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ORIGINAL INVESTIGATION

Morphological evidence of species differentiation within Lepus capensisLinnaeus, 1758 (Leporidae, Lagomorpha) in Cape Province, South Africa

Fernando Palaciosa,�, Chiara Angelonea,b, German Alonsoc, Santiago Reigd

aDepartamento de Biodiversidad y Biologıa Evolutiva, Museo Nacional de Ciencias Naturales (CSIC), C/Jose Gutierrez Abascal 2,

28006 Madrid, SpainbDipartimento di Scienze Geologiche, Universita Roma Tre, Largo San Leonardo Murialdo 1, 00146 Rome, ItalycDepartamento de Ecologıa, Universidad Complutense, Facultad de Ciencias Biologicas, C/Jose Antonio Novais 2,

Ciudad Universitaria, 28040 Madrid, SpaindUnidad de Medicina Experimental, Laboratorio de Imagen, Hospital General Universitario Gregorio Maranon, C/Dr. Esquerdo 46,

28007 Madrid, Spain

Received 10 October 2007; accepted 25 October 2007

Abstract

A morphological study was carried out of the hares (Lepus) from Cape Province previously assigned to thesubspecies L. capensis capensis, L. c. centralis, and L. c. grantii. The purpose of the study was to characterize the speciesL. capensis Linnaeus, 1758. In doing so, it was possible to distinguish two populations which we consider differentspecies, as each shows homogeneous cranial, dental and pelage features. One of them, defined as L. capensis, isdistributed near Cape Town not far from the coast, between Lambert’s Bay and Cape Agulhas. The other species,defined as L. centralis, which includes L. grantii as a synonym, is distributed in central and western Cape Province.L. capensis and L. centralis have a parapatric distribution, with a small area of sympatry in a contact zone inCompagnies Drift area, near Lambert’s Bay. With respect to cranial differences between the two species, L. capensis

has a stronger maxilla and more robust dental series, while L. centralis has larger tympanic bullae. Among dentalcharacters, L. centralis usually has a deeper groove and more abundant cement than L. capensis in the first upperincisor, and its internal lobe is squared, while in L. capensis is rounded. As for pelage color and pattern, L. capensis

shows a more extended white ventral area than L. centralis. Our results are of interest for further research ontaxonomic problems regarding Old World hare populations in which L. capensis is concerned.r 2007 Deutsche Gesellschaft fur Saugetierkunde. Published by Elsevier GmbH. All rights reserved.

Keywords: Lepus capensis; L. centralis; Taxonomy; Syntypes; Museum collections

Introduction

Lepus capensis Linnaeus, 1758 is the hare species forwhich the largest area of distribution has beenrecognized. It has been described as ranging from south

atter r 2007 Deutsche Gesellschaft fur Saugetierku

07.10.013

r. Tel.: +34 91 4111328; fax: +34 91 5645078.

ndo.palacios@mncn.csic.es (F. Palacios).

to north Africa, from Middle East to eastern Asia, andin the Iberian peninsula (Ellerman and Morrison-Scott1951; Ellerman et al. 1953). Petter (1959, 1961)considered L. europaeus as a synonym of L. capensis,thus expanding the range of L. capensis to the Europeanarea of the brown hare.

Since these revisions, L. capensis has been a widelyaccepted taxon in publications about taxonomy, biology,

nde. Published by Elsevier GmbH. All rights reserved.

Mamm. biol. 73 (2008) 358–370

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ecology and genetics of Old World hares and in regionaland country faunas. Nevertheless, L. capensis remains,at a taxonomical level, one of the most poorlycharacterized species, not only in the case of the nominalsubspecies (L. capensis capensis), but also with respect toother populations assigned to the species.

The scanty data available regarding the nominalsubspecies were provided by Roberts (1951) who, in hisbook on South African mammals, included a shortdescription, with data on body and skull dimensions,and pelage color and pattern of several specimens fromEendekuil, Lambert’s Bay and Vredendal. However, thislast locality has to be considered doubtful, becauseRoberts (1951) assigned the two subadult specimensfrom Vredendal of the Transvaal Museum to L. capensis

grantii when describing the body and skull measure-ments, and to L. capensis capensis when describingpelage pattern. Roberts (1951) also postulated theL. capensis capensis distribution area as covering aterritory comprised between Oudtshoorn and OlifantsRiver (SW Cape Province). It is important to note thatRoberts (1951) described some morphological andchorological differences between different populationsof L. capensis, although he did not assign them ataxonomic rank above the subspecies level. Robinsonand Dippenaar (1987), in their morphometric analysesof lagomorphs from South Africa, confirmed thatno specific differences could be recognized withinL. capensis populations. It is worth to remark thatRoberts (1951) and Robinson and Dippenaar (1987)only studied the scanty specimens of L. capensis capensis

from the Transvaal Museum (Pretoria, South Africa),and did not take into account in their analyses the largeseries preserved in the Amathole Museum (KingWilliam’s Town) and the old collection kept in theSouth African Museum (Cape Town). Petter (1959),who studied the collections of the London and Parismuseums, did not include in his revision specimens fromSW Cape Province and accepted most of the Africanforms listed by Ellerman et al. (1953) under L. capensis.

An analysis of the morphological characters definingL. capensis capensis has never been performed, in spiteof the compelling need of comparing the nominalsubspecies with other L. capensis populations and withother species of the genus Lepus. The lack of a precisecharacterization of L. capensis capensis is also a result ofthe absence of holotype material. Moreover, thedescription of several African hare populations assubspecies of L. capensis led to the acceptance of a highmorphological variability within this species and to therecognition of L. capensis in regions so far away fromSouth Africa as Europe or Asia.

