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43 44 bule ranged 15–20 from µm. The tubules open tu single a of lumen the of µm. diameter The 100–300 from ranged lengths 1C). tubule The (Fig. 1B), while the distal part was sinuous (Fig. straight was tubule every of part proximal The the and anterior regionsposterior (Fig. 1A, D). (Fig. cloaca 1B, in detected wereC), few while the of region central the in located were bules the cross-sectioned tubules and openings of tu of Most part. body left and right the between unevenly arranged were tubules The matheca. sper the in tubules 40–45 possessed females studied Both epithelium. desquamation with A-D) connected mucus with filled typically 1 was that (Fig. lumen round a revealed bules F is (Fig. orifice absent inthecloacal it 1A-D). and tube cloacal the in only occurs fold present along the cloaca, while the mid-ventral is fold mid-dorsal The 1C). (Fig. partition of ventral) project into the lumen and form a kind mid- and (mid-dorsal folds conspicuous two tube Y-shaped is (Fig.cloacal men the 1B).In more caudal part of the cloacal chamber the lu cross-sectionthe (Fig.in like 1D), the in while (Fig.cloaca 1A-D). bules opening independently to the walls of the of a simple type, consisting of spermathecal tu the spermathecal tubules. The spermatheca was F regionthe entirewere cloacal prepared. dividual, approximately 100 slides representing Nikon COOLPIX P6000 camera. For each in a using photographed were present. slides The was tubule the which in sections the counting byestimated was tubules the of µm) (in length The cloaca. the into openings of number the Anna Dymek, Jakub Dymek

emale spermatheca emale cloaca morphology N r 2(20)/2018 N rs-etos f h semtea tu spermatheca the of Cross-sections isstar- tube cloacal the of lumen The in sperm lacked females examined Both Results auki P rzyrod nicze ------cytoplasm (Fig.cytoplasm 2A). abundant with nuclei small possessed cloaca did not possess cilia. The epithelium cells of the cells cytoplasm.Those little and nuclei shaped irregular- large had epithelium spermatheca (Fig. part) posterior most the 2A).the of Cells and anterior most (the stratified and part) tral wereaca lined with columnar epithelium (cen cuboidal epithelium, while the walls of the clo directions. both in arranged were cloaca the of part tral arranged towards the back. Tubules in the cen were part caudal the in tubules while rection, di forward the in arranged were cloaca the of lumen (Fig. 1B, C). Tubules in the cephalic cloacal part the to perpendicular located were ings prahc o a ipe ye cnitn of consisting type, simple a of terdigitata spermatheca mandrina e.g. matheca.Salamandridae of species other in As anatomical description of (Fig.melanin) were observed 2C). of granules with (melanophores cells pigment and vessels blood numeroustissue, connective the In (Fig.2C). fibers collagen of consisting tissue connective with filled were spermatheca carbohydrates (PAS-) (Fig. 2B). neutral for negative and (AB+) aminoglycans mucus stained positive for carboxylated glycos uniform.was The walls cloaca inner the lining the epithelium of activity secretory The 2B). (Fig. cells epithelium desquamation the with connected was Additionally, mucus 2B).(Fig. cloaca the of lumen the in clusters formed and (Fig. epithelium 2A). the Mucus covered of the walls basis of the the cloaca in observed were cytoplasm large and nuclei small with cells lar istio vulgaris Lissotriton T The spermatheca tubules were lined with The spaces between the tubules ofthe the tubules between spaces The glandu Additionally,mucus big,round he aa ahrd rvds h first the provides gathered data Discussion , hs pce possesses species this ), , S L. montandoni L. tr . helveticus L. . 42-49 S , sper ala ------

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45 46 to other related species studied to date. In the date.In to studied species related other to SEVER, 1988; BRIZZI AL., 1989, 2004). tube and the cloacal chamber (VERREL AND cloacal roofthe into posterior the of pendently inde opening tubules spermathecal numerous (A): 20µm; (B): 50µm; (C): 40µm. cell; lu: lumen of the spermathecal tubules; mel: melanophore; mu: mucus; s: spermatheca. Scale bar: bv: blood vessel; cf: collagen fibers; co: connective tissue; ct:cloacal tube; e: epithelium; gc: glandular PASstaining.and Abbreviations: blue Alcian (B):staining; trichrome matheca.(A,Mallory`s C): spermatheca;of bules (B): cloaca;(C):the of Secretions sper of tubules Connectivethe and tissue F Anna Dymek, Jakub Dymek

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and and and L. montandoni L. S L. helveticus L. L. vulgaris. L. iegel L. vulgaris L. L. vulgaris vulgaris L. S eae cnitd of consisted females auki tr . Female vent glands vent Female. L. montandoni L. . 42-49 P , 2006). Thus, the L. helveticus L. (OSIKOWSKI L. vulgaris. L. rzyrod (SEVER ET (SEVER than between (VERELL L. vul L. nicze Se are fe ------

