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ORIGINAL ARTICLE

Morphological and scanning electron microscopic studies

of the tongue of the Egyptian fruit bat (Rousettus aegyptiacus)

and their lingual adaptation for its feeding habits

Mohamed M. A. Abumandour & Raafat M. A. El-Bakary

Accepted: 2 May 2013# Springer Science+Business Media Dordrecht 2013

Abstract This study was carried out on the tongues of 12adult normal healthy Egyptian fruit bats of both sexes. The

tongue is protrusible, elongated flat with a rounded apex andits wide and thickness increase gradually toward the lingualroot. There are four types of lingual papillae; two mechanicaland two gustatory. The tongue divided into three parts (ante-rior, middle and posterior), each part subdivides into threeregions; two lateral regions and median region, in addition tothe lingual apex to the anterior region. The lingual papillaeclose to the median region of the tongue were posteriorlydirected toward the pharynx, while theses present on thelateral regions of the tongue are directed medioposteriorly.There are sex subtypes of the filiform papillae; three on theanterior part (small, conical and giant), two on the middle part

(cornflower and leaf-like papillae) while the posterior partcontain rosette shape filiform papillae, in addition to transi-tional papillae and conical papillae. Two gustatory papillaerepresented by; small number of fungiform papillae whichscattered among the filiform papillae on lingual apex andtwo lateral regions of the anterior and middle part of tongue,while the three circumvallate papillae on the posterior partwere arranged in a triangle form.

Keywords Lingual papillae . Egyptian fruit bat. Scanningelectron microscope (SEM)

Introduction

The bat in the second largest mammalian order (Altringham etal. 1996), has an arboreal character (Ogunbiyi and Okon

1976). The bats are the only mammals have ability of flight,in which the anterior member transform into wings (Wilson

and Reeder 1993). Bats are belonged to order Chiroptera,suborder Me g a c h iro p te ra , f am i l y P te ro p o d id a e .

Pteropodidae are feed on fruit, flowers, nectar and pollen,and have Rousettus genus, which have only one species inEgypt called Rousettus aegyptiacus (Altringham et al. 1996).

The tongue is a taste organ in the buccal cavity and withits species-specific lingual papillae on the dorsal surface

plays important role in food intake, digestion in many mam-mals (Iwasaki 2002; Pastor et al. 1993).

In the end, there is a true question, is there a relationshipbetween the morphological structures of the tongue, feedinghabits, geographic position and kinds of the feed taken. This

research was conducted to shows morphostructural studiesof the lingual papillae in the Egyptian fruit bats fed asherbivorous animals and if there is adapted with the geo-graphical distribution. Thus, the results were discussed andcompared with those reported by the literature.

Material and methods

Samples

This study was carried out on the tongues of 12 adult normal

healthy Egyptian fruit bats of both sexes which collect fromfruit farms and old houses near from it from Edfina, Rashid,and Behera Governorate, Egypt.

For gross morphology

Seven bats of both sexes were euthanized to demonstrate thegross morphological features. Two bats were used as freshand five were formalized. The oral cavity was opened; thespecimens were then fixed in 10 % formalin.

M. M. A. Abumandour (*) : R. M. A. El-BakaryAnatomy and Embryology Department, Faculty of VeterinaryMedicine, Alexandria University, Rashid,Edfina, Behera, Post Box: 22785, Egypte-mail: [email protected]

Vet Res Commun

DOI 10.1007/s11259-013-9567-9


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For scanning electron microscopy

A tongue of the five adult bats of both sexes, fixed in (2 %formaldehyde, 1.25 % glutaraldehyde in 0.1 M sodiumcacodylate buffer, pH 7.2) at 4 C. Once fixed, the sampleswere washed in 0.1 M sodium cacodylate containing 5 %sucrose, processed through tannic acid, and finally

dehydrated in increasing concentrations of ethanol (15 mineach in 50, 70, 80, 90, 95 and 100 % ethanol). The sampleswere then critical point dried in carbon dioxide, attached tostubs with colloidal carbon and coated with gold palladiumin a sputtering device. Specimens were examined and

photographed with a JEOL scanning electron microscopeoperating at 15 KV, at the faculty of science, AlexandriaUniversity.