Hoffmann (1998), discussing the taxonomic status ofL. capensis, mentioned that ‘‘there is a state of confusionaround this species which affects to different taxon pairsin eleven geographic contact zones’’. This confusion is due

in part to the emergence and development of criteria todistinguish some local populations within the polytypicL. capensis as valid species, as well as to the failure of theclassic taxonomic criteria set up in general revisions.

As for the validity of classic taxonomic criteria forLepus, the presence of a simple groove in the first upperincisors used by Petter (1959, 1961) to recognizeL. capensis in the three Old World continents hasrevealed to be uneffective. There is evidence thatL. capensis is not the only species showing this feature.Successive studies (Palacios 1996; Palacios and LopezMartınez 1980) showed that hares from Europe(L. castroviejoi, L. granatensis, L. europaeus andL. corsicanus) also are characterized by a simple incisorgroove with a moderate cement filling. The presence ofcement in the incisor groove, used by Robinson (1986) todistinguish L. capensis from L. europaeus, based onForsyth Major’s (1898) statement that L. europaeus lackscement in the incisor groove, also must be considered aninvalid discriminant character. In fact, our data show thatabout 60% of L. europaeus individuals from differentEuropean populations have cement in the incisor groove.

Today, the distribution of L. capensis is considered toinclude Southern, Northern and Eastern Africa, theMiddle East, and Western Asia (Flux and Angermann1990; Hoffmann 1993). Its European distribution isexclusively limited to Sardinia (Amori et al. 1996).

In the light of the confusion over range, distribution,and species limits, it becomes clear that there is apressing need to clarify and define the taxonomy ofAfrican, Asian, and European hares nominally identi-fied as L. capensis. Such an investigation has a specialweight and priority in Cape Province, South Africa,wherein is located the type locality of L. capensis.

The purpose of this paper is the morphologicalcharacterization of Lepus capensis capensis. An examina-tion and definition of the morphological variation in thetype locality would enable its identification fromphenotypic traits. This is necessary in order to comparepopulations putatively belonging to the nominal sub-species with other populations attributed to the samespecies in Africa, Europe and Asia; this study also will bea starting point for further systematic studies of Lepus.

The study area was limited to western Cape Province.In this area are also included the distributional ranges oftwo additional subspecies of L. capensis: L. c. grantii

and L. c. centralis. These subspecies were recognized byRoberts (1951), but have remained poorly defined froma morphological perspective.

Material and methods

General aspects

The studied specimens are held in most cases under the

species name L. capensis in the collections of the Transvaal

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Museum (TM), Pretoria; Amathole Museum, formerly Kaf-

frarian Museum (KM), King Williams Town; National

Museum (NMB), Bloemfontein; South African Museum

(ZM), Cape Town; Natural History Museum, formerly British

Museum of Natural History (BMNH), London; Museum

National d’Histoire Naturelle (MNHN), Paris; Naturhistor-

isches Museum (NMW), Vienna; National Museum of

Natural History, formerly United States National Museum

(USNM), Washington, DC; Carnegie Museum of Natural

History (CMNH), Pittsburgh.

The morphological study was based on skull measurements,

dental characters in occlusal and crossed sections, and

pelage design of adult specimens of relative age IV. Only a

few specimens of relative age between III and IV have been

added to complete the samples. Specimens of relative ages I, II

and III have been studied only for identification. The latter

two age classes, composed of subadults, are abundant in the

studied collections but have not been included in the analyses

to avoid population range overlap in cranial and dental

measurements and in morphological character states. Only

two juvenile specimens of relative age I have been used to

demonstrate the existence of species differences also in young

animals.

Relative age estimations have been made on the

basis of cranial ossification and development (Cabon-

Raczynska 1964; Palacios 1989, 1996), and considering as

adults the specimens showing the following features: sutures

between basioccipitale-exooccipitale and exooccipitale-supraoc-

cipitale ossified; interparietal bones ossified; sutura sagittalis

ossified in its posterior portion; osseous coating of the

mandibular capitulum knitted and polished; well-developed,

non-porous supraorbital bones; well developed, angular-

shaped temporal tubercle; premaxillaries and corpus mandibu-

lae not porous.

The studied specimens’ provenance is western Cape

Province. Some specimens are from the surroundings of

Cape Town, between Saldanha Bay and Cape Agulhas,

and other from the Upper Karroo, Lower Karroo, Little

Namaqualand and Bushman Land (see Roberts 1951).

In particular, the analysis of the collections of the South

African Museum and Amathole Museum allowed us to

substantially increase the amount of L. capensis material from

the Cape of Good Hope and surrounding area with respect to

previous studies (Roberts 1951; Robinson and Dippenaar

1987).

Collection numbers, localities, and relative age of the

studied specimens are listed in Table 1. The complete list of

the examined specimens that includes juveniles, subadults, and

adults may be requested to the authors.

The dental nomenclature used in this study follows

Palacios and Lopez Martınez (1980). The study of teeth

morphology variation was based on occlusal surface

enamel drawings except for the first upper incisor, where

both sectioned and occlusal surface enamel drawings were

used indistinctly. There was considerable consistency in

the incisor enamel pattern in a sample of 19 specimens in

terms of the studied character states in both occlusal surface

and cross-section enamel drawings. Drawings were carried

out using a camera lucida and refracted light on the teeth

surface which was perpendicularly oriented to their own

longitudinal axes.

Measuring protocol

We used a set of 44 cranial, 7 mandibular, 5 dental, and 3

foramina measurements (see Table 2). Description of these

measurements follows Cabon-Raczynska (1964) and Palacios

(1989, 1996). To avoid transcription errors while collecting

data, measurements were recorded to the nearest 0.01mm

using a digital caliper connected to a laptop computer using

the automatic data capturing software EASYCAL (Marcus

and Reig 1991).

