Observations of the
seasonal dietary preference of male Gazella subgutturosa marica Thomas,
1897 (Cetartiodactyla: Bovidae) along foraging trails of central Saudi Arabia
Peter Low Cunningham 1,2
1King Khalid Wildlife Research Center, Thumamah, P.O. Box 61681,
Riyadh, Kingdom of Saudi Arabia
2Zoological Society of London, Conservation Programmes, Regent’s
Park, London, NW1 4RY, United Kingdom
Email: 1,2 pckkwrc@yahoo.co.uk
Date of publication (online): 26
September 2009
Date of publication (print): 26
September 2009
ISSN 0974-7907 (online) |
0974-7893 (print)
Editor: C. Srinivasulu
Manuscript details:
Ms # o2154
Received 04 March 2009
Final received 24 May 2009
Finally accepted 29 August 2009
Citation: Cunningham,
P.L. (2009). Observations of the seasonal dietary preference of male Gazella
subgutturosa marica Thomas, 1897 (Cetartiodactyla: Bovidae) along foraging
trails of central Saudi Arabia. Journal of Threatened Taxa 1(9):
445-449.
Copyright: © Peter Low
Cunningham 2009. Creative Commons Attribution 3.0 Unported License. JoTT allows
unrestricted use of this article in any medium for non-profit purposes,
reproduction and distribution by providing adequate credit to the authors and
the source of publication.
Author Details: Peter Cunningham is an ecologist for the Zoological
Society of London (ZSL) at the King Khalid Wildlife Research Centre at
Thumamah, north of Riyadh in Saudi Arabia, studying the feeding ecology of Sand
Gazelle in protected areas as well as assisting with the management of the
species.
Acknowledgements: My gratitude
extends to H.H. Prince Bandar bin Saud bin Mohammed Al Saud, Secretary General,
NCWCD for his continued support towards conservation efforts in Saudi Arabia
and Ernest Robinson (Director KKWRC) for his assistance in identifying many of
the plant species and for commenting on a draft of this paper.
Abstract: Seasonal
dietary preferences of male Arabian Sand Gazelle were determined using direct
observations of vegetation utilised along foraging trails in central Saudi
Arabia. Twenty-one different plant
species from 12 different families were identified as food items, of which six
species have not previously been recorded in the diet from Saudi Arabia. The importance of forbs in the diet was
confirmed with species selected throughout the year with the highest relative
frequency of use including Farsetia stylosa, Convolvulus lanatusand Neurada procumbens. Knowledge
of the diet and food preference is important for protected area managers.
Keywords:Diet, foraging trails, Gazella subgutturosa marica, reem, Saudi Arabia
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Introduction
Herbivores in arid environments are
frequently faced with problems of variable food quality and quantity which can
be aggravated by overgrazing by livestock and wildlife (El Mouden et al.
2006). In general herbivores select a
broader diet than would seem optimal and eat certain food types on some
encounters but not on others (Owen-Smith 2008). It is thus crucial for species conservation to make sense of this and
possess adequate data on feeding ecology. Little is known about the feeding ecology of Gazella subgutturosa
marica Thomas, 1897 (reem/rheem/rim in Arabic) from
Saudi Arabia (Habibi 1991) although some scientific work has been conducted on
aspects such as genetics (Granjon et al. 1991; Vassart et al. 1993),
ecophysiology (Williamson et al. 1991), population dynamics in captivity
(Rietkerk et al. 1991) and daily activity and social organization
(Launay & Launay 1992).
G. s. marica are
predominantly viewed as grazers, feeding on ephemeral plants after rains but generally
relying on dry grasses, chenopods and desert melons (Thouless et al.
1991). Dwarf shrubs are important browse
(Harrison & Bates 1991), with Wacher (1995) indicating their use of bulbs,
forbs, grasses and occasionally trees - i.e. browse – in the Uruq bani M’arid
Protected Area in southern Saudi Arabia. Haque (n.d.) confirms G. s. marica being grazers as observed in
the Mahazat as-Sayd Protected Area in central western Saudi Arabia, with
grazing preference shown towards Panicum and Stipagrostis grasses. Mowlavi (1978) noted that browsing formed the
largest part of the diet (86%) – increasing during the drier months - in Iran,
while Mohamed & Saleh (1991) state the importance of forbs in their diet
from Bahrain.
Gazella subgutturosa Guldenstaedt 1780
ranges from western China through Mongolia, Afghanistan, Iran and south-eastern
Turkey including the Arabian Peninsula (Harrison & Bates 1991), with the
subspecies G. s. marica confined to the Arabian Peninsula (Thouless et
al. 1991) and the type locality documented as Nejd, central Arabia.