Results

Gross anatomical studies of the tongue and its lingualpapillae

Macroscopically, the tongue is protrusible, elongated flatwith a rounded apex (Figs. 1 and 2b) and lies in the floorof the oral cavity and connected ventrally by the frenulumlinguae (Fig. 2c: 9). The tongue is about 30.25 cm long,0.4 cm in wide at lingual apex, 1 cm in middle part and1.4 cm at the level of circumvallate papillae while thethickness increase gradually toward the pharynx, in whichreach to 0.1 cm at lingual apex, 0.5 cm at the level anterior

part of frenulum linguae, 0.7 cm at middle of intermolar

tubercle and 0.8 cm at the level of circumvallate papillae.The tongue divided into three parts; anterior (free part),middle (l ingual body) and posterior (l ingual root)(Fig. 2a), in which each part is divided into three regions;median and two lateral part regions (Fig. 2a, b and d). Thetongue has four surface; dorsal, ventral and two lateralsurfaces. Tongue extended to out from the limit of the floorof the oral cavity by 0.5 cm (Fig. 2b).

The dorsal surface of anterior part makes up nearly 2/3 ofthe tongue length, which reaches to 1.82 cm in length(Fig. 2b), while the ventral surface of tongue is attached tothe sublingual region by the lingual frenulum of 3 ml thick-

ness which present at the beginning of the anterior end oftongue leaving long free end of tongue, which facilitatesconsiderable freedom of movement, (Fig. 2c: 9). The dorsalsurface of middle part has a shallow intermolar tubercle (notclear in most cases) (Figs. 1 and 2), located close to the

posterior half area of the tongue and reach to 0.8 cm inlength. The dorsal surface of posterior part (lingual root) is awide and short region, which reach to 0.4 cm long and1.4 cm in wide and characterized by the presence of threecircumvallate papillae (Figs. 1, 2, and 3).

Scanning electron microscope of the dorsal surfaceof tongue and its lingual papillae

Four types of lingual papillae; two mechanical (filiform andconical) and two gustatory (fungiform and circumvallate)were recognized, in which the shape, size, number, distri-

bution, nomenclature and orientation of these lingual papil-lae are region specific according and in relation to feedinghabits and food types.

Anterior dorsal part

The anterior part subdivided into four region in form ofU-shape (Fig. 2a and b); lingual apex, two lateral re-gions and median region; lingual apex contain posteri-orly directed small filiform and fungiform papillae, thetwo lateral regions contain medioposteriorly directed

papillae (conical filiform and fungiform), while the me-

dian region contain posteriorly directed giant filiformpapillae only (Fig. 4a).

SEM figures of filiform papillae

The filiform papillae were observed throughout thewhole tongue, in which their shape, size, number andorientation of the papillae itself and its processes variedaccording to their location within the tongue (regionspecific) in relation to the feeding habits and types of

Fig. 1 Dorsal view of the Egyptian fruit bat tongue; 1- lingual apex, 2-intermolar tubercle, 3- hard palate, 4- soft palate, 5- upper canine teeth,6- lower canine teeth

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food particles. The ones close to the middle part oftongue were posteriorly directed toward the pharynxand the base of tongue, while theses present on thelateral edge of tongue are directed medioposteriorly tohelp in the collection and gathering of food particles inthe middle part region of posterior part of tongue thento pharynx (Fig. 6).

There are three subtypes of filiform papillae on the ante-

rior part of tongue; small, conical and giant filiform papillae(Fig. 4a, b and d):-

a- Small filiform papillae:The papillae on dorsal surface of lingual apex and the

tip of anterior part of tongue were small and have differentshape; round or rectangular and its dorsal surface havemicrotubercles and microgrooves, these small filiform

papillae have posterior directed several pointed processesoriginated from all anterior, posterior and lateral margin of

papillae, this processes bearing microtubercles and

microridges and terminated posteriorly by one or twoposterior processes (Figs. 4ac, 6a and 8).

b- Conical filiform papillae:The filiform papillae on dorsal surface of two lateral

regions of the anterior part of tongue were conical-shape, these papillae having posteriorly directed several

pointed processes originated from all margins of thepapillae, in which these originated from anterior andposterior margin are long and large than theses origi-nated from lateral margin which appear as serrated edgewhile the anterior and posterior processes are ended by

one or two terminal processes (Figs. 4d and 6d).c- Giant filiform papillae (trifid or tridentate)

