Downloaded By: [Universiteit van Amsterdam] At: 12:41 19 February 2008 aooi eak ntegenus the with on Colombia, remarks from taxonomic Terricola) Tricladida, (Platyhelminthes, 1990 heterogeneous. 1962), (Hyman, ihntesbaiy rmti esetv new a perspective this From subfamily. systematics the the within in progress hindering poor, relatively orsodne .Crao soad re,Cie Artes, Tel: de Escola Carbayo, F. Correspondence: detailed the the within more knowledge However, morphological . of provided species of have descriptions characteristics gical the of definition genital papilla. the penis accessory the in in as genus) male one ridges only (glandular with the organs is Geoplaninae and descriptions, within The Colombia, original characteristics Peru. on of Panama, combination based and a by Venezuela erected mainly Rica, was genus Tobago, comprises is Costa and it genus Trinidad since from Andean the species North of and distribution Caribbean The 1990. the dubia—including nomina to and inquerendae species addition for group 1990 Kawakatsu, genera—in com- 16 is collective subfamily of This presently eyes. posed 1857 numerous and dorsal testes Stimpson, with species geoplanid includes distribution, in Geoplaninae neotropical Terricola), Tricladida: subfamily (Platyhelminthes: The Introduction muscular the Keywords: the of details to and open mgo that the canalicula subepithelial constituting 8–15 the structures and 11 in from the secretions Differences among derived contains organ. organs mgo both the reproductive each the of one, of of region a longitudinal and of mass central systems underlying consisting central the morphology, an on The unique atrium a papilla. and of male penis coat are the mgo muscle These of circular mgo. musculature or barrel-shaped organs, genital strong accessory very numerous with Colombia, provided from is species land of species new A Abstract Cie Artes, de Escola CARBAYO F. of planarian land of species new A Zoology of Journal Italian O:10.1080/11250000701693255 online DOI: 1748-5851 print/ISSN 1125-0003 ISSN nrcn er,anme fadtoa morpholo- additional of number a years, recent In + 51 0100 Fax: 30911020. 11 55 etoia,tretilfawr,sseais epaia,ml cesr eia organs genital accessory male , systematics, , terrestrial Neotropical, .chiriquii G. niseHmndds nvriaed Sa de Universidade Humanidades, e ˆncias + Pseudogeoplana 51 9300 mi:[email protected] Email: 69439070. 11 55 ac 08 51:85–95 75(1): 2008; March , Hmn 92,and 1962), (Hyman, ge n Kawakatsu, and Ogren # niseHmndds nvriaed Sa de Universidade Humanidades, e ˆncias 08Uin olgc Italiana Zoologica Unione 2008 ge and Ogren Gigantea iatamaupoi Gigantea .sandersoni G. Gigantea is Gigantea oPuo Sa Paulo, ˜o pce rmltrtr n rmtp aeilof material type from and literature from species rprduigacmr uia diinly the Additionally, lucida. camera were the a drawings Mallory/ using specified, or not prepared with When region stained method. pre-pharyngeal Cason were apparatus, the copulatory pharynx, hor- or the transverse graded Sagittal, 7–8 a andizontal paraffin. alcohol in isopropyl in dehydrated with embedded treated were ethanol, body of series the of FAA different from regions with blocks fixed Tissue alcohol. water, in boiling preserved with and and killed made was were it then live the of observations Some Medellı type of the city in the grass in locality, the from collected was holotype The methods and Material chiriquii G. of 1962), material type (Hyman, the conducted in observations author the new by from and from literature Geoplaninae the in known The those with presented. compared species new are is the of mgo, genus organs, genital accessory the male of revision taxonomic of species Puhe 99 exmndhr hwta h eu is genus the that show here reexamined 1949) (Prudhoe, p o. sdsrbdhri.Tepnspplao h new the of papilla penis The herein. described is nov., sp. oPuo v rid eto10,08800Sa 03828-000 1000, Bettio Arlindo Av. Paulo, ˜o oPuo Brazil Paulo, ˜o Hmn 1962). (Hyman, m ge n Kawakatsu, and Ogren Gigantea eilscin fteatro region, anterior the of sections serial m .sandersoni G. sdsrbdhri n a and herein described is n niqi,Colombia. Antioquia, ´n, Puhe 99,and 1949), (Prudhoe, iatabistriata Gigantea oPuo Brazil. Paulo, ˜o .bistriata G. Downloaded By: [Universiteit van Amsterdam] At: 12:41 19 February 2008 rw ntefrt2m ftergtatro ieo h oy h ein nowihtebd a rgetdfrhsooia preparation histological for fragmented was body the which into regions The cm. body. 1 the bar: of Scale side drawn. anterior Free-hand right indicated. the also of mm are 2 first the in drawn 1. Figure accounts Taxonomic supra-intestinal plate. sp, nerve ventral vn, glands; vagina; vitellaria; v, vi, testes; shell t, muscles; sg, mesenchymal transverse net; transverse peripheral sub-muscular sub-intestinal nerve se, p, sb, muscles; papilla; duct; vesicle; mesenchymal penis pp, ovovitelline prostatic pocket; mo, ov, pv, pharyngeal organ; ph, nucleus; ; genital cell pigment genital accessory n, male male mouth; long- ma, mgo, muscles; lm, atrium; m, papilla; sub- muscles; longitudinal penis ls, epidermic the muscles; of subepithelial nucleus itudinal sunk epithelium in, the es, mass; cyanophilic of gonopore; secretion; ill go, il, erythrophilic intestine; atrium; i, genital er, female fa, duct; esophagus; duct; efferent ejaculatory ed, ef, dorso-ventral eyes; dd, e, common dv, muscles; sole; co, mesenchymal muscles; creeping coat; mesenchymal