A measurement error test was performed to assess the

precision of the morphometric characters. Repeatability was

calculated by measuring a series of 10 skulls from a single

locality three times and estimating the variance among skulls

and within repeats (repeatability ¼ s2 among/s2 within+s2

among; Baley and Byrnes 1990). Repeatability was over 92%

in all measurements.

Data analysis

Sample statistics were calculated separately for each group,

pooling together specimens from both sexes. To assess the

significance of potential differences between species in mean

values, Student’s T test was performed on each variable. The

univariate statistical tests were performed to illustrate the main

taxonomic differences between the two species using a wide

variety of variables. Thus, no correction for multiple

comparisons (i.e. Bonferroni) was performed, under the

assumption that the analysis was rather exploratory, and not

intended to confirm or discard the taxonomic differentiation

between the two species. The main conclusion about morpho-

metric differentiation between the two species is drawn from

the multivariate analysis that follows.

Principal component analysis (PCA) was performed to

transform a set of linear measurements into a set factors that

joined those measurements having a similar and independent

variability patterns. The principal components (PC) extracted

served to describe the main patterns of variability within the

whole sample without any a priori hypothesis about the species

assignment of each specimen. Factors were extracted from the

variance–covariance matrix of log-transformed values (Marcus

1990) using PROC PRINCOMP (SAS). Extracted PC were not

rotated to ensure that they represented the main axis of

allometric variability in the data. A subset of 10 traits was

selected for the PCA. This reduction in the number of variables

was aimed to get a better ratio in the number of variables relative

to the number of skulls measured for multivariate analysis, while

providing an adequate depiction of skull morphology. Selection

of the 10 morphometric characters was made according to three

basic criteria: depiction of overall skull dimensions (i.e. trying to

produce an even coverage of skull shape), power to discriminate

among populations or species, and maximum repeatability.

To test for differences between the two species in the

multivariate space of the 10 variables used in the PCA, a

MANOVA was performed using the log-transformed vari-

ables. Prior to the MANOVA analyses, the assumption of

homogeneity of variance–covariance matrices across groups

was tested to p40.05 using PROC DISCRIM (SAS). All the

data analyses were performed using the software developed by

SAS Institute Inc. (1989).

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Table 1. Adult specimens used for the study of skull morphology and univariate and multivariate analyses of skull dimensions and

for the study of teeth morphology

Species Relative age Collection number Locality Skull Teeth

L. capensis IV KM 10486 Compagnies Drift X X

KM 10488 Compagnies Drift X X

KM 10490 Compagnies Drift X X

KM 10493 Compagnies Drift X

KM 10504 Compagnies Drift X

KM 10505 Compagnies Drift X X

KM 10512 Compagnies Drift X X

KM 13040 Gouna, E Calvinia X

KM 30196 Darling X X

ZM 7212 Stellenbosch X X

ZM 13673a Stellenbosch X X

ZM 13673b Stellenbosch X X

ZM 35556 Hopefield, Malmesbury X X

III–IV KM 10487 Compagnies Drift X

KM 10494 Compagnies Drift X

KM 10495 Compagnies Drift X X

KM 10498 Compagnies Drift X

KM 10507 Compagnies Drift X

TM 8859 Eendekuil X

TM 8861 Eendekuil X

TM 41457 Langebaan X

L. centralis IV KM 10482 Eenriet X X

KM 10491 Compagnies Drift X X

KM 10496 Compagnies Drift X X

KM 10503 Compagnies Drift X

KM 10510 Compagnies Drift X

KM 11501 Deelfontein X

KM 11503 Deelfontein X

KM 13020 Calvinia X

KM 13022 Calvinia X

KM 13027 Gouna, E Calvinia X

KM 14850 Sutherland X

KM 14852 Sutherland X

KM 18226 Beaufort West X

KM 29028 W Williston X

KM 29032 Fraserburg X X

ZM 19131 Ezelfontein X X

TM 6022 Calvinia X

TM 27292 Farm Gonas, W Carnarvon X

TM 27293 Farm Gonas, W Carnarvon X X

TM 27296 Farm Gonas, W Carnarvon X

TM 28049 Bitterfontein X

TM 28963 Springbok X

TM 28979 Hutchinson X

TM 28980 Hutchinson X X

TM 28981 Hutchinson X X

TM 28982 Hutchinson X

TM 28983 Hutchinson X

TM 28984 Hutchinson X

TM 32452 Farm Vissenvil, Kenhardt X

TM 37999 Deelfontein X

BMNH 4.2.3.109 Port Nolloth X

BMNH 2.9.1.100 Deelfontein X X

III–IV KM 10483 Eenriet X

F. Palacios et al. / Mamm. biol. 73 (2008) 358–370 361

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Table 1. (continued )

Species Relative age Collection number Locality Skull Teeth

KM 10485 Ezelfontein X

KM 13041 E Calvinia X

KM 23246 Victoria West X

KM 23247 Victoria West X

ZM 8576 Namaqualand X

TM 2156 Lamberts Bay X

TM 7308 Vredendal X

Table 2. Skull measurements and their abbreviations.