The filiform papillae on dorsal surface of median re-gion of the anterior part of tongue (Fig. 4a) were overlapon each other and about 0.81 cm in long and 0.3 cm inwide. These papillae having a wide rectangular bodywhich have 1015 small posteriorly directed anterior

processes, in which each process is small triangular inshape and terminated by only one main process. The bodyterminated posteriorly by three large posteriorly directed

Fig. 2 a and b, d: dorsal view, c; ventrolateral view of the Egyptianfruit bat tongue: A- anterior part of tongue, B-lingual body (intermolartubercle), c- posterior part of tongue, 1-lingual apex, 2- median regionof anterior part of tongue (region of giant filiform papillae), 3- lateralregion of anterior part of tongue, 4-median region of intermolar tuber-cle, 5- lateral region of intermolar tubercle, 6-circumvallate papillae, 7-

median region of posterior part of tongue, 8- lateral region of posteriorpart of tongue, 9- lingual frenulum, 10- ventral surface of tongue

F ig . 3 a Anterior part of tongue; pur ple ar r ow refers tomedioposteriorly directed conical filiform papillae with few fungiform

papillae in the lateral region, while the pink arrow refers to posteriorlydirected giant filiform papillae. b Middle part of tongue; black arrowrefers to medioposteriorly directed cornflower filiform papillae in thelateral region, while green arrow refers to posteriorly directed leaf-likefiliform papillae in the median region. c Posterior part of tongue; redarrow refers to medioposteriorly and medioanterior directed conical

papillae (take V-shaped manner of distributed) in the anterior part of thelateral region, yellow arrow refers to medioposterior directed conical

papillae in posterior part of the lateral region, while blue arrow refers toposteriorly directed rosette-shape filiform papillae in the median region

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posterior processes so called trifid or tridentates papillae(Figs. 4a and 6b: gf). Each one of the trifid posterior

processes in anterior half of median region of anteriorpart of tongue is usually equal in wide until their endsby main posterior process and 2 or 3 accessory process(Fig. 5b and c), while that each one of trifid posterior

processes in posterior half of median region is triangularin shape ending with only main posterior process whichhave ill-developed microridges if compared by theses

present on the trifid posterior processes of giant filiform

papillae in anterior half of median region of anterior partof tongue (Fig. 5d)

SEM figures of fungiform papillae

The very small number of fungiform papillae are scattered

among the filiform papillae on lingual apex (small filiformpapillae) (Fig. 8) and the lateral edges of both anterior (conicalfiliform papillae) and middle part of tongue (cornflower filiform

papillae) (Figs. 4a and 9b), while completely abscent in themedian region of both anterior (region of giant filiform papillae)and middle part (region of leaf-like filiform papillae) of tongue.The fungiform papillae are two types according to its shape;small rectangular fungiform papillae, which present among thefiliform papillae on lingual apex and lateral edges of anterior partof tongue (Fig. 4d and 6d), while the second type is the largeround fungiform papillae, which present among the filiform

papillae on lateral edges of middle part of tongue and character-

ized by its dorsal surface is concave (Fig. 4e). Small rounddepressions on the surface of fungiform papillae for taste buds(2 or 3) and there are many microgrooves,elevated part in whichthe pore of taste bud is in micro-depressed part (Figs. 7fand 8).

Middle part of the tongue

It contains a shallow intermolar tubercle. The middle partsubdivided into three region (Figs. 2a, d and 3c); two lateralregions and median region; the two lateral regions containmedioposteriorly directed papillae (cornflower filiform andsmall number of fungiform papillae), while the median region

contain posteriorly directed leaf-like filiform papillae only.

SEM figures of filiform papillae

a- cornflower filiform papillaeThe filiform papillae on dorsal surface of two lateral

regions of middle part of tongue take the shape of thecornflower (Fig. 4e: ff), theses papillae were characterized

by its two lateral edge was bent on the dorsal surface ofpapillae and ended by small posteriorly directed processesforming a central groove, while the papillae ended by

posteriorly directed round end with a secondary one tothree posterior processes (Fig. 6c: ff). Some of these

papillae were orientated medioposteriorly and some orien-tated medially, in which the papillae were terminated byround end or by main processes and sometimes associatedwith one to three accessory processes.

b- leaf-like filiform papillaeThe filiform papillae on the dorsal surface of median

region of middle part of tongue, which take the shape of theleaf-like (Fig. 6e: if) (Fig. 9a); theses papillae ended with

posteriorly directed main posterior process and posteriorlydirected 2 to 4 accessory lateral processes (Fig. 7a: if).

