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International Journal of Acarology

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Geometric morphometric analyses of the male and

female body shape of Hyalomma truncatum and H.
Marginatum rufipes (Acari: Ixodidae)
a b
E. Pretorius & F.C. Clarke
a
Department of Anatomy, Faculty of Medicine, University of Pretoria, Basic Medical Science
Building, P.O.Box 2034, Pretoria, 0001, South Africa E-mail:
b
Department of Biology, Faculty of Science, P. O. Box 139, MEDUNSA, 0204, South Africa
Published online: 17 Mar 2009.

To cite this article: E. Pretorius & F.C. Clarke (2000): Geometric morphometric analyses of the male and female body shape
of Hyalomma truncatum and H. Marginatum rufipes (Acari: Ixodidae), International Journal of Acarology, 26:3, 229-238

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 Vol. 26, No. 3 Intemat. J. Acarol. 229

GEOMETRIC MORPHOMETRIC ANALYSES OF THE MALE AND FEMALE BODY

SHAPE OF HYALOMMA TRUNCA TUM AND H. MARGINA TUM RUFIPES
(ACARI: IXODIDAE)

E. Pretorius I and F.C. C l a r k e 2

1. Department of Anatomy, Faculty of Medicine, University of Pretoria, Basic Medical Science Building, P.O.Box
2034, Pretoria, 0001, South Africa, e-mail: rpretori@medic.up.ac.za. 2. Department of Biology, Faculty of Science,
Downloaded by [University of Delaware] at 07:23 09 June 2013

P. O. Box 139, MEDUNSA, 0204, South Africa.

A B S T R A C T - A geometric morphometric analysis of the male and female body shape of two species
belonging to the Ixodidae, Hyalomma truncatum (Koch) and H. marginatum rufipes (Koch) was performed.
These two species are very similar in adult appearance, and only the well-trained researcher is able to
distinguish between them. This analysis compares the body shape of the ticks with each other by collecting
16 landmarks from the males and females and analyzing this with a powerful statistical technique, geometric
morphometrics. Relative warp analyses were performed on the data set and the results of the first two relative
warps (for each of the male and female data set) plotted against one another. The first two relative warps
are indicative of the most variation between structures that are studied. Thin-plate-spline analyses were also
performed on the consensus shape of the males and females. Results from thin-plate spline analysis visually
provide input regarding landmarks that are responsible for shape similarities and differences. Geometric
morphometrics is a powerful tool that can be used successfully to identify phenetic relationships between
taxa, and could be an additional tool to be used by tick systernatists.
K e y w o r d s - Acari, Ixodidae, Hyalomma, geometric morphometrics, landmarks, relative warp analysis,
thin-plate spline, consensus shape, South Africa.

INTRODUCTION drier areas of southem Africa (Howell et aL, 1978;

