidia chinensis
var. deliciosa was introduced into New Zealand from China.
Initial domestication was accomplished in New Zealand
and worldwide commercial production began in 1970s by
a single kiwifruit cultivar “Hayward.” As for Actinidia chin-
ensis var. chinensis, tremendous efforts by Chinese kiwifruit
researchers on native germplasm exploration and breeding
successfully domesticated this variety for commercial pro-
duction during the past 35 years. These efforts and accom-
plishments are highly valued worldwide and are a source of
pride for Chinese Actinidia researchers.
I had an experience during my Ph.D. training and
practice in plant genetics and breeding in the United States
from the late 1980s to the 1990s when I marveled at vast
literatures in the university libraries. I became deeply
interested in the many classic works of Burbank (1849-
1926) and Michurin (1885-1935), two great masters of
plant breeding, especially their working notes on exploring
plant natural resources and breeding ideas. Those works
profoundly impressed me with their ways of thinking and
breeding approaches. Thus, the starting point of any plant
domestication and breeding for genetic improvement is
fundamentally based on in-depth understanding and ac-
quired knowledge of basic biology, variation patterns and
processes, and interactions between genetics and the envi-
ronmental. Basic knowledge such as taxonomy, natural dis-
tribution, breeding system, inheritance, selection efficiency,
and hands-on practices are critically important for plant
breeders. China is the second richest country for plant
diversity including many native crop plants. “Learn from
history, learn by analogy” should help propel us forward.
In the late 1990s, I became a principal investigator and
Ph.D. advisor, and began my responsibility of strategic plan-
ning for the multidirectional research of this Chinese en-
demic genus. Thanks to the previous experience in natural
resource surveys guided by my teachers and my footprints
on almost all main natural distribution ranges, although
I still felt that “A leaf before the eye shuts the sight of Mount
Tai,” my expedition and observation notes are quite exten-
sive. Moreover, after many research programs implemented,
progresses achieved, and many graduate students trained,
the information database on Actinidia has been enlarged
extensively, broadening our understanding and knowledge.
Since 2000, I have edited six volumes (Advances in Actinidia
Research I-VI), and published a number of research papers
xiv
and reviews. In the past 10 years, my colleagues and I have
been recognized nationally and internationally in research
on Actinidia resources and taxonomy, population genetics,
breeding, and cultivar improvement. Thus, a monograph
of the genus Actinidia became a priority of my agenda.
However, besides my long-term research in Actinidia,
I have also been a director of botanical gardens for the past
18 years, along with heavy workloads in routine adminis-
tration. Hence, the writing of this monograph was on and
off since 2004 when I drafted the outline. Completion of
the monograph took much longer and was more difficult
than I had initially expected. As such, it took 10 years to
completion owing to indomitable and tireless efforts, in-
cluding those of my students and colleagues. Nonetheless, it
was worth “spending 10 years grinding this sword.”
This book comprises eight chapters directed towards the
purpose of being a special genus monograph of its typical
characteristics of Chinese endemic genera, i.e., the genus
centering in China that is remarkably distributed through-
out much of eastern Asia. From just south of the Equator
in the tropics, to cold temperate regions as far north as lat-
Preface
itude 50° N, the distribution pattern is relatively common
among endemic higher plants in China, representing a con-
stituent of both the Holoarctic and Paleotropic floras. The
book provides thorough information and detailed data of
evolutionary origin, systematics and taxonomy, species de-
scription and characteristics, natural distribution and nat-
ural reserve, domestication and cultivation history, genetic
variation and breeding improvement, cultivars and the ki-
wifruit industry, etc. I have purposely shortened cultivation
and industry aspects of common fruit tree books due to my
designing of the book. I had helped and coauthored with
the late famous kiwifruit expert Mr. Cui Zhixue for his chief
edited book, Actinidia in China, published in 2002, and it
is still largely regarded as the “bible” of genus Actinidia by
many young researchers. But due to lack of information and
data integrity, I felt it has not been regarded as a world-class
monograph for ­genus Actinidia. Publication of Kiwifruit:
The Genus ACTINIDIA has fulfilled my long efforts and
aspiration in this regard, and I hope the monograph will
provide overall and comprehensive information and data
for kiwifruit researchers and botanists in the world.
HUANG Hongwen
Guangzhou, China, March 28, 2014
Acknowledgments
This book, since the first guidelines was drafted, has
taken 10 years to make a reality and involved the great
generosity and support of many people, to whom we are
indebted. First of all, we must thank our senior professors
and colleagues who worked or have been working at the
Actinidia germplasm repository of Wuhan botanical gar-
den in their tenures and made tremendous contributions
of accumulated data and information valuable to the book.
We express our heartfelt thanks to: HUANG Renhuang,
WU Xianwei, HE Zican, LI Jianqiang, ZHANG Zhonghui,
HONG Shurong, SONG Yuanzhen, XU Liyun, HUANG
Hanqian, PENG Fusong, XIONG Zhiting, KE Shanqiang,
CHEN Xuzhong, HU Youmin, ZHANG Suru, ZHANG
Shengju, KANG Ning, CHEN Qian, LI Zheng, WANG
Yehua, WANG Yanchang, YE Qigang, and ZHANG Hong.
This book also benefited greatly from contributions of
many of my students during their studies in past 10 years.
We would like to thank Dr. A.R. Ferguson, a world
authority of kiwifruit, who has devoted many months to
editing the book thoroughly in English and has provided
most valuable advice, suggestions, and outstandingly thor-
ough checking of data, information, and facts used in the
text. He has also provided some photographs. His help has
improved our book and we are most grateful.
Special thanks to CUI Xuechen, JIANG Jingkui, YU
Zhongshu, ZENG Hua, and many others for their generos-
ity of providing information and photographs.
During the endeavor to make Kiwifruit: The Genus
ACTINIDIA as up to date and as accurate as possible,
many experts and individuals from academic institutions,
universities, and kiwifruit enterprises have kindly pro-
vided assistance and help: Guangxi Institute of Botany of
Chinese Academy of Sciences, Institute of Horticultural
Crops of Yunan Academy of Agricultural Sciences,
Institute of Fruit and Tea of Hubei Academy of Agricultural
Sciences, Zhengzhou Fruit Research Institute of Chinese
Academy of Agricultural Sciences, Institute of Botany,
Chinese Academy of Sciences, Horticultural Institute of
Zhejiang Academy of Agricultural Sciences, Horticulture
College of Northwest A&F University, Horticulture &
Landscape College of Hunan Agricultural University,
Horticultural Institute of Hunan Academy of Agricultural
Sciences, College of Horticulture and Forest, Huazhong
Agricultural University, College of Horticulture and Art,
Jiangxi Agricultural University, Lushan Botanical Garden,
Chinese Academy of Sciences, Sichuan Province Natural
Resources Science Academy, Institute of Special Animal
and Plant Sciences of Chinese Academy of Agricultural
Sciences, Jishou University, Horticulture College of Anhui
Agricultural University, Zhongkai University of Agriculture
and Engineering, Sichuan Zhongxin Agricultural S&T com-
pany limited, Beijing Hualin Agricultural S&T company
limited and etc. For their help, we express our heartfelt
gratitude, and we treasure the cooperative spirit within the
Chinese Actinidia research community.
 Introduction
In China, the genus Actinidia Lindl. is known by the name
mihoutao (macaque monkey peach). Throughout the rest of
the world, the most widely used commercial name is kiwi-
fruit. The process of domestication of the kiwifruit started at
the beginning of last century, little more than 100 years ago.
The fruit has been widely accepted because of its unique fla-
vor, its high content of vitamin C, dietary fiber, and a variety
of mineral nutrients, and for its beneficial effects on human
health owing, for example, to its high content of antioxidants
and its alleviation of gastrointestinal illnesses. Domestication
and commercial cultivation of kiwifruit is considered to be
one of the most spectacular examples of successful plant
­domestication in the twentieth century.
Since the first commercial kiwifruit orchard was
e­ stablished in New Zealand by the 1930s, following the
first introduction of Actindia chinensis var. deliciosa into
New Zealand in 1904 (Ferguson and Bollard, 1990), kiwi-
fruit plantings and production have increased, especially
since the late 1970s when commercial production spread
throughout over the world. Today, worldwide kiwifruit
plantings have reached more than 165,000 ha (412,500
acres) and annual production of 2 million tons (Belrose
Inc., 2013).
Over the past 30 years, China has carried out extensive
surveys of the natural resources of the genus Actinidia and
has initiated many studies in taxonomy and systematics,
biogeographical distribution, species and genetic diversity,
habitat ecosystems, etc. The exploration of natural germ-
plasm resources and, more recently, breeding programs have
resulted in the release of many new cultivars to support the
development of the kiwifruit industry. Simultaneously, the
kiwifruit industry in China has developed rapidly, from
essentially nothing in 1978 to current plantings and annual
production accounting for about 45% and 25%, respectively,
of the world total. China has the richest natural resources of
kiwifruit and now it also produces more kiwifruit commer-
cially than any other country. There is no doubt that China
will continue to play an increasingly important role in the
conservation and sustainable use of Actinidia natural re-
sources as well as becoming a more important player in the
world kiwifruit industry in the future.
Kiwifruit: The Genus ACTINIDIA. http://dx.doi.org/10.1016/B978-0-12-803066-0.09999-8
Copyright © 2016 China Science Publishing & Media Ltd. Published by Elsevier Inc. All rights reserved.
☆
Taxonomy, Distribution, and Germplasm
Resources
The genus Actinidia belongs to the family Actinidiaceae.
According to the latest revision (Li et al., 2007a), the ge-
nus has 54 species and 21 varieties, a total of about 75 taxa.
Current commercial cultivation is almost entirely based
on A. chinensis var. chinensis and A. chinensis var. deliciosa
(A. Chevalier) A. Chevalier.
Since Lindley (1836) erected the genus, it has under-
gone four major revisions: Dunn (1911), Li (1952), Liang
(1984), and Li et al. (2007a,d). Early accounts of the genus
separated it into four sections: Leiocarpae (subsequently
split into two series Lamellatae and Solidae), Maculatae,
Stellatae (subsequently split into two series Perfectae and
Imperfectae), and Strigosae. However, these attempts to
subdivide the genus were not satisfactory because indi-
vidual species are often variable, are not readily separated
and the characters used to subdivide the genus were them-
selves not always consistent (Huang and Ferguson, 2007a).
Furthermore, many new taxa were discovered during the
national survey of wild Chinese resources and extensive
genetic studies of natural populations made it obvious that
rather imprecise traits, such as degree of hairness, lamella-
tion of stem pith, and fruit shape are not good criteria for
classification. In particular, the treatment of the section
Strigosae was unsatisfactory. Before the revision of Li et al.
(2007a,d), 76 Actinidia species and 50 sub-specific taxa had
been described (Huang and Ferguson, 2007a), resulting in
considerable confusion. More recent studies by cluster anal-
ysis using both morphological traits and molecular markers
have shown that either dendrogram tree or strict consensus
tree analysis of phylogenetic relationships tends to group
most species into geographic or regional groupings of
northern China, the Yangtze River region, southern China,
southeast China, and southwest China. The revision by Li
et al. (2007a,d) of Actinidia ignored sectional groupings
and clarified the status of some poorly described taxa, even
if further revision is probably required: in total, 17 species,
33 varieties, and 4 forms were combined or treated as syn-
onyms by Li et al. (2007a,d).
 2 Kiwifruit: The Genus ACTINIDIA
The genus Actinidia has a remarkably wide geographic
distribution in eastern Asia, extending from the equator
(tropics) to cold temperate regions as far north as latitude
50° (Liang, 1983; Ferguson, 1990a). Such a distribution
pattern is relatively unusual among higher plants, and it
means that Actinidia is a component of the Holoarctic flora
as well as the Paleotropic flora. However, this pattern is
typical of many Chinese endemic plant genera, i.e., cen-
tered in mainland China but extending to the neighboring
countries. Two Actinidia species occur only in adjacent
countries (Actinidia strigosa Hook. f. et Thoms. in Nepal,
Actinidia hypoleuca Nakai in Japan) and a small number
of Actinidia taxa are found in other countries as well as
China. However, the vast majority of Actinidia taxa are en-
demic to China. There, they occur mostly in the mountains
and hills of south central and southwest China with the
QinLing Mountains forming a northern boundary and the
HengDuan Mountains forming a western boundary. From a
biogeographical viewpoint, the distribution pattern is gen-
erally from southwest to northeast China, structured into
six biogeographical regions:
• southwest China (including Yunnan, Guizhou,
Sichuan, western and southern Tibet);
• southern China (including Guangdong, Hainan,
Guangxi, and southern Hunan);
• central China (including Hubei and eastern
Sichuan, Chongqing, western Hunan, southern and
southwestern Henan, southern Gansu, Anhui, and
southern Shaanxi);
• eastern and south eastern China (including Jiangsu,
Zhejiang, Jiangxi, Fujian, and Taiwan);
• northern China (Hebei, Shandong, Shanxi, Beijing,
and Tianjin); and
• n o r t h e a s t e r n C h i n a ( L i a o n i n g , Ji l i n , a n d
Heilongjiang).
Wild kiwifruit are very abundant. In a survey of wild