Fig. 4 SEM photographs of the dorsal surface of the anterior part ofthe Egyptian fruit bat tongue; slide (a) to show; round lingual apex (1),median region of anterior part (2), lateral region of anterior part (3),small filiform papillae (white arrow), conical filiform papillae (red

star), giant filiform papillae (gf), fungiform papillae (blue head arrow).Slide (b) is high magnification of lingual apex to show; small filiform

papillae (sf) with many lateral edges posteriorly directed small pro-cesses. Slide (c); high magnification of small filiform papillae (sf) withmany lateral edges posteriorly directed small processes (p) which haveon its surface microridges (white arrow) and microtubercles (black

star), while the surface of the small filiform papillae have micro-grooves (blue arrow) and microtubercles (black star). Slide (d) is highmagnification of lateral region of anterior part of tongue to show;conical papillae (cf) with many lateral edges posteriorly directed small

processes (black arrow), very little amount of rectangular fungiformpapillae (fu) distributed among conical papillae (cf). Slide (e) is highmagnification of lateral region of middle part of tongue to show; mediallyand posterior directed cornflower filiform papillae (ff) with laterally(black star) and posterior (white arrow) posteriorly directed small pro-cesses, oval shaped fungiform papillae (fu) with concave surface

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Posterior part of the tongue

The posterior part of tongue subdivided into three region;two lateral regions and median region; the two lateral re-

gions contain medio-anterior directed conical papillae inanterior part of lateral region while have medioposteriorlydirected conical papillae in posterior part of lateral region,so the distribution of papillae on the two lateral regions takethe V-shape manner (Figs. 2d and 3c). The median regioncontains posteriorly directed rosette-shape filiform papillae.The surface of lingual root was relatively flat with someopenings for the posterior salivary glands.

SEM figures of papillae

a- rosette shape filiform papillaeThe filiform papillae on the dorsal surface of median

region of posterior part of tongue take the rosette-shape(Figs. 6fand 7d: rf); theses papillae have round base anditsapex ended with posteriorly directed many small posterior

processes forming a basket-like appearance, whichsurrounded by this many small posterior processes forminga basket-like shape (Figs. 6e and 7b: rf).

b- Transitional papillaeThese papillae represent a transitional stage, which

present between the rosette shape filiform and conicalpapillae in the two lateral region of posterior part near

the median region, which take the tongue shape withcentral groove and posterior pointed end. These papillaeoverlap each other and orientated medioposteriorly(Figs. 6g and 7c: tp).

c- Conical papillae

The papillae on the dorsal surface of two lateral regionof posterior part of tongue (laterally to transitional papillae)take the conical shape (Fig. 6e: cp); theses papillae haveround base with some have transverse direction toward themidline of tongue and some have medioposteriorly direct-ed pointed end, which surround the three circumvallate

papillae laterally and posteriorly (Fig. 7d and e: cp).d- Circumvallate papillae

The three circumvallate papillae were elongatedovoid in shape and present on the lingual root of poste-rior part of tongue and have irregular dorsal surface.These papillae arranged in a triangle form; with apex

directed toward the pharynx and consists of one medianposition papillae, while the base were anterior and con-sists of two laterally position papillae (Figs. 2d and 3c).These papillae consisted of ovoid central bulbsurrounded by a continuous deep groove (Fig. 7d ande: V), the opposite side of which is surrounded by onelayer of 2 segmented circular bad (anterior and posterior

part) while from the lateral side there is one conicalpapillae and medially there is one rosette shape filiformpapillae (Fig. 7e: b). On the surface of papillae there are