Descriptions of ticks that occur in South Africa ap-
peared in the iiterature for over 200 years (Walker, 1991).
Koch (1844) published a historic work on ticks from dif-
ferent parts of the world (including 11 ixodid ticks from
southern Africa) (Hoogstraal, 1956). His research laid part
of the foundation of modem tick systematics with estab-
lishment of 5 genera: Ixodes (Latrielle), Boophilus (Cur-
rice), Rhipicephalus (Koch), Hyalomma (Koch) and Am-
blyomma (Koch). Over the last half century, classification
of the Ixodidae on the generic level has remained remark-
ably stable (Klompen et al., 1997); only one genus,
Anomalohimalaya was added (Hoogstraal et al., 1970).
The genus Hyalomma (one of the smaller genera),
also known as the bont-legged ticks, presently consists of
21 species, of which two species, Hyalomma truncatum
(Koch) and H. rufipes (Koch) (2 subspecies) occur in the
Walker, 1991). Adult ticks are rather large and active,
with dark brown bodies and long, banded legs. They are
drought resistant and have the ability to withstand long
periods of starvation.
Adults of individual Hyalomma spp. often show a
great range of morphological variation. According to
Walker (1991), the failure to recognize this variation, as
well as the fact that the species described by Koch (1844)
remained largely unrecognized, resulted in nomenclato-
rial chaos. This remained so for many years, until ap-
proximately 50 years ago when Hoogstraal (1956) re-
viewed the genus.
Two of the Hyalomma spp., H. marginatum rufipes
and H. truncatum are very similar in adult appearance
(Hoogstraal, 1956). To the naked eye these two species
are similar and only the well-trained researcher is able to
distinguish them (Howell et al., 1978). Hyalomma mar-
 230
ginatum rufipes [syn. H. aegyptium var impressum, mar-
ginatum, H. rufipes (Walker, 1991)], is known as the
large, coarse bont-legged tick. The primary domestic host
of the adults of this tick is cattle (Walker, 1991), but they
also parasitise horses, sheep and goats (Howell et al.,
1978). Wild animals (e.g. rhino, zebra, giraffe, buffalo
and antelope) are also utilized as hosts (Theiler, 1962).
Hyalomma truncatum, known as the small smooth
bont-legged tick, is slightly smaller and narrower than H.
marginatum rufipes, and the scutum is smoother. Many
synonyms for this species exist, including 1-1. transiens, a
name often used in older South African literature (Theiler,
1962). Adult H. truncatum prefers the same hosts as H.
marginatum rufipes, but have also been found on some
medium size and smaller host species (Theiler, 1962; Nor-
val, 1982). The predilection sites of the adults of the two
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species also coincide as they are found mainly near the
anus, genitalia and hooves, and both usually complete a
two-host life cycle. They are more active in summer, but
may be active throughout the year in drier areas. During
unfavorable conditions they become inactive and en-
gorged nymphs may over-winter for as long as 5 months
before molting, while engorged females may survive up
to 8 months (Howell et al., 1978).
The distribution of these two tick species overlaps
partially. Hyalomma marginatum rufipes occurs in all de-
sert and semi-desert regions of southern Africa, at a low
relative humidity. It is present in the greater part of South
Africa and Botswana but absent from the coastal strip in
the east, northeastern part of the Free State Province and
Lesotho, the tropical parts of Namibia and the winter
rainfall areas. The distribution range ofH. truncatum cov-
ers the western and northern parts of South Africa, Bot-
swana and Namibia thus corresponding greatly with that
of H. marginatum rufipes. This species, however, also
occurs in the winter rainfall area of the western Cape
(Howell et al., 1978).
G e o m e t r i c m o r p h o m e t r i c analysis - The main mo-
tivation for the research was the exceptional similarity in
adult body form and also because these two tick species
are so similar in life cycle, predilection sites and distribu-
tion. Although multivariate morphometrics have been
done on acarines before, the aim of this research was to
compare the body shape of males and females of the two
Hyalomma species by using geometric morphometrics.
This will also be the first shape analysis of members of
the Arachnida with this new, powerful statistical method.
Geometric morphometrics is a relatively new field dealing
with the study of shape using either homologous land-
marks (Rohlf and Marcus, 1993) or outlines of a morpho-
logical structure. Landmarks (employed in this study) are
the points at which biological structures are sampled and
these points capture shape changes between the same mor-
Pretorius & Clarke 2000
phological structures in different species. Kendall's defi-
nition of shape space forms the basis of geometric mor-
phometrics (Kendall, 1981, 1984) but this space is non-
linear and non-Euclidean, and therefore conventional lin-
ear multivariate statistical methods cannot be applied to
it. Shape space can, however, be approximated by a linear
space tangent to it. This approximated tangent space now
allows for the performance of standard multivariate statis-
tics on a data set of homologous landmarks (or x,y coor-
dinates) of x number of taxa.
The main purpose of the technique is to permit de-
scription of variability of a morphological structure using
a powerful, comprehensive statistical analysis, and the use
of thin-plate splines to describe the results in terms of
deformations. This produces an exact geometric descrip-
tion of the shape differences between the same structure
in different organisms. Authors like Bookstein (1989a,
1991 ), Rohlf and Slice (1990), Slice (1993), Rohlf (1995)
developed this technique and F. James Rohlf developed
the 'tps series' of programs which performs the statistics
and visualizations of geometric morphometrics. This dy-
namic series of programs is continuously upgraded and
improved by F. James Rohlf, and was used as a tool in the
analysis of the body form of the males and females of H.
truncatum and H. marginatum rufipes.
The ticks used in the present study were identified
by Mr. Arthur Spickett, Division of Parasitology, Agricul-
tural Research Council, Onderslepooit, South Africa. The
voucher specimens are kept in the Onderslepoit Tick Col-
lection. Specimens were obtained from the laboratory
maintained colonies at MEDUNSA (H. truncatum) and
South African Bureau of Standards.
MATERIALS AND METHODS
Determining the l a n d m a r k s - Homologous land-
marks were chosen and their x,y coordinates recorded by
the computer program 'Comp3d' that was linked to a
reflex microscope. The lowest point of each specimen
was taken as the zero point and landmarks data collected
only in the positive area of the x,y axis, This represented
points on half of the bilaterally symmetrical bodies of the
ticks (Table 1). Sixteen landmarks, excluding the zero
point were collected from the dorsal view of the body of
10 males and 10 females of each species. The x,y land-
mark data was then written in tps format, the male and
female coordinates of the two species combined, and the
male and female data sets analyzed separately by the tps
series of programs. (Versions of the individual tps pro-
grams used are mentioned later in the methods section).
Determining w h e t h e r the a m o u n t of variation be-
tween the sexes w a s small enough - Before the two data
sets were analyzed further, it was necessary to determine
 Downloaded by [University of Delaware] at 07:23 09 June 2013

