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The topical collection ‘Twenty years of biodiversity research and nature conservation in the Socotra Archipelago’, in short ‘Socotra biodiversity research and nature conservation’ was conceived at the 18th Friends of Socotra annual meeting and Socotra conference which took place at the Orto Botanico di Palermo, Palermo, Italy, 26–29 September, 2019. In total, 13 research papers are included in the collection, which covers a selection of the latest scientific progress on the fauna and flora of the terrestrial and aquatic environments of the Socotra Archipelago UNESCO World Heritage Site (Yemen). Topics include conservation, taxonomy, ecology, biology and biogeography. The focus is mainly on biodiversity conservation and aimed at identifying current challenges, trends and processes that may impact on local ecosystems and livelihoods, based on analysis of data collected over the last decades. With this collection, we wish to emphasise the importance of taking into account science-based conservation approaches in future strategic steps towards safeguarding the ecosystems of Socotra.
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Rendiconti Lincei. Scienze Fisiche e Naturali
https://doi.org/10.1007/s12210-020-00941-7
SOCOTRA BIODIVERSITY RESEARCH ANDNATURE CONSERVATION
Twenty years ofbiodiversity research andnature conservation
intheSocotra Archipelago (Yemen)
FabioAttorre1 · KayVanDamme2
© Accademia Nazionale dei Lincei 2020
The topical collection ‘Twenty years of biodiversity research
and nature conservation in the Socotra Archipelago’, in
short ‘Socotra biodiversity research and nature conserva-
tion’ was conceived at the 18th Friends of Socotra annual
meeting and Socotra conference which took place at the
Orto Botanico di Palermo, Palermo, Italy, 26–29 Septem-
ber, 2019. In total, 13 research papers are included in the
collection, which covers a selection of the latest scientific
progress on the fauna and flora of the terrestrial and aquatic
environments of the Socotra Archipelago UNESCO World
Heritage Site (Yemen). Topics include conservation, tax-
onomy, ecology, biology and biogeography. The focus is
mainly on biodiversity conservation and aimed at identifying
current challenges, trends and processes that may impact
on local ecosystems and livelihoods, based on analysis of
data collected over the last decades. With this collection,
we wish to emphasise the importance of taking into account
science-based conservation approaches in future strategic
steps towards safeguarding the ecosystems of Socotra.
1 Background
For centuries, islands and their biotas have appealed to biol-
ogists. Covering only 5% of the Earth’s land surface, these
‘living laboratories of evolution’ account for a high propor-
tion of endemics—about 20% of all known vascular plants
and 15% of all amphibians, mammals and birds are found
exclusively on islands (Da Fonseca etal. 2006). However,
due to several factors that are typical to insular ecosystems
such as isolation and small sizes of local populations, over-
whelming evidence has shown that species on islands are
easily driven to extinction. General factors are well known
and include the effects of exotic species, habitat degradation
and deliberate destruction (Sax and Gaines 2008; Caujapé-
Castells etal. 2010; Wood etal. 2017). Therefore, in con-
servation efforts, approaches for islands must be very dif-
ferent to those in the continent (Whittaker 1998). The rich
biodiversity of the Socotra archipelago forms no exception
to the brutal mathematical reality of how rapidly extinction
occurs on islands.
The Socotra Archipelago is situated in the western
Indian Ocean near the Horn of Africa and belongs to
Yemen. It consists of a few islands of which Socotra is
the largest (Fig.1), the latter covering a land area of about
3600 km2. Ever since the first scientific expeditions to
Socotra towards the end of the nineteenth century, biolo-
gists have been fascinated by this place with its strange
umbrella- and bottle-shaped trees. Most likely, Charles
Darwin would have loved visiting the ‘Galápagos of the
Indian Ocean’ instead of the Ecuadorian islands; how-
ever, strong monsoons used to cut Socotra off from the
world for about half a year and therefore opportunities for
research were few. Only in the last 50years has Socotra
been more accessible to researchers, in particular since the
establishment of an airstrip and an increase in traffic and
immigration over the last two decades (Van Damme and
Banfield 2011). The turn of the last millennium is also the
period when major conservation programmes started on
This paper belongs to a Topical Collection “Twenty years of
biodiversity research and nature conservation in the Socotra
Archipelago” edited by Fabio Attorre and Kay Van Damme,
conceived at the 18th Friends of Socotra Conference, 26–29
September, 2019, Orto Botanico di Palermo, Palermo, Italy.
* Kay Van Damme
kay.vandamme@gmail.com
Fabio Attorre
fabio.attorre@uniroma1.it
1 Department ofEnvironmental Biology, Sapienza-University
ofRome, Piazzale Aldo Moro 5, 00185Rome, Italy
2 Senckenberg Research Institute andNatural History Museum
Frankfurt, Senckenberganlage 25, 60325FrankfurtamMain,
Germany
Rendiconti Lincei. Scienze Fisiche e Naturali
1 3
the islands, which in 2008 received recognition as UNE-
SCO World Heritage Site, based purely on the criterion
of harbouring globally important biodiversity and a high
proportion of endemics (Natural WH Site).
About two decades ago, when the most recent episode
of research in Socotra had just started, a charity by the
name Friends of Socotra (FoS) was established. The non-
governmental, non-profit organisation consists of predomi-
nantly researchers and provides a multidisciplinary and
politically neutral platform for discussion, interaction and
synergies related to the conservation of the unique bio-
diversity and culture of these Yemeni islands. Through
annual conferences, a newsletter and small awareness or
pilot activities on the ground together with other organisa-
tions and local NGOs, FoS (www.frien dsofs oqotr a.org) is
the only organisation of its kind that focuses on scientific
efforts related to the unique cultural and natural heritage
of the Socotra Archipelago. The organisation is unusual,
as few islands or island groups in the world have an inter-
national, multidisciplinary group of scientists investing
voluntary time in promoting awareness for local science
and conservation. The group is driven entirely by the enor-
mous efforts by former and current volunteers from all
over the world who have supported the organisation in the
past two decades, and who have a heart for Socotra.
