Bijdragen tot de Dierkunde, 50 (2): 292-302 — 1980

Gential ducts in several species of the cyclophorid

Leptopoma (, Mesogastropoda)

by

K. Jonges

Institute of Taxonomie Zoology (Zoölogisch Museum),

University of Amsterdam, The Netherlands

male and female ducts extend Abstract mass. Both genital

the columellar side of the visceral in along mass; sixteen of the female In species cyclophorid genus Leptopoma the mantle cavity they pass along the right side genital ducts were examined; in four species male genital

ducts were examined as well. In the females, bursa copulatrix beneath the rectum. and seminis found to into the receptaculum were open In female duct into the the the genital opens mantle cavity, close to the longitudinal genital aperture. mantle cavity; in the male it is continued through Bursa copulatrix and receptaculum seminis — when present

— described the which lies behind the in cyclophorid snails, were hitherto having open- penis right cephalic the oviduct Tielecke, Berry, Kasi- ings into pallial (Weber, it within the mantle into tentacle, or opens cavity nathan). a seminal which along the right side A number of anatomical details of the female genital duct groove passes

useful of the head far the of the the in Leptopoma are regarded diagnostic characters for as as tip penis,

the of this second condition group. being regarded the more primi-

tive situation.

In both sexes the of the duct INTRODUCTION palliai part genital

has thick glandular walls: in the male this part is

called in the female this Most terrestrial gastropods are pulmonates, num- prostate gland; glandular

duct can be divided in which is bering about 20,500 species. Land prosobranchs a posterior part

called albumen and an anterior called number about 3650 species which are confined gland, part mainly to the wet tropics. They are derived direct- capsule gland. Two kinds of ly from families of marine snails and survived by sperm pouches are communicating with the female duct: the bursa changes in patterns of behaviour rather than by genital copulatrix which receives and and the changes in structures. The major structural modifi- sperm prostatic fluid,

seminis which cation of the land prosobranchs is the loss of the receptaculum to the sperm pass

inner from the bursa. gills, respiration taking place on the surface of the mantle.

There are both diotocardian and monotocardian CYCLOPHORID GENITAL DUCTS land prosobranchs. The Helicinidae and Hydro- cenidae are diotocardian, the , Po- Cyclophorid genitals have been described by We- matiasidae and Acmidae monotocardian. The ber (1924), Tielecke (1940), Morton (1952), present study deals with the anatomy of the genital Berry (1964) and Kasinathan The de- ducts of several (1975). species of the genus Leptopoma scriptions of Weber ( Cyclopborus ceylanicus (So- belonging to the Cyclophoridae. werby, 1843)), Tielecke (species of ,

Leptopoma and 16 more genera) and Kasinathan

THE and 4 MONOTOCARDIAN (Cyclophorus jerdoni (Benson, 1851) spe-

cies of 3 other REPRODUCTIVE SYSTEM genera) gave an overall comparable picture.

In the vast majority of monotocardian The of the male proso- palliai part genital duct opens branchs the sexes are The testis or via an anterior into seminal separate. gonad, aperture a groove is found in the whorls of the visceral which the ovary, upper runs up the side of head to the tip of BIJDRAGEN TOT DE DIERKUNDE, 50 (2) - 1980 293

the penis. It consists of a hind portion extending receptaculum seminis is missing. The palliai parts

have backwards from the point of entry of the testis of both male and female genital ducts a

terminal duct, and a fore portion leading across the mantle opening.

roof. In Pomatias elegans the male genital duct is a

The palliai part of the female genital duct is closed tube from the testis to the tip of the penis.

described duct with In the female the ovarian duct into as a large glandular an ex- opens a pouch-

tensive lateral opening. Two elongated sacs extend like bursa copulatrix. Anteriorly, the bursa, which

posteriorly from their openings at the inner end of receives the penis during copulation, has a wide

this duct. The larger of these is the bursa copula- opening into the posterior end of the mantle cav-

trix. The smaller is the seminis. The and swollen oviduct is sac receptaculum ity. large palliai open

Weber (1924) did not mention the existence of to the mantle cavity along the greater part of its

a smaller sac in Cyclophorus ceylanicus, but she ventral wall. A pouch-like receptaculum seminis is remarked that in the which but section of one case one big sac, missing, a the ovarian duct some-

must have been the bursa, was containing the re- what posterior to the bursa copulatrix functions as

mains of a spermatophore. Kasinathan (1975) such.

observed in living that spermatophores

the are manufactured in palliai part of the male MATERIALS AND METHODS genital duct and are directly passed into the bursa

copulatrix of the female during copulation. All specimens dissected are from the collection of the Insti- In Murdochia the pallidum (Hutton, 1883) tute of Taxonomie Zoology (Zoölogisch Museum) in Am- duct sterdam and have been collected in Indonesian Archi- male genital is a narrow tube closed through- the pelago. out its whole length, as far as the tip of the penis Of female the genital duct the palliai part alongside the (Morton, 1952). Distinct commu- sperm pouches rectum is figured with the foremost part of the ovarian duct.

