Mem Inst Oswaldo Cruz, Rio de Janeiro, Vol.

98(1): 51-67, January 2003 51

A Bird’s Eye Survey of Central American Planorbid Molluscs

W Lobato Paraense
Departamento de Malacologia, Instituto Oswaldo Cruz-Fiocruz, Av. Brasil 4365, 21045-900 Rio de Janeiro, RJ, Brasil

In the course of two trips to Central America (June 1967 and July-August 1976) I had the opportunity of
collecting topotypic specimens of Planorbis nicaraguanus Morelet, 1849, anatomically defined in this paper, and of
P. yzabalensis Crosse & Fischer, 1879, the identity of the latter with Drepanotrema anatinum (Orbigny, 1835) is
confirmed. The following planorbid species were also found: Helisoma trivolvis (Say, 1817) in Nicaragua, Guate-
mala, Costa Rica and Belize; H. duryi (Wetherby, 1879) in Costa Rica; Biomphalaria helophila (Orbigny, 1835) in
Guatemala, Belize, Nicaragua, Costa Rica and El Salvador; B. kuhniana (Clessin, 1883) in Panama; B. obstructa
(Morelet,1849) in Guatemala, Belize and El Salvador; B. straminea (Dunker, 1848) in Costa Rica; B. subprona
(Martens, 1899) in Guatemala; D. anatinum (Orbigny,1835) in Belize, Guatemala, Nicaragua and Costa Rica; D.
depressissimum (Moricand,1839) in Nicaragua, Costa Rica and Panama; D. lucidum (Pfeiffer, 1839) in Guatemala,
Belize and Nicaragua; D. surinamense (Clessin, 1884) in Costa Rica and Panama; and Gyraulus percarinatus sp. n.
in Panama.
The occurrence of B. kuhniana and D. surinamense is first recorded in Central America, and Gyraulus percarinatus
is the first representative of the genus provenly occurring in the American continent south of the United States.
The following synonymy is proposed: Planorbis declivis Tate, 1870 = Biomphalaria helophila (Orbigny, 1835);
Planorbis isthmicus Pilsbry, 1920 = Biomphalaria kuhniana (Clessin, 1883); Planorbis cannarum Morelet, 1849 and
Segmentina donbilli Tristram, 1861 = Biomphalaria obstructa (Morelet, 1849); and Planorbis yzabalensis Crosse &
Fischer, 1879 = Drepanotrema anatinum (Orbigny, 1835), confirming Aguayo (1933).
Key words: Helisoma duryi - H. trivolvis - Biomphalaria helophila - B. kuhniana - B. nicaraguana - B.obstructa - B. straminea -
B. subprona - Drepanotrema anatinum - D. depressissimum - D. lucidum - D. surinamense - Gyraulus percarinatus - Belize -
Costa Rica - El Salvador - Guatemala - Nicaragua - Panama - synonymy

