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MALACOLOGIA

International Journal of Malacology

| Vol. 45(1) 2003
| }
|

MALACOLOGIA
http:\\malacologia.fmnh.org

EDITOR-IN-CHIEF:
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Copyright © 2003 by the Institute of Malacology
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Marine Biological Station
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Museum d'Histoire Naturelle
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Muséum National d'Histoire Naturelle
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Australian Museum
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Academia Sinica
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The Natural History Museum
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Siedlce, Poland

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Zoologisches Institut der Universitat
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Universidad de Chile
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Museum National d'Histoire Naturelle
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Swedish Museum of Natural History
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Forschungsinstitut Senckenberg
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MALACOLOGIA, 2003, 45(1): 1-40

TOWARD А SYSTEMATIC REVISION ОЕ BROODING FRESHWATER
CORBICULIDAE IN SOUTHEAST ASIA (BIVALVIA, VENEROIDA):
ON SHELL MORPHOLOGY, ANATOMY AND MOLECULAR PHYLOGENETICS
OF ENDEMIC TAXA FROM ISLANDS IN INDONESIA

Matthias Glaubrecht'*, Thomas von Rintelen' & Alexei V. Korniushin?

ABSTRACT

The Indonesian island of Sulawesi, with its central zoogeographical position within the
so-called “Wallacea”, harbors a large number of endemic faunal elements, rendering this
region a biodiversity hotspot. The present paper shows that this holds also true for limnic
molluscs of the family Corbiculidae. Although less species-rich than previously assumed,
we document here that the Indonesian corbiculids exhibit more anatomical and life-history
variation than in the rest of their collective Old World distribution.

As a first step toward a comprehensive revision of the Southeast Asian corbiculids,
morphological characters and molecular genetics are studied in the various taxa described
from Sulawesi and Sumatra. Based on morphological studies of materials collected recently,
especially in the central lakes on the island of Sulawesi and supplemented by historical
museum collections, we conclude that Corbicula is represented on Sumatra by at least
one and on Sulawesi by four endemic taxa. Corbicula javanica (Mousson, 1849), known
from several islands of Indonesia, and C. moltkiana Prime, 1878, sampled in lakes Singkarak
and Manindjau on Sumatra are similar in their anatomical characters and the mode of
brooding to the widely distributed Asian C. fluminea (Múller, 1774), but differ from the latter
in shell form and sculpture. The distinctness of C. linduensis Bollinger, 1914, restricted to
the basin of the Palu River in North Sulawesi is confirmed in finding a peculiar mode of
ovoviviparous reproduction, that is, incubation of embryos in the gills until juveniles are
1.3 mm long. Corbicula matannensis P. Sarasin 8 F. Sarasin, 1898, and C. loehensis
Kruimel, 1913, both occurring within the Malili lake system on Sulawesi, as well as C.
possoensis P. Sarasin & Е. Sarasin, 1898, endemic to Lake Poso, all release small larvae,
a reproductive mode similar to C. fluminea, but they differ from the latter in having broad
siphons with slit-like apertures. Corbicula loehensis differs from C. matannensis in its very
delicate sculpture and hinge, whereas C. possoensis is distinguished from other species
in having big posterior adductors and especially broad inhalant siphon. п addition, only С.
possoensis broods in both demibranchs, whereas all other known brooding corbiculids
incubate in the inner demibranch only. Monoflagellate spermatozoa were observed in all
studied Indonesian taxa except C. javanica, in which sperm structure remains unknown.

Phylogenetic analyses of COI sequences (MP and NJ) including now five Indonesian
taxa studied herein show distinct clades occuring (i) on Sumatra, identified as С. moltkiana,
and (ii) on Sulawesi with two seperate lineages of C. possoensis from Lake Poso being
most distinct from C. matannensis and C. loehensis from the Malili lake system. The analyses
also suggest a close relationship of C. javanica to the Korean C. fluminea within an Asian
cluster, including also the Australian corbiculid. Systematic, biogeographical and evolutionary
implications of these results are discussed.

Key words: freshwater Bivalvia, Corbicula, ovoviviparity, anatomy, systematics,
biogeography, endemics, Sulawesi.

'Museum of Natural History, Department of Malacozoology, Institute of Systematic Zoology, Humboldt University,
Invalidenstrasse 43, D-10115 Berlin, Germany

21. |. Schmalhausen Institute of Zoology, the National Academy of Sciences, Kiev, Ukraine

*Corresponding author: matthias.glaubrecht@rz.hu-berlin.de

2 GLAUBRECHT ETAL.

INTRODUCTION

Limnic bivalves of the family Corbiculidae are
widespread in tropical and subtropical regions
of the Old and New worlds. The genus Cor-
bicula is also widely distributed and abundant in
fresh and brackish waters of Africa, the south-
ern parts of Asia, extending from Turkey and
Israel in the west to China, the Malayan Penin-
sula and the Sunda Archipelago, New Guinea
and eastern Australia. Members of the genus
were also introduced to both Americas and
Western Europe (reviews: Morton, 1986;
Araujo et al., 1993; Pfenninger et al., 2002).

Comprising brackish-water and freshwater
species and, therefore, representing different
stages of adaptation to freshwater environ-
ments, renders Corbicula an important model
organism for evolutionary and ecological stud-
ies not only among molluscs. However, its tax-
опоту and systematics is far from being
resolved and many aspects are still disputable.
A great number of species were described,
especially from Southeast Asia (e.g., Martens,
1897; P. Sarasin & F. Sarasin, 1898; Kruimel,
1913; reviews: Prashad, 1930; Morton, 1979),
resulting in a plethora of named taxa. To date a
comprehensive revision of these corbiculids is
lacking.

While an earlier review (Prashad, 1930) dealt
exclusively with conchological characters, rec-
ognizing many congeneric morphospecies,
some later investigations focused on anatomy
(Britton & Morton, 1979; Harada & Nishino,
1995) and particularly on reproductive biology
(Morton, 1979, 1986), resulting in greatly reduc-
ing the number of species considered valid.
For example, Morton (1986) recognized only
two species, namely C. fluminalis Muller, 1774,
and C. fluminea Muller, 1774. According to this
author, C. fluminalis more frequently inhabits
estuaries, tolerates higher salinity (thus, basi-
cally being a brackish-water representative)
and releases free-swimming veliger larvae. In
contrast, C. fluminea occurs in pure freshwa-
ter only and incubates embryos in the gills,
which are not released before the foot of the
juveniles is well developed. Summarizing the
available biological data, Morton (1986) con-
cluded that both Corbicula species are distrib-
uted throughout the range of the genus and
include a great variety of conchological forms
with overlapping characters.

A different taxonomic concept of Corbicula is
accepted in Japan (Harada & Nishino, 1995).
The estuarine, non-incubating species is re-

ferred to as C. japonica Prime, 1864, and the
local freshwater incubating form as C. leana
Prime, 1864, and C. fluminea is reported from
several Japanese localities (Harada & Nishino,
1995; Komaru et al., 1998). Komaru et al.
(1998) provided morphological characters to
distinguish С. leana from С. fluminea. In addi-
tion, C. sandai Reinhardt, 1878, is recognized
as an endemic species restricted to Lake Biwa
in Japan.

To further complicate corbiculid systematics,
new insights into the genetic structure of Asian
Corbicula are salient to any taxonomic revision.
As shown first by Okamoto & Arimoto (1986),
Japanese taxa have different karyotypes, with
С. japonica and С. sandai being diploid (with 2n
= 38 and 36, respectively), whereas C. /eana is
triploid (3n = 54). Recently, polyploidy has also
been discovered in several taxa from Korea
(Park et al., 2000) and in two color forms of C.
fluminea from China, Sechuan Province (Qiu et
al., 2001). It has been repeatedly reported that
polyploidy is associated with peculiar biflagel-
late spermatozoa and ameiotic reproduction,
resulting in clonality (Komaru & Konishi, 1996,
1999; Komaru et al., 1997, 2000; Konishi et al.,
1998; Siripattrawan et al., 2000; Qiu et al.,
2001; Lee et al. 2002). For example, those in-
vestigations of the reproductive biology of Cor-
bicula showed that biflagellate spermatozoa
observed in C. leana from Japan and C.
fluminea from China and Taiwan are non-re-
ductional, and that these molluscs reproduce
by means of androgenesis, that is, the elimina-
tion of the mother’s genome from eggs and
development of embryos from the genome of
spermatozoon only. Similar biflagellate sper-
matozoa were reported for the C. fluminea
samples from Thailand, Korea and the exotic
forms introduced into the USA. Thus, appar-
ently this taxon is a heterogeneous assem-
blage of variably polyploid and ameiotic clonal
lineages (Siripattrawan et al., 2000). Spermato-
zoa of the Australian C. australis (Lamarck,
1818) are also biflagellate (Byrne et al., 2000),
therefore indicating a clonal structure for this
taxon as well (Siripattrawan et al., 2000).

All these clonal lineages, to our present
knowledge, lack sexually reproducing parental
taxa and, therefore, greatly complicate the
meaningful application of specific names. Con-
sequently, the name C. fluminea has been ap-
plied to multiple genetically distinct clonal
lineages of unknown parentage in recent stud-
ies on European and introduced North Ameri-
can populations. In contrast, there is only one

FRESHWATER CORBICULIDAE FROM INDONESIA 3

documented sexual species of freshwater Cor-
bicula, C. sandai, endemic to the “ancient”
Lake Biwa (Hurukawa & Mitsumoto, 1953). In-
terestingly, at the same time only С. запада!
has monoflagellate spermatozoa, thus allowing
to correlate reproductive mode with sperm
morphology (Konishi et al., 1998; Siripattrawan
et al., 2000). In addition, С. запада! is also the
only known gonochoric freshwater Corbicula,
whereas all other taxa appear to be hermaph-
roditic (Komaru & Konishi, 1996; Byrne et al.,
2000; Siripattrawan et al., 2000).

Surprisingly, only diploid karyotypes were re-
ported for the two introduced European Cor-
bicula morphotypes, traditionally identified as
С. fluminalis and С. fluminea (Pfenninger et al.,
2002). In the absence of direct evidence of
clonality in these morphotypes, hybridization
between those two morphotypes, which was
discovered in this molecular study, might indi-
cate sexual reproduction. Accordingly, clonality
is widely distributed, especially among most
Asian taxa, albeit not the universal feature
among freshwater Corbicula. Therefore, the
genetic structure of these limnic clams needs
further investigation.

Furthermore, not only these new data on ge-
netics, polyploidy and reproduction disagree
with the two-species concept of Asian Cor-
bicula as suggested by Morton (1979, 1986).
Preliminary data from mitochondrial DNA se-
quences utilizing the СО! gene (Siripattrawan
et al., 2000; Lee et al., 2002) indicate that C.
leana, С. japonica and С. sandai are distinct
lineages alongside C. fluminea, whereas two
North American morphotypes (forms A and B)
might have different origin, with the first (form A)
being closer related to Japanese C. leana and
the second (form B) to C. fluminea from Ko-
rea. This analysis, as well as a later one that
included samples from China, Israel and Eu-
rope (Pfenninger et al., 2002), demonstrated
that all studied freshwater Corbicula form one
single clade with poorly resolved relationships,
though, with the exception of С.
madagascariensis Smith, 1882, from Mada-
gascar (erroneously referred to as C. africana
from “Africa” in the latter paper as well as in
GenBank). The modest levels of genetic diver-
gence demonstrated for the freshwater lin-
eages suggested evolutionary recent common
origin (Siripattrawan et al., 2000; Pfenninger et
al., 2002).

Despite these accounts, to date many re-
gions remain poorly investigated with respect
to Corbicula diversity and distribution, in par-

ticular islands of the Sunda Archipelago, such
as the Indonesian islands of Sumatra and
Sulawesi. This island chain is among the bio-
logically most diverse regions in the world, rep-
resenting one of the major hot spots of
biodiversity, areas exceptionally rich in endemic
species and harbouring rare and threatened
species (Myers et al., 2000; Mittermeier et al.,
2000; Reid, 1998). Due to its biogeographically
central position within the so-called “Wallacea”,
in the heart of the complex crossroads of two
continents Asia and Australia, Sulawesi not only
harbours a number of unique and endemic fau-
nal elements, but recently also figured promi-
nently in palaeogeographical research
providing new geological insights (Whitmore,
1981; Hall & Blundell, 1996; Metcalfe et al.,
2001). Consequently, this region became a
central focus of biogeographic interest again
(Whitmore, 1987; Hall 8 Holloway, 1998;
Metcalfe et al., 2001).

Although molluscs have unfortunately only
rarely been considered in biogeographic re-
search (Davis, 1982), especially limnic gastro-
pods from the Sunda region were recently
utilized as models in an approach to synthesize
systematic and geological patterns (overview:
Glaubrecht, 2000). For example, based on the
known distributional pattern found in the con-
stituent taxa for the mainly viviparous
Pachychilidae which are widely distributed
throughout the mainland of Southeast Asia and
the Indo-Malayan Archipelago, reaching as far
east as the Philippine Islands and Sulawesi, it
has been hypothesized that the biogeography of
these limnic snails (i) find their explanation in
palaeogeographical events that go back to the
Cretaceous and early Cenozoic instead of ex-
plaining the distribution as correlated to the
forming of the so-called Sunda- and Sahulland,
respectively, and (ii) that it implies vicariance
over dispersal as causation (Glaubrecht, 2000;
Köhler et al., 2000; Köhler & Glaubrecht, 2001,
2003; Glaubrecht & Rintelen, 2003).

In contrast, according to Siripattrawan et al.
(2000) and Pfenninger et al. (2002), the known
patterns of distribution and genetic divergence
in Corbicula, based on data for continental
Southeast Asia, Japan and Australia, suggest
rather dispersal than vicariance scenario for
these freshwater bivalves. Therefore, it is
promising to test the mentioned scenario by
extending the data set, and to compare the pat-
terns of morphological and genetic divergence
among Indonesian bivalves with that of the
sympatric pachychilid snails. In this context,

4 GLAUBRECHT ETAL.

the corbiculid bivalves provide a second model
group that inhabits the same limnic environ-
ments in this crucial biogeographic region and,
with them incubating eggs and embryos in their
gills, also share a similar reproductive strategy
with the ovoviviparous and viviparous
pachychilid gastropods.

However, any zoogeographical evaluation has
to be based on solid systematic knowledge.
Unfortunately, any modern revision of the
Corbiculidae is still lacking. For example, from
Sulawesi a total of nine endemic corbiculid spe-
cies have been described, especially from its
ancient central lakes (Martens, 1897; Р.
Sarasin & F. Sarasin, 1898; Kruimel, 1913;
Bollinger, 1914). In contrast, Prashad (1930)
recognised only two endemic species on
Sulawesi, assigning all lacustrine taxa to C.
subplanata Martens, 1897. In his revision of the
corbiculids from Sulawesi, Djajasasmita (1975,
1977) recognised four taxa - three endemic
species living in lakes in addition to C.
subplanata as the only riverine form. He also
reported on one widely distributed Asian spe-
cies, C. javanica (Mousson, 1849), as occur-
ring on Sulawesi. From Sumatra, also a total
of nine endemic species of Corbicula have
been described (Prime, 1878; Clessin, 1887;
Martens, 1897, 1900), of which Djajasasmita
(1977) recognized four as valid — С. moltkiana
Prime, 1878, C. gustaviana Martens, 1900, C.
sumatrana Clessin, 1887, and C. tobae Mar-
tens, 1900, and, in addition, recorded four
widely distributed Asian species on the island,
viz. С. javanica, С. pullata Philippi, 1851, С.
rivalis (Philippi, 1850), and C. tumida Deshayes,
1854.

However, none of these studies provided any
sufficiently distinctive characters for the indi-
vidual species. Although shell proportions,
angle between lateral teeth, position of beaks,
shell thickness and sculpture are usually used,
the intraspecific variability of these characters
remained largely unknown and, therefore, the
taxonomic decisions appeared as being highly
arbitrary. Consequently, Morton (1979, 1986)
tentatively suggested conspecificy of the Indo-
nesian species reviewed by Prashad (1930)
and Djajasasmita (1975, 1977) with С.
fluminea, synonymizing the species names
listed above with the latter taxon. However,
Morton did not discuss in detail the taxonomy,
nor did he provide any new data on the mor-
phology or biology of these Indonesian
corbiculids. Thus, not only is the systematics
and phylogenetic relationships of the presum-

ably endemic insular Corbiculidae unknown,
but the anatomy and reproductive biology of
any of the disputable species from this region
has remained undescribed.

The recent discovery of an endemic genus of
Corbiculidae from Lake Poso with its unique
cemented mode of life (Bogan 8 Bouchet,
1998) shows that the molluscan fauna at least
of this lake on Sulawesi is much more specific
than assumed earlier. This stimulated the
present study of other Corbiculidae, namely
species of the genus Corbicula inhabiting Poso
and neighbouring lakes, as was suggested by
Bogan 8 Bouchet (1998). Accordingly, we here
integrate shell characters, new data on the
anatomy, especially the reproductive biology,
and molecular genetics (sequences of COI
mitochondrial gene fragment) for six nominal
taxa of Corbicula from Indonesia, namely C.
javanica, С. moltkiana, С. linduensis, С.
matannensis, С. loehensis and С. possoensis,
based on recent sampling by the authors on
Sumatra and Sulawesi, and we compare with
historical material deposited in the museum
collections. In addition, we report for the first
time for any Indonesian Corbicula principal fea-
tures of sperm morphology, as well as pres-
ence and localization of larvae in gills. We also
provide a preliminary evaluation of the status
and affinities of the studied taxa, but any final
taxonomic decision is postponed until more
molecular data on the diverse morphotypes
from lacustrine and riverine habitats in the area
are available. Some other taxa described by
Martens (1897, 1900) from Sumatra and
Sulawesi are also awaiting revision. However,
because no fresh material on these taxa was
available, these are beyond the scope of our
current study. Nevertheless, we document here
that the corbiculids endemic to the region (Fig.
1), although less species-rich than assumed
before, exhibit more anatomical and life-history
variation than in the rest of their collective Old
World range.

MATERIALS AND METHODS
Material Studied

The material at hand (Fig. 1) was collected
during two field trips to Sulawesi in August
1999 and March 2000 by M. G. and T. v. R., and
on Sumatra in April 2000 by Frank Kóhler and
Sabine Schütt. It is housed in the Malacological
collection of the ZMB, voucher material is also

FRESHWATER CORBICULIDAE FROM INDONESIA 5

© C. javanica

У С. moltkiana

À С. Iinduensis

Ф С. possoensis

® С. matannensis

В
Sulawesi

Ш C. loehensis

С

> Malili lake system

e Matano
o
$
q Mahalona
ys Lontoa
Ye | 4 O à у BE
7 Towuti
\e <
m
2 Мазар!
50 | 10

FIG. 1. The occurrence of freshwater bivalvia of the genus Corbicula on Indonesian islands based
only on material examined in the present study; details are given for C. moltkiana on Sumatra (A)
and four taxa on Sulawesi (В) and the Май! lake System (C), respectively. Numbers refer to the
following locations: 1 - Lake Manindjau, 2 - Lake Singkarak, 3 - Palu River, 4 - Lake Poso. Scale bars
given in km.

6 GLAUBRECHT ET AL.

provided to the Zoological Museum in Bogor.

Type and other historical material represent-
ing Corbicula species from Sulawesi and
Sumatra, including the type specimens of C.
subplanata Martens, C. celebensis Martens, C.
lacustris Martens, C. possoensis P. Sarasin &
Е. Sarasin, and С. matannensis P. Sarasin 8 F.
Sarasin, also housed in the ZMB, were studied
for comparison. This is supplemented by the
relevant type materials recovered in other mu-
seums, including the lectotype of C. fluminea
designated by Araujo et al. (1993), as well as
type lots of C. javanica and C. sumatrana.
Comparative alcohol material of C. fluminea
was kindly provided by С. Ituarte (collected
near Buenos Aires, Argentina).

Shell Morphology and Anatomy

All newly collected material was fixed in 70%
ethanol after cracking the shells of some
specimens per sample; this material is given
as w = wet material in the Material sections
below. Shell measurements were made with a
caliper to a precision of 0.1 mm; size of adduc-
tor scar was measured as the distance be-
tween its uppermost point and junction with
mantle line. Dissections were made under a
Leica MZ 9.5 stereomicroscope, and anatomi-
cal structures illustrated using a camera lu-
cida. Pieces of mantle for the study of
musculature were stained by eosine (water
solution) and mounted in Canada Balsam.

Sperm Morphology

Sperm was obtained from gonads of ethanol-
fixed specimens. Its morphology was studied
by means of interference contrast optics (DIC)
and scanning electron microscopy (SEM), in
the latter case applying hexamethyldisilazane
(HMDS) following the procedure described by
Nation (1983).

Molecular Genetics

DNA was purified from about 1-2 mm? of foot
tissue by CTAB extraction (Winnepenninckx et
al., 1993). Polymerase chain reaction (PCR)
was used to amplify a region of ~710 bp at the
5’-епа of the cytochrom oxidase subunit | gene
(COI). PCR was performed in 25 ul volumes
containing 1X Taq buffer, 1.5 mM MgCl2, 200
UM each dNTP, 1-2.5 U Taq polymerase, ap-
proximately 100 nM DNA and ddH,O up to vol-
ume on a Perkin Elmer GeneAmp 9600 or

2400 thermocycler. After an initial denaturation
step of 3 min at 94°C, cycling conditions were
35 cycles of 1 min each at 94°C, 45-53°C, and
72°C, with a final elongation step of 5 min.

Primers used were LCO 1490 [5’ GCTCAA
САААТСАТАААСАТАТТ 37] and HCO2198 [5'
ТАААСТТСАСССТСАССААААААТСА 31
(Folmer et al., 1994). PCR products were puri-
fied with QiaQuick PCR purification kits
(Qiagen) following the standard QiaQuick PCR
purification protocol. Both strands were cycle
sequenced with the original primers using ABI
Prism BigDyeTM terminator chemistry and vi-
sualized on an ABI Prism 377 automated DNA
sequencer. The resulting sequence electro-
pherograms of both strands were corrected
manually for misreads and merged into one
sequence file using BioEdit Version 5.0.1 (Hall,
1999). Sequences were aligned manually and
checked by translating the DNA sequences
into amino acids in DAMBE 4.0.75 (Xia & Же,
2001) using the genetic code for invertebrate
mitochondrial DNA. The sequences obtained
by this study were analyzed together with Cor-
bicula sequences published by Siripattrawan et
al. (2000) and Pfenninger et al. (2002); the lat-
ter included samples from Hong Kong, which
is near to the type locality (Canton) of C.
fluminea, and from Israel (presumably С.
fluminalis).

Polymesoda caroliniana (Bosc, 1801) and
Neocorbicula limosa (Maton, 1809) were used
as outgroups. The latter taxon needs a nomen-
clatorial commentary. As pointed out by
Parodiz (1996: 265), the generic name
Cyanocyclas Blainville, 1818, is a senior sub-
jective synonym of Neocorbicula Fisher, 1887,
and, therefore, should have priority. Under-
standing that this taxon 1$ in need of a taxo-
nomic revision and possibly formal decision of
the ICZN, we for the time being will use here
the latter generic name, as done in recent mo-
lecular literature (e.g., Siripattrawan et al.,
2000; Pfenninger et al., 2002). GenBank ac-
cession numbers of all sequences used and
ZMB catalogue numbers for original material
are provided in Table 1.

Aligned sequences were processed with
PAUP* 4.0b10 (Swofford, 1998). Corrected se-
quence divergence levels were calculated by
using a General Time Reversible model, to ob-
tain the matrix comparable with that of
Siripattrawan et al. (2000). Phylogenetic trees
were reconstructed using neighbor joining (NJ,
Saitou 8 Nei, 1987) and maximum parsimony
(MP) methods as implemented in PAUP*. NJ

FRESHWATER CORBICULIDAE FROM INDONESIA Y

TABLE 1. Sources of the corbiculid material utilized in this study for СО! sequence data analyses;
numbers in brackets refer to analyses of sequence data as given in Table 4 and Figs. 18 and 19.

Museum GenBank
Taxon Locality data catalog по. accession no. Reference
Corbicula javanica Bogor, Java ZMB 106449 AY275668 This study
(Mousson, 1849)
C. moltkiana L. Singkarak, ZMB 103024 AY275660 This study
Prime, 1878 (1) Sumatra
C. moltkiana (2) L. Singkarak, ZMB 103034 AY275659 This study
Sumatra
C. moltkiana (3) L. Manindjau, ZMB 103025 AY275657 This study
Sumatra
C. moltkiana (4) L. Manindjau, ZMB 103032 AY275658 This study
Sumatra
C. matannensis Sarasin L. Matano, ZMB 103002 AY275663 This study
& Sarasin, 1898 (1) Sulawesi
C. matannensis (2) L. Matano, ZMB 103003 AY275664 This study
Sulawesi
C. matannensis (3) L. Mahalona, ZMB 103009 AY275665 This study
Sulawesi
C. loehensis Kruimel, L. Lontoa, Sulawesi ZMB 103033 AY275667 This study
19131)
C. loehensis (2) Е. Masapi, ZMB 103011 AY275666 This study
Sulawesi
С. possoensis Sarasin & L.Poso, Sulawesi ZMB 190024 AY275661 This study
Sarasin, 1898 (1)
C. possoensis (2) Е. Poso, Sulawesi ZMB 103028 AY275662 This study
C. fluminea Thailand UMMZ AF196270 Siripattrawan et al.,
(Múller, 1774) 266691 2000
C. fluminea Korea UMMZ AF196269 Siripattrawan et al.,
266690 2000
C. fluminea Hong Kong - AY097292 Pfenninger et al.
2002
C. leana Prime, 1864 Japan UMMZ AF196268 Siripattrawan et al.
266668 2000
C. sandai Reinhardt, L. Biwa, Japan UMMZ AF196272 Siripattrawan et al.
1878 266689 2000
C. fluminalis ? Israel - AY097299 Pfenninger et al.,
2002
C. australis NSW, Australia UMMZ AF196274 Siripattrawan et al.,
(Lamarck, 1818) 266662 2000
Corbicula “form А” Michigan, USA UMMZ AF196280 Siripattrawan et al.,
266693 2000
Corbicula “form В” Utah, USA UMMZ AF196278 Siripattrawan et al.,
266695 2000
C. madagascariensis Madagascar UMMZ AF196275 Siripattrawan et al.,
Smith, 1882 255293 2000
С. japonica Prime, 1864 Japan UMMZ AF196271 Siripattrawan et al.
266688 2000
Neocorbicula limosa Argentina UMMZ AF196277 Siripattrawan et al.
(Maton, 1809) 265500 2000
Polymesoda caroliniana Florida, USA UMMZ AF196276 Siripattrawan et al.,
(Bosc, 1801) 265499 2000

8 GLAUBRECHT ET AL.

analyses were conducted using the random
initial seed option to break ties. The robustness
of inferences was assessed through bootstrap
resampling (1000 replicates) (Felsenstein,
1985). In the MP analyses, the heuristic search
algorithm was employed with 10 random addi-
tions of taxa and tree bisection-reconstruction
(ТВК) branch swapping. All other settings
were left at default values. Support for nodes
was estimated by bootstrap resampling (500
replicates) with one random addition per repli-
cate.

Abbreviations Used in Figures

aa — anterior adductor, es — exhalant siphon,
is — inhalant siphon, mc — concentric mantle
musculature, mp — papillae, mr — radial mantle
musculature, p — papillae, pa — posterior ad-
ductor, pss — presiphonal suture, sr — siphonal
retractor.

Museum Acronyms

МЕР - Museo de La Plata, Buenos Aires, Ar-
gentina; MZB — Zoological Museum Bogor, In-
donesia; SMF — Senckenbergsmuseum, Frankfurt/
Main, Germany; UZMC - Universitetets Zoo-
logisk Museum, Copenhagen, Denmark; ZMA —
Zoological Museum Amsterdam, The Nether-
lands; ZMB - Museum für Naturkunde,
Humboldt University, Berlin, Germany (formerly
Zoological Museum Berlin); ZMZ — Zoolo-
gisches Museum, Universitat Zürich, Switzer-
land.

SYSTEMATIC ACCOUNT
Species from the Sunda Islands

Corbicula javanica (Mousson, 1849)
Figs. 2A-C, 3D-F

Cyrena orientalis var. javanica Mousson, 1849:
86, pl. 15, fig. 2.

Corbicula ducalis Prime, 1862: 274; Martens,
1897: 114.

Corbicula javanica — Martens, 1897: 111;
Prashad, 1930: 203, pl. 25, figs. 7-20;
Djajasasmita, 1977: 6.

Type Locality: Tikojia (probably an error for
Tjikoya), Java.

Type Material: Lectotype ZMZ 532199 (Fig. 2A)
from Tijkoya, Java, leg. Zollinger, ex. coll.
Mousson; corresponding to the specimen fig-

ured by Mousson (1849: pl. 15, fig. 2) with the
following measurements: L = 39.5, Н = 33.0,
W/2 = 11.9 mm (present designation, to fix
the status of this specimen as the sole name
bearing type). Paralectotypes (2 specimens)
from the same original lot, ZMZ 532199a.
Paralectotypes ZMZ 532200 (2 specimens),
the same locality and collector.

Other Material Examined: Java: Tjiponnas near
Garut (ZMB Moll. 103054w; leg. М. Schmidt
1902); Bogor (ZMB Moll. 106459; leg. Т. м.
Rintelen, May 2002). Lombok: Narmada (ZMB
Moll. 75535w; leg. Rensch, Sunda Expedi-
tion). Sulawesi: Lake Tempe (ZMB Moll.
103024; leg. Max Weber; originally identified
as Corbicula ducalis Prime, 1862).

Taxonomic Remarks: In respect to the discrep-
ancy of the spelling of the type locality for this
taxon, we follow here Djajasasmita (1977),
who interpreted it as Tjikoya. The latter author
also synonymized C. ducalis Prime, 1862,
which was reported earlier on as a valid spe-
cies from several Indonesian islands (Mar-
tens, 1897; Prashad, 1930) with C. javanica.
Our revision of specimens from Sulawesi
originally identified by Martens as C. ducalis
is consistent with this point of view, although
some minor differences to the typical C.
javanica are discernable. Specimens from
Lombok included in this study (Fig. 2C) also
correspond well to the published descriptions
(Prashad, 1930) and revised type specimens
of C. javanica (Fig. 2A, B).

Description

Shell: Oval or broad triangular, without angles,
somewhat inequilateral, convex. Beaks
broad, protruding, markedly shifted anteriorly.
Periostracum yellow to brown, shiny. Internal
coloration usually white or pale blue, with
purple pattern at lateral teeth. Concentric
sculpture coarse and widely spaced (8-11
ribs per 1 cm), ribs usually not sharp (wave-
like). Hinge plate relatively narrow; cardinal
teeth small; anterior lateral teeth long,
arched. Specimens from Lombok and
Sulawesi up to 20 mm long; Javanese speci-
mens much larger, up to 50 mm long.

Anatomy: Adductors small, oval. Posterior ad-
ductor diameter about 0.13 length of shell
(Table 2). Presiphonal suture not elongated,
length equal to breadth of inhalant siphon. Si-
phons conical, with thick walls (when con-
tracted) and circular or short oval apertures,
both narrow; inhalant siphon with about 30

FRESHWATER CORBICULIDAE FROM INDONESIA Él

FIG. 2. Shells of Corbicula javanica in comparison with C. fluminea: A. C. javanica, Java, lectotype
(ZMZ 532199); В. С. javanica, one of the paralectotypes (ZMZ 532200); С. С. javanica, Lombok (ZMB
Moll.75535); D. C. fluminea, China, lectotype (UZMC). Scale bars = 5 mm.

10 GLAUBRECHT ET AL.

FIG. 3. Anatomy of Corbicula fluminea, Argentina (MLP 5329) (A-C) and C. javanica, Lombok (ZMB
Moll. 75535) (D-F): A. Habitus of soft body; В. Siphons from inside; С, D. Siphons from outside; Е.
Section of siphons; F. Marginal mantle papillae. Scale bars = 1 mm.

FRESHWATER CORBICULIDAE FROM INDONESIA 11

TABLE 2. Morphometric indices (mean and standard deviation provided) calculated from measure-
ments of shells and siphon structure of the Corbicula species studied: L - shell length; H - shell
height; W - shell thickness (two valves); Ц - distance from beak to anterior end; НН - hinge plate
length; А - diameter of the posterior adductor scar; $ - breadth of siphons. Note that for the last two
taxa only range is given for siphons.