kiwifruit in the early 1980s, more than 160,000 tons of fruit
were collected from the wild each year.
A Century of Kiwifruit Domestication
and Industry Development
Although China is the center of origin of the genus
Actinidia and it is the richest in natural Actinidia resources,
few serious attempts at domestication of the genus were
made until it had been successfully domesticated outside
China. According to ancient texts sporadic attempts had
been made. Xin (1983) identified the “Chang Chu” recorded
more than 2000 years ago in the Shijing (a book written be-
tween 1000 and 500 BC) as Actinidia, described as “growing
in moist places.” A more convincing account in ancient texts
is found in a poem by Cen Shen (714-770 CE) of the Tang
Dynasty; the poem—Taibai Dongxi Zhang Laoshe Notes—
that is sent to his niece describes “In [the] middle of garden,
an Actinidia plant climbing on an arbor over a well,” sug-
gesting that as early as 1200 years ago, an Actinidia vine had
been planted in the courtyard. All the various Bencao (great
pharmacopoeias or encyclopedias) in subsequent dynasties
recorded edible and medicinal uses of Actinidia plants and
fruit. There were even occasional attempts, 200 years ago, by
farmers in Huangyan County, Zhejiang Province to collect
Actinidia plants from the wild and plant them around their
homes (Cui et al., 1993).
The process of kiwifruit domestication and commer-
cial cultivation started in New Zealand in 1904 when a
New Zealand school teacher, Isabel Fraser, visiting Yichang
(Ichang), Hubei Province, obtained the seeds of A. chinensis
var. deliciosa and took them back to New Zealand. These
seeds were given to Thomas Allison who then passed to his
brother Alexander Allison, an amateur horticulturist. Plants
from these original seeds were fruiting by 1910. These were
probably the first fruit of A. chinensis var. deliciosa to be
produced outside China. Many European naturalists and
plant hunters introduced Actinidia from China into Europe
and North America countries during the late nineteenth
and early twentieth centuries, but all subsequent important
New Zealand cultivars of A. chinensis var. deliciosa includ-
ing “Hayward,” “Bruno,” “Allison,” “Monty,” “Abbott,” and
“Gracie” can be traced back to those seeds introduced by
Isabel Fraser (Ferguson, 1983, 2004, 2005; Ferguson and
Bollard, 1990). Indeed, these early selected cultivars from
New Zealand dominated world commercial kiwifruit pro-
duction for more than 70 years until the mid-1980s, when a
number of new cultivars of both A. chinensis var. chinensis
and A. chinensis var. deliciosa were selected and released
for orchard production as the result of national survey of
Actinidia resources and cultivar development organized by
the Chinese Ministry of Agriculture. This resulted in note-
worthy changes in the types of kiwifruit cultivars grown,
which significantly changed the world kiwifruit cultivar
structure.
Historical Glance of A. chinensis Taken
Outside China
At the beginning of the twentieth century, China was
a semifeudal society ruled by the decaying Qing Dynasty.
Many western plant hunters and collectors were therefore
able to exploit China's unique and remarkably rich flora,
taking in particular ornamental and garden plants and eco-
nomic plants.
The First Botanical Specimens
The earliest known A. chinensis specimens, cur-
rently held in the French Natural History Museum, Paris,
were collected by d'Incarville in 1740, but examined only
150 years later (Franchet, 1882). The description of the spe-
cies (Planchon, 1847) was instead based on male flowering
specimens collected by Fortune, probably near Ningbo city,
China in 1845 (Ferguson, 1990a,b,c). The first fruiting spec-
imens to be described were collected near Yichang, Hubei
Province by the plant collector Augustine Henry (1857-
1930) in 1886, and sent to the Royal Botanic Gardens, at
 Introduction
Kew, United Kingdom. These fruit were used to prepare
the first European illustration of A. chinensis published
by the then well-known botanist Daniel Oliver (1830-
1916) (Oliver, 1887). These first specimens from Fortune
and Henry were of what we now know as A. chinensis var.
chinensis.
The First Living Plants of A. chinensis Outside
China
The earliest known living plant of A. chinensis grown
outside China was probably that recorded as being in the
arboretum of Maurice de Vilmorin at Les Basrres, Loiret,
France in 1899 (de Vilmorin and Bois, 1904). The seed
from which the plant was raised was probably collected
in Sichuan in 1898 by the French missionary Père Farges.
Because of its geographic origin, it is likely that this
plant belonged to what is now known as A. chinensis var.
deliciosa.
However, credit for the effective introduction and
domestication of kiwifruit should be largely given to the
well-known British plant hunter Ernest Wilson (Ernest
H. Wilson, 1876-1930). In his four adventurous plant col-
lecting expeditions during 1899-1911, Wilson explored
the flora of western China and collected a great number of
Chinese plants, particularly ornamental plants, and sent
them to nurseries (mainly James Veitch & Sons, Ltd) and
botanical research institutions in the United Kingdom and
the United States of America. The plants he introduced
included A. chinensis, Davidia involucrata, Meconopsis inte-
grifolia, Rosa moyesii, Clematis armandii, Clematis ­montana
var. rubens, and species of Cotoneaster, Primula, Acer,
Rhododendron, Viburnum, Hypericum, Lonicera, Berberis,
Spiraea, Ceratostigma, Daphne, Deutzia, Exochorda, Dipelta,
Forsythia, Kolkwitzia, Philadelphus, Syringa, Magnolia, etc.,
as well as 65,000 plant specimens for herbaria in Western
countries (Hillier, 1976; Brigges, 1993). More than 100
genera and more than 1000 species from China enriched
the gardens of Europe and North America and were sub-
sequently used in programs of ornamental flower breeding
and development.
1900: Wilson shipped A. chinensis seed to the United
Kingdom, and subsequently in 1903 a plant of A. chinen-
sis was exhibited at a show of the UK Royal Horticultural
Society. Photographs in the Veitch catalogue the following
year indicate that the plant belonged to A. chinensis var. de-
liciosa (James Veitch and Sons, Ltd., 1904).
1904: A New Zealand school teacher, Isabel Fraser, on
a trip to her sister, Katie Fraser, who was working in the
Church of Scotland mission in Yichang, directly or indi-
rectly obtained A. chinensis var. deliciosa seed from Wilson.
When Isabel Fraser returned to New Zealand in January
1904, she took with her these seeds (Atrkins, 1948), which
were the origin of the world's kiwifruit industry. The cul-
tivar “Hayward,” still accounting for more than 80% of
3
the total world area planted in kiwifruit, the widely grown
cultivar “Bruno” and other early cultivars such as “Allison,”
etc., are the direct descendants of those seeds (Ferguson
and Bollard, 1990).
In July, 1904, the Plant Introduction Experiment Station
at Chico, California of the U.S. Department of Agriculture
(USDA) received through Hankow (Wuhan) plants of
A. chinensis “obtained on the borders of Yunnan by Mr.
Wilson” (Fairchild, 1913). Two “possibly distinct” plants
were registered as USDA Plant Introductions, Number
P.I. 11629 and P.I. 11630 (USDA, 1907a,b). Subsequent
photographs of the plant P.I. No. 11629 indicate that it was
A. chinensis var. deliciosa.
Brief History of the Development
of the Kiwifruit Industry
1904-1924: Introduction of A. chinensis var. deliciosa to
New Zealand. Plants were mostly sold or exchanged between
nurserymen and amateur plant enthusiasts as gifts. By 1917,
New Zealand nurseries were offering to the public seedlings
of A. chinensis var. deliciosa and by 1924 the new plant was
being widely promoted and provoking increasing interest.
1922-1926: Grafted plants became available for sale. All
members of the genus Actinidia are functionally dioecious
and pollination by a male vine is required for fruit produc-
tion. During in the early domestication period, growers
were initially puzzled as to why a female plant with appar-
ently perfect flowers did not produce fruit. Eventually it
was realized that the female, apparently perfect flowers did
not produce viable pollen. Male and female plants could
not be distinguished morphologically at that stage, but once
they had flowered and their gender determined, grafting
onto seedling rootstocks became the standard practice. This
is still the usual practice in New Zealand but in other coun-
tries, vegetative propagation by cuttings or tissue culture
has become widespread.
1930s: Establishment of the first commercial Actinidia
orchard. Augustine Henry (1893) seems to have been the
first to recommend cultivation. His prophetic comment
was, “This fruit might be much improved by cultivation.”
The United States Department of Agriculture also quoted
Frank Meyer as suggesting the possibility of commer-
cial kiwifruit orchards (USDA, 1922). However, the first
A. chinensis var. deliciosa orchard actually established was
that in Wanganui, New Zealand: initially only 14 plants
were grown, but they were producing good-quality fruit by
the early 1930s. Fruit were soon sent to other towns of New
Zealand and readily sold. More planting followed in other
parts of New Zealand, particularly in the Bay of Plenty.
However, these earliest kiwifruit orchards were very small,
generally less than 1 ha.
1950s: For many years, cultivation remained on a small
scale although good prices for the fruit encouraged planting
 4 Kiwifruit: The Genus ACTINIDIA
by growers and the associated development of growing and
management techniques and postharvest storage procedures
(Ferguson, 2011). As production was beginning to exceed
local consumption, trial shipments of the fruit were sent
to Britain in 1952 and Australia in 1954. Exports increased
steadily; in 1952, only 40 boxes were shipped to Britain, in
1954, 563 boxes, and in 1960 18,700 boxes.
1959: Early western botanists and plant explorers often
commented that the flavor of kiwifruit was reminiscent of
the European gooseberry, Ribes uva-crispi (syn. R. grossu-
laria). A. chinensis var. deliciosa was at one stage given the
name “Ichang gooseberry” after the town of Yichang where
E.H. Wilson over-wintered, but this name was seldom used
and in New Zealand the plant was known as the “Chinese
gooseberry” as early as 1917. This remained the most usual
common name until New Zealand fruit were first exported
to the United States in 1959. The exporting firm Turners &
Growers then proposed the name “kiwifruit” after the
kiwi, a native bird emblematic of New Zealand (Ferguson
and Bollard, 1990). The new name was widely promoted
in western markets, and it soon became accepted both
commercially and in the scientific literature. Ironically, for
many years since, people have mistakenly believed that this
new emerging fruit originated from New Zealand, rather
than from China.
1960-1970s: Kiwifruit commercial production in New
Zealand up to the early 1970s was mainly to meet the re-
quirements of the domestic market, but after 1975 exports
began rapidly to exceed local consumption and by 1980
more than 80% of the fruit production was exported. The
increasing demand resulted in a rapid expansion of plant-
ings and more intensive management accompanied by
technological improvements.
(1) Standardization of cultivars and predominance of
“Hayward”: the New Zealand kiwifruit industry was
based on the introduction from China of a small num-
ber of seed of A. chinensis var. deliciosa. Initially plants
were propagated by seed, but associated with propaga-
tion by grafting was the selection of good-fruited types
during the 1920s and 1930s. These types with large fruit,
good taste, and other commercial values include what
are now known as the cultivars “Hayward,” “Bruno,”
“Allison,” “Monty,” and “Abbott,” although they were
often known by a number of different names and not
clearly identified as such. Comparative studies by the
New Zealand Department of Scientific and Industrial
Research (DSIR) resulted in recognition of some better
types. A number of these were grown commercially.
In 1966, the cultivar “Abbott” accounted for c. 50% of
total kiwifruit plantings, “Hayward” for 25%, “Bruno”
for 20%, and “Monty” for c. 5% (Ferguson and Bollard,
1990). However, as exports increased it was found that
fruit of “Hayward” survived the prolonged transport
by ship to Europe much better than fruit of the other
kiwifruit cultivars then available. As exports became
more and more important, growers accepted that it was
fruit quality and consumer satisfaction that were most
important and “Hayward” with its large, good-flavored
fruit with a remarkable storage life replaced the other
cultivars. By 1968, “Hayward” accounted for half the
kiwifruit plantings, in 1973 95% and in 1985, 98.5%
(Ferguson and Bollard, 1990). By then, only “Hayward”
fruit were accepted for export from New Zealand, and
when other countries started growing kiwifruit, they
too grew “Hayward.” For the next 20 years, the kiwifruit
industry throughout the world, with the exception of
China, relied on “Hayward” as its sole fruiting cultivar.
(2) Improvement of orchard management techniques:
“Hayward” is more difficult to manage and crop than
some of the other kiwifruit cultivars. Reliance on
“Hayward” meant that management techniques had to
be modified to suit its requirements. For example, after
many years of research, kiwifruit scientists and growers
in New Zealand understood that for a dioecious plant
such as kiwifruit, adequate pollination is critically im-
portant for achieving high yields of large, uniform fruit.
As “Hayward” became the only fruiting cultivar grown,
the selection of matching male pollenizers became crit-
ical. Two of the first two males selected were “Matua”
and “Tomuri” but later “Chieftain” was widely planted.
In addition, beehives were regularly brought into or-
chards. Specialized companies providing beehives to
ensure kiwifruit pollination became established in kiwi-
fruit growing areas.
(3) Harvesting, packing, storage, and international trans-
portation: the successful expansion of the New Zealand
export industry was possible only because of an em-
phasis on quality control and the standardization of
harvesting, packing, and storage. The development of
appropriate measures of harvest maturity and of con-
ditions for long-term storage (0 °C) were particularly
important (Ferguson and Bollard, 1990; Zespri, 1997;
Ferguson, 2011).
1970-2012: Rise of worldwide kiwifruit industry.
Commercial planting of kiwifruit started in other coun-