The present paper is intended to contribute to the with a forceps, so as to prevent it from retracting back to
knowledge of the distribution of Central American the shell. The shell aperture is kept upward, and the snail
Planorbidae. It deals with samples taken during collecting is gradually plunged into hot water at about 70°C for a
trips to the countries of the region, except Honduras. time proportional to the animal’s size (approximately 45
The zoogeographic importance of Central America as sec for snails about 20 mm in shell diameter). If the shell is
a filter bridge and sweepstakes route is widely recognized plunged to the level of the aperture for the first 15 sec, the
(e.g., Simpson 1953). Accordingly, a better knowledge of animal usually will not retract and then, after being com-
its planorbids (associated, when possible, with data from pletely submerged, will die. Care must be taken not to
the fossil record) is essential for investigating the origin exceed the suitable temperature and time of immersion,
of the mixture that has come to make up the Neotropical since blood and tissue cooking would hinder dissection.
planorbid fauna. The dead animal is plunged into cold water and gently
As noted by Harry (1962), over 250 species of planorbid pulled by the head-foot with a small forceps, so as to
snails were named from the Neotropics between 1798 and disconnect the insertion of the columellar muscle. The
1957; and until three decades ago the systematics of the whole animal is then drawn out of the shell, which re-
Neotropical planorbids was “still in a chaotic state” (Malek mains unbroken and filled with water which dilutes the
1969). Undoubtedly appreciable progress has been made blood residuum avoiding its putrefaction within the shell.
thenceforth,to which we have contributed with studies The animal is fixed in slightly modified Railliet-Henry’s
involving topotypes of 106 nominal species, 23 of which fluid (distilled water 930 ml, sodium chloride 6 g, formalin
are considered valid. 50 ml, glacial acetic acid 20 ml), where it must remain for at
METHODS least 24 h before dissection. One change of the fixative
after the first 24 h is advisable. The amount of fluid should
The snails were collected with a strainer 14 cm in di- be not less than 10 times the volume of material to be
ameter and 7 cm deep, with a 16 mesh wire net. Since at fixed. In specimens with lamellate shell, and in most spe-
that time we used no relaxing agents, the following proto- cies of Drepanotrema, the animal is easily broken when
col was followed. A specimen freely moving about, with attempts are made to pull it from the shell. In such cases
the head and foot well exposed, is carefully picked up the whole specimen (animal within the shell) should be
placed in the fixative after immersion in hot water for about
15 sec; owing to CO2 production, the fixative should be
changed several times to avoid unstopping of the vial
Fax +55-21-2280.5840. E-mail: paraense@ioc.fiocruz.br under gas pressure. The radulae are separated from the
Received 7 August 2002 buccal mass by digestion in a vial with 10% NaOH (left
Accepted 12 September 2002 overnight in the incubator at 56°C). They are then rinsed
52 Central American Planorbids • W Lobato Paraense

in tap water. Those of Biomphalaria nicaraguana were Genus Helisoma Swainson, 1840
mounted in a drop of glycerin on a microscopic slide, with
Helisoma trivolvis (Say, 1817)
the dorsal (toothed) surface upwards as in the living ani-
(Fig. 1)
mal; and those of Gyraulus percarinatus (as well as its
shells) were sputtered with gold to be observed with a Twenty-nine specimens from a pool by the margin of
Zeiss electron microscope (model DSM-940). Measure- Lake Nicaragua at Granada, Granada Department, 9 Au-
ments were made on camera lucida drawings. gust 1976 (CMIOC-2318), together with Biomphalaria
Voucher specimens were deposited in the Malacologi- obstructa and Drepanotrema lucidum. Largest shell: di-
cal Collection of the Oswaldo Cruz Institute (CMIOC). ameter 19 mm, width 7 mm at aperture (5.5 mm at begin-

C
Fig. 1: Helisoma trivolvis from Granada, Nicaragua - A: shell; B: renal region, cross-section; C: reproductive system. Bar = 1 mm
 Mem Inst Oswaldo Cruz, Rio de Janeiro, Vol. 98(1), January 2003 53

