Dental Microwear in Relation to Diet in Blue Duiker and Common Duiker (Cephalophinae , Bovidae, Mammalia) in South Africa

8 Pages·2001·4.8 MB·English

by  Fred Kigozi

DENTAL MICROWEAR IN RELATION TO DIET
BLUE DUIKER AND COMMON DUIKER
IN
(Cephalophinae Bovidae, Mammalia) IN SOUTH AFRICA
,
FredKigozi' & Ric Bernard2
‘DepartmentofMammals,AmatholeMuseum
P.O. Box 1434, King William’s Town, 5600, SouthAfrica.
2DepartmentofZoology& Entomology, Rhodes University,
P.O.Box94, Grahamstown, 6140, SouthAfrica.
ABSTRACT
The blue duiker (Philantomba monticola Thunberg, 1789) and grey or common duiker (Sylvicapra grimmia
Linnaeus, 1758) representtwo ofthe three duikerspecies occurring in southern Africa. Whilst some workhas been
published on the relationship between dental microwear and diet in a number of antelope species, nothing is
documentedforduikers.
Scanningelectronmicroscopy(SEM)analysiswasusedtostudydentalmicrowearonthesecondlowermolarsof
theblueduikerandcommonduikerspecimenscollectedfromtheEasternCapeProvinceofSouthAfricaandhoused
intheShortridgeMammalCollectionattheAmatholeMuseum.
Although there were no significant differences in the dental microwear of the two duiker species, results
confirmedthatthetwoarebrowsing specieswithahigh incidenceofpitsontheirdental surfaces, anattribute dueto
thepresenceoffruitintheirdiet.
Keywords: Dental microwear, scanning electron microscopy (SEM), blue duiker, common duiker, Philantomba
monticola, Sylvicapragrimmia, Bovidae, Cephalophinae.
INTRODUCTION encountered when working with museum material.
&
However, Teaford Runestad (1992) stress the
Variations in dental microwear have yielded importance of using museum specimens collected
insight into a number of oral processes such as, from the same areaat the sametime unless the effects
occlusal relationships and biomechanics of the jaw of spatial and temporal variation are being
&
(Gordon, 1984a, 1984b; Wilkins Cunningham, investigated.
1993), anddietaryhabits (Smith 1984; Teaford, 1986; Several microwear features have been correlated
Taylor & Hannam, 1987). Probably the most with dietary variations (Covert & Kay, 1981; Teaford
important aspect of microwear analysis is the & Runestad, 1992; Lukacs & Pal, 1993). Gordon
possibility ofusing itto deduce the diet ofextinctand (1982) places these features into three categories,
&
fossil forms (Grine, 1981; Daegling Grine, 1987; striations or scratches, pits, and gouges. The
Waddle, 1988; Van Valkenburgh et al., 1990; Lubell distinction between pits and scratches can be made
etal., 1994). throughtheuseofacut-offpointintherangeoflength
Dental microwear analysis has focussed on a to width ratio (Teaford, 1985; Daegling & Grine,
number of herbivorous taxa such as Primates and 1994) or, by subjective determination (Grine, 1986).
&
Hyracoidea(Walkeretal., 1978;Teaford Robinson, Generally, scratches are linear depressions whose
&
1989; Teaford Runestad, 1992), while there have length is always greaterthan breadth (Gordon, 1982).
been a few studies on large antelopes such as Lengths andbreadths ofpits are approximatelyequal,
waterbuck and kudu (Solounias & Hayek, 1993). No and gouges are usually broader, strongly curved, and
such study on duikers or any other species of small oftenS-shaped(Gordon, 1982).
antelopesisdocumented. Microwear features do not necessarily reflect
Dentalmicrowearanalysisisfacilitatedbytheuse specific food items, but rather the mechanical
ofcasts and scanning electron microscopy (Murphy, properties ofthe items orthe constituents ofthe items
1982; Roomans, 1984). Analyses range from (Grine, 1986; Teaford & Robinson, 1989). Therefore,
qualitative to quantitative, and from experimental foods with similar mechanical properties might be
studies using live animals to comparative studies of expected to produce similar microwear patterns
museumcollections(Teaford, 1988). Teaford& Oyen (Daegling & Grine, 1994). Microwear patterns have
(1989) state that the process of taking dental been used to differentiate browsers from grazers
impressions from live animals is a difficult one and (Walkeretal., 1978; Teaford, 1985; Mainland, 1998),
that it presents problems different from those and frugivores from folivores (Teaford & Walker,
Annals ofthe Eastern Cape Museums Vol.2:1-8 2002 1
&
1984;Teaford Runestad, 1992).