2 The topical collection
The topical collection ‘Socotra Biodiversity Research and
Nature Conservation’ materialised as a result of the impor-
tant synergies that are generated at the Friends of Socotra
meetings. In the last two conferences, which took place
in the Kingdom of Bahrain and in the Botanic Garden of
Palermo in Sicily, respectively, we proposed a topical col-
lection which crystallised into the current output. Repre-
sentatives from Yemen were present, and several Socotran
conservationists scientifically contributed to the meet-
ings and to the papers in this collection. At the Palermo
meeting (Van Damme and Livadiotti 2020), an important
awareness activity focusing on Socotra biodiversity was
launched, the UNESCO-FoS campaign entitled Connect-
2Socotra which took place in different scientific institutes
around the world (https ://en.unesc o.org/conne ct2so cotra ).
The topical collection focuses on 20 years of biodiver-
sity research and conservation of the terrestrial environ-
ment of the Socotra Archipelago UNESCO World Heritage
Site. The reasoning behind this collection is to provide
the scientific data and approaches to conservation and to
evaluate what can be learned from assessing the environ-
mental information that was gathered through international
collaborations in recent years. The collection focuses on
trends through the analysis of decades of data and includes
completely new observations as well as introducing new
techniques that can locally be used for biodiversity moni-
toring. Several studies in this topical collection result from
years of continued data acquisition under difficult field
conditions in close coordination with local authorities
and conservationists, with full support by the Ministry
of Water and Environment and the Environmental Protec-
tion Agency (EPA) of Yemen and EPA Socotra Branch,
to whom we are very grateful. The interaction between
conservationists from different cultures working on a com-
mon goal over many years, continuing essential scientific
work for biodiversity during a time of hardship and war in
Yemen, has led to fruitful collaborations.
Studies that examine actual changes in the Socotran
ecosystems over time are few (Van Damme and Banfield
2011; Madĕra etal. 2019). Nevertheless, such studies are
crucial to assess imminent extinction risks and to strate-
gise proposed solutions. The main threats to the terrestrial
environment are well known (Van Damme and Banfield
2011) and have been highlighted since years by the World
Heritage Committee for Socotra. The IUCN Conservation
Outlook (2017) has put the digit at an alarming ‘Signifi-
cant Concern’ for Socotra. This topical collection also
serves the purpose of a warning by the scientific com-
munity: as shown here, there are clearly processes on
Socotra that can lead to imminent extinctions. And the
Fig. 1 Socotra Island is part of a small yet significant Yemeni archi-
pelago, situated in the western Indian Ocean just off the Horn of
Africa. It is the largest island of Arabia. Since 2008, the Socotra
Archipelago has been listed as one of Yemen’s UNESCO World Her-
itage Sites, in particular for its rich biodiversity and high number of
endemic species. Image from Google Earth, modified by Kay Van
Damme
Rendiconti Lincei. Scienze Fisiche e Naturali
1 3
extinction of a single individual or a small population,
whether it belongs to a species of plants or animals, mat-
ters. For example, the loss of a single individual of the
iconic Socotra Dragon’s Blood Tree (Dracaena cinnabari)
would not only affect the entire vegetation that lives shel-
tered below the canopy and the animals associated with it
(Rejžek etal. 2016; Vasconcelos etal. 2020), but it may
also change the amount of water that is part of the island’s
hydrological cycle on which human life depends (Kalivo-
dová etal. 2020). Often, effects that we see today are the
results of trends that have been set in motion over the last
decades, and which are accelerated through global warm-
ing and other, more direct, human interventions.
This is the case for the frankincense tree Boswellia elon-
gata (Fig.3) which is one of the most important endemic
plant species in Socotra. Not only is the most typical Socotra
frankincense tree endangered, but it also plays a key role in
the island’s culture and has potential benefit for economy, as
perhaps it did in a distant past. A close study of individual
trees from aerial photographs now provides the first direct
visual evidence of this species’ decline and indicates the
imminent extinction of its largest subpopulation. Research-
ers witnessed a clear reduction in the largest subpopulation
of Boswellia elongata on Socotra Island between 1956 and
2017, a period of 61years, by about 78% (Lvončík etal.
2020). Through experimental designs and assessment
of the potential causes, the decline has been attributed to
both natural and human-mediated factors. Senescence of
the tree stands and decades of overgrazing by goats (now
roaming more freely because of a shift in culture, i.e. use
Fig. 2 Overmature stands of one of the archipelago’s unique and
endangered frankincense tree species (Boswellia elongata) in an
overgrazed Socotran landscape before (left) and after (right) damage
caused by climate change (cyclones in 2015). Photos from Homhil
Nature Sanctuary, eastern Socotra Island, by Kay Van Damme (May
2010) and Petr Madĕra (2016)
Fig. 3 The analysis of data that
has been collected over decades,
now provides an objective view
on how the immune system of
Socotra’s terrestrial environ-
ment, on which local people
depend, is compromised. Stra-
tegic efforts to increase local
resilience of plant and animal
populations are vital to ensure
that future generations can ben-
efit from the ecosystem services
of the iconic species that have
made the island so famous. The
Socotra Dragon’s Blood Tree,
for example, does not only har-
bour a wide range of endemic
species, it also plays a role in
the island’s hydrological cycle.
Photo by Kay Van Damme,
Socotra Island, 2014
Rendiconti Lincei. Scienze Fisiche e Naturali
1 3
of the rangeland), have made natural regeneration of most
ground-dwelling palatable plants in accessible areas nearly
impossible, in particular in times of drought when grazing
pressures intensify (Miller and Morris 2004; Lvončík etal.
2020). The process is catalysed by non-selective overarching
events such as climate change, which could potentially give
the final nudge for Socotran species that are on the edge at
the downward slope towards extinction (Van Damme and
Banfield 2011).
Global warming recently manifested itself in Socotra in
the form of extreme weather events (cyclones and torrential
rains) which have caused damage to the terrestrial environ-
ment. Physical destruction to the frankincense trees caused
by unusually strong winds could be clearly attributed to two
cyclones in 2015 and are accountable for more than a third
of the individual trees that have disappeared in one of the
island’s largest populations (Fig.2). Weakened by the com-
bination of overgrazing and damage by wind, the popula-
tion was attacked by a local species of bark beetles on top
of that. If the trend continues, the iconic subpopulation of
frankincense trees in a well-known protected area on the
island (Homhil Nature Sanctuary) will be gone in only a
few decades from now (Lvončík etal. 2020). The events are
not localised; we may expect that the majority of exposed
areas where this and other endemic ground-dwelling tree
species occur on the island are facing a similar situation.
