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Contents lists available at ScienceDirect

Agriculture, Ecosystems and Environment

journal homepage: www.elsevier.com/locate/agee

The wild relatives of grape in China: Diversity, conservation gaps and

impact of climate change
Jianfu Jiang a , Shelagh Kell b , Xiucai Fan a , Ying Zhang a , Wei Wei c , Dingming Kang d ,
Nigel Maxted b , Brian Ford-Lloyd b , Chonghuai Liu a, *
a
Zhengzhou Fruit Research Institute, Chinese Academy of Agricultural Sciences, Henan, Zhengzhou 450009, PR China
b
School of Biosciences, University of Birmingham, Edgbaston, Birmingham, B15 2TT, UK
c
Institute of Botany, The Chinese Academy of Sciences, Beijing 100093, PR China
d
College of Agronomy and Biotechnology, China Agricultural University, Beijing 100193, PR China

A R T I C L E I N F O A B S T R A C T

Article history: China is one of the major diversity centres of grape (Vitis spp.) and is therefore one of the most abundant
Received 20 April 2014 sources of Vitis germplasm in the world. Grape wild relative species (GWRs) represent a potentially
Received in revised form 11 March 2015 important source of valuable traits for the improvement of cultivated grape varieties and have signicant
Accepted 20 March 2015
characters for resistance to biotic and abiotic stress factors. We studied the ecogeographic diversity of
Available online xxx
GWRs, conservation gaps and impact of climate change on GWRs in China, based on a wide range of
distribution data sourced from germplasm and herbarium specimens, eld surveys and other literature.
Keywords:
Results show that there are 39 species, 1 subspecies and 14 varieties of GWRs native to China and that 19
Grape
Crop wild relative
species and 9 varieties are the closest wild relatives to cultivated grape according to the Taxon Group
Vitis Concept. GWRs are distributed in nearly all provinces in China except for Xinjiang, but they are
Diversity particularly abundant in Jiangxi and Hunan provinces. The richest regions for GWRs are the Qinling, Daba,
Conservation Wuling, Nanling and Wuyi mountains. Around 22% of GWR species are found in natural reserves (NRs)
Climate change and are well protected, but 15 species are not found in NRs and require further strengthening of both
protection and collection. The potential distribution of GWRs at the present and predicted future climate
was compared using BIOCLIM. The results showed that simulated current distributions matched actual
distribution ranges. Under the future climate scenario with doubled CO2 concentration, suitable areas for
continued survival of 21 GWRs could be reduced. Our results will therefore be extremely valuable for the
development of a complementary conservation strategy for Vitis in China.
2015 Elsevier B.V. All rights reserved.

1. Introduction
Grapes are economically among the most important fruit in the
world, there are 800010,000 grape cultivars existing worldwide
today (Ramezani et al., 2009), and most production cultivars are
derived from only one species V. vinifera, because of its high berry
qualities. However, V. vinifera is highly susceptible to fungal
diseases, which causes heavy losses in grape production (Wan
et al., 2008a). On the other side, V. vinifera had narrow genetic
background, grape breeders are making efforts to use wild
germplasms to improve resistance of cultivars and to breed new
resistant cultivars (Alleweldt and Possingham, 1988; Brown et al.,
1999).
* Corresponding author. Tel.: +86 371 65330966; fax: +86 371 65330987.
E-mail address: liuchonghuai@caas.cn (C. Liu).
China is one of the major centres of diversity of grape (Vitis spp.)
and is therefore one of the most abundant sources of Vitis
germplasm in the world. Grape wild relative species (GWRs) are a
potentially important source of valuable traits for the improve-
ment of cultivated grape varieties and have signicant characters
for resistance to biotic and abiotic stresses such as cold, drought,
pests and diseases. Grape breeding has proved that it is important
to use wild germplasm resources carrying a range of resistance
genes, to counter the shortage of the resistance genes within the
cultivated species. Currently, many elite cultivars have been
developed using GWRs such as V. amurensis, V. heyneana,
V. pseudoreticulata, V. davidii and V. bryoniaefolia. In addition,
GWRs and their hybrids have been used for wine production in
China (Wan et al., 2008b).
With the environment pollution and ecoystem degradation of
recent years, many wild animals and plants are endangered or
faced with extinction and GWRs are no exception. The loss of a
species means the loss of the genes that it carries. It is recorded in

http://dx.doi.org/10.1016/j.agee.2015.03.021
0167-8809/ 2015 Elsevier B.V. All rights reserved.