ning of outer whorl), whorls 5.75. For description, see Nicaragua, 9 August 1976 (CMIOC-2322A), sharing a
Paraense (1976b: 188). swamp with B. nicaraguana and D. depressissimum, to
Additional specimens of H. trivolvis were collected be dealt with below.
from Lake San Cristóbal (or Petencito), near Cobán, Alta B. helophila was collected by Dr R Milward de
Verapaz Department, Guatemala, 5 August 1976, largest Andrade from Lake Coatepeque, El Salvador, April 1963
shell 16.5 mm diameter (CMIOC-2317); from Coris river, (CMIOC-757), 17 specimens. Largest shell: diameter 5.5
Cartago Province, Costa Rica, 12 August 1976, largest mm, width 2 mm at aperture (1.5 mm at beginning of outer
shell 14 mm (CMIOC-2336); and from a pond at Boston whorl), whorls 4.5, lamellate.
village, Belize District, 30 July 1976, largest shell 15 mm B. helophila was studied by Tate (1870: 159, no fig-
(CMIOC-2298). ure) under the name of P. declivis, as follows:
In Tate’s (1870) paper on the Nicaraguan molluscs this 35. Planorbis declivis, Tate. Shell orbicular, depressed, light
species is called Planorbis tumidus Pfeiffer, 1839, as a horn colored, somewhat shining, strongly and regularly
dweller of Lake Nicaragua and San Juan river. Pfeiffer’s wrinkled across; whorls four to five, rounded, suture deep,
original “description” (1839: 354) merely says: “Specimina the last two whorls flat above and below, concealing the
incompleta, Pl. fragili affinia”, so that P. tumidus must be others; the spire and umbilicus deeply and abruptly im-
considered a nomen nudum. Moreover, similarity to P. pressed; last whorl suddenly deflected at the aperture, which
is horizontal and nearly circular; peristome united all round;
fragilis cannot be determined since, in Martens’ (1899: outer lip slightly reflected.
389) words, “P. tumidus, L. Pfeiffer (1839), also undefined, Dimensions of adult shell: greater diameter .25 inch [6.3
is said to resemble P. fragilis, a species undescribed at mm]; height .075 inch [1.9mm].
that time.” Really, the first published description and fig- Marshy pools San Augustin near Acoyapa,living in com-
ures of P. fragilis were given by Dunker (1850: 46-47, Pl. pany with Planorbis kermatoides and Velletia excentrica.
10, Figs. 41-43), and referred to a Mexican Helisoma.
Tate’s description applies well to Orbigny’s syntypes
In spite of the indefinition of P. tumidus Pfeiffer, sub-
(in Paraense 1996a, Fig. 1).
sequent authors treated this name as valid. Dunker (1850:
FC Baker (1945: 85) placed P. declivis in the genus
39-40) described its shell as having 9 lines (19 mm) in
Tropicorbis Brown & Pilsbry, 1914, subgenus Obstructio
diameter and 3 lines (6.3 mm) in width at the aperture, and Haas, 1939, the latter characterized by the presence of
compared young specimens to a flat variety of P. trivolvis apertural lamellae during some stage of growth of the shell.
Say. Strebel (1874: 40-42) gave a detailed description of He also figured (Plate 133, Figs 25-27) a Nicaraguan shell
shells regarded by him as P. tumidus Pfeiffer, collected from the collection of the US National Museum (no. 24006),
together with P. trivolvis at Veracruz, Mexico, and fig- closely resembling our specimen of Fig. 2A. Four years
ured one specimen from Cuba considering it very closely before (Baker 1941) he had proposed the new name
related to trivolvis. If the International Code is to be fol- Tropicorbis tatei for P. declivis, as the latter was preoc-
lowed, P. tumidus Pfeiffer 1839 is not an available name cupied by P. declivis Braun, a fossil planorbid from Ger-
and must be replaced by P. tumidus Dunker, 1850. many mentioned by Genth (1848). This nomenclatural
Helisoma duryi (Wetherby, 1879) change was overlooked by the editor of his posthumous
monograph on the family Planorbidae.
Fifteen specimens from Coris, Costa Rica, were re- Other homonyms are Planorbis declivis Sowerby
ceived from Dr Rodrigo Zeledón (INCIENSA) in August (1877, Pl. 4, Fig. 29) and P. declivis “Gould” Clessin (1884:
1981 (CMIOC-2720); largest shell: diameter 15 mm, width 142, pl. 17, Fig. 13). The former, from unknown locality,
9 mm at aperture (5.5 at beginning of outer whorl), whorls about 14 mm in diameter and 5 mm in width at the aperture,
5. In shell and anatomic characters they are indistinguish- is a wider shell resembling Helisoma. The latter, from Cuba,
able from the specimens studied by Paraense (1976a). 5 by 1 mm in the description but 3.5 mm in the figure,
Genus Biomphalaria Preston, 1910 seems to represent a young specimen of D. lucidum
(Pfeiffer, 1839).
Biomphalaria helophila (Orbigny, 1835)
(Fig. 2) Biomphalaria kuhniana (Clessin, 1883)
(Fig. 3)
Ten specimens from a marsh at El Prado, Izabal De-
partment, Guatemala, 4 August 1976 (CMIOC-2312), to- Thirty specimens from a “Chinese well” in a farm at
gether with Drepanotrema anatinum. Largest shell: di- La Isleta (environs of Panama City), 11 June 1967
ameter 5 mm, width 1.5 mm at aperture and at beginning of (CMIOC-1637). Largest shell: diameter 8.5 mm, width 3
outer whorl, whorls 4.5. In shell and anatomic characters mm at aperture (2.5 mm at beginning of outer whorl), whorls
they are indistinguishable from topotypic specimens from 4.75. In shell and anatomic characters they are indistin-
Peru studied by Paraense (1996a), who also collected this guishable from topotypes from Paramaribo (Surinam) stud-
species in Belize, Nicaragua and Costa Rica (Paraense ied by Paraense (1988).
1996a). As shown by Paraense (1996a), Planorbis dentiens Additional specimens of B. kuhniana were collected,
Morelet, 1849 from Belize, and Tropicorbis shimeki F.C. on the same day, at other localities near Panama City:
Baker, 1945 from Ometepe, Nicaragua, are synonyms of swamps at Las Cumbres, largest shell 7mm (CMIOC-1635);
B. helophila. drainage ditches near the hippodrome, largest shell 8 mm
After publishing those findings I realized that B. (CMIOC-1639); and at the grounds of the National Uni-
helophila also occurs at Acoyapa, Chontales Department, versity, Tocumen, largest shell 8 mm (CMIOC-1642).
54 Central American Planorbids • W Lobato Paraense

