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17th Annual Sahelo-Saharan Interest Group Meeting
2 days of talks on biodiversity conservation in the Sahara and in the Sahel
Desert refugia under a changing climate : spatial ad temporal patters
of botanical diversity in a hyper-arid mountain system
Peter COALS et al., Researcher – WildCRU, University of Oxford
May 4 – 6, 2017
Desert	refugia under	a	
changing	climate
Spatial	and	temporal	patterns	of	botanical	
diversity	in	a	hyper-arid	mountain	system	
Peter	Coals	1*
Avi Shmida 2,	Amiel Vasl 3,	Nasr	Mansour	
Duguny 4 &	Francis	Gilbert	1
1.		School	of	Life	Sciences,	University	of	Nottingham,	Nottingham	NG7	2RD,	UK
2.		Dept.	of	Ecology,	Evolution	&	Behavior,	Institute	of	Life	Sciences,	The	Hebrew	University	of	
Jerusalem,	Jerusalem,	Israel.
3.	Leon	Blaustein Ecology	Lab,	University	of	Haifa,	Haifa	31905,	Israel.
4.	Abu	Seila,	St	Katherine,	South	Sinai,	Egypt.
Outline
• Background
• Global	climate	change
• Biodiversity	loss
• Finger-prints	of	climate	change;	range	contraction
• Knowledge	gaps	and	research	needs
• Research	question
• Study	site
• Data	collection
• Results
• Conclusions
• Conservation	implications	and	future	directions
Background
• Global	climate	change:
• Rapidly	recent	temperature	increases
• Predicted	to	continue	
Dufresne,	J.	L.,	Foujols,	M.	A.,	Denvil,	S.,	Caubel,	A.,	Marti,	O.,	Aumont,	O.,	...	&	Bony,	S.	2013.	Climate	change	projections	using	the	IPSL-CM5	Earth	System	Model:	from	
CMIP3	to	CMIP5.	Climate	Dynamics,	40(9-10),	2123-2165.
Time	evolution	of	the	
global	mean	surface	
air	temperature	
anomaly	(Dufresne	
et	al.	2013)
Climate	change	&	biodiversity	loss
Bellard, C., Bertelsmeier, C., Leadley, P., Thuiller, W. & Courchamp, F. 2012. Impacts of climate change on the future of biodiversity. Ecology Letters 15(4):
365-377.
• Climate	change	is	a	significant	driver	of	current	and	predicted	
biodiversity	loss	across	taxa.	
Projections	of	
biodiversity	
losses	driven	by	
climate	change	
(Bellard et	al.	
2012)
Finger-prints	of	climate	change
• Shifting	climatic	zones	reduce	suitable	habitat	area,	leading	to	
isolation	of	populations	and	subsequent	mountain-top	extinctions
• Range	restricted	species	are	especially	vulnerable	to	range	
contraction,	upslope	movement,	and	mountain-top	extinction.
Parmesan,	C.	2006.	Ecological	and	evolutionary	responses	to	recent	climate	change.	Annual	Review	of	Ecology,	Evolution	and	Systematics 37:	637-669.
Knowledge	gaps;	range	shifts
• Plants;	mostly	studied	in	temperate	regions,	little	from	subtropical	
and	arid	regions.
• Lower	altitudinal	range	limits	are	under	researched,	and	rarely	
considered	together	with	upper	limits.	
• We	present	the	first	such	study	from	the	Middle	East.	
Research	effort	
(publications,	N)	
on	climate-related	
range	shifts	since	
1850s	for	
terrestrial	
ecoregions	
(Lenoir	&	
Svenning 2015).
Lenoir, J. & Svenning, J.-C. 2015. Climate-related range shifts - a global multidimensional synthesis and new research directions. Ecography 38: 15-28.
Research	questions
• What	are	the	altitudinal	patterns	of	diversity	in	a	desert	
mountain	floral	refuge?
• Is	there	evidence	of	recent	altitudinal	range	shifts	in	a	
hyper-arid	Middle	Eastern	desert	mountain	flora?
• How	do	directions	of	shift	for	upper	and	lower	altitudinal	
range	limits	of	plants	vary?
Location
• St	Katherine	Protectorate	(SKP),	South	Sinai,	Egypt	
4350	km2 of	the	southern	
peninsula	of	Sinai
Altitude	to	2643m	(Egypt’s	
tallest	mountain)
SKP	Flora
• 19	of	Egypt’s	33	endemic	plant	species.