47 48 ademic Press. UK. Selection. Sexual Competitionand Sperm ni ( zone hybrid newt a in variation traits secondary male and for University A Jagiellonian of Society dents Stu PhD by supported partly was work The slides. glass making in help her for Podkowa useful for comments on and the D.manuscript byas reported SEVER (2002). females, polyandrous of progeny in advantage male last the to lead could phenomenon This fromthe cloaca, the last male. first meet sperm through oviduct the from passing Eggs,when glands. anterior more filling mating next the thefill glands, most posterior from with sperm may insemination first the from uation, sperm males. various divide from sperm to opportunity an provide spermatheca type H sperm of competitionfemale choice occurrence (BI and cryptic the for conditions ate R MØLL AND (BI newts in polyandry common 1997 ( newts smooth the and Carpathian the between barrier iological mucus composition could act as a natural phys B Bi J. Ba Anna Dymek, Jakub Dymek

r AFI nna Dymek. 2003.markers,DN Nuclear mitochondrial ). MolecularEcology. 12, 1913–1930. EAD N r zi . D R., izzi b kh ). r k . Szy W., ik , 2001; OSIK 2001; ŃSKI, 2(20)/2018 N W The presence offemale sperm storage and A e D MØLL AND a cknowledgments e would like to thank to like would d Triturus vulgaris Triturus References . . M R., T. ER, 1998; OSIK ER,1998; e l mur f i Mi no ER, 1998). The simple The 1998).ER, auki J M, Ra M., J. a c ø . Call G., , 2003) cre 2003) OWSKI, aa E al., ET halak ll P x er rzyrod OWSKI T. montando T. A.Osikowski n such In a such sit

A R P. . KH f on nicze i 1998. ńs AND EAD R C. i A K ki ki c A ------lution. 51, 2045–2050. montandoni T. species, newt two tween helveticus dae). A Triturus newt, palmate the of anatomy Cloacal 2004. 310–322. Salamandridae). phibia: salamander, 1989. Female cloacal anatomy in the spectacled Salamandridae), in the breeding season. breeding the in Salamandridae), newt, ian 2013 Science. 29(6), 390–395. ( ( newt smooth the species, newt closely-related two in pheromones courtship for preferences logica. Krakow. 55, 109–114. alpestris the in insemination ter sis. Kraków. Poland. Triturus).genus The of PhD newts female in Triturus rodzaju z traszek samic u poliandrii kwencje of Linnean Society. Journal 67(3), 343–352. Biological Salamandridae). dela, vulgaris species, newt two between isolation Mi 1 J. Mi B O O O O . montandoni . vulgaris L issotriton L r s s s s zi . D R., izzi c c ik ik ik ik 9. et fr eul noptblt be incompatibility sexual for Tests 997. . Cloacal anatomy of the male Carpath male the of anatomy Cloacal . halak P., halak Ra P.,G halak ows ows ows ows mphibia-Reptilia. 25, 233–245. and (Biological consequences of polyandry , Caudata, Salamandridae). Folia bio Folia Salamandridae). Caudata,, issotriton montandoni issotriton L ki ki ki ki ki T. montandoni T. aaadia terdigitata Salamandrina : No choice mating design. Evo A e ) (Salamandridae). ) A A A l . . . , Ci ., 07 Sem rnpr af transport Sperm 2007. f (A 2012. 03 Booize konse Biologiczne 2003. ) and the newt Carpathian i r no f z ern i pii, Salamandri mphibia, lpine newt ( newt Alpine es S ńs Triturus vulgaris Triturus . Ta G., tr symmetric female female Asymmetric reooia 45, Herpetologica. ik J., Ra J., ik ki J. ki (A . 42-49 iak- mphibia, Uro mphibia, 1999. Sexual 1999.Sexual (A Zu nter Z oological oological mphibia, f zia zia Triturus Triturus Triturus (A i ńs Z G. i and oo m ki ki K ------.

WWW.NAUKOWCY.ORG.PL 2 (20)/2018 WWW.NAUKOWCY.ORG.PL 2 (20)/2018 WWW.NAUKOWCY.ORG.PL 2 (20)/2018 WWW.NAUKOWCY.ORG.PL 2 (20)/2018 WWW.NAUKOWCY.ORG.PL 2 (20)/2018 WWW.NAUKOWCY.ORG.PL 2 (20)/2018 H 82(1), 49–56. Zoologica.season.Acta breeding the after cae garis ( of the smooth in newt females Enfield, NH. Urodela.of Publishers, Phylogeny Science and 292, 165–179. Zoology. Experimental of amphibians. Journal Anatomical Records. 232, 229–244. Caudata).(Amphibia: salamanders of cloacae the of phylogeny and level.ily Herpetologica. 47, 165–193. (Amphibia: Caudata salamanders of cloacae the of phylogeny and dridae). Amphibia-Reptilia. 6, 11–22. montandoni newt, Montandon’s the of haviour 5–7. is in the smooth newt,dynamics andSociobiology. Ecology ioral 49, 145–149. ( montandoni newt Montandon’s female of cess suc reproductive increases insemination tiple Science.logical 30(9), 748–753. O S t S S S S S P P y g o l o t s i e e e e e e , in the laboratory.Herpetology.,the of in Journal 2, ec ec E C, A C., E. y s v v v v v v ik er er er er er er i i L.): o o o w . R A., A. 199 D. M.,H H M.,D. 2003. M. D. D. M. 2002. 1992 M. D. 1991. M. D. . Ultrastructure of the spermathe the of Ultrastructure .II

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Buegr (adt: Salaman (Caudata: (Boulenger) Caudata ra t h a 2. e fiń C ñ nsemination and egg laying laying egg and Insemination a a o t a p r a lli I).. Evolution at the fam lli . 2001. B. a s , F Cmaaie anatomy Comparative . oprtv anatomy Comparative Salamandridae Reproductive Biology Biology Reproductive IV i J. ki fiń emale sperm storage in storage sperm emale da h da . Salamandridae.The . n a i , U , s y y T., y T., y M 95 Sxa be Sexual 1985. ki J. 20 J. ki

a l e d o r t w e n Triturus vulgar Triturus pr storage Sperm Triturus v. vul v. Triturus Wa ( L ) ) or 01. Mul 01. ). Behav t a m r e p s n o t i r t o s s i ig Triturus Triturus Triturus i hts ar ------h

e c

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