Thedietoftheblueduiker variesthroughoutits
&
range (Faurrie Perrin, 1993) and comprises mainly
fallen leaves and fruits (Bowland, 1990; Hanekon &
Wilson, 1991). Whereas, Dubost (1984) found a high
occurrence of fruit in its diet and described them as
frugivorous, Bowland(1990) consideredblue duiker
to be folivorous on account oftheir diet comprising
mainly leaves. In a recent dietary classification of
African bovidae, Gagnon & Chew (2000) state that
with the exception of common duiker, all duiker
speciesarefrugivorous.
The diet of common duiker consists mainly of
forageofvariousdicots, twigs, flowersand somefruit
&
(Wilson Clarke, 1962; Boomker, 1983; Allen-
&
Rowlandson, 1986,Skinner Smithers, 1990).
The aim ofthis study was to determine ifdental
microwear is a sufficiently refined tool to detect the
small differences in the diets ofthe blue duiker and Figure 2. The lowerjaws ofblue duiker(foreground)
common duiker. In addition, patterns of dental andcommon duiker(background) showing the
microwearcouldbecomeavaluabletaxonomictool in positioning ofthe second lowermolar.
identification ofskulls, jawbones and loose teeth of
thetwospecies.Accurateidentificationofspeciesisof mammalscollection(Appendix 1). Thissamplesizeis
fundamental importance in ecological monitoring, comparableto thetenusedbyTeaford(1985), andthe
assessment, impact and conservation work as it often eightofGordon(1984d).
underpins the data from which subsequent analyses The second lower molars (Skinner & Smithers,
andinterpretationsaremade. 1990)wereselectedforSEManalysisbecausetheyare
placedbetweentwoothermolars(Fig.2)andtherefore
MATERIALS AND METHODS the occlusal function is the same on both sides
(Rensberger, 1973). They are also smaller in width
SAMPLINGOFSPECIMENS thantheuppermolars andtheyoccludeovertheentire
surface whereas the upper ones have only part ofthe
Eleven skullsfromeachofthetwo duikerspecies,
crownoccludedbyallthe loweronesas aresultofthe
overlap(Butler, 1978).
PREPARATIONOFDENTALREPLICAS
Withtheuseofcottonwooltheteethsurfaceswere
cleaned with water, then ethanol and finally acetone
(Rose, 1983). This freesthe surface ofany dirt, glue,
loose matrix or grease (Rose, 1983). Vigorous
scrubbing which may create artefactual scratcheswas
&
avoided (Teaford Oyen, 1989). After allowing the
surfacetoairdry(Rose, 1983), athincoatingofacetic
acid was applied to it, and a wall ofBostik Prestik
sticky stuff(Genkem Ltd, England) was built around
the tooth. Latex was then poured on top of the acid
layer. The acetic acid facilitates rapid and proper
setting of the impression material and together with
Figure 1. The skulls ofblue duiker(left) andcommon thebarrierprevenStEseMepageofthelatex(Ryan, 1979).
Preliminary analysis indicatedthat latex did
duiker(right).
not pick up the dental impressions, and subsequently
its use was abandoned. Molds ofthe teeth were then
collected from two neighbouring districts in the made with a mixture ofan equal amount ofAquasil
Eastern Cape Province, and with the closest possible smart wetting impression material (S.W.I.M) “base”
dates of collection, were used for the study. The and Aquasil S.W.I.M “ hardener catalyst” (Dentsply/
specimenswereobtainedfromtheAmatholeMuseum Caulk,MilfordDE,USA). AsrecommendedbyRose
2 Kigozi & Bernard: Blue & Common Duiker Dental Microwear
(1983)theimpressionmaterialwasappliedtotheteeth alwayspossibletorecordeveryfeature.