Abbreviation Measurement

AH Alveolar height, between external margin of

alveoli dentales and the top of bulla alveolaris

maxillae

ANW Anterior nasal width

BCL Brain-case length (Acrocranium—

intersection of frontal suture with the line

joining the two anterior borders of incisura

postorbitalis)

BCW Brain case width

BL Basal length (Prosthion–Basion)

CBL Condylobasal length (Prosthion—posterior

plane of the occipital condyle)

CSL Coronary suture length

DFZ Diameter of foramen zygomaticus

DLFP Diameter of lateral foramen palatinum

DPFP Diameter of posterior foramen palatinum

DIRIMS Distance between first upper incisor

posterior end and sutura incisivomaxillaris

EDMA External diameter of meatus acusticus

ENL External nasal length

FIL Frontoincisive (facial) length (Prosthion—

intersection of frontal suture with the line

joining the two anterior borders of incisura

postorbitalis)

FSL Frontal suture length

FTL Facial tubercle length

FUISW First upper incisor section width

FUISL First upper incisor section length

HBC Height of the brain case between the

basisphenoideum and ossa parietalia

HCMM1 Height of corpus mandibulae at lower first

molar level

HDFM Horizontal diameter of foramen magnum

HPF Height between processus palatinus and the

nasal processes of frontal bone

IFL Foramen incisivum length

IFW Incisive foramina width

INL Internal nasal length

LCTRL Lower cheek-tooth row length

LMRML Length of ramus mandibulae

LP3SL Lower third premolar section length

LP3SW Lower third premolar section width

MCL Mandible capitulum length

MLAP Mandible length, taken from the angular

process

Table 2. (continued )

Abbreviation Measurement

MLCP Mandible length, taken from the capitulum

MH Mandible height between inc. varosum and

the capitulum

MMPH Height of processus muscularis mandibulae,

between the anterior margin of processus

articularis and the tip of the process,

perpendicular to the margin

NPW Narrowing of the palatine bridge

ONL Occipitonasal length (anterior nasal tip—

Acrocranion)

PL Palatal length

PNW Posterior nasal width

PPW Postpalatal width

POW Preorbital width

PZW Posterior zygomatic width

RIC Radius of first upper incisor curvature

RW Rostral width

SFW Smallest frontal width

SOOL Length of medial part of pars parietalis of

squama ossis occipitalis

SOOW Width of medial part of pars parietalis of

squama ossis occipitalis

SSL Sagittal suture length

SSOL Sagittal suture ossification length

TBL Tympanic bullae length

TBW Tympanic bullae width

TL Total skull length (Prosthion–Acrocranion)

TW Tympanic width

UCTRL Upper cheek–tooth row length

UDL Upper diastema length

UP4SL Upper fourth premolar section length

VDFM Vertical diameter of foramen magnum

WSP Width between supraorbital processes

WFT Width between facial tubercles

ZAL Zygomatic arch length

F. Palacios et al. / Mamm. biol. 73 (2008) 358–370362

Results

Two hare populations were differentiated on the basisof their morphological features. Specimens in thesepopulations are in both cases hares showing simpleincisor grooves, previously identified as characteristic ofLepus capensis. The distinction between the two

ARTICLE IN PRESSF. Palacios et al. / Mamm. biol. 73 (2008) 358–370 363

populations is further supported by the homogeneity ofthe morphological characters within their respectiveareas of distribution and by the presence in somelocalities of specimens of the two populations. This factreflects the existence of partial sympatry in the limit oftheir areas of distribution, although due to the scarcityof available specimens it has not been possible todetermine if this situation also exists in additionalcontact areas.

The results enabled us to establish that the twopopulations belong to distinct species. One of them fitsin the distribution area of L. capensis capensis as definedby Roberts (1951). For this reason it is designated fromhere on as L. capensis. The second population fits withinthe areas of distribution of L. capensis centralis Thomas,1903 and L. capensis grantii Thomas and Schwann, 1904and is designated henceforth as L. centralis as that namehas taxonomic priority.

Fig. 2. Skull posterior view of adult specimens (relative age

IV) of L. capensis KM 10490 Compagnies Drift (left) and L.

centralis KM 28963 Springbok (right).

Skull morphology

Adult individuals of the two species show markedlydifferent skull morphologies (Figs. 1 and 2). Forinstance, the maxillary bone and the masticatoryapparatus are more robust in L. capensis than inL. centralis. Other differences can be noticed in thepalatine bridge, where the lamina horizontalis is wider in

Fig. 1. Skull dorsal and ventral views of adult specimens (relative

(above) and L. centralis (below). L. capensis specimens: ZM 35556 H

specimens: KM 18226 Beaufort West (adult); KM 13021 Calvinia (

L. capensis. This latter species also shows more massivefacial tubercles.

Lepus centralis is characterized by a more gracile skullin its rostral part, due to the narrowness of thepremaxillary region and a lesser development of thefacial tubercles. On the other hand, L. centralis shows agreater development of the posterior part of the skullwith respect to L. capensis, due to larger dimensions ofthe tympanic bullae and of the squama ossis occipitalis

(Fig. 2) as well as a larger width of the braincase.Lepus capensis also shows more developed nasal

bones lying in a more advanced and higher position

age IV) and juvenile specimens (relative age I) of L. capensis

opefield (adult); KM 18322 Bredasdorp (juvenile). L. centralis

juvenile).

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with respect to the premaxillary bone if compared toL. centralis. As far as this feature is concerned,L. capensis is more similar to the L. victoriae–L. saxatilis

group than to L. centralis, even if in L. capensis the nasalprotrusion never reaches that of the specimens of theabove mentioned group. The shape of the nasal bonesalso distinguishes L. capensis and L. centralis: theposterior border is squared in the former and roundedin the latter. Moreover, in L. capensis the external edgeof the parietal bone is rounded, while in L. centralis it isstraight and parallel to the internal edge.

The distinctive cranial characters of both species areevident since the first ontogenetic stages, as shown bythe young individuals of relative age I of both species(Fig. 1). It is worth remarking that the L. centralis

specimen shows much more developed tympanic bullae

in spite of its lesser size than the L. capensis specimen.The difference in the posterior contour of nasal bones isalso evident in Figure 1.