No. of
measured
specimens
Species (dry/wet) H/L W/H U/L HH/H A/L S/L

C. fluminea (lectotype) 1 0.93 0.80 0.48 0.100 0.14 not

measured

C. javanica 5/0 0.83 0,75 0.43 0.059 0.14 not
ZMZ 532199, 532200 +0.059 +0.066 +0017 +0.005 +0.012 measured
(type lot)

C. javanica 3/0 0.82 0.81 0.39 0.057 0.13 not
ZMB 75535 +0.035 +0.022 +0021 +0006 +0.009 measured

C. javanica 10/5 0.89 0:73 0.42 0.069 0.12 0.12
ZMB 106459 +0.041 +0.020 +0.018 +0.004 +0.013 +0.010

C. moltkiana 10/2 0.78 0.68 0.47 0.080 0.14 0.24
ZMB 54370, 103058 +0:056 =0.020 -+0.022 20.016 °=z0.007 +0.05

C. moltkiana SMF 5994 10/0 0.95 03 0.46 0.099 0.18 not
(paratypes of C. +0.049 +0.065 +0.047 +0.007 +0.020 measured
sumatrana)

C. moltkiana 6/0 0.94 0.80 0.44 0.111 0.18 not
ZMB 170000-01 +0.126 +0.033 +0.069 +0.016 +0.043 measured
(syntypes of С. lacustris)

C. moltkiana 5/4 0.91 0.69 0.47 0.079 0:17 0.18
ZMB 103024 +0.029 0.0039 +0.040 +0.004 +0.015 +0:013

C. moltkiana 7/4 0.84 0.69 0.50 0.073 not 0.19
ZMB 103032 +0.049 +0.039 +0.032 +0.008 measured +0.030

C. linduensis 10/4 0.77 0.63 0.49 0.081 0.14 0.18
ZMB 103016 +0,022 =0.039 =0.029 _=0.007 +0015 +0.033

C. matannensis 10/6 0.84 0.64 0.42 0.098 0.15 0.24
ZMB 103002 +0.043 +0.035 +0.022 +0.005 +0.010 +0.015

C. matannensis 9/7 0.83 0.59 0.39 0.96 0.13 0.26
ZMB 103009 30.045. - 0.041... 20031 +0013 =0.017 +0.034

C. matannensis 4 0.96 0.70 0.42 0.111 0.15 not
ZMB 103006 +0.050 +0.090 +0.052 +0.016 +0.016 measured

C. loehensis 5 0.88 0.68 0.44 0.078 0.14 not
ZMB 103010 +0.028 +0.034 +0.011 +0.015 +0.023 measured

С. loehensis 8/3 0.88 0.68 0.47 0.075 0.12 01832
ZMB 103011 +0.040 +0.026 +0.046 +0.011 +0.010 0.032

C. loehensis 3/2 0.87 0.60 0.44 0.070 0.12 0.27-0.33
ZMB 103033 +0.047 +0.040 +0.026 +0.014 +0.004

С. possoensis 10/7 0.93 0.65 0.41 0.115 0.23 0.32
ZMB 103028 +0028 =0.033 +0.033 +0:007 0.019 +0.022

С. possoensis 3/2 0.89 0.75 0.31 0.120 0.21 0.30-0.31
ZMB190024 +0.070 +0.037 +0.010 +0.012 +0.022

A A Е Е A Е НЕЕ Е
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papillae. Black pigment concentrated in rings rings. Siphonal muscles strong, arranged in
internally at base of both siphons (Fig. ЗЕ); broad bands. Papillae on outer surface of

outer surface of siphons white or with pale presiphonal suture arranged in two rows (Fig.

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FRESHWATER CORBICULIDAE FROM INDONESIA 13

3D). Marginal mantle papillae well developed,
densely arranged (Fig. 3F). Radial mantle
muscles strong, arranged in band, individual
bundles not separated.

Reproductive Biology: Only eggs were found in
all dissected specimens, therefore sperm
morphology remained unknown. One speci-
men from Java contained in its inner
demibranchs several hundred small larvae
(0.13-0.15 mm long) with uncalcified shells.
Since these larvae were apparently not fully
developed, the final size of released young
remains unknown. Other studied specimens
were not brooding.

Distribution and Ecology: According to
Djajasasmita (1977), this taxon is widely dis-
tributed in Southeast Asia, from the Malay
Peninsula to Timor and Aru Islands, as well
as to the Philippines. It occurs in several dif-
ferent types of lentic and lotic habitats, that
is, rivers, creeks and irrigation canals as well
as lakes and ponds (Djajasasmita, 1977).

Remarks: This taxon is recognizable as a dis-
tinct morphotype, differing from the typical
form of C. fluminea in its inequilateral shell,
anterior shift of beaks, and relatively narrow
hinge plate (Fig. 2, Table 3). Anatomical
characters described here well agree with
those reported for C. fluminea (Britton &
Morton, 1979; Harada & Nishino, 1995) and
observed in specimens of the latter species
studied here for comparison (Fig. ЗА-С),
with the only difference of a darker pigmen-
tation of the outer surface of siphons. In
fact, we anticipate that C. javanica might
probably just be a variety of C. fluminea.
However, this supposition should be con-
firmed by investigating the reproductive biol-
ogy and molecular genetics of the relevant
forms in more detail.

Corbicula moltkiana Prime, 1878
Figs. 4-6, 7A, В, 8A, В

Corbicula moltkiana Prime, 1878: 43, pl. 2, fig.
2a, b, с; Prashad, 1930: 200, pl. 25, figs. 17-
22; Djajasasmita, 1977: 4.

Corbicula sumatrana Clessin, 1887: 78, pl. 3,
fig. 7; Prashad, 1930: 198, pl. 25, figs. 1-8;
Djajasasmita, 1977: 7.

Corbicula verbecki Clessin, 1887: 79.

Corbicula moltkeana [sic] Prime — Martens,
1897: 111, pl. 7, figs. 1-6.

Corbicula lacustris Martens, 1897: 118, pl. 7,
figs. 20-24.

Type Locality: “Sumatra”, not exactly specified.

Type Material: Type specimens of C. moltkiana
Prime located in UZMC are presumed to be
lost (T. Schiotte, pers. comm.). Presumably
syntypes (although catalogued as paratypes)
of С. sumatrana Clessin, “Lake Singkarah”
[sic!], leg. Verbeck 1880 (SMF 5994, 5995).
Holotype of C. verbecki, Clessin, same local-
ity data (SMF, not numbered). Syntypes of C.
lacustris Martens, Lake Singkarah, leg. We-
ber (ZMB Moll. 170000, 170001, 170002).

Other Material Examined: Sumatra: Lake
Manindjau (ZMB Moll. 54370w, 103058,
103059; leg. Max Weber; originally identified
as C. moltkiana by E. v. Martens); Lake
Manindjau, at shore near Manindjau (ZMB
Moll. 103024, 103034; leg. Köhler & Schütt ,
April 2000); Lake Singkarak (0°32.89'S,
100°31.92’Е) (ZMB Moll. 103025, 103032;
leg. Köhler & Schütt, April 2000) (Fig. 1).

Taxonomic Remarks: We have at our disposal
one alcohol lot and two dry lots coming from
the collection of Eduard von Martens, bearing
his identification. Obviously, the specimens
are those cited in Martens' (1897) monograph
as “moltkeana” (evidently a misspelling of
Prime's original name). While characters of
these specimens (Fig. 4E) well agree with
the original description and figures of Prime
(1878), we retain the original identificaton as
C. moltkiana. Comparison of type speci-
mens of the three taxa described from Lake
Singkarak, namely C. sumatrana Clessin
(Fig. 4B), C. verbecki Clessin, and C.
lacustre Martens (Fig. 4C), as well as recent
collections of Corbicula from this lake (Fig.
4D) with the available figures and material of
C. moltkiana did not provide convincing dis-
tinctive characters and, thus, suggests that
these taxa are conspecific.

Description

Shell: Variable in shape, but usually triangular
or trapezoid, from high to markedly elongated,
compressed. Posterior margin somewhat
truncate, with characteristic posteroventral
right or obtuse angle. Beaks narrow, central
or somewhat shifted anteriorly, not protrud-
ing. Periostracum yellow, dark green or dark
brown to black, shiny. Internal shell surface
from white to dark purple. Concentric sculp-
ture of variable spacing (9-12 ribs per 1 cm),
ribs sharp. Hinge plate moderately broadened
to broad (Table 2, 3). Cardinal teeth well de-
veloped; anterior lateral teeth thick, straight or
slightly arched. Up to 30 mm long.

14 GLAUBRECHT ET AL.

FIG. 4. Shells of Corbicula moltkiana from Sumatra with synonyms as suggested in the present paper:
A. Original figure from Prime (1878), locality unknown (not to scale); В. One of the syntypes of С.
sumatrana, Lake Singkarak (SMF 5995); С. One of the syntypes of С. lacustre, Lake Singkarak
(ZMB Moll. 170000); D. С. moltkiana, Lake Singkarak (ZMB Moll. 103024); E. C. moltkiana, Lake
Manindjau (ZMB Moll. 54370). Scale bar = 5 mm.

FRESHWATER CORBICULIDAE FROM INDONESIA 19

B

FIG. 5. Anatomy of Corbicula moltkiana, Sumatra, Lake Manindjau: А. Habitus of soft body; В. Siphons
from outside; C-D. Papillae of inhalant siphon (С - ZMB Moll. 54370, D - ZMB Moll. 103034); EF.
Marginal mantle papillae (E - ZMB Moll. 54370, F - ZMB Moll. 103034). Scale bars = 1 mm.

Anatomy: Adductors small, oval (Fig. 5A, Table
2). Presiphonal suture longer than aperture of
inhalant siphon. Siphons conical, narrow in
small specimens and rather broad in full
grown ones, with thick walls and circular or
oval apertures; number of inhalant siphon pa-

pillae varies from 30 to about 80, arranged in
one or two rows (Figs. 5B-D, 6B), in largest
specimens additional row of small papillae
may appear. Black pigment concentrated in
rings at base of papillae in both siphons, but
patches of pigment also seen around siphons

16 GLAUBRECHT ET AL.

FIG. 6. Corbicula moltkiana, Sumatra, Lake
Singkarak (ZMB Moll. 103025), view of mantle
and gill: А. Siphons from inside, В. Section of
siphons; С. Gill with incubated larvae, from
inside. Scale bars = 1 mm.

(Fig. 6A, B), in some specimens internal sur-
face of siphons almost entirely pigmented.
Some papillae of inhalant siphon with dark
rings. In specimens from Martens' collection
pattern of pigmentation indistinguishable.
Siphonal muscles strong, arranged in broad
bands (Fig. 6A). Papillae on outer surface of
presiphonal suture arranged in single row,
sometimes in two rows. Marginal mantle pa-
pillae well developed, densely arranged. Ra-
dial mantle muscles strong, arranged in
band, their bundles indistinguishable in
smaller specimens but distinct in large ani-
mals (Fig. 7A, B).

Reproductive Biology: Gonads of the dis-
sected animals contained either sperm or
eggs. Spermatozoa (Fig. ЗА-В) monoflagel-
late; head length 11.5 + 0.58 um (n = 7).
Eight out of 15 specimens collected in Lake
Singkarak (ZMB Moll. 103026) were brooding
and their inner demibranchs contained sev-
eral hundred larvae of approximately equal
size (0.25-0.3 mm long). The only gravid

specimen found in Lake Manindjau (ZMB
Moll. 54370) contained larger larvae (about
0.35 mm long).

Distribution and ecology: To date known from

several localities on Sumatra and the Malay
Peninsula, according to Djajasasmita (1977).
Material revised by this study was collected in
lakes only (Fig. 1A), but the species was also
recorded in rivers and ditches (Djajasasmita,
1977).

Remarks: This species is remarkably variable

both in shell and anatomical characters.
Specimens from Lake Manindjau differ from
those collected in Lake Singkarak (type local-
ity of C. sumatrana and С. lacustris) in their
narrower hinge plate and densely arranged
ribs. Moreover, the old lots from Lake
Manindjau are distinguished from the new
collections from the same lake by their elon-
gated shells, broadened siphons, and weaker
mantle muscles; anatomical differences
might be associated with the larger size of
animals collected by Weber, in comparison
with those from our collections (26-28 and
15-18 mm, respectively). Furthermore, the
purple form found in both sampled lakes
(stored separately as ZMB Moll. 103032,
103034) alongside the yellow one (ZMB Moll.
103024, 103025) showed also somewhat
more delicate ribs and darker internal pig-
mentation of siphons. The form described as
C. lacustre is characterized by smaller size
(up to 18 mm), high, thick-walled shell, and
especially coarse sculpture; it is probably a
deep water variety of the same species
(Djajasasmita, 1977). While differences be-
tween the extreme forms from the localities
discussed here are rather pronounced (Fig.
4), intermediate forms could be also found.
All forms assigned here to С. moltkiana can
be recognized by their angulate compressed
shell, narrow not protruding beak, relatively
broad hinge plate and sharp ribs. These
characters distinguish it from the widely dis-
tributed Southeast Asian taxon, C. javanica
(Table 3), as well as from the typical form of
C. fluminea. The differences from C. javanica
in shell elongation and convexity, position of
beaks, and relative breadth of hinge plate
(Table 2) were significant at p < 0.05. Note-
worthy, one specimen of C. javanica was
found in Weber's lot from Lake Manindjau
(ZMB Moll. 103.058), being well distinguish-
able from the sympatric С. moltkiana by its
protruding beak, narrow hinge and widely
spaced smoothened ribs.

FRESHWATER CORBICULIDAE FROM INDONESIA 17

FIG. 7. Mantle musculature of Indonesian Corbicula: A. С. moltkiana, Sumatra, Lake Singkarak (ZMB
Moll. 103024); В. С. moltkiana, Sumatra, Lake Manindjau (7МВ Moll. 54370); С. С. matannensis,
Sulawesi, Lake Matano (ZMB Moll. 103002); D. С. possoensis, Sulawesi, Lake Poso (ZMB Moll.
103028). Scale bar = 1 mm. mc - concentric musculature, mr - radial musculature.

Large specimens of С. moltkiana are similar
in some anatomical characters (form of
siphons and patterns of mantle musculature)
to the lacustrine taxa from Sulawesi (С.
matannensis and C. loehensis). Dark rings
of pigment seen at base of both siphons in C.
moltkiana are similar to those of С. fluminea
(Britton & Morton, 1979; Harada & Nishino,
1995), but the internal pigmentation of siphons
in the former taxon 1$ generally more intense
than in the latter.

Sperm and eggs were not found in the same
animal. We understand as functional males
those animals producing sperm, while we did
not find the relatively large eggs when in-
specting the gonad. Although the exact ex-
pression of sexuality in this species remains
to be verified by detailed seasonal observa-
tions and histological study of gonads, we
anticipate that C. moltkiana is not a simulta-
neous hermaphrodite. In this aspect, it might

be similar to С. sandai from Lake Biwa, but
apparently differs from the other freshwater
taxa studied so far (Konishi et al., 1998;
Byrne et al., 2000; Qiu et al., 2001). It should
be stressed again here, that our finding of
monoflagellate sperm in this taxon provides
an indication of meiosis and sexual reproduc-
tion (Siripattrawan et al., 2000).

Species from Sulawesi

Corbicula linduensis Bollinger, 1914
Figs. 8C, 9A, В, 10, 11

Corbicula moltkiana var. linduensis Bollinger,
1914: 575, pl.-18; fig. 12.

Corbicula linduensis Bollinger — Djajasasmita
1975: 84, fig. 1.

Corbicula lindoensis
Djajasasmita, 1977: 4.

Type Locality: Lake Lindu, Sulawesi.

[sic!] Bollinger —

18 GLAUBRECHT ET AL.

FIG. 8. Sperm morphology (SEM) of Indonesian Corbicula exhibiting monoflagellate spermatozoa: A,
B. C. moltkiana (ZMB Moll. 103024); C. C. linduensis (ZMB Moll. 103016); D. C. matannensis (ZMB
Moll. 103003); Е. С. loehensis (ZMB Moll. 103010); Е, С. С. possoensis (ZMB Moll. 103028). Scale
bars = 2 ит.

Type Material: Syntypes deposited at the Natu-
ral History Museum Basel (Switzerland) are
reported to be lost (Ц. Wüest, pers. comm.).

Material Examined: Sulawesi: river at the road
from Palu to Gimpu, basin of the Palu River
(01°13.75'$, 119%56.69E) (ZMB Moll.
103016w; leg. Brinkmann 8 Rintelen, March
2000) (Fig. 1).

Description

Shell: Oval, usually markedly elongated, with
rounded posteroventral angle (Fig. 9A, B).

Periostracum yellow to brown. Internal col-
oration white or purple. Beaks central, narrow
and not protruding. Surface sculpture with
widely spaced, low ribs (n = 10-12 ribs per
1 cm); pronounced folds of periostracum no-
ticeable between ribs. Hinge plate moderately
broad. Cardinal teeth delicate; lateral teeth
relatively short, straight. Largest specimen
available for this study was 17 mm long. Ac-
cording to the literature, the specimens from
Lake Lindu were on average 23 mm long.
Anatomy: Adductors small, round (Fig. 10A,
Table 2). Presiphonal suture not elongated,

FRESHWATER CORBICULIDAE FROM INDONESIA 19

FIG. 9. Shells of Corbicula taxa from Sulawesi (right valve from outside, left valve from inside): А, В.
Corbicula linduensis, Palu River system (ZMB Moll. 103016); С. Syntype of С. matannensis, Lake
Matano (ZMB Moll. 50799); D. C. matannensis, Lake Matano (ZMB Moll. 103002); E. C. matannensis,
Lake Mahalona (ZMB Moll. 103009); Е. С. matannensis, Lake Towuti (ZMB Moll. 103006); С. Juvenile
specimen of the same species from Lake Towuti (ZMB Moll. 103007). Scale bars = 5 mm.

20

GLAUBRECHT ETAL.

|

№

K

O

3

O
SS

de _

FIG. 10. Anatomy of Corbicula linduensis, Sulawesi (ZMB Moll. 103016): A. Habitus of soft body; B.
Siphons from outside; C. Inhalant siphon papillae; D. Marginal mantle papillae; E. Clutch of juveniles
from a gill. Scale bars = 1 тт (С and D to same scale).

length equal to diameter of inhalant siphon.
Siphons conical, thin-walled, apertures circu-
lar or short oval, both somewhat broadened;
inhalant siphon with about 50 papillae ar-
ranged in two rows (external row with shorter
papillae) (Figs. 10B, C, 11A, B). Internal pig-
mentation of siphons weak, but pale internal
ring sometimes noticeable around exhalant
siphon. Papillae not pigmented. Siphonal
muscles rather strong, arranged in broad
bands. Papillae on outer surface of
presiphonal suture arranged in two uneven
rows (Fig. 10B). Marginal mantle papillae well

developed, densely spaced (Fig. 10D). Ra-
dial mantle muscles strong, arranged in
band, with only anterior bundles distinct,
separated from each other.

Reproductive Biology: Gonads of the dissected

animals contained either sperm or eggs.
Spermatozoa (Fig. 8C) monoflagellate, rela-
tively small (head length 8.8 + 0.27 um, n=
7). Eight of 12 studied specimens were
brooding. Some of them contained several
hundred larvae of usual size for Corbicula.
However, most of the gravid animals carried
in each inner demibranch 10 to 35 juveniles

FRESHWATER CORBICULIDAE FROM INDONESIA 21

FIG. 11. View of mantle and gills of Corbicula
linduensis, Sulawesi (ZMB Moll. 103016): A.
Siphons from inside; B. Section of siphons; C.
Gill with incubated larvae, from inside. Scale
bars = 1mm.

with up to 1.5 mm long shells (Fig. 10E,
11C).

Distribution and Ecology: Apparently restricted
to the lake and river of the Palu basin. To date
only known from lacustrine habitats
(Djajasasmita, 1975, 1977). Our recent find-
ings extend the known distribution from the
lake proper to the Palu valley, though, since
the specimens studied here were collected in
a small river, on muddy bottom with vegeta-
tion.

Remarks: This species is similar in its elon-
gated shell to some forms of C. moltkiana.
However, the sculpture (smoothened ribs)
and siphonal characters of C. linduensis are
more similar to that in C. fluminea and C.
javanica (Table 3). Sperm and eggs were not
found in the same animal. Taking into ac-
count the monoflagellate type of sperm, we
suggest that this species is meiotic,
similarily to С. moltkiana. It is distinguished
from the other taxa studied by the significantly
smaller spermatozoa (p < 0.01, t-test). Char-
acteristics of its brooding process, i.e. having

the largest incubated juveniles known, are
unique among Corbicula species. According
to Djajasasmita (1975) the population from
Lake Lindu has dramatically decreased since
1950, thus rendering conservation strategy
for this unique bivalve an urgent task.

Corbicula matannensis P. Sarasin &
F. Sarasin, 1898
Figs. 7C, 80, 9C-G, 12, 13

Corbicula matannensis P. Sarasin & F. Sarasin,
1898: 92, pl. 11, figs. 158—160; Kruimel, 1913:
231; Djajasasmita, 1975: 84, fig. 3;
Djajasasmita, 1977: 4.

Corbicula towutensis Kruimel, 1913: 231, pl. 4,
fig. 3.

Corbicula mahalonensis Kruimel, 1913: 231, pl.
4, fig. 4.

Corbicula subplanata (partim) Martens —
Prashad, 1930: 203, pl. 26, figs. 7-9, 13.

Type Locality: Lake Matano, Sulawesi.

Type Material: Syntype of C. matannensis P.
Sarasin & F. Sarasin (ZMB 50799), from Lake
Matano (Fig. 11A). Syntypes of C. towutensis
Kruimel and C. mahalonensis Kruimel (ZMA)
[vidi].

Other Material Examined: Sulawesi: Lake
Matano: $ shore, small bay (02°28.04'S,
121°14.04'E) (ZMB Moll. 103000w; leg.
Glaubrecht & Rintelen, 15 August 1999); S
shore (02°28.44'S, 121°15.78'E) (ZMB Moll.
103001w; leg. Glaubrecht & Rintelen, 15 Au-
gust 1999); E bay, at outlet of Petea River
(02°32.06’S, 121°28.50E) (ZMB Moll.
103002w; leg. Glaubrecht & Rintelen, 16 Au-
gust 1999); $ shore, at Salonsa (02°30.49'S,
121°19.96'Е) (ZMB Moll. 103003w; leg.
Glaubrecht & Rintelen, August 1999); NW
shore (2°26.01’S, 121°13.03’E) (ZMB Moll.
103004w; leg. Glaubrecht & Rintelen, 11-12
August 1999). Lake Towuti: W shore, bay at
outlet of Larona River (02°46.09'S,
121°21.57°E) (ZMB Moll. 103006; leg.
Glaubrecht & Rintelen, 18 August 1999); N
shore, swamp W of Mahalona inlet, lake side
of sand-bar (02°40’S, 121°31.8’E) (ZMB Moll.
103007w; leg. Bouchet, 1991). Lake
Mahalona: at mouth of outlet (02°36.88'S,
121°30.98’E) (ZMB Moll. 103008; leg.
Glaubrecht 8 Rintelen, 24 August 1999); E
shore, cape (02°35.58’S, 121°30.68'E) (ZMB
Moll. 103009w; leg. Glaubrecht & Rintelen,
24 August 1999) (Fig. 1).

Taxonomic Remarks: Comparison of the avail-
able material of C. matannensis to the

22

GLAUBRECHTETAL.

es

FIG. 12. Anatomy of Corbicula matannensis from Sulawesi, Lake Matano (ZMB Moll. 103002): A.
Habitus of soft body; B. Siphons from outside; C. Papillae of inhalant siphon; D. Marginal mantle
papillae. Scale bars = 1 mm (C and D to the same scale).

syntypes of C. subplanata Martens, 1897
(ZMB Moll. 103017, from Minralang River, near
Tempe, Sulawesi), confirmed the differences
in patterns of sculpture between these taxa
mentioned by Djajasasmita (1975). However,
the status of the latter taxon remains unclear,
until its soft parts are available for anatomical
and molecular study. Given that the differ-
ences between the respective taxa concern
only shell proportions, which proved to be
variable in Corbicula (Morton 1979, 1986;
Harada & Nishino 1995), we accept here the
synonymization of C. mahalonensis and C.
towutensis under С. matannensis as sug-
gested by Djajasasmita (1975).

Description

Shell: Circular in young specimens and tetrago-
nal in fully grown ones, with obtuse postero-

ventral angle (Fig. 9C-G). Periostracum from
pale yellow to dark violet in small shells and
usually black in large ones, dull. Internal-col-
oration from white to deep purple, usually
darker on outer margin. Beaks central in
young shells but markedly shifted forward in
adults, narrow, not protruding. Concentric
sculpture pronounced, densely spaced (15—
20 ribs per 10 mm), ribs sharp. Hinge plate
usually broad; cardinal teeth well developed;
lateral teeth straight. Length up to 32.5 mm
(syntype from Lake Matano).

Anatomy: Adductors small, oval (Fig. 12A,

Table 2). Presiphonal suture relatively long.
Siphons cylindrical, thin-walled, with broad
slit-like apertures; inhalant siphon somewhat
broader than exhalant, with 55 to 70 papillae
arranged in two rows (internal row of long, ex-
ternal of short papillae) (Figs. 12B, С, 13A-
C). Both inner and outer surface of siphons

FRESHWATER CORBICULIDAE FROM INDONESIA 23

FIG. 13. Corbicula matannensis from Sulawesi, view of mantle and gill: A. Siphons from inside, Lake
Matano (ZMB Moll. 103002); B. Section of siphons from specimen of the same locality; C. Section of
siphons, Lake Towuti (ZMB Moll. 103006); D, E. Gill with incubated larvae, Lake Matano (ZMB Moll.
103002). Scale bars = 1 mm.

and papillae densely pigmented; with dark
median internal stripe along presiphonal su-
ture. Siphonal muscles weak, forming two
narrow bands and dispersed fibers below
and above these bands (Figs. 12A, 13A). Ar-
rangement of papillae on outer surface of
presiphonal suture variable: sometimes ar-
ranged in several rows, sometimes in single
row or dispersed (Fig. 12B). Marginal mantle
papillae small, widely spaced (Fig. 12D). Ra-
dial mantle muscles weak, forming separate
bundles (Fig. 7C).

Reproductive Biology: Gonads of the dis-
sected animals contained either sperm or
eggs. Spermatozoa monoflagellate, head
length 11.1 + 0.30 ит (п = 7). Brooding speci-
mens were found in three samples (two from
Lake Matano and one from Lake Mahalona),

representing a total of eight gravid specimens
out of 30 specimens dissected. Numerous
larvae (0.30-0.33 mm long) were located
only in inner demibranchs (Fig. 13D, E).

Distribution and Ecology: Occurring in the

larger lakes of the Май! system, that is,
Lake Matano, Mahalona and Towuti. This
species is known from lacustrine habitats
only so far.

Remarks: This species is distinguished from

C. subplanata, as described by Martens
(1897), by its sculpture. Corbicula
matannensis has much more densely placed
ribs, and they are narrower and more sharp;
because spacing between ribs in Corbicula
increases with age, equally sized specimens
should be compared. The latter taxon is also
characterized by rather peculiar anatomical

24 GLAUBRECHT ET AL.

FIG. 14. Shells of Corbicula from Sulawesi: А. С. loehensis, Lake Masapi (ZMB Moll. 103010); В. С.
loehensis, Lake Lontoa (ZMB Moll. 103005); C. Syntype of Corbicula possoensis, Lake Poso (ZMB
Moll. 50798; D. Corbicula possoensis, Lake Розо (ZMB Moll. 190024). Scale bars = 5 mm.

FRESHWATER CORBICULIDAE FROM INDONESIA 25

и о

—

NI

S

|

Le
SEO
7"

B

es

pod

20 Sn

u... а Ont
AT AIR

``
Фо

©
0 © ©

Ge

1 Teese

Pre. 2;

000

FIG. 15. Anatomy of Corbicula loehensis, Lake Masapi (ZMB Moll. 103010): A. Habitus of soft body;
B. Siphons from outside; C. Siphons from inside. Scale bars = 1 mm.

characters (broad siphons with strong inside
pigmentation, and widely spaced marginal pa-
pillae of mantle). One sample from Lake
Towuti contained small shells (up to 16 mm
long) with delicate sculpture (Fig. 9G) that
show some similarity to C. loehensis (see be-
low). However, none of these specimens were
brooding, therefore, we conclude that all of
them were young and could represent C.
matannensis, in which juveniles have more
delicate sculpture than adults. Noteworthy, the
doubtful specimens from Towuti were similar
to young Corbicula from Lake Matano and dis-

tinguished from C. loehensis of Lake Masapi
and Lake Lontoa (older name: Wawontoa) by
their broad hinge plate. The shells from Lake
Matano are the largest (length up to 35 mm)
characterised by well developed sculpture (n =
14-16 ribs per 10 mm in the middle and
about 12 at the outer margin, near the beaks
ribs are fine) and strong hinge. The speci-
mens from Lake Towuti are very similar to that
from Lake Matano, but they never reach such
a large size (largest specimen found was 23
mm long) and are more round and convex
(Fig. 11D, Table 2). This form 1$ similar to

26 GLAUBRECHT ETAL.

FIG. 16. Corbicula loehensis and С. possoensis, view of siphons and gills: A. С. loehensis, Lake
Lontoa (ZMB Moll. 103005), section of siphons; В. С. possoensis, (ZMB Moll. 190024), siphons from
inside; C. С. possoensis (ZMB Moll. 190024), section of siphons; D. С. loehensis, Lake Masapi (ZMB
Moll. 103010), gill with incubated larvae from outside; Е. С. possoensis (ZMB Moll. 190024), anterior
portion of gill from inside (inner demibranch partly removed). Scale bars = 1mm.

small specimens of С. possoensis, but can Corbicula masapensis Kruimel, 1913: 232, pl.

be readily distinguished by the size of the ad- Ata

ductors (see below). The corbiculids from Corbicula subplanata Martens (part) —
Lake Mahalona are more elongated (espe- Prashad, 1930: 203, pl. 26, figs. 11-12.
cially posterior part) and flat (see Fig. 9E, Type Locality: SE shore of Loeha Island, Lake
Table 2). However, variation within one and Towuti.

the same lake (Matano) 1$ also considerable Type Material: syntypes ZMA [vidi].

(Fig. 11C, D, Table 2). Other Material Examined: Sulawesi: Lake

In features of reproductive biology (1.е., in
presence of sperm and eggs in different ani-
mals, sperm morphology and mode of
brooding) this species is similar to С.
moltkiana from Sumatra.

Corbicula loehensis Kruimel, 1913
Figs. 8E, 14A,B, 15, 16A,D

Corbicula loehensis Kruimel, 1913: 232, pl. 4,
figs. 2, 3; Djajasasmita, 1975: 84, fig. 3;
Djajasasmita, 1977: 4.

Masapi: S shore (02%50.84'S, 121°21.09'E)
(ZMB Moll. 103011w; leg. Brinkmann 8
Rintelen, 30 March 2000); Lake Masapi, local-
ity not specified (ZMB Moll. 103010w; leg.
Bouchet, 1991). Lake Lontoa (= Wawontoa):
SW shore (02°39.6’$, 121°44.8’E) (ZMB
Moll. 103005w; leg. Bouchet, October 1991);
W shore (02%39.90'S, 121°43.46'E) (ZMB
Moll. 103033w; leg. Brinkmann 4 Rintelen,
Mar 2000) (Fig. 1).

Taxonomic remarks: Djajasasmita (1975)
synonymised С. masapensis with С.

FRESHWATER CORBICULIDAE FROM INDONESIA a

loehensis. This point of view is tentatively ac-
cepted here, but we suggest to study Lake
Towuti populations carefully before a final de-
cision on the taxonomic status of the forms
under consideration is possible. Identity of
the form present in Lake Lontoa to C.
loehensis was first shown by Djajasasmita
(1975).