The rate of extinction is specific to biological attributes of
the species, however, studies show similar trends for other
endemics, like Dracaena cinnabari, albeit over centuries
rather than decades (Madĕra etal. 2019). First estimates of
age indicate that many trees of the latter species can easily
be several centuries old (Madĕra etal. 2020). In some spe-
cies, relatively less palatable to goats, regeneration is good,
such as in the endemic Dendrosicyos socotranus, Socotra’s
famous cucumber tree. Habrová etal. (2020) found regenera-
tion for this species in 77% of the locations studied. How-
ever, to avoid an entirely skewed landscape that is mainly
dominated by grazing-tolerant species, efforts are needed
because ground-rooting species are truly at risk. The grazing
impacts on vegetation is stronger during dry periods when
there is less choice for alternatives, as exclosure experiments
show (Lvoncik etal. 2020).
Ecosystems dominated by trees in the Socotran land-
scape are not as widespread as it would appear to the out-
side. Satellite image analysis using available state-of-the-art
approaches combined with ground-trouthing indicates that
the Socotra highlands and mountains harbour the majority
of the trees (Riccardi etal. 2020). Statistical analysis sug-
gests that the overall distribution of trees on the island is still
largely governed by natural factors such as soil availability
and local climate, less so by direct human impacts. For now.
As the population grows, these impacts may increase further
as well. Such impacts can be monitored also using remote
sensing technologies. Vegetation productivity in Socotra was
assessed over ca. 20years using NVDI (normalised differ-
ence vegetation index) (Rezende etal. 2020). The results
show that overall the island seems very dynamic in terms
of vegetation productivity, however the mountain areas and
wadi slopes show a clear decrease over the past decades and
in particular over the last few years (Rezende etal. 2020).
The decrease could be attributed to the long-term effects
of grazing combined with the effects of extreme weather
events, leading the loss of vegetation, as in the Boswellia
case study, and the loss of soil linked to erosion.
Plants are also important for many endemic vertebrate
species on the island, such as birds and reptiles that are
strongly dependent to the availability of healthy stands of
woody vegetation. About 90% of the terrestrial reptiles are
endemic to the archipelago, and several of those rely on the
availability of the trunks and branches of trees or bushes
(Fasola etal. 2020). The endemic gecko Hemidactylus dra-
caenacolus, assessed as critically endangered in the IUCN
Red List, is even found exclusively on tree trunks of the
endemic Dragon’s Blood Tree (Vasconcelos etal. 2020).
Therefore, the declines or the disappearance of vegetation
through a combination of factors, may accelerate the direct
extinction of associated species in the absence of vegetation
cover. Considering the rapid changes in vegetation over the
last decades (Lvoncik etal. 2020; Rezende etal. 2020), the
fact that some endemic reptiles have a total area of occupa-
tion of less than 10 km2 (Fasola etal. 2020) and the pres-
ence of invasives such as rats and cats, extinction scenarios
for these island endemics are realistic if there is no direct
intervention.
On an island, everything is closely connected, and so are
people to their environment. The vegetation does not only
play a key role in creating niches for other plant and animal
species, but also may provide another important ecosys-
tem service as part of the hydrological cycle of the island.
Kalivodová etal. (2020) measured the amount of horizontal
precipitation caught by individual Dragon’s Blood Trees.
The authors estimated that at the highest altitudes (ca.
950–1550m a.s.l.), the mean annual horizontal precipitation
below the tree crown is about 790mm, exceeding 40% of the
total annual precipitation. Considering the relatively slow
growth of Dragon’s Blood Trees (Madĕra etal. 2020), con-
servation strategies to preserve these assets will need local
involvement for the duration of more than a human lifetime.
The freshwater environments also show a decline in
indicator species. A first analysis of 123years of drag-
onfly records in the archipelago suggests a decline in the
relative richness of these conspicuous insects in the area of
the coastal capital of Socotra, Hadiboh (Van Damme etal.
2020). Although species richness in this animal group is
not high, the pattern is clear and it affects several species.
The latter is relevant to the protection of local wadis, but
Rendiconti Lincei. Scienze Fisiche e Naturali
1 3
particularly shows the need to protect coastal lagoons, which
harbour a rich diversity of birds, fish and arthropods. A con-
spicuous dragonfly called the phantom flutterer (Rhyothemis
semihyalina), disappeared from Socotra’s coastal lagoons
in the east and the west. Even though the latter study is not
based on standardised surveys, it is the first long-term study
assessing changes over time in animal species richness on
Socotra. In this case the decline was attributed primarily
to habitat alteration through urban development. The study
shows also that there is a cultural dimension to these insects
locally—the people of Socotra recognise dragonflies as a
sign of good water and they have no less than six terms in
the unique language to describe them. Such positive inter-
action towards invertebrates can be a very valuable asset in
local conservation, for example to protect the iconic beauti-
ful Socotra bluet (Azuragrion granti), the archipelago’s only
endemic damselfly, Near Threatened in the IUCN Red List
(Van Damme etal. 2020).
The above, habitat degradation and climate change, are
all well-known factors that impact on biotas worldwide.
However, arguably the worst known drivers of extinctions
on islands are the invasive alien species. Exotic species
are a well-known threat to biodiversity on Socotra, but the
arrival of new and unwanted guests may strongly impact
on local economy and culture as well. Witt etal. (2020)
present the first records of detrimental plant pests, the red
palm weevil (Rhynchophorus ferrugineus) and the dubas bug
(Ommatissus lybicus) on Socotra, two invasive alien spe-
cies which are known to strongly reduce date palm yields
in the region. Another new invasive on Socotra is recorded
in Hula and Niedobová (2020), the Mediterranean recluse
spider, of which the bite can pose a risk to children. Based
on successful eradication programmes in other places in the
world, Witt etal. (2020) suggest that more measures are
urgently needed to protect the island and its biodiversity
and livelihoods, from exotics. Such measures would include
phytosanitary approaches, quarantine facilities at the port
and airport, and most importantly a good communication
between all stakeholders to mutually arrive to the best strate-
gies (Witt etal. 2020). Even though such suggestions have
been made repeatedly in recent years and despite efforts as
part of ongoing projects, and a priority issue as identified by
the World Heritage Committee, such measures were unfor-
tunately never achieved on the ground. However, the delays
in implementation increase the risk of more pest incursions
as evident in the presence of the red palm weevil and other
invasive alien species. Therefore, such exotics pose huge
risks, not only towards the unique biodiversity of Socotra,
but also for local livelihoods, the unique culture and human
health. More invasives may be present on the island yet
remain undetected.