Skull dimensions

Univariate analysis

The results of the univariate statistical analysisperformed on cranial measurements of adult specimensof both populations are reported in Table 3. In spite ofthe general size of the skull being similar in the twospecies (see TL and PZW values), L. capensis dimen-sional values are significantly higher than those ofL. centralis in the following cranial measurements:HPF, AH, UCTRL, LCTRL, RW, WFT, POW, ENL,PNW, NPW, HCMM1, and in particular in dentalmeasurements UP4SL, LP3SW and FUISW. On theother hand, L. centralis shows significantly higher valuesof skull measurements FIL, TBL, TBW, DIRIMS andSOOW than L. capensis. Moreover, the ratio ONL/TLis higher in L. capensis due to its more prominentnasal border with respect to the premaxillary anterioredge. These results confirm the morphological observa-tions highlighted in the section on skull morphology(Figs. 1 and 2).

Multivariate analysis

A PCA of skull measurements was performed.Although five PC were necessary to explain 90% ofthe variance, we focused on the description of the firstthree, which seem to convey the maximum informationabout morphometric differences between species. Thefirst three principal components explain 79%(50,5+17,9+10,5%) of the analyzed species variability(Table 4). The first PC describes the most evidentdifferences between the two species and can be inter-preted as the result of the combined variation of SOOWand in a lesser extent of TBL and TBW, as well as thatof NPW, AH and UCTRL in the opposite direction.

The second PC describes the combined variation ofSOOW and NPW (positive), and of TBL and TBW(negative). The third component describes variation ofgeneral size, particularly in PNW, HPF, TBW, TBL,AH and WFT. Bivariate plots of PC 1 versus PC 2 orPC 3 show a marked separation of the two species alongthese PC axes (Figs. 3 and 4). Thus, most of thevariability (80%) explaining differences between the twospecies is due to shape rather than size. This translatesinto a higher probability that such variables have ataxonomic rather than ecological significance (i.e. sizegradients related with climate or environmental factors).The MANOVA test performed on the set of 10 variablesindicated highly significant differences between the twospecies (F(10,38) ¼ 30.6, po0.00001).

Dental morphology

Data on the character states allowing the discrimina-tion between L. centralis and L. capensis are shown inFigure 5 and Table 5. The most important differencesbetween the two species are concentrated in the firstupper incisor, where L. centralis shows a narrow,squared I1/ internal lobe, slightly more prominent thanthe external one; its posterior contour is flat or slightlyconcave, and the incisor groove is very deep andcompletely filled with cement. L. capensis shows a wide,rounded and more prominent I1/ internal lobe; theposterior contour of the I1/ is generally concave and theincisor groove usually is more shallow and has littlecement.

A higher degree of crenulation and complexity in theenamel of P2/, P3/ and P/3 entrants is also noticeable inL. capensis, especially in the number of loops in theanterior side of the P3/ hypoflexus. In P/3 morphology,it also stands out that the length and width of theocclusal section in L. centralis are approximately thesame and the internal contour is straight, while inL. capensis the tooth section is longer and its internalcontour is rounded.

Pelage pattern

White pelage is more extended in the underparts ofL. capensis than in L. centralis (Fig. 6). In the former,the white pelage reaches the posterior part of the breastand the base of the anterior limbs, while in L. centralis itis reduced to a patch along the centre of the belly,separated from breast and limbs by the transitionalvinaceous-pinkish pelage of the sides. These resultsagree with the pelage description of the subspecies ofL. capensis capensis and L. capensis centralis by Roberts(1951). Nevertheless, the study of pelage color deservesfurther investigation as it is necessary to determine theextent of potential intrapopulation variability.

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Table 3. Sample statistics of the skull measurements of adult specimens of L. centralis and L. capensis from western Cape Province