The shell of this snail is similar to that of P. isthmicus 112. Pl. nicaraguanus.
Pilsbry, 1920, collected from so called “Chinese wells” at T. discoidea, depressa, obtuse angulata, tenuis, confertissime
Panama City, and here considered a junior synonym of B. striatula, rufo-cornea, nitens, utrinque concava; anfr. 6 sensim
kuhniana. crescentes, sutura profunda discreti; apertura obliqua,
lunato-subangularis. Labrum simplex, tenue.
Biomphalaria nicaraguana (Morelet, 1851) Diam. 16. - Altit. 5.
(Fig. 4) H. lacum Nicaraguanensem.
Fifty-six specimens from a swamp at Acoyapa The type of P. nicaraguanus is shown in Fig. 4A;
(Chontales Department), near Lake Nicaragua, 9 August although larger than the specimen of Fig. 4B, the likeness
1976 (CMIOC-2322), together with B. helophila and D. between them becomes evident on the drawing.
depressissimum. Largest shell: diameter 10 mm, width Smaller shells of our specimens are light horn in color
3 mm at aperture (2.5 mm at beginning of outer whorl), and barely translucent, but owing to adsorption of envi-
whorls 5.25 (Fig. 4B). Reviewing the descriptions of Neo- ronmental material the shell grows dark to become black
tropical planorbids I identified these specimens, by the in full-grown specimens. The growth lines are moderately
shell characters, with Planorbis nicaraguanus, originally marked. The whorls, delimited by deep sutures, increase
described by Morelet (1851: 14, no figure, type locality regularly in width and are roundly protuberant on both
Lake Nicaragua): sides, more on the left. The inner whorl is deep sunken on

Fig. 2: Biomphalaria helophila from El Prado, Guatemala - A: shell; B: reproductive system. Bar = 1 mm
 Mem Inst Oswaldo Cruz, Rio de Janeiro, Vol. 98(1), January 2003 55

the right side and less so on the left, where it is more The melanin pigment of the pulmonary wall is distributed
clearly visible. The periphery is rounded and somewhat into irregular spots which tend to coalesce at the roof of
shifted rightward.The aperture is roundish, with a sharp the cavity. There is no renal ridge.
peristome and a white callus on the parietal wall. There The reproductive system is shown in Fig. 4C. The
are no apertural lamellae. ovotestis is composed of about 60 diverticula, most of
The cephalopedal mass is diffusely brownish gray. which are unbranched. A minutely dissected ovotestis

Fig. 3: Biomphalaria kuhniana from La Isleta, Panama - A: shell; B: reproductive system. Bar = 1 mm (unless otherwise stated)
56 Central American Planorbids • W Lobato Paraense

Fig. 4: Biomphalaria nicaraguana - A: type, Natural History Museum, London (1893.2.4.594); B-E: specimen from Acoyapa, near Lake
Nicaragua. B: shell; C: reproductive system; D: longitudinal section of penial complex. Bar = 1 mm. E: radular teeth; numbers indicate
position in transverse row; C: central; 1, 5: laterals; 8: intermediate; 15, 18: marginals
 Mem Inst Oswaldo Cruz, Rio de Janeiro, Vol. 98(1), January 2003 57