Please cite this article in press as: J. Jiang, et al., The wild relatives of grape in China: Diversity, conservation gaps and impact of climate change,
Agric. Ecosyst. Environ. (2015), http://dx.doi.org/10.1016/j.agee.2015.03.021
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China Species Red List that V. yunnanensis, V. wenchouensis and V.
hui are in serious danger with fewer than 5 remaining populations
subjected to continuing decline. There is only one known
remaining site for both V. bashanica and V. mengziensis and so
their populations are under very serious threat (Wang and Xie,
2004).
Changes in climatic conditions on earth, including global
warming, are no longer contested. According to the Fourth
Assessment Report of the Intergovernmental Panel on Climate
Change (IPCC), global mean surface air temperature has increased
approximately 0.8  C since the start of 20th century, and will
continue to rise between 1.1 and 6.4  C by the end of this century
(Betts and Hawkins, 2014), ooding and droughts will be more
frequent and severe, some soils will degrade, and climatic
extremes will be more likely to occur (Jones et al., 2007; Ainsworth
et al., 2008). As some species have already responded to a
temperature increase of 0.6  C, it is clear that more substantial
effects on species and ecosystems will occur in the future (Root
et al., 2003), especially for those species with a restricted
distribution pattern (Midgley et al., 2002) and also CWRs (Maxted
et al., 2013).
Predicting the current or future distributions of species has
principally been conducted using bioclimatic models that assume
the climate ultimately restricts species distributions. These models
summarise a number of climatic variables within the known range
of a species, thus generating a bioclimatic envelope. The models
can then be used to: (a) identify the species current potential
distribution, that is, all areas with climatic values within the
species bioclimatic envelope and (b) assess whether these areas
will remain climatically suitable under future climate scenarios
(Pearson and Dawson, 2003).
In China, there are 3 national repositories for grapevine
(Zhengzhou, Zuojia and Taigu), which specialize in grape
germplasm collection, conservation, as well as identication and
utilization. Nine species have been protected in national reposito-
ries, but they are all common species (Ren et al., 2012). Natural
reserves play an important role in protecting wildlife including
GWRs, but research on the distribution of GWRs has been
restricted to certain regions with truly systematic studies absent
in past years (Zuo and Yuan, 1981). The number of GWRs in China is
unknown together with their geographical distribution. It is
therefore not clear whether GWRs are receiving adequate
protection. Knowledge is absent on which districts have the
richest grape resources and which species are endangered.
Furthermore, there is no reporting undertaken on the impact of
future climate change on the GWR resources. Such information is
crucial for research, protection and usage of the GWRs in China and
indeed globally.
The present study aims to investigate (1) the number of GWRs
in China and their relation with cultivars for priority utilization; (2)
the distribution and conservation gaps of GWRs in China; (3) the
impact of future climate change on GWRs in China.
2. Materials and methods
2.1. Data collection
2.1.1. Species data
We have set up a distribution database of the GWRs in China by
surveying various sources of information: (1) 155 specimen records
of wild grapes since 1990 from the major domestic specimen
museums through the Chinese Virtual Herbarium (CVH, http://
www.cvh.org.cn); (2) Flora of China (VITACEAE) (Ren and Wen 2007),
as well as valid provincial and regional oras, such as Flora of Yunnan
(Kunming institute of Botany, Chinese academy of science, 2000),
Flora of Zhejiang (Flora of Zhejiang editorial board,1993), A checklist
Please cite this article in press as: J. Jiang, et al., The wild relatives of grape in China: Diversity, conservation gaps and impact of climate change,
Agric. Ecosyst. Environ. (2015), http://dx.doi.org/10.1016/j.agee.2015.03.021
of Vascular Plants of Guangxi (Zuo and Liu, 2010) and Dabieshan
sylva (Zi and Zhang, 2006); (3) the scientic survey reports on natural
reserves (NRs) in recent years, totaling 380 related reports on
domestic NRs covering all mainland provinces in China except
Hongkong, Macao and Taiwan areas; (4) the research summaries on
wild Vitis published in various academic journals and the relevant
literature of the wild Vitis records; and (5) recent eld investigations
in Henan, Hunan, Guangxi, Zhejiang, Jiangxi and Tibet.
Subsequently, we cross-checked, veried and where necessary
corrected the species names and geographic references according
to Chinese Ampelography (Kong, 2004). Controversial species are
not included in the database.
Ultimately, we obtained 2675 unique localities for 38 species 1