L. centralis L. capensis P

N MEAN MAX MIN N MEAN MAX MIN

ONL 35 85.4 89.7 82.2 20 86.8 90.7 83.6 0.008373

TL 36 87.2 91.6 84.0 20 87.4 91.7 84.0 ns

CBL 36 77.9 82.4 74.8 20 78.0 81.6 75.2 ns

BL 35 70.9 74.7 67.3 20 70.8 74.5 68.4 ns

PL 36 34.3 37.0 32.9 20 35.3 38.0 32.9 0.000792

FIL 36 57.1 60.5 54.5 20 59.1 62.0 56.3 0.000014

BCL 36 43.9 47.4 40.4 20 43.7 45.3 41.7 ns

INL 35 28.9 32.2 23.7 20 30.4 32.9 27.5 0.006030

ENL 35 36.2 40.3 34.1 20 38.0 40.5 35.3 0.000040

UCTRL 36 14.9 16.2 13.6 20 16.0 17.0 15.1 0.000000

UDL 36 25.8 28.8 24.0 20 25.7 27.8 24.3 ns

IFL 36 22.6 24.7 20.9 20 22.4 24.4 20.9 ns

ZAL 36 33.9 36.8 31.6 20 35.1 37.1 33.1 0.000873

MLAP 30 61.0 65.2 57.8 17 61.9 64.3 58.3 ns

MLCP 33 64.1 68.5 61.5 18 65.6 67.6 63.3 0.001757

LCTRL 35 15.8 17.1 14.5 20 17.2 18.2 16.4 0.000000

BCW 36 30.4 32.4 28.2 20 30.3 32.4 28.3 ns

POW 34 16.7 20.9 14.4 14 18.9 20.6 16.4 0.000009

SFW 36 12.1 14.0 9.7 20 12.4 14.2 10.6 ns

PZW 34 40.1 41.8 37.3 20 40.2 42.3 47.7 ns

WSP 32 29.1 32.5 26.9 14 29.7 32.1 27.2 ns

ANW 36 10.5 12.1 9.0 20 11.1 12.1 9.8 0.008527

PNW 36 19.0 22.0 17.3 19 20.3 22.1 18.8 0.000030

RW 36 23.6 25.1 22.2 20 24.6 25.6 23.5 0.000004

WFT 34 35.8 37.8 33.8 19 37.7 40.3 35.5 0.000004

PPW 36 8.6 9.7 7.3 20 8.6 9.5 7.8 ns

IFW 36 10.0 11.4 8.2 19 9.8 11.3 8.6 ns

NPW 36 5.5 7.1 4.5 20 6.1 7.0 5.4 0.000062

HBC 36 25.4 27.9 24.3 20 25.0 26.4 23.4 ns

HPF 36 20.1 22.0 19.0 20 21.7 23.2 20.4 0.000000

MH 33 38.7 41.5 35.7 19 40.0 42.1 37.0 0.005221

VDFM 31 12.4 14.2 10.9 19 12.0 13.7 10.7 ns

HDFM 35 10.6 11.6 9.6 20 10.7 11.8 9.9 ns

TBL 36 13.1 14.7 12.1 20 11.7 12.8 10.9 0.000000

TBW 36 9.6 10.7 8.4 20 8.8 9.9 7.9 0.000001

AH 36 14.6 15.9 13.5 20 15.8 17.6 14.7 0.000000

FSL 36 33.4 38.0 28.4 20 34.8 37.4 31.2 0.012851

SSL 36 19.7 23.1 17.4 20 20.3 22.1 17.9 ns

CSL 36 23.4 25.3 21.7 20 23.9 26.5 21.8 0.048712

TW 27 35.6 38.1 33.5 16 34.7 37.4 31.8 0.029613

EDMA 33 6.7 7.7 6.0 16 6.3 7.6 5.6 0.010062

FTL 29 7.5 9.4 6.4 17 7.7 9.2 5.5 ns

MMPH 33 2.3 3.1 1.7 19 2.5 6.8 1.6 ns

SSOL 35 10.0 14.9 4.5 19 10.1 13.6 6.5 ns

DIRIMS 36 �1.0 1.5 �2.9 20 �0.1 1.3 �1.5 0.000346

SOOL 35 14.9 18.2 12.5 20 14.5 15.5 13.2 ns

SOOW 35 11.0 13.5 8.6 19 9.6 11.3 7.5 0.000040

FUISW 36 2.4 2.7 2.2 20 2.6 2.8 2.5 0.000001

FUISL 36 1.7 2.0 1.6 20 1.8 2.1 1.7 0.000524

UP4SL 34 2.2 2.4 2.0 19 2.4 2.6 2.3 0.000000

LP3SL 35 3.2 3.5 2.8 20 3.5 3.8 3.2 0.000001

LP3SW 35 2.9 3.2 2.6 20 3.1 3.4 2.8 0.000000

MCL 34 8.3 9.6 7.3 19 8.5 9.5 7.7 ns

LMRML 31 24.6 26.7 21.3 19 24.2 26.1 22.6 ns

HCMM1 35 12.9 14.2 11.3 20 13.7 15.5 12.7 0.000026

F. Palacios et al. / Mamm. biol. 73 (2008) 358–370 365

ARTICLE IN PRESS

Table 3. (continued )

L. centralis L. capensis P

N MEAN MAX MIN N MEAN MAX MIN

RIC 36 9.6 10.8 8.3 19 9.5 11.3 8.2 ns

DFZ 26 0.7 1.8 0.0 20 0.9 2.0 0.3 ns

DPFP 26 0.6 0.7 0.4 20 0.5 0.7 0.4 ns

DLFP 26 0.2 1.2 0.0 19 0.1 0.4 0.0 ns

N, number of analyzed specimens; MEAN, mean values; MAX, MIN, maximum and minimum values; P, statistical significance of the differences

between the two groups (Student’s T test); ns, not significant. List of studied specimens in Table 1. Alphabetical list of skull measurements in Table 2.

Table 4. Eigenvectors and variable components for the first

three PC extracted from the variance–covariance matrix of the

10 morphological characters selected

PC1 PC2 PC3

AH 0.234 0.051 0.251

UCTRL 0.238 0.008 0.185

TBL �0.358 �0.273 0.351

TBW �0.274 �0.336 0.396

HPF 0.199 0.081 0.371

ENL 0.159 �0.008 0.195

PNW 0.178 �0.021 0.577

WFT 0.146 0.054 0.253

NPW 0.378 0.581 0.068

SOOW �0.651 0.680 0.213

% Var. expl. 50.5 17.9 10.5

Eigenvectors are not rotated and scaled to unit length, to facilitate

interpretation of character loadings within each PC.

-0.40

-0.30

-0.20

-0.10

0.00

0.10

0.20

0.30

0.40

-0.30 -0.20 -0.10 0.00 0.10 0.20 0.30

PC2

PC

1

L. capensis

L. centralis

g

c

Fig. 3. Scatter plot of PC 1 and PC 2 scores for the

whole sample of the populations from western Cape

Province of L. capensis and L. centralis. Holotypes of

L.centralis and L. centralis grantii, identified with the names

L. capensis centralis and L. c. grantii at the BMNH collection,

are marked with c and g.

-0.40

-0.30

-0.20

-0.10

0.00

0.10

0.20

0.30

0.40

-0.20 -0.10 0.00 0.10 0.20

PC3

PC

1

L. capensis

L. centralis

g

c

Fig. 4. Scatter plot of PC 1 and PC 3 scores for the whole

sample of the populations from western Cape Province of

L. capensis and L. centralis. Holotype specimens of L. centralis

and L. centralis grantii are marked with c and g.