with the cases of nicating female genital duct are absent. In all a figure is given the columellar side of the

palliai oviduct with the bursa copulatrix and receptaculum A duct section of the ovarian functions as recepta- seminis lying in the plane of the paper. In most cases also a culum seminis. is which figure given shows the ventral aspect of the palliai

In oviduct with the posteriormost of the albumen to- Opisthostoma retrovertens Tomlin, 1938, a part gland gether with bursa, and ovarian duct in seminis receptaculum lying pouch-like receptaculum is present, but a the plane of the paper. distinct bursa copulatrix is missing (Berry, 1964).

The of the male duct palliai part genital opens ABBREVIATIONS USED IN THE FIGURES via slit into anteriorly a a seminal groove.

a — anus

— albumen ag gland

be — bursa copulatrix GENITAL DUCTS IN POMATIASIDAE

eg — capsule gland AND ACMIDAE fcg — folds of capsule gland

ga — genital aperture

hg — hypobranchial gland Creek (1951, 1953) described the genitals of Po-

— of lga lips genital aperture matias ele and Acme fusca gans (Muller, 1774) lpo — lumen of palliai oviduct

(Montagu, 1803). In Acme fusca the male genital m — mantle

mc — mantle cavity duct opens within the mantle cavity into a seminal me —• mantle edge which extends the side of the head groove along od — ovarian duct

— and the whole length of the penis. In the female p penis

pr — prostate a "common" duct links the bursa copulatrix, r —- rectum ovarian duct and the of the glandular palliai part rs — receptaculum seminis

duct with each from — seminal genital other. Apart con- sg groove

— it has sp spermatophore necting these parts an opening into the t — tentacle mantle thus to enter the cavity, permitting sperm td — testis duct

bursa — copulatrix during copulation. A pouch-like wpo glandular walls of palliai oviduct 294 K. JONGES - GENITAL DUCTS OF LEFTOPOMA

Leptopoma diplochilus Sykes, 1903 cavity. Before reaching the palliai oviduct it makes

(Figs. 1-5) one complete coil. The palliai oviduct is a large

glandular duct with an extensive lateral opening; Sykes, 1903: 66, pl. VI figs. 16-18 (type locality: "Wai- pressed against the columellar side lie two sacs, giou"); Schepman, 1909: 197 (Waigeu I.: Piapis Bay); L.

the one of which is the bursa and diplochilus + vars. erythrostoma, typica, maculata, unicolor larger copulatrix (type localities of Schepman, 1915: 28, fig. 7 the four vars.: the smaller the receptaculum seminis. Posteriorly Waiu-waigé, along river Rabiai, Beo, Jefjum, Waimenil, and these sacs extend beyond the palliai oviduct Van Kaiawat, all on Waigeu I.); L. diplochilus: Benthem lie the visceral Jutting, 1963: 673 (West New Guinea: Arguni Bay, in Muka on mass.

entrance of Arguni Bay, near village of Ton- Both the bursa and into Arguni area; receptaculum open geronu). the the oviduct and narrow space between palliai

Specimens dissected. Two SS (figs. 1-3, 4-5) the columellar muscle, thus in fact into the mantle

and one 9, Waigeu I.: Beo (leg. De Beaufort, 1910). cavity; their openings are close to the lateral

opening of the palliai oviduct. Male genital duct. — From the testis In one of the females studied, the bursa con- which completely fills the apical whorls of the tained a spermatophore (fig. 16; in situ: fig. 15). visceral mass, the straight testis duct leads forward In the other female the bursa was filled with a to the posterior end of the mantle cavity. The which vast quantity of a jelly-like substance was prostate is a broad tube with thick glandular walls; for a part bulging out of it. In the second case in it can be divided a fore portion, which has an of mobilization disso- probably a process sperm by and extensive lateral opening, a hind portion. An- lution of one or more spermatophores was going seminal leads the teriorly a groove from genital on. aperture to the tip of the penis. The penis is not a The free edges or "lips" of the genital aperture closed tube but a laterally folded structure inside are somewhat thickened; by their shapes and po- which the seminal groove has its continuation. sitions they keep the genital aperture closed for the

greater part. Female — genital duct. The recepta- On the ventral side of the oviduct tube is palliai a culum seminis is quite large; it nearly as long runs alongside the rectum; anteriorly it into as the bursa copulatrix. Other details are congruent opens the palliai oviduct. It is not clear where this tube with those in the next species, L. gianellii, the originates posteriorly and what function it has. female genital duct of which will be discussed is in and This tube indicated by an arrow figs. 7 more extensively. 12.