subspecies and 13 varieties of GWRs that were then used for
further analysis.
2.1.2. Climate data
The climate data used in this research are based on the gridded
spatial database available from the World Climate website
(http://www.worldclim.org/). The climate database, with
30 arc-second resolution (ca. 1 km at the equator), was developed
using monthly mean temperature and monthly precipitation at
climatic stations (ca. 7000 stations for temperature and
20,000 stations for precipitation) between 1950 and 2000
(Hijmans et al., 2005). This database is more accurate and
precise, as it used altitude as one of parameters when
interpolating the climatic variables, compared to the 2-dimen-
sional interpolation used in previous estimations (Hijmans et al.,
2005).
2.2. Diversity analysis
Taxonomical review was undertaken of all GWR species related
to the cultivated grape, and analysis of their relationship to the
domesticated species using the Taxon Group Concept established
by Maxted et al. (2006).
2.3. Distribution, richness and gap analysis
The database in Section 2.1.1 mainly consists of species names,
collection localities (with longitude and latitude), habitats and
altitudes. For some specimens localities and published distribu-
tional data had only county names rather than detailed informa-
tion of location such as geographic coordinates. Therefore it was
necessary to determine each locality with latitude and longitude
coordinates by referring to The Gazetteers of China (Department of
Gazetteer, Institute of Topographic Science, National Survey and
Drawing Bureau, 1983), which has been successfully used for
analogous analyses (Lei et al., 2003; Xu et al., 2008; Wang and Ni,
2009; Wu and Zhou, 2012). There are 23 provinces, 5 autonomous
regions and 4 municipalities in China. A province is equivalent to
an autonomous region and municipality in terms of administrative
level; thus we refer to all provinces and autonomous regions as
province for the convenience of discussion (Huang et al., 2011).
We have used ARC GIS 10.0 software (ESRI) to describe the
geographical distribution of GWRs, and the number of GWRs
distributed in each province in China.
Using DIVA-GIS 5.4 software (Hijmans et al., 2001), we
produced a 1 1 decimal degree size grid map and statistic of
the species richness in each grid. By using GIS overlap techniques
we produced the distribution map in detail so as to show the
regions where more species are distributed and hence the
distribution centre of GWRs.
Based on the geographical distribution of GWRs, using ARC GIS
10.0 software (ESRI), we identied gaps in ex situ conserved
germplasm by comparing the layer of GWRs distribution with
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Chinese natural reserves (NRs). If the locality is in the reserves
layer, it indicates this species has been well protected, if not, it
shows there are gaps in ex situ conserved diversity and a need for
further protection in future (Maxted et al., 2008a). The layer of
Chinese NRs was downloaded from the World Database on
Protected Areas website (http://www.wdpa.org).
2.4. Potential distribution of GWRs and prediction of the future
distribution
Bioclimatic models are widely used as tools for assessing
potential responses of species to climate change. One commonly
used model is BIOCLIM (Busby, 1991) as implemented in DIVA-GIS
(www.diva-gis.org), which summarises up to 35 climatic param-
eters throughout the known range of a species, and assesses the
climatic suitability of habitat under current and future climate
scenarios (Beaumont et al., 2005).
We have used this Bioclim model to predict the current and
future geographic distribution of each GWR under study. The
environmental data consisted of climate surfaces for present and
projected future conditions. For present climate we used
WorldClim climate surfaces (Hijmans et al., 2005) because of
their high spatial resolution (with 30 arc-second) and global
extent. Projected future climate data are available from Govindas-
amy et al. (2003) which has the highest spatial resolution available.
They used the CCM3 model at a 50 km spatial resolution and for a
concentration of CO2 in the atmosphere of 600 ppm (two times
pre-industrial). This CO2 concentration (including other green-
house gasses expressed as CO2 equivalents in terms of their
warming potential) might occur around 2100. In order to match the
1 km resolution of the current climate conditions, a downscaling
procedure was applied to the CCM3 data by calculating the
predicted change in monthly means from the CCM3 model. These
modied data were then downscaled to 1 km resolution using
smoothing (spatial interpolation), and added to the current
WorldClim climatic surfaces.
We have selected climatic variables associated with tempera-
ture and rainfall distribution of 19 bio-climatic variables which
have important implications for species (Xu et al., 2008), including
annual mean temperature, mean monthly temperature range,