tries, including the United States about 1960 with the first
commercial crops in 1965, Italy in 1966, France in 1967,
and Japan around 1977 (Ferguson and Bollard, 1990).
Commercial production further expanded to Chile in
South America and Iran in Middle East during the 1980s.
Meanwhile, the Chinese kiwifruit industry developed from
nothing to the world's largest over the past 30 years. In
2012, it is estimated that total kiwifruit plantings in China
were c. 75,000 ha, whereas Italy had c. 27,000 ha, New
Zealand c. 14,000 ha, and Chile 12,000 ha (Belrose Inc,
2013). China therefore had just over half of the total world
plantings of kiwifruit, accounting for about one quarter of
total world production. As young orchards mature, the pro-
portion of kiwifruit produced in China can be expected to
increase.
 Introduction
Domestication of A. chinensis var. chinensis
and Development of Industry in China
For various reasons China lagged behind New Zealand
and other western countries in investigating its native
Actinidia resources and in attempting commercial kiwifruit
cultivation in investigating its native Actinidia resources.
More than 50 years ago, in 1955, the Nanjing Botanical
Garden, Chinese Academy of Sciences, grew Actinidia
plants selected from the wild and studies on their biolog-
ical characteristics were initiated; however, the program
was not sustained and the plants introduced into culti-
vation were abandoned. Institute of Botany, The Chinese
Academy of Sciences, Beijing introduced A. chinensis var.
deliciosa from Mt Taibai of the Qinling Mountains, Shaanxi
Province in 1957, and from Mt Funiu, Henan Province in
1961. Cultivation experiments and basic biological studies
followed, including extensive work on morphology, growth
and development, and reproductive biology. Important ba-
sic data were collected especially on propagation techniques
such as seed germination, softwood cuttings, budding,
and top-working allowing the conversion of wild plants to
grafted cultivars. These field experiments continued for
more than 30 years and, although Beijing is climatically un-
suitable for kiwifruit cultivation, the information and the
experience obtained later proved very valuable for the devel-
opment of the Chinese kiwifruit industry. At about the same
time, the Wuhan Botanical Garden, the Lushan Botanical
Garden, the Hangzhou Botanical Gardens, the Northwest
Agricultural College, and other institutes started programs
of kiwifruit introduction and cultivation (Beijing Botanical
Garden, Chinese Academy of Sciences, Zhengzhou Fruit
Institute, Chinese Academy of Agricultural Sciences, 1978;
Huang, 2009a).
Soon after the founding of the People's Republic of
China, Chinese pomologists or horticulturists began a
number of sporadic surveys of China's natural Actinidia
resources. In 1958, the Central China Agricultural College
and the Hubei Fruit Institute conducted extensive survey
of A. chinensis resources in the Wudang mountains and re-
corded the main types of natural variants. A similar effort
by the Nanping Agricultural Institute, Fujian Province in
1959 surveyed natural resources over 14 counties in north-
ern Fujian (Guangze, Shaowu, etc.), and the Huangyan
Citrus Research Institute, Zhejiang Province documented 14
valuable groups of natural ecotypes from a local survey of
wild resources in 1960. More extensive and more thorough
surveys and documentation of natural resources began after
the early 1970s, particularly in Henan, Guangxi, and Hubei
Provinces. In 1975, the Zhengzhou Fruit Institute, Chinese
Academy of Agricultural Sciences initiated an extensive sur-
vey of wild Actinidia resources in cooperation with the Xixia
Forest Institute (Xixia County, Henan Province) and also
with other research institutes during 1976-1978. The survey
first started in Xixia County for purpose of evaluation and
selection of large-fruited genotypes and then extended to
5
the whole of Henan Province to prepare a complete inven-
tory of natural resources and reserves of wild Actinidia. At
the same time, the Guangxi Forest Institute and the Central
China Agricultural College at Yichang conducted useful
preliminary surveys of Actinidia resources in the moun-
tainous areas of Guangxi and Hubei Provinces in 1974 and
1978, respectively, and a number of large-fruited plants were
selected. These efforts provided baseline data and meth-
odologies that directly led to a subsequent national survey.
Using the increasing information accumulated from these
surveys, Xixia County, Henan Province began propagation
of seedlings and cultivation trials in 1977. Xixia County
was thus a pioneer of the infant kiwifruit industry in China
and it greatly promoted the initial steps in the industry's
development.
National Survey of Wild Actinidia Resources
in China and Expansion of the Chinese
Kiwifruit Industry
The national survey of wild Actinidia resources was
initiated at a national Actinidia research forum held at
Xinyang, Henan Province, August 1978, organized by the
Chinese Ministry of Agriculture and the Chinese Academy
of Agricultural Sciences. A group of 45 people from re-
search institutes and universities, and marketing, process-
ing, production and management sectors and representing
the main Actinidia distribution areas of 16 provinces (or
autonomous regions) attended the meeting. The Chinese
Academy of Sciences and the China Supply and Marketing
Cooperative Agency also sent representatives to the meet-
ing and participated in the strategic planning of scientific
research and industrial development; a prelude to China's
national level of scientific research and to the development
of a Chinese kiwifruit industry. The experiences and infor-
mation obtained since 1955 in surveys and the results of at-
tempts at domestication were discussed and summarized at
the meeting. Furthermore, based on the analysis of scientific
Actinidia research and industrial development in foreign
countries, a strategic plan for scientific research in China
for 1978-1985 was suggested with the clear aim of catching
up with the kiwifruit industry in the rest of the world. The
overall research plan emphasized an inventory of Actinidia
resources, selection and breeding, propagation and orchard
techniques, storage and marketing techniques, development
of processed products, possible medicinal uses, with special
emphasis on taking advantage of China's rich wild resources
and their potential contribution to the developing kiwifruit
industry.
Subsequently, a national cooperative group for Actinidia
research was established with the late Cui Zhixue, a well-
known fruit-tree germplasm expert, as general coordinator.
This national cooperative program promoted long-term
and systematic investigation of Actinidia resources. By 1992,
apart from Xinjiang, Qinghai, and Ningxia Provinces, 27
 6 Kiwifruit: The Genus ACTINIDIA
provinces (or autonomous regions) had completed or par-
tially completed surveys of Actinidia resources and a base-
line inventory was obtained. Furthermore, selection and
breeding for cultivar improvement was launched and re-
sulted in more than 1450 superior genotypes selected from
wild populations of A. chinensis var. chinensis, A. ­chinensis
var. deliciosa, and Actinidia arguta (Cui et al., 1993). This
is probably the best example in the modern history of fruit
breeding of cultivar improvement being directly based
on large-scale selection from wild populations of natural
distribution range with the richest native resources. The
cultivars developed from these selected genotypes have had
a profound impact on the types of kiwifruit cultivars grown
throughout the world, as well as on the development of
the Chinese and the world kiwifruit industry over the past
20 years.
Impact of New Cultivar Development
on the Kiwifruit Industry
One of the most important aims of Chinese horticul-
turists in initiating the national Actinidia survey was to
take advantage of the rich natural Actinidia germplasm re-
sources, with selection and breeding focused in particular
on A. chinensis var. chinensis to get fruit superior to those
of the cultivar “Hayward” (A. chinensis var. deliciosa)—­
essentially the only fruiting kiwifruit cultivar then widely
grown commercially anywhere in the world. Subsequently,
over the next decade (from about 1978 to 1990) much ef-
fort was expended on extensive evaluation, experimental
trials, regional tests and pilot orchard trials of the 1450 su-
perior genotypes selected from the wild. By the early 1990s,
a number of new cultivars had been released, mainly of
A. chinensis var. chinensis (46 cultivars) but also some of
A. chinensis var. deliciosa (11 cultivars) with more than an-
other 200 good-fruited selections still under evaluation. At
the same time, kiwifruit commercial cultivation started in
China with 1 ha in 1978, developed rapidly to 4000 ha in
1990, and soared to 40,000 ha in 1996. The cultivars planted
in this rapid expansion were mainly those selected from
the wild, with the important exception being “Hayward,”
introduced from New Zealand. Cultivars of A. chinensis
var. chinensis account for more than 25% of the Chinese ki-
wifruit plantings (Huang and Ferguson, 2003). By 1985, a
number of new Chinese yellow-fleshed kiwifruit cultivars
(A. chinensis var. chinensis) selected from the wild popula-
tions had been released, such as “Wuzhi No. 3” (October,
1985), “Ganmi No. 1” (syn. “Zaoxian”) (November 1985),
“Ganmi No. 2” (syn. “Kuimi”) (November, 1985), “Ganmi
No. 3” (syn. “Jinfeng”) (November, 1985), Lushan xiang
(November 1985), and “Yixiang” (November, 1985).
Thus, the domestication of A. chinensis var. chinensis
was successfully accomplished in China whereas that of
A. chinensis var. deliciosa had occurred in New Zealand
more than 50 years earlier. The domestication of A. chin-
ensis var. chinensis from wild to commercial production
over little more than 20 years has significantly changed the
world kiwifruit industry, especially in the types of cultivar
grown. One of the more obvious changes was the arrival of
yellow-fleshed kiwifruit as well as those with a red center
as all the cultivars of A. chinensis var. deliciosa previously
grown had green fruit flesh. The first yellow-fleshed kiwi-
fruit to enter into international trade was launched in 2000.
New Zealand had also introduced material of A. chinensis
var. chinensis from China through informal channels from
1977 onward and a yellow-fleshed cultivar, “Hort16A,” was
the result of a crossing program between plants from differ-
ent parts of China (Muggleston et al., 1998). “Hort16A” and
indeed “Hayward” are only a few generations removed from
the wild and there is no fundamental difference between
them and the new Chinese cultivars selected directly from
wild populations.
The rise of Actinidia research and the kiwifruit in-
dustry in China have already had and will continue to
have far-reaching impacts on the world kiwifruit industry.
Over the past 30 years, selection from the wild Chinese
genetic resources and breeding programs have resulted in
the cultivar composition of the Chinese kiwifruit industry
being very different to that in the rest of the world. The
yellow-fleshed kiwifruit cultivars developed in China are
beginning to be more widely grown outside China and this
will encourage kiwifruit market and consumption diversi-
fication (Huang and Ferguson, 2003, 2007a). For example,
the cultivar “Jintao” (“Golden peach”; A. chinensis var.
chinensis) is being promoted in Europe and South America
through propagation licensing agreements. Chinese horti-
culturists successfully achieved domestication of A. chin-
ensis var. chinensis from the wild to large-scale commercial
production in little more than 20 years. The parallel de-
velopment of the yellow-fleshed A. chinensis var. chinensis
cultivar “Hort16A” in New Zealand and its release onto in-
ternational markets mean that the world kiwifruit industry
has changed from the situation in which a single cultivar of
A. chinensis var. deliciosa, “Hayward,” with its green-fleshed
fruit, was essentially the only type of kiwifruit available in
most countries to the current situation in which perhaps
10% of the world kiwifruit production is of cultivars of A.
chinensis var. chinensis and 90% of A. chinensis var. deliciosa
cultivars. The recent rapid expansion of plantings of the
yellow-fleshed, red-centered cultivar, such as “Hongyang”
(selected from seedlings of wild populations of A. chinensis
var. chinensis collected in Henan in the 1990s, Wu and Li,
1993; Wang et al., 2003a,b) and the increased production of
its fruit is now changing the potential world kiwifruit mar-
ket into green-fleshed, yellow-fleshed, and red-fleshed fruit,
a more diversified pattern than just the usual green.
World kiwifruit production and research are now un-
dergoing major and profound changes. China, as the home-
land of the genus Actinidia, has been the cradle of cultivars
for the world. During the initial stages of the introduction
and domestication of kiwifruit, China was well behind the
major players such as New Zealand and Italy in both sci-
entific research and commercial production. Even 20 years
 Introduction
ago, Chinese information of Actinidia was either nonexis-
tent or hardly to be found in the scientific literature. Now
China is an acknowledged center of excellence for scientific
studies on many aspects of kiwifruit biology. In a recent
major review on kiwifruit biology (Huang and Ferguson,
2007a), more than a quarter of the references cited were
by Chinese authors, particularly in basic biology, taxon-
omy and systematics, population genetics, ecology, and
habitat and biogeographical biology considered as the re-
search center of the world. The increasing sophistication of
Chinese research on Actinidia is remarkable. Achievements
in exploring Actinidia genetic resources and new cultivar
7
development in China will lead Actinidia research and
contribute significantly to role in sustainable development
of the world kiwifruit industry in future. One example will
suffice: scientists in China were the first to discover races of
diploids and tetraploids among wild populations of A. chin-
ensis var. chinensis, as well as hexaploid and tetraploid
races of A. chinensis var. deliciosa. These discoveries have
greatly enhanced our understanding of the abundant wild
resources of kiwifruit and have allowed the development
of more targeted selection techniques for further cultivar
improvement to produce larger and higher-quality fruit
(Huang and Ferguson, 2007a).
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 C H A P T E R