F. Palacios et al. / Mamm. biol. 73 (2008) 358–370366

Distribution

Localities in SW South Africa of the two species areshown in Figure 7, with distribution of the vegetationtypes following Keay et al. (1958). In W Cape Province,L. capensis occupies a small area that may beprovisionally established in accordance with the local-ities of collection specimens between Cape Agulhas andLamberts Bay. Some of the most remarkable localitiesof L. capensis by their proximity to the Cape of GoodHope are Bredasdorp, Swellendam, Stellenbosch, Mal-mesbury, Hopefield, West Coast National Park. Therange of L. capensis overlaps with the Cape Fynbos, avegetation type defined as Cape Macchia by Keay et al.(1958), which is distributed from the Atlantic to theIndian Ocean in the area near Cape Town.

The range of L. centralis stretches through the CapeProvince portion lying to the south of Orange riverand between the Atlantic coast and three localities inthe east: Philipstown, Richmond and Graaf Reinet.

ARTICLE IN PRESS

L. c

a pe n

sis

P2/I1/

1 mmL

.ce n

t ral

is

P3/ P/3

Fig. 5. Comparison of teeth morphologies in L. capensis (above) and L. centralis (below): I1/, upper first incisor; P2/, upper second

premolar; P3/, upper third premolar, P/3; lower third premolar. See text and Tables 1 and 5 for further details. Drawings of L.

centralis correspond to specimen KM 10491 Compagnies Drift; those of L. capensis to ZM 13673a Stellenbosch, except for P/3

which corresponds to KM 10504 Compagnies Drift.

Table 5. Comparison of some selected characters of occlusal and crossed dental sections in L. capensis and L. centralis

Dental

element

Character L. centralis L. capensis

I1/ General shape Somewhat squared Somewhat rectangular

Internal lobe shape Narrow, squared, partially higher than

external one

Broad, rounded, fully elevated

Posterior contour Usually flat Slightly concave

Groove depth Very deep Deep

Cement Very abundant Not abundant

P2/ Crenulation depth on paraflexus

anterior edge

Absent or very shallow undulation Shallow to medium depth

Loop number on paraflexus

anterior edge

0–1 Usually 1 or more

P3/ Hypercones relative thickness Mesial hypercone approximately as thick as

distal hypercone

Mesial hypercone thicker than

distal hypercone

Loop density on hypoflexus

anterior edge

Usually less than 7 Usually 7 or more

P/3 Lingual anteroconid outline Smooth Wavy

Paraflexid depth (when present) Deep Very deep

Crenulation posterior edge

hypoflexid

Smooth or slightly undulated Usually slightly undulated

Lingual contour Straight Rounded

I1/, upper first incisor; P2/, upper second premolar; P3/, upper third premolar; P/3, lower third premolar. List of studied specimens in Table 1.

F. Palacios et al. / Mamm. biol. 73 (2008) 358–370 367

The southern localities of L. centralis near the range ofL. capensis are, from east to west, Beaufort West,Sutherland, Kruidfontein, Calvinia and Lamberts Bay.In the latter two localities (specifically in CompagniesDrift and Gouna), sympatry between the two species hasbeen detected. However, it is noticeable that nocollection examined holds L. centralis specimens fromthe L. capensis exclusive area, between Cape Agulhasand Cape Columbine, suggesting that the two speciesalso have distinct areas of parapatry.

The range of L. centralis overlaps with vegetationtypes defined by Keay et al. (1958) as Subdesert Steppe(Karoo shrub and grass) and Karoo Succulent Steppe.

Discussion

The two hare populations differ distinctly in theircranial and dental size and morphology, pelage colorpattern, and geographical distribution. Moreover, they

ARTICLE IN PRESS

Fig. 6. Outline of mounted skins of a group of specimens in ventral view taken from a photograph, showing belly white pelage

patches. Specimens from left to right are: L. centralis: KM 13027 Gouna, KM 13041 Gouna, KM 10482 Eenriet; L. capensis: KM

10488 Compagnies Drift, KM 10495 Compagnies Drift, KM 10486 Compagnies Drift.

CAPE PROVINCE

AtlanticOcean

IndianOcean

Orange River

29

15 15

152

29

2928

32

30

Kenhardt

Williston

FarmVissenvil

FarmGonas

CarnarvonDeelfontein

Victoria West

Hutchinson

Beaufort West

FraserburgGouna

Sutherland

Porth Nolloth

Namaqualand

Eenriet

Springbok

Ezelfontein

Bitterfontein

VredendalCalvinia

Compagnies Drift

Lamberts Bay Graafwater

Langebaan

Darling

Hopefield,Malmesbury

Stellenbosch

Bredasdorp

Eendekuil

2520

Fig. 7. Geographic distribution of the localities of the studied specimens of L. capensis and L. centralis in Cape Province and

distribution of the Cape vegetation. (Black circles) Localities of L. capensis; (empty circles) localities of L. centralis; (2) temperate

and subtropical evergreen forest; (15) Cape macchia; (20) undifferentiated: relatively dry types; (25) wooded steppe with abundant

Acacia and Commniphora; (28) Karoo succulent steppe; (29) Subdesert steppe: Karoo, shrub and grass; (30) subdesert steppe:

transitional and mixed Karoo; (32) desert.

F. Palacios et al. / Mamm. biol. 73 (2008) 358–370368

maintain homogeneous characters in their respectivedistribution areas even in the small area of sympatrythat they share in Lambert’s Bay and Calvinia. Ittherefore follows that they should be considered asdistinct species.

On the basis of the specimen localities of the twospecies and the distributional data of Roberts (1951), wedesignate the species whose distribution correspondsto the neighborhood of the Cape of Good Hope asL. capensis, while the species living to the north and

northeast as L. centralis. The latter includes L. grantii

as a synonym, as both show similar morphologiccharacters and the skull of the holotype of L. grantii

fits well within L. centralis with respect to variability(Figs. 3 and 4).