mean annual temperature range, maximum temperature of the
warmest month, minimum temperature of the coldest month,
isothermality, temperature seasonality, mean temperature of
wettest quarter, mean temperature of driest quarter, mean
temperature of warmest quarter, mean temperature of coldest
quarter, annual precipitation, precipitation of wettest month,
precipitation of driest month, precipitation seasonality, precipita-
tion of the driest quarter, precipitation of wettest quarter,
precipitation of warmest quarter and precipitation of coldest
quarter (Jarvis et al., 2008). The predictive results of Bioclim are
represented by 5 categories: grid cells within the 02.5 percentile
have a low climatic suitability, those within the 2.55 percentile a
Table 1
Taxon group concept applied to GWRs in China.
TG1 TG2
same species same series or section as crop
as crop
There is no V. amurensis,V. amurensis var.dissecta, V. amurensis var. funiushanensis, V.
TG1 for V. balanseana, V. balanseana var. tomentosa, V. balanseana var. cifolioides, V.
vinifera luochengensis, V. luochengensis var.tomentoso-nerva, V. exuosa, V. betulifolia,
V. wilsonae, V. pseudoreticulata, V. yunnanensis, V. mengziensis, V. fengqinensis,
V. hekouensis, V. piloso-nerva, V. chunganensis, V. chungii, V. piasezkii, V.
piasezkii var. pagnucii, V. 1anceolatifoilosa, V. davidii, V. davidii var. cyanocarpa,
V. davidii var. ferruginea, V. adstricta, V. bryoniifolia var. ternata, V. zhejiang-
adstricta
Please cite this article in press as: J. Jiang, et al., The wild relatives of grape in China: Diversity, conservation gaps and impact of climate change,
Agric. Ecosyst. Environ. (2015), http://dx.doi.org/10.1016/j.agee.2015.03.021
3
medium, those within the 510 percentile a high, those within
the 1020 percentile a very high and those cells within the
20100 percentile have an excellent climatic suitability (Hijmans
et al., 2001).
Only species for which we had at least 10 distinct localities of
occurrence were included in the analysis (Andy et al., 2008),
resulting in a study of 26 individual species (including subspecies
and varieties) that were analysed. Hernandez et al. (2006) reported
a 30% prediction success using Bioclim with less than 10 samples.
The prediction increases to over 80% when 75 samples are used,
but this threshold would have limited our study to just 12 species
with more than unique localities.
We used the Bioclim model to estimate current and future
species ranges, and subsequently used to calculate extinction risk.
The change metrics of suitable areas (contain medium, high, very
high and excellent) were achieved by ARC GIS 10.0 software (ESRI).
3. Results
3.1. The diversity of GWR in China
Sixty eight Vitis species have been recognised in China (see
Supplementary Appendix A1), although one-third (22 species) are
controversial. Chinese Ampelography (Kong, 2004) is a major
authority on grape research, and it records 39 species, 1 subspecies
and 14 varieties of GWRs native to China, but this authority does
not recognise sections or series. The systematic study of Vitis in
China by Wang and Zhu (2000) indicated the existence of
42 species, 1 subspecies and 12 varieties of wild grape, which
included 1 subgenus, 5 sections and 4 series. We have taken into
account both viewpoints, and suggest that 19 species and 9
varieties are actually the closest wild relatives to cultivated
grape according to the Taxon Group Concept; there is no taxon
group 1 (TG1) for cultivated grapes, and the GWRs in China belong
to TG2 and TG3 (Table 1).
3.2. The distribution patterns
There are great differences in distribution for the different
species. V. heyneana, V. exuosa, V. amurensis, V. davidii, and V.
bryoniaefolia have the widest distributions, found in more than
20 provinces with more than 200 sites identied in this study.
There are 26 species distributed across fewer than 10 site, namely
V. wenchouensis, V. yunnanensis, V. hui, V. piloso-nerva,
V. wuhanensis, V. erythrophylla, V. longquanensis, V. fengqinensis,
V. luochengensis, V. hekouensis, V. jinggangensis, V. ruyuanensis,
V. amurensis var. dissecta, V. zhejiang-adstricta, V. bryoniaefolia var.
ternata, V. shenxiensise, V. luochengensis var. tomentoso-nerva,
V. amurensis var. funiushanensis, V. balanseana var. cifolioides,
V. bashanica, V. wuhanensis var. arachnoidea, V. menghaiensis,
V. mengziensis, V. romaneti var. tomentosa, V. balanseana var.
TG3
same subgenus as crop
V. heyneana, V. heyneana subsp. cifolia, V. bellula, V. bellula var. pubigera, V.
retordii, V. hui, V. longquanensis, V. menghaiensis, V. sinocinerea, V. romaneti, V.
romaneti var. tomentosa, V. shenxiensise, V. wuhanensis, V. silvestrii, V.
wenchouensis, V. tsoii, V. ruyuanensis, V. jinggangensis, V. erythrophylla, V .
hancockii, V. davidii var. cyanocarpa, V. davidii var. hispida, V. wenxianensis, V.
wuhanensis var. arachnoidea, V. bashanica
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Fig. 1. Geographical distribution of grape wild relatives in China.