cr, genital muscle diagonal duct; accessory common ovovitelline male cm, glandular the ; of muscle organ canaliculus barrel-shaped b, c, body; the coat; of end anterior a, abbreviations Figure chiriquii G. of material type 86 ia obl-hpd spau rsn.Inner present. Esophagus bell-shaped. cylind- to Pharynx region. cephalic rical the layer in muscle present mesenchymal only Sub-neural life. in green Flattened Diagnosis .Carbayo F. Gigantea iatamaupoi Gigantea eeas analyzed. also were , Gigantea EIT rslu 1928–33 Bresslau, SERIATA epaia tmsn 1857 Stimpson, Geoplaninae ERCL alz 1892 Hallez, TERRICOLA iatamaupoi Gigantea iatabistriata Gigantea ge n aaas,1990 Kawakatsu, and Ogren a mln.Dru olive Dorsum long. cm 6 ca. p o.Dra imn atr n itiuino h yso h ie ooye o lrt h yswr not were eyes the clarity For holotype. fixed the of eyes the of distribution and pattern pigment Dorsal nov. sp. p nov. sp. , .sandersoni G. ,and ertera a ams m (6 8 km. Palmas, Las road the Near locality Type 80%. ethanol in apparatus: preserved are posterior copulatory body and the 4 and of Region end slides. pharynx 51 on sections slides; sagittal sections 9 transverse region: sagittal on pre-pharyngeal transverse 2: on sections slides; region horizontal behind: 3: 7 1); region (Figure 8/XII/2002, slides; mm 16 8.5 slides on sections locality. 19 to on type tip sections the anterior from in Andre am grass Mauricio the 10:00 on Crawling at 220. PL. MZUSP Holotype. material Type locality. type the from known Only papilla penis Distribution the of canalicula. mgo several of with each wall typically atrium; presents dorsal Penis one. the male female from the the emerging as obliquely long papilla as apparatus atrium Genital Male some long. fibers. with layer sube- longitudinal circular a longitudinal interspersed esophagus by followed one-fiber-thick the layer of pithelial a and of pharynx composed the of musculature 75 olco’ ae,Muii Andre the of Mauricio combination free names, a is collector’s epithet specific The Etymology the to native is species the if region. confirm was to It gardens possible forests. not eucalyptus grass-covered and conifer and by surrounded buildings by occupied Medellı eteo owo h pce aei dedicated. is name species the whom to Restrepo, u 33 9 27 n niqi tt,Clmi.Teae is area The Colombia. State, Antioquia ´n, 0 ) a 90mattd,Mncplt of Municipality altitude, m 1900 ca. W), sMun ´s o Restrepo, ˜oz leg. sMun ´s ein1, Region u 13 9 10 0 ˜oz N, Downloaded By: [Universiteit van Amsterdam] At: 12:41 19 February 2008 etal.I h ehlcrgo,teeihlu is epithelium the region, cephalic 6 the In ventrally. rw tie oreaogtefrt1 m The the mm. in 10 dark borders mm. brown first medial 8 with first yellow, of the pale pair is along side A ventral course brown millimeters. are stripes six body brown the the first of the of margins in The turn color brown. contents dark intestinal ground the it but The converge yellowish, is rounded, pointed. dorsum is borders extremity posterior the anterior the the this extremities, curve; From a the maximum 1). flat forming (Figure the to region very with pharynx and region 10%), the lanceolated in of is width ratio body length The (width: width. in from (Translated scarcer. Spanish.) become in original they of end the body posterior in to the the up towards mm the 7 externally stripe; abundant middorsal of discoloration black are region eyes green The the light length. to of longitudinal appearance up the a has dark it lighter where initially apparatus, grows then copulatory anterior it very and posteriorly, the body; green in the color. black of in is black end the region is which in middorsal body, except The the green of light region is are anterior intestinal dorsum the borders by the the caused branches; of partially is end appearance maybe green, head The posterior olive The pointed. The is width. very crawling. in specimen is when mm 6 living curled and dorsally length The in notes: mm 70 collector’s From morphology External ntemda eino h oslpre-pharyngeal dorsal the of 25 region region, medial the the at 17 in is width epithelium body The sole, region. of creeping pre-pharyngeal 90% the approximately in has only which ciliated is epithelium The secretions subepidermic and body Epidermis the of 60% distance body 73%. to a of gonopore, at equivalent the is 14% length; tip mouth The to anterior row. (equivalent from single a body in the length) of region 1910 width. of invagina- simple total 30 are A body tions pits body. Sensory the the counted. of were end eyes of posterior the one-third to progressively narrows up eyes with with band mm. occupy- the 4 band dorsal, Posteriorly, to become a they up mm 30 ing biserial to become mm 4 Behind eyes a From 1). (Figure the in millimeter region, first region this the cephalic in the row single bordering halos, without m h ysaemnlbtd 30 monolobated, are eyes The mm 6 and length in mm 60 is worm preserved The ih oslyadventrally. and dorsally high, m m ep etal urudn h anterior the surrounding ventrally deep, m m nteltrlrgo n 17 and region lateral the in m m ndiameter, in m m high m m m hc)lctdudrtedra sub-muscular dorsal the under located thick) osldaoa ae ihdcsaefbr (37 fibers decussate a abundant. with layers: most layer three in diagonal the gather directions, dorsal fibers are various mesenchymal dorso-ventral in mesenchymal Other the arranged the are while region, fibers pre-pharyngeal muscle the In musculature