Gazella subgutturosa is classified
as Vulnerable (A2ad: population size reduction over 30% over the last 10 years
or three generations; a – direct observation; d – actual or potential levels of
exploitation) by the IUCN (2008) with 35% of the global population share
associated with Saudi Arabia (Dunham et al. 2001) and “endangered” by Flamand
(1995). The subspecies marica is
categorized as Vulnerable as the global population of Arabian Sand Gazelle is
estimated to be less than 10,000 mature individuals with no subpopulation
containing more than 1,000 mature individuals, and the largest subpopulation in
Oman undergoing a continuing decline (Antelope Specialist Group 2008).
Although this study was limited to a
single sex with a small sample size enclosed in a relatively small area without
the entire range of potentially available vegetation (e.g. trees and larger
shrubs), the aim of this paper is to quantify for the first time the diet of G.
s. marica from central Saudi Arabia using back tracking as a technique.
Materials and Methods
Two adult male G. s. marica kept
under “natural” conditions in an enclosure of approximately 400ha in size were
followed for one or two mornings each month from January 2008 to January 2009. The distance followed depended on the
visibility of the tracks being followed, with a minimum of 5m used arbitrarily
and a mean of 458 ± 38.2 m per month. Additional food – alfalfa – was supplied on a daily basis and the
gazelle had free access to water. The
gazelle were released into the pen in 2004 as part of a publicity/public
relations exercise.
The technique used can be described as
“back tracking”. The area was
crisscrossed until a fresh clearly visible foraging trail was intercepted. This foraging trail was then back tracked and
all green plants within a 1m width along the foraging trail (i.e. transect)
were recorded and inspected for signs of feeding. This included clear bite
damage to a plant, plant remains observed at a damaged plant and changes in the
rhythm of the tracks followed indicating foraging (e.g. tracks closer together
or local milling together with plant damage as clear evidence of foraging).
All the green – i.e. potentially
palatable – plants along the transect were recorded and plants utilised – i.e.
showing bite damage – were identified in situ or samples taken and
identified with the use of the available literature (Collenette 1985) and the
assistance of the King Khalid Wildlife Research Centre (KKWRC) resident
botanist, Dr E.R. Robinson.
Absolute frequency of use was determined
as the percentage of the food plant actually utilised in relation to the total
amount of that specific plant. Relative
frequency of use was determined as the percentage of the food plant actually
utilised in relation to the total number of food plants potentially
available. This method of analysis is
based on that of Cavallini & Serafini (1995).
A shortcoming of this study is that only
two male gazelle were kept in the enclosure consequently limiting the
statistical analysis and potentially indicating a bias in information only for
males. However, as additional food
(alfalfa) and water were supplied daily, the dietary selection could be viewed
as true preference – i.e. food plant selection was not out of necessity as
gazelle had access to alfalfa, but rather a true selection for plant species
and plant parts.
Study
area
The KKWRC is located approximately 70km
north of Riyadh in central eastern Saudi Arabia in an area known as Thumamah
(Figure 1). The limestone Tuwaiq Escarpment borders the area towards the east
with sandy-gravel plains and a belt of sand dunes towards the west (25013’N
& 46037’E elevation 600m) (Child & Grainger 1990). The overall size of the general Thumamah
protected area is 350km² (fenced) with the KKWRC fenced enclosure within this
being approximately 20km². The fenced
enclosure in which the feeding observations were made is approximately 400ha in
size. The pen originally housed a
variety of wild ungulates with the last internal pens having been dismantled by
2007 and the vegetation having recovered naturally since the original occupants
were removed. Thumamah falls within the
phytogeographical region known as the Eastern Najd (Migahid 1978) or the
floristic region known as the Saharo-Sindian (Guest 1966) or Sudanian
(Mandaville 1984) and characterised by dwarf shrubland. The rainfall is highly variable and the
average for several years is 107.8mm per annum (Robinson 2008a). The vegetation in the study area is
representative of the general area although having been severely degraded
(overgrazed) due to the high densities of ungulates previously kept in the
area, but since then having recovered with species previously locally scarce
(although not viewed as rare) due to overgrazing having returned (E.R. Robinson pers. comm.). No other ungulates shared the study area
although other mammals in the study area include a variety of small rodents,
Ethiopian hedgehog (Paraechinus aethiopicus) and red fox (Vulpes
vulpes).
Results
A total of 3,914 green (i.e. potential
palatable) plants (spring-360, summer-1,161, autumn-247, winter-2,146) were
encountered along the various foraging trails which extended over a total
distance of 5,500m. These consisted of
32 plant species (6 grasses and 26 forbs) of which 21 species were utilised to
some extent and 11 species (34.4% - total; 2 grasses [33.3%] & 9 forbs
[34.6%]) avoided (not observed being utilized) throughout the year. The Gramineae utilised comprised four species
and the forbs utilised comprised 11 families with Compositae having the most –
i.e. three species – followed by Boraginaceae, Chenopodiaceae, Convolvulaceae
and Cruciferae with two species each. All other families had one species each (Table 1).