tomentos. Consequently their future survival is threated and they
deserve strengthening of protection and collection (Fig. 1).
GWRs are found between 97 S and 133 S being mainly
distributed unevenly in the southeast of China. The highest
numbers of species are found in Jiangxi and Hunan provinces,
where more than 20 species occur. Further from these regions,
fewer species occur (Fig. 2). There are 2 species (V. betulifolia and V.
heyneana) in the southeast part of Tibet and the eastern region of
Qinghai, respectively. There is only one species (V. amurensis) in
Heilongjiang, Jilin and Inner Mongolia, and no GWR in Xinjiang.
Yunnan province has the most endemic species for China
(V. hekouensis, V. menghaiensis and V. mengziensis), followed by
Zhejiang (V. wenchouensis and V. zhejiang-adstricta), and Guang-
dong has one endemic species (V. ruyuanensis). Further analysis
shows that several regions with the highest species richness occur
in mountainous areas, such as Qinling, Daba, Wuling, Nanling and
Wuyi mountains. In contrast, species richness is low in the plain
and desert regions (Fig. 3).

Fig. 2. Numbers of grape wild relatives distributed in provinces of China.

Please cite this article in press as: J. Jiang, et al., The wild relatives of grape in China: Diversity, conservation gaps and impact of climate change,
Agric. Ecosyst. Environ. (2015), http://dx.doi.org/10.1016/j.agee.2015.03.021
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Fig. 3. Species richness of grape wild relatives in China (1 1 grid).

3.3. The conservation gaps
The conservation gaps for GWRs in China were identied by
comparing the maps of GWRs distribution and the Chinese NRs
layer (Fig. 4 and Table 2). We identied 2675 effective sites for 52
GWRs, 22.3% (596 sites) occurred in NRs and have been well
protected. Fifteen species including V. heyneana, V. exuosa,
V. davidii, V. adstricta, V. betulifolia, V. amurensis, V. wilsonae,
V. chunganensis, V. piasezkii, V. pseudoreticulata, V. romaneti,
V. heyneana subsp. cifolia, V. sinocinerea, V. bellula, V. chungii have
received good protection, distributed in more than 10 NRs each,
and particularly for V. heyneana, which is found in 78 NRs. Ten
species were found to have more than 3 locations in NRs, namely V.
adenoclada, V. piasezkii var. pagnucii, V. balanseana, V. retordii, V .
hancockii, V. davidii var. cyanocarpa, V. davidii var. ferruginea,
V. silvestrii, V. tsoii, V. bellula var. pubigera. There was only 1 site in
NRs for the 12 species V. jinggangensis, V. yunnanensis, V. amurensis
var. funiushanensis, V. hekouensis, V. erythrophylla, V. 1anceolatifoi-
losa, V. longquanensis, V. hui, V. luochengensis, V. romaneti var.
tomentosa, V. wenxianensis and V. wuhanensis. Of greatest concern,
15 species could not be found in NRs at all, and they were
V. fengqinensis, V. luochengensis var. tomentoso-nerva, V. balanseana
var. cifolioides, V. piloso-nerva, V. wuhanensis var. arachnoidea,
V. menghaiensis, V. mengziensis, V. balanseana var. tomentosa,
V. ruyuanensis, V. bryoniifolia var. ternata, V. shenxiensise,
V. amurensis var. dissecta, V. davidii var. hispida, V. wenchouensis,
V. zhejiang-adstricta. Hence these species in particular need urgent
action in terms of protection and collection.

Fig. 4. In situ protected GWR species of China.

Please cite this article in press as: J. Jiang, et al., The wild relatives of grape in China: Diversity, conservation gaps and impact of climate change,
Agric. Ecosyst. Environ. (2015), http://dx.doi.org/10.1016/j.agee.2015.03.021
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Table 2
Number of sites and accessions conserved, and predicted change in suitable habitat.