Description

Shell: Round to ovate, with somewhat obtuse
posterior edge (Fig. 14A, B). Periostracum
yellow in specimens from Lake Masapi and
dark violet in those from Lake Lontoa, with
silky glitter. Internal coloration white and
purple, respectively. Beaks subcentral.
Sculpture very fine, formed by delicate ribs
(30-40 per 1 cm). Hinge plate moderately
broad; cardinal teeth delicate; laterals rela-
tively short, straight. The largest examined
specimen was 18 mm long, but according to
literature data the species may reach a
length of 25 mm (Djajasasmita, 1975).
Anatomy: Adductors small, oval (Fig. 15A,
Table 2). Siphons cylindrical, rather thin-
walled, with broad oval apertures, inhalant si-
phon somewhat broader than exhalant, with
about 50 papillae arranged in two rows. Inter-
nal surface of siphons, papillae and
presiphonal suture usually strongly pig-
mented (Figs. 15B, C, 16A). Marginal mantle
papillae small, widely spaced. Mantle muscu-
lature weak, muscle bundles well distin-
guishable and separated.

Reproductive biology: Gonads of the dissected
animals contained either sperm or eggs.
Spermatozoa monoflagellate, head length 9.1
+ 0.21 ит (п = 7). The largest of the studied
specimens was brooding. Larvae located in
its inner demibranchs (Fig. 16D) were about
0.24 mm.

Distribution: Recorded ashore in Lake Towuti
(Kruimel, 1913; Djajasasmita, 1975) and its
satellite lakes Masapi and Lontoa.

Remarks: Anatomical characters of C.
loehensis and C. matannensis are similar
(Table 3), as well as are juvenile shells. How-
ever, the characters of juvenile shells in the
former species (very fine sculpture and deli-
cate hinge) are also retained in more ad-
vanced individuals of the latter species,
which might indicate distinct developmental
trends. Differences in coloration between the
shells from Lakes Masapi and Lontoa are
also noteworthy.

This species is similar in features of repro-
ductive biology (structure of gonads, sperm
type and mode of brooding) to С. moltkiana
and C. matannensis, but differs in having sig-
nificantly smaller spermatozoa (р < 0.001).

Corbicula possoensis P. Sarasin & F.
Sarasin, 1898
Figs. 7D, 8F-G, 14C, D; 16B, С, E; 17

Corbicula possoensis P. Sarasin & F. Sarasin,
1898: 92, pl. 11, figs. 161-162; Kruimel, 1913:
231.

Corbicula subplanata Martens (part) -
Prashad, 1930: 203, pl. 26, figs. 10, 17-20.
Corbicula matannensis P. Sarasin & F. Sarasin
(part) - Djajasasmita, 1975: 84;

Djajasasmita, 1977: 4.

Type Locality: Lake Poso, Sulawesi.

Type Material: syntype ZMB Moll. 50798 (Fig.
14C).

Other Material Examined: Lake Розо: S shore,
Tentena, beach at Hotel “Pamona Indah”
(01%45.92'S, 120°38.42'E) (ZMB Moll.
190024w; leg. Glaubrecht 8 Rintelen, Sep-
tember 1999); at Hotel “Mulia” (02*%03.91'S,
120°41.50'E) (ZMB Moll. 103028w; leg.
Brinkmann & Rintelen 23 September 2000);
SW shore, Matawai (02°02.4’S, 120°38.1’E)
(ZMB Moll. 103012w; leg. Bouchet, Septem-
ber 1991); Е shore, Tolambo Bay (01°59.8’S,
120°42.1'E) (ZMB Moll. 103013w; leg.
Bouchet, 1991); М shore, Saluopa
(01*46.6'S, 120°32.9'E) (ZMB Moll. 103014w;
leg. Bouchet, 1991) (Fig. 1).

Description

Shell: Triangular or short tetragonal, often with
blunt keel at posterior end (Fig. 14C, D).
Periostracum dark violet to black, some-
times yellowish in very young specimens,
shiny. Internal coloration purple. Posterior
margin truncate (diagnostic feature). Beaks
markedly shifted forward, in small specimens
relatively narrow while in large ones this can
not be seen because shells are eroded.
Hinge plate relatively broad, even in young
specimens. Cardinal teeth well developed;
lateral teeth somewhat shortened, straight.
Sculpture: 25-30 delicate ribs per 1 cm.

Anatomy: Adductors large, posterior larger

than anterior one (Figs. 14, 17A). Diameter of
posterior adductor more that '/, length of shell
(Table 2). Presiphonal suture relatively short,
not exceeding diameter of inhalant siphon.

28

GLAUBRECHT ETAL.

es

FIG. 17. Anatomy of Corbicula possoensis, Lake Poso: A. Habitus of soft body (ZMB Moll. 103014); В.
Siphons from outside (ZMB Moll. 190024); С. Papillae of inhalant (ZMB Moll. 190024); D. Marginal
mantle papillae (ZMB Moll. 190024). Scale bars = 1 mm (С and D to same scale).

Siphons broad, cylindrical, inhalant siphon
markedly broader than exhalant, both with
slit-like apertures; inhalant siphon with 60 to
70 papillae (depending on age), arranged in
one or two rows (papillae of outer row al-
ways relatively smaller) (Figs. 16B, C, 17A,
B). Both inner and outer surface of siphons
and papillae densely pigmented, presiphonal
suture also pigmented internally. Papillae on
the outer surface of presiphonal suture
scarce, unevenly arranged (Fig. 17B). Mar-
ginal mantle papillae small and widely
spaced (Fig. 17D). Mantle musculature
weak, muscle bundles separated, dispersed
(Fig. 7D).

Reproductive Biology: Gonads of the dis-
sected animals contained either sperm or
eggs. Spermatozoa monoflagellate, head
length 10.7 + 0.37 um (n = 6). Branchial incu-
bation was observed in two samples contain-

ing larger specimens (exceeding 18 mm); 6
of 15 dissected specimens were brooding
and contained larvae in both demibranchs of
each gill (Figs. 16E), one gravid specimen
had larvae in inner demibranch only, though.
Larvae were 0.26-0.30 mm long.

Distribution: Restricted to Lake Poso in Central

Sulawesi.

Remarks: Conchologically, this species 1$ dis-

tinguished from С. matannensis by its trun-
cate posterior edge, fine sculpture (more
than 20 ribs per 10 mm) and larger posterior
adductor scar (Table 3). Beaks in large shells
are somewhat broader and placed more an-
teriorly than in all previous species; also the
hinge plate is relatively higher. Corbicula pos-
soensis differs from С. matannensis also in
having broad siphons extending over about
'/, of the body length in the former and '/, in
the latter species. Differences in relative

FRESHWATER CORBICULIDAE FROM INDONESIA 29

height of hinge plate, adductor size, siphons
breadth and number of ribs were confirmed
by t-test (p < 0.001). These diagnostic char-
acters were consistent among all examined
specimens, which is, in concert with our mo-
lecular data, the argument against synony-
mization of С. possoensis with С. matannensis
as was earlier on suggested by Djajasasmita
(1975).

Corbicula possoensis differs from other stud-
ied congeners in characteristics of brooding,
because it is the only Corbicula species
known that incubates in both demibranchs,
instead of only the inner demibranch. Slight
but significant (p < 0.05) difference in sperm
size between this taxon and C. matannensis
is also noteworthy. Other reproductive fea-
tures of C. possoensis are similar to those
shown for the taxa from Sumatra and
Sulawesi (Table 3).

MOLECULAR PHYLOGENETICS

Of a total of 614 base pairs included in the fi-
nal alignment, 103 were parsimoniously infor-
mative. Heuristic search recovered 56 most
parsimonious trees of 368 steps (CI = 0.742,
RI = 0.685). The strict consensus tree (Fig. 18)
shows that all newly sequenced taxa included
in the present molecular study form a well зир-
ported monophyletic clade with those freshwa-
ter taxa of Corbicula studied earlier, with the
exception of C. madagascariensis. Within this
clade three distinct Indonesian taxa are sup-
ported in this study, occuring (i) on Sumatra,
identified as С. moltkiana, and (ii) on Sulawesi
with C. matannensis and C. loehensis from
the Malili lake system. In addition, two different
sequences with unresolved relationships were
obtained from specimens identified morphologi-
cally as C. possoensis from Lake Poso. Fur-
thermore, one other of the Indonesian taxa, C.
javanica, is shown in the parsimony analysis
as closely related to C. fluminea from Korea
and the North American Corbicula “form В”, а
clade that is also well supported. The second
sequence attributed to C. fluminea, originating
from material from Thailand, a sequence of C.
australis from Australian, and C. cf. fluminalis
from Israel is also clustering with this clade;
however, the bootstrap support for the joint
group 1$ weak.

The NJ tree (Fig. 19) is more resolved than
the consensus tree recovered by the maximum
parsimony analysis, but the clades supported

by high bootstrap values are basically the
same in both reconstructions. The two differ-
ent morphotypes of C. moltkiana from Lake
Manindjau cluster together, as well as the two
different morphotypes from Lake Singkarak, ir-
respective of the morphological similarity be-
tween the corresponding morphotypes found in
each of the two lakes; thus, the four samples
from Sumatra cluster according to geography
instead of morphology. The NJ analysis also
indicates an outstanding position of C.
moitkiana and a sister relationship between С.
loehensis and C. matannensis, although the
support of the relevant clades is below the 50%
level. Heterogeneity of C. possoensis is found
in this analysis as well.

Distance analyses show remarkable similar-
ity in СО! sequences (divergence levels not
exceeding 1%) between the samples from the
adjacent lakes Manindjau and Singkarak on
Sumatra, as well Matano and Mahalona on
Sulawesi (Table 4). The divergence between
sequences obtained from different samples in
Lake Matano was approximately of the same
level as the difference between the samples
from Matano and Mahalona. Only minor differ-
ences were recovered for C. loehensis from
Masapi and Lontoa, while divergence level be-
tween this taxon and C. matannensis reaches
3.5%. The distance between two sequences of
C. possoensis comprised 3.8%. Most of the
pairwise sequence divergence levels calcu-
lated for the freshwater Corbicula taxa by this
study did not exceed the level of 4.1% reported
by Siripattrawan et al. (2000); only C. moltkiana
showed greater distances, especially when
compared with C. possoensis (up to 5.7%) and
C. australis (up to 5.2%).

In conclusion, the molecular data agree with
the morphological comparisons presented
above in showing (i) distinctness of three
lacustrine taxa from Sumatra (C. moltkiana)
and Sulawesi (С. matannensis and С.
loehensis), respectively, (ii) distinctness of
these Indonesian taxa with respect to known
continental Asian lineages, and (iii) the relation-
ship of C. javanica and C. fluminea. However,
the results concerning С. possoensis are
equivocal, because no morphological charac-
ters correlated with the observed sequence
differences were found. Plotting sperm mor-
phology data on the trees based оп СО! se-
quence data suggests polyphyletic origin of the
biflagellate condition, since no close relation-
ship between the lineages sharing this state
could be found.

30 GLAUBRECHT ETAL.

Polymesoda caroliniana
Neocorbicula limosa
Corbicula japonica, Japan
C. madagascariensis, Madagascar
C. possoensis, Poso (1) |
Sulawesi
100 C. possoensis, Poso (2)
C. sandai, Biwa [| Japan
C. loehensis, Masapi
84 99 C. loehensis, Lontoa
C. matannensis, Matano (2) Sulawesi
73 C. matannensis, Matano (1)
51 C. matannensis, Mahalona (3)
C. leana, Japan

93

Е Corbicula "form А", USA Asia

. fluminea, China

. moltkiana, Singkarak (1)

. moltkiana, Singkarak (2) |
Sumatra
100 . moltkiana, Manindjau (3) 4

. moltkiana, Manindjau (4)

. australis, NSW N Ausiralia

ооо оо оо

. fluminea, Thailand

C. fluminalis?, Israel

C. javanica, Java Asia
C. fluminea, Korea

70
Corbicula "form В", USA

FIG. 18. Strict consensus of 56 maximum parsimony trees (368 steps, CI = 0.742, RI = 0.685)
obtained for the corbiculid COI extended dataset. The numbers below branches show bootstrap
support (if more than 50%).

FRESHWATER CORBICULIDAE FROM INDONESIA 31

Polymesoda caroliniana

Neocorbicula limosa

Corbicula japonica, Japan

C. madagascariensis, Madagascar
C. moltkiana, Singkarak (1)
C. moltkiana, Singkarak (2)

100 | + Sumatra

С. moltkiana, Manindjau (3)

511 С. moltkiana, Manindjau (4)

C. possoensis, Poso (2) | Sulawesi

Corbicula “form A", USA

100| С: /eana, Japan Asia

C. fluminea, China

C. australis, NSW [| Australia
C. fluminea, Thailand

С. fluminalis?, Israel

Corbicula "form В", USA Asia

C. javanica, Java

C. fluminea, Korea

C. sandai, Biwa [] Japan

C. possoensis, Poso (1)

C. loehensis, Masapi

C. loehensis, Lontoa
100

С. matannensis, Matano (2) re
77|- С. matannensis, Matano (1)
691 с. matannensis, Mahalona (3)

— 0.01 substitutions/site

FIG. 19. А neighbor joining tree obtained for the corbiculid СО! extended dataset. The numbers below
branches show bootstrap support (if more than 50%).

GLAUBRECHT ET AL.

32

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FRESHWATER CORBICULIDAE FROM INDONESIA 33

DISCUSSION
Taxonomy and Systematics

Previous phylogenetic investigations sug-
gested a common, monophyletic origin of all
Old World freshwater Corbicula, with the es-
tuarine C. japonica being the sister taxon
(Siripattrawan et al., 2000; Pfenninger et al.,
2002). This is confirmed in the present study
by the inclusion of new sequence data from In-
donesian taxa.

In an attempt to clarify the systematics of
these Indonesian corbiculids, mainly of those
species inhabiting Sumatra and Sulawesi, five
taxa were found to possess specific distinct-
ness and identity, based on shell morphology,
anatomy and the features of reproductive biol-
ogy (including brooding), as well as on molecu-
lar data. While the corbiculids studied from
Sumatra are identified as C. moltkiana, four
distinct taxa were identified on Sulawesi: С.
possoensis (with two distinct lineages revealed
in the NJ analysis) endemic to Lake Poso and
C. matannensis plus C. loehensis occurring in
the central Malili lake system. Suitable material
of the fourth species, С. linduensis from the
Lindu River, was not available for molecular in-
vestigation.

The species distinctness of C. javanica from
the island of Java remains doubtful in the ab-
sence of data on sperm morphology and, thus,
there is the possibility of clonality. Remarkable
is its close affinity to Asian lineages of C.
fluminea found in our study. However, the taxo-
nomic status of the latter is also problematic,
because it was shown to be an assemblage of
several clonal lineages, probably of different
origin, that also includes allochthonous popula-
tions introduced into North America and Eu-
rope (Siripattrawan et al., 2000; Pfenninger et
al., 2002). Therefore, any synonymizations in
this stage of investigation seem to be prema-
ture.

Corbicula taxonomy has historically been
plagued by a plethora of nominal taxa de-
scribed from numerous, at least partly
ecophenotypic shell morphs. The occurrence
of polyploidy, unisexual reproduction, hermaph-
roditism, and androgenesis recently reported
for certain Corbicula populations (see Introduc-
tion) suggest that variation observed in these
freshwater bivalvia could be the result of
clonality and not necessarily imply species level
differences, as discussed in Siripattrawan et
al. (2000). Because polyploidy is prevalent in

some freshwater molluscs, especially among
bivalves, it is assumed to have played an im-
portant role in shaping their diversity and can
pose significant challenges to reconstruct phy-
logenetic evolution also in Corbiculidae (Lee et
al., 2002).

While the determination of presence or ab-
sence of meiosis is a rather laborious task ap-
plicable only in adequately fixed material,
fortunately in Corbicula there is an exception-
ally convenient morphological marker to help
distinguish clonal forms, since, to the present
knowledge, studied ameiotic lineages all have
biflagellate spermatozoa (Komaru & Konishi,
1999; Siripattrawan et al., 2000; Qiu et al.,
2001). Our preliminary observations suggest
that Indonesian corbiculids reproduce sexually
and have monoflagellate spermatozoa (with the
exception of C. javanica, in which sperm
structure remains unknown). In this respect,
they appear to be similar to C. sandai from the
“ancient” Lake Biwa in Japan, which is known
to be diploid and reproduces sexually with
monoflagellate sperm (Hurukawa & Mitsumoto,
1953; Okamoto & Arimoto, 1986). Interestingly,
our NJ analysis revealed that С. sandai clus-
ters together with the Sulawesi clade, whereas
the topology of the MP tree is not resolved in
this respect.

Given the notorious variability of morphologi-
cal characters applied traditionally in Corbicula
taxonomy, which is also shown in this study,
and the incompleteness of data on reproduc-
tive biology as well as molecular genetics, any
final decision about the systematics and taxo-
nomic status of different forms (morphotypes)
distributed across Australasia is still not pos-
sible. Uncertainties remain in particular for the
Javanese form and the question concerning the
existence, distribution and identity of C.
fluminea. However, several of our systematic
conclusions based on the new data on Indone-
sian taxa are in agreement with some of those
already reached in previous revisions, mainly
those by Djajasasmita (1975, 1977).

Implications from Morphology

Generally, confirming the presence of en-
demic taxa in several lacustrine habitats on
Sumatra and Sulawesi, this study adds sup-
port to the recognition of the following taxa: C.
moltkiana, C. linduensis, C. possoensis, C.
matannensis and C. loehensis. The first of
these taxa recorded in the lakes on Sumatra is
similar to the widely distributed Asian C.

34 GLAUBRECHT ET AL.

fluminea (Britton & Morton, 1979; Morton, 1986;
Агацо et al., 1993; Chen et al., 1995; Harada &
Nishino, 1995; Komaru et al., 1997, 2000), in
anatomical characters and the features of
brooding; however, its species status is sup-
ported by sperm morphology and COI se-
quence data.

Taxonomic distinctness of Corbicula
linduensis from North Sulawesi, first suggested
by Djajasasmita (1975), is supported here by
characteristics of brooding (limited number of
large juveniles incubated in gills) not known in
any other Corbicula. However, in the absence
of molecular data for this taxon, its relation-
ships remain unknown. Noteworthy, brooding in
C. linduensis seems to be somewhat similar to
the South American Neocorbicula limosa, as
described by Ituarte (1994). Since the latter rep-
resents an independent lineage within the fam-
ily, judging from morphological study (Dreher
Mansur & Meier-Brook, 2000) and molecular
data (Siripattrawan et al., 2000; Figs. 18, 19),
its mode of reproduction has apparently
evolved independently.

The taxa inhabiting the Malili lake system on
Sulawesi differ from their congeners not only in
shell parameters, but also in anatomical char-
acters, for example in the form and pigmenta-
tion of siphons. Although diagnostic
applications of these characters in Corbicula is
hindered by the considerable intraspecific vari-
ability (see, for example, the description of C.
moltkiana), in this particular case anatomical
differences are supported by molecular data.
As shown in this study, the distinctness of C.
matannensis occurring in Lake Matano and
Lake Mahalona (connected through the Petea
River; Fig. 1) from C. loehensis inhabiting the
satellite lakes Masapi and Lontoa of Lake
Towuti (but both connected via separate river
systems) is in agreement with the taxonomy
used by Djajasasmita (1975). Unfortunately, no
molecular data are available to date for the lat-
ter species from Lake Towuti proper, where
both taxa possibly live sympatrically according
to Djajasasmita (1975).

The data on C. possoensis restricted to Lake
Poso are controversial. While morphological
observations show similarity of all available
lots, molecular data suggest their heterogene-
ity. Although Corbicula from Lake Poso appar-
ently needs further study, the outstanding
position of C. possoensis in relation to
corbiculids inhabiting the Malili lake system
found in the present study is consistent with
recent results on endemic pachychilid gastro-

pods from Lake Poso, which exhibit a similar
isolated position among the Tylomelania clade
in morphological and molecular phylogenies
(Rintelen & Glaubrecht, 1999, 2002; Rintelen et
al., submitted).

Some anatomical characters, for example
the broad cylindrical (fringe-like) form of si-
phons and the number and arrangement of the
exhalant siphon papillae, which are common in
Corbicula species inhabiting Lake Poso and the
Malili Lakes on Sulawesi were also reported for
the Japanese estuarine species C. japonica
(Harada & Nishino, 1995). However, the internal
coloration of siphons and papillae is remarkably
similar in all Sulawesi taxa but differ from that
of C. japonica, which is also very distinct
karyologically (Okamoto & Arimoto, 1986), in its
non-brooding reproduction (reviewed by
Morton, 1986) and its molecular genetics
(Siripattrawan et al., 2000). Therefore, any
similarity in form of siphons between the fresh-
water corbiculids in question and their probable
estuarine sister taxon are unlikely to be
synapomorphic.

Spermatozoan Morphology

The new data on sperm morphology shown
here for Indonesian taxa suggest that
monoflagellate spermatozoa are more com-
mon among freshwater Corbicula than as-
sumed in previous studies (Komaru 4 Konishi,
1996, 1999; Byrne et al., 2000; Siripattrawan et
al., 2000). Interestingly, the monoflagellate type
is known to occur in species inhabiting lacus-
trine habitats, such as, for example, Lake Biwa
in Japan (С. sandai), lakes Singkarak and
Manindjau on Sumatra (C. moltkiana), Lake
Poso (C. possoensis) and the Malili lake sys-
tem (C. matannensis, C. loehensis) on
Sulawesi.

In addition, while all Corbicula with biflagellate
spermatozoa are simultaneous hermaphro-
dites (Komaru 8 Konishi, 1996, 1999; Konishi
et al., 1998; Byrne et al., 2000), the Indonesian
corbiculids with monoflagellate sperm appar-
ently have a different expression of sexuality.
Since monoflagellate sperm is reported for the
gonochoric Corbicula sandal (Siripattrawan et
al., 2000, and literature cited therein), we hy-
pothesize that the Indonesian taxa also have
separate sexes.

However, sperm morphology of many riverine
corbiculids, especially those occurring on other
Sunda Islands, 1$ still not studied; therefore, it
is too early to judge on this habitat-sperm mor-

FRESHWATER CORBICULIDAE FROM INDONESIA 99

phology correlation. Our phylogenetic recon-
struction does also not reveal a close relation-
ship between lineages sharing the biflagellate
type of sperm, because the latter occurs in
clonal Corbicula within both Asian clades found
in the analyses (Figs. 18, 19).

The diversity of head size in spermatozoa of
Corbicula is remarkable, although no correla-
tion between size and the monoflagellate/bi-
flagellate type was found. The biflagellate
spermatozoa of the Chinese C. fluminea and
Japanese С. leana are distinguishable by their
large size of 16-25 um (Komaru 8 Konishi,
1996; Qiu et al., 2001), whereas biflagellate
spermatozoa of C. australis are relatively small
with 9.3 um on average (Byrne et al., 2000).
The latter are, thus, similar in size to the
monoflagellate spermatozoa found here for C.
loehensis (9.1 um). Biflagellate spermatozoa of
another Japanese form, C. aff. fluminea, are
reported to be also relatively small compared
with the sympatric C. leana (13.9 um and 16.9
um, respectively) (Konishi et al., 1998). Size
difference in corbiculid sperm observed in taxa
from China was found to be correlated with
ploidy (Qiu et al., 2001) and in taxa from Japan
with number of mitochondria (Konishi et al.,
1998).

Brooding

Observations presented above on reproduc-
tion in Indonesian corbiculids agree with the lit-
erature data in showing prevalence of brooding
among freshwater Corbicula (Morton, 1986;
Byrne et al., 2000; Siripattrawan et al., 2000). To
date, within the genus only the estuarine (i.e.,
brackish-water) sister taxon С. japonica is
non-brooding and characterized by the devel-
opment with free-swimming veligers (Byun 8
Chung, 2001) Among the freshwater
corbiculids the endemic С. запада! from Lake
Biwa with its benthic egg masses with direct
developing young (Hurukawa & Mitsumoto,
1953) remains the only known exception of an
ovoviviparous reproductive mode.

However, we here documented a greater di-
versity of brooding characteristics in taxa
particularily from Sulawesi than was witnessed
earlier for the rest of the collective Old World
range of Corbicula. Remarkable is the pres-
ence of large juveniles being incubated in the
gills of C. linduensis and the brooding utilizing
both demibranchs in С. possoensis, which 1$
both not known in any other congeners.

Historical Zoogeography

According to the phylogenetic systematics
discussed above, two groups of taxa can be
distinguished among Indonesian corbiculids.
On one hand, there 1$ at least one common
widespread clade in Asia that includes popula-
tions identified as C. fluminea occuring in Ko-
rea and Thailand, as well as those populations
from the Sunda Islands Java and Lombok as-
signed here tentatively to C. javanica and C.
australis in Australia.

On the other hand, Sumatra and Sulawesi
seem to harbour Corbicula species with fairly
restricted occurrences that cluster according
to their distribution not only on but within these
islands. To the present knowledge, all lacustrine
forms described herein are endemic to their
respective lakes and lake systems with three
separate regions to be distinguished (Fig. 1): (1)
Northwest Sulawesi with the graben or basin of
the Palu River and Lake Lindu where C.
linduensis occurs, (ii) Lake Poso with the en-
demic С. possoensis, and (iii) the central lakes
of the Malili system with C. matannensis
(mainly in Lake Matano and Lake Mahalona)
and С. loehensis (in the satellite lakes of and in
Lake Towuti). Another species, C. subplanata
was described based on shells only from a
fourth region in southwest Sulawesi (area of
Minralang), but its specific identity and status
remains to be substantiated by anatomical and
molecular data. In contrast, sampling on
Sumatra 1$ to date too scarce to allow for any
solid judgement of an equally restricted occur-
rence of C. moltkiana in lakes Manindjau and
Singkarak only. In addition, the specific identity
and affinity of C. tobae endemic to Lake Toba in
northern Sumatra remains unresolved.

The pattern of endemic occurrences of
lacustrine corbiculids strongly correlates with
the distribution recently studied in detail for
pachychilid gastropods of the endemic
Tylomelania clade on Sulawesi (Rintelen &
Glaubrecht, 1999, 2002; Rintelen et al., submit-
ted), as well as with the biogeography of Indo-
nesian Ancylidae (Glaubrecht, unpub. data),
and is, therefore, no artefact of insufficient data
on the range of limnic molluscs in Southeast
Asia. In case of the evolution of endemic
corbiculid bivalves in separate areas within the
geologically complex island of Sulawesi, it re-
mains to be tested, based on further sampling
and detailed molecular studies, whether this
biogeographic pattern finds its historical expla-

36 GLAUBRECHT ETAL.

nation in the spatial isolation over longer geo-
logical time in concert with the composite na-
ture of this odd shaped island that formed by
fusion of several microplates (terranes) in Late
Miocene-Early Pliocene (palaeogeographical
background: Whitmore, 1981; Hall & Bundell,
1996; Metclafe et al., 2001).

Using other limnic molluscs as models, it has
recently been hypothesized that, for example,
the phylogeny and biogeography of pachychilid
gastropods of Brotia, sensu lato, in Southeast
and Austral Asia reflect palaeogeographical
events since the Cretaceous/Cenozoic rather
than more recent geological history
(Glaubrecht, 2000; Glaubrecht & Rintelen,
2003; Kóhler et al., 2000; Kóhler & Glaubrecht,
2001, 2003). The latter comprise, for example,
those events related to the formation of
Sundaland and its drowning during the Plio-
Pleistocene. Accordingly, the distribution of
taxa of the Brotia, sensu lato, complex might
represent an ancient vicariance pattern caused
by plate and terrane tectonics that has not
been obscured subsequently, presumably due
to comparatively restricted dispersal abilities of
these viviparous snails in conjunction with eco-
logical factors.

In contrast, the available evidence in case of
the East Asian and Australian freshwater
corbiculids was regarded incompatible with an
ancient vicariance scenario. Above all, an as-
sumed late Cenozoic origin of freshwater Cor-
bicula restrict applicability of the studied group
as indicator of a long and complex geological
history and biogeography within the so-called
“Wallacea” (as transitional zone between the
Australian and Oriental region). Second, the
mitochondrial COI sequences generated for
those corbiculids collectively distributed from
the Japanese Archipelago to Australia indicated
a phylogenetically shallow polytomy, suggesting
an evolutionary recent common origin to
Siripattrawan et al. (2000) and Pfenninger et al.
(2002). Showing rather low levels of genetic
distances between different lineages of Asian
freshwater Corbicula, our analyses including
now the Indonesian taxa in general support this
scenario of rather late divergence of freshwater
lineages. Nevertheless, the higher sequence
distances in particular shown by C. moltkiana
on Sumatra may indicate that this divergence
started earlier than Pleistocene age suggested
by Pfenninger et al. (2002).

In summary, three statements of biogeo-
graphical importance are implied by the
present study: (1) presence of distinct and, in

relation to other Asian forms, old taxa on
Sumatra and Sulawesi, (ii) a remarkable diver-
sity of Corbicula on the island of Sulawesi with
at least three distinct lineages and taxa, re-
spectively, and (iii) presumably a relatively late
colonization of the Sunda Islands Java and
Lombok by С. javanica with its strong affinity to
C. fluminea.

Evolutionary Ecology

The СО! sequence data in conjunction with
the new finding of exceptional life history and
anatomical characteristics, including features
of sperm morphology and incubation, pre-
sented herein for Indonesian Corbicula suggest
an evolutionary ecology hypothesis on their ori-
gin (theoretical background, reviews:
Glaubrecht, 1996; Streit et al., 1997). In par-
ticular, the diversity of Corbicula in the so-
called “ancient” lakes on Sulawesi deserve
such an explanation, while there is only one riv-
erine corbiculid which has been collectively
assigned to C. subplanata (see Introduction).

Accordingly, we anticipate colonization of
Sumatra and Sulawesi by an early, sexual re-
producing and incubating corbiculid ancestor
with monoflagellate spermatozoa and subse-
quent radiation by speciation of individual
corbiculids in situ particularly in Lake Poso and
the Malili lake system, respectively, once these
special habitats open up and provided new
ecological opportunities. A time frame for this
process can be given very tentatively only, with
an estimated age of Lake Poso and the Malili
lakes of about 1-2 myr (Rintelen et al., submit-
ted).

Apparently, local ancient lakes with their tem-
porally stable habitats facilitated an endemic
radiation of specialized forms in case of C.
possoensis in Lake Poso and C. matannensis
and C. loehensis inhabiting the Malili lakes. The
latter two lineages might originate from
intralacustrine divergence within (at least tem-
porarily) separated lakes and/or independent
colonizations of the Malili system. In contrast,
this specific intralacustrine speciation in an
ancient lake setting is more unlikely in case of
C. linduensis (for which riverine localities are
also reported here for the first time), because
Lindu is not known to fulfill the criteria of being
an ancient lake. Nevertheless, the outstanding
mode of brooding in C. linduensis may indicate
rather long isolation.

Exceptional cases for lacustrine speciation
and adaptive radiation on Sulawesi are pro-

FRESHWATER CORBICULIDAE FROM INDONESIA 37

vided by gastropods of known incidences in
Pachychilidae (Tylomelania), Ancylidae
(Protancylus) and Lymnaeidae (Miratesta)(P.
Sarasin & F. Sarasin, 1898; Rintelen &
Glaubrecht, 1999; Rintelen et al., submitted),
and among bivalves also by the evolution of the
endemic corbiculid genus Possostrea in Lake
Poso (Bogan & Bouchet, 1998). On a more
subdued scale, such a process that involves
the evolution of several adaptations unknown
for long in other freshwater congeners seems
to have occurred only in Corbicula sandai of
Lake Biwa. Although not brooding but laying
benthic eggs masses with direct developing
young, this endemic Japanese corbiculid share
certain reproductive features (i.e., the
monoflagellate sperm) with taxa endemic to
Sulawesi, to which also its sequence data ex-
hibit a certain affinity.

CONCLUSION

As shown in the present study, peculiarities
of shell morphology, anatomy, sperm morphol-
ogy and the brooding process, as well as avail-
able molecular data support the presence of
several endemic Corbicula taxa on Sumatra
(C. moltkiana) and particularly on Sulawesi (C.
linduensis, C. possoensis, C. matannensis
and C. loehensis). These taxa apparently rep-
resent relatively old and distinct genetic lin-
eages which show no particularly close
relationship to any previously studied Corbicula
from the Japanese islands, Asian mainland or
Australia. In contrast, C. javanica that is sup-
posed to be widely distributed across the
Sunda Archipelago, appears closely related to
a Korean lineage identified as C. fluminea
within an Asian cluster, and might be a later
migrant in this region. Future additional mor-
phological, biological and molecular investiga-
tions may provide more decisive information
concerning the evolutionary pathways along
which Corbicula species colonized freshwater
habitats in Southeast and Austral-Asia.