Mesenchymal in region. occurs cephalic fibers the the of arrangement height same The body region. to relative subepider- ( The thickness I). musculature (Table bundles mic in layer arranged longitudinal a also diagonal then a and by bundles, a decussate followed Geoplaninae: with is of layer layers circular three subepithelial typical the are There musculature Subepidermic are cyano- observed. region were granulous cells pre-pharyngeal only secretory philic and the ill-stained of and artifactually epithelium tissues ventral The closed open margins. granules body the sparse onto xanthophilic of Scarce cells dorsally. secretory only Granulous open one. cells marginal secretory epithe- the acidophilic dorsal onto the scarcely and onto lium open respectively, rhabdito- cells, and gen cyanophilic granulous cells, secretory . land Froehlich several by for reported (1955) bundles previously in as run ducts, type parallel necks erythrophilic of this The of margins. cells most the of at ventral absent and being dorsal type, epithelium, the fourth onto A the discharge border. with sensory erythrophilic, between xanthophilic, the located and is sole openings creeping type and scarcer granules region: respec- sparse third, cephalic cyanophilic, A the indistinctly tively. in and epidermis xanthophilic entire the onto ucelyrTikesin Thickness layer Muscle al .Tikeso ueiemcmsuauei h pre- the in musculature subepidermic of of region pharyngeal Thickness I. Table eta oa 5– – 2–5 ? 1–2 35 2–5 1–2 15–30 10 22 3 33 total 8 Ventral 23 2 longitudinal Ventral diagonal Ventral circular Ventral total Dorsal longitudinal Dorsal diagonal Dorsal circular Dorsal 5 ntepepayga ein w ye of types two region, pre-pharyngeal the In discharge cells secretory granulous of types Two C,o ch s91 ntepre-pharyngeal the in 9.1% is mc:h) or MCI, e epaia rmColombia from Geoplaninae New iatamaupoi Gigantea m p nov. sp. m ubro fibers of Number e bundle per m 87 m Downloaded By: [Universiteit van Amsterdam] At: 12:41 19 February 2008 ftepayxi aea iwo h ooye cl a:1mm. 1 bar: Scale holotype. the of view lateral in pharynx the of 3. Figure Scale holotype. the of region mm. 1 pre-pharyngeal bar: the of half of section 2. Figure (125 supra- ones, transverse intestinal two and net, nerve peripheral 88 ufe rebre ftepayx hsrgo is region This pharynx. the the in of outer ciliated border free cuboidal, The ruffled is layer. epithelium the subepithelial one-fiber- pharyngeal a longitudinal of around composed thick is columnar musculature its mouth; non-ciliated, squamous, esophagus: short a long. is mm 0.4 There pocket. of to pharyngeal three-quarters cylindrical the pharyngeal occupying long, of 3), mm (Figure 3.9 middle bell-shaped is the pharynx in The located pocket. is mouth The system Digestive the extinct. body, become the and of and number apex in fibers diminish anterior fibers the the dorso-ventral than to apparent sub- Close the more former. are and layer that diagonal supra- dorsal so the than the scarce others, intensely region more the this become fibers In first intestinal the mm. in 1.5 occurs It ca. fibers. transverse by constituted ( neural layer. sub-intestinal sparsely the of more those the somewhat than of distributed are fibers The layer bundles. present supra-intestinal other not The do each. layers fibers is two 1–4 layer of bundles diagonal by dorsal constituted The 2). (Figure respectively h iigeihlu ftepayga oktis pocket pharyngeal the of epithelium lining The sub- layer, fourth a is there region, cephalic the In .Carbayo F. iatamaupoi Gigantea 5 iatamaupoi Gigantea eo h eta ev lt,25 plate, nerve ventral the below m ) n u-netnl(100 sub-intestinal and m), p o.Darmai reconstruction Diagrammatic nov. sp. p o.Darmai transverse Diagrammatic nov. sp. m m m), m) ae.Tepottcvscei opsdo cuboidal of composed is vesicle circular prostatic one-fiber-thick The a layer. of composed is muscularis lumen. papilla atrial The male the fixation. of longitudinal during most a occupies caused shows insertion It probably dorsal one. fold the ventral with the to large, posterior is papilla. papilla vesicle penis penis the inside continues The The duct and ejaculatory sinuous distally. coat a muscle as tubular common proxi- the sinuous rounded penetrates is and which dorso- laterally portion, mally open bent unpaired which the are plane, into sagittal branches the paired to anteriorly Its common the the coat. 4). outside (Figure located muscle of vesicle is vesicle prostatic part prostatic the The proximal of the branches oriented with paired dorso-anteriorly communicate are and portions prostatic ventrally. sub- distal slightly the bend transversal Their and at narrow the 2); they (Figure level, of vesicle layer fibers muscle efferent intestinal between The side. run body same ducts with the of testis duct each efferent the obliquely commu- or two) dorso-ventrally body (rarely the nicates ductule of 55.5% A and respectively. relation 25.4% length, at in end, located, anterior are the Most to testes the 3). (Figure posterior between pharynx and body the anterior of the root of the side and ovaries single- each a on in row arranged to-triple its are between They 2). partially (Figure intestine, diverticles supra-intest- and layer between muscle position, inal in dorsal diameter, ue.Tersaei one,250–300 rounded, their is in spermatozoa shape with Their mature, lumen. are testes The system reproductive Male pharynx