1
Systematics and Genetic Variation of Actinidia

1.1 Taxonomic Revision 9 1.2.17 Actinidia linguiensis R.G. Li & X.G. Wang and
1.1.1 Actinidia Lindl. 12 A. wantianensis R.G. Li & L. Mo 24
1.1.2 Clematoclethra (Franch.) Maxim. 16 1.2.18 Actinidia longicarpa R.G. Li & M.Y. Liang and A. rubrafilmenta
1.1.3 Saurauia Willdenow 16 R.G. Li & J.W. Li 24
1.2 Treatment of Taxa in the Most Recent Revision of Actinidia 17 1.2.19 Actinidia petelotii Diels and A. rudis Dunn 24
1.2.1 Three Varieties of Actinidia arguta (Siebold & Zuccarini) 1.2.20 Actinidia pentapetala R.G. Li & J.W. Li 24
Planchon ex Miquel 20 1.2.21 Actinidia zhejiangensis C.F. Liang and Actinidia persicina
1.2.2 Actinidia melanandra Franchet, Varieties and Synonyms 21 R.H. Huang & S.M. Wang 24
1.2.3 Actinidia kolomikta (Ruprecht & Maximowicz) Maximowicz 21 1.2.22 Actinidia vitifolia C.Y. Wu and A. rubus H. Léveillé 24
1.2.4 Actinidia tetramera Maximowicz 21 1.2.23 Actinidia rubricaulis Dunn and A. fasciculoides C.F. Liang 24
1.2.5 Actinidia valvata Dunn 21 1.2.24 Actinidia ulmifolia C.F. Liang, A. sorbifolia C.F. Liang,
1.2.6 Actinidia callosa Lindley and A. umbelloides C.F. Liang 21 A. stellatopilosa C.Y. Chang, A. obovata Chun ex C.F. Liang,
1.2.7 The Actinidia chinensis Planchon Complex 21 and A. grandiflora C.F. Liang 25
1.2.8 Actinidia eriantha Bentham 22 1.3 Paleopolyploid Origin, Biogeographic Patterns, and
1.2.9 Synonomy of Actinidia fortunatii Finet & Gagnepain, Reticulate Evolution 25
A. glaucophylla F. Chun, A. henryi Dunn and A. gracilis 1.3.1 Paleopolyploid Origin and Evolution of Actinidia 25
C.F. Liang 23 1.3.2 Biogeographic Patterns and Reticulate Evolution 27
1.2.10 Actinidia fulvicoma Hance, A. cinerascens C.F. Liang and 1.4 Actinidia Palynology 30
A. fulvicoma f. hirsuta (Finet & Gagnepain) C.F. Liang 23 1.4.1 Pollen Size and Morphology 30
1.2.11 Actinidia hemsleyana var. kengiana (F.P. Metcalf) 1.4.2 Phylogenetic Relationships in Actinidia as Revealed by
C.F. Liang and A. kengiana F.P. Metcalf 23 Pollen Morphology 31
1.2.12 Actinidia carnosifolia C.Y. Wu, A. carnosifolia var. glaucescens 1.4.3 Applications of Palynology 32
C.F. Liang, and A. henryi var. polyodonta Handel-Mazzetti 23 1.5 Flower Morphology and Sex Variation 35
1.2.13 Actinidia flavofloris H.Z. Jiang 24 1.6 Ploidy Variation and Breeding Applications 37
1.2.14 Actinidia leptophylla C.Y. Wu, A. maloides H.L. Li and 1.6.1 Basic Chromosome Number 37
A. maloides f. cordata C.F. Liang 24 1.6.2 Ploidy Races and Interploidy Hybridization 37
1.2.15 Actinidia jiangkouensis S.D. Shi & Z.S. Zhang and 1.6.3 Nuclear DNA Content and Flow Cytometry 42
A. laevissima var. floscula S.D. Shi 24 1.6.4 Ploidy Variation for Kiwifruit Breeding 43
1.2.16 Actinidia guilinensis C.F. Liang 24 1.7 Genomic Diversity 43

1.1 Taxonomic Revision
Subdivision of the genus Actinidia has long been
contentious. An early taxonomic treatment by Gilg
(1893) split eight species into two groups, Monanthe and
Pleianthe, based on the type of inflorescence, solitary
or cymose, but he ignored some of the other Actinidia
species then known. His classification was subsequently
shown to be inadequate as it could separate male and
­female plants of an individual species.
Many new Actinidia species were described during the
next two decades and in the first systematic revision of the
genus, Dunn (1911) recognized 24 species, some further
­divided into varieties and forms. He established four sections
Vestitae, Maculatae, Ampullif,erae, and Leiocarpae, based on
the degree of pubescence, the shape of ovary, and the pres-
ence or absence of lenticels on the fruit surface (maculate vs.
immaculate). In the next major revision, Li (1952) empha-
sized the structure of leaf hairs and discarded the ambiguous
characters of ovary shape and degree of ­pubescence as used

Kiwifruit: The Genus ACTINIDIA. http://dx.doi.org/10.1016/B978-0-12-803066-0.00001-0

Copyright © 2016 China Science Publishing & Media Ltd. Published by Elsevier Inc. All rights reserved.
 10 Kiwifruit: The Genus ACTINIDIA
by Dunn. Li divided the section Vestitae into two sections
Stellatae (with stellate hairs) and Strigosae (with simple hairs),
merged the section Ampulliferae into the section Leiocarpae,
and retained the section Maculatae. Li listed 36 species and
14 varieties in China and neighboring countries.
The revision by Liang (1984) considered only the Actinidia
species in China, but these are nearly all of the species in the
genus. He modified Li's division of the four sections by fur-
ther subdividing two series Lamellatae and Solidae within the
section Leiocarpae, taking into account whether the pith was
lamellate or solid, and two series Perfectae and Imperfectae
within the section Stellatae by taking into account the struc-
ture and, abundance of the stellate hairs and whether they
were shed. Subsequently, Li and Li (2010) proposed revision of
some of the section names, replacing section Maculatae with
section Actinidia and section Stellatae with section Vestitae.
Liang's treatment significantly increased the number of
taxa to a total of 51 species, 35 varieties, and 6 forms (Liang,
1984). Subsequently, many inter- and infraspecific variants
were found as the national survey of Actinidia resources
in China progressed during 1978-1992 and about 20 new
species or new varieties were published. By 2005, as many
as 76 species and infraspecific 50 taxa had been described
(Huang and Ferguson, 2007a).
There were, however, serious problems. The infrage-
neric subdivisions were questionable and difficult to use
(Huang et al., 1999; Li et al., 2000). There were also doubts
as to whether many of the new species could be justified. A
number of new species and taxa ignored the fact that where
exists interspecific hybrids in natural populations and sym-
patric distribution of different species in natural ranges.
The ignorance has resulted individually different plants or
ecotypes to be treated as new species and taxa.
The sections Actinidia (formerly Maculatae) and Strigosae
seem to be heterogeneous and many of the characters de-
limiting these groups are found elsewhere in the genus
(Condon, 1991). Thus individual species within the section
Actinidia (Maculatae) may have lamellate or solid pith, and
the lenticellate fruit (possession of which is used to separate
the section Actinidia (Maculatae) from the Leiocarpae) are
also found in the sections Strigosae and Vestitae (formerly
Stellatae). The species within the Strigosae as defined by Liang
lack common morphological features and they have scattered
geographic distributions (Liang, 1983). The need to separate
the Imperfectae from the Perfectae of the Vestitae (Stellatae)
(based on the structure, abundance, and persistence of the
hairs) indicate that possession of stellate hairs is not a par-
ticularly reliable character for subdividing the genus. The
degree of hairiness can be a subjective character and the dis-
covery of new taxa has shown that it is also a relative factor.
Furthermore, there does not seem to be a clear distinction
between simple and stellate hairs, but transitions between
them (e.g., in A. cylindrica var. reticulata and A. sabiifolia, both
placed in the section Actinidia (Maculatae) (Condon, 1991;
He et al., 2000a,b). Solid pith is also found in some species
outside the Solidae, so it too is not a definitive characteristic
and in some taxa seems to vary with the particular specimen
studied (e.g., A. callosa, A. glaucocallosa, A. latifolia).
Detailed morphological comparisons of the species thus
indicate that the infrageneric subdivisions of the genus as
modified by Liang (1984) are not sustainable. Using clus-
ter analysis of 50 morphological characters, Huang et al.
(1999) proposed subdividing the genus into three sections:
Leiocarpae retaining all species with smooth-skinned fruit,
Maculatae including the species with spotted (lenticellate)
fruit and Vestitae, including all species with leaf hairs, which
would be further divided into two series, Stellatae for spe-
cies with stellate hairs on the undersides of the leaves and
Strigosae for the species with simple and coarse leaf hairs. Li
et al. (2000) suggested subdividing Actinidia into two subge-
nera, Leiocarpae and Maculatae, based on cladistic analysis
of 22 morphological characters. Using solely morphological
criteria, the Leiocarpae with their smooth-skinned, hairless
fruit, seem consistently to form a reasonably coherent group-
ing. Phylogenetic analysis based on the microscopic structure
of the leaf hairs likewise suggested that the Leiocarpae is a
monophyletic group but that the Maculatae and Stellatae are
polyphyletic (He et al., 2000a,b). However, the various subdi-
visions as proposed by Huang et al. (1999) and Li et al. (2000)
left the Maculatae very heterogeneous, with individual species
varying in type of leaf hair and frequency of lenticels on the
fruit skin, making it difficult to delimit the sections.
Molecular studies have also confirmed that the subdi-
vision of Actinidia as proposed by Liang is artificial. Most
studies agree with the conclusions from morphology that
the Leiocarpae are, in general, distinct from the remainder
of the genus, whereas the other three sections cannot be
sustained. Isozyme polymorphism in eight enzyme systems
(Testolin and Ferguson, 1997) indicated that three species in
the Leiocarpae, series Solidae: A. macrosperma, A. polygama,
and A. valvata, formed a natural grouping as did three species
in the Leiocarpae, series Lamellatae studied: various forms of
A. arguta, A. hypoleuca, and A. melanandra. The exception
was A. kolomikta, usually placed in the Lamellatae because
of morphological similarities, even though it has yellow an-
thers whereas A. arguta, A. melanandra, and closely related
species all have black anthers. Isozyme patterns suggest that
A. kolomikta is only distantly related to the other species of
the Lamellatae. This separation of A. kolomikta is supported
by studies of leaf flavonoids: A. kolomikta is very different to
the other species within the Leiocarpae studied (A. arguta, A.
hypoleuca, A. macrosperma, A. polygama, and A. valvata) or,
indeed, any other Actinidia species so far examined. RFLP
analysis (restriction fragment length polymorphism analysis)
of four regions of chloroplastic DNA likewise indicates that
A. kolomikta is best treated as distinct from other members
of the Lamellatae (Testolin et al., 1997; Cipriani et al., 1998).
In contrast, however, RAPD analysis (random amplified
polymorphic DNA analysis) supports retaining A. kolomikta
within the Leiocarpae (Huang et al., 2002a; Kim et al., 2003).
 Chapter 1 Systematics and Genetic Variation of Actinidia
A problem with such molecular approaches is that often only
a single representative from a taxon has been examined (Li
et al., 2011) or that the same representative of a taxon is not
necessarily used in different studies.
It is clear that for elucidation of the relationships within the
genus, evidence from a number of different approaches needs
to be combined. Although the accumulated evidence supports
treating the Leiocarpae (apart possibly from A. kolomikta) as
a monophyletic group, the section Maculatae appears to be a
polyphyletic group with most members either as stand-alone
single species or more closely related to individual members of
the Stellatae or the Strigosae than to each other (Huang et al.,
2002a). Conversely, individual species within the Strigosae or
the Stellatae have a closer affinity to species in the other sec-
tions than to the others in the section in which they have been
placed. A consensus is emerging that the relationships between
the different groups of Actinidia taxa are more easily under-
stood if geographic distributions are considered.
The topographic complexity typical of southwestern
and central China, with sites at varying altitudes and micro-
climates at any given latitude influenced by the mountain
chains has probably encouraged speciation in Actinidia.
Geographic barriers are among the most important mech-
anisms involved in the evolution of new plant species (allo-
patric speciation) and it seems the greater the environmental
diversity, the greater the number of Actinidia taxa (Zhao and
Liu, 1996). Ecological separation and geneflow dynamics
within and between species would account for regional clus-
tering of taxa. In the treatment of the genus by Liang (1984),
the various sections had characteristic geographic distribu-
tions, e.g., the Leiocarpae were to be found mainly in north
China, north of the Yellow River. More recent studies involv-
ing use of cluster analysis dendrograms and strict consensus
trees associate many of the Actinidia into geographic regions
corresponding to north China, the Yangzi River region,
south China, southeast China, and southwest China. Thus
Huang et al. (2002a) and Li et al. (2003a) considered that A.
callosa var. strigillosa, A. chinensis var. chinensis, A. chinensis
var. deliciosa, A. grandiflora, A. hubeiensis, and A. lijiangensis
make up a grouping mainly centered on the Yangzi River,
and A. chrysantha, A. cylindrica, A. farinosa, A. glaucophylla
(now treated as probably being in A. fortunatii), A. lianggu-
angensis, A. rufotricha, and probably A. indochinensis another
grouping centered in south China in Guangxi and Guizhou,
extending into Guangdong and Hunan.
Recently, Li et al. (2007a) made a systematic revision
of Actinidiaceae for the new English version of The Flora
of China. They did not attempt a separation of Actinidia
species into infrageneric groupings. They recognized 52
species and 16 varieties of Actinidia with 17 species, 33
varieties, and 4 forms being treated as synonyms, and two
new combinations being proposed.
They concluded that many of the characters previously
used to separate taxa were not sufficiently constant or
11
c­ orrelated with each other. Some of the recently described
new taxa had been based on a single or a very small num-
ber of specimens (Li et al., 2011), possibly anomalous, and
the insufficient attention given to interspecific hybrids in
natural populations and the sympatric distribution of dif-
ferent species had resulted in individually different plants
or ecotypes being treated as new taxa. Some new taxa were
not validly published in that the location of types was not
given. The difficulties are discussed further in Section 1.2.
More work, especially more comprehensive fieldwork, is
required to determine whether all the species so far described
are sufficiently distinct to justify separation at the specific
level or whether, it is more useful to revert to broader species
concepts. Differences between species or between infraspe-
cific taxa are not always large or consistent and there has been
debate as to the level at which the differences observed justify
separation of taxa (Li et al., 2011). A good example would be
A. arguta and closely related taxa (Dunn, 1911; Nakai, 1933;
Li, 1952; Liang, 1984; Li et al., 2007a, 2011). Furthermore,
there may be consistent, if generally small, morphological dif-
ferences between male and female plants of the same taxon,
and there can also be considerable morphological variation
in, for example, leaf size, shape, and pubescence even within
a single plant (Dunn, 1911). Transitional forms suggest
considerable hybridization between taxa with overlapping
geographical distributions (Huang and Ferguson, 2007a; Liu
et al., 2008), and some of the more recently described species
may be natural hybrids, especially those whose descriptions
are apparently based on a single genotype (Li et al., 2003a). It
seems unwise to generalize conclusions from studies of only
one or two individual genotypes of a particular taxon. Species
that are widespread are often polymorphic and rather than
attempt fine distinctions, it might be better simply to accept
that some species may be morphologically quite variable.
Male and female plants of a particular taxon usually cluster
closely in terms of genetic similarity (Li et al., 2003b) as do
varieties within a species, although the lack of clustering of all
varieties in species such as A. callosa and A. fulvicoma, both
recognized as being morphologically polymorphic, indicates
a need to reconsider some species boundaries (Huang et al.,
2002a; Li et al., 2003a,b).
Actinidiaceae 猕猴桃科Mihoutao Ke (based on J.-Q. Li, X.-W.
Li, and D.D. Soejarto. 2007. Actinidiaceae. pp. 334-360. In:
Z.-Y. Wu, P.H. Raven, and D.-Y. Hong (eds.), Flora of China,
vol. 12, Science Press, Beijing; Missouri Botanical Garden
Press, St Louis.)
Trees, shrubs, or woody vines. Leaves alternate, simple,
shortly or long petiolate, not stipulate. Flowers bisexual or
unisexual or plants polygamous or functionally dioecious,
usually fascicled, cymose, or paniculate. Sepals (2 or 3 or)
5, imbricate, rarely valvate. Petals (4 or) 5, sometimes more,
imbricate. Stamens 10 to numerous, distinct or adnate
to base of petals, hypogynous; anthers 2-celled, versatile,
­dehiscing by apical pores or longitudinally. Ovary superior,
disk absent, locules and carpels 3-5 or more; placentation
axile; ovules anatropous with a single integument, 10 or
 12 Kiwifruit: The Genus ACTINIDIA