surfaces with disposable plastic syringes. Once the The number of microwear features per field of
mouldswerepeeledofftheteeththeywereallowedto examination was compared between samples using
sit in a dust free environment atroom temperature for the Mann-Whitney statistic or t- test. Chi-square
about eight hours, in order to permit total degassing analysis was used to test for interspecific differences
(Grine 1986). Thisisnecessaryinordertopreventany intheproportionsofpitsandscratches(Zar, 1996).
artefactual pitting (Gordon 1984 c). Initially epoxy-
resincastsweremadefromthemouldsaswasthecase RESULTS
in several studies (Gordon, 1984 d; Bullington, 1988;
Teaford & Runestad, 1992). However, this was Threetypesof features;scratches,pitsandgouges
discontinued, as a result of failure of the casts to were identified as occurring on all teeth of the blue
separate cleanly from the moulds, after setting. Rose duikerand common duiker. Representative images of
(1983) states that clean separation is an important casts from the teeth ofthe blue duiker and common
requirement for suitability of any casting material, duikerareshowninthemicrographs(Figs.3AandB).
whichifnotmetmayresultintoformationofartefacts.
ThisledtoacomparativeexaminationofSEM images
ofteeth and images oftheirAquasil S.W.I.M moulds.
Basedon thesimilarityoftheseimagesitwasdecided
to use the original Aquasil moulds which once made
and allowed time to degass were sputter-coated with
SEM
gold priorto analysis(Echlin, 1978).
SCANNING ELECTRON MICROSCOPY (SEM)
ANALYSIS
Approximatelytwohoursaftercoatingthemoulds
they were carefully orientated in a particular angle,
(long axes ofmoulds uniformly placed) and mounted
on marked stubs. Care was taken to avoid any direct
contactwiththemouldsurfaces(Rose, 1983).
They were examined in a Jeol JSM 840 scanning
electron microscope. Teeth were rotated in various
positions to have an overview of features, and
comparative micrographs were taken (Crompton &
Kielan-Jaworowska, 1978). Sets of micrographs at
magnificationsof130xand450x weretaken,butonly
thelatterwereusedforanalyses.
DATA COLLECTION AND ANALYSES
Data were collected from eleven micrographs for
each ofthe two duiker species. These were a total of
twenty-two micrographs taken from similar occlusal
facets ofthe second lower molars. With the use ofa
B41420/3 illuminated magnifier all identifiable pits
and scratchesinanareaof16 cm2atthecentreofeach
micrograph (of 450 X magnification) were counted
and recorded. Gouges were subsumed as scratches
(Grine, 1986). Pits and scratches were identified
independently by subjective determination following
themethodofGordon(1982),ratherthanbyimposing B
an arbitrarily set length to width ratio on the features.
In order to ensure that features are correctly Figure 3. Scanning electron micrographs taken fromthe
categorised, only those which could clearly be occlusal surfaces ofthe second lowermolars ofcommon
identified were recorded. It has been stated (Gordon, duiker(A) and blue duiker(B). The horizontal arrow
1982; Teaford and Walker, 1984) that, because ofthe indicates a scratch and the vertical arrow a pit.
overlapandlargenumbersoffeaturesperfielditisnot Magnification is450X.
Annals ofthe Eastern Cape Museums Vol.2:1-8 2002 3
1 1
Table 1. Numbers, ratios, percentages andresults ofstatistical comparisons ofmicrowearfeatures onteeth ofblue and
common duiker.
Dentalmicrowearfeatures Blueduiker(n=11) Commonduiker(n=11) Statisticalresults
Meannumberofscratches 79(15%) 84(14.9%)
per 16cm2
Meannumberofpits 449(85%) 481(85.1%)
per 16cm2
RatioofPits: Scratches 5.7: 5.5: InsignificantInterspecificdiff.
=
(Chi-Square;p 0.96)
The total number and percentages of pits and whichischaracteristicof browsers.