Fig. 5 SEM photographs of the dorsal surface of the median region ofthe anterior part of the Egyptian fruit bat tongue; slide ( a) to show;giant filiform papillae (gf) with trifid posterior processes (black star),and many anterior small processes (black arrow), small filiform papil-lae (sf). Slide (b) magnification of the median region of the anterior

part to show; giant filiform papillae with trifid posterior processes(black star) on it microridges (white arrow). Slide (c) magnificationof one of the trifid posterior processes of giant filiform papillae inanterior part of median region to show; that the each one of the trifid

posterior processes is usually equal in wide until their end by mainposterior process (p) and 2 or 3 accessory process (black arrow), whitearrow show the microridges. Slide (d) magnification of the trifid

posterior processes of giant filiform papillae in posterior part of medianregion to show; that the each one of the trifid posterior processes istriangular in shape ending with only main posterior process (r) whichhave ill-developed microridges (red arrow), blue arrow show anterior

processes

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clear micro-elevation and micro-depression area andnumerous taste buds were located in the epithelium of

papillae (Fig. 7g).

Discussion

The elongated flat surface tongue with long free anteriorpart, in addition to protrusible character of tongue with thicklingual frenulum; all this facilitating the movement oftongue while swiping the extracts of fruit pulp, agree with(Birt et al. 1997; Mqokeli and Downs 2013).

The lingual prominence presents in many animals; bankvole (Jackowiak and Godynicki 2005), rabbits (Nonaka et al.

2008) and most artiodactylas (most are grass-eating) (Zhengand Kobayashi 2006). Carnivores and Pigs (omnivorousartiodactylas) not have a lingual prominence (Emura et al.2006; Kumar and Bate 2004). While (Pastor et al. 1993) incommon European bat reported that there is a prominent ofintermolar tubercle, but (Jackowiak et al. 2009) agree with ourobservation, that no typically intermolar tubercle. The lingualmedian sulcus is a characteristic feature in some mammals;(Jackowiak and Godynicki 2005; Nasr et al. 2012). But ourstudy agree with (Emura et al. 2001; Jackowiak et al. 2009;

Gregorin 2003; Mqokeli and Downs 2013), that there is nolingual median sulcus in all bats.

Lingual papillae are species-specific (differing in shape,size, number, orientation and distribution) among mammalianspecies, so these differences may depend on dissimilarities indiet, feeding habits and handling of food in mouth (Emura et al.2002), similar in Egyptian fruit bat, this confirm by presence offour types of lingual papillae; two mechanical and two gusta-tory as in; bats (Selim et al. 2008; Abayomi et al. 2009); othermammals have four lingual papillae; one mechanical (filiform)and three gustatory as rats (Nasr et al. 2012), bank vole(Jackowiak and Godynicki 2005), dog and fox (Emura et al.2006). However, three types of papillae (filiform, fungiformand circumvallate) as in; bats (Pastor et al. 1993; Emura et al.

2001; Park and Lee 2009a; Masuko et al. 2007), however,there are two types only of papillae (filiform and fungiform)in hematophagous bats (Masuko et al. 2007).

The morphological structure of filiform papillae differsfrom; simple in rodents to compound structure in artiodactyls.The filiform papillae have many divisions, such as classifiedas seven group in bat by (Park and Hall 1951), our study and(Kobayashi and Shimamura 1982) make six group. Fivegroup in; lesser dog-faced fruit and nectarivorous bats (Emuraet al. 2001; Masuko et al. 2007). Four groups in; frugivorous

Fig. 6 SEM photograph of the dorsal surface of the Egyptian fruit battongue to show different types of filiform papillae (slide a to g) andconical papillae (slide e); small filiform papillae (sf) on lingual apex,giant filiform papillae (gf) on median region of anterior part, cornflow-er filiform papillae (ff) on lateral edge of middle part of tongue with

bent processed lateral edge (e) forming a dorsal central groove (g),fungiform papillae (fu), conical filiform papillae (cf) on lateral edge of

anterior part of tongue, leaf-like filiform papillae (lf) on median regionof middle part, rosette shape filiform papillae (rf) on median region of

posterior part, transitional conical filiform papillae (tp) with centralgroove (white arrow), conical papillae (cp) on lateral edge of posterior

part and posterior to circumvallate papillae, layer of bad (b) aroundcircumvallate papillae (V)

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bats (Masuko et al. 2007; Mqokeli and Downs 2013). Whilethree groups in; bats (Jackowiak et al. 2009; Park and Lee2009a; Pastor et al. 1993). Functionally in Egyptian fruit bat,

the numerous filiform papillae groups, suggests the role offiliform papillae in food and liquid transport toward pharynx,in which this papillae increase the adhesion of food to thesurface of tongue; because of eating habit during flight, agreewith general information noted by (Ojima et al. 2000).