the fibers of stroma muscle the oriented in run radially and Longitudinally ueihla ae olwdb iclrlayer circular the a of by and followed (170 pharynx layer longitudinal one-fiber-thick the the subepithelial a of in of consists The esophagus found epithelium. musculature this cells the with inner secretory associated of two are granulous same that esophagus The of to distally. similar types squamous and proximally and of is esophagus, erythrophilic epithelium pharynx inner cells, the The secretory respectively. types cyanophilic, two granulous by crossed and of irregular, being surface cells apical the the of of ciliated, epithelium cuboidal, is The esophagus one. the circular two-fiber-thick a followed layer the com- by longitudinal is outside one-fiber-thick a musculature located of outer posed are The which stroma. of cells, pharyngeal bodies secretory respec- the granules, abundant tively, erythrophilic of and types cyanophilic with two by crossed h feetdcsaedsal ulo pra Its sperma. of full distally are ducts efferent The m )wt oeitrpre ogtdnlfibers. longitudinal interspersed some with m) m min Downloaded By: [Universiteit van Amsterdam] At: 12:41 19 February 2008 nyo rtrpii yecosn h ra.Saebr 50 bar: Scale penis organ. the the are of crossing represented type secretions organ erythrophilic the genital of clarity, only For accessory holotype. male the of a papilla of section itudinal 5. Figure 140–180 ca. as each mgo, organs, protuberances, genital conical accessory male numerous arranged. longitudinally fibers abundant and possesses radially papilla penis layer the of muscle is stroma circular papilla not subepithelial penis was (8 a the papilla of penis of composed the muscularis of The cells examined. cells, epithelial secretory the wall granulous The of respectively. of erythrophilic, and types (6 cyanophilic two layer and by infra-nucleated circular is crossed papilla a penis the of scarce of muscu- epithelium composed Its by a is cells. laris crossed has secretory granulous duct epithelium the erythrophilic ejaculatory ciliated portion; The columnar (25 proximal layer fibers. a very intermingled of its composed is with of muscularis vesicle, prostatic exception whole the the into secre- Scale open vesicle. granulous prostatic cells the tory and Erythrophilic duct efferent epithelium. the between ciliated limit the shows line dashed The papilla. penis the mm. 1 of bar: fold a indicate lines dashed 4. Figure h ufc ftepnspplai rvddwith provided is papilla penis the of surface The m )floe yalniuia n (8 one longitudinal a by followed m) iatamaupoi Gigantea iatamaupoi Gigantea p o.Darmai eosrcino ouaoyaprtsi aea iwo h ooye h double- The holotype. the of view lateral in apparatus copulatory of reconstruction Diagrammatic nov. sp. p o.Rcntuto falong- a of Reconstruction nov. sp. m hc)of thick) m m m ) The m). ) The m). m min m m. 50 epithelial mgo. papilla the the under of penis inconspicuous surface the is layer of long- this layer since and subepithelial of fibers circular circular sunken likely the The most are coat. fibers muscle of itudinal muscle base circular Some under cross the muscle each. arranged, Abundant coat. longitudinally fibers muscle fibers, circular the 2–8 overlain with of interspersed of are those with fibers composed oblique bundles and is longitudinal cyanophilic of coat ill muscle layers an with The filled with barrel-shaped space mass. and strong inner apparent very granules an a and erythrophilic most coat is The muscle mgo circular a section. of histological structure 20– of per maximum a 35 visible being abundant, are They 100–130 and diameter aaiuaoe etal notesraeo the of surface the into centrally open canalicula ooye o lrt,tescein ersne r nyof only are 50 represented bar: Scale secretions organ. the the crossing the type of clarity, papilla erythrophilic penis the For of organ genital holotype. accessory male a of section 6. Figure nmocnan –5urgtcnlcl,35– canalicula, upright 8–15 contains mgo An m ep lgtydltdi t ai.The basis. its in dilated slightly deep, m iatamaupoi Gigantea e epaia rmColombia from Geoplaninae New p o.Rcntuto fatransverse a of Reconstruction nov. sp. m nhih Fgrs5–7). (Figures height in m m m. 89 Downloaded By: [Universiteit van Amsterdam] At: 12:41 19 February 2008 xenlt h feetdcs eidtefemale dorsum the the to Behind curve and ducts. dilated become efferent they slightly atrium the ovaries, run the to They as is height external spermatozoa. same ducts of the at ovovitelline full posteriorly and the dilated of slightly first portion region. (external) the proximal dorso-lateral from its The in arise ovaries body ducts of of third 25.4% ovovitelline to from equal The distance body a length. the at of lay end anterior They the plate. nerve ventral and layer the muscle sub-intestinal The the (1 between maturation. position, elongated of are stage incipient ovaries in are Vitellaria system reproductive Female oe fasbpteilcrua ae (7 layer circular (7 layer subepithelial longitudinal a a by followed com- of is and muscularis two posed cyanophilic The cells, of respectively. secretory openings erythrophilic, granulous receives of and types epithelium to ciliated possible not was them. in it material necks the but secretory cyanophilic discern canaliculi, out muscle by send the the They delimited towards between coat. the also space of and layers the coat are nuclei muscle inside Cell circular canalicula. located