ACKNOWLEDGEMENTS

We are grateful to Frank Kohler for his
assistence during field work in Indonesia, and
to Sabine Schütt (both ZMB) for material col-
lecting on Sumatra in 2000. We thank
Ristiyanti Marwoto (MZB) for logistical help dur-
ing the field work, МР! for providing research

and collecting permits, and Ahmat Munandar
for field assistence in August 1999 on Sulawesi.
We thank E. Wásch, V. Heinrich and N.
Brinkmann for various technical help with pho-
tographs, as well as |. Kilias for her help with
literature research. We also thank Philippe
Bouchet (Paris), R. Araujo (Madrid) and C.
Ituarte (Buenos Aires) for providing material for
comparison, and Diarmaid O Foighil and two
anonymous reviewers for valuable comments
on an earlier version that helped to significantly
improve the manuscript. A. K. was funded
through a travel grant of the Deutsche For-
schungsgemeinschaft (September to Decem-
ber 2000) and an Alexander von Humboldt
Research Fellowship (March 2002 to February
2003) allowing for research visits to the ZMB.
The 1999/2000 field work in Indonesia has
been funded through grants of the Deutsche
Forschungsgemeinschaft to M.G. (Gl 297/1 and
297/4).

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Revised ms. accepted 28 February 2003

MALACOLOGIA, 2003, 45(1): 41-100

THE APPLE SNAILS OF THE AMERICAS
(MOLLUSCA: GASTROPODA: AMPULLARIIDAE:
ASOLENE, FELIPPONEA, MARISA, POMACEA, POMELLA):
ANOMENCLATURAL AND TYPE CATALOG

Robert H. Cowie’ & Silvana С. Thiengo*

ABSTRACT

Ampullariidae are freshwater snails predominantly distributed in humid tropical and sub-
tropical habitats in Africa, South and Central America and Asia. This catalog is concerned
only with the American species, the majority of which are placed in the genus Pomacea.
Species of Pomacea are found throughout most of South and Central America and the
Caribbean, with a single species extending into southeastern USA. The other American
genera are Asolene, Felipponea, Pomella and Marisa, all South American. The taxonomy
of the group is heavily based on shell morphology but the true number of valid taxa re-
mains unknown, pending revisionary work. This catalog provides the rigorous nomencla-
tural base for this future work by bringing together all the available and unavailable
genus-group and species-group names that have been applied to American ampullariids,
indicating their current nomenclatural status (species, subspecies, synonyms, etc.). The
catalog lists 14 published genus-group and 307 published species-group names for Ameri-
can ampullariids. Of these, 7 genus-group (including 2 subgeneric) and 141 species-group
(including 23 infraspecific) names are currently valid. There are 4 genus-group synonyms,
133 species-group synonyms, and 11 species-group homonyms. Also listed are 3 unavail-
able genus-group and 23 unavailable species-group names. The catalog provides biblio-
graphic details for all names, details of type localities and locations of type material, and
geographic distribution as far as can be ascertained given the confused state of the tax-
onomy. The catalog is a work of nomenclature; it is not a revisionary work of taxonomy.

Key words: Ampullariidae, freshwater snails, nomenclature, type material, North America,
South America, Central America.

INTRODUCTION

Ampullariidae are freshwater snails predomi-
nantly distributed in humid tropical and sub-
tropical habitats in Africa, South and Central
America and Asia. They include the largest of
all freshwater snails (Pomacea urceus Can at-
tain a shell height of 145 mm — Burky, 1974; Р
maculata can exceed 155 mm - Pain, 1960)
and frequently constitute a major portion of the
native freshwater mollusk faunas of these re-
gions. Among the seven to ten genera usually
recognized, the two largest are Pomacea, per-
haps with about 50 real species (but 117 nomi-
nally valid species recognized herein), and Pila,
with about 30 (Berthold, 1991). Snails in these

two genera particularly are frequently known as
“apple snails” because many species bear
large, round, often greenish shells. The term
“apple-shell” was first used by Perry (1810c) in
his introduction of the name Pomacea, for his
new species P. maculata, because of “its gen-
eral resemblance to ... an apple” (“Ротит” in
Latin), and not from the Greek Пора, which
means an operculum.

Ampullariidae (junior synonym Pilidae; Cowie,
1997а; ICZN, 1999a) are operculate snails.
They are most closely related to the
Viviparidae, together with which they form the
superfamily Ampullarioidea in the orders or su-
perorders (depending оп classification)
Mesogastropoda of earlier authors and Caeno-

‘Corresponding author. Center for Conservation Research and Training, University of Hawaii, 3050 Maile Way,
Gilmore 408, Honolulu, Hawaii 96822, USA; cowie@hawaii.edu
2Instituto Oswaldo Cruz, Av. Brasil 4365, 21045-900 Rio de Janeiro, RJ, Brasil; sthiengo@ioc.fiocruz.br

42 COWIE & ТНЕМСО

gastropoda of more recent authors (Ponder &
Warén, 1988; Berthold, 1989; Bieler, 1992;
Ponder & Lindberg, 1997).

Traditional subdivision of the family, by vari-
ous authors, has been into seven to ten gen-
era, with the form of the siphon and the
operculum considered diagnostically significant
(e.g., Michelson, 1961). Pain (1972) briefly re-
viewed the history of taxonomic work on the
family. More recently, Berthold (1991: 245-250)
recognized ten genera (and three subgenera)
with approximately 120 species. His detailed
anatomical account treated representative spe-
cies from each of these generic groupings. He
divided the family into two subfamilies: the
Afropominae (containing just a single Recent
African species in the genus Afropomus); and
the Ampullariinae, which he subdivided into the
tribes Sauleini (one genus, Saulea, containing
two African species, one Recent, one fossil)
and Ampullariini (the remainder). He further
subdivided the Ampullariini into the groups
Heterostropha and Antlipneumata, but these
divisions and names have been criticized by
Bieler (1993), who reanalyzed Berthold's data
using cladistic techniques. Bieler's reanalysis
showed that the various groupings of genera
remained more or less similar to those of
Berthold, but the relationships among these
groups were inconsistent. Given these incon-
sistencies, it seems unwise to force the vari-
ous clades into a traditional hierarchy of
family-group names. One of the ten genera
recognized by Berthold (1991), Pseudo-
ceratodes (African, fossil only), was included in
the family only tentatively. Of the remaining nine
genera, six contain fewer than six species
each: Afropomus and Saulea are African;
Asolene, Felipponea, Pomella and Marisa are
South American. The three genera Lanistes
Montfort, Pila (Ampullaria Lamarck and
Ampullarius Montfort are junior synonyms;
Cowie, 1997a; ICZN, 1999a) and Pomacea,
containing 21, about 30, and about 50 species,
respectively, comprise the great majority of
species in the family. Lanistes (distinguished
by its hyperstrophic and hence superficially sin-
istral shells) is African (including Madagascar).
Pila is African and Asian. Pomacea is South
and Central American.

This catalog is concerned only with the
American species, the majority of which are
placed in the genus Pomacea. Species of
Pomacea are found throughout most of South
and Central America and the Caribbean, with a
single species, Pomacea paludosa, extending
into the southeast USA. The genus is divided

into two subgenera, Pomacea sensu stricto
and Pomacea (Effusa). Berthold (1991) con-
sidered Pomacea to be monophyletic, with
Marisa as its sister-group, and the similarities
between Marisa and Pomacea subgenus
Effusa to be convergent (see also Pilsbry,
1933: 72-73). However, the relationships
among the two subgenera of Pomacea and the
genus Marisa are not well resolved (Bieler,
1993) and, at least in terms of shell morphol-
ogy, the three taxa intergrade. The distinctions
among these and the other American genus-
group taxa have been generally not well under-
stood. This catalog follows Berthold (1991:
248—250) regarding validity and status of ge-
nus-group names, without necessarily implying
support for his taxonomic views.

One or more species of Pomacea, introduced
to Southeast Asia and islands of the Pacific, in-
cluding the Hawaiian Islands, have become
major agricultural pests, notably in rice and taro
but also in other crops (Cowie, 2002). However,
the true identity of the species involved is uncer-
tain, having been treated variously as Pomacea
canaliculata (Smith, 1992; Hendarsih et al.,
1994), Р. lineata (Cheng, 1989; Laup, 1991), Р
gigas (see Guerrero, 1991), P “insularis” (see
Acosta & Pullin, 1991), Pomacea cf.
canaliculata (Ng et al., 1993), simply Pomacea
sp. (Acosta & Pullin, 1991), a “hybrid [of]
Ampullaria canaliculata and Ampullaria cuprina”
(Anderson, 1993), and even “Ampularius sp. a
hybrid of undetermined origin” (Lacanilao,
1990). Keawjam & Upatham (1990) recognized
three species of Pomacea introduced in Thai-
land: Р canaliculata, P. insularum and an uni-
dentified species of Pomacea. Mochida (1991)
indicated that as well as P. canaliculata (which
he considered frequently to have been
misidentified as Р insularum) two other species
of Pomacea have also been introduced to the
Philippines: Р gigas and Р cuprina (the latter
possibly a misidentification of P bridgesii, a spe-
cies that has been carried all over the world by
the domestic aquarium trade — Cowie, 1995). In
Japan, three “strains of Pomacea canaliculata”
have been identified, differing in shell colour and
pattern, salinity tolerance, and in aspects of re-
production and growth (Brand et al., 1990; Fujio
et al., 1991). In the Philippines, the snails have
even been identified as species of Pila (see
Guerrero, 1991). In Hawaii, where four
ampullariid species are recorded (Cowie, 1995),
snails in an aquaculture project have been re-
ported as hybrids of Pomacea canaliculata and
P. paludosa (Nishimura et al., in Tamaru,
1996).

NEW WORLD AMPULLARIIDAE 43

Ampullariid species-level taxonomy has been
heavily reliant on shell morphology, yet snail
shells, and especially ampullariid shells, ex-
hibit much intraspecific variation. The tax-
onomy and systematics of most species have
not been adequately worked since their original
descriptions. The pest species (even if it turns
out to be more than one species) in Southeast
Asia nevertheless appears to belong to a rela-
tively well circumscribed group of more or less
closely related species from South America.
However, within this group, the species and
their relationships are very poorly understood.
The group comprises a large number of nomi-
nal species, including P. canaliculata. From
time to time, some of the species within this
“canaliculata group” have been formally syn-
onymized, informally linked together, distin-
guished as separate species, and so on. This
confusion was discussed but not resolved by
Alderson (1925), the most recent author to re-
vise Pomacea and Pila widely (referring to the
two genera together as “Ampullaria”). He im-
plicitly recognized most of the species in the
“canaliculata group” as a more or less closely
knit group. Within this group he further recog-
nized a number of rather vaguely defined as-
sociations of species, for instance explicitly
linking Pomacea immersa, P. amazonica and
Р haustrum, although without formally synony-
mizing them; and informally referring to an-
other subset of the group as “the /ineata
group”. However, he did retain most species
as valid. It is quite possible that, just as for the
large number of Central American species
synonymized under Pomacea flagellata by
Pain (1964), many other “species” of
Ampullariidae, including those in the
“canaliculata group”, do not deserve distinct
specific status (Pain, 1960; Cazzaniga, 1987,
2002). A modern revision, involving not only
conchology but also internal anatomy and mo-
lecular characters, might reduce the
“canaliculata group” to as few as three spe-
cies, possibly Р canaliculata, P. lineata and P.
gigas (= maculata — see main catalog). Until
such work is undertaken, however, the status
of these various nominal species will remain
obscure.

The purpose of this catalog, then, is to pro-
vide a rigorous base for this revisionary work
by bringing together all the available and un-
available genus-group and species-group
names in the large genus Pomacea and the
other much smaller South American genera of
Ampullariidae, indicating their current nomen-

clatural status (species, subspecies, syn-
onyms, etc.) generally according to the most
recent revisions; a total of 14 genus-group and
307 species-group names (Table 1). The cata-
log also provides bibliographic details for all
names, details of type localities and locations
of type material, if known, and geographic dis-
tribution as far as can be ascertained given the
confused state of the taxonomy.

EXPLANATORY INFORMATION
Scope

This catalog lists all published genus-group
and species-group names found in the litera-
ture, whether available or unavailable accord-
ing to the International Code of Zoological
Nomenclature (ICZN, 1999b), that have been
applied to the Ampullariidae of North, Central,
and South America, and the Caribbean region.

Arrangement and Treatment of Taxa

The sequence of genera 1$ alphabetical.
Subgenera appear in alphabetical order within
genera. Genus-group synonyms are listed
chronologically under the genus-group head-
ing. All species-group names (valid and in-
valid, available and unavailable) are listed
alphabetically within genera/subgenera. Incor-
rect spellings are listed only if confusion might
be caused by their omission; they may also
be mentioned in Remarks sections. Treatment
of species-group names follows the major
authoritative revisions, although few of these
are recent. Names proposed as “forms”, “va-

TABLE 1. Summary of the numbers of names of
American ampullariid taxa (including names that
are incerae sedis) treated in this catalog.

Available Unavailable
Genus-group Valid genus 5
names Valid subgenus 2 3
Synonym 4
Species- Valid species 17
group names Valid infra- 23
specific 23
Synonym 123
Homonym* 11

*Includes homonyms considered to represent valid
species (10) and valid infraspecific taxa (1).

44 COWIE & THIENGO

rieties”, etc., and neither already synonymized
nor raised to subspecific status are simply
treated, along with subspecies, as infraspe-
cific. Treatment of genus-group names also
follows the most recent authoritative revisions.
In some instances in which we treat a name
as a junior synonym, one or more other
names automatically become synonyms of the
senior synonym because they had already
been treated as synonyms of the junior syn-
onym. In some cases this results in the intro-
duction of a new synonymy, indicated in
boldface by “N. syn.”. However, no other revi-
sionary work has been attempted and no new
taxonomic decisions have been made.

Typographical Treatment of Names

Family and genus-group headings are cen-
tered in upper case type. Valid genus-group
names are listed flush left in boldface upper
case type. Valid, available species-group
names are listed flush left in boldface, in-
fraspecific names preceded by a “+”. Syn-
onyms are listed in italics flush left, upper case
for family and genus-group names, lower case
for species-group names. In the species-
group, junior homonyms are also listed in ital-
ics flush left, or in boldface italics flush left if
new or replacement names have not been pro-
vided. Nomenclaturally unavailable names are
listed in plain type, flush left.

Taxonomic References

The citation for the original proposal of a ge-
nus-group name follows the name. The refer-
ence consists of author(s), date of publication
and page number. For species-group names,
on the line following the name and indented,
the name is given in its original generic combi-
nation (including subgenus if in the original de-
scription, and using the original orthography,
even if incorrect) and with its original status in-
dicated (e.g., subspecies, “var.”, as neces-
sary). The name is followed by its author(s),
date of publication, page number, and plate/fig-
ure number(s). When an author published the
same name as new for the same taxon in
more than one place, the later citation is given
in square brackets following the first citation.

The author/date citation acts as a reference
to the work as listed in the Literature Cited
section. If an author published more than one
work in the same year, a suffix (a, b, с, etc.),
indicating chronological order of publication, is

attached to the date in both the catalog text
and the Literature Cited. Authors’ names con-
taining the terms “де”, “4”, “von”, if being of
European continental origin, are cited and
alphabetized in the Literature Cited by the
main name, e.g., “Ampullaria guadelupensis
Martens, 1857” in the main body of the catalog
and “Martens, E. von. 1857” in the Literature
Cited.

The page number cited is that on which the
name first appeared. In some instances, the
name first appeared on different pages, for in-
stance in a list or key, with the actual descrip-
tion beginning on a subsequent page. In such
cases, both page numbers are cited.

If the current status of a species-group
name differs from that in the original descrip-
tion, this is indicated, with appropriate refer-
ences, in a Remarks section below the
standard entry for the species.

Type Species

For nomenclaturally available genus-group
names, the type species and its method of
fixation (following Code Arts. 66-70) are given
following the literature citation.

Homonyms and Replacement Names

Homonymy of species-group names is indi-
cated in the Remarks section under the name.
In many cases, the junior homonym has al-
ready been synonymized with another earlier
name, or a replacement name has already
been provided. In cases in which a replace-
ment name appears necessary, no replace-
ment name is here provided, pending further
research. We have not made an exhaustive
search for possible senior homonyms.

Unavailable Names

Unavailable names are listed with full citation
and a statement of why the name is unavail-
able, e.g., “nom. nud.”, “first published as a jun-
ior synonym of ...”, etc. No other information is
provided except for explanatory details in the
Remarks section, if necessary. Obviously in-
correct spellings are not listed but may be
mentioned in annotations.

Misidentifications

Misidentifications are not formally listed. No
genus-group misidentifications are mentioned.

NEW WORLD AMPULLARIIDAE 45

Species-group misidentifications are noted in
square brackets or in Remarks sections, if
necessary for clarity.

Miscellaneous Annotations

Under each genus-group heading, explana-
tory and other useful information is given im-
mediately under the genus-group synonymy.
Annotations other than those indicated in the
above paragraphs are placed in square brack-
ets immediately following the item to be clari-
fied or, if the annotations are more extensive,
placed in a Remarks section following the
standard entry for the species.

Type Localities and Type Material

The type locality (“the geographical ... place
of capture, collection or observation of the
name-bearing type” [Code Art. 76]) is given for
each available species-group name immedi-
ately following the author and citation. The lo-
cation is given verbatim as published by the
author, without translation. If no locality was
given by the author, this is simply stated, in
square brackets. Any additional intepretive or
explanatory information regarding the type lo-
cality is placed in square brackets following
the originally published locality, or, if extensive,
in the Remarks section. However, an exhaus-
tive attempt to determine the exact collection
locality has not been made.

Location and catalog numbers of type mate-
rial, if known, are given, following the type lo-
cality information. In many instances, the
original descriptions did not designate a holo-
type or even identify a type series. Even
though many of these descriptions were prob-
ably based on single specimens, it is rarely
possible to determine this with certainty.
Therefore, in most cases, the material known
to have been used in describing a new spe-
cies should be designated as a lectotype (or
lectotype and paralectotype(s) if more than
one specimen is present in the inferred type
series) (Code Rec. 73F). Rather than desig-
nating lectotypes here, we consider such
specimens to be syntypes, pending further
study. The information provided is derived
from the literature (citations given), and from
enquiries made to numerous museums and
our own study in various museums (see Ac-
knowledgments); an exhaustive search for
type material has not been made.

Distributions

The distribution (if known) of each species is
given following the type locality and type material
information. In most cases this information is
not detailed, providing simply the country or re-
gion from which the species has been recorded
in the literature. Citations for the sources of this
information are provided, unless the only infor-
mation available is the type locality, for which the
reference has already been provided.

Museum Collection Acronyms and Type Mate-
rial Holdings

The following acronyms are used for the vari-
ous museum collections referred to in the
catalog. The number of American taxa repre-
sented by type or possible type material in
each collection, as known to us or referred to in
the literature, is indicated in parentheses. Re-
search will undoubtedly uncover additional type
material in many of these collections.

AMNH = American Museum of Natural History,
New York, U.S.A. (4)

ANSP Academy of Natural Sciences, Philadel-
phia, U.S.A. (22)

BMNH The Natural History Museum, London,
О.К. (83)

CAS California Academy of Sciences, San
Francisco, U.S.A. (1)

CMNH Carnegie Museum of Natural History,

Pittsburgh, U.S.A. (1)
HUJ Hebrew University of Jerusalem, Israel (5)

IMLA Fundación e Instituto Miguel Lillo,
Universidad Nacional de Tucumán, Ar-
gentina (1)

MACN Museo Argentino de Ciencias Natu-

rales, Buenos Aires, Argentina (1)
MCZ Museum of Comparative Zoology, Har-
vard University, Cambridge, U.S.A. (29)

MCSN Museo Civico di Storia Naturale,
Milano, Italy (0)

MHNG Muséum d'Histoire Naturelle, Genève,
Switzerland (8)

MHNS Museo de Historia Natural La Salle,
Caracas, Venezuela (1)

MMUE The Manchester Museum, University of
Manchester, Manchester, U.K. (1)

MNCN Museo Nacional de Ciencias Natu-
rales, Madrid, Spain (3)

MNHN Muséum National d’Histoire Naturelle,

Paris, France (30)
MNHNS Museo Nacional de Historia Natural,
Santiago, Chile (possibly 21)
ММК] Museu Nacional, Rio de Janeiro, Brasil
1
Museu de Zoologia da Universidade de
Säo Paulo, Brasil (0)

MZUSP

46 COWIE & ТНЕМСО

NMW National Museum of Wales, Cardiff,
U.K. (10)

NZSI Zoological Survey of India, National
Zoological Collection India, West Ben-
gal, Calcutta, India (1)

RMNH Nationaal Natuurhistorische Museum,
Leiden, Netherlands (2)

SMFD Forschungsinstitut und Naturmuseum
Senckenberg, Frankfurt-am-Main,
Germany (possibly 2)

UF Universisty of Florida, Florida Museum
of Natural History, Gainesville, U.S.A. (1)

UMMZ University of Michigan, Museum of Zo-
ology, Ann Arbor, U.S.A. (5)

USNM National Museum of Natural History,
Washington D.C., U.S.A. (11)

ZMHB Museum für Naturkunde der Humboldt-
Universitát, Berlin, Germany (24)

ZMUH Universität von Hamburg, Zoologisches
Institut und Zoologisches Museum,
Hamburg, Germany (0)

ZMZ Zoologisches Museum der Universität,
Zürich, Switzerland (1)

ZSM Zoologische Staatssammlung Múnchen,
Germany (14)

Abbreviations

The following abbreviations are used
throughout the catalog.

Art(s). Article(s) (of the Code)

Code International Code of Zoological
Nomenclature (ICZN, 1999b)

fig(s). figure(s)

ICZN International Commission on
Zoological Nomenclature

nom. nud. nomen nudum

N. syn. New synonymy

pl(s). plate(s)

p. page oo

pers. comm. personal communication

Rec. Recommendation (of the Code)

s.l. sensu lato

spm(s) specimen(s)

Susie sensu Stricto

subg. subgenus

SYSTEMATIC CATALOG

Family AMPULLARIIDAE Gray, 1824

AMPULLARIIDAE Gray, 1824: 276. Type ge-
nus Ampullaria Lamarck, 1799 [= Pila Réding,
1798].

PILIDAE Preston, 1915: 96. Type genus Pila
Röding, 1798.

ICZN (1999a), following Cowie (1997a: 83-
88), confirmed the name Pilidae as a junior syn-
onym of Ampullariidae and invalid.

Genus ASOLENE Orbigny, 1838

ASOLENE Orbigny, 1838d: 364. Type species:
Helix platae Maton, 1811, by subsequent des-
ignation of Gray (1847: 148).

AMPULLOIDEA Orbigny, 1841e: 379. New
name for Asolene Orbigny, 1838 (see Pilsbry,
1933: 74).

AMPULLOIDES Orbigny, 1842c: 1. Incorrect
spelling of Ampulloidea Orbigny, 1841.

ASOLENA Herrmannsen, 1846a: 84. Incorrect
spelling of Asolene Orbigny, 1838.

AMPULLAROIDES Gray, 1847: 148. Incorrect
spelling of Ampulloidea Orbigny, 1841 (see
Pilsbry, 1933: 74).

Treated as a full genus following Berthold
(1991: 23). Orbigny (1841e: 379) replaced
Asolene with Ampulloidea, treating only platae
Maton, 1811, but without explicitly saying that
this was the only species and hence not desig-
nating it as the type. Previously (Orbigny,
1838d: 364), he had included two species
(platae Maton, 1811, and celebensis Quoy &
Gaimard, 1834 [the latter is now placed in Pila
Röding, 1798]).

brownii

Ampullaria Brownii Jay, 1839: 112, pl. 1, fig. 4.
River Amazon [= Brasil]. Syntype: AMNH
56107 (Boyko 8 Cordeiro, 2001: 16) [labeled
as “figd type” in Jay's handwriting (P. M.
Mikkelsen, pers. comm. to RHC, 7 May
2002)]; possible additional syntype material:
MCZ. Distribution: Brasil.

Remarks: Pain (1960: 430) stated that
“[t]hrough the kindness of Dr. J. С. Bequaert
and Dr. W. J. Clench [both of the MCZ], | have
been able to examine the type”. This is not
considered an inadvertent lectotype designa-
tion because it is not specific as to the speci-
men examined (Boyko 8 Cordeiro, 2001: 16).
Synonym of crassa Swainson, 1823, teste
Philippi (1852a: 34), Paetel (1887: 477),
Kobelt (1913Е 189) and Pain (1960: 429).

crassa
Ampullaria crassa Swainson, 1823a: pl. 136,
upper and lower figs. [No locality given.]
Syntypes (“the only specimen | have” and the
specimen in “the figure of Martini” (Swainson,
1823a: pl. 136): possibly MMUE (Dean, 1936:
232; H. McGhie, pers. comm. to RHC, 28
July 2002), not found by us in ВММН (cf.
Dance, 1986: 227). Distribution: Brasil, Bo-
livia, Peru, Equador, Colombia, Venezuela,

NEW WORLD AMPULLARIIDAE 47

Guyana, Surinam, French Guiana (Baker,
1914: 661; Pain, 1960: 430; Geijskes & Pain,
1957: 45).

Remarks. Placed in Limnopomus Dall by
Pain (1952: 31, 1960: 429) and Geijskes 8
Pain (1957: 45), but in Asolene Orbigny by
Tillier (1980: 21), followed here.

crassa
Ampullaria crassa Orbigny, 1835a: 33. Rio
Parana (republica Argentina). Syntypes:
BMNH 1854.12.4.332 (13 spms.) [labeled
“roissyl”].

Remarks. Junior primary homonym of
crassa Swainson, 1823, and crassa
Deshayes, 1830 [= Melantho ponderosa
(Deshayes, 1825), teste Paetel (1887: 478);
not Ampullariidae], replaced by roissyi
Orbigny, 1841. Synonym of pulchella Anton,
1838, teste Gaudion (1879: 38), lhering
(1898: 50, 1919: 337) and Hylton Scott (1958:
310).

cyclostoma

Ampullaria cyclostoma Spix, in Wagner,
1827: 4, pl. 4, fig. 5. Brasilia. Syntype: ZSM
20012075 (E. Schwabe, pers. comm. to
RHC, 28 July 2002; see also Fechter, 1983:
221). Distribution: Argentina, Paraguay, Uru-
guay, Bolivia, Brasil (Paraguay-Parana drain-
age) (Pain, 1960: 430).

Remarks. Authorship is given here as “Spix,
in Wagner”, following Cowie et al. (in prep.).
Synonym of platae Maton, 1811, teste Hylton
Scott (1958: 308) and S. C. Thiengo (unpub-
lished), followed here, although contrary to
Pain (1960: 430), who retained it as a valid
species in Limnopomus Dall, 1904, which 1$
here treated as a synonym of Pomacea
Perry, 1810.

exumbilicata
Helicina exumbilicata Spix, in Wagner, 1827:
4, pl. 5, fig. 4. In aquis Provinciae Bahiensis.
Type material: probably lost (S. C. Thiengo,
unpublished).
Remarks: Authorship is given as “Spix, in
Wagner”, following Cowie et al. (in prep.), who
also explain the publication history of this
work. Spix illustrated exumbilicata as a valid
species, but Wagner, in writing the description,
treated it as a variety of crassa Swainson,
1823. Synonym of crassa Swainson, 1823,
teste Philippi (1852a: 73) and Pain (1950b:
72), although the latter cited Spix's pl. 4, fig. 2.
fasciolata
Helix fasciolata Spix, in Wagner, 1827: 4, pl.
5, fig. 1. In aquis Provinciae Bahiensis. Type

material: probably lost (S. C. Thiengo, unpub-
lished).

Remarks. Authorship is given here as “Spix,
in Wagner”, following Cowie et al. (in prep.),
who also explain the publication history of this
work. Spix illustrated fasciolata as a valid spe-
cies, but Wagner, in writing the descriptions,
treated it as a variety of crassa Swainson,
1823. Synonym of crassa Swainson, 1823,
teste Pain (1950b: 72), although he cited pl.
5, IG: 2

+ gallardoi

Ampullaria pulchella Gallardoi lhering, 1919:
337. curso inferior del río Paraná hasta
Corrientes, del Chaco Argentina y del señor
А. de W. Bertoni, de la Asunción. Type mate-
rial: not found by us in MACN, not found by us
in MZUSP (cf. Dance, 1986: 214). Distribu-
tion: Argentina, Bolivia (Hylton Scott, 1958:
311)

granulosa

Ampullaria granulosa Sowerby, 1894: 49, pl.
4, fig. 24. Cayenne. Lectotype (Pain, 1949b:
pl. 2, figs. 5, 6; see also Tillier, 1980: 20):
BMNH 1894.6.11.1. Distribution: Guyana,
Surinam, French Guiana (Vernhout, 1914a:
43; Pain, 1952: 30; Geijskes & Pain, 1957:
46; Tillier, 1980: 20)

Remarks. Placed in Limnopomus Dall by
Pain (1952: 31) and Geijskes 8 Pain (1957:
45). Placed here in Asolene following Tillier
(1980: 17). The original description was not
explicitly based on a single shell, so the
specimen figured by Pain as the “type” must
be considered a lectotype (Code Art. 74.6,
Rec. 73F).

impervia

Ampullaria impervia Philippi, 1851: 17, pl. 4,
fig. 7 [1852b: 21]. Brasilien. Syntype: ZSM
20012067 (E. Schwabe, pers. comm. to
RHC, 28 July 2002). Distribution: “Brésil”
(Gaudion, 1879: 31), “Bolivia, etc.” (Sowerby,
1909a: 353).

Remarks. Synonym of crassa Swainson,
1823, teste Pain (1960: 429).

monticola

Ampullaria crassa var. monticola Vernhout,
1914b: 47, pl. 1 [not pl. 2, as indicated in the
text], fig. 15a, b. Mount Cottica on the right bank
of the Lawa ... altitude of 450 m ... French
Guyana. Holotype: RMNH; paratype(s): RMNH.
Distribution: French Guyana (Tillier, 1980: 21).

Remarks. Three specimens were men-
tioned in the text, but only two were figured.
These were RMNH no. 132 (fig. 15a) and no.

48 COWIE & ТНЕМСО

131 (fig. 15b), but neither was specified as
being the holotype. Synonym of crassa
Swainson, 1823, teste Tillier (1980: 21).
naticoides

Ampullaria naticoides Orbigny, 1835a: 33. Un-
available name; first published as a junior syn-
onym of platae [as “Platea”] Maton, 1811, not
made available before 1961 (Code, Art. 11.6).
Syntypes: BMNH 1854.12.4.337 (7 spms.).

Remarks. Locality given by Orbigny (1835a:
33) as “Rio de la Plata, provincia Buenos-
Ayres (republica Argentina)”. Treated as a
synonym of platae Maton, 1811, by Orbigny
(1841e: 379), Paetel (1887: 480), Sowerby
(1909a: 356), Pilsbry (1933: 74) and Hylton
Scott (1958: 308).

+ nubila
Ampullaria nubila Reeve, 1856c: pl. 14, fig.
65. River Salomoens. Syntypes: BMNH
20020672 (2 spms.). Distribution: Brasil, Bo-
livia, Peru (Pain, 1960: 430).

Remarks. Subspecies of crassa

Swainson, 1823, teste Pain (1960: 430).
oblonga

Ampullaria (Pomus) crassa var. oblonga

Nevill, 1884: 11. Brazil; Amazon Rv. Type ma-

terial: possibly NZSI, not found by us in BMNH

(cf. Dance, 1986: 220). Distribution: Brasil.

Remarks. Junior primary homonym of
oblonga Swainson, 1823. Synonym of crassa
Swainson, 1823, teste Paetel (1887: 480).

olivieri
Ampullaria Olivieri Deshayes, 1830a: 31.
Cayenne. Type material: probably lost (Тег,
1980: 16). Distribution: French Guiana.

Remarks. Synonym of crassa Swainson,
1823, teste Deshayes (1838: 548), Philippi
(1852a: 34), Paetel (1887: 480) and Sowerby
(1909a: 347).

ormophora
Ampullaria ormophora Morelet, 1857: 30.
Nová Caledoniá [error]. Syntype: BMNH
1893.2.4.1805. Distribution: Brasil (from
syntype label).