Species Number of distribution site Number of sites in reserves Conserved ex situ Predicted change in suitable habitata
V. heyneana 346 78 268 0.064
V. exuosa 319 72 247 0.089
V. davidii 224 53 171 0.052
V. bryoniaefolia 213 38 175 0.114
V. amurensis 205 36 169 0.063
V. heyneana subsp. cifolia 121 26 95 +0.029
V. chunganensis 121 29 92 0.050
V. wilsonae 120 36 84 0.088
V. pseudoreticulata 111 26 85 0.001
V. betulifolia 108 37 71 0.072
V. piasezkii 103 27 76 0.063
V. romaneti 76 26 50 +0.083
V. balanseana 64 8 56 0.200
V. sinocinerea 63 15 48 0.205
V. chungii 60 11 49 +0.021
V. tsoii 57 4 53 0.174
V. piasezkii var. pagnucii 48 8 40 0.075
V. bellula 45 11 34 0.278
V. hancockii 39 6 33 0.259
V. retordii 31 7 24 0.341
V. davidii var. yanocarpa 27 6 21 +0.055
V. adenoclada 26 9 17 0.148
V. 1anceolatifoilosa 24 1 23 0.075
V. davidii var. erruginea 21 6 15 0.120
V. bellula var. pubigera 18 3 15 0.277
V. silvestrii 12 4 8 +0.441
V. wenchouensis 8 0 8
V. yunnanensis 7 2 5
V. hui 5 1 4
V. piloso-nerva 5 0 5
V. wuhanensis 4 1 3
V. erythrophylla 4 1 3
V. longquanensis 4 1 3
V. fengqinensis 3 3
V. luochengensis 3 1 2
V. hekouensis 3 1 2
V. jinggangensis 3 2 1
V. ruyuanensis 3 0 3
V. amurensis var. dissecta 3 0 3
V. zhejiang-adstricta 3 0 3
V. bryoniaefolia var. ternata 2 0 2
V. shenxiensise 2 0 2
V. luochengensis var. tomentoso-nerva 2 0 2
V. amurensis var. funiushanensis 1 1 0
V. balanseana var. cifolioides 1 0 1
V. bashanica 1 0 1
V. wuhanensis var. arachnoidea 1 0 1
V. menghaiensis 1 0 1
V. mengziensis 1 0 1
V. romaneti var. tomentosa 1 1 0
V. balanseana var. tomentosa 1 0 1
V. wenxianensis 1 1 0
a
+ means increase in suitable area and means reduce. There are no data if the distribution site is less than 10 because of the low accuracy.

3.4. The simulation of potential distributions and prediction of future
distributions under climate change
Potential distributions of 26 GWRs under the present (Fig. 5 left,
only lists 5 species here, for others see Supplementary
Appendix B1) and predicted future climate (Fig. 5 right column)
were compared using BIOCLIM. Simulated present distributions
matched actual distribution ranges. The centre of the present as
well as future potential distributions is in south China, the main
distribution of V. heyneana is in Guizhou province, but under the
future climate scenario with doubled CO2, the suitable area is
expected to shift northward and show trends of fragmentation.
Under future climate change, not all suitable areas for GWRs
will reduce. The suitable area for 5 species will become larger as a
result of climate change (Table 2), especially for V. silvestrii, where
the suitable area will increase by 44.1%. However, suitable area for
21 GWR species will decrease, and particularly for V. adstricta,
V. sinocinerea, V. balanseana, V. bellula, V. hancockii, V. adenoclada,
V. tsoii, V. retordii, V. davidii var. ferruginea and V. bellula var.
pubigera, for which the suitable areas will decrease by more than
10%. The situation will be even more serious for V. retordii, for
which the suitable area will be reduced by 34.1%.
4. Discussion
Application of the Taxon Group Concept assumes that
taxonomic relatedness is an indicator of the degree of genetic
relatedness, and for practical purposes, classical taxonomy
remains an extremely useful means of estimating genetic relation-
ships. The taxon group concept can be applied to all crop and CWR
taxa, and can be used to dene CWR relatedness for 78% of crop and
CWR taxa where there is insufcient scientic information to apply
the gene pool concept (Maxted et al., 2006), as long as the existing
classication for the genus contains an infrageneric structure.

Please cite this article in press as: J. Jiang, et al., The wild relatives of grape in China: Diversity, conservation gaps and impact of climate change,
Agric. Ecosyst. Environ. (2015), http://dx.doi.org/10.1016/j.agee.2015.03.021
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Fig. 5. Present and future potential distribution maps for 5 GWRs in China, predicted by DIVA-GIS. The left (a,c,e,g,i) gures represent the present potential distribution range;
those on the right (b,d,f,h,j) represent the future potential distribution range. Four grey degrees (05%, 510%, 1020%, 2040%) indicate that the darker of the color, the more
probable will the species been distributed. A, B: V. heyneana; C, D: V. exuosa; E, F: V. bryoniaefolia; G, H: V. wilsonae; I, J: V. davidii.