through the also of also and and portion organ base distal the through the of its atrium surface male epithelial the through of the the cavity organ outside the each into the located discharge are around penetrate nuclei mgo, and whose erythrophi- granules, of canalicula lic cells secretory the fibers Abundant muscle canaliculus. between not Some do it. and run under papilla penis musculature are the nuclei possess into its sunk and deeply the centrally, more thinner of is surface region epithelial papilla The protuberance. conical right). the (on contraction muscle after and left) the 7. Figure 90 h aearu a uodlt columnar to cuboidal a has atrium male The .Carbayo F. iatamaupoi Gigantea p o.Shmtcrpeetto ftepooe oeo cino aeacsoygntlogn eoe(on before organ, genital accessory male a of action of mode proposed the of representation Schematic nov. sp. 6 . m n eta in ventral and mm) 0.1 m m). m m) hc h rgnldsrpinwsbsd The in based. 13 figure was in description reconstructed apparatus original on copulatory specimens the three from known which only is species This ogtdnladdaoa iesadevlp the 4). envelops (Figure (25–70 atria and female layer fibers and a male diagonal by and longitudinal constituted is It coat muscular Common is (4 atrium one. longitudinal layer the a circular by of subepithelial followed a the muscularis by of The constituted epithelium atrium. the with female filled cross are erythrophilic secretions with lacunae amorphous cells the The Scarce to granules. atrium. cyanophilic female dorsal epithelium adjacent the of the epithelium of of surface that The to distally similar apical is convex. gonopore the and cell non- with columnar, each lacunar columnar, is to tall cuboidal atrium proximally female ciliated, the of epithelium falyro nemnldfbr (6 fibers intermingled the of into layer a glands composed is of shell muscularis its portion; of curved anteriorly openings and epithelium its as is narrow atrium male’s. that female the The as folds portion. long distal some the funnel-shaped of at lumen is extension, exception its It the of rest with atrium. the proximal in female elongated the and is proximally the opens and of It coat, region muscle oriented. common forward the ventrally inside vagina located The is duct. There is ovovitelline 4). (Figure join glandular vagina common and the no anteriorly enter then and and other each plane, sagittal the and iatabistriata Gigantea h vvtliedcshv alcbia ciliated cuboidal tall a have ducts ovovitelline The Hmn 1962) (Hyman, m hc) The thick). m m hc)of thick) m m m) Downloaded By: [Universiteit van Amsterdam] At: 12:41 19 February 2008 h toao h ei ail.Cnee nthe in of epithelium Centered the papilla. beneath penis immediately inside the bundles and in sheath of joined stroma run fibers the muscle Part a 9). these Figure apparently of the of in muscles (inset consists orientated of longitudinally Prudhoe, sheath by penis called pyriform-to-globose organs mgo, the An pyriform of mgo. the small epithelium of half openings present ventral papilla distal epithelium dorsal the The consistent and (1949). is Prudhoe mgo by seen the that of is with description (1949) The Prudhoe 9. Figure in in 4 figure holotype in the reconstructed of apparatus copulatory The examined. ftesceoyclswr o bevd No observed. not were organ. the cells with associated the observed secretory was cells, nuclei musculature secretory The the aspect. of spherical a necks of shows cell which its of the whole in by pyriform crossed half and the inner unnucleated of thin, wall is The invagination penis infranucleated. the is of 10–12 epithelium papilla penis The a the inset). of 8, (Figure of epithelium papilla the consists of histological as invagination one organ it, pyriform in penis of organs An side the 1–4 ventral with section. in 8 the Figure in 8. in organs Figure only seen in located genital detail are accessory papilla in Nr. outlined male FMNH also paratype The is the drawn). (free-hand of which papilla penis that 3147, the is of organ (1962) genital accessory Hyman male a of section longitudinal a of mm. 1 reconstruction bar: a Scale inset, the In 3147. Nr. 8. Figure h nywr nw rmteseiswsre- was species the from known worm only The iatasandersoni Gigantea iatabistriata Gigantea Puhe 1949) (Prudhoe, Hmn 92.Darmai eosrcino ouaoyaprtsi aea iwo h aayeFMNH paratype the of view lateral in apparatus copulatory of reconstruction Diagrammatic 1962). (Hyman, m deep m .Fohih 1978, approaching Froehlich, 1990, canal M. are Kawakatsu, They genital and below. Ogren female from or a horizontally penis and new a the with papilla provided with apparatus share copulatory Geoplaninae a of species genera 16 group the collective of the to addition In species new the of position Taxonomic Discussion genital accessory Male seen. not (1962). also were analysis Hyman organs the with same of consistent are the those worm obser- is the The of 10). 