Remarks. No ampullariids are known from
New Caledonia, which has therefore been
considered in error (Crosse, 1871: 185). Ten-
tatively place in Asolene Orbigny, based on
the syntype label, on which 1$ written “=
nubila Reeve”.

petiti
Ampullaria Petiti Crosse, 1891: 214, pl. 4, fig.
2. in flumine Amazonidum, Americas теп-
dionalis. Type material: MNHN (Sowerby,
1909b: 363) [not found by из]; topotype:
MNHN (coll. Jousseaume”; Тег, 1980: 19).

Remarks. Sowerby (1909a: 356) thought it to
be “perhaps a variety of A. impervia, Phil.” but
subsequently (Sowerby, 1909b: 363) consid-
ered the two taxa distinct. Synonym of crassa
Swainson, 1823, teste Pain (1960: 429), but
treated here as a valid species, following
Berthold (1991: 250; see also Tillier, 1980: 21).

platae
Helix Platæ Maton, 1811: 331, pl. 24, figs. 16,
17. America australi ... Rio de la Plata. Type
material: location not known to us. Distribu-
tion: Paraguay (Martens, 1857: 200; Paetel,
1873: 65, 1888: 481); Paraná (Pilsbry, 1933:
74); La Plata (Sowerby, 1909a: 356; Pilsbry,
1933: 74); La Plata and southern Brasil,
“sistema del río Paraná” (Ihering, 1919: 333).

pulchella
Ampullaria pulchella Anton, 1838: 50. [No lo-
cality given]. Type material: location not
known to us. Distribution: Rio Parana, La
Plata, Bolivia (Sowerby, 1909a: 348).

Remarks. Synonym of cyclostoma Spix, in
Wagner, 1827, teste Pain (1960: 430), but
treated here as a valid species in Asolene,
following Hylton Scott (1958: 310) and
Berthold (1991: 250).

roissil
Ampullaria roissii Orbigny, 1838c, pl. 52, figs.
1-3. Unavailable name; incorrect original
spelling of roissyi Orbigny, 1838.

roissyi
Ampullaria roissii Orbigny, 1838c, pl. 52, figs.
1-3 [given as “Roissyi by Orbigny (1841e:
377)]. New name for crassa Orbigny, 1835,
non Swainson, 1823, non Deshayes, 1830.
Distribution: Rio Parana, Argentina (Orbigny,
1835a: 33).

Remarks. The name roissyi, as given by
Orbigny (1841e: 377), was an incorrect sub-
sequent spelling (Code, Art. 33.3); it was not
an emendation, as it was not demonstrably
intentional (Code, Ан. 33.2). However, “roissy/’
is in prevailing use, attributed to Orbigny, and
is therefore deemed the correct original spell-
ing (Code, Art. 33.3.1). Variety of cyclostoma
Spix, 1827, teste Sowerby (1909a: 348). Syn-
onym of cyclostoma Spix, 1827, teste Pain
(1960: 430). Synonym of pulchella Anton,
1838, teste Philippi (1852a: 33), Ihering (1898:
50, 1919: 337) and Hylton Scott (1958: 310),
followed here.

sloanii
Ampullaria Sloanii Férussac, 1827: 413. Un-
available name; nom. nud.

Remarks. Listed as from “Cayenne” by
Férussac (1827: 413), Jay (1836: 47; 1839:

NEW WORLD AMPULLARIIDAE 49

65; 1850: 283), Drouët (1859: 84), Gaudion
(1879: 40) and Paetel (1888: 481). Listed as
a synonym of crassa Swainson, 1823, by
Tillier (1980: 21).
solida

Ampullaria solida Busch, 1859: 168. Ecuador.
Syntypes: BMNH 20020683 (2 spms.). Distri-
bution: Ecuador (Miller, 1879: 149).

Remarks. Synonym of crassa Swainson,

1823, teste Pain (1960: 429).

sowerbyi
Ampullaria sowerbyi Vernhout, 1914a: 29, pl.
1, fig. 13 [holotype]. Lawa. Holotype: RMNH.
Distribution: Surinam.

Remarks. The description is explicitly
based on only a single specimen. The “type”
(i.e., the single specimen) is indicated as be-
ing in RMNH, and a specimen, which must be
this single specimen, is figured. Synonym of
granulosa Sowerby, 1894, teste Pain (1952:
30), Geijskes & Pain (1957: 45) and Tillier
(1980: 20).

spixii
Ampullaria Spixii Orbigny, 1838d: 376, pl. 52,
figs. 7, 8. New name for zonata Orbigny,
1835, non Spix, 1827. Distribution: “Sistema
del Plata у del rio Paraná ... Puerto Bertoni,
Alto Paraná, en Río Grande do Sul ... y en el
rio Uruguay” (Ihering, 1919: 336).

Remarks. Synonym of cyclostoma Spix,
1827, teste Sowerby (1909a: 348), but treated
here as a valid species, following Hylton Scott
(1958: 312) and Berthold (1991: 250).

storeria
Ampullaria Storeria Jay, 1839: 112, pl. 1, fig.
5. River Amazon [= Brasil]. Probable syntype:
AMNH 56107 (Boyko 8 Cordeiro, 2001: 16)
[labeled as “figd type” in Jay's handwriting (P.
M. Mikkelsen, pers. comm. to RHC, 7 May
2002)]. Distribution: Brasil.

Remarks. Considered a variety of platae
Maton, 1811, by Jay (1850: 283). Philippi
(1852a: 34, 63) could not decide its status as
a real species or a synonym of platae Maton,
1811. Treated here as a synonym of platae
Maton, 1811, following Martens (1857: 210)
and Gaudion (1879: 37).

zonata

Ampullaria zonata Orbigny, 1835a: 32. Rio
Parana (republica Argentina)...Lacubus
provincias Corrientes (republica Argentina).
Syntypes: ВММН 1854.12.4.327-329 (28
spms.) [labeled “зрих!”], MNHN (2 lots, 8
spms.), MCZ (2 spms.) [labeled as para-
types].

Remarks. Junior primary homonym of zonata
Spix, 1827; replaced by spixii Orbigny, 1838.

Genus FELIPPONEA Dall, 1919

FELIPPONEA Dall, 1919: 10. Type species:
Ampullaria (Felipponea) neritiniformis Dall,
1919, by monotypy.

Considered a synonym of Asolene Orbigny,
1838, by Pilsbry (1933: 74), but treated here as
a full genus with three included species, fol-
lowing Hylton Scott (1958: 317) and Berthold
(1991: 23, 250).

elongata
Ampullaria (Felipponea) elongata Dall, 1921:
133. Uruguay River, Dept. of Paysandu. Holo-
type: USNM 333024. Distribution: Uruguay.

Remarks. Junior primary homonym of

elongata Orbigny, 1842, which is here listed
under Pomacea Perry, 1810.

iheringi
Asolene iheringi Pilsbry, 1933: 73, pl. 2, figs.
7 [paratype], 8 [paratype], 9 [holotype], Эа [ho-
lotype]. Rapids of Butni, Rio Uruguay, be-
tween San Borja and Uruguayana, Rio
Grande do Sul, Brazil. Holotype [figured and
distinguished by measurements given in the
text]: ANSP 124615 ["124615a” (Baker, 1964:
168)]; paratypes: ANSP 365363 (3 spms.).
Distribution: Southern Brasil.

neritiniformis
Ampullaria (Felipponea) neritiniformis Dall,
1919: 10. Rio Uruguay, Department of
Paysandu. Holotype: USNM 332780;
topotype: ANSP 141211 (Pilsbry, 1933: 76).
Distribution: Uruguay River basin: Argentina,
Brasil, Uruguay (Hylton Scott, 1958: 318;
Faraco et al., 2002).

Genus MARISA Gray, 1824

MARISA Gray, 1824: 276. Type species: Helix
cornuarietis Linnaeus, 1758, by subsequent
designation of Gray (1847: 148).

CERATODES Guilding, 1828: 537, 540. Type
species: Helix cornuarietis Linnaeus, 1758,
by original designation.

Gray (1824: 276) established Marisa “for a
genus of shells which has been confused with
Ampullaria, but which differs from it in having a
horny operculum and simple peristome”. That
is, he was establishing Marisa for American
ampullariids, distinguishing them from Old
World species with a calcified operculum,
which are now placed in Pila Réding, 1798.
[Ampullaria Lamarck, 1799, is a junior objective
synonym of Pila Róding, 1798 (Cowie, 1997a;

50 COWIE & ТНЕМСО

ICZN, 1999a; and see below under Pomacea
Perry, 1810)]. Martens (1899: 424) took Gray's
Marisa to have been “intended for all American
Ampullariæ”, but incorrectly considered it jun-
ior to Ceratodes Guilding, 1828, because of
the date of establishment of cornuarietis
Linnaeus, 1758, as the type species. Dall
(1904: 52) misinterpreted Ampullaria as refer-
ring to American species with a horny opercu-
lum, and Marisa to “cover Ampullaria s.s.”
[what would now be called Pomacea Perry,
1810], and, incorrectly, to exclude
cornuarietis, which he considered, again in-
correctly, to have “so persistently and inaccu-
rately been asserted to be the type of Marisa”.
Modern usage restricts Marisa Gray, 1824, to
those species related to the type, cornuarietis
Linnaeus, 1758, and places the majority of the
remaining American species in Pomacea
Perry, 1810 (with a small number in Asolene
Orbigny, 1838, Felipponea Dall, 1919, and
Pomella Gray, 1847).

The type species of Marisa Gray, 1824, has
been considered as Marisa intermedia Gray,
1824, by monotypy (e.g., Pilsbry & Bequaert,
1927: 169; Baker, 1930: 11; Berthold, 1991:
249), whereas in fact cornuarietis Linnaeus,
1758, was also an originally included species,
as Pilsbry (1933: 72) realized. Dall (1904: 52)
misinterpreted Marisa Gray as not including
cornuarietis Linnaeus, 1758. Gray (1847: 148)
designated cornuarietis Linnaeus, 1758, as the
type of Marisa Gray. Ceratodes Guilding, 1828,
was established with two species included:
fasciatus Guilding, 1828, and cornuarietis
Linnaeus, 1758; the latter was explicitly desig-
nated as the type, contrary to the statement of
Berthold (1991: 249) that the type was estab-
lished by monotypy. Hence, Ceratodes
Guilding, 1828, is a junior objective synonym of
Marisa Gray, 1824.

Berthold (1991: 25, 159) included only two
species (cornuarietis Linnaeus, 1758, and
planogyra Pilsbry, 1933) in his summary of the
genus, but although he illustrated (рр. 12-13) a
third species, chiquitensis Orbigny, 1838, he
apparently considered this (p. 249) to be part of
the wide range of morphological variation in
cornuarietis Linnaeus, 1758.

chiquitensis
Ampullaria Chiquitensis Orbigny, 1838d: 367,
pl. 48, figs. 10, 11. sud-est de la province de
Chiquitos (république de Bolivia) ... entre les
Missions de San-Miguel e de San-José ... а
peu pres de la première Mission, dans le lac

de los Migueleños, et dans les marais des
environs. Syntypes: BMNH 1854.12.4.326 (9
spms.), ММНМ (1 spm.). Distribution: Bolivia
(Berthold, 1991: 13).

Remarks. Described in genus Ampullaria,
subgenus Ampullaria s. str., and within a sec-
tion (Ceratodes) composed of “Espèces
déprimées”, but as the binomen “Ceratodes
Chiquitensis”. Thus, the original combination
could be considered to be with either
Ampullaria or Ceratodes. Since the species
was explicitly described within the genus
Ampullaria, we prefer Ampullaria chiquitensis
as the original combination (cf. Petit 8
Harasewych, 1990: 69). Retained as a dis-

_ tinct species by Pilsbry (1933: 72) but here
considered a synonym of cornuarietis
Linnaeus, 1758, following Sowerby (1909a:
359) and Berthold (1991: 249).

contrarius
Planorbis contrarius Múller, 1774: 152. [No
locality given]. Syntypes: the specimens fig-
ured by Seba, as cited by Múller; locations

unknown, possibly Uppsala University
(Dance, 1986: 225).
Remarks. Synonym of cornuarietis

Linnaeus, 1758, teste Orbigny (1835a: 30),
Anton (1838: 50) and Gaudion (1879: 27).
cornuarietis

Helix Cornu arietis Linnaeus, 1758: 771. O.
Europæo [еггог; “probably Venezuela, but
certainly somewhere between the Guianas
and Colombia” (Pilsbry, 1933: 71)]. Type ma-
terial: the specimen(s) figured by Lister, re-
ferred to by Linnaeus (1758: 771) (Pilsbry,
1933: 71); lost (Dance, 1967: 21). Distribu-
tion: northern South America, including Co-
lombia, Venezuela, Guyana, Surinam [?
error; Geijskes 8 Pain (1957: 47)], French
Guiana, Bolivia, Brasil, Trinidad and Tobago
(Pilsbry, 1933: 71-72).

Remarks. lhering (1919: 333) appears to
have incorrectly recorded this species south
of the Amazon basin, from the Río Paraguay,
Río de la Plata, and Río Grande do Sul.

fasciatus
Ceratodes fasciatus Guilding, 1828: 540, pl.
supp. 28, figs. 4—7 in fluviis Атепсае æquinoc-
tialis. Type material: not found by us in BMNH
(cf. Dean, 1936: 234; Dance, 1986: 213).

Remarks. Synonym of cornuarietis
Linnaeus, 1758, teste Guppy (1866: 44), and
here retained as such despite being consid-
ered a subspecies (as knorrii Philippi, 1852)
of cornuarietis Linnaeus, 1758, by Baker
(1930: 26). Junior secondary homonym of

NEW WORLD AMPULLARIIDAE 51

fasciata Roissy, 1805, when both species are

placed т Pomacea (e.g., Baker, 1930: 26).
intermedia

Marisa intermedia Gray, 1824: 276. Brazils.

Syntype: ВММН 1895.11.6.1 [although

Berthold (1991: 249) stated “Typus

verschollen ist”].

Remarks. Pilsbry (1933: 72) considered
intermedia Gray, 1824, to be “doubtless an
Effusa, but ... unrecognizable specifically”.
Berthold (1991: 249) treated intermedia Gray,
1824 [Marisa] as different from intermedia
Férussac [Pomacea subgenus Effusa], con-
sidering the former (incorrectly) as the type of
the genus Marisa Gray. Either a junior syn-
onym of cornuarietis Linnaeus, 1758, or a
senior synonym of planogyra Pilsbry, 1933,
teste Berthold (1991: 249).

knorrii
Ampullaria Knorrii Philippi, 1852a: 57, pl. 18,
fig. 3 [1852b: 28]. die Insel Trinidad. Type
material: probably MNHNS. Distribution: Ven-
ezuela, Guyana, Surinam, Trinidad, Colom-
bia, Panama (Baker, 1930: 26).

Remarks. Synonym of cornuarietis
Linnaeus, 1758, teste Guppy (1866: 44) and
Sowerby (1909a: 359), or of fasciatus
Guilding, 1828, teste Philippi (1852a: 57) and
Baker (1930: 26).

planogyra

Marisa planogyra Pilsbry, 1933: 70, pl. 2, figs.
2-5a. Santa Rosa, in the Descalvados region
of Matto Grosso. Holotype: ANSP 158776
[158776a (Baker, 1964: 168)]; paratypes:
ANSP 158780 (16 spms.), 158787 (19
spms.), 365366 (3 spms., from holotype lot),
MCZ (1 spm.). Distribution: Brasil.

rotula
Ampullaria rotula Mousson, 1869: 183.
unteren Magdalenstrome [= Lower

Magdalena river, Colombia]. Syntypes: ZMZ
525321 (3 spms.). Distribution: Panama,
Costa Rica (Pilsbry, 1933: 71), Colombia
(Martens, 1899: 425).

Remarks. Mousson (1873: 19) placed it in
Ceratodes Guilding, 1828. Synonym of
cornuarietis Linnaeus, 1758, teste Sowerby
(1909a: 359), and retained here as such de-
spite being considered a distinct, closely re-
lated species or subspecies of cornuarietis
Linnaeus, 1758, by Pilsbry (1933: 71).

Genus POMACEA Perry, 1810

POMACEA Perry, 1810c: [unnumbered plate
and text] [= pl. 12 (Mathews & lredale, 1912:

11; Clench & Turner, 1956: 120; Geijskes &
Pain, 1957: 42; R. E. Petit, pers. comm. to
RHC, 16 October 2000); not pl. 11, as stated
by Cowie (1997a: 84)]. Type species:
Pomacea maculata Perry, 1810, by mono-

typy.

CONCHYLIUM Cuvier, 1816: 426. Type spe-
cies Nerita urceus Muller, 1774, by subse-
quent designation of Pilsbry & Bequaert
(1927: 170) [as Bulimus urceus Bruguiere].

LIMNOPOMUS Dall, 1904: 52. Type species:
Ampullaria columellaris Gould, 1848, by origi-
nal designation.

The status of the genus-group names
Ampullaria Lamarck, 1799, and Ampullarius
Montfort, 1810, both frequently used incorrectly
in combination with names of species of
Pomacea, have been clarified by Cowie
(1997a) and ICZN (1999a) as junior objective
synonyms of Pila Róding, 1798 (see also Pain,
1956b: 79). Pomus “Humph.” Gray, 1847, is
also a junior objective synonym of Pila Róding,
1798. Limnopomus Dall, 1904, is here treated
as a synonym of Pomacea Perry, 1810, follow-
ing Berthold (1991).

The distinction between the two subgenera
Pomacea s. str. and Pomacea (Effusa) is not
clear. Only those species that have been ex-
plicitly placed in subgenus Effusa are listed
under that heading. Others whose placement
is uncertain are listed under Pomacea s. Str.,
pending further research. Many of the more
obscure species-group names have never be-
fore been placed in combination with the ge-
nus-group name Pomacea, because of the
traditional but incorrect use of the genus-group
name Ampullaria for these American species
(Cowie, 1997a; ICZN, 1999a). Hence many of
the species listed here are probably new com-
binations with Pomacea.

Subgenus EFFUSA Jousseaume, 1889

EFFUSA Jousseaume, 1889: 255. Type spe-
cies: Helix glauca Linnaeus, 1758, by subse-
quent designation of Baker (1930: 11).

Baker (1930: 20) considered /uteostoma
Swainson, 1823, to be distinct from glauca
Linnaeus, 1758, but the /uteostoma of most
other authors, including Jousseaume (1889:
255), to be misidentifications of glauca
Linnaeus, 1758. Subsequent authors have syn-
onymized luteostoma Swainson, 1823, with
glauca Linnaeus, 1758 (see below).

52

baeri

Ampullaria Baeri Dautzenberg, 1902: 312, pl.
9, figs. 12, 13. Rio Mixiollo [= Misciotto;
Berthold, 1991: 13], province de Huallaga,
Pérou. Lectotype (Fischer-Piette, 1950: 170):
ММНМ; paralectotypes: ANSP 99328 (1
spm.), MCZ (1 spm.), UMMZ 46767 [? error],
ZMHB 59269 (1 Spm.); possible
paralectotypes: ZMHB 63631 (2 spms.),
109517 (1 spm.) (M. Glaubrecht, pers.
comm. to RHC, 1 March 2003). Distribution:
Peru (Berthold, 1991: 13).

Remarks. Probably synonymous with
glauca Linnaeus, 1758, teste Boss &
Parodiz (1977: 116), but not definitively syn-
onymized.
balteata
Ampullaria balteata Philippi, 1851: 21, pl. 5,
fig. 7 [1852b: 22]. [No locality given. Trinidad
“chosen” by Baker (1930: 25).] Lectotype
(Baker, 1930: 25): the specimen illustrated in
“Philippi's first figure” [= pl. 5, fig. 7], probably
MNHNS. Distribution: “Trinidad; also shells
from Venezuela ..., Colombia ..., Tobago ...,
and Martinique ... that are intermediate be-
tween this form and neritina” (Baker, 1930:
25); also “Venezuela - Guyane - Maroni -
Orenoque” (Gaudion, 1879: 24).

Remarks. Baker (1930: 25) explicitly ex-
cluded the later figures of Philippi (1852a: 55,
pl. 17, fig. 4). Although the precise origin of
the designated lectotype is unknown, Baker's
choice of Trinidad as the type locality follows
Code Rec. 76A.1.4. Synonym of /uteostoma
Swainson, 1823 [= glauca Linnaeus, 1758],
teste Paetel (1887: 477) and Alderson (1925:
6), but treated as a form of glauca Linnaeus,
1758, by Sowerby (1909a: 350) and Baker
(1930: 25). Synonym of glauca Linnaeus,
1758. N. syn.

castanea

Ampullaria castanea Deshayes, 1830a: 31.
[No locality given.] Syntype: ММНМ. Distribu-
tion: “Orinocco” (Philippi, 1852a: 41); “La
Guyane - Haut-Brésil = Haut-Amazone”
(Gaudion, 1879: 26); “Guyana” (Paetel, 1873:
64, 1887: 477); “unknown” (Baker, 1930: 22).
Remarks. Synonym of luteostoma
Swainson, 1823 [= glauca Linnaeus, 1758],
teste Jay (1850: 36), or possibly of neritina
Gmelin, 1791 [= glauca Linnaeus, 1758],
teste Baker (1930: 22). Synonym of glauca
Linnaeus, 1758. N. syn.
chlorostoma
Ampullaria chlorostoma Sowerby, 1825: 44.
Unavailable name; first published as a junior

COWIE & THIENGO

synonym of /uteostoma Swainson, 1823, not
made available before 1961 (Code Art. 11.6).

cingulata

Ampullaria cingulata Philippi, 1851: 19, pl. 5,
fig. 3 [1852b: 22]. [No locality given.] Syntype:
ZMHB 1376 (1 spm.) (M. Glaubrecht, pers.
comm. to RHC, 1 March 2003). Distribution:
Venezuela (Martens, 1857: 203; 1873: 202).

Remarks. Sowerby (1909a: 347) consid-
ered it a “doubtful species which may possi-
bly be young of A. gigas”, but it was
considered a valid species and placed in
subgenus Effusa by Baker (1930: 10). Baker
(1930: 10) considered “Lago de Valencia,
Ven. [= Venezuela]” as the type locality. This
was probably correct because Philippi (1851:
19) described the species from material in
the Berlin Museum, and Martens (1873: 202)
gave this as the only known locality for mate-
rial in the Berlin Museum. However, Martens
(1873: 202) also indicated that some of the
material was without locality data. Strictly
then, the type locality probably includes the
above location but may not be restricted to it.

conica

Ampullaria effusa variety conica Guppy,
1866: 44 [by bibliographic reference to
Guppy, 1864: 244]. Trinidad. Type material:
probably Victoria Inst., Trinidad [destroyed],
not found by us in BMNH (cf. Dance, 1986:
213). Distribution: Trinidad.

Remarks. Junior primary homonym of
conica Lamarck, 1804 [also 1822] (now
placed in family Naticidae), conica Swainson,
1823 [= virens Lamarck, 1822 (Philippi
(1852а: 73)] (now placed in Pila Róding,
1798), and conica Wood, 1828 [Ampullaria
conica selected as the correct original com-
bination by Cowie (1997b: 4)] (now placed in
Pila Róding, 1798). Synonym of neritina
Gmelin, 1791 [= glauca Linnaeus, 1758],
teste Baker (1930:22). Synonym of glauca
Linnaeus, 1758. М. syn.

crocostoma

Ampullaria crocostoma Philippi, 1852а: 42, pl.
12, fig. 3 [1852b: 26]. Caraccas. Possible
syntypes: ZMHB 109501 (3 spms.) (M.
Glaubrecht, pers. comm. to RHC, 1 March
2003) [the largest shell is very similar to the
original figure (F. Kóhler, pers. comm. to RHC,
6 March 2003)]; possibly also MNHNS. Distri-
bution: Venezuela, Guyana (Pain 1950b: 71).

Remarks. Synonym of glauca Linnaeus,
1758, teste Baker (1930: 18) and
Starmühlner (1988: 253), followed here, al-
though treated as a variety of that species by

NEW WORLD AMPULLARIIDAE 53

Pain (1950b: 69). See also Boss & Parodiz
(197-72 116).

cuprina

Ampullaria cuprina Reeve, 1856e: pl. 1, fig. 1.
[No locality given] Syntypes: BMNH
20020652 (2 spms.). Distribution: unknown.

Remarks. Synonym of glauca Linnaeus,
1758, teste Starmúhlner (1988: 253), fol-
lowed here, although considered a variety of
that species by Sowerby (1909a: 351).
dubia
Ampullaria dubia Guilding, 1828: 539, pl.
supp. 27, figs. 7, 8. in fluviis Americæ
æquinoctialis ... small river in the Gulph of
Paria ... canals of Demerara [Baker (1930:
15-16), in designating the lectotype, re-
stricted the type locality to the “Gulf of Paria,
probably one of distributaries of Rio
Orinoco”]. Lectotype (Baker, 1930: 15): the
specimen in Guilding's fig. 7, not found by us
in BMNH (cf. Dean, 1936: 234; Dance, 1986:
213). Distribution: Guyana, Surinam, Rio
Orinoco, St. Lucia (Lesser Antilles),
Guadeloupe (Baker, 1930: 16).

Remarks. Synonym of /uteostoma
Swainson, 1823 [= glauca Linnaeus, 1758],
teste Sowerby (1909a: 350). Synonym of
glauca Linnaeus, 1758, teste Starmúhlner
(1988: 254), followed here (see also Baker,
1930: 12, 15), despite Alderson (1925: 3) and
Pain (1950b: 70) considering it unidentifiable.
effusa
Nerita effusa Müller, 1774: 175. [No locality
given. Rio Yaracuy, Ven. [= Venezuela] cho-
sen by Baker (1930: 17).] Syntypes: the
specimens figured by Seba and Geve, as
cited by Múller, and the specimens *In Museo
Moltkiano” (Múller, 1774: 176; see also Baker,
1930: 17) [not in the Copenhagen Museum
(O. S. Tendahl, pers. comm. to RHC, 18 April
2002)]; not the specimen illustrated by Lister,
as cited by Müller, nor ANSP 50596 (Baker,
1930: 17). Distribution: French Guiana [? er-
ror], Surinam [? error] (Drouét, 1859:79),
Martinique (Saulcy, 1854: 141; Paetel, 1887:
478), Venezuela (Baker, 1930: 17), Guyana
(Pain, 1950b: 65).

Remarks. Baker (1930: 17) considered the
locality of ANSP 50596 as the type locality,
but although this action was not a valid neo-
type designation (Code Art. 75) and the origin
of the true type material is unknown, his
choice of the type locality appears to follow
Code, Rec. 76A.1.4. Synonym of glauca
Linnaeus, 1758, teste Swainson (1823a: pl.
157), Philippi (1852a: 43) and Starmühlner

(1988: 253), followed here (see also Baker,
1930: 12, 17), contrary to Sowerby (1909a:
350) who treated it as a variety of that spe-
cies. Gmelin (1791: 3626) listed effusa
Múller, 1774, as variety “y” of ampullacea
Linnaeus, 1758, which is now placed in Pila
Röding, 1798. However, Philippi (1852a: 43)
and Starmühlner (1988: 253), in listing
Gmelin's variety as a synonym of glauca
Linnaeus, 1758, gave it as “ampullaria var. y.
Gm” and “Helix ampullaria var. |", respec-
tively; and Gaudion (1879: 29), in listing it as
a synonym of effusa Múller, 1774, gave it as
“Helix ampullaria var Gmel”. These usages
of “ampullaria” are misspellings of
“ampullacea”.

expansa

Ampullaria expansa Miller, 1879: 152, pl. 15,
fig. 6. Rio Santiago prope Playa de oro, in
provincia Esmeraldas. Type material: location
not known to us. Distribution: Ecuador (Miller,
1879: 152; Sowerby, 1909a: 349).

Remarks. Placed in subgenus Effusa fol-
lowing Kobelt (1913a: 147), who placed it in
his “Formenkreis der Ampullaria glauca L.
(Effusae Martens)”. Junior primary homonym
of expansa Nevill, 1877, which is now placed
in Pila Róding, 1798.

geveana

Ampullaria Geveana Philippi, 1852a: 26.
Unjustified emendation of gevesensis
Deshayes, 1838.

Remarks. Philippi (1852a: 26) explicitly
made the emendation. However, the original
name is here considered a result of incorrect
latinization, which is not treated as an inad-
vertent error and therefore not a justification
for emendation (Code Art. 32.5.1). As an
emendation, geveana Philippi, 1852, is avail-
able and a junior objective synonym of
gevesensis Deshayes, 1838 (Code Art.
33.2.3.), and hence a synonym of glauca
Linnaeus, 1758, as indicated by Starmühiner
(1988: 253).

gevesensis

Ampullaria Gevesensis Deshayes, 1838:
541. [No locality given.] Syntype: ММНМ. Dis-
tribution: French Guiana (Sowerby 1909a:
350), Guyana, Venezuela “in all probablility …
from Venezuela in the north, southwards to
the Amazon Valley” (Pain, 1950b: 71),
Surinam (Pain, 1952; 31; Geijskes 8 Pain,
1957: 44).

Remarks. Variety of glauca Linnaeus, 1758,
teste Sowerby (1909a: 350 [as
“Geveanensis”]), Pain (1950b: 69) and

54

Geijskes & Pain (1957: 44). Зупопут of
effusa Müller, 1774 [= glauca Linnaeus,
1758], teste Baker (1930: 17). Synonym of
glauca Linnaeus, 1758. N. syn.

glauca

Helix glauca Linnaeus, 1758: 771. [No locality
given. Rio Tuca, near Tucacas, Venezuela
“chosen” by Baker (1930: 12, 18).] Type mate-
rial: lost (Dance, 1967: 21). Distribution
(glauca Linnaeus, 1758, and its varieties):
Brasil, Bolivia, Colombia, Venezuela, Guyana,
Surinam, French Guiana, Trinidad, Grenada,
Barbados, Guadeloupe, Dominica, Martinique,
St. Lucia (Vernhout, 1914a: 43; Pain, 1950b:
69; Geijskes & Pain, 1957: 44; McKillop &
Harrison, 1980: 271; Tillier, 1980: 24;
Starmühiner, 1984: 89-91, 1988: 254).

Remarks. The designation by Baker (1930:
19) of Knorr’s figure as a neotype [“type”] was
invalid (Code Art. 75). However, although the
origin of the type material is unknown, Baker's
choice of the type locality appears to follow
Code, Rec. 76A.1.4. Baker (1930: 12-13) rec-
ognized nine “forms” of glauca Linnaeus, 1758,
considering that “at least the first six of these
are not geographic subspecies”. Some of
these “forms” are here treated as synonyms,
others as undetermined infraspecific taxa. Be-
cause glauca Linnaeus, 1758, is highly variable
(e.g., Arias, 1952: 64) revisionary study would
probably synonymize all nine “forms”.
guadelupensis
Ampullaria guadelupensis Martens, 1857:
199. Caripe auf Guadeloupe. Syntypes:
ZMHB 1385 (2 spms.) (M. Glaubrecht, pers.
comm. to RHC, 1 March 2003); no type mate-
rial found by us in BMNH or MCZ (cf. Dance,
1986: 218). Distribution: Guadeloupe.

Remarks. Synonym of glauca Linnaeus,
1758, teste Baker (1930: 18) and Star-
mühlner (1988: 253).
intermedia
Ampullaria intermedia Férussac, in Quoy &
Gaimard, 1825d: 489, pl. 68, figs. 1-3. Brésil.
Syntypes: MNHN (2 spms.). Distribution:
Brasil (Berthold, 1991: 12).

Remarks. Synonym of sordida Swainson,
1823, teste Orbigny (1835a: 31), Philippi,
(1852a: 38), Sowerby (1909a: 357) and
Thiengo (1989: 351), followed here, although
contrary to Berthold (1991: 23), who treated it
as a valid species in subgenus Effusa.
luteostoma
Ampullaria luteostoma Swainson, 1823a: pl.
157, top and bottom figs. [No locality given.]
Type material: possibly MMUE (Dean, 1936:

COWIE & THIENGO

232; H. McGhie, pers. comm. to RHC, 29
July 2002), not found by us in BMNH (cf.
Dance, 1986: 227). Distribution: Venezuela,
Guyana, French Guiana, Martinique,
Guadeloupe (Pain, 1950b: 71).