A crop wild relative is a wild plant taxon that has an indirect use belong to GP1B, or TG1b and TG2 may be considered close to CWR
derived from its relatively close genetic relationship to a crop; this and demand higher conservation priority, while those in GP2 or
relationship is dened in terms of the CWR belonging to gene pools TG3 and TG4 are more remote and may be afforded lower priority.
1 or 2, or taxon groups 14 of the crop. Therefore, taxa which Those in GP3 or TG5 would be excluded from being considered

Please cite this article in press as: J. Jiang, et al., The wild relatives of grape in China: Diversity, conservation gaps and impact of climate change,
Agric. Ecosyst. Environ. (2015), http://dx.doi.org/10.1016/j.agee.2015.03.021
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8 J. Jiang et al. / Agriculture, Ecosystems and Environment xxx (2015) xxxxxx
wild relatives of that particular crop. Therefore, application of the
gene pool and taxon group concepts provides a pragmatic way of
establishing the degree of CWR relatedness and thus assists in
establishing conservation priorities (Maxted et al., 2008a,b).
Although China is one of the major centres of diversity of wild
grape species, most commercial cultivars are derived from V.
vinifera, and originated from South Europe and West Asia. In China
there is, therefore, no TG1 for V. vinifera, but there are 19 species
and 9 varieties in TG2 which are the closest wild relatives to
cultivated grape according to the Taxon Group concept.
Zuo (1981) and Kong (2004) have reported the distribution of
the wild grapes in China, principally taking references of the
specimen data and other information from Flora of China
(Vitaveae). More recently, realizing the importance of germplasm
resources, scientists have started to investigate crop wild relatives
in more detail, and in recent years, new species have been added to
the original distributions. We updated the distribution data for
GWRs in China, and propose further protection policies for those
GWRs. V. heyneana, V. exuosa, V. amurensis and V. bryoniaefolia
distributed mostly in China, the areas with richest CWRs were in
Jiangxi and Hunan provinces, which is basically consistent with
previous researches (Kong, 2004).
The establishment of natural reserves is really important for the
protection of wildlife including CWRs in China. The total number of
reserves in China was 2640 covering an area of 1497 km2 by the end
of 2011 (Department of Nature and Ecology Conservation, 2012).
But the reserves have an unbalanced distribution. The reserves are
centralized in the provinces of Guangdong, Inner Mongolia,
Heilongjiang and Jiangxi, etc. In terms of area of the reserves,
they are principally distributed in provinces in the west including
Tibet, Qinghai, Xinjiang, Inner Mongolia, Sichuan and Gansu, etc.
and importantly those regions have relatively few CWRs including
grape wild relatives. The area of the natural reserves in Zhejiang,
Hebei, Fujian, An'hui, Henan and Guangdong provinces in the
eastern part are all lower than 5% of the land area, and
consequently does not match the distribution of CWRs in general,
let alone grape wild relatives.
Since 2001, the state began to implement the construction of
the original habitat protection area (point) of the wild plants. Up to
2009, 116 original habitat protection points of the wild kindred
plants have been established including the wild apple, wild
kiwifruit and wild citrus; however, the GWRs were not yet
included. Although they were abundant in China, but some species
have been facing extinction problem and deserve for protection.
Upon that, the country needed to set up the natural reserves or the
original habitat protection sites for the 15 GWRs species that are
not found in NRs.
To cope with the impacts of future climate change, it is
imperative that we can condently predict the current and future
potential distributions of species, particularly crop wild relatives,
so that their conservation can be planned effectively. Species
distribution models have a broad range of applications, and have
been used to assess the potential threat of pests or invasive species
(Ungerer et al., 1999; Sutherst et al., 2000), to obtain insights into
the biology and biogeography of species (Anderson et al., 2002).
Such predictions have been used to identify hotspots of endan-
gered species (Godown and Peterson, 2000), to prioritise areas for
conservation (Chen and Peterson, 2002), and to establish suitable
locations for species translocations or cultivation (Jovanovic et al.,
2000; Beaumont et al., 2005). Wang and Ni (2009) investigated the
actual geographical distribution patterns and modelled the
potential distribution ranges of 5 Caragana species in northern
China. The results showed that under a future climate scenario, the
5 Caragana species all shifted northward and reduced their areas.
Wang et al. (2011) predicted present and future potential
distribution of Malus baccata, and results show that the overall
Please cite this article in press as: J. Jiang, et al., The wild relatives of grape in China: Diversity, conservation gaps and impact of climate change,