1962) (Figure vations by herein (Hyman, drawn drawn and 20 examined figure holotype, in of Hyman apparatus copulatory The Remarks 12 approximately is to 100–120 sheath opened muscle cavity The pyriform atrium. male a the is there papilla penis the hog h pteimo h aeo h cavity. the of granular base cavity the of cells the of of into epithelium mass open necks the a through They also secretions. are with granular there filled with mass, the is In cavity secretion. the and iatachiriquii Gigantea m ih h pc ewe h ucesheath muscle the between space The high. m m ndaee;teprfr aiy 8– cavity, pyriform the diameter; in m e epaia rmColombia from Geoplaninae New Hmn 1962) (Hyman, ..Fohih 1978, Froehlich, E.M. Gigantea Pseudogeoplana Enterosyringa , ,5 91 E. Downloaded By: [Universiteit van Amsterdam] At: 12:41 19 February 2008 a:1mm. 1 bar: iue10. Figure mm. 1 bar: Scale drawn). (free-hand papilla penis the of organ genital accessory male a of section longitudinal a of reconstruction a inset, 9. Figure 92 .Carbayo F. iatasandersoni Gigantea iatachiriquii Gigantea Hmn 92.Darmai eosrcino ouaoyaprtsi aea iwo h ooye Scale holotype. the of view lateral in apparatus copulatory of reconstruction Diagrammatic 1962). (Hyman, Puhe 99.Darmai eosrcino ouaoyaprtsi aea iwo h ooye nthe In holotype. the of view lateral in apparatus copulatory of reconstruction Diagrammatic 1949). (Prudhoe, Downloaded By: [Universiteit van Amsterdam] At: 12:41 19 February 2008 eu epciey n asdtegopt generic to group the raised rank. and respectively, Peru, Kawakatsu the and to 1955) them Ogren morphological added between Later, (1990) their differences 1967). minor in (Froehlich show others or to subgenus aspects gradually formal pass a the to providing status avoided Froehlich iil otenkdeeo h pdri fthe of epidermis the on body; the eye of region naked ventro-lateral anterior the are which to papillae, extensible of visible tip the at in located absent are characteristics present maupoi Gigantea genera these of all relc 15)icue ihnthe within group: included (1955) Froehlich of species for proposed was group The (1967). formal as: (1990) Kawakatsu and Ogren by terized nlcnlta pn otevnrlsd erthe near side ventral the to complex. opens copulatory that and canal mesenchyme, inal the into Enterosyringa sunk partially culature Gusana and h genus The into only fits species new The .sandersoni G. 1939), (Hyman, n,perhaps, and, the and structures, idaia papilla, these G. with penis provided the species following on ridges glandular .gigantea G. Trinidad. from and extends Peru to group a Rica with The Costa also atrium. some papilla, female gland- penis with dilated the provided on species ridges from the ular coming of or most horizontal below, canal female the with h : (mc body broad large, [ of Geoplaninae eaeatu iae.Lclte nPeru, Trinidad. in and Panama, below; Localities Rica, Costa from dilated. Colombia, approaching antrum or female (but in horizontal as papilla absent penis canal typically be the present, may on ridges papilla ridges penis glandular with 15%); is 1951, 5 Xerapoa C]vleof value MCI] oss h ueiemclniuia mus- longitudinal subepidermic the possess .gigantea G. es relc (1955): Froehlich sensu d .R acs 1951), Marcus, B.-R. (du rsnsasnl ueiemcintest- subepidermic single a presents relc,15.Excepting 1955. Froehlich, .gigantea G. Gigantea Gigantea .bistriata G. ,and , p o. h esr isin pits sensory the nov.: sp. .cameliae G. .vongunteni G. .picadoi G. gopcetdb Froehlich by created -group .idaia G. .chiriquii G. ru eas otspecies most because group a rce rmtenon- the from erected was ru,fo aaaand Panama from group, and Fhmn,1914). (Fuhrmann, Bacap 1912), (Beauchamp, .unicolor G. d .R]Marcus, B.-R.] [du G Fhmn,1914), (Fuhrmann, . Gigantea chiriquii ,bothwithout .gigantea G. .montana G. Liana Gigantea ;female ); (Hyman, charac- , Xerapoa and - , uthrznal rse h ei papilla. penis the ejaculatory the crosses and chiriquii horizontal, horizontally is duct penis two includes the into group subdivided which the first here of A is heteroge- genus groups. discussion a the comparative species, constitute a new For to group. seem neous 2005 Sterrer, and h rxmlpr satpclpnspapilla penis the into typical species the Froehlich a of gigantea inclusion E.M. the as Consequently, questioned 1978). part (Froehlich with proximal in continuity which its atrium, displays the male only the sections of median wall of the whole consisting the one, typical of a fold not a is shape, in oblique as idaia groups: Gigantea the anteriorly. of and group. first the downwards into canal genital female the running Regarding female of the of region atrium exception proximal the the enters the canal of genital With course unknown. gouvernoni the charac- the is taxonomic and with important teristic, genus, an consistent non- ducts, the not its ovovitelline of is since atrium caution, definition female some species with dilated The here group. included this to belong duct ejaculatory gouvernoni their G. and postero-downwards. atrium, dorsal male the runs the from emerging of wall obliquely penis a possess montana e pce sdsigihdfrom distinguished gouvernoni is species new genus. heterogeneous the the for reinforces suggested groups status species the these of any gathering interpreted into of be difficulty must The organs the male cautiously. the and of of interpretation anatomy contracted the the everted, strongly partially is is papilla penis known worm duct only ejaculatory the since and Nevertheless, atrium, oriented). male postero-backwards the of of the wall from characteristics emerging dorsal obliquely canal male (penis group the genital second the and dorsal-to- female the aspect), (the from coming horizontal group atrium female first the enters the of istics running atrium female the anteriorly. canal of and upwards genital region female duct proximal the ovovitelline species, (common these In group. hs pce oehrwith together species