• One	of	the	most	important	centres	of	plant	diversity	in	the	Middle	
East	(IUCN,	1994).	
• Many	species	of	plants	exhibit	disjunct distributions	of	Holarctic	
species	found	more	commonly	further	north,	suggesting	that	these	
species	are	relics	of	a	more	humid,	colder	past.	
IUCN.	1994.	Centres	for	plant	diversity:	a	guide	and	strategy	for	their	conservation.	IUCN,	Cambridge,	UK.
http://flora.org.il/en/plants/CRASIN/
Crataegus x	
sinaincus
Refugial pattern	of	diversity
• Temperate	flora	remnant	species	remain	only	at	higher	altitudes.
• Hill	numbers	diversity	indices:
• (a)	0D:	Increasing	species	richness	with	altitude.
• (b)	1D:	Decreasing	typical	species	with	altitude.
• (c)	2D:	Increasing	common	species	with	altitude.	
• Communities	become	more	uneven	at	higher	altitudes	with	a	few	
species	showing	increasing	levels	of	dominance.	
• Thus	increased	species	richness	at	high	altitudes	consists	mainly	of	
rare	species.
Endemics
• Endemics	peak	in	density	at	high	altitudes
• (a)	Plantago	sinaica
• (b)	Polygala	sinaica	
• (c)	Silene	schimperiana
Data	collection
• Field	surveys:
• October	to	mid-December	2014	
• Surveys	were	carried	out	in	mountainous	areas	
predominantly	within	an	igneous	ring-dyke	area	over	an	
altitude	range	of	1324	m	to	2629	m.		
• Locations	of	transects	was	chosen	to	cover	all	major	
habitat	types.	
• Quadrats	of	area	100	m2 every	50	m	change	in	elevation.	
• 36	sites;	283	quadrats;	28300	m2.	
• Abundances of	all	species	recorded.
• Comparison	of	field	data	with	a	1970s	dataset	compiled	by	Arbel &	
Shmida (1979)	within	the	St	Katherine	ring-dyke
• Method	replicated	by	2014	surveys.
• Lower	resolution;	quadrats	placed	every	200m	change	in	altitude.	
• Raw	data	unavailable	from	1970s	only	upper	and	lower	altitudinal	
limits	of	species	occurrences	available.		
• Paired	species	2014-1970s:
• 81	species;	upper	altitudinal	limits	
• 25	species;	lower	altitudinal	limits	
• More	than	10	individuals	recorded	during	the	2014	field	surveys:
• 63	upper	limits		
• 22	lower	limits
Arbel O.	&	Shmida A	.1979.	The	vegetation	of	the	high	mountains	of	South	Sinai.	Society	for	the	Protection	of	Nature.	Tel	Aviv.	67	pp.	[in	Hebrew].
Results
• Is	there	evidence	of	recent	altitudinal	range	
shifts	in	a	hyper-arid	Middle	Eastern	desert	
mountain	flora?
• Predictions:
• Upper	altitudinal	range	limits	shift	upslope.
• Lower	altitudinal	range	limits	shift	upslope.
Results;	upper	limits
• Predicted	increase	in	mean	upper	altitudinal	limit	1970s-2014:
• Significant	increase in	mean	upper	altitudinal	limits (mean	2014:	
2228.6	± 294.5	m,		mean	1970	2125.2	± 350.2	m:	1-tailed	paired	t	=	
3.37,	df =	61,	p<0.001).
Species	limit	
shifts
• Significantly	greater	
numbers	of	species	upper	
range	limits	moved	
upslope	(resolution	over	
100	m):
• Movement	of	over	100	m	
(26/40,	binomial	test	
p=0.04).	
• Movement	of	over	250	m	
(16/18,	binomial	test	
p<0.001).
Results;	lower	limits
• Predicted	increase	in	mean	lower	altitudinal	limit	1970s	to	2014:
• Significantly	reduced mean	lower	altitudinal	limit	(current	
mean	1568.0	± 162.1	m,	past	mean	1668.2	± 166.6	m;	paired	t	
=	3.02,	df =	20,	p=0.0064).
Species	limit	
shifts
• Significantly	greater	
number	of	species	
decreased	their	individual	
lower	altitudinal	limits	
than	did	not	(17/22,	
binomial	test	p=0.008)	
• Movement	over	100	m	
(12/13,	binomial	test	
p=0.002).
Summary
• Mean	upper	limits	and	majority	of	species	showed	upslope	
movement.
• In	line	with	predictions.