The walls of the palliai oviduct become much Leptopoma gianellii Tapparone Canefri, 1886 convoluted anteriorly. The lumen of this part of (Figs. 6-16) the palliai oviduct can be dilated considerably

when its walls are straightened out. L. gianellii + var. a Tapparone Canefri, 1886: 183, pl. II

10-11 "Katau del fiume figs. (type locality: e rive Fly, costa In the other of the meridionale della Nuova Guinea"); Schepman, 1919: 177 species Leptopoma studied,

(West New Guinea: Lorentz River, Digul River, Van der structure of the female genital duct is basically the

Sande River, Bivak 1., Sabang, Resi Ridge, Van Weel's same as in L. gianellii. Therefore, in the discussion Van Camp); L. gianellii + forma a: Benthem Jutting, 1963: of the following species, remarks will be made 668, 669 (West New Guinea: Sermovai River, Sorong, Digul when differences in River). only appear some details.

Specimens dissected. Two 9 9 (figs. 6-13, 14-16),

West New Guinea: Lorentz River (New Guinea Exped., Leptopoma perlucidum 1909). (De Grateloup, 1840) (Figs. 17-23)

Female duct. — From the genital ovary in the whorls of the visceral where it Cyclostoma perlucida De Grateloup, 1840: 442, pl. Ill fig. upper mass, Iles 13 (type locality: "Manille; les Malaises"); Cyclostoma is embedded in tissue of the digestive gland, the vitrea (non Cyclostoma vitreum Draparnaud, 1801) Lesson, ovarian duct leads forward to the mantle straight 1831: 346, pl. XIII fig. 6 (type locality: "la Nouvelle-Gui- - 295 BIJDRAGEN TOT DE DIERKUNDE, 50 (2) 1980

Figs. 1-5. Male genital ducts in Leptopoma diplochilus: 1-3, 4-5 (Waigeu I.).

Figs. 6-23. Female genital ducts within the genus Leptopoma: 6-13, 14-16, L. gianellii (New Guinea); 17-19, L. perlucidum

(Java); 20, L. perlucidum (Enggano I.); 21-23, L. perlucidum (Buru). 296 K. JONGES - GENITAL DUCTS OF LEPTOPOMA

dissected. Three née"); Cyclostoma vitreum: Mousson, 1849: 57, pl. VI fig. Specimens 9 9 (figs. 25-27) and

Talaud Salibabu I. 4 (Java: Kurissan); Leptopoma vitreum: Von Martens, 1867: two $ $, Archipelago: Lirung on (leg.

145 (East Java: Wonosari, Banjuwangi); Van Benthem Jut- Lam, 1926).

ting, 1948: 566, fig. 23 (Java: Tjisolok, Wijnkoopsbay,

Udjung Genteng, Bandjar, Semarang, Gunung Kidul, Wlingi, Male genital duct. — There are no diag- Kali Mrawan, Malang); Leptopoma perlucidum: Van Ben- nostic of difference with L. diplochilus. them Jutting, 1963: 664 (West New Guinea: Jamur, Sorong, points

Klamono, Ajamaru, Merauke, Schouten I., Biak I., Owi I.).

Female genital duct. — The bursa Specimens dissected. Three 9 9 (figs. 17-19) and copu-

two $ $, West Java: Wijnkoopsbay (leg. Lieftinck, 1934); latrix is very large, extending far beyond the pal-

I.: one $ (fig. 20), Enggano Boeah-Boeah (leg. De Jong, liai oviduct posteriorly. The opening of the bursa 1956); one 9 (figs. 21-23), Bum: Wa Katin (leg. Toxo- far lies forward. The posterior part of the palliai peus, 1921). has oviduct a rectangular shape.

Male genital duct. — Like in L. diplo-

chilus the prostate has an extensive lateral opening,

and also in other the situation is the regards same Leptopoma niasense Fulton, 1907

as in that species. (Figs. 28-30)

Female duct. — In all females genital Fulton, 1907: 155, pl. X fig. 6 (type locality: "Nias Id."); examined the posterior part of the palliai oviduct Degner, 1928: 335 (Mentawei Is.); Van Benthem Jutting, 1959a: 184, 185, 187 (Satellite islands of Sumatra: Simalur has a more or less rectangular shape compared I., Pulu Babi, Nias I., Mentawei Is.). in L. In with a more rounded shape gianellii. the Specimen dissected. One $ (figs. 28-30), Nias I.: females from Java the bursa is a small pouch com- wood and ladangs near cave Tegindawa (leg. Van der Meer in size with the 17- parable receptaculum (figs. Mohr, 1931).

19). In the female from Enggano I. (fig. 20) the bursa is twice as long as the receptaculum, and in Female genital duct. —- The bursa copu- the female from Burn (figs. 21-23) the bursa latrix extends far beyond the palliai oviduct exceeds the size of the it contained receptaculum many times. posteriorly; a spermatophore (fig. 29).