The variation in filiform morphology may be related tomasticatory methods and dietary habits, as (Yoshimura et al.2009), this confirmed in Egyptian fruit bat, small filiform

papillae with its several pointed posterior directed smallprocesses [as in (Jackowiak and Godynicki 2004)], and

microtubercles, microridges and microgrooves with the con-ical filiform papillae on lateral sides of anterior part oftongue help in catching and fixing of food materials during

the flight. while the giant filiform papillae with its posteri-orly directed small anterior and large trifid processes inmedian region of anterior part of tongue help in posteriordirection of the catched food materials with the anteriordirected canine; so become clear that these filiform papillaeon the anterior part of tongue compensate the absence ofupper and lower incisive teeth which leaving the space

between the canine teeth, this space is adapted for allowingthe elongated tongue to move freely when feeding, this isagree with nectar-feeding bat (Mqokeli and Downs 2013).

Fig. 7 SEM photographs of the dorsal surface of the posterior part ofthe Egyptian fruit bat tongue; slide (a, b, c) to show three types offiliform papillae; leaf-like filiform papillae (lf) ended with main pos-terior process (white star) and 2 to 4 accessory lateral processes (whitearrow) on median region of middle part; rosette shape filiform papillae(rf) with many small posterior processes (p) forming a basket-like (c)on median region of posterior part, transitional papillae (tp) between(filiform and conical papillae) with central groove (black arrow). Slide(d, e) to show; rosette shaped filiform papillae (rf); conical papillae

(cp) surrounding the circumvallate papillae (V) which surrounded byone layer of 2 segmented circular bad (b) leaving circular groovearound the circumvallate papillae (V). slide (f, g) to show the taste

bud on the surface of fungiform papillae (F) and circumvallate papillae(G); in which on fungiform papillae there are microgrooves (whitearrow), elevated part (blue star) in which the pore of taste bud inmicro-depressed area while on surface of circumvallate papillae thereare clear micro-elevation (red star) and micro-depression (white star)in which the pore of taste bud in the micro-elevated area

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The greater protrusible tongue with the retention of caninesof S. australis; suggest that it may be more efficient than

Pteropus species when collecting nectar from flowers, (Birtet al. 1997).

The differences in the filiform morphology may be relat-ed to type of food eaten, which varied according to geo-

graphical distribution; this clear in same animal; as in thisstudy of Egyptian fruit bat (captured from Egypt), there aredifference in shape, size, orientation and number of thelingual papillae; there are six groups of filiform papillae inaddition to transitional papillae, while (Jackowiak et al.

2009) in Egyptian fruit bat (captured from Poland), notedthat there are three types only. or this variation may reflectthe different in subspecies as; certain P. poliocephalusadapted to a nectar diet and others to fruit diet (Eby 1991).This is clear in case of, P. scapulatus having long pointed,and dense giant papillae on a large area of anterior part oftongue, suggest that P. scapulatus prefers a diet of nectar

(Eby 1995), with its migratory behaviour to area with no orlittle flowering, so P. scapulatus consume fruit (Eby 1995),so some variation in giant papillae morphology.

The lingual adaptation of in Egyptian fruit bat; firstly, thetongue tip not have giant filiform papillae and instead a

blunt small filiform papillae, in which this distributionmay have important role in feeding habits, this confirmed

by (Hall et al. 1995), while (Paton and Collins 1989) innectar-feeding bats, noted that the functions of giant filiform

papillae on the tongue tip to increase the surface area tocollect nectar. The another adaptation, the giant filiform

papillae were rough to touch to help in power catching by

piercing the skin of soft fruits to consume fruit and press itbetween the tongue the ridged hard palate to release thejuices (Bonaccorso and Gush 1987). Also, among the lin-gual papillae morphology the orientation; in Egyptian fruit

bat, cornflower filiform papillae on two lateral regions ofmiddle part; some orientated medioposteriorly and someorientated medially while, in lateral region of posterior part;having the medio-anterior directed conical papillae in ante-rior part and medioposteriorly directed conical papillae in

posterior part, while (Pastor et al. 1993) in common Euro-pean bat, at junction of posterior and intermolar eminence, atuft of filiform papillae was oriented anteriorly in opposition

to all the others .There are three classifications of fungiform papillae;

according to its function, distribution and shape; Accordingto its function there are three types; first (gustatory) type; asin Egyptian fruit bat and (Kilinc et al. 2010), second (mixed)