These w pce eanta ontftceryit any into clearly fit not do that remain species Two eadn h pce ftescn ru,the group, second the of species the Regarding sandersoni Gigantea includes group second A , ru,adhnei h genus. the in hence and group, Fgr 10), (Figure .picadoi G. nalseiso h ru,tefemale the group, the of species all in , and , , e epaia rmColombia from Geoplaninae New .maupoi G. h ei ail,oiial described originally papilla, penis the .idaia G. .vongunteni G. and .gigantea G. rsnstefml character- female the presents p o. and nov., sp. .unicolor G. and , .bistriata G. Gigantea + Gigantea .gouvernoni G. aia nesthe enters vagina) .sandersoni G. Gaf 1899), (Graff, yterelatively the by .gouvernoni G. .cameliae G. .cameliae G. pce,which species, eogt this to belong , .vongunteni G. .cameliae G. pce,in species, Jones .In , fits G. G. G. G. 93 G. is , Downloaded By: [Universiteit van Amsterdam] At: 12:41 19 February 2008 sandersoni pce,wihrsml hs of those resemble which species, 94 enin seen nopeeydfeetae n ontpresent 1978). not glands of primordia do (Froehlich even not and muscles, differentiated differentiated incompletely non Geoplaninid: the vongunteni in G. in present distinguished mgo, muscularized by can gland- of nature. neither muscular organs, nor has genital ular, species accessory the male that no reporting (1914) description Fuhrmann original by the following misinterpretation cameliae G. of sion ercilla mgo: picadoi with G. species geoplaninid bistriata eight are There in organs genital Geoplaninae accessory male of Diversity comparative A follows species Geoplaninids organs. in below. mgo the genital on the account of accessory structure be of basis male could the on solely species Geoplaninae of new is from the muscularis distinguished its Furthermore, and epithelium absent. lacunar in non muscu- a atrium a with female by The underlain atrium epithelium laris. female the lacunar the of species a details new shows the histological in in atrium: other female each apparatus, from copulatory long differ similarly a present three which these in apparatus genital species. the of length shorter yeisedo uclroe ic hncomparing of when organs since the one, muscular glandular of a instead of be type might 1939), (Hyman details reveal of sections gical deviation the mgo: of of type because common division the from this in included ocrigtemsuaue w ye fmgo of types two musculature, the Concerning h tutrsosre yHmni h histolo- the in Hyman by observed structures The nls oelrecls hc a egadcells gland to epithelium be 1939). seem may (Hyman which regular bundles cells, The large the … some enclose modified. surface, be to the appears bundles muscle to these Where come this papilla. elevating small sometimes at a penis, into the crossed reach of which surface papilla] fibers, the muscle of penis bundles by the intervals [of epithelium Fohih17) ntecmaaiediscus- comparative the In 1978). (Froehlich .Carbayo F. iatamontana Gigantea , iatabistriata Gigantea .idaia G. .idaia G. h raso h om of worms the of organs The . , .sandersoni G. shvn g,wiheietywsa was evidently which mgo, having as and , .montana G. .montana G. , , .maupoi G. .ercilla P. .bistriata G. .maupoi G. iatamontana Gigantea , ya 13)rfre to referred (1939) Hyman , .vongunteni G. ihohrseisHyman species other with and hc r ndqaeto inadequate are which , n uclrzdones, muscularized and ; .bistriata G. .bistriata G. p nov., sp. n rmalspecies all from and .maupoi G. p o. and nov., sp. .vongunteni G. and , .montana G. , .picadoi G. scovered is a not was .ercilla P. p nov. sp. are , G. G. , , ail nelce ihtefml timo the of atrium transfer. female sperm the penis during their partner with inter- maintain interlocked to the the individual papilla stimulate an of might help or mgo epithelium course The the papilla. through penis are they projected since function entirely copulatory a with associated role. and vlto eea ie ihnTricladida. of within mgos times several independent of of evolution evidence position possibly is and the type that adenodactyl morphology reported in (2006) and al. variation et Bipaliidae Grant Paludicola Maricola. families the and in the and in (Terricola), Rhynchodemidae reported also were reach may be should fibers species The the epithelium. re-examined. the turbellarians of surface in the the that beneath usually 78) states p. (1951, 1991) Hyman al. however, membrane; et basement (Rieger layer epidermalcells the under entirely is Turbellarians for musclefibers of position The 1939). (Hyman type glandular of structures the to vongunteni nature muscular a attributed isr(98 eae h raso hsspecies this of organs Marcus the individual with together related for Bois-Reymond be function (1998) a Winsor suggest Du probably not did to (1951) organs. thought prostatic of been it. organs the considered have (1949), have Prudhoe authors After few and geoplaninids, they that suggest homologous. also not are Geoplaninae in existing mgo papilla. penis the into basal deeply each more the continue surrounds organs in onto that while epithelium attach mgo, the and of coat membrane barrel-shaped the this muscle of of base mgo circular the the across of run fibers species the new mgo, that each fact in the canalicula to is addition canaliculus In the maupoi nature. G. of lamellar wall in a the sunk of the not is and sheath papilla, of muscle penis the the papilla, wall penis in the the contrast, from In