Leptopoma intermedium Von Martens, 1867

(Fig. 24) Leptopoma minus Von Martens, 1867

(Figs. 31-32) L. vitreum 8 intermedium Von Martens, 1867: 144, pl. IV

4 fig. (type locality: "Amboina, Buru"); L. vitreum var intermedium Van Benthem Lek- L. vitreum minus Von 1867: IV 6 : Jutting, 1927: 10 (Buru: E Martens, 144, pl. fig.

L. sula, plain of the mountain lake Rana, mountains surround- (type locality: "Amboina"); vitreum var. minus: Van

ing lake Rana). Benthem Jutting, 1928: 154 (Sumba: Laora); L. vitreum

minus: Zilch, 1954: 145 (Ambon). dissected. Specimen One 9 (fig. 24), Buru: sur- roundings of lake Rana (leg. Toxopeus, 1921). Specimen dissected. One 9 (figs. 31-32), Ambon,

in secondary wood (leg. Lieftinck, 1949).

Female — L. genital duct. Like in per- lucidum the posterior part of the palliai oviduct has a rectangular shape. Leptopoma pulicarium Pfeiffer, 1861

(Figs. 33-36)

Leptopoma concinnum (Delessert, 1841) Pfeiffer, 1861: 29, pl. III fig. 7 (type locality: "Batchian"); (Figs. 25-27) Reeve, 1862: Conch. Icon. 13, Leptopoma, pl. V fig. 28

(Batjan); Kobelt, 1901: 444, pl. LV fig. 7 (Batjan). Cyclostoma concinnum Delessert, 1841: pl. XXXVIII fig. 14

(type locality: unknown ("Habite...?")); Sowerby, 1843: Specimens dissected. One $ (figs. 33-35), Batjan: 134, pl. XXIX figs. 223, 224 Minda- 1953); (Philippines: Bohol, Wajaua (leg. Wegner, one 9 (fig. 36), Obi: near nao, Camiguing). Telaga Obi (leg. Wegner, 1953). BIJDRAGEN TOT DE DIERKUNDE, 50 (2) - 1980 297

24-42. Female ducts within the L. intermedium L. concinnum Figs. genital genus Leplopoma: 24, (Buru); 25-27, (Talaud

Archipelago); 28-30, L. niasense (Nias I.); 31-32, L. minus (Ambon); 33-35, L. pulicarium (Batjan); 36, L. pulicarium

(Obi); 37-38, L, massena (Halmahera); 39-42, L. bicolor (Ceram). 298 K. JONGES - GENITAL DUCTS OF LEPTOPOMA

— bursa Leptopoma massena (Lesson, 1831) Female genital duct. The copu-

(Figs. 37-38) latrix, extending far beyond the palliai oviduct

is broad. posteriorly, very

Cyclostoma massena Lesson, 1831: 346, pl. XIII fig. 7 (type

locality: "havre de Doréry, à la Nouvelle-Guinée"); Lepto-

massenae Tapparone 1878: 274 Port Dorey ) poma : Canefri, ( ; Leptopoma papuanum Dohrn, 1862 id., 1883: 261 (New Guinea); id., 1886: 171 (West New (Figs. 47-55) L. callichloros + 1886: Guinea); var. a Tapparone Canefri,

178, pl. II figs. 16-17 (type locality: "Ramoi, di fronte all' Dohm, 1862: 181 (type locality: "In Nova Guinea"); L. isoletta Sorong"); L. crenilabre Strubeil & Boettger in Stru- aurantiellum + vars. a and ß 1886: bell, 1892: 49 (type locality: "Halmahera"); Kobelt, 1897: Tapparone Canefri, II Tan- 176, pJ. figs. 4-5 (type locality: "Kulo-Kadi presso 23, pl. IV fig. 7 (Halmahera); L. callichloros: Smith, 1897: gion Bair; Tangion Bair, costa occidentale della Nuova Gui- 415 (New Guinea: Kapaur); L. massenae: Fulton, 1899: nea"); L. papuanum: Van Benthem Jutting, 1958b: 303 213 (Batjan); L. crenilabre : Kobelt, 1901: 412, pl. LII figs. (Misool: 10 km N. of Lelintah, environs of Fakal, between 1-2 (Halmahera); Zilch, 1954: 142, pl. XIV fig. 15 (Halma- and environs of rainforest Waigama Waima, Solal, near hera; Halmahera: Patani); Van Benthem Jutting, 1959b: 40

Ake Gam, village gardens near Lenmalas); Van Benthem Jutting, (Halmahera: Lamo on the Goal Plains, Mt. Sembilan, 1963: 667 localities in West New L. (many Guinea). Toliwang); massena massena: Van Benthem Jutting, 1963: 666 (many localities in West New Guinea). Specimens dissected. One 5 (figs. 47-50), West