Despite the challenges, Socotra is still at this moment,
a treasure trove of biodiversity. While freshwater
environments are largely well studied, the terrestrial envi-
ronments still harbour new species that rest in existing col-
lections. For example, new species are being described from
collections from the first large (among the recent biological)
expeditions, now 20 years ago. People who describe spe-
cies in animal and plant groups are part of an endangered
population themselves; however they keep carrying out this
important and tedious taxonomical work. In this topical col-
lection, malacologists Neubert and Bochud (2020) describe
two new endemic species of molluscs from the terrestrial
environments of Socotra. Also, in the flora of Socotra, still
a large amount of work remains to be done, in particular in
assessing the genetic diversity of plant populations that may
or may not harbour cryptic species, although much more
research is needed for single species (Giovino etal. 2020).
All these taxonomical works can help to understand patterns
of diversity, areas of higher richness, and help to even look
into biogeographical patterns of richness and evolution, as
Purchart etal. (2020) have started for butterflies, darkling
beetles and spiders in the archipelago. Comparing such pat-
terns in diversity and comparison to other islands can be key
to global research in biodiversity and biogeography.
3 Outlook
As the publications in this topical collection contribute to
our knowledge of the terrestrial and aquatic environments in
the Socotra archipelago, they thereby may help to identify
the challenges that other fragile insular ecosystems and arid
lands in the world face. These challenges are global and have
been identified on Socotra years ago, before the effects were
as visible as now. It is clear from the studies herein that if
the current trend continues, the Socotran ecosystems may
actually rapidly lose iconic endangered species and a per-
haps equally fragile endemic culture over the next decades
or centuries (Fig.3). This trend is realistic and should be met
with matching efforts in conservation.
Socotra has changed. The changes in the terrestrial eco-
systems now have become tangible and seem to manifest
themselves clearly when examining processes over a few
decades. Such processes can only be observed through rig-
orous scientific approaches and targeted monitoring, cover-
ing years of data. Such research, which only started at the
turn of the millennium in Socotra, is vital to strategise con-
servation priorities, while practical activities are needed to
focus on applied science that assesses positive effects from
interventions. Primarily environmental factors, more than
human factors, seem to currently drive the dynamics of the
terrestrial ecosystems in Socotra, sometimes in pulses of
increased intensity, and these effects likely will continue to
catalyse changes on the islands and its inhabitants in the
years to come. Climate change impacts have also clearly
Rendiconti Lincei. Scienze Fisiche e Naturali
1 3
revealed the sensitivity of local endemic plant species, which
will need to retain a strong resilience and positive human
interventions to avoid extinction. However, the new studies
in this topical collection reveal that not only climate change,
but also cultural and socio-economic effects play a role in
affecting the landscapes, albeit through perhaps slower and
sometimes more localised processes. These include effects
of (i) overgrazing, leading for example to overmature tree
populations of palatable species, (ii) urbanisation, altering
coastal areas including lagoons and wadis, and (iii) new
invasive alien species which are more likely to arrive now
due to an increase of import and lack of control measures
at entry pathways, and which may have a devastating effect,
not only on biodiversity, but also on food security and local
livelihoods. The combined effect of overgrazing and climate
impacts, seem to also lead to loss of vegetation in the moun-
tains, Socotra’s local biodiversity hotspot. Even though we
have an image of the Socotran people as being stewards of
their environment for centuries (Van Damme and Banfield
2011), new challenges arise and cultural shifts have been
disrupting the bond between people and their environment
since decades, as is evident in the overgrazing effects. The
local capacity to cope can benefit from strategic and cost-
effective solutions, which can be helped by science.
The immune system of Socotra is now compromised. The
current status of the ecosystems is unravelled by impacts that
painstakingly reveal slower underlying mechanisms. Extinc-
tion is a relatively slow process; however as it is known to
be accelerated strongly in island biotas, we may soon be
running out of time to protect Socotra’s most iconic flagship
species and the biota associated with them. The causes and
pathways for extinction are well known, and if not countered
by immediate strategic conservation action involving local
communities, several typical species will clearly not survive
the coming centuries. If the trend continues, future genera-
tions might be able to visit a Socotran frankincense tree only
in a botanical garden, accompanied by a little plaque saying
‘extinct in the wild’.
We believe there is time to turn the tide. Human impacts
on nature can be reduced as shown in other places in the
world. It needs all crew aboard working together and some
positive currents to avoid a ship from crashing. In order
for the ecosystems to be resilient in the face of upcoming
challenges, it will require all hands on deck. Science-based
strategies, cooperation and community involvement are key
and have shown to be effective in other highly valuable bio-
diversity areas in the world.
As Socotra is recognised by UNESCO as a Natural World
Heritage Site, there is a large interest in supporting the local
environment, materialised in several ongoing conservation
projects. If such efforts are managed properly and in full
support of the local authorities and local communities to
help against the coming challenges, major challenges can
be faced. Improved biosecurity measures are an example of
cost-effective priority measures to avoid loss of future local
livelihoods and biodiversity, and to avoid more costly future
interventions to eradicate invasives. To keep promoting the
communication of scientific outputs towards a wider pub-
lic, providing science-based options to conservation, and to
keep spreading awareness about the island’s uniqueness, an
organisation like FoS and collaboration with local environ-
mentalists and authorities are key. The latter is illustrated by
the fact that no less than five papers in this topical collection
include indigenous co-authors from Socotra Island.
We hope that these scientific contributions related to
conservation science in Socotra, presented in this topical
collection, can help to strategise future activities and help
reverse some adverse impacts from climate change and other
processes that drive species to extinction.
Acknowledgements Several studies of the Topical Collection were
partially funded by the UNE-GEF Project #5347 which has been locally
executed by the Ministry of Environment (Yemen) and the Environ-
mental Protection Agency (Yemen): “Support to the Integrated Pro-
gram for the Conservation and Sustainable Development of the Socotra
Archipelago”. A special thanks is due to Friends of Socotra (FoS)
and its volunteers, for the enabling environment that stimulated this
collaborative effort. This topical issue could not have been realised
without the efforts of all contributors, including authors, reviewers
and colleagues helping in the field surveys. We also thank the Italian
Agency for Development Cooperation for the financial and institutional
support for the organization of the Friends of Socotra meeting that took
place at the Botanic Garden of Palermo, Italy, 26–29 September, 2019.