2
to

;>
t')

Figs. 1-2. M a l e s - 1. Hyalomma marginatum rufipes with sixteen landmarks. 2. Hyalomma truncatum without to
landmarks.
 Downloaded by [University of Delaware] at 07:23 09 June 2013

3 4
tO

tO
Figs. 3-4. Females - 3. Hyalomma truncatum with 16 landmarks. 4. Hyalomma marginatum rufipes without 0
0
landmarks (extreme example of each species was used for illustration purposes). 0
 Vol. 26, No. 3 Internat. J. Acarol. 233

I
5 6
M I
fg
I m
[] • m
m m
H. m. rufipes -I" • N H. truncatum
H. m. rufipes
N
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Fig. 5. Relative warp 1 (x-axis) and 2 (y-axis) computed and plotted against one another to indicate positions of the
males of Hyalomma marginatum rufipes and H. truncatum relative to one another and to the reference configuration
(situated at the origin). Fig. 6. Relative warp 1 (x-axis) and 2 (v-axis) computed and plotted against one another to indicate
positions of the females of Hyalomma marginatum rufipes and H. truncatum relative to one another and to the reference
configuration (situated at the origin).

whether the observed variation in shape is sufficiently
small that the distribution of points in the tangent space
can be used as a good approximation of their distribution
in shape space. The 'tpsSmall' program (F. James Rohlf,
Version 1.12) was used to determine whether the amount
of variation in shape in each data set is small enough to
permit statistical analyses to be performed in the tangent
space (which is linear) approximate to Kendall's shape
space (which is nonlinear). In other words, 'tpsSmall'
helps one to assess the accuracy of the approximation of
shape space by the tangent space. The least-squares re-
gression slope (through the origin) and the correlation
(uncentered) between the Procrustes distances and the
Euclidean distance between all pairs were computed to
measure their deviation from a linear relationship.
'TpsSmall' can plot the distances in the tangent space
versus Procrustes distances and if the scatter is close to a
straight line, it implies that the approximation is good
enough for the data set.
Analysis of the variation between taxa using the
nonaffine shape variation (relative warps) - In order to
relate trends in shape of the two sexes, relative warp
analyses (RWA) were performed. An RWA is a principal
components analysis (PEA) of the covariance matrix of
the partial warp scores and is performed by the program,
where a = 0 (Bookstein, 1991). In other words, relative
warps with (x = 0 is just a PCA of shape, and the shape is
expressed in terms of partial warps, but not dependent
upon them (Rohlf, 1993 and personal communication).
An alternative is also to perform a PCA of the alligned
specimens; this will result in an identical ordination. Rohlf
(1993) suggested that a = 0 should be used in these types
of analyses, because a = 0 presents morphometric differ-
ences at all scales. When ot is greater than 0, geometri-
cally small-scale variation is given less weight than the
large-scale variation, the result is that of reducing the
weight given to regions having more landmarks relative
to the weight given to regions having fewer and therefore
more widely spaced landmarks (Rohlfet al., 1996). Rela-
tive warps are computed, using 'tpsRelw' (F. James
Rohlf, Version 1.16), to summarize the variation among
the sexes (with respect to their partial warp scores) in as
few dimensions as possible. Each relative warp can be
plotted as a deformation of the space of the reference
configuration of landmarks. The relative warp analyses
are based on the nonaffine components of shape variation,
which, according to Rohlf et al. (1996) dominate, depend-
ing on the data, the estimate of overall morphological
relationships. The first two relative warps are usually
indicative of most of the variations between members of
the data set, and therefore their scatter plot presents the
most visual information about variation between the
sexes. Shape changes implied by variation along the first
two relative warp axes can be shown as deformations
using thin-plate splines.
Obtaining consensus configurations and thin-
plate splines for each sex of the two species - Because
shape differences between the two sexes of the two spe-
cies were studied, the average or consensus configuration
of landmarks for the 10 specimens for each sex was com-
puted using 'tpsSuper' (F. James Rohlf, Version 1.03).
Orthogonal least-squares Procrustes average configura-
tion of landmarks was computed using generalized or-
thogonaI least-squares procedures. The consensus con-
 234 Pretorius & Clarke 2000

