de Bavay, John

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PAP. & PROC. ROY. Soc. TASMANIA, 1950 (15TH DECBMBER, 1951)Notes on the Female Urogenital System of Tarsipes spenserae(Marsupialia) .1ByJ. M. DE BAVAYTasmanian Museum(With 9 Figures)SUMMARYTwo specimens of the fe'male urogenital system of Tarsipcs S]JCYU)CTae aI'€' described. One of theseis parous, the other non-parous. In both examples the cul-de-sac is very long and in the paroussp€.dmen a patent connexion exists between it and the urogenital sinus. A patent connexion in thenon-parous spe'cimen is doubtful. The parous urogenital system displays a singular exam_pIe ofpostponed regression of the Wolffian duct.In 1900 the female urogenital system of Tarsipcs TostTaius (') was describedfor the first time by Hill. Hill based his description upon two females, one parousand the other probably parous, both of which, had a permanent connexion betweenthe cul-de-sac and the urogenital sinus. This represents a condition unique amongthe Phalangeridae and, with the exception of certain Macropods, among theMarsupialia generally. As this is a point of considerable significance it wasdecided to re-examine the question (").The information obtained in this way completely confirms Hill's observationsand in some respects supplements them.The literature on TaTsipcs is meagre and apparently no full descriptioill of itsstructure exists. Described originally by Gray (1842) and almost simultaneouslyby Gervais and Verr,eaux (1842) Ta,Tsipcs has, despite its structural peculiarities,received little attention from comparative anatomists. However, sufficient isknown to indicate that TaTSipes occupies a special position among the Phalangeridae,being in some ways the most specialized member of that family.The snout is long and curved and, accordingly, well adapted to searchingflowers for the pollen and nectar upon which it chiefly subsists. Its degeneratedentition, too, reflects a diet which needs little mastication, the cheek teeth beingvariable in number, haplodont in form and completely vestigial (Bensley, 1903).Parker (1890) noted several aberrant features in the skull and concluded 'in thegeneral form of the skull TOTsipcs differs from all the other Phalan-gistidae '.(l) The investigations dealt with in this paper hayE' been assisted by a grant provided equally bythe Trustees of the Commonwealth Science and Industry E,ndowme'nt Fund and the Tasmanian StateGovernment.(2) The spedfic name spenserae is now considered to have priority over rostratus.e) This was made possible through the kindness of the Director of the National Museum,Melbourne,143144 FEMALE I.:ROGENITAL SYSTEMIn its pes, Tarsipes shows the greatest modifications of any of the Phalangeridae.Bensley (1903) says "the second and third digits being greatly reduced, andalmost completely enclosed in a common integument; while the fourth digit isgreatly enlarged, and like the fifth is provided with a nail instead of a claw,as in the placental Primates. In addition to these obviously progressive characters,Tarsipes presents other peculiarities. The terminal pad of the hallux is not swollento the same extent as is usual in arboreal forms. The hallucal planter pad isshort. The second digital plantar pad is subdivided and its outer portion hasmigrated to the fourth digit".In its female urogenital system, too, Tarsipes would appear to differ from theother members of its family.Two spirit specimens may years old provided the material for this investigationand have been numbered 316 and 317 ('). Their measurements are as follows:-1\To. 316 No. 31727 mm. 24'5 fi'lm.37 mm. 36 mm.93 mm. 95 fi1m.94 mm. 95 mm.11 mm. 11 mm.15 mm. 14'5 mm.give the measurements of his specimens so aHead (snout to ear-notch)Head (D.C.L.)"Head and body (D.C.L.)TailEarPesUnfortunately Hill does notcomparison cannot be made.The pouch of No. 316 is very definite and possesses four nipples, two of themvery prominent; the other two are small and insignificant. This specimen is almostcertainly parous. The pouch of