Remarks. Variety of glauca Linnaeus, 1758,
teste Pain (1950b: 69, 71). Synonym of
glauca Linnaeus, 1758, teste Boss &
Parodiz (1977: 116), followed here.

+ minuscula

Pomacea (Effusa) glauca form minuscula
Baker, 1930: 24, pl. 30, fig. 8. Quebrada
Sucremo, a small, swampy brook in heavy
forest near Boquerón, Venezuela (station
number “H, VIII, b, 29”). Holotype: UMMZ
92069; paratypes: ANSP 147706 (2 spms.),
MCZ (1 lot, 3 spms.). Distribution: Venezu-
ela.

Remarks. Retained as a distinct infraspe-
cific taxon, following Baker (1930: 12, 24) and
pending further research.

neritina

Helix neritina Gmelin, 1791: 3638. [No locality
given. Belmont, near Port of Spain, Trinidad
“chosen” by Baker (1930: 22).] Holotype: the
specimen illustrated in “Kaemerer Conch.
Rudolst. p. 185. n. 2. t. 11. f. 7.” (cited by
Gmelin), location not known to us. Distribu-
tion: ? Colombia, Venezuela, Trinidad, To-
bago, St. Lucia, Martinique, Guadeloupe
(Baker, 1930: 22).

Remarks. Synonym of glauca Linnaeus,
1758, teste Philippi (1852a: 43), Paetel
(1887: 480) and Starmühlner (1988: 253)
(see also Baker, 1930: 12, 22). Although the
locality of the holotype is unknown, Baker’s
choice of the type locality appears to follow
Code, Rec. 76A.1.4.

oculuscommunis

Helix oculus communis Gmelin, 1791: 3621.
[No locality given. Rio Yaracuy, Venezuela,
“chosen” by Baker (1930: 14).] Lectotype
(Baker 1930:14): the specimen illustrated by
“Seba (Thes., pl. 40, figs. 3-5)”; paralecto-
types: the specimens illustrated in the other
figures cited by Gmelin (1791: 3621). Distri-
bution: Venezuela, Guyana, French Guiana
(Baker, 1930: 14).

Remarks. Baker (1930: 14) designated the
lectotype; and, although the locality of this
specimen is unknown, his choice of a type
locality appears to follow Code Rec. 76A.1.4.
Synonym of glauca Linnaeus, 1758, teste
Philippi (1852a: 43), Sowerby (1909a: 350)
and Starmühlner (1988: 253) (see also Bak-
er, 1930: 12, 14). Synonym of gevesensis

NEW WORLD AMPULLARIIDAE 55

Deshayes, 1838 [= glauca Linnaeus, 1758],
teste Pain (1950b: 69).
oligista

Pomacea (Effusa) oligista Pilsbry & Olsson,
1953: 98, pl. 6, fig. 6. on the road ... from
Cartagena to Barranquilla ... a freshwater lake
known as the Cienaga de Luruaco. Holotype:
ANSP 189546 ["189546a” (Baker, 1964: 168)];
paratypes: ANSP 189547 (10 spms.), 365367
(3 spms., figured). Distribution: Colombia.

Remarks. Pilsbry & Olsson (1953: 98) gave
measurements of the “type” and the largest
“paratype” but gave no catalog numbers.
Synonym of planorbula Philippi, 1852, teste
Pain (1956a: 76-77).

orinoccensis

Ampullaria orinoccensis Troschel, 1848: 548.
am обет Pomeroon. Syntypes: ZMHB 1384a
(1 spm.), 1384b (3 spms.), 1384c (2 spms.)
(M. Glaubrecht, pers. comm. to RHC, 1
March 2003); possible syntypes: MCZ [la-
beled as paratypes, but as “oronocoensis
Reeve”]; type material possibly also in the
Dohrn collection, Stettin Museum [destroyed;
Dance, 1986: 210: 229]; no type material
found by us in BMNH (cf. Dance, 1986: 210).
Distribution: Guyana, Surinam, Venezuela
(Vernhout, 1914a: 43; Pain, 1952: 30-31;
Geijskes & Pain, 1957: 45), French Guiana
(Tillier, 1980: 27).

Remarks. Name attributed to Ziegler by
Troschel (1848: 548). Variously misspelled,
e.g., as “Oronocensis” by Reeve (1856b: pl.
10, fig. 45) and “Orinocensis” by Martens
(1873: 204). Synonym of dubia Guilding,
1828 [= glauca Linnaeus, 1758], teste Baker
(1930: 15). Variety or subspecies of glauca
Linnaeus, 1758, teste Pain (1950b: 70; 1952:
31), Geijskes & Pain (1957: 44) and Tillier
(1980: 26). Synonym of glauca Linnaeus,
1758, teste Starmühlner (1988: 253 [as
“oronocensis Reeve”]), followed here.

pachystoma
Ampullaria pachystoma Philippi, 1849: 17.
Brasilia [? error]. Type material: probably
MNHNS. Distribution: Brasil [? error].

Remarks. Synonym of /uteostoma
Swainson, 1823 [= glauca Linnaeus, 1758],
teste Paetel (1887: 480) and Alderson (1925:
6). Variety of glauca Linnaeus, 1758, teste
Sowerby (1909a: 350). Retained as a distinct
species by Baker (1930: 16-17). Synonym of
glauca Linnaeus, 1758, teste Starmühlner
(1988: 253), followed here.

pattersoni
Pomacea (Effusa) pattersoni Boss &
Parodiz, 1977: 112, figs. 7-9. Vicinity of

Yarina (6°172’ $; 75°172’ W), upstream from
Isla Navarro, close to Río Huallaga, Depart-
ment of San Martín, Peru. Holotype: MCZ
272900; paratype: MCZ 272918. Distribution:
Peru.
philippiana

Pomacea (Effusa) glauca form philippiana
Baker, 1930: 14. canal near Georgetown,
British Guiana. Holotype ANSP 70016
["170016a” (Baker, 1964: 168)]; paratypes:
ANSP 365368 (14 spms.). Distribution:
Surinam to Orinoco and Guadeloupe (Baker,
1930: 14).

Remarks. Synonym of glauca Linnaeus,

1758 (see Baker, 1930: 12, 14). N. syn.

+ planorbula

Ampullaria planorbula Philippi, 1852a: 26, pl.
7, fig. 3 [1852b: 23]. [No locality given.]
Syntype: ZMHB 2131 (M. Glaubrecht, pers.
comm. to RHC, 1 March 2003). Distribution:
“Рауа” (Paetel, 1888: 481), “Para.”
(Sowerby, 1909a: 359).

Remarks. Retained as a distinct infraspe-
cific taxon, following Baker (1930: 12, 24) and
pending further research. Pilsbry (1933: 72)
considered it “to be the young stage of some
variety of P. (Effusa) glauca (L.)".

prunulum
Ampullaria prunulum Reeve, 1856c: pl. 18,
fig. 82. New Granada [in 1856 = present-day
Colombia and Panama]. Syntypes: BMNH
20020679 (3 spms.). Distribution: Colombia
and/or Panama.

Remarks. Synonym of glauca Linnaeus,
1758, teste Starmühlner (1988: 253), fol-
lowed here, although considered a variety of
that species by Pain (1950b: 69).

quinindensis
Ampullaria quinindensis Miller, 1879: 151, pl.
15, fig. 5. Rio Quinindé qui influit in fluminem
Esmeraldas. Type material: location not
known to us. Distribution: Ecuador (Miller,
1879: 152; Sowerby, 1909a: 357 [as
“quinquidensis’]).

rhodostoma
Ampullaria rhodostoma Appun, 1871: 141,
548. Unavailable name; nom. nud.

Remarks. Treated as a synonym of
luteostoma Swainson, 1823, by Alderson
(1925: 6).

suprafasciata
Ampullaria geveana var. suprafasciata
Kobelt, 1913b: 157, pl. 57, figs. 7, 8. [No lo-
cality given.] Type material: possibly SMFD,
ZMHB (Dance, 1986: 215), but not found in
ZMHB (M. Glaubrecht, pers. comm. to RHC,
1 March 2003).

56

Remarks. Synonym of glauca Linnaeus,
1758, teste Baker (1930: 18).
tamsiana
Ampullaria Tamsiana Philippi, 1852a: 51, pl.
16, figs. 1, 2 [1852b: 27]. Puerto Cabello.
Syntype: ZMHB 109502 (1 spm.) [= pl. 16, fig.
2], 109503 (2 spms.); possible syntypes
109503 (2 spms.) [one of these ? = pl. 16,
fig. 1] (M. Glaubrecht, pers. comm. to ВНС, 1
March 2003; Е. Kóhler, pers. comm. to RHC,
6 March 2003), possibly also ММНМ$. Distri-
bution: Venezuela (Berthold, 1991).

Remarks. Name attributed to Dunker by
Philippi (1852a: 51; 1852b: 27). Synonym of
glauca Linnaeus, 1758, teste Baker (1930:
18, 20).
teres
Ampullaria teres Philippi, 1849: 19. [No local-
ity given.] Syntype: ZMHB 109504 (M.
Glaubrecht, pers. comm. to RHC, 1 March
2003); type material possibly also in MNHNS.
Distribution: Cuba [? error] (Gaudion, 1879:
40; Paetel, 1888: 482), “La Plat.” [? error]
(Paetel, 1873: 65).

Remarks. “Form” of glauca Linnaeus,
1758, teste Pilsbry (1927a: 251). Synonym of
neritina Gmelin, 1791 [= glauca Linnaeus,
1758], teste Baker (1930: 22) (see also
Baker, 1930: 12). Synonym of glauca
Linnaeus, 1758. N. syn.
tristis
Ampullaria effusa variety tristis Guppy, 1866:
44. Trinidad [in title of publication]. Type mate-
rial: probably Victoria Inst., Trinidad [de-
stroyed], not found by us in BMNH (cf. Dance,
1986: 213). Distribution: Trinidad.

Remarks. Synonym of neritina Gmelin, 1791
[= glauca Linnaeus, 1758], teste Baker
(1930: 22) (see also Baker, 1930: 12). Syn-
onym of glauca Linnaeus М. syn.

villata

A. villata Sowerby, 1909a: 350. Unavailable
name; first published as a junior synonym of
gevesensis Deshayes, 1838 [as “деуеапеп-
sis”], not made available before 1961 (Code
Art. 11.6).

Remarks. Name attributed to Martens by
Sowerby (1909a: 350) and listed as a syn-
onym of gevesensis Deshayes, 1838 [=
glauca Linnaeus, 1758]. Not listed under Mar-
tens’ authorship by Ruhoff (1980: 564), the
Zoological Record or Kabat 8 Boss (1997:
365).

Subgenus POMACEA Perry, 1810

Details as for genus Pomacea Perry, 1810.

COWIE 8 THIENGO

acuta

Ampullaria acuta Paetel, 1873: 64 [1887:
476]. Unavailable name; nom. nud.
Remarks. Name attributed to Menke by
Paetel (1873: 64, 1887: 476), and by Gaudion
(1879: 23), with locality “Vera Cruz”. Not
listed by Sowerby (1916: 70), Sherborn
(1922-1933) or Ruhoff (1980: 123).

aldersoni

Pila (Pomacea) aldersoni Pain, 1946a: 180;
pl. 6, figs. 1, 2. Ecuador, in a marsh near
Santa Barbara, about 170 miles S.E. of
Quito. Holotype and paratype (distinguished
as such in the text and the only two speci-
mens on which the description was based):
BMNH 1946.6.24.25 (1 spm.), possibly
NMW.Z.1981.118.00091 (1 spm.) ог
NMW.1955.158.02411 (Melvill-Tomlin collec-
tion, 1 spm.). Distribution: Ecuador.

Remarks. The original label of NMW.
1955.158.02411 states that it was collected in
November 1939, whereas the type series
was collected in January 1939; however, it
does say “co-type”. NMW.Z.1981.118.00091 is
small compared to the type dimensions and is
not the specimen figured (H. Wood, pers.
comm. to RHC, 30 October 2001).

amazonica

Ampullaria Amazonica Reeve, 1856b: pl. 12,
fig. 55. River Amazon. Syntype: BMNH
20020645. Distribution: Amazon (Sowerby,
1909a: 346).

angulata

Ampullaria angulata Jay, 1836: [85 (explana-
tion of pl. 3)], pl. 3, fig. 7. Mexico [error].
Syntype: AMNH 56108 (Boyko & Cordeiro,
2001: 16) [labeled as “figd pl. 3, fig. 7” in
Jay’s handwriting (P. M. Mikkelsen, pers.
comm. to RHC, 7 May 2002)].

Remarks. Synonym of scalaris Orbigny,
1835, teste Jay (1839: [116]), Martens (1857:
202), Gaudion (1879: 23) and Ihering (1898:
48), which has never been found as far north
as Mexico. We therefore consider Jay’s local-
ity to be incorrect.

angulata

Ampullaria angulata Dunker, 1845: 188.
reipublicae Argentinae. Type material: not in
ZMHB (Е. Kóhler, pers. comm. to RHC, 6
March 2003), not found by us in BMNH (cf.
Dance, 1986: 210).

Remarks. Junior primary homonym of
angulata Jay, 1836.

angulata

Ampullaria angulata Deshayes, 1850: 45, pl.
72, fig. 23. [No locality given.] Type material:
possibly Ecole des Mines, not found by us in

NEW WORLD AMPULLARIIDAE 57

ВММН or ММНМ (cf. Dance, 1986: 210). Dis-
tribution: unknown.

Remarks. Junior primary homonym of
angulata Jay, 1836, and angulata Dunker,
1845. Probably a synonym of scalaris
Orbigny, 1835, and hence retained here as a
South American species.
angulata
Pomus angulata H. Adams & A. Adams,
1854c: 347. Unavailable name; пот. nud.

Remarks. Name attributed to Jonas by H.
Adams 8 A. Adams (1854c: 347). Not listed
ру Sowerby (1916: 70), Sherborn (1922-
1933) or Trew (1992: 16). Perhaps the attri-
bution to Jonas was in error, or Jonas'
concept of the species was а mis-
identification of one of the three taxa listed
above.

arata

Ampullaria malleata var. Arata Fischer &
Crosse, 1890: 235 [1888: pl. 44, fig. 6d, бе;
plate published without name]. in Laguna de
los Cocos, provinciæ Vera Cruz ..., in
paludibus prope Palizada et San Geromino,
provinciæ Yucatan ..., in paludibus fluminis
Usumasinta, prope Balancan, provincias
Tabasco. Type material: Sallé collection, not
found by us in BMNH, ММНМ, etc. (cf. Dance,
1986: 209, 225). Distribution: Mexico.

Remarks. Synonym of flagellata Say, 1829,
teste Baker (1922: 37) and Pain (1964: 227).
+ archimedes
Ampullaria Archimedes Spix, in Wagner,
1827: 1, pl. 2, fig. 2. [No locality given.] Type
material: probably lost (Philippi, 1851: 10;
Fechter, 1983: 221; S. C. Thiengo, unpub-
lished). Distribution: unknown.

Remarks. Authorship is given here as “Spix,
in Wagner”, following Cowie et al. (in prep.),
who also explain the publication history of this
work. Spix illustrated archimedes as a full
species, but Wagner, in writing the descrip-
tions, treated archimedes “Spix” as a variety
of zonata “Wagner”. We retain it as an in-
fraspecific taxon of zonata Spix, 1827, follow-
ing Philippi (1851: 10) and Sowerby (1909a:
359), though they were synonymized by Mar-
tens (1857: 202).
armeniacum
Ampullaria armeniacum Hupé, 1857: 69, pl.
13, fig. 5. le fleuve des Amazones. Distribu-
tion: “Haut-Amazone” (Gaudion, 1879: 24).
Type material: not found by us in MNHN (cf.
Dance, 1986: 214).
aulanieri
Ampullaria Aulanieri Deville & Huppé, 1850:
642, pl. 15, fig. 4. lac de Cruz Playa, sur la

rivière de Ucayali (Pérou). Syntypes: MNHN
(3 lots, 15 spms.). Distribution: Peru (Pain,
1960: 424).

auriformis

Ampullaria auriformis Reeve, 1856e: pl. 28,
fig. 133a, b. Honduras. Syntype: BMNH
20020646. Distribution: Honduras.

Remarks. May be a variety of hopetonensis
Lea, 1834, teste Sowerby (1909a: 346), but
note the skepticism of Pain (1964: 225) re-
garding this.

aurostoma

Ampullaria aurostoma Lea, 1856: 110.
Carthagena. Syntypes: USNM 106299 [fig-
ured by Lea (1866, pl. 22, fig. 4), labeled in
the USNM as the holotype], USNM 106273 (11
spms.) [labeled as paratypes], MCZ (1 lot) [la-
beled as paratypes]. Distribution: Mexico [?
error] (Paetel, 1887: 477), Colombia, Ven-
ezuela (Baker, 1930: 9; Pain, 1956a: 78).
Remarks. Also published by Reeve (1856e:
pl. 28, fig. 131a, b), who said “Lea MS”. Re-
tained as a distinct species following Baker
(1930: 8) and Pain (1956a: 78), contrary to
Sowerby (1909a: 347) and Kobelt (1913a:
145) who synonymized it with cerasum
Hanley, 1854. Placed in subgenus
Limnopomus Dall by Baker (1930: 8).

australis

Ampullaria australis Orbigny, 1835a: 32.
lacubus Pampas meridionalibus Buenos-
Ayres (republica Argentina). Syntype: BMNH
1854.12.4.335. Distribution: Argentina.

Remarks. Synonym of insularum Orbigny,
1835, teste Sowerby (1909a: 353), or a vari-
ety of hopetonensis Lea, 1834 [= paludosa
Say, 1829], teste Sowerby (1909a: 352; but
see Pain, 1964: 225). Synonym of
canaliculata Lamarck, 1822, teste Hylton
Scott (1958: 299) and Cazzaniga (2002:
73);

autumnalis

Ampullaria autumnalis Reeve, 1856a: pl. 4,
fig. 16. [No locality given.] Syntype: BMNH
20020647. Distribution: unknown.

Remarks. Synonym of sordida Swainson,
1823, teste Sowerby (1909a: 357) and Kobelt
(1913h: 206).

avellana

Ampullaria avellana Sowerby, 1909a: 346
[name], 360 [description], text fig. Lagunella,
Venezuela. Syntypes: BMNH 1909.10.19.34
(1 spm.) (see also Sowerby, 1909a: 359),
HUJ 21519 (1 spm.) (H. Mienis, pers. comm.
to RHC, 4 August 2002), MHNG 1093/99 (2
spms.) (Y. Finet, pers. comm. to RHC, 22
August 2002). Distribution: Venezuela.

58

Remarks. Junior primary homonym of
avellana Lamarck, 1822 [not Ampullariidae
(Sowerby, 1825: 44; Jay, 1850: 294)].
batabana
Ampullaria batabana Paetel, 1887: 477. Un-
available name; nom. nud.

Remarks. Listed as from Cuba by Paetel
(1887: 477).
belizensis
Ampullaria Belizensis Crosse & Fischer, in
Fischer 4 Crosse, 1888: [explanation of] pl.
45, fig. 2, 2a-c [Crosse 8 Fischer, 1890: 110;
see also Fischer & Crosse (1890: 231, pl. 48,
fig. 9, 9a)]. [No locality given. in coloniá
anglicá Belize (Crosse 8 Fischer (1890:
110).] Syntypes: MNHN (5 lots, 36 spms.)
(see also Sowerby, 1909b: 363). Distribution:
Belize.

Remarks. Synonym of flagellata Say, 1829,
teste Pain (1964: 227).
brasiliensis
Ampullaria Brasiliensis Paetel, 1887: 477.
Unavailable name; nom. nud.
bridgesii
Ampullaria Bridgesii Reeve, 1856b: pl. 11,
figs. 50, 51. Rio Grande, Bolivia. Lectotype
(Pain, 1960: 425): BMNH 20010487 (shell fig-
ured as no. 50 by Reeve); paralectotype:
BMNH 20010488. Distribution: Brasil (Baker,
1914: 660), Bolivia.

Remarks. Pain (1960: 425) considered the
nominotypical subspecies of bridgesii Reeve,
1856, to be a rare and local form, with the
subspecies diffusa Blume, 1957, being much
more widespread. Sometimes synonymized
with scalaris Orbigny, 1835 (e.g., lhering,
1898: 48), but almost certainly incorrectly.
bulla
Ampullaria bulla Reeve, 1856d: pl. 22, fig.
104a, b. [No locality given but type material
labeled as from Mexico.] Syntypes: BMNH
20020648 (2 spms.). Distribution: Ecuador [?
error] (Paetel, 1887: 477), Mexico (Sowerby,
1909a: 346).
buxea
Ampullaria buxea Reeve, 1856e: pl. 23, fig.
112. [No locality given.] Syntype: BMNH
1907.10.28.210. Distribution: Colombia
(Paetel, 1887: 477), Jamaica (Sowerby,
1909a: 346).

Remarks. Synonym of fasciata Roissy,
1805, teste Pilsbry (1927a: 247), although
considered а possible synonym of
hopetonensis Lea, 1834, by Martens (1857:
203).

COWIE & THIENGO

caliginosa

Ampullaria caliginosa Reeve, 1856e: pl. 25,
fig. 118. [No locality given.] Type material: not
found by us in ВММН. Distribution: Florida
(Walker, 1918: 124).

Remarks. Synonym of paludosa Say, 1829,
teste Sowerby (1916: 70), followed here,
though Pilsbry (1927a: 250) was not certain
of this synonymy.

camena

Pomacea camena Pain, 1949a: 258; pl. 13,
figs. 5, 6. shallow stream near Lagunella,
Venezuela, at 800 metres. Holotype: BMNH
1946.10.2.4 (Pain, 1949a: 258; incorrectly cit-
ing BMNH 1946.10.2.3, which is the number

-ofthe holotype of vickeryi Pain, 1949, in both

his paper and the ВММН register); paratypes
(1 only mentioned by Pain (1949a: 258)):
HUJ 21516 (1 spm.) (H. Mienis, pers.
comm. to RHC, 4 August 2002),
NMW.Z.1981.118.00108 (Pain collection, 2
spms.), NMW.1955.158.02412 (Melvill-Tomlin
collection, 1 spm.), MCZ. Distribution: Ven-
ezuela.

Remarks. The holotype in the BMNH is not
a close match to the specimen illustrated by
Pain (1949a: figs. 5, 6), although the other
two holotypes illustrated in Pain’s paper are of
the BMNH specimens (P. B. Mordan, pers.
comm. to RHC, 2 November 2001, 7 Febru-
ary 2003).

canaliculata

Ampullaria canaliculata Lamarck, 1822a: 178.
les rivieres de la Guadeloupe [? error; per-
haps Lago Guadeloupe, Argentina, not the
Caribbean island of Guadeloupe (Pain,
1946b: 58; Hylton Scott, 1958: 300; Thiengo
et al., 1993: 68; Cazzaniga, 2002: 74)]. Pos-
sible holotype: MHNG 1093/91 (Sowerby,
1909b: 363; Mermod, 1952: 88; Y. Finet, pers.
comm. to RHC, 24 October 1994, 22 August
2002). Distribution: Argentina, Bolivia, Brazil,
Paraguay, Uruguay (Hylton Scott, 1958: 301—
303) [but ascertaining the true distribution of
this variable species depends on detailed
taxonomic study; e.g., Cazzaniga (1987)].
Remarks. Lamarck (1804: 32) also de-
scribed a marine fossil from the Eocene of
France as Ampullaria canaliculata, this spe-
cies being the type species of Amauropsina
Chelot, 1885, which is either in the Naticidae
(Kabat, 1991: 426) or Ampullospiridae
(Tracey et al., 1996: 116). An application
(Cowie et al., 2001) was submitted to the
ICZN to retain both names as valid (Code Art.

NEW WORLD AMPULLARIIDAE 59

23.9.5), and this was so ruled by the ICZN
(2002: 137).

cassidiformis
Ampullaria Cassidiformi Reeve, 1856b: pl.
12, fig. 56. Lake of Maracaibo, Venezuela.
Syntype: BMNH 20020649. Distribution: Ven-
ezuela.

Remarks. The original spelling, as given
above, is considered incorrect, as it was
clearly an inadvertent error [Code Art. 32.5.1]
inasmuch as other species published by
Reeve at the same time with similarly
formed names did not lack the “5”. The index
to Reeve's work has “cassidiformis”, but this
was published later and therefore is to be
considered as evidence external to the origi-
nal publication. Synonym of eximia Dunker,
1853, teste Baker (1930: 6).

castelloi

Ampullaria Castelloi Sowerby, 1894: 48, pl. 4,
fig. 22. River Meta, S.E. of Bogota. Lectotype
(Pain, 1949b: pl. 1, figs. 1, 2): BMNH
1893.5.29.3 [possibly part of the type series
but not the specimen originally figured by
Sowerby; P. B. Mordan, pers. comm. to
RHC, 7 February 2003]; possible paralecto-
types: MCZ (1 lot) [labeled “? paratypes”]. Dis-
tribution: Colombia, Surinam (Vernhout,
1914a: 29, 41, 43) [? error; Geijskes & Pain,
1957: 45).

Remarks. Placed in Limnopomus Dall by
Pain (1949b: 39). Sowerby (1894: 48) based
his description on more than one shell (*... in
some specimens [the umbilicus is] com-
pletely closed”). Therefore, the specimen fig-
ured by Pain (1949b: pl. 1, figs. 1, 2) as the
“type” must be considered a lectotype (Code
Art. 74.5, Rec. 73F).

castelnaudii
Ampullaria castelnaudii Hupe, 1857: 65, pl.
11, fig. 1. le fleuve des Amazones. Syntypes:
MNHN (4 lots, 7 spms.). Distribution: “Haut-
Amazone” (Gaudion, 1879: 26).
catamarcensis
Ampullaria catamarcensis Sowerby, 1875:
600, pl. 72, fig. 4. Catamarca (on the Andes
of Peru) [? = Cajamarca (Peru); Cazzaniga
(1987: 59-61)]. Syntypes: BMNH 1875.4.19.2
(2 spms.). Distribution: Peru.

Remarks. Placed in subgenus Limnopomus

Dall by Cazzaniga (1987: 59-61).

+ catemacensis
Ampullaria patula catemacensis Baker,
1922: 39, pl. 14, figs. 2-4, pl. 15. fig. 7. Lake
Catemaco. Holotype (Baker, 1922: pl. 14, fig.

2): UMMZ 31850; paratypes: UMMZ 31850 (5
spms., not separated from the holotype),
ANSP 133680 (4 spms.); topotypes: MCZ. Dis-
tribution: Lake Catemaco, Mexico (Naranjo-
García 8 Garcia-Cubas, 1986: 603).

cerasum

Ampullaria cerasum Hanley, 1854: [unnum-
bered page], Ampullaria pl. 2, fig. 7. [No lo-
cality given.] Syntype: BMNH 1907.11.21.83.
Distribution: Mexico (Martens, 1899: 421;
Sowerby, 1909a: 347).

Remarks. Not listed among Hanley's taxa
by Norris & Dance (2002: 370).

+ chamana

Ampullaria lattrei chamana Hinkley, 1920: 53
[1921: pl. 4, fig. 5]. Guatemala [in publication
title]. Lectotype (Baker, 1964: 168): ANSP
“46231” [error; correctly 46321] ; para-
lectotypes: Bryant Walker collection, Mu-
seum of the lllinois University, Hinkley
collection (Hinkley, 1920: 54), ANSP 76238 (2
spms.; originally 3 spms. in this lot), MCZ.
Distribution: Guatemala.

chaquensis

Pomacea canaliculata chaquensis Hylton
Scott, 1948: 242. Madrejón de Ingeniero
Juárez, Chaco salteño. Formosa. Syntypes
[“Cotipos”]: IMLA. Distribution: Argentina, Bo-
livia (Hylton Scott, 1958: 304).

Remarks. Synonym of canaliculata
Lamarck, 1822, teste Cazzaniga (1987: 56).

chemnitzii

Ampullaria Chemnitzii Philippi, 1852a: 39, pl.
10, fig. 5[1852b: 25]. [No locality given. 4.5
kilometers south ... from Tucacas “chosen”
by Baker (1930: 5)] Lectotype: the specimen
illustrated in “Philippi's figure” (Baker, 1930:
5) Type material: probably MNHNS. Distribu-
tion: Ecuador, Colombia, Venezuela (Pain,
1956a: 75).

Remarks. Although the origin of the desig-
nated lectotype is unknown, Baker's (1930: 5)
type locality choice (the locality being that of
his own material) appears to follow Code Rec.
76А.1.4. Synonym of lineata Spix, 1827, teste
Sowerby (1909a: 354) and Vernhout (1914a:
27), but retained here as a distinct species,
following Baker (1930: 5).

chiapasensis

Ampullaria malleata var. Chiapasensis
Fischer & Crosse, 1890: 235, pl. 48, fig. 5. in
paludibus prope Las Playas, in provincia
Chiapas. Type material: Morelet collection, not
found by us in BMNH or ММНМ, not found in
MHNG by Y. Finet (pers. comm. to RHC, 5

60

August 2002) (cf. Dance, 1986: 219). Distri-
bution: Mexico.

Remarks. Synonym of livescens Reeve,
1856, teste Pain (1964: 228).
cincta
Ampullaria cincta Cristofori & Jan, 1832:
[Section Ila, Pars la] 7, [Mantissa] 3. Jamaica.
Type material: formerly MCSN [destroyed; A.
Garassino, pers. comm. to RHC, 5 Septem-
ber 2002]. Distribution: Jamaica.

Remarks. Synonym of fasciata Roissy,
1805, teste Pilsbry (1927a: 247).
citreum
Ampullaria сйгеит Reeve, 1856e: pl. 24, fig.
116a, b. [No locality given.] Syntype: BMNH
1907.11.21.83. Distribution: “Western Hemi-
sphere” (Sowerby, 1909a: 347).
columbensis
Ampullaria Columbensis Jay, 1836: 47 [1839:
65; 1850: 282]. Unavailable name; nom. nud.

Remarks. South America given as locality
by Jay (1836: 47).
columbiensis
Ampullaria columbiensis Philippi, 1851: 20,
pl. 5, fig. 5. [No locality given; “West-
kolumbien” on label in ZMHB] Syntypes:
ZMHB 1343 (2 spms.) (M. Glaubrecht, pers.
comm. to RHC, 1 March 2003), possibly also
MNHNS. Distribution: possibly “Rio Pastasa
in Andibus orientalibus” (Miller, 1879: 150),
Colombia.

Remarks. Name attributed to Sowerby by
Philippi (1851: 20). Miller (1879: 150) dis-
cussed his material under “Ampullaria aff.
Columbiensis Phil.”. Alderson (1925: 53) con-
sidered it “unrecognized” but discussed it
under interrupta Sowerby, 1909, which was
placed in Limnopomus Dall, 1904, by
Sowerby (1909a: 361). We therefore include
it tentatively in Pomacea Perry, 1810.
columbiensis
Ampullaria columbiensis Reeve, 1856b: pl. 5,
fig. 25. Chiriqui, Veragua [Colombia].
Syntypes: BMNH 20020650 (2 spms.), MNHN
(3 spms.).

Remarks. Name attributed to Sowerby, MS.
Junior primary homonym of co/umbiensis
Philippi, 1851; replaced by martensiana
Nevill, 1884. Also replaced by tristrami
Fischer & Crosse, 1890, Fischer & Crosse
(1890: 245) apparently being unaware of
Nevill, 1884. Synonym of flagellata Say, 1829,
teste Pain (1964: 227).
columellaris
Ampullaria columellaris Gould, 1848: 74.
Province of Maynas, Peru. Lectotype

COWIE & THIENGO

(Johnson, 1964: 57): USNM 5547. Distribu-
tion: Peru, Bolivia, ? Ecuador (Pain, 1960:
429).