Table 1. Description of landmarks used (males and females) in the geometric morphometric study.

LANDMARKS DESCRIPTION
MALES FEMALES
Points on midline perpendicular to Posterior tip of scutum
1. 2
Widest point on the male body Widest point on the female body
2.
A scutal groove Anterior point of marginal groove
3. of alloscuttma
Eye Widest point on scutum (where
4. scutum and alloscutum meet)
Point where the cervical groove Eye
5. closely approaches the mid line
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Point on midline perpendicular to Point where the cervical groove

6. the posterior end of cervical closely approaches the midline
groove
Point on midline perpendicular to Point on midline perpendicular to
7. the most anterior point of cervical the posterior end of cervical
field groove
Most anterior point of scutum on Halfway between the cervical pit
8. midline and the midline
Intersection - midline and dorsal Anterior point of cervical field
9. ridge of basis capituli where lateral and cervical grooves
originate (cervical pit)
Fusion point on midline of Medial margin of scapula -
10. outer cheliceral sheath posterior point
Anterior lateral comer of basis Most anterior point of scutum on
11. capituli midline
Anterior tip of scapula Intersection - midline and dorsal
12. ridge of basis capituli
Medial margin of scapula - Medial margin of scapula -
13. anterior point anterior point
Medial margin of scapula - Anterior tip of scapula
14. posterior point
Midpoint between 14 and lateral Anterior lateral comer of the basis
15. body wall capituli

Anterior point of cervical field Fusion point on midline of

16. where lateral and cervical grooves outer cheliceral sheath
originate (cervical pit)

figuration for each sex of the species is the mean landmark
configuration for the particular sex and therefore a consis-
tent allignment is needed. In order to visually determine
how the landmarks (and which of the landmarks) vary
between the two male and two female consensus configu-
rations of the two species; the "plot consensus" option of
'tpsSuper' was activated.
Thin-plate splines of the variation between the sexes
of the species can also be drawn, and shape variations,
because of positional movement of landmarks, visualised.
'TpsSplin' (F. James Rohlf, Version 1.14) was used in the
analysis of the two consensus landmark positions of each
of the two sexes because it allows visual determination of
which landmarks are responsible for the differences in
shape.. The computations involve the calculations of a
GLS (Generalized least squares) reference for the males
and females and this reference is represented by a perpen-
dicular Cartesian grid. The shape differences of, e.g., the
female thin-plate spline of each consensus of the two
species can now be visualized by studying the deforma-
tions of each of the consensus configuration thin-plate
splines.
 Vol. 26, No. 3 Internat. J. Acarol. 235

7 a:, 1
O.I-~
q~6 7
,
I0
11 ~°@o ~o
%6 I 8%~12--16 QI~6~BI41K313 11

3 4 i ~ I~ ~ 1 2 ~ )
0 2 OI dllP 5 c~

0 H. truncanum
@ 0 H. truncanum
H. m. rufipes
I~ H. m. rufipes
Inearl
Downloaded by [University of Delaware] at 07:23 09 June 2013

• mean

Fig. 7. Diagram indicating the difference in land- Fig. 8. Diagram indicating the difference in land-
mark position of the consensus configuration of land- mark position of the consensus configuration of land-
marks of females of Hyalomma marginatum rufipes and marks of males of Hyalomma marginatum rufipes and 11.
H. truncatum as well as the mean. truncatum as well as the mean.