No. 317 is scarcely discernible and the nipplesquite indistinguishable. This specimen is quite definitely non-parous. These twofemales, then, nearly indentical in size differ in their sexual condition and thisdifference might be expected to be demonstrated in their respective reproductivesystems.Serial transverse sections 10,u in thickness were cut of each specimen.Despite their long immersion in spirit the histology of both specimens issurprisingly good, especially that of No. 317.While Hill's account generally is scarcely to be improved upon, for the sakeof completeness some repetition will be made here. Following is a table of thesignificant measurements of No. 316 and No. 317.No. 316 0/0 a.f· No. 317 % a·f·Total length of urogenital system 24·83 mm. 19·86 mm.Length a.f.(') - combined length of-vaginal system and urogenital sinus 15'01 mm. 11·99 mm.Length of vaginal system (') 4·43 mm. 29·51 3·11 mm. 26Length of urogenital sinus (8) 10·58 mm. 70·55 8·88 mm. 74·06Length of cloaca(') 4·86 mm. 32·38 3·63 mm. 30·27Length of urethra (10) 2·22 mm. 14·79 1·37 mm. 11·43Length of ute'rine papilla 890,u 5'93 660", 5·6(4) From specimens R12267 and R13016 respectively, belonging to the National Museum, Melbourne.(') D.C.L. = Dorsal contour length.(6) See Figure 3.(7) The vaginal length is taken as extending from the cranial extremity of the lateral vaginaeto the anterior limit of the urogenital sinus.(S) The tip of the clitoris is taken as marking the posterior extremity of the urogenital sinus.(V) The cloaca is' taken as beginning from the point of junction of rectum and urogenital sinusand extending to the' paste,rior limit of the vent.(10) The urethra is regarded as comn1encing from the point of entry of the ureters into thebladder and extending to the anterior limit of the urogenital sinus.J. M. DE BAVAY 145OvariesThese are small triangular to ovoid bodies about 1·05 mm. long by 0·93 mm.broad, situated near the mid-line, immediately dorsal to and in close contact withthe uteri.Large, apparently mature, ova are present in the ovaries of both No. 316and No. 317, but those of the latter contain a relatively greater number of verysmall ova.Fallopian TubesThese are quite short and little complicated. In No. 317 the proximal part ofthe tubes runs forward a short distance within the wall of the uterus and parallelto the uterine cavity. The lining of the Fallopian tubes is a single layer ofcubical cells with dense cytoplasm.Uteri and uterine necksThe uteri are pearshoped bodies dorso-ventrally compressed. In No. 316 theepithelium is compact and colummar with a vacuolated cytoplasm and has mostlyseparated from the endometrium. The uterine glands are long and well developed.There appears to be a physiological difference between the left and right uteri.The uterine gland cells of the left uterus have deeply staining nuclei, contrastingstrongly with those of the right uterus which are scarcely visible, and even with thoseof the histologically superior No. 317. The left uterus also contains considerablestrongly staining debris, much of it membranous and possibly foetal.The uterine epithelium of No. 317 is similar to that of No. 316 but morccubical and has remained attached to the endometrium. There is comparativelylittle debris in the uterine cavities and no differentiation between right and leftuterine glands.The uterine necks, distinguished from the uteri by the absence of uterine glands,contract gradually to run caudally within a common sheath of connective andmuscular tissue. They open upon a single very prominent uterine papilla ofconical form which projects backwards into the median vagina (figs. 3, 5) by twoapertures III No. 317 but by only one in No. 316. This later case is undoubtedly dueto a certain amount of destruction of the intervening tissue. Although notinsisted upon, it is quite possible, despite it being called so here, that this conicalprojection does not represent the uterine papillae, but rather the wall of theanterior vaginal canal. Pearson (1950) defines the cuI-dc-sac (median vagina) ashaving its anterior limits at the level of the ora uterorum. Extensions anterior tothe cul-de-sac correspond, then, to Hill's 'anterior vagina:l canals' and it is thisview which is adopted in the present case. In No. 316 the width of the lumen inthe left uterine neck is about eight times that in the right. It is probable thatthis feature and the common os uterorum in this specimen are the result offoetal passage. (fig. 5).Cui-dc-sacThis is a dorso-ventrally compressed chamber surrounding anteriorly theuterine papilla and from the sides of which arise the lateral vaginae. Its epitheliumconsists of a single layer of small cubical cells which in parts are very muchelongated and flattened.There is a complete absence of a septum in both specimens, except for avestige at the base of the uterine papilla.The cuI-dc-sac passes caudally as a wide shallow chamber which, posteriorly,lies between the lateral vaginae and the urethra, communicating, in the case ofNo. 316, by a narrow fissure with the short posterior vaginal sinus. The latter146 FEMALE UROGENITAL SYSTEMitself opens into the urogenital sinus a few sections later. It will be seen thenthat in No. 316 the cul-de-sac has become a true median vagina, thus confirmingHill's observations made fifty years earlier.In the case of No. 317, however, the posterior end of the cul-de-sac is separatedfrom the urethra by a solid plate of cells (fig. 8), but within this solid plate thereis evidence of a future perforation that will ultimately convert the cul-de-sacinto a functional median vagina having a permanent opening into the urethra.Fi..9. 1 2··;:··..,,,··,,·,,·:+·:·:·, :, .....+.3 ,..: , ):: ~ :A e, IC Pv"..:ti·cJ.E FIi'IG. l.---Diagrammatic sagitta] section showing the relations of the posterior vaginal sinus (1) andmedian vagina (2) with the' urethra (3) and urogenital sinus (4).FIG. 2.-Transverse s€'ctions illustrating successive levels as shown in fig. 1.Lateral VaginaeThe lateral vaginae arise from the cul-de-sac at about the level of the orauterorum. They curve gently caudally, converging to form a short posteriorvaginal sinus before finally joining the urogenital sinus. The posterior part ofeach lateral vagina and a portion of the posterior vaginal sinus are occluded 1Ilboth specimens, the occlusion in No. 317, however, being much greater. (figs. 4,7, 8.)Because they are arranged one on top of another, the true relationship betweenlateral vaginae, cul-de-sac and urethra is difficult to demonstrate by means ofdrawings. Figs. 1 and 2, show in diagramatic form the disposition of thecul-de-sac with respect to the urethra and lateral vaginae.In the parous specimen No. 316 the lateral vaginae present a most unusua~feature. (figs. 4, 5, 6 and 7.) From the posterior third of each lateral vagina along duct passes cranially within the walls of the lateral vaginae, anterior vaginalcanals and uterine necks to terminate as a small cyst or swelling.Each duct possesses a distinct lumen for most of its length and in its courserotates around the lateral vagina in a loose spiral. It should be noted that thisduct, undoubtedly the Wolffian duct, is absent in No. 317. The lateral vaginae ofNo. 316 have a less strongly developed muscular sheath than those of the non-parous No. 317, and present in comparison with the latter a relatively embryonicappearance.SUl\tIMARY OF RESULTS AND DISCUSSIONsalient that emerge froln this study Tarsipesby Hill, the existence in a parous female of a true median vagina, andof Wolffian duct in a female of a highlyin his doubtfully of Tarsipesconnexion between cul-de-sac sinus concluded, bycertain Macropods, that since passage was open the animalpreviously. He contended that' Tarsipes thus agrees withspecies of the Macropodinae the possession direct and, afterfirst parturition a permanent passage for the birth of the young'. Theevidence supplied by the present supports this conclusion. Thus out oftwo specimens described here, and the two described by Hill, three possess amedian vagina while the fourth