&
scratches recordedon the teeth ofthe blue duikerand Covert Kay, (1981) and Peters (1982) attribute
commonduikerweresimilarandforbothspeciesthere the differential wear on teeth of grazers (many
were more pits than scratches on the dental surfaces scratches and few pits) to the opaline phytoliths in
(Table 1). grasses. Bakeretal., (1959)andKay&Covert,(1983)
The chi-square analysis showed no significant demonstrated that wear caused by the opaline
interspecific differences intheproportions ofpits and phytoliths and that of gritty diets are essentially
scratchesbetweenduikerteeth(p=0.96). similar. Blueduikereatfallenleaves, fruit, andflower
Interspecific comparison ofthe mean number of (Skinner & Smithers, 1990; Faurie & Perrin, 1993;
features per field indicated no significant differences Bowland & Perrin, 1998) which are ingested along
(p = 0.53, Mann-Whitney). Intriguingly however, with accompanying debris. As a result, it might be
significant intraspecific differences were shown to expectedthat blue duikerdental wearwouldbe closer
exist between the number of pits and scratches per tothatofgrazers,buttheresultsfromthepresentstudy
field in both the blue duiker (P < 0.0001, t-test) and suggest that this is not the case. It is possible that
comTmhoenldouwieksetrr(ePco<r0d.e0d0n0u1m,tb-etersto)f.featuresina 16cm uTteialfiosartdio&noWfaflrukietrb,y(b1o9t8h4)duiaknedrTsepaecfioersde&xplRauinnsestthiasd.
squarefield(fromamicrographof450X)oncommon (1992) relate ahigh incidence ofpits to a frugivorous
duikerteethwas 21 comparedwith 9 ina similarfield diet. It is therefore likely that the dental effects of
onblue duikerteeth. Thehighestnumberrecorded, in eating fruits in both duiker species overshadows any
the same size of fields for common duiker and blue minorfeedinghabitdifferences.
duikerwere 94and97respectively. The significant intraspecific differences (between
similar sites on the molars ofthe same species) in the
DISCUSSION number offeatures per field is interesting. Although
everyeffortwasmadetoensurethatspecimensforthe
It is established that the diets of grazers and study were collected from the same area at the same
browsers result in different patterns of molar time,thiswasnotentirelypossible. Allspecimenshad
microwear (Walker et al., 1978; Teaford, 1985; beencollected fromPeddie andKing William’s Town
Solounias&Hayek, 1993;Mainland, 1998).Browsers districts, which are neighbouringjurisdictional areas
are characterised by many pits and few scratches, oftheEasternCapeprovince.Atthetimeofcollecting
while grazers have many scratches and few pits the specimens, both these districts were described as
(Solounias&Hayek, 1993). Afrugivorousdietresults having similar vegetation types, which was a
in teethwith ahighdensityofpits (Teaford&Walker, combination ofvalley bushveld and Eastern province
1984; Teaford& Runestad, 1992)andhardfruiteaters thomveld (Comins, 1962; Acocks, 1975). All
have wider pits than soft fruit eaters (Teaford, 1985; specimens were collectedbetween 25 May 1948 and
Teaford& Runestad, 1992). Blueduikerandcommon 5 April 1949. However, most specimens were
duiker have been described as browsers by Boomker collected during autumn and winter (Appendix 1).
(1981),Wilson(1966),Bowland(1990)and Bowland Despite the use ofonly adult specimens (ofunknown
& Perrin (1998) and this is supported by the type of ages)inthestudy, agewasnotfullycontrolledfor,and
microwear exhibited on their dental surfaces. The therefore may also have been a variable determining
molars of both species are predominantly pitted the results. Although the factor of gender is not
4 Kigozi & Bernard: Blue & Common Duiker Dental Microwear
mentioned in the literature, in this study only three counts. Microwear feature densities and relative
specimens were of unknown sex, and males and abundance have widely been used to detect dietary
females were spread evenly throughout all categories differences among closely related species (Teaford &
&
(speciesandlocalities)(Appendix 1). Walker, 1984; Teaford Runestad, 1992; Solounias
Gordon (1982) postulates that the recognised & Hayek, 1993) however, feature dimensions are also
types of microscopic abrasion features are not equally important (Gordon, 1982). Robson & Young,
intrinsically different, but rather manifestations of (1990) state that microwearfeature dimensions rather
different degrees ofshear and compression subjected thanfeaturedensitiesandrelativeabundances,maybe
totheagentswhichproducemicrowear. Accordingto the most suited for investigating diet differences of
thisview,pitsandscratchesarefoundatoppositeends closely related species. On the other hand, Teaford &
ofa continuum ofsurface wear phenomena, such that Runestad (1992) describe scratch widths as poor
the decision about where to make the division is indicatorsofdietarydifferences.