Agric. Ecosyst. Environ. (2015), http://dx.doi.org/10.1016/j.agee.2015.03.021
range of distribution reduces or even disappears. In our study, not
all suitable areas for GWRs will reduce, but several GWRs (21 out of
26) will experience potential range contractions, some (5 out of 26)
will show range extending, mainly because of the response of the
species to weather variables and seasonal response, echoing the
results of Warren et al. (2001) and Peterson et al. (2007) who have
found that species display a range of responses to climate change,
both positive and negative.
Species distribution model is an important tool for studying the
potential areas of species, simulating the biological invasion and
establishing the protection strategy of endangered species.
Compared with other statistic model, Bioclim model only needs
present date and more effective to deal with data, but there are
some imperfections in Bioclim model. First, the 19 climatic
variables were with equal importance. Second, the model is based
on the climatic variables only, ignoring the factors such as soil,
vegetation and human factor. Morano and Walker (1995) revealed
that three North America Vitis species varied in soil environments
and associated plant communties, among which V. berlandieri
adapts to drier habitats than the other species. Jin et al. (2013)
showed that the prediction of Phyllostachys edulis distribution in
China using both climate and soil predictors performed with
higher efciency and the climate factors played a driving role in the
simulation of the potential habitat, while soil factors mainly
impacted as limiting factors. And it was predicted that the intensity
of both the potential habitat expending and migrating was less
than using the climate factors only. If considering the factors of soil
constraints, it could be inferred that the distribution range of GWRs
may decreased much signicantly, and their suitable habitat
distribution scope will be reduced.
GWRs are light-demanding plant and disperse over long
distances through birds, monkeys and other animals in forests,
mainly distributed in mountain areas, such as the Qinling, Daba,
Wuling, Nanling and Wuyi mountains. Their distribution is
currently threatened by environmental destruction and human
activities. Under future climate change, the suitable area of most
GWRs will be further reduced and appropriate strategies should be
taken to mitigate this situation. The endangered mechanism
should be analyzed and population dynamics should be monitored
for endangered and rare GWRs, and then the impact of climate
change on endangering processes should be evaluated and
determined. Moreover, we can use complementary (in situ and
ex situ) conservation approaches to secure GWRs according to
prioritization purpose of protection and the level of endangering.
The extinction rate of some endangered species could be
accelerated by climate change, which can be preserved through
ex situ conservation. In addition, international cooperation should
be strengthened and international successful experiences should
be referred to promote the protection and utilization of GWRs in
China.
5. Conclusions
There are 39 species, 1 subspecies and 14 varieties of grape wild
relative species native to China and 19 species and 9 varieties are
the closest wild relatives to cultivated grapes according to the
Taxon Group concept. GWRs are distributed in every province in
China except for Xinjiang, but they are particularly abundant in
Jiangxi and Hunan provinces, the richest regions being mainly in
the Qinling, Daba, Wuling, Nanling and Wuyi mountains.
More than 22% of GWR can be found in natural reserves and are
therefore well protected, but 15 species are not found in any
natural reserves and require urgent action in terms of protection
and germplasm collection.
By comparing the potential distribution of GWRs under the
present and predicted future climate senerios, it is found that
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J. Jiang et al. / Agriculture, Ecosystems and Environment xxx (2015) xxxxxx 9

under future climate scenarios with doubled CO2, the suitable area
for 5 species will become larger as a result of climate change, but 21
GWR species will reduce signicantly, adding to the urgency for
renewed conservation protection of these valuable resources.
Acknowledgments
This research was supported by China Agriculture Research
System (CARS-30), the Agricultural Science and Technology
Innovation Program (CAAS-ASTIP-2015-ZFRI) and partially sup-
ported by the ChinaEuropean Union cooperation special project
of the Ministry of Science and Technology of China (0807) and the
ChinaUK Sustainable Agriculture Innovation Network (SAIN
project: CWR China). We thank Hongmin Zhang from Temple
University for his assistance with English editing and the
anonymous reviewers for helpful comments and suggestions
which improved the manuscript.
Appendix A. Supplementary data
Supplementary data associated with this article can be found, in
the online version, at http://dx.doi.org/10.1016/j.agee.2015.03.021.
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Agric. Ecosyst. Environ. (2015), http://dx.doi.org/10.1016/j.agee.2015.03.021