Remarks. Unless it can be determined that
Gould based his description on only a single
specimen, Johnson's (1964: 57) listing of the
“holotype” in fact designated a lectotype
(Code Art. 74.6, Rec. 73F). The type species
of Limnopomus Dall, 1904, which is here
considered a synonym of Pomacea Perry,
1810.

commissionis

Amp. decussata var. commissionis lhering,
1898: 51. Iguape. Syntypes: ZMHB 109513
(1 spm.), 109514 (2 spms.) (M. Glaubrecht,
pers. comm. to RHC, 1 March 2003); no type
material in MZUSP (cf. Dance, 1986: 214).
Distribution: Brasil (Pilsbry, 1933: 74).
Remarks. Raised to full species level and
placed in Asolene Orbigny, 1838, by Kobelt
(1913h: 202), but removed from Asolene
Orbigny, 1838, by Ihering (1919: 341).

conoidea

Ampullaria conoidea Martens, 1899: 423, pl.
24, figs. 10, 11. Costa Rica. Possible
syntypes (description based explicitly on 2
spms. only): ZMHB 21857 (2 spms.) (M.
Glaubrecht, pers. comm. to RHC, 4 May
2002, 1 March 2003). Distribution: Costa
Rica.

consolatrix

Ampullaria consolatrix |hering, 1919: 338. FI.
Uruguay, prope Itaquy, Río Grande do Sul.
Type material: not found by us in MZUSP (cf.
Dance, 1986: 214). Distribution: Brasil.

contamanoensis

Ampullaria contamanoënsis Preston, 1914:
527. Contamano, Rio Ucayali, Eastern Peru.
Syntypes: BMNH 1915.1.6.84,
NMW.Z.1981.118.00096 (Pain collection, 1
spm.), not in HUJ, not found by us in UMMZ
(cf. Dance, 1986: 206, 222). Distribution:
Peru.

Remarks. Synonym of aulanieri Deville &
Huppé, 1850, teste Pain (1960: 424).

cornucopia

Ampullaria cornucopia Reeve, 1856e: pl. 1,
fig. 4. Columbia [= Colombia]. Syntype:
BMNH 20020651. Distribution: Colombia
(Sowerby, 1909a: 347).

costaricana

Ampullaria costaricana Martens, 1899: 418,
pl. 24, figs. 14-17. Nicaragua: Lake of Nicara-
gua ... N.W. Costa Rica: Rio Saveyre, at
Boca Culebra ... S.W. Costa Rica: Palmar,
south of the Rio Grande de Terraba ... М.

NEW WORLD AMPULLARIIDAE 61

Panama: Chiriqui. Syntypes: ZMHB 109507
(1 spm.) [= fig. 16], 109508 (1 spm.) [= fig.
15], 109509 (1 spm.) [= fig. 17], 109510 (1
spm.) [= fig. 14], 109511 (7 spms.), 109512 (1
spm.) (M. Glaubrecht, pers. comm. to RHC,
1 March 2003; F. Kôhler, pers. comm. to
RHC, 6 March 2003); no type material found
by us in BMNH or MCZ (cf. Dance, 1986:
218). Distribution: Nicaragua, Costa Rica,
Panama.

Remarks. Additional localities in Costa Rica
given by Martens (1901: 644).
cousini
Ampullaria Cousini Jousseaume, 1877: 185,
pl. 3, fig. 3. la République de l'Equateur.
Syntypes: ММНМ (2 lots, 3 spms.). Distribu-
tion: Ecuador (Sowerby, 1909a: 347).

crosseana

Ampullaria Crosseana Hidalgo, 1871: 206
[1872: 142, pl. 7, fig. 1]. in fluvio Amazonum
dicto, Americæ meridionalis. Syntypes:
MNCN 15.05/11485 (1 spm., figured), 15.05/
1047 (1 spm.) (Villena et al., 1997: 75). Dis-
tribution: River Amazon (Sowerby, 1909a:
348).

Remarks. Synonym of maculata Perry,
1810, teste Pain (1960: 423).
cubensis
Ampullaria cubensis Morelet, 1849: 24. prov.
borealis insulæ Cuba. Syntypes: ВММН
1893.2.4.1675-6 (2 spms.). Distribution:
Cuba [? error; Pilsbry, 1927a: 252].

Remarks. Synonym of teres Philippi, 1849,
teste Paetel (1887: 478), and according to
the syntype labels. However, Pilsbry (1927a:
252) conclusively demonstrated that it is not
teres Philippi, 1849 (nor cubensis Reeve,
1856), but he was unable to locate the type
material (see also cubensis Reeve, 1856) and
hence considered the species too poorly
known to place it in the synonymy of any
known species.
cubensis
Ampullaria Cubensis Reeve, 1856c: pl. 18,
fig. 83a, b. Cuba. Type material: not found by
us in BMNH. Distribution: Cuba.

Remarks. Junior primary homonym of
cubensis Morelet, 1849, replaced by poeyana
Pilsbry, 1927. Treated as a variety of glauca
Linnaeus, 1758, by Sowerby (1909a: 350)
and placed in the “Formenkreis” of glauca
Linnaeus, 1758, by Kobelt (1913a: 150). Syn-
onym of glauca Linnaeus, 1758, teste
Starmühlner (1988: 253 [as “culemsis’]).
Pilsbry (1927a: 251-252), followed here, re-
tained poeyana Pilsbry, 1927, as a distinct

species not in the “same section of the ge-
nus [as glauca Linnaeus, 1758)” (i.e., subg.
Effusa). It is possible that the syntypes listed
under cubensis Morelet, 1849, although la-
beled “Morelet”, are syntypes of cubensis
Reeve, as we found no type material labeled
“cubensis Reeve” in the BMNH. If Pilsbry had
thought this to be the case, it would explain
his inability to find Morelet's material. How-
ever, neither specimen matches Reeve’s fig-
ures.

cumingii

Ampullaria Cumingii King & Broderip, 1831:
344. in Sinu Paname, (Island of Saboga, in a
small hill-stream). Type material: not found by
us in BMNH (cf. Dance, 1986: 215); topotype:
USNM 4673 (Morrison, 1946: 6). Distribution:
Mexico [? error], Panama, Ecuador [? error]
(Sowerby, 1909a: 348).

Remarks. Confusion over the type locality
was clarified by Morrison (1952: 105-106),
who considered the locality as originally pub-
lished (“Saboga”) to be correct.

dacostae

Ampullaria Da Costee Sowerby, 1909a: 348
[name], 359 [description], text fig. Costa
Rica. Syntype: BMNH 1909.10.19.35 (see
also Sowerby, 1909a: 359). Distribution:
Costa Rica.

decussata

Ampullaria decussata Moricand, 1836: 445,
pl. 2, figs. 26, 27. Bahia [in publication title;
Lake Baril, Brasil, according to the label as-
sociated with the MHNG syntypes]. Syntypes:
HUJ 21518 (2 spms.) (H. Mienis, pers.
comm. to RHC, 4 August 2002), MCZ (2 lots;
one of them is 141866), MHNG 33484 (9
spms.) (Y. Finet, pers. comm. to RHC, 26
August 2002), MNHN (2 lots, 13 spms.),
ZMHB 109516 (2 spms.) (M. Glaubrecht,
pers. comm. to RHC, 1 March 2003). Distri-
bution: Brasil (Sowerby, 1909a: 348).

delattrei

Ampullaria Delattrei Fischer & Crosse, 1890:
246, pl. 48, fig. 7, Та [1888: pl. 45, fig. 4, 4а].
Unjustified emendation of /attrei Reeve, 1856.

Remarks. Although the name was given as
“Delattrei’ by Fischer & Crosse (1888: [expla-
nation of] pl. 45, fig. 4), this was not explicitly
an emendation. The emendation was explicit
in Fischer & Crosse (1890: 246), and there-
fore dates from 1890. Reeve (1856b: pl. 5,
fig. 22) gave the collector's name incorrectly
as “Lattre” and spelled the species name as
“Lattrei’. However, there is no evidence in the
original publication of an inadvertent error

62

(Code Ан. 32.5.1) that would justify the етеп-
dation. Although accepted by some (e.g.,
Martens, 1899: 419; Alderson, 1925: 31), the
emendation has not been accepted by others
(e.g., Hinkley, 1920: 53; Pain, 1964: 229) and
so cannot be considered to be in prevailing
use; it is therefore an unjustified emendation
(Code Art. 33.2.3.1) and a junior objective syn-
onym of /attrei Reeve, 1856.

depressa

+

Ampullaria depressa Say, 1824: 264, pl. 14,
fig. 2. East Florida ... tributary to St. John’s
river, and on the plantation of Mr. Fatio ... Lake
George. Lectotype (Clench & Turner, 1956:
121; see also Baker, 1964: 168): ANSP
50580; paralectotype: ANSP 365373.

Remarks. Junior primary homonym of
depressa Lamarck, 1804 [not Ampullariidae].
Replaced by paludosa Say, 1829.
diffusa
Pomacea bridgesi diffusa Blume, 1957: 1,
[unnumbered text figs.; holotype]. Lagune
mitten in der Stadt St. Cruz, Bolivia. Holotype:
ZSM 20011991; paratypes: ZSM 20011990 (4
spms.) [? ex coll. Blume]; possible
paratypes: ZSM 20011989 (c. 110 spms.) [?
the “alle anderen Paratypoide” (Blume, 1957:
2)]; Pain collection (1 spm.) (Blume, 1957: 2;
E. Schwabe, pers. comm. to RHC, 28 July
2002). Distribution: Brasil, Peru, Bolivia (Pain,
1960: 425).

Remarks. Possibly a valid species (F.
Naggs, pers. comm. to RHC, 9 July 2002).
The true identity and origin of the snails cur-
rently referred to widely as bridgesii Reeve,
1856, in the domestic aquarium trade (Perera
8 Walls, 1996) is not known; they may be
more correctly referred to diffusa Blume,
1957.

dilatata

Ampullaria fasciata variété dilatata Orbigny,
1842c: 4. Cuba [in publication title]. Type ma-
terial: not found by us in BMNH (nor listed by
Gray, 1855: 27—29) or MNHN; not in MHNG (Y.
Finet, pers. comm. to RHC, 20 August 2002)
(cf. Dance, 1986: 219, 220). Distribution:
Cuba.

Remarks. Synonym of paludosa Say, 1829,
teste Pilsbry (1927a: 250).

disseminata

Ampullaria disseminata De Kay, 1843: 124.
Unavailable name; first published as a junior
synonym of paludosa Say, 1829, not made
available before 1961 (Code Art. 11.6).
Remarks. DeKay (1843: 124) attributed the
name to Say as a manuscript name, but the

COWIE & THIENGO

name does not occur in Say’s published writ-
ings (Binney, 1858: [237], 1865: 5).

dolioides

Ampullaria Dolioides Reeve, 1856c: pl. 16,
fig. 75a, b. Bombay [error]. Syntypes: BMNH
20020653 (2 spms.). Distribution: Guyana,
Surinam, French Guiana, Venezuela (Pain,
1950b: 65; Geijskes & Pain, 1957: 43; Tillier,
1980: 29).

Remarks. For a history of the interpretation
of this species see Prashad (1925: 83; 1931:
167). Considered a synonym of lineata Spix,
1827, by Pain (1952: 31) but in error accord-
ing to Geijskes & Pain (1957: 43), followed
here, who treated it as a valid species, as did
Tillier (1980: 29).

dolium

Ampullaria dolium Philippi, 1852a: 40, pl. 11,
fig. 1 [1852b: 25]. Guyana, namentlich der
Orinoco. Type material: probably MNHNS.
Distribution: Guyana.

Remarks. Synonym of urceus Múller, 1774,
teste Gaudion (1879: 41), Sowerby (1909a:
358), Alderson (1925: 10) and Baker (1930: 2).

dorbignyana

Ampullaria Dorbignyana Philippi, 1852a: 65,
pl. 21, fig. 4. [The locality of Orbigny's speci-
men; “die La Plata Staaten” (Philippi, 1852a:
66)]. Holotype: the specimen illustrated in
Orbigny’s (1835a) figure (pl. 4, fig. 4), loca-
tion not known to us. Distribution: “Brésil —
Parana — Plata” (Gaudion, 1879: 35).
Remarks. Philippi (1852a: 65) explicitly de-
scribed this species on the basis of
Orbigny's (1835a) figure. Philippi (1852a: 49)
also named it “d’Orbignyf’ [= dorbignyi. We
select dorbignyana, the heading of Philippi's
description and the more widely used alter-
native (e.g., Alderson, 1925: 21), as the cor-
rect original spelling. Synonym of canaliculata
Lamarck, 1822, teste lhering (1898: 49) and
Sowerby (1909a: 348; 1909b: 363).

dorbignyi

Ampullaria d'Orbignyi Philippi, 1852a: 49. In-
correct original spelling of dorbignyana
Philippi, 1852.

+ dysoni

Ampullaria Dysoni Hanley, 1854: [unnum-
bered page], Ampullaria pl. 2, fig. 5. Hondu-
ras. Syntype: BMNH 1907.11.21.65.
Distribution: Honduras (Martens, 1899: 417;
Pain, 1964: 230).

Remarks. Subspecies of flagellata Say,
1829, teste Pain (1964: 230). Not listed
among Hanley's taxa by Norris & Dance
(2002: 371).

NEW WORLD AMPULLARIIDAE 63

electrina
Ampullaria electrina Reeve, 1856c: pl. 20, fig.
95a, b. [No locality given.] Syntypes: BMNH
20020654 (3 spms.). Distribution: unknown.

Remarks. Sowerby (1909a: 349) placed it in

Pomacea [аз Ampullaria].

elegans
Ampullaria elegans Orbigny, 1835a: 33. Rio
Piray, provincia Santa Cruz de la Sierra (re-
publica Boliviana). Syntypes: BMNH
1854.12.4.330 (3 spms.), MNHN (4 lots, 14
spms.). Distribution: Bolivia.

Remarks. Synonym of cyclostoma Spix,
1827, teste Pain (1960: 430).

elongata
Ampullaria fasciata variété elongata Orbigny,
1842c: 4. Cuba [in publication title]. Type ma-
terial: not found by us in BMNH (nor listed by
Gray, 1854: 17) or MNHN, not in MHNG (Y.
Finet, pers. comm. to RHC, 20 August 2002)
(cf. Dance, 1986: 220). Distribution: Cuba.

Remarks. Synonym of paludosa Say, 1829,

teste Pilsbry (1927a: 250).

+ erogata

Ampullaria erogata Crosse 8 Fischer, in
Fischer 8 Crosse, 1888: [explanation of] pl.
46, figs. 6, ба, 7 [Crosse & Fischer, 1890:
113; see also Fischer 8 Crosse (1890: 251)].
[No locality given. Peten, Guatemalae ...
Cacoprieto, in isthmo de Tehuantepecensi,
reipublicae Mexicanae (Crosse & Fischer,
1890: 113)]. Holotype: the specimen illus-
trated by Crosse 8 Fischer, in Fischer 8
Crosse (1888, fig. 6, 6a), not found by us in
MNHN. Distribution: as for flagellata Say,
1829 (Pain, 1964: 230).

Remarks. Treated by Pain (1964: 229) as
an “ecological race” of flagellata Say, 1829,
occupying the same geographic range. De-
termining its true taxonomic status requires
further study.

erronea
Ampullaria erronea Nevill, 1877: 17. S.
America. Holotype: NZSI. Distribution: South
America.

erythrostoma
Ampullaria erythrostoma Reeve, 1856c: pl.
13, fig. 59. Zanzibar [error (Sowerby, 1909a:
349; Pain, 1950b: 68)]. Type material: not
found by us in BMNH. Distribution: Peru
(Sowerby, 1909a: 349).

Remarks. Synonym of guyanensis
Lamarck, 1822, teste Sowerby (1909a: 349),
Kobelt (1913f: 186) and Pain (1960: 427), and
hence of urceus Müller, 1774 (see Tillier,
1980: 27). N. syn.

eumicra

Ampullaria eumicra Crosse & Fischer, 1890:
113 [see also Fischer & Crosse (1890: 243,
pl. 48, fig. 10, 10a)]. provincia Oajaca dicta,
reipublicae Mexicanae. Syntypes: MNHN (4
spms.). Distribution: Mexico.

Remarks. Synonym of flagellata Say, 1829,
teste Pain (1964: 227).

exculpta

Ampullaria malleata var. Exculpta Fischer &
Crosse, 1890: 235 [1888: pl. 44, fig. 6, 6a-c;
plate published without name]. in Laguna de
los Cocos, provinciæ Vera Cruz ..., in
paludibus prope Palizada et San Geromino,
provinciæ Yucatan ..., in paludibus fluminis
Usumasinta, prope Balancan, provinciæ
Tabasco. Type material: Sallé collection, not
found by us in BMNH, ММНМ, etc. (cf. Dance,
1986: 209, 225). Distribution: Mexico.
Remarks. Synonym of flagellata Say, 1829,
teste Baker (1922: 37) and Pain (1964: 227).

eximia

Ampullaria eximia Dunker, 1853: 93. die
Provinz Coro am See von Maracaybo,
Republik Venezuela. Syntypes: MCZ 125225,
ZMHB 4039 (3 spms.) (M. Glaubrecht, pers.
comm. to RHC, 4 May 2002, 1 March 2003).
Distribution: Venezuela (Baker, 1930: 6).

falconensis

Pomacea falconensis Pain & Arias, 1958: 6,
pl. 1, figs. 1-4, pl. 2, figs. 1-7. 5 km. SW de
Chichiriviche, Estado Falcon, Venezuela
(68 152’ W; 10 502’ N). Holotype: MHNS
4000 (female); paratypes: MHNS 3499 (7
spms.), NMW.Z.1981.118.00116 (Pain col-
lection, 4 spms.), MCZ 224267 (2 spms.);
ZSM 20012070 (1 spm.) (E. Schwabe, pers.
comm. to RHC, 28 July 2002). Distribution:
Venezuela.

fasciata

Ampullaria fasciata Roissy, 1805: 374. les
rivières de la Jamaïque, de la Guadeloupe et
de Saint-Domingue [Haiti]. Type material: lo-
cation not known to us. Distribution: Jamaica
only, Guadaloupe and Haiti being incorrect
(Pilsbry, 1927a: 248).

Remarks. Pilsbry (1927a: 247) clarified the
status of this species as being from Ja-
maica, as Roissy had stated, and that it is
not an Asian species in the synonymy of
ampullacea Linnaeus, 1758 (which is now
placed in Pila) as Sowerby (1909a: 354) had
considered; that is, Roissy (1805: 374; see
also Schumacher, 1817: 200) mistakenly in-
cluded ampullacea Linnaeus, 1758, in his
synonymy.

64 COWIE 8 THIENGO

fasciata
Ampullaria fasciata Reeve, 1856b: pl. 9, fig.
41. [No locality given.] Lectotype (Kobelt,
1914b: 220): BMNH 20020655.

Remarks. Junior primary homonym of
fasciata Roissy, 1805, and fasciata Lamarck,
1816 [incertae sedis in family Ampullariidae].
Synonym of insularum Orbigny, 1835, teste
Sowerby (1909a: 353). Kobelt (1914b: 220)
designated the specimen in Reeve’s pl. 9, fig.
41 as a lectotype, copying the figure as his
own pl. 77, fig. 1.

ferruginea
Ampullaria ferruginea Martens, 1857: 205.
Unavailable name; nom. nud.

Remarks. Attributed to “R. pl. 14” [= Reeve,
1856b, pl. 14] with locality “Laplata”. However,
the name “ferruginea” does not appear in
Reeve's work. Gaudion (1879: 29) also listed
the name and said “Reev [sic] Hab: La Plata”
but with no other information. Not listed by
Sowerby (1916: 71), Ruhoff (1980: 271) or
Kabat 8 Boss (1997: 208).

figulina
Ampullaria figulina Spix, in Wagner 1827: 3,
pl. 4, fig. 4. [Type locality as for lineata Spix,
1827]. Syntypes: ZSM 20012063-5 (3 lots, 4
spms.) (E. Schwabe, pers. comm. to RHC,
28 July 2002; cf. Fechter, 1983: 221). Distri-
bution: Brasil (Baker, 1914: 659).

Remarks. Authorship is given here as “Spix,
in Wagner”, following Cowie et al. (in prep.),
who also explain the publication history of this
work. Spix illustrated figulina as a full species,
but Wagner, in writing the description, treated
figulina “Spix” as a variety of lineata
“Wagner’. Wagner at first sight appears to
have also treated it as a synonym of his own
new species-group name “minor”. Cowie et
al. (in prep.) discuss why this is not the case
and why “minor” is not an available name.
Alderson (1925: 29) considered figulina Spix,
1827, impossible to identify with certainty.
Synonym of lineata Эрих, 1827, teste Pain
(1960: 422), followed here, although he cited
pl. 6, fig. 4.

flagellata
Ampullaria flagellata Say, 1829c: 260. Mexico
... a Short distance below Vera Cruz. Lecto-
type (Pilsbry, 1891a: 325-326): ANSP 50645
[50645а” (Baker, 1964: 168)]; paralectotype:
ANSP 50645; topotypes: MCZ 139677. Distri-
bution: Central America, from central Mexico
to Panama, extending into northern Colombia
(Magdelana drainage area) (Pain, 1964: 228;

Naranjo-Garcia & Garcia-Cubas, 1986: 603).
Remarks. Although Baker (1964: 168) con-
sidered the type as fixed by monotypy [“TOM”
(Baker, 1964: 149)], he also noted that this
specimen was the “smaller and fresher of
type lot’, implying that there were additional
specimens. Thus, Pilsbry (1891a: 325-326)
in referring to a single specimen as “Say’s
type”, and providing dimensions for it, desig-
nated a lectotype (Code Art. 74.6), for which
Baker (1964: 168) gave the catalog number.
flatilis
Ampullaria flatilis Reeve, 1856b: pl. 7, fig. 31.
Tabasco, Mexico. Syntype: BMNH 20020656.
Distribution: Mexico.

Remarks. Synonym of flagellata Say, 1829,

teste Pain (1964: 227).

flava

Pomacea paludosa flava Smith, 1937: 147.
canals near Pinecrest on the Tamiamai Trail.
Central Everglades and near Miami, Florida.
Lectotype (Baker, 1964: 168): ANSP 188992
[as “188992a’]; paralectotype: ANSP 365372
(2 spms.). Distribution: Florida.

Remarks. Synonym of paludosa Say, 1829,

teste Clench 8 Turner (1956: 120).

fumata
Ampullaria fumata Reeve, 1856e: pl. 26, fig.
124a, b. Province of Chiapes [= Chiapas],
Mexico. Type material: not found by us in
BMNH. Distribution: Mexico.

Remarks. Synonym of flagellata Say, 1829,
teste Pain (1964: 227). Discussed briefly by
Strebel (1873: 32).

+ garciae
Pomacea paludosa garciae Richards, 1933:
169, fig. 21. swamp near the town of
Mendoza (or Paso Real) about five kilome-
ters from the terminus of the Ferro-Cariles
Unidos de la Habana at Guane, Pinar del Rio,
Cuba. Holotype: ANSP 160873 ["160873a”
(Baker, 1964: 168)]; paratypes: ANSP 365371
(2 spms.); topotype: MCZ. Distribution: Cuba.

georgil
Ampullaria Georgii Williams, 1889: 47.
marshes near the La Plata, at Buenos Ayres,
in the Argentine Republic. Type material: loca-
tion not known to us. Distribution: Argentina.

Remarks. Synonym of insularum Orbigny,
1835, teste Sowerby (1909a: 353).

ghiesbreghti
Ampullaria Ghiesbrechtii Reeve, 1856e: pl.
26, fig. 123. Province of Chiapes [= Chiapas],
Mexico. Syntype: BMNH 20020657. Distribu-
tion: Mexico, Guatemala (Pain, 1953: 222).

NEW WORLD AMPULLARIIDAE 65

Remarks. Originally spelled “Ghiesbrechti”
but explicitly emended to “Ghiesbreghti by
Fischer & Crosse (1890: 233). Although there
is no evidence in the original publication of an
inadvertent error (Code Art. 32.5.1) that would
justify the emendation, the emendation is in
prevailing use attributed to Reeve (e.g.,
Pilsbry, 1893: 338; Alderson, 1925: 44; Pain,
1953: 222, 1964: 228) and is therefore
deemed to be a justified emendation (Code
Art. 33.2.3.1). Synonym of livescens Reeve,
1856, teste Pain (1964: 228).
gigantea
Ampullaria ? gigantea Barbosa Rodrigues,
1892: 52. avec ГЕту$ macrococcygeana ...
à la même époque geologique ... ; dans les
ravins des environs du Rio Nanay; Loreto-
Yacu, dans l'étage tertiaire [probably near
Loreto on the upper Amazon in Peru, above
the junction with the Rio Javari (Boss &
Parodoz, 1977: 111)]. Type material: may
have been lost (Patterson, 1936: 50; Boss &
Parodiz, 1977: 111). Distribution: Peru.

Remarks. Fossil, probably Pliocene (Boss
& Parodiz, 1977: 111). Junior secondary
homonym of giganteus Tristram, 1864.
giganteus
Pomus giganteus Tristram, 1864: 414. Lake
Peten, Vera Paz [Guatemala]. Type material:
not found by us in BMNH (cf. Dance, 1986:
229). “Paratype” (Pain, 1953: 222) [? =
syntype]: NMW.Z.1981.118.00125 (Pain col-
lection, 1 spm.). Distribution: Guatemala.

Remarks. Synonym of livescens Reeve,
1856, teste Pain (1964: 228).
gigas
Ampullaria gigas Spix, in Wagner, 1827: 1, pl.
1, figs. 1, 2. In flumine Amazonum. Type ma-
terial: formerly in ZSM but probably lost
(Alderson, 1925: 16; Fechter, 1983: 221; S.
C. Thiengo, unpublished); possible syntype:
MHNG 33489 (1 spm.) (Y. Finet, pers. comm.
to RHC, 26 August 2002). Distribution: Brasil
(Sowerby, 1909a: 350; Baker, 1914: 659).

Remarks. Authorship is given here as “Spix,
in Wagner”, following Cowie et al. (in prep.).
Synonym of maculata Perry, 1810, teste
Pilsbry (1927b: 63), Pain (1956a: 79, 1960:
423), Geijskes 8 Pain (1957: 42) and Boss &
Parodiz (1977: 112), contra lhering (1919:
334), who synonymized it with insularum
Orbigny, 1835.
gossei
Ampullaria Gossei Reeve, 1856c: pl. 20, fig.
ЭЗа, b. Jamaica. Syntypes: ВММН 20020658

(3 spms.). Distribution: Jamaica (Sowerby,
1909a: 351; Pilsbry, 1927a: 249).

guaduasensis

Ampullaria guaduasensis Anderson, 1928:
23, pl. 1, figs. 19, 20. near San Juan de Rio
Seco, on the east border of the upper valley
of the Magdalena River, Colombia. Holotype:
CAS 2721. Distribution: Colombia.

Remarks. Pleistocene fossil (Boss 4
Parodiz, 1977: 118).

gualtieri

Amp. Gualtieri Orbigny, 1835a: 32. Unavail-
able name; first published as a junior syn-
onym of canaliculata Lamarck, 1822, not
made available before 1961 (Code, Art. 11.6).

Remarks. Name attributed to Sowerby by
Orbigny (1835a: 32) but we have been un-
able to find it in any Sowerby work.

guatemalensis

Ampullaria flagellata var. guatemalensis Mar-
tens, 1899: 413, pl. 22, fig. 11, 11a. N. Guate-
mala: Panzos ... Cahabon ... W. Guatemala:
Paso Antonio, in the lower part of the Rio
Michatoya, near the Pacific coast ... Cerro
Zunil. Syntypes: MCZ [labeled as paratypes],
ZMHB 109505 (4 spms.), 109506a (8 spms.),
109506b (1 spm.) (M. Glaubrecht, pers.
comm. to RHC, 1 March 2003; F. Kohler,
pers. comm. to RHC, 6 March 2003); no type
material found by us in BMNH (cf. Dance,
1986: 218). Distribution: Guatemala.
Remarks. Synonym of flagellata Say, 1829,
teste Sowerby (1909a: 352 [as “Morelet ?”]).

guyanensis

Ampullaria Guyanensis Lamarck, 1822a:
176. les rivieres de la Guyane. Lectotype
(Tillier, 1980: 27): MHNG 1093/90 (Y. Finet,
pers. comm. to RHC, 27 August 2002) (see
also Sowerby, 1909a: 349; Mermod, 1952: 84;
Pain, 1960: 427); probable paralectotypes:
MNHN (2 spms., “coll. Lamarck”; 1 spm.,
“coll. Buffon”; see also Tillier, 1980: 27). Dis-
tribution: Brasil, Peru, Colombia, Venezuela,
Guyana, French Guiana (Pain, 1960: 427).
Remarks. Subspecies of urceus Muller,
1774, teste Pain (1960: 426) and Geijskes &
Pain (1957: 47). Synonym of urceus Muller,
1774, teste Tillier (1980: 27), followed here.

haemastoma

Ampullaria heemastoma Reeve, 1856b: pl. 7,
fig. 34. Peru. Syntype: BMNH 20020659. Dis-
tribution: Peru.

Remarks. Synonym of guyanensis
Lamarck, 1822, teste Sowerby (1909a: 351;
1909b: 363) and Alderson (1925: 12), and

66

hence of urceus Müller, 1774 (see Тег,
1980: 27). N. syn.

hanleyana

Ampullaria hanleyana Alderson, 1926: 42.
Type material: lost [“Hanley’s type” is lost
(Pain 1951: 146)]. Distribution: Brasil (Pain,
1960: 424).

Remarks. Introduced as а new name for

swainsoni Hanley, 1854. However, Hanley
(1854: [unnumbered page], Ampullaria pl. 1,
fig. 1) clearly indicated that he thought he was
illustrating swainsoni Philippi, 1852, not a new
species. Thus, swainsoni Hanley, 1854, is not
a homonym of swainsoni Philippi, 1852, as
Alderson (1926: 42) thought, but а
misidentification. Alderson's (1926: 42) new
name and citing of Hanley's figure therefore
constitutes the original description of this spe-
cies. Not listed by Norris 8 Dance (2002: 377).
ратеу!
Ampullaria Натеу! Reeve, 1856e: pl. 23, fig.
113. [No locality given.] Type material: not
found by us in BMNH. Distribution: Rio
Parana (Ihering, 1919: 336).

Remarks. Synonym of pulchra Griffith &
Pidgeon [as “Gray”], 1834, teste Alderson
(1925: 33, 1926: 42).
haustrum
Ampullaria haustrum Reeve, 1856b: pl. 5, fig.
23. River Maranon. Possible syntype: BMNH
20020660. Distribution: Brasil, Bolivia, Peru
(Pain, 1960: 422-423).

Remarks. Synonym of canaliculata
Lamarck, 1822, teste Ihering (1898: 49) and
Thompson (1997: 91), but here retained as a
distinct species because of its reported pro-
duction of green eggs, in contrast to the pink
eggs of canaliculata Lamarck, 1822 (Cowie,
2002).
hollingsworthi
Pila (Pomacea) hollingsworthi Pain, 1946a:
180; pl. 6, figs. 3-5. Colombia, in a swiftly
flowing stream with a rocky bed near Bogota.
Holotype: BMNH 1946.6.24.24; paratype:
NMW.Z.1981.118.00198 (H. Wood, pers.
comm. to RHC, 30 October 2001). Distribu-
tion: Colombia.

Remarks. Belongs in Limnopomus Dall,
1904, teste Pain (1946a: 181), although Pain
agreed with Alderson (1925: 1) that
Limnopomus Dall, 1904, is not a distinguish-
able taxon.
hondurasensis
Ampullaria Hondurasensis Reeve, 1856a: pl.
3, fig. 15. Honduras. Syntypes: BMNH
20020662 (2 spms.). Distribution: Honduras,

COWIE & THIENGO

Guatemala (Nevill, 1884: 9), Nicaragua (Mar-
tens, 1899: 420).

Remarks. Synonym of flagellata Say, 1829,
teste Pain (1964: 227).

hopetonensis

Ampullaria Hopetonensis Lea, 1834: 115, pl.
19, fig. 84. Hopeton, near Darien, Georgia.
“Paratypes”: MCZ 151580 (Clench & Turner,
1956: 121). Distribution: USA (Georgia).

Remarks. Synonym of paludosa Say, 1829,
teste Alderson (1925: 29), Pilsbry (1927a:
249) and Clench & Turner (1956: 120).

immersa

Ampullaria immersa Reeve, 1856b: pl. 11, fig.
52. Rio Grande, Bolivia. Syntypes: BMNH
20020663 (1 spm.), MCZ [labeled as
“cotypes”]; topotypes: MCZ. Distribution: Bo-
livia (Sowerby, 1909a: 351).