New Guinea: Pionierbivak on the Mamberamo River (Dutch Specimen dissected. One 2 (figs. 37-38), Halma- New Guinea Exped., 1920); two 9 9 (fig. 51 and figs. 53- hera: Ake Lamo on the Goal Plains ("Molukken Exped.", with dark be- a specimen a peristome), N.W. Miscol: 1951). 55,

tween and Waima Waigama (leg. Lieftinck, 1948); one 9

(fig. 52), N. Misool: surroundings of Solal (leg. Lieftinck, 1948). Leptopoma bicolor (Pfeiffer, 1852)

(Figs. 39-42) Leptopoma melanostoma (Petit de la Saussaye, Cyclostoma bicolor Pfeiffer, 1852: 145, pl. XIII fig. 9 1841) ("Locality unknown"); Pfeiffer, 1853: 374, pl. XLYIII figs. (Figs. 56-58) 25-27 (locality unknown); Leptopoma bicolor: Reeve, 1862:

Conch. Icon. 13, Leptopoma, pl. II fig. 13 (locality un-

vitreum latilabre known); L. γ Von Martens, 1867: 144, pi. Cyclostoma melanostoma Petit de la Saussaye, 1841: 308

IV "Ceram L. fig. 5 (type locality: bei Wahai"); latilabre: (type locality: "Nouvelle Guinée"); Pfeiffer, 1846: 173, pl. Von Moellendorff, 1897: 32 (Ceram: Wahai); Kobelt, 1906: XXV figs. 12-15 (New Guinea); Leptopoma melanostoma 451, LVI 1-2 Van Benthem pl. figs. (Ceram); Jutting, 1953: + var. ß Tapparone Canefri, 1883: 258 (type locality of the 284 (Ambon); Zilch, 1954: XVI 42 150, pl. fig. (Ceram: var. ß: "Is. Aru, Vokan"); L. nigrilabrum Tapparone Cane- Wahai) perlucidum forma latilabre: ; L. Van Benthem Jut- fri, 1886: 182, pl. II figs. 12-13 (type locality: "Kapaor, costa 1959b: 44 Goal Plains). ting, (Halmahera, occidentale della Nuova Guinea"); L. melanostoma + var.

rufolabiata: Schepman, 1919: 177 (West New Guinea: Specimens dissected. One 9 (figs. 39-42), West Moaif); L. vitreum melanostomum: Boettger, 1922: 396, pi. Ceram: Honitetoe (leg. De Beaufort, 1910); one 9, Ceram: XXII figs. 28-29 (Aru Is.); Zilch, 1954: 146 (Aru Is.: surroundings of Wahai (collector unknown). Seltutti, Kobroor) ; L. melanostoma + forma ß + forma

rufolabiata: Van Benthem Jutting, 1963: 669, 670 (many

localities in West New Guinea).

Leptopoma sp. Specimen dissected. One 9 (f'gs. 56-58), S.W. (Figs. 43-46) New Guinea: Aen Dua (leg. Lundquist, 1941).

Specimen dissected. One 9 (figs. 43-46), Morotai

Female duct. —- The I.: Goegoeti (leg. Lam, 1926). genital receptacu- seminis lum is a relative large sac, exceeding the The five of the lot have thin specimens a trans- bursa copulatrix in length. lucent whitish shell with eight extremely fine

white bands at the of the weak spiral places very

spiral keels. The shells resemble most those of Leptopoma moussoni Von Martens, 1865

L. cinctellum Pfeiffer, 1861; differences are a (Figs. 59-61)

larger size of the shells and a relatively larger size Von Martens, 1865: 52 (type locality: "Timor bei Kupang"); of the shell apertures, the last whorl being more Rensch, 1933: 495 (Sumba: Mao Marru); L. moussoni mous- globose, soni: Zilch, 1954: 151, pl. XVI fig. 45 (Timor: Kupang); BIJDRAGEN TOT DE DIERKUNDE, 50 (2) - 1980 299

43-65. Female ducts within the Figs. genital genus Leptopoma: 43-46, Leptopoma sp. (Morotai I.); 47-50, L. papuanum (New

Guinea); 51, 52, 53-55, L. papuanum (Misool); 56-58, L. melanostoma (New Guinea); 59-61, L. moussoni (Sumba); 62-65, L. fulgurans (Batjan). 300 K. JONGES - GENITAL DUCTS OF LEFTOPOMA

and Kasinathan bursa L. moussoni: Van Benthem Jutting, 1958a: 91 (East Sumba: ry (1964) (1975), copu-

Prai Rende Baing, Mao Marru; West Sumba: Jawang, Wai, and seminis into the latrix receptaculum open pal- Waimangura; Central Sumba: Langgaliru). liai oviduct. Female genitals in Lep topoma are dis- Specimens dissected. Two 9 9 (figs. 59-61), East similar in this regard. In all species of Leptopoma Sumba: edge of wood near Wai Lekabe Valley (Sumba discussed in this bursa copulatrix and Exped., 1949). paper, recep- seminis found the taculum were to open against wall of the palliai oviduct into the mantle cavity; Leptopoma fulgurans Dautzenberg, 1902 their openings are near the longitudinal genital (Figs. 62-65) aperture.