more per locule; styles as many as carpels, distinct or con- in China. The name kiwifruit is used for commercially im-
nate (then only one style), generally persistent. Fruit a berry portant selections of Actinidia chinensis var. chinensis and
or leathery capsule. Seed not arillate, usually with large em- A. chinensis var. deliciosa. Kiwifruit originated in central
bryos and abundant endosperm. China and are now widely cultivated throughout the world.

Three genera and c. 357 species: Asia and the Americas; For additional information, see the papers by Li et al.
three genera (one endemic) and 66 species (52 endemic) (2007a) and Liang et al. (1984).
1a. Trees or shrubs; flowers bisexual or plants functionally dioecious.................................................................................................... 3. Saurauia ....................

(水东哥属)
1b. Woody vines; flowers bisexual, plants monoecious or dioecious.
2a. Ovary 15-30-loculed and -styled, styles distinct; fruit a berry without a ridge; seed numerous; stamens 15-130.............. 1. Actinidia ...

2b. Ovary 5-loculed and -styled, styles connate; fruit berrylike or a dry leathery capsule, 5-ridged; seed usually 5;...................................
(猕猴桃属) stamens 10.......................................................................................................................................................... 2. Clematoclethra
....................................

(藤山柳属)

pink, red, yellow, or green, bisexual, plants polygamous

1.1.1 Actinidia Lindl. or functionally dioecious. Sepals (2-)5 (or 6), distinct or
猕猴桃属 mihoutao shu connate at base, imbricate, rarely valvate, persistent or
TYPE: Actinidia callosa Lindl. not. Petals (4 or) 5 (or more than 5), imbricate. Stamens
Lindl., Nat. Syst. ed. 2, 439. 1836; Benth. & Hook. f., Gen. Pl. numerous, in functionally female flowers often with
1: 177.1862; Dunn, J. Linn. Soc. Bot. 39: 394. 1911; H.-L. shorter filaments and smaller sterile anthers; filaments
Li, J. Arnold Arbor. 33: 1.1952; C.Y. Wu, Fl. Yunnan 1: slender; anthers yellow, brown, purple, or black, versatile,
54. 1977; C.F. Liang, Fl. Reipubl. Popularis Sin. 49(2): attached at middle, 2-celled, dehiscing lengthwise, usually
196. 1984. J.-Q. Li et al., Fl. China 12: 334. 2007. divaricate at base. Disk absent. Ovary ovoid, cylindrical,
Trochostigma Sieb. & Zucc., Abh. Akad. Wiss. Muench. iii. or bottle-shaped, glabrous or hairy, many loculed; ovules
726, t. 2, f. 2. 1843; Endl., Gen. Pl. Suppl. iii. 94. 1843. numerous per locule; styles as many as carpels (15-30),
Kolomikta Regel, Bull. Phys. Math. Acad. Petersb. xv. 219. 1857. usually reflexed, persistent, radiating, in functionally
female flower elongating after anthesis; rudimentary
Climbing shrubs, glabrous or hairy, indumentum of ovary in functional male flower very small, with minute
stellate or simple hairs; pith solid or lamellate. Branches styles. Fruit a berry, globose, ovoid, or oblong, spotted
usually with linear, lengthwise lenticels; winter buds small, with lenticels or not, glabrous or hairy. Seed numerous,
enclosed in swollen base of petiole or exposed. Leaves oblong, small, immersed in pulp; testa cartilaginous,
often long petiolate; stipules minute, obsolete, or absent; ­reticulate-pitted; albumen copious and abundant; embryo
leaf blade membranous, papery, or leathery, venation pen- comparatively large, cylindrical, straight, at center of albu-
niveined, veinlets reticulate, usually in cross-bars, margin men; cotyledons short.
serrate or dentate, rarely entire. Inflorescences cymose,
axillary, often pseudo-umbellate, few- or many flowered, Fifty-four species in total: occurrence in East and
or flowers solitary; bracts present, minute. Flowers white, Southeastern Asia, 52 species (44 endemic) in China.

1a. Fruit not spotted with lenticels.

2a. Pith solid, white; flowers white; sepals 2-5; petals 5-12.
3a. Sepals (4 or) 5; petals 5; leaves adaxially sparsely strigillose........................................................................................ 33. A. polygama .....................

(葛枣猕猴桃)
3b. Sepals 2 or 3; petals 5-12; leaves adaxially not strigillose.
4a. Fruit ovoid to obovoid, rostrum on apex ± conspicuous; seed c. 3 mm, c. 2.5 mm in diam.; sepals (2 or) 3; petals 5-9;
anthers oblong to linear, 2.5-4 mm.............................................................................................................................. 49. A. valvata
....................

(对萼猕猴桃)
4b. Fruit globose, rostrum on apex inconspicuous; seed 4-5 mm, c. 3 mm in diam.; sepals 2 or 3; petals 5-12; anthers ovoid,
1.5-2.5 mm............................................................................................................................................................ 27. A. macrosperma
........................

(大籽猕猴桃)
2b. Pith lamellate, white or brown; flowers greenish, white, or red; sepals 4-6; petals 5.
5a. Ovary bottle-shaped; flowers white or greenish; apex of fruit rostrate; pith white to brown; leaves abaxially glaucous or not.
6a. Leaf blade abaxially usually not glaucous, ovate to broadly ovate or orbicular, sometimes ovate-oblong, membranous to
papery..................................................................................................................................................................................1. A. arguta
.............................

(软枣猕猴桃)
6b. Leaf blade abaxially usually glaucous, ovate-lanceolate, ovate-oblong, oblong, or ovate, occasionally orbicular, papery to
leathery.................................................................................................................................................................... 28. A. melanandra
........................