A B

L / I f I ~ t l t j
" " -- I l I I l I l
I I , , ' I I %
I I

I-- " "1" - - I - ~t -- -I- - -I- - - - I - -- 1, - - - I - - -I

i_. I I i I ji I I I

o i . .i . . .I . . J. . . i . T--~-- }--J--" q 1:::

"El
o
I~ -/- _l__l---l--J--4-1--)~ @ L_, J._, )_, . I.
i
. I .
i
. I ~! .1_ _,1,~,~ _ ', .~
•

, _ . . . . . I~ _ q_ _ ._~ ,'-- ~i" - - : - -+ - ' - + - ~ . ,. -~ -*

I----I'---i---1 I-- -I- -i- 4,,i

'--,- '" - -- !- ?;! -! ~-I-- - I

=,.7 - - i - ~ - -- - 1
L__I.__I_._L__k~.L,._I__I__I~.j
L _ _L. __t__ J__ L~--I. - t -- - L - J- ~ -I

Fig. 9. Shape variation, shown as deformations of the GLS reference, using thin-plate splines (Landmark positions
on a grid) - A. Female Hyalomma marginatum rufipes. B. Female H. truncatum.

10 B
A
I I .I_. l._l.~.l_-p-_i-~_l
I-----I----L _l .1_ I I ~ I I t I I I I , I I 6 I

r''-1"--I----I---i---4---/---L---l-~-I I | | I I _ I I ~ .i ~ - ¢-.-~- ~ : . ~ .I
- _ ~ i i i i | 41~ ' i q

~- -; ~ , - ~ "
r-,--r-
¢=" " - - i - ~ - , ~ - ~
!2_
.
I
~ . . _ i _
-~ i

I----!----L---I-~ L ~ ~ I I I
I I I I ~ I I I I

F - --I" - " I'- " - - l - ~ - 1-" - - - I - ~ -!-- - _t. _ _ ~ . _ -1' I - - Jr - - - I -

I i I
- 1
i
. . . . : • ~ 1
t ~-
= a -

, ! i I i ~, . . . . ' 2-~-! I I I I ~ ~ I l t I
r-- ~ I.I- - = i - - - "I = "1 "II I I i I "I