In Pomatias elegans and Acme fusca a pouch- Dautzenberg, 1902: 3, figs. 6 (t/pe locality: "Obi"); id., like seminis is of the 1903: 15, pl. I figs. 10-11 (Obi); Van Benthem Jutting, receptaculum missing, a part 1959b: (Batjan; 42 Batjan near Wajaua). duct In ovarian functioning as such. both species,

is Specimens dissected. Two $9 (figs. 62-65), however, a pouch-like bursa copulatrix found Batjan: Wajaua (leg. Wegner, 1953). which is situated posteriorly (Creek, 1951, 1953). In Pomatias elegans the bursa, which receives the

Female duct. — Thebursa genital copu- penis during copulation, has a wide opening into latrix shows a particularity. The posterior part of the posterior end of the mantle cavity. In Acme it is in fact which via a separate pouch opens an fusca a "common" duct, linking ovarian duct and outlet into a "corridor". This corridor (indicated bursa copulatrix with the palliai oviduct, has an by an arrow in fig. at the normal 63) opens posi- into opening the mantle cavity as well, which al- wall of the tion against the palliai oviduct near lows sperm to enter directly into the bursa during the opening of the receptaculum seminis. copulation. Thus, in having separate openings into In one of the females examined the corridor the mantle for and cavity the copulatory egg-pro- contained a spermatophore (fig. 62); in the other ducing parts, the female genital system in species female the posterior pouch was completely filled of Leptopoma resembles that in Pomatias elegans substance with a jelly-like of which a thin strand and Acme fusca. the was present in corridor as well. The occurrence of spermatophores in members

of the Cyclophoridae was already mentioned by

Weber and Kasinathan In the Leptopoma decipiens Pfeiffer, 1861 (1924) (1975).

(not figured) bursa copulatrix of three female specimens of Lep-

topoma spermatophores were found (figs. 16, 29,

Pfeiffer, 1861: III 10 29, pl. fig. (type locality: "Batchian"); 62). Reeve, 1862: Conch. Icon. 13, Leptopoma, pl. IV fig. 20 In male of the (Batjan). specimens Leptopoma prostate

has, like the palliai oviduct, an elongated lateral Specimens dissected. One $ and one 9, Obi: Ang- opening, while in other snails, de- gai (leg. Wegner, 1953). cyclophorid scribed in the literature, the prostates have an an- Male genital duct. — There are no diag- terior opening. Probably the relatively large size nostic points of difference with L. diplochilus. of the spermatophores makes an elongated lateral

opening necessary.

Femalegenitalduct. — The long slender Kasinathan (1975) mentioned sizes of the bursa copulatrix extends far the beyond palliai of the spherical egg capsules cyclophorid species oviduct posteriorly. The left of the lip genital he studied. is aperture laterally folded, forming a sort of tube. The diameter of these is egg capsules large enough to explain the large lateral opening of the

DISCUSSION palliai oviduct in these species. In Pomatias ele-

female is slit gans the genital aperture a ventral of According to descriptions cyclophorid female comparable to the elongated lateral opening of the

Weber Tielecke Ber- genitals by (1924), (1940), palliai oviduct in Leptopoma, and the great extent BIJDRAGEN TOT DE DIERKUNDE, 50 (2) - 1980 301

naturf. be correlated also with the of , 1958a. Landmollusken von Sumba. Verh. Ges of it may expulsion Basel, 69: 90-117, pis. MI. the when large egg capsules which, fully formed, 1958b. Non-marine Mollusca of the island of Misool , fill the oviduct completely palliai (Creek, 1951). Nova Guinea, (N.S.) 9: 293-388.

the female duct 1959a. Catalogue of the non-marine Mollusca of Su- Basically the structure of genital -, matra and of its satellite islands. Beaufortia, 7: 41-191, stud- is the same in the 16 species of Leptopoma pis. I-II. in details differences be ied; however, some can 1959b. Non-marine Mollusca of the North Moluccan ,

islands Halmahera, Ternate, Batjan and Obi. Treubia, recognized. These details, which may be regarded 25: 25-87. useful diagnostic characters, are: ( 1 ) the shape of , 1963. Non-marine Mollusca of West New Guinea. the of the oviduct — the albu- posterior part palliai Part 2, operculate land shells. Nova Guinea, (Zool.)