(黑蕊猕猴桃)
Another random document with
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„Delcommune“ als erledigt an. Die Besatzung von „Hedwig von Wißmann“
überraschte am 27. Februar 1915 einen belgischen Posten bei Tembwe und
erbeutete dessen Maschinengewehr. Ein belgischer Offizier und 10 Askari fielen,
ein schwer verwundeter belgischer Offizier und ein Engländer wurden gefangen.
Bei uns fiel ein Askari, ein Europäer wurde tödlich, ein Askari schwer verwundet.
Im März 1915 nahmen die Belgier in Ubwari, dessen Bewohner sich deutsch-
freundlich gezeigt hatten, Massenverhaftungen vor und hängten eine Anzahl Leute
auf.
Nach aufgefangenen Funksprüchen sind im Juni auf dem Tanganjika mehrere
belgische Walfischboote fertiggestellt gewesen; an einem neuen belgischen
Dampfer, dem „Baron Dhanis“, wurde gearbeitet. Deutscherseits wurde am 9. Juni
1915 Dampfer „Goetzen“ fertiggestellt und von der Truppe übernommen. Er hat
bei den Truppenverschiebungen auf dem Tanganjika wertvolle Dienste geleistet.
Bei Bismarckburg war die dortige Polizeitruppe unter dem tüchtigen Verwalter des
Bezirks, Lt. d. Res. Haun, zur Schutztruppe übergetreten. Es kam zu kleineren
Scharmützeln auf feindlichem Gebiet, und auch hier gelang es, den Feind im
wesentlichen fernzuhalten.
Erst Anfang Februar 1915 rückten mehrere hundert feindliche Askari in
Abercorn ein, und Teile derselben drangen bis in die Gegend der Mission Mwasye
vor, zogen dann aber wieder ab.
Mitte März wurde dann die Truppe unter Lt. d. Res. Haun am Kitoberge durch
eine englisch-belgische Abteilung im Lager überfallen. Lt. Haun geriet schwer
verwundet in Gefangenschaft und mehrere Askari fielen. Oberleutnant Aumann
wurde mit einer Abteilung, die später als Kompagnie formiert wurde, von
Hauptmann Falkenstein, dem Führer der 5. Feldkompagnie (Langenburg),
abgezweigt und deckte in der Gegend von Mbozi die deutsche Grenze. Dort waren
im Februar 1915 mehrfach mehrere hundert Mann starke Abteilungen in
deutsches Gebiet eingedrungen; Ende März wurden in Karonga Europäer in
unbekannter Zahl, in Fife und in anderen Orten der Grenze etwa 800 Mann
gemeldet. Der Feind schien also einen Angriff vorzubereiten. Er streifte bis in die
Gegend von Itaka vor, und Anfang April wurde Kituta am Südende des
Tanganjikasees als von den Belgiern verschanzt gemeldet. Major von Langenn,
der nach Wiederherstellung von seiner schweren Verwundung — er hatte ein
Auge verloren — am Russissi tätig war, wurde mit der Führung der Operationen in
dem ihm bekannten Gebiet Bismarckburg-Langenburg betraut. Außer seiner
früheren 5. Feldkompagnie, die bei Ipyana und in der Gegend von Mbozi stand,
wurden ihm hierzu die etwa kompagniestarke Abteilung Bismarckburg und drei
Kompagnien unterstellt, die von Kigoma und Daressalam herangezogen wurden.
Während des Seetransportes nach Bismarckburg fanden östlich dieses Ortes
einige erfolgreiche Zusammenstöße unserer Patrouillen gegen 50 bis 250 Mann
starke feindliche Streifabteilungen statt.
 Major von Langenn hatte am 7. Mai 1915 4 Kompagnien bei Mwasye
versammelt, eine gegenüberstehende belgische Abteilung ging zurück. Am 23.
Mai warf Patrouille Oberleutnant von Debschitz eine belgische Kompagnie zurück,
von der 2 Europäer, 6 Askari fielen. Am 24. Mai erging Befehl an Langenn, mit 3
Kompagnien nach Neu-Langenburg gegen den dort als bevorstehend gemeldeten
Angriff abzurücken. Den Befehl in Gegend Bismarckburg übernahm General
Wahle. Dieser traf am 6. Juni in Kigoma ein und sammelte bei Bismarckburg die
als 29. Feldkompagnie formierte Abteilung Bismarckburg und die von Daressalam
herangezogene 24. Feldkompagnie und halbe Europäerkompagnie.
Am 28. Juni griff General Wahle mit 2½ Kompagnien die Farm Jericho an,
brach aber das Gefecht ab, als er erkannte, daß die feste Stellung ohne Artillerie
nicht zu nehmen sei. Bei uns fielen 3 Europäer, 4 Askari, verwundet wurden 2
Europäer, 22 Askari. General Wahle wurde durch 2 Kompagnien von Langenburg
her verstärkt.
Seit dem 25. Juli 1915 belagerte General Wahle mit 4 Kompagnien und 2
Geschützen C/73 den bei Jericho stark befestigten Gegner. Von Abercorn aus
unternommene Entsatzversuche wurden abgeschlagen, am 2. August 1915 aber
die Belagerung aufgehoben, da mit der vorhandenen Artillerie eine Wirkung nicht
zu erzielen war. General Wahle fuhr mit 3 Kompagnien zurück nach Daressalam.
Die 29. Kompagnie blieb bei Jericho, die 2 Geschütze in Kigoma.
Am 19. Juni war durch „Goetzen“ der bei Kituta auf Strand liegende Dampfer
„Cecil Rhodes“ abgeschleppt und versenkt worden.
Während des September und Oktober kam es nun zu dauernden
Patrouillengefechten an der Grenze von Bismarckburg; bei Abercorn drangen
wieder belgische Verstärkungen ein. Am 3. Dezember wurde bemerkt, daß die
Befestigungen von Jericho verlassen und geschleift waren. Ein neues, nordöstlich
Abercorn erbautes Fort beschoß Oberleutnant Franken am 6. Dezember mit 100
Gewehren und einem Maschinengewehr und brachte dem Feinde dabei
anscheinend Verluste bei.
Die englische Marineexpedition, deren Anmarsch über Bukama-Elisabethville
seit langem beobachtet wurde, hatte am 22. Oktober 1915 die Lukugabahn
erreicht. Die aufgefangenen Notizen, daß für die Deutschen eine Überraschung
auf dem Tanganjika vorbereitet würde, brachten mich auf den Gedanken, daß wir
hier mit besonders konstruierten kleinen Fahrzeugen, die vielleicht mit Torpedos
ausgerüstet wären, zu rechnen haben würden. Es handelte sich also um eine sehr
ernst zu nehmende Gefährdung unserer Herrschaft auf dem Tanganjika, die auf
unsere gesamte Kriegführung von ausschlaggebendem Einfluß sein konnte. Die
gleichzeitig mit diesen Vorbereitungen stattfindenden feindlichen
Truppenverschiebungen in der Richtung auf den Kiwusee und auf Abercorn zu
bewiesen, daß Hand in Hand eine beabsichtigte Landoffensive gehen sollte. Um
hierbei den Feind möglichst noch während seiner Versammlung zu schlagen, griff
Hauptmann Schulz am 27. September 1915 bei Luwungi die Belgier an und
brachte ihnen schwere Verluste bei.
Der Dampfer „Kingani“ überfiel in der Nacht vom 28. Oktober eine belgische
Telegraphenbaukolonne und machte einige Beute. In der Lukugamündung wurde
ein fahrender Eisenbahnzug festgestellt. „Kingani“ kehrte von einer
Erkundungsfahrt zur Lukugamündung nicht zurück und war nach einem
belgischen Funkspruch vom 31. Dezember verlorengegangen. Vier Europäer, acht
Farbige sollen tot, der Rest gefangen sein. Augenscheinlich war der günstige
Zeitpunkt, die feindlichen Vorbereitungen zur Herrschaft auf dem Tanganjika zu
stören, verstrichen.
Am 9. Februar 1916 wurde dann noch einer unserer armierten Dampfer durch
den Feind genommen.
Auf dem Nyassasee war der deutsche Dampfer „Hermann von Wißmann“, der
vom Ausbruch des Krieges nichts wußte, am 13. August 1914 von dem englischen
Regierungsdampfer „Gwendolin“ überrascht und fortgenommen worden.
Hauptmann von Langenn war mit seiner in Massoko bei Neu-Langenburg
stehenden 5. Feldkompagnie am 9. September 1914 gegen die englische Station
Karonga vorgegangen. Beim Kampfe gegen die in fester Stellung befindlichen
Engländer wurde Hauptmann von Langenn selbst schwer verwundet. Die beiden
Kompagnieoffiziere gerieten, gleichfalls schwer verwundet, in englische
Gefangenschaft. Die deutschen Unteroffiziere und die Askari schlugen sich sehr
brav, mußten aber doch einsehen, daß sie gegen die Schanzen des Feindes
nichts ausrichten konnten und brachen deshalb das aussichtslose Gefecht ab.
Über 20 Askari waren gefallen, mehrere Maschinengewehre und leichte
Geschütze verlorengegangen. Aus Iringa und Ubena trafen nun umgehend
Verstärkungen der 2. Kompagnie ein; auch mehrere hundert Wahehehilfskrieger
wurden aufgeboten. Nach und nach stellte sich heraus, daß der Feind auch starke
Verluste erlitten hatte. Er hütete sich vor größeren Unternehmungen gegen den
Bezirk Langenburg, so daß dieses fruchtbare, für uns so wichtige
Verpflegungsgebiet uns anderthalb Jahre lang erhalten blieb.
Später rückte unsere 5. Kompagnie bei Langenburg mit ihrem Hauptteil wieder
näher an die Grenze zur Mission Ipyana vor. Am 2. November 1915 fand am
Lusirafluß ein Vorpostengefecht statt und dem Dampfer „Gwendolin“ auf dem
Nyassasee wurden einige Geschütztreffer beigebracht.
Anfang Dezember 1914 fanden nördlich Karonga, am Ssongwefluß,
Patrouillenzusammenstöße statt. Oberarzt Dr. Gothein, der Anfang Mai 1915 aus
englischer Gefangenschaft zurückgeliefert worden war, erzählte, daß in dem
ersten Gefecht bei Karonga, am 9. September 1914, der Feind an Toten 6
Europäer und 50 Askari, an Schwerverwundeten 7 Europäer und über 50 Askari
verloren hatte. Die Engländer unterhielten eine rege Spionage, besonders durch
den „Wali“, einen eingeborenen Verwaltungsbeamten am Ssongwe.
An der Grenze kam es im Mai 1915 zu einigen für uns günstigen Überfällen.
Die Regenzeit verzögerte sich, so daß der südliche Teil des Bezirkes Langenburg
bis Ende Juni gegen einen Angriff als geschützt gelten konnte.
Im Juni 1915, als Major von Langenn mit seinen Verstärkungen eingetroffen
war, kam es entgegen der Erwartung zu keinen größeren Gefechten. Die Zeit
wurde benutzt, um auf englischem Gebiet einen Telegraphen ab- und auf
deutschem Gebiet in Richtung auf Ubena wieder aufzubauen. Auch im August
bewahrheitete sich die Nachricht von einem geplanten feindlichen Angriff nicht.
Am 8. Oktober erst trafen stärkere feindliche Truppen, Europäer und Askari, in Fife
ein. Auch an dieser Grenze kam es zu zahlreichen kleinen Scharmützeln. Gegen
Ende des Jahres wurde das Eintreffen neuer Verstärkungen auch bei Ikawa
festgestellt. Hauptmann Aumann wies am 23. Dezember 1915 dort den Überfall
einer feindlichen Abteilung von etwa 60 Europäern und zwei Maschinengewehren
ab.
Am Njassasee kam es nur zu unbedeutenden Zusammenstößen. Am 30. Mai
landeten die Engländer bei Sphinx-Hafen 30 Europäer, 200 Askari mit zwei
Geschützen und zwei Maschinengewehren. Sie erlitten durch unsere 13 Gewehre
und ein Maschinengewehr anscheinend über 20 Mann Verluste und fuhren nach
Zerstörung des Wracks des „Hermann von Wißmann“ ab.

Askarikopf
 Zweites Buch
Der konzentrische Angriff der
Übermacht
(Vom Eintreffen der südafrikanischen Truppen bis
zum Verlust der Kolonie)
 Safari

Erster Abschnitt
Feindlicher Vorstoß am Oldoroboberge[4]

Ö stlich des Oldorobo zeigte der Feind nun mehrfach stärkere

Truppenmassen von 1000 und mehr Mann, die sich auf weiter
Entfernung gegen den Berg entwickelten, aber nicht herankamen.
Es waren also Übungen, durch welche die jungen südafrikanischen
Europäertruppen für die Bewegungen und den Kampf im Pori
ausgebildet werden sollten. Anfang Februar ging der Feind mit
mehreren Regimentern von Osten her gegen den Oldorobo vor. Für
uns war es erwünscht, wenn er dort so fest anbiß, daß er nicht
wieder fortkommen konnte; es bestand die Hoffnung, ihn dann durch
einen Gegenangriff mit der bei Taveta lagernden Abteilung des
Hauptmann Schulz zu schlagen. Andere, mehrere Kompagnien
starke deutsche Abteilungen standen westlich von Taveta an der
Straße nach Neu-Moschi und an der nach Kahe, bei der Pflanzung
Neu-Steglitz.
Am 12. Februar ging wiederum ein auf mehrere Regimenter
geschätzter europäischer Gegner bis auf 300 m gegen den Oldorobo
vor. Das Kommando in Neu-Moschi war in dauernder telephonischer
Verbindung mit Major Kraut, hielt nun den günstigen Augenblick für
gekommen und gab Befehl zur Feuereröffnung. Die Wirkung unserer
Maschinengewehre und unserer 2 leichten Geschütze war als gut
gemeldet worden, als das Kommando im Automobil Neu-Moschi
verließ, um sich auf das Gefechtsfeld zu begeben. Abteilung Schulz
erhielt Befehl, von Taveta aus hinter der Abteilung Kraut, gedeckt
gegen das Feuer der feindlichen schweren Artillerie, entlang zu
marschieren und den feindlichen rechten (nördlichen) Flügel
entscheidend anzugreifen. Die bei Neu-Steglitz stehenden Truppen
rückten nach Taveta vor. Es lagen einige wilde Meldungen vor über
feindliche Panzerautomobile, die durch die Buschsteppe fahren
sollten. Die Phantasie der Farbigen, denen diese Panzerautomobile
etwas ganz Neues und Überraschendes waren, hatte die Leute
Gespenster sehen lassen. Am Oldorobo angekommen, wurde das
Kommando telephonisch orientiert, daß der Feind, der gegen unsere
stark verschanzte Front vorgegangen war, mit schweren Verlusten
abgewiesen worden war, und daß Abteilung Schulz voll entwickelt
gegen seine rechte Flanke vorging. Die zahlreichen in unsere
Stellung auf dem Oldorobo einschlagenden Geschosse der
englischen Haubitzen taten fast keinen Schaden, obgleich sie recht
gut lagen. Dem großen Munitionsaufwande der feindlichen Artillerie
gegenüber mußten sich unsere leichten Geschütze darauf
beschränken, besonders günstige Ziele auszunützen, nicht nur, weil
die Munition knapp war, sondern auch, weil wir keine Schrapnelle
hatten. Der Feind ging in Auflösung durch das Pori zurück. Über 60
Europäer wurden durch uns beerdigt. Nach Gefangenenaussagen
und nach den erbeuteten Papieren waren drei Regimenter der 2.
südafrikanischen Infanteriebrigade im Gefecht gewesen. Es war also
den Engländern tatsächlich gelungen, die militärischen Kräfte der
südafrikanischen Union für ihre imperialistischen Ziele nutzbar zu
machen; nach erbeuteten Papieren scheint die Aussicht auf
Pflanzungen und Farmen bei der Anwerbung der Leute als Zugmittel
benutzt zu sein. Die plötzliche Erkrankung des zur Übernahme des
Oberbefehls in Ostafrika schon auf der Ausreise befindlichen
britischen Generals Smith-Dorrien dürfte den Engländern nicht
ungelegen gekommen sein; denn die Übertragung des Oberbefehls
an einen Südafrikaner, den General Smuts, hat auf die Werbung
einen günstigen Einfluß ausgeübt. Die Ausbildung dieser neu
geschaffenen Verbände war gering, und es war an dem Verhalten
der vielfach sehr jugendlichen Europäer zu erkennen, daß viele noch
niemals an einem ernsthaften Kampf teilgenommen hatten. Nach
dem Gefecht am Oldorobo beobachteten wir aber, wie der Feind die
Lücken in seiner Ausbildung sehr gründlich zu verbessern suchte.
Trotz dem Nachdrücken durch die Abteilung Schulz und
mehrfachen Beschießen sich sammelnder feindlicher Abteilungen
entkam bei der Schwierigkeit und Unübersichtlichkeit des Geländes
der Feind noch in seine befestigten Lager.
Interessant war es, daß in mehreren aufgefundenen
Tagebüchern der ausdrückliche Befehl (order) verzeichnet stand,
daß keine Gefangenen gemacht werden sollten. Tatsächlich hatte
der Feind ja auch keine gemacht, aber es schien doch angezeigt, an
den britischen Befehlshaber eine Anfrage zu richten, damit wir unser
Verhalten den englischen Gefangenen gegenüber danach einrichten
konnten. Es liegt kein Grund vor, die Mitteilung des Brigadegenerals
Malleson, daß ein solcher Befehl nicht gegeben sei, in Zweifel zu
ziehen; dieser und manche spätere Fälle zeigen aber, was für ein
Unsinn in den privaten Tagebüchern steht. Es ist deshalb verkehrt,
wenn der Feind deutsche Aufzeichnungen, die in seine Hände fielen,
ohne eingehende Prüfung für Ernst nehmen will.
Zu dieser Zeit waren auch die feindlichen Truppen auf dem
Longidoberg erheblich verstärkt worden. Dieser Berg, der von dem
Feinde wahrscheinlich aus Nachschubschwierigkeiten geräumt
worden war, wurde neuerdings wieder von ihm besetzt. Unsere
Patrouillen hatten den dicht bewachsenen Felsen mehrfach
erstiegen und die feindlichen Lager aus nächster Nähe beobachtet.
Wenn es an sich schon schwer ist, Truppenzahlen richtig
einzuschätzen, so ist das in dem buschigen Gelände, wo immer nur
wenige Leute gleichzeitig zu sehen sind, und wo das Bild sich
dauernd ändert, ganz unmöglich. Die Angaben von Eingeborenen
waren zu ungenau. Aber allgemein mußte aus der Gesamtlage
sowie aus der Steigerung des Nachschubes, der andauernd mit
Ochsenwagen zum Longido vom Norden her gebracht wurde,
geschlossen werden, daß der Feind sich erheblich verstärkte.
 Seine Streifzüge in das Kilimandjarogebiet waren blutig
abgewiesen worden. Als eine Eskadron von indischen Lancers
zwischen Kilimandjaro und Meruberg hindurch sich nach Süden
vorbewegte, wurde sie von einer unserer berittenen Patrouillen unter
Oberleutnant Freiherrn von Lyncker sogleich energisch angegriffen.
Unsere Askari, die den hohen Wert von Reittieren für unsere
Kriegführung begriffen hatten, stürzten sich mit dem Ruf: „Wahindi
kamata frasi“ (Es sind Inder, fangt die Pferde) auf den abgesessenen
Feind. Dieser war so überrascht durch die Schnelligkeit unserer
Leute, daß er in wilder Flucht davonlief und einen Teil seiner Pferde
stehenließ. Unter anderem war der brave europäische Führer tot
liegengeblieben; es war ihm nicht gelungen, die Kopflosigkeit seiner
Leute zu verhindern.
Ich möchte überhaupt betonen, daß in dieser ersten Zeit des
Krieges das Verhalten der britischen Berufsoffiziere durchweg ein
ritterliches war, und daß die Achtung, die sie uns zollten, voll
erwidert wurde. Aber auch unsere Askari gewannen durch ihr braves
Verhalten im Gefecht und durch ihre Menschlichkeit die Achtung des
Feindes. Der schwerverwundete englische Oberleutnant Barrett fiel
am 10. März in die Hände unserer Askari; auf Grund falscher
Schilderungen glaubte er, daß seine letzte Minute gekommen sei,
und war erstaunt, als unsere Askari, bei denen sich kein Europäer
befand, ihn so gut es ging verbanden und zum Arzt trugen.
Verwundert äußerte er: „Ihre Askari sind ja Gentlemen.“ Bis zu
welchem Maße die Vorstellung der englischen Soldaten irregeleitet
war, zeigte mir am 12. Februar ein junger am Oldorobo gefangener
Südafrikaner, der fragte, ob er nun erschossen werden würde. Wir
lachten ihn natürlich aus. Gewiß kommen in einem langen Kriege
Fälle von Roheit und Unmenschlichkeit vor. Das ist aber auf beiden
Seiten der Fall und darf nicht, wie es von der englischen Presse
geschehen ist, verallgemeinert und zu einer unwürdigen Hetze
ausgenutzt werden.