had 77 diverticula, of which 69 were simple, 6 bifurcate (7-13 against 16-33 in nicaraguana), besides having a
and 2 trifurcate. The seminal vesicle is beset with vaginal pouch and a thinner distal loop of the vas defer-
well-developed diverticula varying in shape from knoblike ens.
to fingerlike. Voucher specimens were deposited at the Natural His-
The oviduct is straight in larger specimens, somewhat tory Museum, London.
tortuous in smaller ones, emptying into a moderately de-
Biomphalaria obstructa (Morelet, 1849)
veloped oviducal pouch. The nidamental gland has no (Figs 5, 6)
taxonomically significant characteristics. The vagina is
short and smooth-walled (without vaginal pouch). The Fifty-five specimens from Lake Dueñas, near San
spermatheca varies in shape according to the amount of Miguel Dueñas, Sacatepéquez Department, Guatemala, 3
its contents: club-shaped when empty, when full it shows August 1976 (CMIOC-2307), together with H. trivolvis
an elongated ovoid body about as long as the duct. The and D. lucidum. Largest shell: diameter 10 mm, width 3
caudal portion of the female duct (from the carrefour to mm at aperture (2.5 mm at beginning of outer whorl); whorls
the middle of the oviducal pouch) is about half as long as 5; 22 shells with apertural lamellae. Both shell (Fig. 5A, B,
the cephalic portion (from the middle of the oviducal pouch C) and reproductive system (Fig. 5D) answer perfectly to
to the vaginal opening) in adult specimens, the ratio be- the description of topotypic B. obstructa by Paraense
tween their lengths (caudal:cephalic) varying from 0.42 to (1990). One shell, 8 mm in diameter, with four rest marks
0.57. The ratio between the lengths of the cephalic por- (growth rings), indicative of growth stop (Fig. 5B), showed
tion and of the penial complex varies from 2.85 to 4.76. on dissection six apertural lamellae, not outwardly appar-
The spermiduct gives off a series of 16-33 prostatic ent, a little behind the second rest mark (Fig. 5C), formed
diverticula. They are short and divided into small inter- when the shell was 6.5 mm in diameter.
mingling branches which give the prostate a compact ap- Another sample of B. obstructa was collected from a
pearance, making difficult to ascertain the number of di- pond at Boston village, Belize, together with H. trivolvis
verticula. The foremost diverticulum encircles, and some- (see above): 94 specimens, largest shell 11 mm in diam-
times overlaps, the apex of the spermathecal body. The eter, whorls 5.5, with a set of apertural lamellae in the third
vas deferens, chiefly its distal loop, is specially wide in whorl, 30 July 1976 (CMIOC-2297). Specimens about 7 mm
comparison with the other Biomphalaria species, and in diameter (Fig. 6B) agree with Morelet’s description of
about as wide as the penis sheath. The latter is uniformly P. cannarum from Belize (1849: 16, no figure):
cylindric, about one and a half times as long as the pre- 35. P. cannarum
puce (penis sheath:prepuce ratio 1.53-1.80), and a little Testa discoidea, superne subtiliter perforata, subtus vix
narrower. Owing to shouldering of the preputial upper excavata, striata, corneofuscescens; anfr. 5-6 obtuse angulati,
end (Fig. 4D) it has a swollen appearance. The penial com- supra semiplanulati, inferne semiteretes; apertura oblique
plex (penis sheath plus prepuce) is about 3 to 4 times lunaris.
shorter than the cephalic portion of the female duct (ratio Diam. 7. - Altit. 2.
cephalic portion:penial complex 2.85-4.76). The penis, H. Belise, in littore Hondurasano.
about as long as the penis sheath, tapers to a point where Fig. 6A shows a type specimen of P. cannarum. The
is the outlet of the penis canal (Fig. 4D, pe). Histologically reproductive system of a specimen from Boston is shown
it shows an erectile tissue surrounding the penis canal in Fig. 6C.
and enveloped by a double muscular coat, of which the B. obstructa was collected by Dr R Milward de
inner layer is longitudinal and the outer one circular, the Andrade at San Salvador, El Salvador, April 1963
whole invested by the outer epithelium. As usual with (CMIOC-756): 18 specimens - 1 lamellate, 6.5 mm diam-
Biomphalaria, there are two main extrinsic muscles – a eter; largest shell: diameter 12.5 mm, width 3.5 mm at aper-
retractor and a protractor of the penial complex – in- ture (3 mm at beginning of outer whorl), whorls 5.
serted into the junction of the penis sheath with the pre- B.obstructa is a senior synonym of Segmentina
puce, besides a variable number of secondary retractors donbilli, described by Tristram (1861: 232, no figure) as
and protractors attached to the preputial wall. follows:
In one of the larger specimens the radula had 96 trans-
verse rows of teeth, with the formula 25-1-25 (7 laterals, 6 31. SEGMENTINA DONBILLI, nov. spec. T. compressa,
albido-cornea, tenuiter striata, superne et infra similiter
intermediates, 12 marginals). Radular teeth are shown in
umbilicata, convexiuscula, anfract. 5, lente accrescentes,
Fig. 4E. rotundati: apertura obliqua, rotundata: perist. intus
This snail resembles B. obstructa, from which it dif- albo-labiatum, dentes 6 intus ostendens, quator in pariete
fers in having more protuberant whorls and forming no externa duo in par. interna.
apertural lamellae, but conclusively in anatomic charac- Diam. maj. 9, min. 7 mill., alt. 21/2 mill.
ters: absence of vaginal pouch; prostatic diverticula Hab. Lake of Dueñas.
crowded together and hardly distinguishable; vas defer-
ens, chiefly its distal loop, notably wider; penis sheath Biomphalaria straminea (Dunker, 1848)
longer and a little narrower than the prepuce (compare The occurrence of this species in the Coris river,
with Fig. 5D). Its prostate resembles that of the Cartago Province, Costa Rica (Paraense et al. 1981) corre-
extralacustrine form of Biomphalaria prona (see Paraense sponds to its northernmost record in the American conti-
et al. 1992), but the latter has fewer prostatic diverticula nent (12 August 1976, CMIOC-2335).
58 Central American Planorbids • W Lobato Paraense