In view of the fact that there is no type material ofL. capensis in collection holdings, we propose thefollowing specimens as syntypes: ZM 7212 adult male,ZM 13673a adult female, and ZM 13673b adultmale (specimens from Stellenbosch), ZM 35556

ARTICLE IN PRESSF. Palacios et al. / Mamm. biol. 73 (2008) 358–370 369

adult (Hopefield, Malmesbury), KM 18322 juvenile(Bredasdorp), and KM 30196 adult female (Darling).

This work constitutes a morphological characteriza-tion of L. capensis in the area in which the species’type locality is putatively included, such that theconcrete characters which define L. capensis canhenceforth be used comparatively in analysis of otherspecies and populations. Until now, the results ofprevious taxonomic work, both the classical studies(Roberts 1951) and the recent morphometric studies(Robinson and Dippenaar 1987), included all themedium sized hares from South Africa showing a simplegroove in the first upper incisor in the speciesL. capensis, thus accepting an excess of morphologicvariability in a single species. This consequently led tomorphologic range overlap.

In Europe, where the scarce knowledge on haretaxonomy has been similar in the past decades, somehare populations referred to L. capensis by Petter (1961)have been recently recognized as independent species. Inparticular: L. granatensis (Iberian peninsula), L. corsi-

canus (Italian peninsula and Sicily), and L. europaeus

(widespread in continental Europe) have been consid-ered valid species on the basis of morphology andchorology (Palacios 1983, 1989, 1996), in concordantcombination with biochemical (Bonhomme et al. 1986)and molecular analyses (Perez-Suarez et al. 1994;Pierpaoli et al. 1999; Estonba et al. 2006).

Further research will determine whether the dis-tribution of L. capensis spreads to the east of theCape of Good Hope, and whether it is bigger thanthe small area that surrounds the type locality. It mayalso be possible to determine whether other haremetapopulations from Southern Africa belong toL. capensis.

L. centralis is provisionally recognized as a fullspecies, so that the appropriate distinction can beestablished with L. capensis in the study area. Newinvestigations taking into account material from neigh-boring areas will allow the recognition of a definitivename for that population considering synonyms andpriorities. Future research should also encompassmolecular studies to shed further light on the character-ization of L. capensis in the type locality that can be usedcomparatively in taxonomic analyses of other speciesand populations.

Acknowledgements

We thank the curators, collection managers andtechnicians charged of the studied mammal collectionsfor their help: Lloyd Wingate, Fred Kigozi, AmatholeMuseum (King William’s Town); Stephan du Toit, NicoAvenant, Johan Eksteen, National Museum (Bloemfon-tein); Liz Herzholdt, Naas Rautenbach, Gary Bronner,

Teresa Kearney, Duncan MacFadyen, Transvaal Mu-seum (Pretoria); Michael Cluver, Denise Hammerton,Melissa Stander, South African Museum (Cape Town);Paulina Jenkins, Richard Harbord, Natural HistoryMuseum (London); Francis Petter, Michel Tranier,Jacques Cuisin, Museum National d’Histoire Naturelle(Paris); Friederike Spitzenberger, NaturhistorischesMuseum (Vienna); Michael Carleton, National Museumof Natural History (Washington); and Duane Schlitter,Steve Williams, Carnegie Museum of Natural History(Pittsburgh). We also thank Rainer Hutterer for theabstract translation to German. The comments of LuisRuedas, the anonymous reviewer, and the editor werevery helpful to improve the manuscript and areacknowledged. Financial support is greatly acknowl-edged from the Consejerıa de Educacion, Community ofMadrid, and the Direccion General de Investigacion,Secretarıa de Estado de Polıtica Cientıfica y Tecnologicaof Spain.

Zusammenfassung

Morphologische Hinweise auf artliche Differenzierung

innerhalb des Taxons Lepus capensis Linnaeus, 1758

(Leporidae, Lagomorpha) in der Kapprovinz Sudafrikas

Die Hasen (Lepus) der Kap-Provinz, die bisher denUnterarten L. capensis capensis, L. c. centralis, und L. c.

grantii zugerechnet werden, wurden morphologischuntersucht. Das Ziel der Studie war, die Art L. capensis

Linnaeus, 1758 naher zu charakterisieren. Dabei konn-ten zwei distinkte Populationen unterschieden werden,die wir als verschiedene Arten ansehen, da innerhalbbeider Populationen homogene Schadel-, Zahn- undFellmerkmale existieren. Die eine Art, die wir alsL. capensis definieren, kommt bei Kapstadt in Kusten-nahe, zwischen Lambert’s Bay und Cape Agulhas, vor.Die andere Art, die wir als L. centralis definieren (mitgrantii als Synonym), ist in der zentralen und westlichenKap-Provinz verbreitet. L. capensis und L. centralis sindparapatrisch verbreitet mit einem kleinen Sympatrie-Areal in der Kontaktzone im Compagnies Drift Gebietnahe Lambert’s Bay. Differenzen im Schadelbau beiderArten sind bei L. capensis durch eine kraftigere Maxillaund robustere Zahnreihen gegeben, wahrend L. centralis

großere Ohrblasen aufweist. Der erste obere Schneide-zahn von L. centralis ist in der Regel tiefer gefurcht undhat mehr Zement als bei L. capensis, auch ist der innereLobus bei der erstgenannten Art quadratisch und beiL. capensis abgerundet. In der Fellfarbung istL. capensis durch ein ausgedehnteres weißes Bauchfellvon L. centralis unterschieden. Unsere Resultate sindrelevant fur alle weiteren taxonomischen Untersuch-ungen an Hasen der Alten Welt, sofern Populationenvon L. capensis betroffen sind.

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