• Mean	lower	limits	and	majority	of	species	showed	downslope	
movement.
• Contrary	to	predictions.
• Suggests	that	factors	in	addition	to	temperature	are	influencing	
lower	limit	shifts.	
• Scope	of	study	limited	by	lack	of	historical	data.	
• Check	it	isn’t	annual	variation…
Shrubs	&	Trees	
• Support	full	data	for	larger	movements:
• Higher	mean	upper	limits	(present	mean	2219.1	± 311.2	m,	past	
mean	2139.5	± 353.3	m:	paired	t	=	2.30,	df =	36,	p=0.027).
• Lower	limits	of	shrubs	and	trees	were	also	significantly	lower	in	the	
present	data	(1585.7	± 145.7	m)	than	in	the	1970s	(1725.0	± 171.8	
m:	paired	t	=	5.27,	df =	12,	p=0.0002).	
Upper range		limits Lower	range	limits
All	species No	majority	upslope	
(21/38,	p=0.31)
Majority	downslope
(14/16,	p=0.006)
Movement	>	100 m No	majority	upslope
(15/22,	p=0.07)
Majority	downslope	
(9/9,	p=0.006)
Movement	>	250 m Majority	upslope
(7/8,	p=0.04)
N.A.
Species’	paired	limit	movements
Species
Limit	movement	patterns
Range	size	change
Lower	limit Upper	limit
Alkanna	orientalis down stationary expanded
Astragalus	echinus down down no	change
Calipeltis	cucullaris stationary up expanded
Colchicum	guessfeldtianum up down contracted
Cotoneaster	orbicularis stationary up expanded
Crataegus	x	sinaica stationary stationary no	change
Globularia	arabica down up expanded
Nepeta	septemcrenata stationary down contracted
Origanum	syriacum down stationary expanded
Phlomis	aurea down up expanded
Polygala	sinaica down stationary expanded
Pterocephalus	sanctus stationary stationary no	change
Pulicaria	undulata stationary up expanded
Rubus	sanctus down stationary expanded
Salvia	multicaulis down down expanded
Scariola	orientalis down down expanded
Silene	leucophylla down up expanded
Silene	schimperiana stationary down contracted
Stipa	parviflora stationary up expanded
Thymus	decussatus down down expanded
Verbascum	decaisneanum stationary up expanded
Verbascum	sinaiticum down up expanded
Endemics:
Polygala	sinaica
gained	~200m	
lower	range	limit.
Silene schimperiana
lost	~100m	
altitudinal	range	at	
high	range	limit.
Conclusions
• Simple	predictions	of	upslope	shifts	are	and	
contracting	ranges	are	not	met.
• Wider	climate	change,	including	changes	in	
water	balance,	area	of	bare	soil	surface	and	
elevated	atmospheric	carbon	dioxide	levels	may	
influence	range	shifts	in	plants.	
• Reliable	climatic	and	environmental	information	
is	lacking	in	South	Sinai.		
• Further	research	of	drivers	of	range	shifts	in	arid	
lands	are	required.
Conservation	for	desert	mountain	
flora	under	changing	climate
• Many	species	show	expansions	of	their	altitudinal	ranges.
• Low	present	threat	posed	by	range	contraction.
• However,	the	Sinai	endemic	Silene schimperiana has	contracted	in	
altitudinal	range.	
• Risk	must	be	considered	on	a	case-by-case	basis	for	Sinai’s	endemic	
and	rare	species.	
• Further	similar	studies	required	across	other	desert	regions.			
• Whilst	we	cannot	conclusively	state	that	observed	point	to	
environmental	change	that	may	pose	ecological	and	conservation	
issues	for	the	future.	
• Highlight	the	necessity	of	increasing	the	comprehensiveness	and	
quality	of	the	region’s	environmental	monitoring	programmes.
• Only	through	the	collection	and	use	of	detailed	fine-scale	data	can	
the	underlying	causes	and	conservation	implications	of	observed	
range	shifts	be	determined.
Our	thanks	to	the	Egyptian	Environmental	Affairs	Agency	for	
permission	to	carry	out	the	2014	work,	and	we	are	very	grateful	to	Mr	
Mohamed	Kotb and	the	rangers	of	the	St	Katherine	Protected	Area	for	
their	support	for	our	work	in	this	and	other	projects.	We	are	hugely	
grateful	to	Ibrahim	ElGamal whose	botanical	and	terrain	expertise	
significantly	enhanced	the	quality	of	this	work.

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