Fig. 9 Transverse section ofmiddle (b) and posterior part (a)of Egyptian fruit bat tongueshowing the leaf-like filiform

papillae (lf), fungiform papillae(fu) showing taste buds (blackhead arrow) (Haematoxylin

and Eosin100)

Fig. 8 Transverse section of anterior part of Egyptian fruit bat tongueshowing the small conical filiform papillae (sf), fungiform papillae (fu)showing taste buds (black head arrow) (Haematoxylin and Eosin100)

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type, means some papilla having taste buds and some nothave any taste buds as in; Australian Megachiroptera (Birt etal. 1997), while third (mechanical) type is (no taste buds) asin; goat (Kurtul and Atalgin 2008). Second classificationaccording to distribution, there is four types according to(Chung and Kwun 1977); first type, distribution on thewhole area of dorsal surface as (Park and Lee 2009a),

second type, the distribution on the side edges, tip and rearend section as in S. species (Park and Lee 2009b). Thirdtype, the distribution only in the tip and side edge areas asin; our study and (Emura et al. 2001), rat (Nasr et al. 2012),while fourth type, indicates a center and tip distribution as in

bank vole, (Jackowiak and Godynicki 2005). Third classifi-cation according to shape; in Egyptian fruit bat, have twosubtypes; rectangular (on anterior part), round (on middle

part), while in other bats there is only one type, elliptical orcircular (Park and Lee 2009a). Dome-shape (Emura et al.2006), round; (Emura et al. 2007), Mushroom-shaped (Nasr2012; Kurtul and Atalgin 2008), fungus-shaped in; rat (Din

et al. 1995).The Egyptian fruit bat considered as herbivorous animal,

so fungiform and three circumvallate papillae and their tastebuds compensated the absence of foliate papillae, and con-firm that the increase in the number of circumvallate papil-lae corresponds to increase sensitivity in the taste, while thespecial distribution of fungiform confirm to increase taste

perception in this area before food is swallowed.The number of circumvallate papillae in bats is known to

depend on the types of food species consume; none in blooddrinking Desmodus rotundus (Greenbaum and Phillips 1974)and hematophagous bats (Masuko et al. 2007), two papillae as

in common European bat (Pastor et al. 1993); Korean greaterhorseshoe bat (Son et al. 2000); lesser dog-faced fruit andFlying Fox (Emura et al. 2001, 2002); free-tailed bat(Gregorin 2003); Korean long-fingered bats (Park and Lee2009a). Three papillae; in Egyptian fruit bat, in our study and(Jackowiak et al. 2009); frungivorous bats (Emura et al. 2001,2002), S. australis and Pteropus (Birt et al. 1997); E.wahlbergi (Mqokeli and Downs 2013). While four papillaein long-nosed bat and in frugivorous and nectarivorous bats,(Masuko et al. 2007); N. Robinsoni by (Birt et al. 1997).

Our study noted that, the oval three circumvallate papil-lae were consisted of ovoid central bulb surrounded by a

continuous deep groove, as (Ciuccio et al. 2010). (Nasr et al.2012) in rat, circumvallate papillae were large circular,dough-nut-shaped papillae, resembled those of bats as;

Pteropus vampyrus and Myotis macrodactylus (Emura etal. 2002). The number of pad layers around the circumval-late papillae varied among species; in primates (Kobayashiet al. 2004) had only one layer of pads, while in Egyptianfruit bat study, there is one layer of 2 segmented circular

bad. But, circumvallate papillae of S. caecutiens had twodistinct layers of pad, (Park and Lee 2009b).

Our study revealed that, dorsal surface of circumvallatepapillae is somewhat irregular, similar to (Emura et al. 2006)in dog and fox, while it is smooth; in fox (Jackowiak andGodynicki 2004). Our study agrees with (Mqokeli andDowns 2013; Jackowiak et al. 2009) in that the distributionof three circumvallate papillae in the form of a triangle.

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