~... _I.__L_ _I___L_ _1.~_1~- / _ _ L ~ . I . J.. _1. _ J __ L . J _ _L _ .t_ . _l- -- J

Fig. 10. Shape variation, shown as deformations of the GLS reference, using thin-plate splines (Landmark positions
on a grid) - A. Male Hyalomma marginatum rufipes. ]3. Male H. truncatum.
 236 Pretorius & Clarke
RESULTS
D e t e r m i n i n g the l a n d m a r k s - The males and fe-
males of each of the species are illustrated in Figs. 1 - 4.
(Sixteen landmarks were chosen and the landmark posi-
tions indicated by labelled dots on the male of H. m.
rufipes (Fig. 1) and female of//. truncatum (Fig. 3).
D e t e r m i n i n g w h e t h e r the amount of variation be-
tween the species w a s small enough - The correlation
(uncentered) between the tangent space, Y, regressed onto
Procrustes distance (geodesic distances in radians) for the
males as well as the females is 1.000000, implying that
the approximation is excellent for these data.
Relative w a r p analysis - The first two relative
warps (using ~ = 0) of the male landmarks accotmt for
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68.76% of the variation among the specimens, while these
two relative warps also account for 52.58% of variation
in the females, computed by a singular-value decomposi-
tion of the weight matrix (Rohlf, 1993). The first two
relative warps of data set 1 (male) were plotted against
each other, to indicate variation along the two axes (this
was also done for the female data set). The males of the
two species are found overlapping each other on the scat-
ter plot, indicating that the male body shape is very simi-
laro except for individual variation. The females of the
species form two more distinct clusters, with all the mem-
bers of one species grouped together, but with a few
specimens from each species situated close to one another,
appearing to form a continuum. Figures 5 and 6 respec-
tively show a two-dimensional ordination of the overall
diversity of the males and females of the two species of
Hyalomma in terms of their nonaffine components of
shape variation.
Consensus configurations and thin-plate splines
- Figs. 7 and 8 show the differencse in landmark positions
of the consensus males and consensus females of each
species relative to each other (the mean configuration is
also indicated). Figs. 9 and 10 indicate the thin-plate spli-
nes of the consensus configuration of each of the sexes
(GLS was specified as the reference).
DISCUSSION
The relative warp analysis of the male body form of
the two species indicates that there is a clear overlap of
body shape (Fig. 5). This is, however, less clear in the
female relative warp analysis, although it seems as if a
continuum is present (Fig. 6).
The average or consensus configuration for each sex
was also computed, and the consensus landmark positions
of each sex compared with 'tpsSuper' (Figs. 7 and 8).
From this analysis the "movement" of landmarks can be
2000
seen, and the landmarks responsible for the shape differ-
ences pinpointed.
Landmarks 1, 2, 6 and 15 vary the most between the
female consensus configurations of the two species (Fig.
7). Landmark 1, the posterior tip of the scutum varies the
most between the two species. The landmark is situated
higher in H. m. rufipes, indicating that the alloscutum
covers a larger portion of the body. (The lowest point on
the body of all the specimens was entered as the zero point
for the x, y axis. Landmark 1 is therefore a function of
this zero point).
Landmark 2 is the widest point on the female body.
The body of//. m. rufipes is wider that that ofH. trunca-
tum and the landmark 2 also echoes that. Landmark 6, the
point where the cervical groove closely approaches the
midline, is slightly lower in 1t. m. rufipes. The area be-
tween landmarks 1, 2 and 6 is therefore smaller in H. m.
rufipes than inH. truncatum. Landmark 15 is the anterior
lateral comer of the basis capitulum and is situated higher
in 11. rn, rufipes. The rest of the landmark positions vary
slightly, and landmarks 5, representing the eye, landmark
6 and landmark 12, where the midline intersects with the
posterior edge of the basis capituli, are situated at the same
place in the females of both species.
The positional differences of the consensus land-
marks of the males of the two species do not vary as much
as those of the females, although the displacement seems
more constant between different landmarks (Fig. 8).
Landmark 3 representing a scutal groove, 4, the eye and
5, the point where the cervical groove approaches the
midline, vary the most between the males. Landmark 2
representing the width of the body varies much less than
is the case with the female body width. Hyalomma m.
rufipes, however, appears to be longer than 11. truncatum.
Thin-plate spline analysis (Fig. 9a,b), a more visual
aid to pinpoint the differences in shape indicated that the
spline of the consensus configuration of female H. m.
rufipes (Fig. 9a) tends to widen at the bottom while that
of female H. truncatum (Fig. 9b) narrows slightly.
The spline of the consensus configuration of male
H. m. rufipes (Fig. 10a) indicates that the anterior area of
the scutum is compressed, while the same area in the
consensus configuration of male H. truncatum (Fig. 10b)
is extended. The posterior area of the scutum of H. m.
rufipes is, however, wider than that of H. truncatum.
From the shape analysis of the body of the two tick
species the following can be concluded: relative warp
analysis indicates that, although there are very slight shape
differences between the sexes of the two species, the male
body shape overlaps, while the female body shape forms
a continuum. The consensus configuration comparison
and thin-plate spline analysis, however, indicate that, al-
though there are some shape differences, this is caused by
 Vol. 26, No. 3 Internat. J. Acarol.
very slight landmark movement. Thin-plate spline analy-
sis indicate that the shape of the body of males and fe-
males ofH. m. rufipes (Figs. 9a, 10a) tend to be wider and
extended anterolaterally, while this is the opposite in both
males and females of//. truncatum (Figs. 9b, 10b).
The classification system of the Ixodidae was rela-
tively stable over the past century and this has extended
towards hypotheses of relationship between genera. One
of the aims of this research was to reopen the debate on
the taxonomic and perhaps also the phylogenetic place-
ment of Hyalomma ticks. The main aim of this study was,
however, to present a new technique to analyze morpho-
logical characters with, and, for the first time, to use
members of the Ixodidae. Because geometric morphomet-
rics visually compares shape differences, it presents a new
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and different perspective to an old taxonomic problem that
has not recently been addressed. Although the results of
this research are based only on evidence from one type of
analysis, there is no doubt to the presence of useful evo-
lutionary information within the body shape of ticks.
Geometric morphometric analysis should therefore be
used as another source of information, especially if cladis-
tic data are particularly homoplastic. Hence, we are not
advocating the use of geometric morphometrics as the
only evidence on which to base hypotheses of natural
relationships. A large suite of data should be used in
phylogenetic studies to give the best estimate of evolu-
tionary history.
Our conclusion, therefore, based only on body
shape, is that there are probably not two individual spe-
cies, but rather two subspecies, with speciation still in the
process of taking place. This hypothesis is also substanti-
ated by the great similarities in adult appearance, hosts,
life cycle (Theiler, 1962; Norval, 1982) and distribution
(Howell et al., 1978) of the two subspecies.
ACKNOWLEDGEMENT
The authors would like to thank M. Tumer, EM
Unit, MEDUNSA for the preparation of the electronmi-
croscope phtos.
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