[4] Skizze IV.

 Zweiter Abschnitt
Vorrücken des Feindes und Kampf bei Reata

I n jener Zeit wurden nun die ersten feindlichen Späherpatrouillen

beobachtet und teilweise auch festgenommen. Es waren dies
„Schensi“ (harmlos erscheinende Eingeborene), welche als Zeichen,
daß sie das Objekt ihres Auftrages wirklich erreicht hatten,
bestimmte Gegenstände, z. B. Teile vom Bahnkörper der
Usambarabahn, mitbringen mußten. Das Gesamtbild, das man sich
zu machen hatte, zeigte, daß der Feind das Gebiet der
Usambarabahn und die Anmarschwege zu derselben eingehend
erkundete. Ein Blick auf die Karte lehrt nun folgendes. Ein
gleichzeitiges Vordringen des Feindes vom Oldorobo und Longido
her in Richtung auf Neu-Moschi mußte den Verlust des wirtschaftlich
wertvollen Kilimandjarogebietes zur Folge haben. Wenn wir aber vor
einem überlegenen Feind auf unsere Hauptnachschublinie
allmählich ausweichen wollten, so führte dies zu einer Bewegung
unserer Hauptkräfte längs der Usambarabahn, also fast spitzwinklig
zu der Richtung eines vom Oldorobo her erfolgenden Angriffes. Die
Gefahr für uns, von dieser unserer Hauptnachschublinie durch den
Feind abgeschnitten zu werden, war sehr groß. Ging der Feind
nördlich des Djipesees vor, so war er in seiner Bewegungsfreiheit
beschränkt durch den Kilimandjaro und das Steilmassiv des
Nordparegebirges. Es ist klar, daß dann sein Vordrücken unmittelbar
gegen Kahe für uns am unangenehmsten war und, wenn erfolgreich,
unsere rückwärtige Verbindung, die Usambarabahn, durchschnitt.
Noch bedenklicher aber war es für uns, wenn der Feind südlich des
Djipesees durch das Tal vordrückte, welches zwischen Nordpare und
Mittelpare südlich Lembeni an die Nordbahn heranführte. Schließlich
konnte er auch zwischen Mittelpare und Südpare durch das Tal bei
Same die Bahn erreichen. Beim Anmarsch aus Lembeni und Same
konnte der Feind sich schnell und streckenweise ohne
Vorbereitungen auf freier Ebene einen brauchbaren Weg für seine
Kraftfahrzeuge schaffen und auf diese seine Unternehmungen
basieren.
Bei unserer geringen Truppenzahl in der Gegend des
Kilimandjaro — etwa 4000 Gewehre — war es unmöglich, die
Streitkräfte zur Sicherung gegen diese verschiedenen
Anmarschmöglichkeiten zu zersplittern. Schon aus rein defensiven
Gründen mußten wir die Kräfte zusammenhalten und dicht am
Feinde bleiben, um ihn dort festzuhalten, wo wir waren, und
hierdurch seine Bewegungen zu überwachen. Ob es glücken würde,
die beiden aus Richtung Longido und Makatau im Vormarsch gegen
das Kilimandjarogebiet zu erwartenden Hauptgruppen des Feindes,
deren jede allein uns erheblich überlegen war, nacheinander einzeln
zu schlagen, war von Anfang an sehr fraglich. Eine Aussicht hierzu
bot sich nur, wenn unsere Truppen blitzartig erst gegen die eine und
dann ebenso schnell gegen die andere feindliche Gruppe bewegt
werden konnten. Die Vorbereitungen hierzu wurden getroffen, und
auf Grund persönlicher Erkundung wurden eine Anzahl
Kolonnenwege, die mit Namen und Bezeichnungen versehen waren,
in dem zerklüfteten Waldgelände nördlich der großen Straße
festgelegt, die von Neu-Moschi nach Westen führt. Zur Benutzung
dieser Wege in größerem Umfange ist es nicht gekommen. Man
durfte sich eben nicht scheuen, eine Sache neunundneunzigmal
vergeblich zu versuchen, wenn das hundertste Mal Aussicht auf
Gelingen bot. Mit der Befolgung dieses Grundsatzes sind wir nicht
schlecht gefahren.
Die Tätigkeit des Feindes steigerte sich, und der Gegner zeigte
bei den vielen kleinen Zusammenstößen eine gute Ausbildung. Er
hatte auch zahlreiche neue Askaritruppenteile aufgestellt, die sich
zum großen Teil aus den begabten Stämmen des Njassagebietes
rekrutierten. Das dem Kilimandjaro nach Nordwesten zu
vorgelagerte und nur spärlich mit Busch bestandene Steppengebiet
war seiner großen Übersichtlichkeit wegen zu überraschenden
Offensivunternehmungen für uns nicht günstig; mehr Aussicht hierzu
bot das dichte Buschgelände zwischen Kilimandjaro und Meru
selbst, das der Gegner, der vom Longido kam, voraussichtlich
durchschreiten würde. Hier wurde nun eine aus fünf ausgesuchten
Askarikompagnien bestehende Abteilung von rund 1000 Gewehren
versammelt. Bei der Unübersichtlichkeit des Geländes glückte es
aber dieser Abteilung nicht, eine der zahlreichen Anfang März nach
Süden vordrückenden feindlichen Kolonnen entscheidend zu fassen.
Auch für den Gegner war die Schwierigkeit, sich zu orientieren, groß,
und erst durch einen indischen Meldereiter, der eine Meldung statt
zu seiner eigenen Truppe versehentlich zu uns brachte, erfuhren wir,
daß sich hier die erste ostafrikanische Division unter General
Steward befand. Da diese Zusammenstöße sich in der Gegend von
Geraragua und südwestlich davon abspielten, war es für ein
Eingreifen mit unseren bei Neu-Steglitz und Himo stehenden
Reserven zu weit (Geraragua-Neu-Moschi etwa zwei Tagemärsche).
Ehe dieser Moment eintrat, drückte der Feind auch von Osten her
vor. Die Flugrichtung der feindlichen Flieger zeigte das
augenscheinliche Interesse des Gegners für die Gegenden ein bis
zwei Stunden nördlich von Taveta. Man mußte auf den Gedanken
kommen, daß der Feind aus seinen östlich Oldorobo gelegenen
Lagern sich nicht ein zweites Mal an diesem Berge blutige Köpfe
holen wollte, sondern beabsichtigte, diese Stellung nördlich zu
umgehen und das Wasser des Lumiflusses, eine Stunde nördlich
von Taveta, zu erreichen. Am 8. März wurden vom Oldorobo aus
mächtige Staubwolken beobachtet, die sich vom feindlichen Lager
aus in der angegebenen Richtung bewegten. Auch wurden
zahlreiche Automobile festgestellt. Vom Ost-Kitovo aus, einem 6 km
westlich Taveta gelegenen Berge, beobachtete auch das Kommando
diese Bewegungen. Unsere Kampfpatrouillen, die Gelegenheit
hatten, die feindlichen Kolonnen mit Erfolg zu beschießen und auch
eine Anzahl Gefangene zu machen, stellten einwandfrei fest, daß an
dieser Stelle die feindlichen Hauptkräfte anrückten und General
Smuts anwesend war.
Noch am Nachmittag des 8. 3. beobachtete das Kommando
starke feindliche Kolonnen von Europäern am Dzhallasee, die von
dort in breit ausgedehnter Schützenlinie ein Stück in Richtung auf
den Ostkitovo vorgingen. Bei dem Mangel an Artillerie mußten wir an
diesem Tage und häufig auch noch später ruhig mit ansehen, daß
der Feind in nicht zu großer Entfernung vor unserer Front wenig
geschickte Bewegungen ungestraft ausführen durfte. Es war aber
klar, daß diese umfassende Bewegung des Feindes die Stellung am
Oldorobo, der wir im Verlaufe des Krieges eine Anzahl glücklicher
Gefechte verdankten, nunmehr unhaltbar machte. Ich beschloß, die
Truppen auf den Bergen, die westlich von Taveta die Lücke
zwischen dem Nordparegebirge und dem Kilimandjaro sperrten, zu
neuem Widerstand zu entwickeln. Abteilung Kraut erhielt
telephonisch Befehl, am Wege, der von Taveta nach Neu-Steglitz
führte, auf den Reata-Latema-Bergen Aufstellung zu nehmen.
Nordwestlich des Latema-Berges, an der Straße, die von Taveta
nach Himo führt, besetzte Abteilung Schulz die Berge von
Nordkitovo und sicherte den Abmarsch der Abteilung Kraut. Diese
Bewegungen wurden in der Nacht ungestört vom Feind ausgeführt.
Auf unserem äußersten linken Flügel, an den Südosthängen des
Kilimandjaro, sperrte die Kompagnie des Hauptmann Stemmermann
die Straße, die von Mission Rombo nach Himo und Neumoschi
führte. Mission Rombo wurde vom Feinde besetzt. Ein Teil der
Eingeborenen machte kein Hehl daraus, daß sie nunmehr Leute der
Engländer wären. Hierdurch wächst die Wahrscheinlichkeit dafür,
daß in dieser Gegend schon seit langem englische Spionage und
Beeinflussung der Eingeborenen tätig waren und daß die oft am
Ostabhange des Kilimandjaro beobachteten Lichtsignale hiermit in
Zusammenhang standen.
Die von uns eingenommene Bergstellung war durch das Gelände
sehr begünstigt, hatte aber doch den großen Nachteil, daß unsere
paar tausend Askari viel zu wenig waren, um die etwa 20 k m breite
Front wirklich zu füllen. Es konnten nur einige Punkte der vorderen
Linie besetzt werden; die Hauptmasse der Truppe wurde bei Himo
zur Verfügung gehalten, um sie, je nachdem sich die Lage
entwickelte, einzusetzen. Es war eine Zeit großer Spannung. Vor
uns hatten wir den weit überlegenen Feind, hinter uns, aus Richtung
Longido nach Süden vordrückend, einen gleichfalls überlegenen
Gegner, und unsere rückwärtige Verbindung, die zugleich unser
Abmarschweg war, wurde, wie oben geschildert, in so empfindlicher
Weise vom Feinde bedroht. Mit Rücksicht auf das Gelände, unsere
Bekanntschaft mit diesem und in Anbetracht der augenscheinlich
nicht allzu gewandten taktischen Führung des Feindes hielt ich es
aber nicht für aussichtslos, diesen mindestens in einem seiner Teile
gründlich zu schlagen. Die Stellungen der Linie Reata-Nordkitovo
sollten deshalb zu zähem Widerstand ausgebaut werden. Von Tanga
wurde eines der dort montierten Königsberggeschütze mit der Bahn
heranbefördert. Mit Recht wird der Leser fragen, warum dies nicht
längst geschehen war. Aber das Geschütz war ohne Räder und
schoß von feststehendem Pivot aus, war also sehr unbeweglich. Es
ist daher erklärlich, daß sein Einsatz so lange hinausgezogen wurde,
bis über die Stelle kein Zweifel mehr bestand.
Die Lage entwickelte sich nun so schnell, daß das Geschütz
gegen Taveta nicht mehr eingesetzt werden konnte. Es ist daher an
der Bahn bei Kahe, am südlichen Ufer des Panganiflusses
aufgebaut worden, von wo es später in den Gefechten von Kahe
ausgezeichnet gewirkt hat.
 Verpflegung bringende Eingeborene