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... Therefore, it is undoubtedly a global hotspot for biodiversity. It offers beautiful wildlife scenery characterized by colorful coral reefs, coastal fish, and amazing crustaceans [16,17]. ...
... The proposed management plan for Socotra recommends that community-based tourism, such as small-scale and community-driven development, should be a thrust in formulating tourism development [12]. The plan capitalizes on including the local communities in tourism activities, ensuring that tourism profits are equitably available to all the stakeholders and are favorable for the environment [15,17]. ...
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Socotra Island, a UNESCO World Heritage Site, faces unique challenges in balancing tourism development with conservation efforts. This research focuses on the conservation of Socotra through tourism development. It explores the concept of ecotourism and its significance in promoting sustainable practices. This paper addresses ecotourism development's social, economic, and environmental impacts and highlights its benefits for local communities. This study analyzes the status of tourism in Socotra and evaluates its effectiveness, considering the perspectives of the local community, tour guides, tourism companies, and tourists. Finally, this research highlights the potential for sustainable tourism development on Socotra Island. It addresses key challenges, including effective tourism management, coordination, waste disposal, and community involvement. This research contributes to the growing body of knowledge on sustainable tourism development in ecologically sensitive regions. By advocating for a collaborative approach, this research offers valuable insights for policymakers, conservation practitioners, and tourism stakeholders working in Socotra and similar destinations.
... Taking into account the ongoing decline of the terrestrial ecosystems on Socotra for various reasons, ecological restoration and reforestation activities of this endemic tree species are urgently needed [12,30,31]. To identify the most suitable areas for such interventions, a model, based on the Dragon tree as a case study and applicable to other species, integrating suitability and accessibility analyses is proposed in this study. ...
... The unique umbrella-shape of the crown constitutes a fundamental natural system that traps humidity brought by the ocean breeze and helps recharge the shallow aquifers of the island [18]. Therefore, the disappearance of these dragon's blood trees would have a significant impact on the freshwater resources [18], biodiversity [58], and touristic attractiveness of Socotra [30], in addition to the potential loss of key human-nature linkages that the island is known for. ...
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We examine the distribution of Dracaena cinnabari, the Socotran Dragon’s Blood Tree, an endangered species endemic to the island of Socotra (Yemen)—and we propose an accessibility approach to its conservation, taking the proximity of local communities and land users into account. Using the present occurrence of D. cinnabari, we applied a machine learning algorithm (random forest classifier) to estimate the potential distribution of the species across the island (overall validation accuracy of 0.91) based on available climatic and physiographic parameters. In parallel, we used an accessibility methodology to generate a map of the energy cost of accessing potential areas from the villages. This community-focused accessibility map, combined with the potential distribution map of Dracaena cinnabari, could contribute to decision-making processes related to long-term ecological restoration and reforestation activities. With our case study, we wish to emphasize that user-focused efforts and the implementation of sustainable land practices should play key roles in conserving endangered tree species.
... Socotra Island, the largest in the archipelago, is home to several iconic endemic tree species, including frankincense (Boswellia spp.), myrrh (Commiphora spp.), and the Socotran dragon's blood tree (Dracaena cinnabari Balf.f.). These trees hold not only ecological significance but also cultural importance (Miller and Morris 2004;Attorre and Van Damme 2020). However, despite their value, these endemic species are currently facing severe threats, with the primary concern being a lack of natural regeneration associated with overgrazing Hamdiah et al. 2024). ...
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Question Plant–plant facilitation is a critical ecological mechanism in arid environments, influencing biodiversity and ecosystem resilience globally. Shrubs often serve as nurse species, enhancing tree regeneration and sheltering plant communities, particularly in overgrazed or degraded habitats. In this study, we examine the role of shrubs as nurse species in the Socotra Archipelago (Yemen), a biodiversity hotspot in which several endemic tree species, including the iconic frankincense ( Boswellia spp.), myrrh ( Commiphora spp.), and Socotran dragon's blood ( Dracaena cinnabari ) trees, are threatened. This is largely due to a lack of natural regeneration caused by the combined effects of overgrazing by goats, sheep, and climatic events such as extreme droughts and cyclones. To aid conservation of threatened trees in arid regions, nature‐based solutions are urgently needed to help tree regeneration. Effective nurse plants have this potential, particularly in arid environments. We therefore examined the role of thorny, poisonous, and/or unpalatable shrubs as nurse plants in sheltering threatened plant communities, with a focus on woody species in an arid insular context. Study Area The Socotra Archipelago (Yemen) situated in the western Indian Ocean, east of the Horn of Africa. It is a biodiversity sanctuary and a UNESCO Natural World Heritage Site. Methods Vegetation surveys were conducted in 144 paired plots under six common shrub species and adjacent open areas. Community data, environmental variables, and functional traits were analysed using RLQ and fourth‐corner analyses, while Linear Mixed Models (LMMs) evaluated the effects of environmental variables and nurse species on key functional traits based on Community Weighted Means (CWMs). Facilitation effects were quantified using the Relative Interaction Index (RII). Results Our analysis revealed significant variations in species composition and functional traits between under‐canopy and open‐area plots. Certain shrubs, such as Cebatha balfourii , facilitated significantly higher species richness under its canopy compared to open areas. Elevation and grazing pressure influenced these interactions, with notable effects on functional traits like the occurrence of legumes and tree growth forms. Buxus hildebrandtii was less effective in supporting species richness, while C. balfourii, Lycium sokotranum , and two Cissus species exhibited significant positive facilitation. The LMMs confirmed the importance of environmental variables and nurse shrub characteristics in shaping plant community dynamics. Conclusions The results highlight differences in the facilitative potential of the studied species, with some showing a stronger ability to act as nurse shelters through microhabitat amelioration and protection from herbivory. The presence of tree species under shrubs is confirmed, and the role of these nurse species in supporting diverse plant communities provides critical insights for conservation strategies, supporting biodiversity resilience and sustainable management in degraded landscapes like Socotra Island and other arid environments. Future efforts should focus on leveraging nurse shrubs to mitigate environmental pressures and enhance ecological restoration, in particular to help conserve range‐restricted and threatened plant species.
... The Western Indian Ocean Islands, delimited at the north by Socotra and at the south by Madagascar, have been identified as a global biodiversity hotspot (Myers et al. 2000;Attorre and Van Damme 2020;Agnarsson et al. 2015). Although extremely variable in size, and geological origin and history, these islands harbour a terrestrial fauna notable for striking patterns of radiation and endemicity (Legros et al. 2020;Bergsten and Biström 2022;Sherlock et al. 2024). ...