shows signs of acquiring one.The long duct which runs alongside each lateral vagina in No. 316, is almostcertainly a vestige of the W olffian duct. In Inost fen1ale marsupials the W olffian ductdegenerates long before maturity is attained, but it has been found in the coursethe investigations which been proceeding in the Tasn1anian Museum fornumber of years that among the Dasyuridae a vestige of the W olffian ductpersists as a very short solid rod or hollow diverticululTI towards the posterior end ofeach lateral vagina, but never before in the course of this work has a Marsupialurogenital systen1 shown such a striking example of postponed regression ofembryonic structure. Its persistence in such a specialized animal is undoubtedlyabnormal.The posterior part of the lateral vaginae is feature nowthe different Marsupial families. I t is mentionedfor the Taslnanian Museum has indicated that it isquestion significant enough to require that a separate paper be devoted toto acknowledge the advice and encouragementpaper by Director the Taslnanian148 FEMALE UROGENITAL SYSTEMREFERENCESDENSLEY, B. A., 1903.-0n the evolution of the Australian Marsupials with remarks on therelationships of the lVlarsupials in general. Trans L£nn. Soc. Lond. (2), 9, part 3,pp. 83-217.GERVAIS, P. and VERREAUX, J.-Description du Tarsi]Jes rostrah-ls. Magas. de Zoologie Paris 1842mammif. 12 pp. 3 pI.-------June', 1842.--Tarsipes rostratu8. Froe. Zool. Soc. Lond. 1842, p.l.GRAY, March, 1842.-Tarsipes spenserae. Ann Mag. Nat. llist. IX., p. 40. ex Gervais M.S.HILL, J. P., 1900.-0n the Female Genital Organs of Tarsipes rostratus. Prof:. Linn,. Soc. N.S. W.25, pp. 523-525.PARKER, W. K., 1890.-0n the skull of Tarsipes rostratu8. Studies from the ZVIuseum of Zoology inUnivcr8itJ! College Dundee. 1 pp. 79-84.ROTENBERG, D., 1928.~Notes on the Male Generative Apparatus of Ta-rsipcs SlJcnserac. Journ. Roy.Soc. W.A. 15 pp. 9-14.EXPLANATION OF FIGURESa.f.~Combined length of the vaginal system andthe urogenital sinus.bl.-Bladder.b.s.~Blood sinus.cl.--Clitoris.conn.~Connexion between the median vagina andurogenital sinus.d.~Debris within uterine ne'ck.d.gr.~Dorsal groove in the clitoris.ect.t.-Erectile tissue.l.v.-Lateral vagina.l.l.v.-Left lateral vagina.l.ur.:-'Left ureter.?n.v.c.-Median vaginal cul-de-sac.l.ur.-Left ureter.m.'/}.c.-Median vaginal cul-de-sac.oc.l. v.-Occluded lateral vagina.oc.p.v.s.-Occluded posterior vaginal sinus.os.ut.~Common os uterorum.r.l.v.~-Right lateTal vagina.1".uT.-Right ureter.8.~Septum.s.conn.-801id connexion of posterior vaginal sinusand urethra.s.l.-Suspensory ligament of the clitoris.t.cu.-Terminal cyst of the Wolffian duct.ur.-Ureter.'urcth.-Urethra.u.g.s.-Urogenital sinus.ut.-Uterus.1d.n..~Uterine' neck.ut.lJ.~Uterine papilla.W.d.-Wolffian duct.IIIIIq.f.--__u.g.s. _---- c!. ----_----dgp'.-----___ut.n.~__os.ui.----m.v.c.~t----l.v.M'-""'"'-----w.d.EXPLANATION OF PLATEFIG. 5.-No. 316. Transverse section through 4 showing the distended right ute'rine(ut.n.) containing a certain amount debris (d.). At this level Wolffian(W.d.) runs within the floor of the anterior vaginal canal (a.v.c.). X6.-No. 316. Transverse section through fig. showing the intimate' associationlateral vagina (l.v.) with the Wolffian ducts (W.d.). The left ureteropening into the bladder (bI.).316. Transver8e section through C in fig. 4 showing fusion of the right(W.d.) with the, right lateral vagina (r.1.v.) and thE' occlusion of the(occ.l.v.). X 60317. Transverse section corresponding to a section through D inand left late'ral vaginae have fused to form a posterior vaginal sinusis largely occluded. The cul-de-sac (m.v.c.) is also partially occludedthe urethra (ureth.) by cells (s.conn.). 60.316. Transve'rse section through the clitoris showing the deep dorsal X

Notes on the female urogenital system of Tarsipes spencerae (Marsupialia)

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Notes on the female urogenital system of Tarsipes spencerae (Marsupialia)

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