During winter (92.3%) and summer (67.4%)
forbs dominated the vegetation encountered along the foraging trails (Fig.
2). Forbs were more frequently utilised
than grasses throughout the seasons (35.6% versus 15.4%).
Absolute frequency of use indicates the
importance of the grass Stipagrostis plumosa, widespread in the area,
throughout the year (Table 1). Furthermore grasses were utilised more often throughout the year except
during summer and winter when the forbs Neurada procumbens (46.7%) and Plantago
boissieri (67.7%) were utilised more often. The only grass to exceed a 50% relative frequency of use throughout the
year was during spring when Panicum turgidium reached 65.5% ranking it
the seventh most important food species. Panicum turgidium consistently had the highest relative frequency
of use of the grasses (Table 1).
According to the relative frequency of
use the importance of forbs is evident. Farsetia
stylosa, Convolvulus lanatus and Neurada procumbens (ranked 1
- 3) were selected throughout the year and were consistently utilised above 60%
(Table 1). Neurada procumbens was
also a dominant plant (high absolute frequency of use) associated with summer
rainfall. Other seasonally important
forbs with high relative frequency of use were Centuarea pseudosinaica(spring - 80%), Convolvulus oxyphyllus (summer - 68.2%), Kochia
indica (spring - 84.6% & summer - 55.5%) and Heliotropium
rammosisima (winter - 41.9%). Another forb utilized throughout the year albeit never at more than 50%
relative frequency of use was Polycarpea repens.
Discussion and Conclusion
According to Owen-Smith & Novellie
(1982) foraging performance is more sensitive to changes in food quality than
quantity, and a “clever ungulate” is defined as a short-term optimizer for
performance alone. This study using foraging
trails revealed that G. s. marica fed on a wide variety of plants (21
species in 12 families) of which six species (Cakile arabica,Centuarea pseudo sinaica, Kochia indica, Launaea cassiniana, Plantago
boissieri & Rumex vesicarius) have not previously been recorded
in the diet from Saudi Arabia. The
importance of forbs in the diet from central Saudi Arabia supports the findings
of Mohamed & Saleh (1991) from Bahrain. Important food plants (high relative frequency use) such as Farsetia
stylosa and Convolvulus lanatus are ubiquitous although not dominant
in the area. The nutrient content of the
various plant species selected by G. s. marica during this study was not
analysed although Farsetia sp. and Convolvulus lanatus are
generally viewed as valuable fodder plants (Collenette 1985; Al-Zoghet
1989). It is interesting to note that
Fabaceae were not observed to be utilised although this is known to be an
important family and often utilised by herbivores due to their high nutritional
value (El Mouden et al. 2006). This is
probably due to the absence of Acacia species in the enclosure as well
as the typical legumes in the area being small and prostate forbs making it
difficult to notice if utilisation has occurred.
Rainfall for 2008 was low with only 29.7mm
received during spring resulting in only 76 species of plants recorded for May
2008 although the average is 84.5 plant species thus indicating that the number
of plant species is directly influenced by rainfall (Robinson 2008b). Vegetative response to rainfall, result in
changes in plant selection as indicated by the increase in the use of Neurada
procumbens during summer and autumn after the late spring rains and Plantago
boissieri as a result of early winter rains. The diet and selection of plant and plant
parts is thus affected by the intensity and seasonal distribution of rainfall
indicating the importance to wildlife managers of monitoring rainfall events,
intensity, distribution and related vegetative response.
Although G. s. marica feed on a wide
variety of plants these species are also preferred by domestic stock indicating
that where gazelle and domestic stock (e.g. goats & camels) occur together
in unfenced protected areas there is likely to be intense competition. This is
especially important for Protected Areas in Saudi Arabia with a mainly gravel
substrate as Gallacher & Hill (2006) indicate that intense grazing by
camels (which are viewed as major competitors of wild ungulates including
gazelle in Saudi Arabia (Robertson 1993)) affect the plant ecology most on such
substrate with heavy grazing furthermore reducing species richness and
diversity and could affect the recruitment of perennial species (Gallacher
& Hill 2008). In future it would be useful
to expand this study to include both sexes as well as different age classes of
gazelle. Knowledge of the diet and plant
preference of G. s. marica is useful for protected area managers and can
assist with future envisaged reintroductions and be used as supportive argument
to reduce the domestic stock grazing pressure in protected areas in Saudi
Arabia.
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