Am 10. März erkundete der Feind gegen unsere gesamte Front.
Berittene Abteilungen von etwa 50 Mann ritten heran, saßen dann ab
und gingen in weit ausgedehnter Schützenlinie, die Pferde am Zügel,
weiter vor, bis sie Feuer erhielten. Dies war ihr Zweck. Das Feuer
verriet ihnen unsere Stellungen, wenn auch unvollkommen. Uns bot
diese Art der Erkundung Gelegenheit zu Teilerfolgen, die dem Feind
eine Anzahl Menschen kostete und uns einige 20 Pferde
einbrachten. Von unserem Berge von Nordkitovo aus beobachteten
wir genau, wie Teile unserer Schützenlinien, schwache Momente
beim Gegner erkennend, rasch vorgingen und die feindlichen
Erkundungsabteilungen von mehreren Seiten unter Feuer nahmen.
Mir schien der Kräfteeinsatz dieser feindlichen Unternehmungen zu
groß, um sie nur mit einer Erkundungsabsicht zu erklären; es
machte mir den Eindruck, daß es sich um ernsthafte, aber etwas
verunglückte Angriffsbewegungen handelte. Über die Richtung des
weiteren feindlichen Hauptangriffes konnte noch keine Klarheit
gewonnen werden. Eine Umfassung unseres linken (nördlichen)
Flügels bot für den Feind sehr viel weniger taktische
Schwierigkeiten, beraubte ihn aber des wirksamen Druckes auf
unsere rückwärtigen Verbindungen. Die für uns unangenehmste
Richtung von Taveta über Reata gegen Kahe bedingte für den Feind
einen schweren und auch bei großer Überlegenheit wenig Erfolg
versprechenden Frontalangriff gegen die befestigten Höhen von
Reata und Latema. Es schien mir angezeigt, hinter der Abteilung des
Majors Kraut, die auf der Höhe von Reata-Latema lag, den
Hauptmann Koehl mit zwei Kompagnien so dicht heranzuschieben,
daß er zum schnellen Eingreifen auch ohne Befehl in der Lage war.
Die telephonische Verbindung mit unseren Abteilungen war für den
Augenblick sichergestellt. Man mußte aber damit rechnen, daß sie
zum mindesten sehr erschwert wurde, sobald eine Abteilung sich
von den augenblicklichen Drahtlinien entfernte. Material zu
schnellem Bau eines Kabels, das den Bewegungen hätte folgen
können, war nicht verfügbar. Ebenso fehlte es uns an leichten
Funkenstationen, durch die später die Engländer mit Erfolg die
Bewegungen ihrer Kolonnen im Busch regelten.
Am 11. März früh erschien wieder ein Flugzeug über Neumoschi
und warf einige Bomben. Ich sprach gerade mit einem alten Buren
über das Gefecht vom 12. Februar und daß es von den Engländern
doch unverantwortlich sei, eine solche Menge junger und in den
Tropen ganz unerfahrener Leute den Einflüssen unseres Klimas und
des Tropenkrieges so rücksichtslos auszusetzen. Von Reata her
meldete Major Kraut, daß sich starke feindliche Truppen von Taveta
her auf seine Stellungen zu bewegten. Bald erfolgte auch ein
kräftiger feindlicher Angriff von mehreren tausend Mann gegen
unsere in ihrer Stellung eingenisteten 3 Kompagnien. Unsere 3
leichten Geschütze konnten den Geschützkampf gegen die schwere
Artillerie natürlich nicht aufnehmen und mußten sich wie bei dem
Gefecht am Oldorobo darauf beschränken, mit ihren wenigen
Granaten die günstigen Momente gegen feindliche dichtere
Truppenmassen auszunutzen. Ich hielt den Angriff in dem mir
bekannten schwierigen Gelände für wenig aussichtsvoll, doch
wurden die zwei hinter der Abteilung des Majors Kraut zur Verfügung
stehenden Kompagnien des Hauptmanns Koehl zum Angriff
angesetzt. Hauptmann Koehl, der ursprünglich einen der Lage
entsprechenden und entscheidenden Flankenstoß gegen den Feind
beabsichtigt hatte, mußte sich in dem unbekannten und dichten
Busch von dessen Unausführbarkeit überzeugen. Zeit und Ort und
damit die Wirksamkeit seines Angriffes hätten ganz dem Zufall
überlassen werden müssen. Richtigerweise marschierte er deshalb
an Major Kraut zu dessen unmittelbarer Unterstützung heran. Auf
Grund meiner Beobachtung von Nordkitovo aus und nach den
einlaufenden Meldungen gewann ich das Bild, daß der Feind unsere
Front von Reata bis Kitovo beschäftigen wollte, seine entscheidende
Bewegung aber um unseren linken Flügel ausholte. Dorthin
bewegten sich zunächst große Reitermassen, die auf den Höhen
und Schluchten des südöstlichen Kilimandjaroabhanges
abwechselnd sichtbar wurden und wieder verschwanden. Die 11.
Kompagnie, Hauptmann Stemmermann, die sich oberhalb dieser
Reiter am Hange befand, verhinderte, daß der Feind diesen erstieg.
Im Laufe des Nachmittags hatte sich die Spitze der Reiter durch die
dichten Bananenpflanzungen bis in die Gegend von Marangu
vorgearbeitet. Anscheinend waren die Reiter sehr erschöpft. Es
wurde beobachtet, daß sie sich zum Teil mit den vorgefundenen
unreifen Bananen verpflegten.
Im Laufe des Nachmittags wurde es klar, daß ein starker
Frontalangriff des Feindes sich gegen die Abteilung Kraut auf dem
Reata- und Latemaberg richtete. Die eingehenden telephonischen
Meldungen lauteten aber günstig: der Feind hatte augenscheinlich
schwere Verluste; Hunderte von Tragbahren waren bei ihm zum
Abtransport Verwundeter in Tätigkeit. Abends waren alle Angriffe
des Feindes gegen unsere Front mit schweren Verlusten für ihn
abgeschlagen. Bei Dunkelheit hatten die beiden Kompagnien der
Abteilung Koehl energisch nachgedrückt und den sich von neuem
setzenden Feind unter Maschinengewehrfeuer genommen. Ich hatte
mich abends nach Himo zurückbegeben und war gegen 11 Uhr
damit beschäftigt, den Befehl zum Angriff auf die bei Marangu
festgestellten feindlichen Reiter in der Morgenfrühe des 12. März
auszugeben. Da telephonierte Leutnant Sternheim, der Führer der
Geschütze der Abteilung Kraut, daß der Feind in der Nacht noch
einmal angegriffen habe und mit starken Massen in die Stellung bei
Reata eingedrungen sei. Die Meldung ließ es als wahrscheinlich
erscheinen, daß dieser starke Feind von Reata nun weiter in der
Richtung auf Kahe drücken und uns von unserer Verbindung
abschneiden würde. Dieses Risiko zu übernehmen und trotzdem
den Angriff auf den Feind bei Marangu durchzuführen, erschien mir
zu bedenklich. Ich gab daher Befehl zum Abmarsch der bei Kitovo
und Himo stehenden Truppen noch während der Nacht an die
Straße Kahe-Reata. Als Sicherung sollte in Himo die Kompagnie
Stemmermann verbleiben. Der befohlene Abmarsch der Truppe
mußte die unangenehme Folge haben, daß im günstigsten Falle für
Stunden jede Verbindung des Kommandos mit den einzelnen
Truppenteilen aufhörte. Wer diese Nachtmärsche im dichten Busch
kennt, der weiß, wie leicht außerdem Teile ganz abreißen und dann
auf absehbare Zeit überhaupt nicht wieder zu erreichen sind.
Glücklicherweise war mir das Gelände wenigstens einigermaßen
bekannt, als wir querfeldein auf die neue Straße herüberrückten und
hierbei andauernd lebhaftes Feuer auf den Reata- und
Latemabergen hörten. Einige Versprengte, die sich im Busch
verlaufen hatten, kamen uns entgegen; unserer Angabe, daß wir
Deutsche wären, glaubten sie nicht und verschwanden wieder. An
der neuen Straße angekommen, trafen wir auf den Verbandplatz.
Auch hier waren die Angaben der vielen Verwundeten so
widersprechend und unklar, daß man nur den Eindruck eines sehr
heftigen Nahkampfes im Busch hatte, aber über dessen eigentlichen
Zusammenhang und Verlauf man nichts erfuhr. Allmählich gelang es,
telephonische Verbindung mit Major Kraut zu erhalten. Dieser
befand sich mit einem Teil seiner Abteilung am Südwesthange des
Reataberges an der Straße Kahe-Taveta. Das Feuer auf den Höhen
war allmählich verstummt, und seine Patrouillen hatten auf dem
Reataberge nichts mehr vom Feinde gefunden. In der Morgenfrühe
des 12. fand Major Kraut auf den Bergen einen Teil seiner Abteilung
in den alten Stellungen wieder; der Feind war nach Taveta
zurückgegangen.
Als ich gegen 6 Uhr früh auf dem Reataberge eintraf, wurde die
zahlreich vorhandene Beute gesammelt. Das Durcheinander des
nächtlichen Nahkampfes im Busch war sehr groß gewesen.
Englische Gefallene, die weit hinter der Front der Abteilung Kraut im
Busch lagen, zeigten an, wie weit einzelne Abteilungen des Feindes
geraten waren. Feindliche Einzelschützen, die versteckt in den
Felsen eingenistet waren, unterhielten ein gutgezieltes Feuer und
waren nicht auszumachen. Es war klar, daß der feindliche Angriff mit
schweren Verlusten für den Gegner abgewiesen worden war. Der
Abtransport der eigenen und feindlichen Verwundeten ging glatt
vonstatten, ebenso der der Gefangenen. Mit den aus der Gegend
von Himo an die Straße Kahe-Reata durch dichten Busch
heranrückenden Abteilungen bestand keine Verbindung, und es war
nicht zu erwarten, daß diese vor Ablauf mehrerer Stunden
hergestellt werden würde.
Bei dieser Lage war es zu bedauern, daß ich die Truppen
unseres linken Flügels, die in der Gegend Kitovo-Himo gestanden
hatten, an die Straße Kahe-Reata herangezogen hatte. Bei der
Preisgabe der Höhen unseres linken Flügels war die Stellung von
Reata auf die Dauer nicht zu halten, um so weniger, als unsere
Stellung kein Wasser hatte, das eine Stunde weit von rückwärts
herangebracht werden mußte. Um die in Marsch befindlichen
Abteilungen des linken Flügels wieder abzudrehen und von neuem
in die Gegend von Himo und Kitovo in Marsch zu setzen, dazu fehlte
in diesem Augenblick die Verbindung, und es war, wie erwähnt,
anzunehmen, daß sie nicht vor Ablauf von Stunden erreicht werden
würde. Ich beschloß deshalb, die Stellung von Reata zu räumen,
und ging nach Aufräumung des Gefechtsfeldes mit der am weitesten
nach dem Feinde zu gelegenen Gefechtslinie bis an das südwestlich
des Reataberges gelegene Wasser zurück. Im Laufe des Tages
trafen dahinter die anderen Abteilungen an verschiedenen Punkten
der Straße Kahe-Reata ein und bezogen Lager.