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... Several challenges to Socotra's biodiversity affect entire habitats including the shrublands, woodlands and forests, the latter of which are relatively limited in distribution . The rich terrestrial flora of Socotra is strongly impacted by overgrazing which has started centuries ago and which has partly shaped the plant communities by counter-acting regeneration of palatable species, leading to a general decline of habitat-and soil quality (Van Damme and Banfield, 2011;Attorre and Van Damme, 2020;Maděra and Van Damme, 2022). Affected by overgrazing, environmental factors such as soil and climatic factors play major roles in shaping the current distribution of trees in Socotra . ...
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Societal Impact Statement Conserving frankincense trees (Boswellia) is crucial for both ecological and socio‐economic reasons. Surveying these trees in the field and using remote sensing unmanned aerial vehicles in the Socotra Archipelago, we found that Socotran frankincense trees are threatened by forest fragmentation, overgrazing, and increasingly frequent extreme climate events. A better understanding of the distribution and the threats of these important insular species will improve the conservation policy of the local authorities and benefit local communities in the Socotra Archipelago. At the same time, this work serves as a good practice example to guide conservation efforts for other culturally important threatened tree species around the world, therefore helping to sustain local livelihoods, fostering ecological resilience, and supporting socio‐economic stability. Summary Globally, frankincense trees (Burseraceae: Boswellia) are increasingly under threat because of habitat deterioration, climate impacts, and the olibanum trade. Despite harboring nearly half of the species in the genus, up‐to‐date insights are lacking for the insular endemic frankincense trees of the Socotra Archipelago UNESCO (United Nations Educational, Scientific and Cultural Organization) World Heritage Site (Yemen). We combined georeferencing of individual trees in the field with remote sensing applying unmanned aerial vehicles (UAVs) to evaluate Boswellia distribution and (sub)population sizes in the entire Socotra Archipelago. We counted 17,253 trees across all 11 taxa and we surveyed almost 55% directly in the field, collecting individual information on threats and health indicators. We estimate that the current total population sizes of the relatively common Socotran Boswellia taxa (Boswellia elongata, Boswellia popoviana, and Boswellia ameero) consist of a few thousand mature individuals with fragmented distribution of which a large proportion occurs in highly disjunct relictual stands, while the more range‐restricted species survive only through a few hundred (Boswellia nana and Boswellia samhaensis) to fewer than a hundred trees (Boswellia scopulorum). Our field data show that the Socotran frankincense trees are threatened by fragmentation and overgrazing resulting in a lack of natural regeneration, in combination with effects of extreme climate events (e.g., higher frequency and intensity of cyclones and prolonged drought) and potential future infrastructure developments; the species are less impacted by resin collection. We provide recommendations to strategize urgent protection of the declining Socotran frankincense trees, and we update their conservation status, resulting in an endangered status for seven and a critically endangered status for four taxa.
Chapter
The Socotra Archipelago is universally renown because of its biodiversity, nonetheless the cultural heritage—such as the centuries-old culture of small fishing communities—is not of minor importance. Even though Socotra Island is still hardly accessible to mass tourism, the number of tourists visiting it has been increasing and fishing tourism is growing, attracting amateurs and professionals from all over the world. Our research, based on semi-structured interviews and observation during fieldwork, aims at investigating if and how tradition-based fishing tourism could be a source for extra income for fishermen of Socotra and at the same time how it could incentivize sustainable tourism development on the island. The paper analyses fishermen’s tangible and intangible heritage, as well as the current fishing tourism offer in order to outline a concrete proposal of community-based tourism.
Chapter
To learn about the traditional and indigenous practices that have maintained Soctora’s rich ecosystem, short-term qualitative field research was conducted during a two-week excursion to Socotra in May 2022. Using a local guide, data was collected through the authors’ observations of human interactions with the environment; engagement in local activities (e.g., hiking, snorkeling); interviews with key informants (e.g., government officials, university faculty, community leaders) and community members; and review of government reports. Sustainable indigenous practices were identified related to (1) food supply and storage; (2) water collection, use and storage; (3) land use; and (4) waste disposal. Socotra’s indigenous people have cared for and lived off the island’s resources for centuries, perhaps millenia, using traditional practices that maintained the fragile and unique biodiversity that exists today (Damme and Banfield in Zoology in the Middle East 54:31–88, 2011). Currently, there is a tension between conservation of Socotra’s unique biodiversity; development for basic infrastructure (i.e., water, waste, energy); development for tourism; and local and indigenous people and practices. As the population, tourism and development increase, the delicate balance that has maintained the environment and locals for millennia, is threatened. In addition, climate change and environmental degradation threaten Socotra’s ecosystems. Although it is too simple to say that indigenous practices are the solution to environmental degradation (Briggs in Progress in Development Studies 5:99–114, 2005), scholars have suggested we look to traditional practices used by local and indigenous peoples for ideas that may offer some hope (Hernandez in Fresh banana leaves: healing indigenous landscapes through indigenous science, North Atlantic Books, 2022; Nelson & Shilling in Traditional ecological knowledge. Cambridge University Press, 2018; Singh et al. in Journal of Ethnobiology and Ethnomedicine 17(50), 2021). This research attempts to fill part of that need.