Fig. 5: Biomphalaria obstructa from Lake Dueñas, Guatemala - A: shell; B: shell with four growth rings and outwardly inapparent inner
lamellae; C: specimen of Fig. B dissected to disclose inner lamellae; D: reproductive system. Bar = 1 mm
 Mem Inst Oswaldo Cruz, Rio de Janeiro, Vol. 98(1), January 2003 59

Fig. 6 - A: type specimen of Planorbis cannarum (= Biomphalaria obstructa) from Belize; Natural History Museum, London (93.2.4.600-
3); B: shell of Biomphalaria obstructa from Belize (CMIOC-2297); C: reproductive system of B. Bar = 1 mm
60 Central American Planorbids • W Lobato Paraense

Biomphalaria subprona (Martens, 1899) Other samples of D. anatinum were collected at Belize
(pond at Boston village), 30 July 1976, largest shell 2.5 mm
Originally described from Amatitán, State of Tabasco,
(CMIOC-2301); Helado river at Taxisco, Santa Rosa De-
Mexico, this species was found by Paraense (1996b) in
Lake Amatitlán, Guatemala Department, Guatemala. partment, Guatemala, 2 August 1976, largest shell 3 mm
(CMIOC-2302); swamp at El Prado, Izabal Department,
Genus Drepanotrema Fischer & Crosse, 1880 Guatemala, 4 August 1976, largest shell 3 mm (CMIOC-
Drepanotrema anatinum (Orbigny, 1835) 2313); marsh at Rivas, Rivas Department, on Lake Nicara-
(Fig. 7) gua, 10 August 1976, largest shell 3.5 mm (CMIOC-2324);
swamp at Catalina, Guanacaste Province, Costa Rica, 12
Nine specimens from a swamp by the margin of Lake August 1976, largest shell 4 mm (CMIOC-2330); and drain-
Izabal at Mariscos, Izabal Department, Guatemala, 4 Au- age ditch at the grounds of the National University,
gust 1976 (CMIOC-2310), together with D. lucidum. Larg- Tocumen, Panama Province, 12 June 1967, largest shell
est shell: diameter 4 mm, width 1.5 mm at aperture (1.25 mm 3.5 mm (CMIOC- 1644).
at beginning of outer whorl), whorls 5.5. In shell and This species was described under the name of Plan-
anatomy it exactly corresponds to the description of orbis yzabalensis by Crosse & Fischer (1879), based on
D. anatinum by Paraense & Deslandes (1956a). shells from Balancán (Mexico) and Lake Izabal, so that