nature. and lamellar sandersoni a organ, of is the canaliculi wall, the atrium of under the immediately epithelium of located part is from musculature their and papilla penis the in details: cytological sandersoni oelkl,temoin mgo the likely, More h g in mgo The They Geoplaninae. to restricted not are mgo The eyltl skono h ucino h g in mgo the of function the of known is little Very the of morphology and position in differences The .vongunteni G. ifri h oiin n itlgcland histological and position, the in differ , ecie yFhmn 11)a of as (1914) Fuhrmann by described , and p o.i h oeseishvn various having species sole the is nov. sp. .montana G. .idaia G. .maupoi G. .sandersoni G. ihapoal cocoon-forming probable a with .idaia G. , .maupoi G. , p o. h g project mgo the nov., sp. .picadoi G. .maupoi G. h g rjc from project mgo the ucefbr fthe of fibers muscle p o. and nov., sp. , .sandersoni G. .sandersoni G. .idaia G. p o.are nov. sp. but , G. G. G. , Downloaded By: [Universiteit van Amsterdam] At: 12:41 19 February 2008 .Gre FN)terucmlctdadi the ( in the care bistriata their aid in with specimens uncomplicated Dr. of their loan and helping (FMNH) (NHM) Gerber for Harris J. E.A. (PUCRS) sections; Souza histological G. To Acknowledgements flat- and geoplaninid the in hypothesizing worms. Geoplaninae for to tools relationships contribute phylogenetic within might useful and are Tricladida, studies longitudinal characteristics interspersed comparative Both some a by with layers fibers. followed one a muscle layer is, circular subepithelial that the longitudinal epithelium; pharyngeal of thin inner mesenchymal and the organization region, beneath transverse inverted cephalic the sub-neural the to restricted a pre- musculature the of value: taxonomic sence of that characteristics noteworthy logical is no It show natural. maupoi genera being existing of and species state evidence poor the a of in knowledge remains morphological while avoided was here change taxonomic any proposing canalicula. genus, the mgo. papillar of the retracting out the pressure, reduce would driven internal systems muscle the be these of of would Relaxation secretions of and cells fibers, lumen the muscle surrounding the the shortening the the from of for narrow increased was internal basis also which Additional canalicula, papilla. a would a as pressure provides mgo into of and sinking protrusion from mesenchym, papilla the the prevents coat the muscle a underlying of as muscles shortening subepithelial it longitudinal Simultaneous the enlarging 7). (Figure and papilla organ increases the long mgo in the pressure of internal coat muscle barrel-shaped the ugsin na al eso ftetx;J Hesson J. text; the of version and early an comments on valuable suggestions the for (IB-USP) Froehlich letta hr seiec fhtrgniyi the in heterogeneity of evidence is there that Albeit of fibers the of contraction that hypothesized is It p o.peet w nomnmorpho- uncommon two presents nov. sp. and .chiriquii G. epciey;D.E.M. Dr. respectively); , .sandersoni G. and , G. G. isrL 98 set ftxnm n ucinlhsooyin 1991. histology functional and GE. of Aspects 1998. L. Rieger Winsor JPS, Smith S, Tyler RM, the Rieger from and roundworms Some 1949. S. Prudhoe Caribbean family the of from species the to planarians Index 1990. M. land Kawakatsu RE, Ogren Some and 1962. Platyhelminthes LH. Invertebrates: Hyman The 1951. LH. Hyman Central, North, from flatworms of species New 1939. LH. Hyman Australian of Biodiversity 2006. D. Blair R, Sluys LJ, Grant Me Colombie. de terrestres Planaires 1914. O. Fuhrmann landplanarians. chilean of collection a On of 1978. EM. zoogeography Froehlich the to contribution A 1967. CG. Froehlich References 06/ No. (Proc. was (150692/2003-0). FAPESP CNPq work and by 04788-8) This supported acknowledged. partially gratefully reviewers also anonymous Two are English. with help his for relc G 95 So 1955. CG. American Froehlich South On 1951. E. Marcus Bois-Reymond Du ersra ltom Tildd:Trioa.Pedobiologia Terricola). (Tricladida: 42:412–432. flatworms 3. terrestrial Vol. invertebrates, pp Wiley-Liss. of York: 7–140. New Nemertinea. anatomy and BJ, Platyhelminthes Bogitsh Microscopic FW, Harrison editors. In: Turbellaria. Zoological Platyhelminthes: planarians. land 281:420–433. Society IV Linnean the Surinam. of Journal and Indies I: West Part 28:79–166. College Women’s Terricola) Fuji of Tricladida, Bulletin Geoplaninae. (Turbellaria, Geoplanidae 2110:1–25. Novitates Museum American countries. Hill. McGraw York: New 2. vol. Rhynchocoela. National States 86:419–439. United and Museum the of Proceedings America. Systematics South of and review Australia. Tricladida: a in and 4:435–471. localities, Biodiversity (Platyhelminthes: distribution and species paludicolan New planarians Paludicola): freshwater 5:748–792. Neuchatel de Naturelles Sciences des Societe Cie la de Filosofia Se de Paulo, Sao Faculdade de Universidade de 23: Boletim Lilloana Zoologica Acta planarians. 153–162. land Se neotropical Paulo. Sao de Cie 195–279. Filosofia Universidade de Faculdade Letras, de Boletim Geoplanidae. ers nvriaed a al.Se Paulo. Sao Cie Filosofia de 217–255. de Universidade Faculdade Letras, de Boletim geoplanids. e epaia rmColombia from Geoplaninae New bemrooi aooi das taxonomia e morfologia ˆbre reZooi 3:7–80. Zoologia ´rie reZooi 19: Zoologia ´rie reZooi 16: Zoologia ´rie niseLetras, e ˆncias nise ˆncias nise ˆncias ´moires 95