Article
Ethnopharmacological relevance: Due to the long biogeographical isolation since ancient times, Soqotra Island (southern Yemen) is rich in medicinal endemic plants. However, information on their traditional therapeutic knowledge is scarce in scientific literature and still not completely documented. Aim of the study: (1) Documentation and search the native traditional therapeutic knowledge focusing on the medicinal plants which are endemic to Soqotra (2) Identification of the important endemic plants treating diseases by quantitative analysis of the collected data. Materials and methods: Ethnobotanical data were collected from 105 local informants through open interviews, semi-structured questionnaires and field survey in February 1990 to document the traditional medicinal plants in Soqotra Island. Of them we reported in this study the data of the endemic species, due to their wide biodiversity in the island. The quantitative data were analyzed for use reports (URs) and the informant consensus factor (ICF). The reported diseases were classified according to the International Classification of Primary Care-2 (ICPC-2). New reported medicinal plants in this study were identified compared to literature review. The level of traditional Soqotri knowledge among Islanders was observed during 1990-2003. Results: In the study, 86 medicinal endemic plants with 7654 therapeutic URs were explored for 69 ailments types belongs to 11 (ICPC-2) groups. The highest ICF value (0.99) were calculated for both eye and ear diseases categories, followed by general and unspecified ailments (0.98), digestive and skin (0.97 for each), and musculoskeletal ailments (0.96) among all diseases categories. While the highest URs (2276) were identified for skin diseases, followed by digestive diseases (URs = 2146). All the 86 recorded medicinal plant species were endemic to Soqotra, belonging to 57 genera and 29 families. Euphorbiaceae and Burseraceae were the dominant families, followed by Apocynaceae, Fabaceae and Asteraceae. Shrubs were the dominant growth form (41 species, 47.67%), followed by herbs (27; 31.4%), and trees (14; 16.28%). Leaves (45 species; 31.25%) were the most frequently used parts, followed by stems (34 species; 23.61%) and barks (species 23; 15.97%). The most cited preparation method was paste for 51 species followed by liquid exudates (48 species) and powder (26 species). Based on the comparative literature review, we recorded 36 new endemic medicinal plants and reported new traditional therapeutic uses for the remaining 50 endemic plants for the first time. On the other hand, all the reported endemic medicinal plants have been enlisted in the IUCN Red List of Threatened Species, of them 21endemic species enlisted as highest threatened species. Conclusion: According to the quantitative data analyzed, Soqotra Island has a valuable traditional knowledge in several medicinal endemic plants that used by local people in the primary health care until the study time at 1990. This study emphasizes the importance of early time in documenting traditional indigenous knowledge before losing and to protect and conserve the endemic plants against highly endangered. Consequently, these ethnomedicinal data could be used in the future as guide for pharmacological and phytochemical studies to discover new therapeutic drugs.
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Dragonflies are important bioindicators of aquatic ecosystem health. We discuss the distribution and richness of the 22 dragon- and damselfly species known from the Socotra Archipelago UNESCO World Heritage Site (Yemen) in the western Indian Ocean. We compiled 841 records from literature, field data and museum collections, covering a surveying period of 123 years (1896–2019). Distribution of the only endemic, the Socotra Bluet Azuragrion granti (McLachlan, 1903), an indicator of pristine lotic waters, is updated. Relatively higher dragonfly species richness occurs in eastern Socotra where suitable aquatic habitats are present all year. We observed a significant drop of the number of taxa over several decades in areas in the Hadiboh Plain where human development impacts are strongest. We discuss the indigenous people’s perception of dragonflies to assess the potential of these invertebrates as local flagships for aquatic biodiversity conservation. Our knowledge of the Socotran Odonata remains incomplete, yet they are now among the best studied animal groups in the archipelago. A new record of an Oriental Anax indicates that the island’s dragonfly fauna may yet harbour surprises. Our compilation of odonate records is the first for Socotra and serves as a baseline for future monitoring of the archipelago’s vital aquatic habitats. We converted all records into a geodatabase which we uploaded to an online, freely available web mapping server and mobile application to facilitate research and conservation of the Socotran dragonflies ( www.dragonfliesofsocotra.com ). Graphic abstract
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The Socotra Archipelago is a unique hotspot of biodiversity, particularly for terrestrial reptiles (28 endemic out of 31 native species) whose ecology is known only in general terms. We describe the ecological preference and the daily activity of 21 species of terrestrial reptiles, geckos, lizards, skinks, snakes and chameleon, from the largest island of the archipelago. The database includes 6950 records collected by 260 standardized transects from 2007 to 2013. The sister species Haemodracon riebeckii–H. trachyrhinus with very distinct body sizes, overlapped in their distribution but segregated by habitat and substrate. Three pairs of sister species with great phylogenetic affinity and similar morphology (Pristurus insignis–P. insignoides; Pristurus guichardi–P. obsti; Hemidactylus dracaenacolus–H. granti) were only partially segregated by distribution, habitat, substrate and body size, in the order of decreasing frequency. On the contrary, the other, phylogenetically more distant species of geckos, widely distributed and less similar in morphology, overlapped completely in their habitat use and distributional range. Such patterns suggest that strictly related species reduced their competition by segregation in different habitats and temporal niches and by non-overlapping distribution within the relatively small island of Socotra. On the other hand, the activity rhythm was similar within each genus. The unique biodiversity of Socotra endured up to present, thanks to a human impact lower than for most other inhabited islands, but the pace of change has accelerated recently. Conservation measures are urgent, particularly for the species of endemic geckos with a distribution range as small as 10 km2.
Article
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The presence of Loxosceles rufescens (Dufour, 1820), commonly known as the Mediterranean recluse spider, is recorded from Socotra Island (Yemen) for the first time. This originally circum-Mediterranean species is potentially harmful to humans, considering the strong cytotoxicity of the venom and the species often inhabiting human settlements. It has been shown that L. rufescens can easily be introduced to non-native areas with transported goods, and is hence an alien invasive species to nearly all other continents and many Islands. From the region, it was recorded from mainland Yemen, Saudi Arabia and the United Arab Emirates. We conducted a wide survey and recorded the species across the entire Island of Socotra, in both natural and artificial habitats.
Article
We assessed seven decades of change in the largest known population of the endangered endemic Boswellia elongata Balf. F. (Burseraceae) on Socotra Island (Yemen). To quantify the population change we evaluated tree number and locations on digitized images from various sources in the period 1956–2017 and combined this with direct field measurements of the population between 2011 and 2017. Our study reveals that the Homhil Nature Sanctuary B. elongata population shows a continuous decline since 1956. The steady but slow natural decline was strongly accelerated by two catastrophic cyclones in November 2015, when 38% of the trees were directly destroyed by strong winds. During the following 2 years 29% of the remaining trees died additionally. The remaining population has a bell-shaped size distribution; most trees are around 40 cm in diameter (range 18 to 70 cm). Tree ring analysis of 11 dead trees with a diameter of 29 to 44 cm without bark, resulted in estimated tree ages between 80 and 101 years. We estimate that similar-sized trees showing strong signs of senescence have a maximum age of a little over 100 years. The age structure of the Homhil population is, therefore, unbalanced with large sized trees prevailing. Natural regeneration is absent for decades. Viable seeds are available and have been shown to germinate, but the development of seedlings into saplings is a bottleneck. If the decline continues at the current rate, only 30 trees will remain there in 2036. Protection, planting and awareness activities are needed to keep this unique frankincense tree in Homhil Nature Sanctuary.