B
Fig. 7: Drepanotrema anatinum from Lake Izabal at Mariscos, Guatemala - A: shell; B: reproductive system. Bar = 1 mm
 Mem Inst Oswaldo Cruz, Rio de Janeiro, Vol. 98(1), January 2003 61

our material is topotypic. Aguayo (1933) listed P. A single specimen of D. depressissimum, 5 mm diam-
yzabalensis as a synonym of D. anatinum; by the way, eter, was collected from a marsh at Rivas, on Lake Nicara-
the likeness between the two forms was stressed by gua, 10 August 1976 (CMIOC-2326); and a sample of 14
Fischer & Crosse (1880: 75-76). specimens was taken from a swamp at Catalina,
Guanacaste Province, Costa Rica, 12 August 1976, larg-
Drepanotrema depressissimum (Moricand, 1839)
est shell 8 mm (CMIOC-2332).
(Fig. 8)
After describing his Planorbis declivis (B. helophila,
Two specimens from a swamp at Acoyapa, Chontales see above), collected near Acoyapa, Tate (1870) mentions
Department, Nicaragua, in syntopy with B. helophila its being accompanied by P. kermatoides Orbigny, 1835.
and B. nicaraguana, 9 August 1976 (CMIOC-2323). Larg- In all probability he was really dealing with D. de-
est shell: diameter 9 mm, width 1 mm at aperture (0.75 mm pressissimum, which is present throughout South and
at beginning of outer whorl), whorls 5.5. A detailed de- Middle America (including the West Indies), whereas the
scription of this species was given by Paraense and presence of D. kermatoides outside South America needs
Deslandes (1957). confirmation on anatomical grounds.

Fig. 8: Drepanotrema depressissimum from Acoyapa, near Lake Nicaragua - A: shell; B: reproductive system. Bar = 1 mm
62 Central American Planorbids • W Lobato Paraense

Drepanotrema lucidum (Pfeiffer, 1839) Drepanotrema surinamense (Clessin, 1884)

(Fig. 9) (Fig. 10)
Twenty-six specimens from a swamp by the margin of Ninety-seven specimens from a highly polluted drain-
Lake Izabal at Mariscos, Izabal Department, Guatemala, age ditch at Juan Diaz, about 6 km to the northeast of
4 August 1976 (CMIOC-2309), together with D. anatinum. Panama City, 12 June 1967 (CMIOC-1641). Largest shell:
Largest shell: diameter 6 mm, width 2 mm at aperture (1.5 diameter 7 mm, width 2 mm at aperture (1.5 at beginning of
mm at beginning of outer whorl), whorls 5. Shell and outer whorl), whorls 5.5. For description, see Paraense
anatomy as in Paraense and Deslandes’ description (1956b) and Deslandes (1960).
as D. melleum (Lutz, 1918). Additional samples were collected around Panama City,
Other samples of D. lucidum were collected at Belize from a marsh at Las Cumbres, largest shell 6 mm
District, from a pool beside the road to the international (CMIOC-1636); and from drainage ditches at the hippo-
airport (Air Port Mile 8), together with B. helophila, 30 drome Presidente Remón, largest shell 7 mm (CMIOC-1640)
July 1976, largest shell 7 mm (CMIOC-2294); from marshes and at Tocumen, largest shell 5 mm (CMIOC-1645).
by the margin of Lake Nicaragua at Granada, 9 August D. surinamense was also found in Costa Rica, 12 Au-
1976, largest shell 5 mm (CMIOC-2320), together with gust 1976: in a marsh at Catalina, Guanacaste Province,
H. duryi and B. obstructa; and at Rivas, Nicaragua, 10 largest shell 7.5 mm (CMIOC-2331), and in a ditch con-
August 1976, largest shell 3.5 mm (CMIOC-2325), together nected with the Coris river at Cartago, largest shell 4 mm
with D. anatinum and D. depressissimum. (CMIOC-2337).
The occurrence of D. lucidum in Costa Rica is men- This species is indistinguishable by shell characters
tioned by Taylor (1993). from D. lucidum (compare Figs 9 and 10), but the two

Fig. 9: Drepanotrema lucidum from Lake Izabal, Guatemala - A: shell; B: reproductive system. Bar = 1 mm
 Mem Inst Oswaldo Cruz, Rio de Janeiro, Vol. 98(1), January 2003 63