always arbitrary (Gordon 1988). Different cut-off Inconclusion, dentalmicrowearcannotbe usedto
points have been used. Daegling & Grine (1994) separate the blue duiker and common duiker.
definedpitsasthose featureswith alength-widthratio However, the abundance of pits on the occlusal
&
of4:1 or below, while Teaford Walker (1984) and surfaces supports the observation that both are
Teaford (1985) assigned to pits a ratio of 10:1 and browsers,withfruitaspartoftheirdiets.
anything above to scratches. Subsequent assessment
of the features indicated that, those features ACKNOWLEDGEMENTS
recognised as pits in this study possessed length to
width ratios ofabout 4:1 and below. In an analysis of Financial support from the National Research
feature dimension ratios, Solounias & Hayek (1993) Foundation (NRF) is very much appreciated. Thanks
concluded that the best diagnostic method of tooth to Mr Robin Cross the director of the electron
microwearanalysis utilizesthenumberofpits smaller microscope unit of Rhodes University for the
than orequal to the ratio four(length overwidth), the technical assistance. All the collections managers and
numberofscratchesbetween fourandonehundred in curators of the Natural History Museums who
length to width ratio, and that ofgouges greater than responded to our enquiry about duiker specimens in
onehundred inlengthtowidthratio. theirrespectivecollectionsarealsothanked.
The data for this study were from microwear
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Annals ofthe Eastern Cape Museums Vol.2:1-8 2002 7
APPENDIX
1
Information accompanyingthe blueandcommon duikermuseumspecimenswhosemolarswereusedinthe dental microwear
analyses.
Specimen CatalogueNumber Sex Districtoforigin Datecollected
KM M
Blueduiker 15056 KingWilliam’sTown 25April 1948
Blueduiker KM 15060 ? KingWilliam’sTown 25May 1948
KM
Blueduiker 15082 F KingWilliam’sTown 30July 1948
KM M
Blueduiker 15037 KingWilliam’sTown 27June 1949
Blueduiker KM 15055 M KingWilliam’sTown 06May 1948
Blueduiker KM 15558 ? KingWilliam’sTown 08May 1949
KM
Blueduiker 15559 ? KingWilliam’sTown 27June 1949
KM M
Blueduiker 15052 Peddie 04July 1948
KM M
Blueduiker 15053 Peddie 01 July 1948
KM M
Blueduiker 15062 Peddie 04July 1948
KM M
Blueduiker 15081 Peddie 19July 1948
Commonduiker KM 15098 F KingWilliam’sTown 02March 1949
Commonduiker KM 15099 F KingWilliam’sTown 05April 1949
Commonduiker KM 15100 F KingWilliam’sTown 07April 1949
Commonduiker KM 15101 F KingWilliam’sTown 17January 1949
Commonduiker KM 15102 M KingWilliam’sTown 17January 1949
Commonduiker KM 15103 M KingWilliam’sTown 09July 1948
Commonduiker KM 15129 M KingWilliam’sTown 16May 1948
Commonduiker KM 15138 M Peddie 28August 1948
Commonduiker KM 15162 M Peddie 02August 1948
Commonduiker KM 15163 M Peddie 08August 1948
Commonduiker KM 15164 F Peddie 08August 1948
KEY
KM
AmatholeMuseum(FormerlyKaffrarianmuseum)
? Notgiven
F Female
M
Male
KingWilliam’sTown(3327CD)andPeddie(3327AA)districtsareintheEasternCapeProvinceofSouthAfrica.
<5 Kigozi & Bernard: Blue & Common Duiker Dental Microwear