23: XXVII-XXX. — which is 653-726, men gland rounded or rectangular; ( 2 ) pis. BERRY, A. J., 1964. The reproduction of the minute cyclo- the proportions of the sizes of bursa copulatrix and phorid snail Opisthostoma (Plectostoma) retrovertens the form of the lips of receptaculum seminis; (3) from a Malayan limestone hill. Proc. zool. Soc. Lond.,

142: the genital aperture (in most species they are thin; 655-663. BOETTGER, C. R., 1922. Die Landschneckenfauna der Aru- in in a number of species however, the left lip und Kei-Inseln. Abh. senckenb. naturforsch. Ges., 35: is folded or particular thickened, being laterally 355-417, pis. XXI-XXIII. over a part of its length even completely tube- CREEK, G. A., 1951. The reproductive system and embryo- logy of the snail Pomatias elegans (Müller). Proc. malac. like). Soc. Lond., 21: 599-640. A condition of the bursa is particular copulatrix 1953- The morphology of Acme fusca (Montagu) with , found in L. fulgurans. In this species the posterior special reference to the genital system. Proc. malac. Soc. Lond., 29: 228-240. part of the bursa copulatrix is a separate pouch DAUTZENBERG, PH., 1902. Description de mollusques nou- which via an outlet into a "corridor". This opens veaux provenant de l'île Obi (Moluques). Naturaliste, condition the might have some implications on 1902: 247-248.

1903. Contribution à la faune malacologique terrestre generic level. , et d'eau douce des îles Obi et Iolo. J. Conch. Paris, 51: of male ducts in four Comparison genital species 5-20, pl. I. of did not reveal 1928. von den Mentawei- Leptopoma any diagnostic points DEGNER, E., Binnen-Mollusken

Inseln. Treubia, 10: 319-390. of difference. Therefore, morphologic characters DELESSERT, B., 1841. Recueil de coquilles décrites par La- in of the male genital duct seem less Leptopoma marck dans son histoire naturelle des animaux sans

for the of this vertèbres et non encore I-XL (J. appropriate taxonomy group. Shapes figurées: 1-99, pis Roth- schild, Paris). and sizes of spermatophores, manufactured in the DOHRN, H., 1862. Descriptions of new operculated land

are characters. prostate, probably good diagnostic shells. Proc. zool. Soc. Lond., 1862: 181-184.

FULTON, H. G, 1899. A list of the species of landmollusca

collected by Mr. W. Doherty in the Malay Archipelago;

ACKNOWLEDGEMENTS with of and descriptions some supposed new species

varieties. Proc. malac. Soc. Lond., 3: 212-219, pl. XI.

I to Dr. H. E. Coomans and Mr. R. G. Moolen- of am grateful 1907. Descriptions new species of Trochomorpha, , beek for the facilities offered at the department of Malacolo- Cochlostyla, Amphidromus, Bulimulus, Drymaeus, Pla- of gy the Institute of Taxonomie Zoolog/. Mr. J. Zaagman costylus, Stenogyra, Leptopoma, Cyclophorus, Cyclotus, of the Institute is thanked same kindly for his assistance . Ann. Mag. nat. Hist., (7) 19: 149-157, pis. during the preparation of the figures. IX-X.

1840. Mémoire GRATELOUP, J. P. S. DE, sur plusieurs espè-

de nouvelles de ces coquilles ou peu connues mollusques

REFERENCES exotiques, vivants, terrestres, fluviatiles et marins. Act.

Soc. linn. Bordeaux, 11: 389-455, pis. I-IV.

BENTHEM JUTTING, W. S. S. VAN, 1927. Fauna Buruana, KASINATHAN, R., 1975. Some studies of five cyclophorid

Mollusca. Treubia, 7: 1-33, pis. I-II. snails from peninsular India. Proc. malac. Soc. Lond.,

1928. Non-marine Mollusca of Sumba. Treubia, 10: 41: , 379-394.

153-162. KOBELT, W., 1897. Land- und Süsswasserkonchylien. Abh

, 1948. Critical revision of the Javanese operculate land- senckenb. naturforsch. Ges., 24: 19-92.

shells of the families Hydrocenidae, Helicinidae, Cy- , 1901-1906. Die gedeckelten Lungenschnecken (Cyclo-

clophoridae, Pupinidae and Cochlostomatidae. Treubia, stomacea). In: Martini & Chemnitz, Syst. Conch. Cab.,

19: 539-604. 1 (19, N.F.): 401-488, pis. LI-LXI (Bauer und Haspe,

1953. Annotated list of the non-marine Mollusca of , Nürnberg).

the Moluccan islands Ambon, Haruku, Saparua and LESSON, R. P., 1831. Mollusca. In: Voyage autour du Nusa Laut. Treubia, 22; 275-318. monde. . . sur la Corvette "La Coquille" . . . 1822-1825, 302 K. JONGES - GENITAL DUCTS OF LEPTOPOMA