Fig. 10: Drepanotrema surinamense from Juan Diaz, Panama - A: shell; B: reproductive system. Bar = 1 mm
64 Central American Planorbids • W Lobato Paraense

species can be easily separated by the anatomic features. pressed on both sides, a little more on the right. Body
whorl widely convex, with a wide, slightly concave de-
Genus Gyraulus “Agassiz” Charpentier, 1837
pression on the right at the apertural region. Growth lines
Gyraulus percarinatus sp. n. slightly prominent and crossed by deeper spiral lines, so
(Figs 11, 12) that the surface shows a finely reticulate appearance (Fig.
(Latin per: very, extremely; carinatus: keeled) 12B). In nondried shells the surface is beset with lines of
Nineteen specimens from a drainage ditch on the thin hairs running over the spiral lines and more conspicu-
grounds of the Faculty of Agronomy, Panama University ous on the periphery (Fig. 12A). Periphery strongly keeled.
at Tocumen, 12 June 1967 (CMIOC-1643). Largest shell: Aperture narrowly ogival, with a sharp peristome and a
diameter 5 mm, width 1.25 mm at aperture and beginning very thin callus on the parietal wall.
of outer whorl, whorls 5. Cephalopedal mass diffusely brownish gray. Melanin
Shell (Fig. 11A) thin, extremely flattened, honey yel- pigment concentrated in the axis of the tentacles and
lowish, moderately translucent, with slowly increasing arrranged in speckles throughout the pulmonary wall,
whorls plainly visible on both sides and delimited by shal- accumulating on the walls of the pulmonary and renal
low but well-marked sutures. Inner whorls widely de- veins. Kidney margins straight.

Fig. 11: Gyraulus percarinatus from Tocumen, Panama - A: shell; B: reproductive system; C: penis with stylet. Bar = 1 mm (except 11C)
 Mem Inst Oswaldo Cruz, Rio de Janeiro, Vol. 98(1), January 2003 65

Reproductive system shown in Fig. 11B, C. Ovotestis ticula, 21 to 32 in the present sample, ovoid to oblong and
composed of about 30 pear-shaped diverticula, usually unbranched (the foremost one sometimes bifurcate). Vas
unbranched and sometimes bifurcate (28 unbranched and deferens with a descending limb running cephalad to the
2 bifurcate in a minutely dissected ovotestis). Seminal level of the vaginal opening, and turning back into an
vesicle swollen and convoluted, with some small ascending limb about as long as the descending one (with-
outpocketings at the cephalic end. Oviduct short, con- out reaching the level of the male opening as, for instance,
tinuing into a long flat nidamental gland. Vagina short, in Biomphalaria) to open into the proximal end of the
smooth-walled. Spermathecal body usually obovoid, vary- penis. Penis sheath subcylindric, a little swollen at the
ing from club-shaped to spheroidal according to the proximal portion, about as long as the prepuce and a little
amount of its contents. Spermathecal duct about as long narrower. Penis proximally swollen, narrowing gradually
as the body. Spermiduct parallel to the prostatic duct, the toward its distal end, where the sperm canal opens
two organs joining a little beyond the foremost prostatic subterminally. Penis tip furnished with a pin-like stylet.
diverticulum to form the vas deferens. Prostatic diver- Prepuce, somewhat wider than the penis sheath, without

Fig. 12: Gyraulus percarinatus from Tocumen, Panama - A: shell showing hair-like projections; B: shell surface showing reticulate
sculpture; C: radular teeth (central and laterals); D: radular teeth (marginals). SEM prepared by Dr Ortrud Monika Barth.
66 Central American Planorbids • W Lobato Paraense

special characteristics. A retractor muscle is attached to pp: prepuce; pr: prostate; prd: prostatic diverticula; ps
the cephalic end of the penis sheath. Penial complex (pe- penis sheath; pv: pulmonary vein; rm: retractor muscle of
nis sheath plus prepuce) about one-third the length of penial complex; rr: renal ridge; rt: renal tube; rv: renal vein;
the female duct. sd: spermiduct; sp: spermatheca; spd: spermathecal duct;
Jaw fragmented with many plates, as usual in the ge- sv: seminal vesicle; ur: ureter; ut: uterus; va: vagina; vc:
nus. vaginal corrugation; vd: vas deferens; vp: vaginal pouch
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