Article
The importance of populations of the Socotra dragon’s blood trees (Dracaena cinnabari Balf.f., 1882) for horizontal precipitation capture was estimated for the first time. Dragon’s blood trees (Asparagaceae) inhabit semiarid cloud forests with low levels of rainfall that strongly depend on additional moisture from fog and clouds. Our estimation was based on a detailed description of the aboveground biomass structure of stands of the dragon’s blood trees on Socotra Island. The aboveground biomass was described using non-destructive methods. An example of a well-preserved forest was measured in terms of the basic biometric characteristics of each tree, and the trees were divided into three classes based on crown projections. For each class, we took detailed measurements of one representative tree. All measured and counted growth characteristics were converted to the stand level. The tree population at the whole island level was investigated using remotely sensed data. The estimation of horizontal precipitation was performed using detailed and precise evaluation of the available climate data in half-hour intervals over the period of June 2009 and June 2010 and the ambulatory direct measurements (12.6.2009–26.6.2009). The mean annual horizontal precipitation below the crown of the dragon’s blood tree was estimated to be 792 mm in the highest (951–1545 m a.s.l.), 373 mm in the middle (601–950 m a.s.l.) and 46 mm in the lowest (180–600 m a.s.l.) altitudinal zones. Our model showed that the horizontal precipitation exceeded 40% of the total annual precipitation. The decline in the dragon’s blood tree forest could therefore cause the loss of additional water from the island’s hydrological cycle.
Article
Socotra, the largest of four islands in the Socotra archipelago (Yemen), is characterized by high geo-morphological and biological diversity. Traditional management of land resources has been impacted by cultural shifts, globalization and urban expansion. In addition, the island has been affected by climate impacts, such as droughts and recent cyclones causing significant economic and environmental damage. Our study presents an analysis of the dynamics of the vegetation on the island for the period 2001-2019 and highlights the status of different landscapes. We used Earth Observation techniques and the Normalized Difference Vegetation Index to assess land productivity dynamics basing on vegetation indexes derived from average resolution satellite images. The results show that although the arid landscapes of Socotra seem resilient, over 30% of the central mountainous areas of the Hageher (hosting a unique and high biodiversity) are decreasing in productivity , whereas in the lowlands (where plant diversity is relatively lower) productivity has increased significantly for the same time period, up to 40% in some areas. Given the difference in biological diversity of the sites, this study can help prioritize conservation areas and activities, which are crucial for Socotra's terrestrial environment. The methodology applied could be replicated for continuous monitoring in the future.
Article
We reviewed and compared the occurrence and biogeographic origins of three arthropod groups, darkling beetles (Coleoptera, Tenebrionidae), spiders (Araneae) and butterflies (Papilionoidea) on the Socotra archipelago (Yemen). More than half of the darkling beetles (80.3%) and spiders (51.5%) are endemic to the Socotra archipelago, in contrast with only 5 species (18.5%) of butterflies. The number of non-endemic taxa is higher among those distributed widely in the Afro-Oriental region, less of taxa from Afro-Arabian or African origin. These species colonised the Archipelago by natural means, but we document also introduction through human activity. The exact distribution and evolutionary patterns still need thorough research (e.g. molecular, phylogenetic, and phylogeographic analyses).
Article
We conducted a study throughout Socotra Island (Yemen) to investigate the distribution, population structure and regeneration potential of the endemic and threatened tree species Dendrosicyos socotranus, the Cucumber Tree. In total, 163 live trees (excluding seedlings) were found in all (24) investigated locations, covering an area of 96 km2. The height, stem circumference, fertility and coordinates of each tree were measured and data regarding the ecotope, land cover and natural regeneration were recorded. We estimated that there could be more than 6400 Cucumber Tree individuals among the populations on Socotra Island. The majority of the trees occur in the “high Shrubland with succulents” and “wadi” land cover classes. In general, the fertility of the trees is good, and natural regeneration occurred in 77% of the locations with adult trees; it was found that 16% of the individual trees regenerated. Our study, collecting data on the regeneration of the species and estimating the size of local populations for the first time, can be used for local conservation and future monitoring of this important, endangered Socotran endemic.
Article
Socotra Island, in the western Indian Ocean, harbors high biodiversity and endemism and makes up the largest part of the Socotra archipelago UNESCO World Heritage site. Its climatic, pedological, and geomorphological characteristics, together with the long geological isolation and inaccessibility, led to the flourishing of unique tree diversity, with great cultural and ecological value. Lately, trees on Socotra are facing new threats linked to the abandonment of traditional management practices, climate change impacts, and growing human pressure. This study assesses the relative influence of environmental factors and human activity as drivers of tree cover and density, which can be used to support land management policies for the conservation of this key local resource. Tree cover and density were assessed on a total of 3600 plots of 0.5 ha each, laid on a regular grid covering the entire area of the island through Collect Earth, a free open source software tool that allows an augmented visual interpretation of high-resolution satellite images. A total of 770,000 trees, with an average of 2.14 trees/ha, were estimated on the island. The collected data was validated by ground surveys in 26 plots. The relative influence of selected predictors (seven environmental, two anthropic) on tree cover and density was analyzed through logistic and survival regression. Tree cover and density were found to be impacted positively by steeper slopes, higher rainfall, and marked precipitation seasonality and negatively by relatively warmer temperatures and a higher sand content in the soil. Distance of each plot from the nearest road and settlement was calculated and neither of these anthropic predictors emerged as significant, indicating that the limited number of trees present are not currently as affected by anthropic pressure, but, instead, their presence is mostly limited by the harsh climatic and pedological factors. However, as human population and development increase, periods of drought rise, and extreme and random climatic events intensify, these additional stresses on the environment could negatively impact the availability of natural resources in the terrestrial environment. Lastly, potential recommendations to the existing conservation policies and Socotra Conservation Zoning Plan are discussed based on the results.
Article
The endemic fauna of achatinoid terrestrial molluscs for the Socotran Archipelago (Yemen) is revised. The main group falls into two genera, viz. Riebeckia E. v. Martens, 1883 with five species and Balfouria Crosse, 1884 with seven species, two of which are here described as new to science, i.e. Balfouria nitida n. sp., and Balfouria minima n. sp. Another globally distributed species could be found, and the endemic Zootecus species of the outer island of Abd el-Kuri is illustrated. The highly specialised shell structure of Balfouria is discussed, which is used to camouflage the animals with a mixture of clay and mucus protecting the shell during the individual lifetime of a specimen.