2 (1): 1-471, Moll. pis. I-XVI (A. Bertrand, Paris). boga Exped., Monogr. XLIX lb ( = livr. 43): 109-232,

E. 1865. Ost- X-XVI. MARTENS, VON, Über neue Landschnecken aus pis.

indien und über zwei Seesterne von Costa Rica. Mber. 1915. On a collection of land and freshwater and •——,

dt. Akad. Wiss. Berl., 1865: 51-59. marine Mollusca from Waigeu, Ceram, Puluh Weh and

1867. Die Landschnecken. In: Die Preussische Dierk., 20: 17-33. , Expe- Java. Bijdr.

dition nach Ost Asien, 2: i-ix, 1919. On a collection of land, freshwater and marine (Zool.) 1-447, pis. , I-XXII (Königlichen geheimen Ober-Hofbuchdruckerei Mollusca. Nova Guinea, (Zool.) 13: 155-196, pis. IV- (R. v. Decker), Berlin). VIII.

und kri- E. collection of land-shells from New MOELLENDORFF, O. VON, 1897. Diagnosen neuer Smith, A., 1897. On a

tischer Landdeckelschnecken. Nachr.-Bl. dt. malakozool. Guinea. Ann. Mag. nat. Hist., 20: 409-420, pl. IX.

of Ges., 29: 31-45. Sowerby, G. B., 1843. Monograph the genus Cyclostoma.

A the land Thesaurus 1: XXIII-XXXI MORTON, J. E., 1952. preliminary study of oper- Conchyliorum, 89-156, pis.

culate Murchodia pallidum (Cyclophoridae, Mesogastro- (Sowerby, London).

poda). Trans. R. Soc. N. Z., 80: 69-79. STRUBELL, B., 1892. Landschnecken aus Halmahera. Nachr.-

MOUSSON, A., 1849- Die Land und Siisswasser-Mollusken Bl. dt. malakozool. Ges., 24: 41-50.

von Java: i-iv, 1-126, pis. I-XXII (F. Schultess, Ziirich). SYKES, E. R., 1903. Descriptions of new landshells from the

10: PETIT DE LA SAUSSAYE, S., 1841. Description d'une nouvelle Austro-Malayan region. J. Malac., 64-67, pl. VI.

de zool. des espèce Cyclostome. Revue (Soc. Cuv.), 4: 308- TAPPARONE CANEFRR, C., 1878. Catalogue coquilles rap-

de la Nouvelle-Guinée M. Bull. Soc. 309. portées par Raffray.

PFEIFFER, L., 1846-1853. Die gedeckelten Lungenschnecken zool. Fr., 3: 244-277, pl. VI.

(Helicinacea et Cyclostomacea). In: Martini & Chem- 1883. Fauna malacologica délia Nuova Guinea e delle , nitz, Syst. Conch. Cab., 1 (19): 1-400, pis. A, I-L isole adiacenti. Annali Mus. civ. Stor. nat. Giacomo

(Bauer und Raspe, Nürnberg) . Doria, 19: 1-313, pis. I-XI.

1852. Descriptions of fourteen new of land- 1886. Fauna délia Nuova Guinea e delle , species -, malacologica

shells, from the collection of Hugh Cuming, Esq. Proc. isole adiacenti. Suppl. I. Annali Mus. civ. Stor. nat.

zool. Soc. Lond., 20: 135-147, pl. XIII. Giacomo Doria, (2) 4: 129-200, pl. I.

1861. Descriptions of new of land- TIELECKE, H., 1940. Anatomie, und Tiergeogra- , forty-seven species Phylogenie

shells, from the collection of H. Cuming, Esq. Proc. phie der Cyclophoriden. Arch. Naturgesch., (N. F.) 9:

zool. Soc. Lond., 1861: 20-29, pis. II-III. 317-371.

L. 1862. the Die Mantel- und REEVE, A., Monograph of genus Leptopoma. WEBER, L., 1924. Geschlechtsorgane von

Conchologia Iconica, 13: pis. I-VII (L. Reeve & co., Cyclophorus ceylanicus (Sowerby). Jena. Z. Naturw.,

London). 60: 397-438.

des Natur-Mu- RENSCH, B., 1933. Zur Landschneckenfauna von Sumba. ZILCH, A., 1954. Die Typen und Typoide

Sber. Ges. naturf. Freunde Berl., 1933: 492-508. seums Senckenberg, 13: Mollusca, Cyclophoridae, Cyclo-

141- SCHEPMAN, M. M., 1909. The Prosobranchia of the Siboga- pborinae-Cyclophoreae (1). Arch. Molluskenk., 83:

Expedition. Part II: Taenioglossa and Ptenoglossa. Si- 157, pis. XIII-XVI.

Received: 19 June 1980.