Remarks. Synonym of canaliculata
Lamarck, 1822, teste Ihering (1898: 49). Syn-
onym of haustrum Reeve, 1856, teste Pain
(1960: 422).

innexa

Ampullaria innexa Crosse & Fischer, in
Fischer & Crosse, 1888: [explanation of] pl.
44, fig. 7, 7a—c [Crosse & Fischer, 1890: 111;
see also Fischer & Crosse (1890: 242)]. [No
locality given. Monte de Mistan, prope
Coapan, in provincia Oajaca, reipublicee
Mexicanæ (Crosse & Fischer (1890: 111).]
Type material: not found by us in MNHN. Dis-
tribution: Mexico.

Remarks. Synonym of flagellata Say, 1829,
teste Pain (1964: 227).

insularum

Ampullaria insularum Orbigny, 1835a: 32. Rio
Parana (republica Argentina). Syntypes:
BMNH 1854.12.4.309-313 (7 spms.), MNHN
(3 lots, 5 spms.), MHNG 33487 (2 spms.) (У.
Finet, pers. comm. to RHC, 26 August 2002).
Distribution: Argentina, Brasil (Baker, 1914:
659).

Remarks. Synonym of gigas Spix, 1827,
teste Ihering (1898: 49) and $. С. Thiengo
(unpublished), followed here, although con-
trary to various authors (e.g., Baker, 1914:
659), who treated it as a valid species.

interrupta

Ampullaria interrupta Sowerby, 1909a: 353
[name], 361 [description], text fig. Laguna
Urao, Venezuela. Syntype: BMNH
1909.10.19.33 (see also Alderson, 1925: 52;
Pain, 1950a: 110). Distribution: Venezuela.
Remarks. Placed in Limnopomus Dall,
1904, by Sowerby (1909a: 361) and dis-
cussed as such by Pilsbry (1933: 75).

NEW WORLD AMPULLARIIDAE 67

intropicta
Ampullaria intropicta Reeve, 1856d: pl. 21,
fig. 101a, b. [No locality given.] Syntypes:
BMNH 20020664 (3 spms.). Distribution:
Brasil (syntype label).

Remarks. Synonym of decussata

Moricand, 1836, teste Sowerby (1909a: 348).
labiosa

Ampullaria labiosa Philippi, 1852a: 58, pl. 18,

fig. 5 [1852b: 28]. [No locality given.] Holotype

[“das einzige Exemplar” (Philippi, 1852a: 58)]:

Koch collection, location not known to us.

Distribution: unknown.

Remarks. Philippi (1852a: 58) attributed the
name to Koch. Synonym of flagellata Say,
1829, teste Pain (1964: 227), though listed
as from India by Paetel (1887: 479) and
“Indes orientales” by Gaudion (1879: 32).

lamarckii
Ampullaria Lamarckii Philippi, 1852a: 67, pl.
21, fig. 5. [No locality given.] Type material:
probably MNHNS. Distribution: unknown.

Remarks. Synonym of flagellata Say, 1829,
teste Pain (1964: 227).

lattrei
Ampullaria Lattrei Reeve, 1856b: pl. 5, fig. 22.
Coban, Guatemala. Syntypes: BMNH

20020665 (2 spms.). Distribution: Guatemala
(Martens, 1899: 419; Sowerby, 1909a: 354;
Pain, 1964: 229).

Remarks. See delattrei Fischer & Crosse,
1890.

lemniscata

Ampullaria lemniscata Crosse & Fischer, in
Fischer 8 Crosse, 1888: [explanation of] pl.
44, fig. 5, 5a—c [Crosse & Fischer, 1890: 112;
see also Fischer & Crosse (1890: 248)]. [No
locality given. coloniâ апайса Belize (Crosse
& Fischer (1890: 112).] Syntypes: MNHN (4
spms.) (see also Sowerby, 1909b: 363). Dis-
tribution: Belize, Mexico (Sowerby, 1909a:
352).

Remarks. Synonym of flagellata Say, 1829,
teste Pain (1964: 227).

leucostoma

Ampullaria leucostoma Swainson, 1823a: pl.
175. [No locality given.] Type material: possi-
bly MMUE (Dean, 1936: 232; H. McGhie,
pers. comm. to RHC, 29 July 2002), not
found by us in BMNH (cf. Dance, 1986: 227).
Distribution: Venezuela (Paetel, 1887: 479).

Remarks. Synonym of urceus Müller, 1774,
teste Philippi (1852a: 54), Gaudion (1879:
41), Sowerby (1909a: 358) and Alderson
(1925: 10).

levior

Ampullaria levior Sowerby, 1909a: 354
[name], 361 [description], text fig. Amazon
River. Syntype: BMNH 1909.10.19.36. Distri-
bution: Amazon River; Surinam, Brasil
(Vernhout, 1914a: 28, 43).

Remarks. Зупопут of lineata Spix, 1827,
teste Pain (1960: 422).

lineata

Helix lineata Spix, in Wagner, 1827: 3, pl. 5,
fig. 2. in aquis Provinciae Bahiensis, e.g. in
fluvio Itahype [see also Thiengo (1987: 563)].
Syntypes: ZSM 20012054 (1 spm.),
20012066 (1 spm.), 20012074 (1 spm.) (E.
Schwabe, pers. comm. to RHC, 28 July
2002; cf. Fechter, 1983: 221), MNHN (2
spms.). Distribution: Brasil, Guyana, French
Guyana, Surinam (Sowerby, 1909a: 354;
Baker, 1914: 660; Vernhout, 1914a: 43)
[Brasil only, teste Pain (1960: 422)].

Remarks. Authorship 1$ given here as “Spix,
in Wagner”, following Cowie et al. (in prep.),
who also explain the publication history of this
work. Pain (1950b: 72) listed “Helix liniata
Spix” in the synonymy of crassa Swainson,
1823 (although he cited Spix’s pl. 5, fig. 1),
but subsequently (Pain, 1960: 422), followed
here, treated /ineata Spix, 1827, as a valid
species. Misspelled “/ineolata” by Deshayes
(1850: 44).

linnaei

Ampullaria Linnaei Philippi, 1852a: 62, pl. 20,
fig. 6 [1852b: 29]. [No locality given.] Holotype
[eines ... Exemplares” (Philippi, 1852a: 62)]:
probably MNHNS. Distribution: unknown.

Remarks. Synonym of lineata Spix, 1827,
teste Sowerby (1909a: 354).

+ livescens

Ampullaria livescens Reeve, 1856b: pl. 5, fig.
21. [No locality given.] Syntype: BMNH
1986214. Distribution: Tabasco and Chiapas,
Mexico; Lake Petén, northern Guatemala
(Pain, 1964: 228).

Remarks. Subspecies of flagellata Say,
1829, teste Pain (1964: 228).

lutea

Poomacea [sic] paludosa Say var. lutea
Farfante, 1942: 51. Unavailable name; nom.
nud.

Remarks. Listed as a synonym of paludosa
Say, 1829, by Clench & Turner (1956: 120).

lymnaeaeformis

Ampullaria lymnææformis Reeve, 18565: pl.
8, fig. 39. River Marañon. Syntypes: BMNH
20020666 (2 spms.). Distribution: Peru.

68

Remarks. Зупопут of aulanieri Deville &
Huppé, 1850, teste Pain (1960: 424). Fre-
quently spelled “/ymnaeformis”.
maculata
Pomacea maculata Perry, 1810c: [unnum-
bered plate and text] [= pl. 12 (Mathews &
Iredale, 1912: 11; Geijskes & Pain, 1957: 42;
К. Е. Petit, pers. comm. to RHC, 16 October
2000)]. the South Sea [error; Mathews &
Iredale, 1912: 11]. Type material: not found by
us in BMNH (cf. Dance, 1986: 221). Distribu-
tion: Brasil, Peru (Pain, 1960: 423).

Remarks. Possibly a synonym of urceus
Müller, 1774, teste Berthold (1991: 248).
malleata
Ampullaria malleata Jonas, 1844: 35 [1846:
122, pl. 10, fig. 11, 11a, 11b]. Juxta Tabasco,
urbem Mexicanum. Lectotype [“le type de
Jonas” (Fischer & Crosse, 1890: 237)]: prob-
ably ZMHB 109515 (M. Glaubrecht, pers.
comm. to RHC, 1 March 2003; F. Kohler,
pers. comm. to RHC, 6 March 2003, 20
March 2003); possible paralectotypes [la-
beled paratypes]: MCZ. Distribution: Mexico.

Remarks. Fischer & Crosse (1890: 237)
stated that the “type” of Jonas was collected
in Tabasco by Fokkes. Martens (1899: 412)
mentioned two specimens, one collected
from Tabasco by Fokkes and given by Jonas
to Dunker, and another, from the Dunker col-
lection, illustrated by Martens (1899: pl. 22,
fig. 10) but with no mention of its collector or
whether it had ever been in Jonas’ posses-
sion. Although Martens (1899: 412) sug-
gested that the latter specimen might be
Jonas’ “type”, it seems more likely that the
former is the “type” and that it is the speci-
men indicated as such by Fischer & Crosse
(1890: 237). Martens’ figure and that of Jonas
(1846: fig. 11) are almost identical. Jonas
gave his collection to ZMUH, which would ex-
plain the statement of Fischer & Crosse
(1890: 237) that Jonas’ “type” was there. Mar-
tens, however, stated that both his speci-
mens were from the Dunker collection, in
ZMHB, and the specimen ZMHB 109515 al-
most perfectly matches Jonas’ figure, even
to the small depression in the lower rim of
the aperture (Е. Köhler, pers. comm. to RHC,
20 March 2003). This probably came about
through donation or exchange, as Dunker
certainly exchanged material with collectors
in Hamburg, because there is material of
other taxa from him that ZMUH obtained from
the Altonaer Museum (another museum in

COWIE & THIENGO

Hamburg) after the ZMUH collections were
destroyed in the Second World War (B.
Hausdorf, pers. comm. to RHC, 10 March
2003). However, it yet could be that Fischer &
Crosse (1890: 237) and Martens (1899: 412),
although both referred to the “type”, were ac-
tually referring to different shells. Synonym of
flagellata Say, 1829, teste Baker (1922: 37)
and Pain (1964: 226).

manco

Pomacea manco Pilsbry, 1944: 145, pl. 11,
figs. 31, 32. collecting station 161, on the
Pachitea River, about one mile upstream
from Quebrada Sungarillo. Holotype: ANSP
Invertebrate Paleontology 4596 [“4596a”
(Baker, 1964: 168)]; paratypes: ANSP In-
vertebrate Paleontology 78898 (2 spms.).
Distribution: Peru (Boss & Parodiz, 1977:
110).

Remarks. Fossil. Placed in Limnopomus
Dall, 1904, by Parodiz (1969: 110).

manetou

Pila Manetou Róding, 1798: 145. [No locality
given.] Type material: possibly Art and Natural
History Museum, Gotha (Stewart, 1930: 35;
Dance, 1986: 206). Distribution: unknown.

Remarks. Synonym of urceus Muller, 1774,
teste Baker (1930: 2).

+ marginatra

Ampullaria marginatra Jonas, 1845: 169. [No
locality given.] Type material: “in Museo hon.
Gruner” (Jonas, 1845: 169), ZMHB 29964
(lost; M. Glaubrecht, pers. comm. to RHC, 1
March 2003). Distribution: unknown.

Remarks. Variety of zonata Spix, 1827,
teste Philippi (1851: 10; 1852a: 63, 74) and
Sowerby (1909a: 359).

martensiana

Ampullaria (Pomus) martensiana Nevill, 1884:
10. New name for columbiensis Reeve, 1856;
non Philippi, 1851. Distribution: Colombia.

Remarks. Synonym of flagellata Say, 1829,
teste Pain (1964: 227).

martinezi

Ampullaria Martinezi Hidalgo, 1866: 345, pl.
14, fig. 5. Santa-Rosa, Reipublicæ Æquatoris
[Ecuador]. Lectotype: MNHN (Fischer-Piette,
1950: 68); paralectotypes: MNHN (1 spm)
(Fischer-Piette, 1950: 68), MNCN 15.05/7524
(1 spm.), 15.05/12306 (7 spms.) (Villena et
al., 1997: 75). Distribution: Ecuador (Miller,
1879: 151; Sowerby, 1909a: 354).

melanocheila

Ampullaria melanocheila Reeve, 18565: pl. 5,
fig. 24. [No locality given.] Syntype: BMNH

NEW WORLD AMPULLARIIDAE 69

20020667. Distribution: Brasil (Paetel, 1887:
480).

Remarks. Зупопут of sordida Swainson,
1823, teste Sowerby (1909a: 357).
melanostoma
Ampullaria reflexa Var. melanostoma Philippi,
1852a: 35, 58, pl. 18, fig. 4. [No locality given.]
Syntype: ZMHB 109500 (M. Glaubrecht, pers.
comm. to RHC, 1 March 2003); type material
possibly also in MNHNS. Distribution: un-
known.

Remarks. Philippi (1852a: 35) attributed the
name to “Parr. in litt.” Synonym of malleata
Jonas, 1844, teste Martens (1857: 189, 207),
but a variety of flagellata Say, 1829, teste
Martens (1899: 411). Treated here as a syn-
onym of flagellata Say, 1829. N. Syn.
meridaensis
Pomacea (Limnopomus) meridaensis Pain,
1950a: 109. Merida, Venezuela. Holotype
(Pain, 1950a: 110): the specimen figured by
Alderson (1925: pl. 11, fig. 7); paratypes:
Alderson collection (Pain, 1950a: 110), HUJ
21517 (1 spm.) (H. Mienis, pers. comm. to
RHC, 4 August 2002), MCZ 171558 (2
spms.), FMNH (1 spm.), ZSM 20012068 (1
spm.; ex Alderson collection) (E. Schwabe,
pers. comm. to RHC, 29 July 2002). Distribu-
tion: Venezuela.

Remarks. Synonym of camena Pain, 1949,
teste Pain (1957: 175).
mermodi
Ampullaria mermodi Sowerby, 1919: 152, [un-
numbered text figure]. Central America.
Syntypes: MHNG 33490 (3 spms.) (Y. Finet,
pers. comm. to RHC, 26 August 2002; see
also Tillier, 1980: 19 [as “Sowerby, 1905”]).
Distribution: ? Guyana, ? Central America
(Pain, 1950b: 72).

Remarks. Pain (1950b: 72) stated that this
species was founded on a single specimen.
However, this was not made explicit by
Sowerby (1919: 152-153), who described it
from “photographs” sent to him by Mermod.
Tillier (1980: 19) indicated that Sowerby's
(1919: 152) figure illustrated the largest of
three syntypes.
meta
Ampullaria meta lhering, 1915: 12, pl. [3], figs.
6, 7. Cidade da Barra, Rio S. Francisco River,
Bahia. Holotype: ММК). Distribution: Brasil.

+ metcalfei

Ampullaria Metcalfei Reeve, 1856e: pl. 25,
fig. 119a, b. [No locality given.] Type material:
not found by us in BMNH. Distribution: Ven-
ezuela (Baker, 1930: 4).

Remarks. Possibly a synonym of vexillum
Reeve, 1856, teste Alderson (1925: 14). Sub-
species of swainsoni Philippi, 1852, teste
Baker (1930: 4), although he noted collec-
tions containing both forms and a “good se-
ries of intermediates”.

mexicana

Ampullaria Mexicana Martens, 1857: 207.
Unavailable name; nom. nud.

Remarks. Listed by Martens (1857: 207) as
a manuscript name of Philippi; also listed by
Gaudion (1879: 33). Treated as a synonym
of malleata Jonas, 1844, by both these au-
thors. Also listed by Paetel (1873: 65, 1887:
480).

+ miamiensis

Ampullaria miamiensis Pilsbry, 1899: 365
[1927a: 252, pl. 22, figs. 5 (lectotype), 6, 7].
creek flowing from the Everglades near Mi-
ami, Dade County, in southeastern Florida.
Lectotype (Pilsbry, 1927a: 253; see also
Baker, 1964: 168): ANSP 77369;
paralectotypes: ANSP 361441 (59 spms.,
uncounted juveniles), CMNH 62.19966 (1
spm.), 62.33743 (1 spm.) (Parodiz & Tripp,
1988: 141), USNM (1 spm.) [labeled as
“cotype”], MCZ (“paratypes”; Clench &
Turner, 1956: 122). Distribution: Florida.

Remarks. Treated by Clench & Turner
(1956: 122) as a “race” or “local population” of
paludosa Say, 1829, but not formally synony-
mized.

miltocheilus

Ampullaria miltocheilus Reeve, 1856e: pl. 25,
fig. 120a, b. Province of Chiapes [= Chiapas],
Mexico. Lectotype [“Le type”; Fischer &
Crosse (1890: 248)]: ВММН 20020668/1;
paralectotypes: 20020668/2-5 (5 spms.);
based on our study of the BMNH material.
Distribution: Mexico.

Remarks. Variety of ghiesbreghti Reeve,
1856, teste Martens (1899: 418). Synonym of
cumingii King & Broderip, 1831, teste
Sowerby (1909a: 348). Probably a synonym
of quitensis Busch, 1859, teste Alderson
(1925: 44). Not a synonym of ghiesbreghti
Reeve, 1856, teste Pain (1953: 223), who
also remarked on its shell “resembling spe-
cies of Limnopomus”.

miltochilus

Ampullaria miltochilus Fischer & Crosse,
1890: 247. Unjustified emendation of
miltocheilus Reeve, 1856.

+ minor

Ampullaria (Pomus) gigas var. minor Nevill,
1884: 9. les environs de Corrientes, et sur les

70

rivages de la Plata, près de Buenos-Ayres [lo-
cality given by Orbigny (1838d: 372) for the
variety illustrated in his pl. 50, fig. 5]. Holo-
type/syntypes: the specimen(s) illustrated by
Orbigny (1838e: pl. 50, figs. 5, 6) [the two fig-
ures probably illustrate a single (live) speci-
men but this is not certain], location not
known to us. Distribution: La Plata (Orbigny,
1838d: 372), Rio Parana (Nevill, 1884: 9).

Remarks. Name proposed by bibliographic
reference to Orbigny (1838e: pl. 50, figs. 5, 6).
Junior primary homonym of minor Nevill, 1877,
which is now placed in Pila Roding, 1798.
modesta
Ampullaria modesta Busch, 1859: 168. Ecua-
dor. Type material: location not known to us.
Distribution: Ecuador (Miller, 1879: 150).
monachus
Ampullaria monachus Crosse & Fischer, in
Fischer & Crosse, 1888: [explanation of] pl.
46, fig. 5, 5a [Crosse & Fischer, 1890: 112 [as
“monacha’]; see also Fischer & Crosse
(1890: 250)]. [No locality given. Santa
Efigenia, in Isthmo Tehuantepecensi,
reipublicae Mexicanae (Crosse & Fischer
(1890: 113)]. Holotype: the specimen illus-
trated by Crosse & Fischer, in Fischer &
Crosse (1888, fig. 5, 5a), not found by us in
MNHN. Distribution: Mexico.

Remarks. Synonym of flagellata Say, 1829,
teste Pain (1964: 227).

monstrosa

Ampullaria fasciata var. monstrosa Sowerby,
1825: 44. Unavailable name; nom. nud.
Remarks. Not listed by Sherborn (1922-
1933).
nais
Pomacea nais Pain, 1949a: 257; pl. 13, figs.
3, 4. small stream on the south bank of the
Amazon near Obidos, Brasil. Holotype:
BMNH 1947.2.3.1 [not 1946.2.3.1, as stated
by Pain (1949a: 257)]; paratype:
NMW.Z.1981.118.00093 (Pain collection, 1
spm.). Distribution: Brasil (Pain, 1960: 424).
Remarks. May be a “local race” of lineata
Spix, 1827, teste Pain (1960: 424).
nigrilabris
Ampullaria nigrilabris Philippi, 1852a: 65, pl.
21, fig. 2[1852b: 29]. [No locality given.] Type
material: probably MNHNS. Distribution: “Rio
Janeiro” (Gaudion, 1879: 34; Sowerby, 1909a:
355; see also Paetel, 1873: 65, 1887: 480).
nobilis
Ampullaria nobilis Reeve, 1856a: pl. 2, fig. 8.
River Marañon. Possible syntype: BMNH

COWIE & THIENGO

20020669; topotypes: ANSP 120276 (Baker,
1930:3). Distribution: Venezuela (Baker,
1930: 3), East Peru (Sowerby, 1909a: 354),
Brasil (Baker, 1914: 660).

Remarks. Synonym of guyanensis
Lamarck, 1822, teste Pain (1960: 427), and
hence of urceus Müller, 1774 (see Tillier, 1980:
27). М. Syn.

notabilis

Ampullaria notabilis Reeve, 1856c: pl. 14, fig.
63. [No locality given.] Syntype: BMNH
20020670. Distribution: Peru (Sowerby,
1909а: 355) [? error; Alderson, 1925: 45;
Pilsbry, 1927a: 250].

Remarks. Зупопут of nubila Reeve, 1856,
teste Paetel (1887: 480). Possibly a syn-
onym of paludosa Say, 1829, teste Alderson
(1925: 45) and Pilsbry (1927a: 250). Re-
tained here as a distinct species, pending fur-
ther study.

novaegranadae

Ampullaria novee-granadee Busch, 1859: 169.
New Granada [in 1859 = present-day Colom-
bia and Panama]. Syntypes: ВММН
20020671 (2 spms.). Distribution: Colombia
and/or Panama.

oajacensis

Ampullaria malleata var. Oajacensis Fischer
& Crosse, 1890: 235 [1888: pl. 46, fig. 3, За,
3b; plate published without name]. Monte de
Mistam, prope Coapam, provincias Oajaca.
Type material: Sallé collection, not found by
us in BMNH, MNHN, etc. (cf. Dance, 1986:
209, 225). Distribution: Mexico.

Remarks. Synonym of flagellata Say, 1829,
teste Pain (1964: 227).

oblonga

Ampullaria oblonga Swainson, 1823a: pl.
136, middle figs. [No locality given.] Syntypes:
“in the late Mrs. Bligh's collection”
(Swainson, 1823a: pl. 136), location not
known to us. Distribution: Venezuela (Philippi,
1851: 21; Sowerby, 1909a: 355), Guadeloupe
(Gaudion, 1879: 35).

Remarks. Synonym of urceus Müller, 1774,
teste Pain (1960: 426), but retained here as a
valid species based on our own observations
($. С. Thiengo, unpublished).

ocanensis

Ampullaria (aurisformis var. ?) ocanensis
Kobelt, 1914b: 222, pl. 77, figs. 4, 5[1914e:
177]. Ocaña in Neu-Granada [= Colombia].
Figured specimen (Kobelt, 1914b: figs. 4, 5):
ZMUH [destroyed; B. Hausdorf, pers. comm.
to RHC, 3 May 2002]. Distribution: Colombia.

NEW WORLD AMPULLARIIDAE 71

occlusa

Ampullaria occlusa Crosse & Fischer, in
Fischer & Crosse, 1888: [explanation of] pl.
45, fig. 3, 3a-c [Crosse & Fischer, 1890: 111;
see also Fischer & Crosse (1890: 244)]. [No
locality given. Tanesco, Guatemalæ (Crosse
& Fischer (1890: 112).] Syntypes: ММНМ (2
lots, 11 spms.) (see also Sowerby, 1909b:
363). Distribution: Guatemala.

Remarks. Synonym of flagellata Say, 1829,
teste Pain (1964: 227).
ochracea
Ampullaria ochracea Jay, 1836: [85 (explana-
tion of pl. 3)], pl. 3, fig. 8 [1839: [explanation of
pl. 3, fig. 8]. Spanish Maine [= isthmus of
Panama to mouth of Orinoco River].
Syntypes: AMNH 56106 (1 spm.) [labeled as
“figd type” in Jay’s handwriting (P. M.
Mikkelsen, pers. comm. to RHC, 7 May
2002)], 56106A (1 spm.); additional 6
syntypes [Jay (1839: 116) mentioned 8 spms.
in total]: location not known (Boyko &
Cordeiro, 2001: 16).

Remarks. Synonym of flagellata Say, 1829,
teste Martens (1899: 405) and Pain (1964:
226).

+ olivacea

Ampullaria olivacea Spix, in Wagner, 1827: 2,
pl. 3, fig. 1. in fluminibus Amazonum,
Solimoes, Japurá alliisque in interiore
continente Brasiliae aequatorialis. Type mate-
rial: probably lost (Fechter, 1983: 221; $. С.
Thiengo, unpublished). Distribution: Brasilian
Amazon (Pain, 1960: 428).

Remarks. Authorship is given here as
“Spix, in Wagner”, following Cowie et al. (т
prep.). Junior primary homonym of
Ampullaria olivacea Lamarck, 1816. How-
ever, Lamarck (1822a: 178), followed by
Philippi (1852a: 28), placed olivacea
Lamarck, 1816, in the synonymy of
Ampullaria guinaica Lamarck, 1822, which
has long been placed in the African genus
Lanistes (e.g., Nevill, 1884: 14). Thus, be-
cause olivacea Spix, 1827, and olivacea
Lamarck, 1816, have not been considered
congeneric after 1899, no replacement
name is provided and the case must be re-
ferred to the ICZN for a ruling (Code, Art.
23.9.5). Wagner (1827: 2) listed the older
name guyanensis Lamarck, 1822, in syn-
onymy. However, Pain (1960: 427), followed
here, treated olivacea Spix, 1827, as a sub-
species of urceus Muller, 1774, and distinct
from guyanensis Lamarck, 1822.

oviformis

Ampullaria oviformis Deshayes, 1830a: 34.
Cayenne. Syntypes: MNHN (2 lots, 2 spms.).
Distribution: French Guiana (Sowerby,
1909a: 355) [? error; Tillier, 1980: 16].

Remarks. The two syntypes are clearly two
different species, indicating the need for fur-
ther study to clarify this species’ true identity
(see also Tillier, 1980: 16).

palmeri

Ampullaria palmeri Marshall, 1930: 4, pl. 1,
figs. 5, 8. small stream in dense jungle, 13
kilometers south of Puerto Santos, Province
of Santander del Norte, Republic of Colom-
bia. Holotype: USNM 380696; paratypes:
USNM 380697. Distribution: Colombia.

paludosa

Ampullaria paludosa Say, 1829c: 260. New
name for depressa Say, 1824, non Lamarck,
1804. Distribution: USA (Alabama, Georgia,
Florida), Cuba (Clench & Turner, 1956: 122;
Cowie, 1997b: 5).

papyracea

Ampullaria papyracea Spix, in Wagner, 1827:
3, pl. 4, figs. 1, 2. in fluviis et stagnis
Provinciarum Bahiensis, Pernambucanae et
Piauhiensis. Syntypes: ZSM 20012059 (2
spms.) (E. Schwabe, pers. comm. to RHC,
28 July 2002; see also Fechter, 1983: 221).
Distribution: Brasil, Peru, Venezuela, Guyana,
Surinam, French Guiana (Pain, 1950b: 66,
1960: 429).

Remarks. Authorship is given here as “Spix,
in Wagner”, following Cowie et al. (in prep. ).

patula

Ampullaria patula Reeve, 1856d: pl. 21, fig.
100a, b. [No locality given.] Syntypes: BMNH
20020673 (3 spms.). Distribution: Amazon,
Brasil, New Granada [= Colombia and
Panama] (Walker, in Baker, 1922: 39).
Remarks. Junior primary homonym of
patula Lamarck, 1804, which is now placed in
the family Naticidae (see also Lamarck,
1822b: 549). Not listed by Sowerby (1912: 72).

pealiana

Ampullaria pealiana Lea, 1838: 16, pl. 23, fig.
77. Turbaco, Colombia, South America. Lecto-
type [as “Figured holotype”) (Abbott, 1955: 126,
pl. 4, fig. 2): ANSP 192933; paralectotypes:
MCZ 161600. Distribution: Ecuador, Colombia
(Pain, 1956a: 78), Venezuela (Paetel, 1887:
480), Panama (Martens, 1899: 423).

Remarks. We treat “pealeana” Philippi,
1852 (1852a: 62) as an incorrect subsequent
spelling.

72 COWIE & ТНЕМСО

penesma
Ampullaria penesma DeKay, 1843: 124. Un-
available name; first published as a junior
synonym of paludosa Say, 1829, not made
available before 1961 (Code Art. 11.6).

Remarks. DeKay (1843: 124) attributed the
name to Say as a manuscript name, but the
name does not occur in Say's published writ-
ings (Binney, 1858: [237], 1865: 5).

periscelis
Pila periscelis Róding, 1798: 146. [No locality
given.] Type material: possibly Art and Natu-
ral History Museum, Gotha (Stewart, 1930:
35; Dance, 1986: 206).

Remarks. Possibly a synonym of
chemnitzii Philippi, 1852, teste Baker (1930:
a).

peristomata
Ampullaria peristomata Orbigny, 1835a: 33.
Guarayos (republica Boliviana). Syntypes:
BMNH 1854.12.4.331 (10 spms.), MNHN (2
lots, 6 spms.). Distribution: Brasil (Baker,
1914: 660), Peru (Paetel, 1888: 481), Bolivia.

Remarks. Synonym of cumingii King &
Broderip, 1831, teste Sowerby (1909a: 348)
and Kobelt (1912h: 141), and of elegans
Orbigny, 1835, teste Gray (1855: 29), but
treated as a valid species by Baker (1914:
660), followed here. The BMNH and MNHN
syntype lots are clearly two different species,
indicating the need for further study to clarify
this species’ true identity.

pernambucensis
Ampullaria Pernambucensis Reeve, 1856d:
pl. 22, fig. 103. Pernambuco. Syntypes:
BMNH 20020674 (3 spms.). Distribution:
Brasil.

+ pertusa
Ampullaria pertusa Sowerby, 1894: 48, pl. 4,
fig. 22. [No locality given.] Holotype (the single
specimen on which the description was ex-
plicitly based): ВММН 20020675 (figured also
by Pain, 1949b: pl. 1, figs. 3, 4). Distribution:
Venezuela (Sowerby, 1909a: 355, Pain,
1949b: 39).

Remarks. Variety of castelloi Sowerby,

1894, teste Pain (1949b: 39).

phaeostoma
Ampullaria phaeostoma Philippi, 1852a: 45,
pl. 13, fig. 3 [1852b: 26]. [No locality given.]
Type material: probably MNHNS. Distribution:
“Haut-Amazone” (Gaudion, 1879: 37) [? er-
ror].

Remarks. Synonym of flagellata Say, 1829,
teste Pain (1964: 227).

physis

Ampullaria physis Hupe, 1857: 67, pl. 12, fig.
2 [two figs.]. le fleuve des Amazones.
Syntypes: MNHN (2 lots, 4 spms.). Distribu-
tion: Amazon River (Sowerby, 1909a: 356).

Remarks. Synonym of lineata Spix, 1827,
teste Pain (1960: 422).

physoides

Ampullaria Physoides Reeve, 1856d: pl. 22,
fig. 107a, b. Pernambuco. Syntypes: BMNH
20020676 (4 spms.). Distribution: Brasil, India
[error; one of the BMNH syntypes has а horny
operculum, indicating its New World origin]
(Paetel, 1888: 481).

picta

Ampullaria picta Reeve, 1856e: pl. 24, fig.
117a, b. [No locality given.] Syntypes: BMNH
1907.11.21.91-92 (2 spms.). Distribution:
Mexico (Mazatlan) (Sowerby, 1909a: 356).

pinei

Ampullaria Pinei Dall, 1898: 75. Homosassa
River, Florida. Possible syntype: USNM
152699 [labeled as the figured “type”, al-
though Dall (1898: 75-76) did not designate
or figure a type]. Distribution: USA.

Remarks. Synonym of paludosa Say, 1829,
teste Clench & Turner (1956: 120).

роеуапа

Атри/апа poeyana Pilsbry, 1927a: 251, pl. 21,
figs. 7, 8 ["Type”], 9. New name for cubensis
Reeve, 1856, non Morelet, 1849. Distribution:
Cuba.

Remarks. Pilsbry (1927a: 251, 253) provided
this name as а “п. sp.” and designated а “ho-
lotype” (ANSP 50618) ["50618a” (Baker, 1964:
168)]. Two specimens from the same lot are
now ANSP 365370. However, Pilsbry was sim-
ply providing a replacement name for
cubensis Reeve, 1856, so the type material of
this species is Reeve's and Pilsbry's designa-
tion of a holotype is invalid. Although cubensis
Reeve, 1856, has been considered a variety
or synonym of glauca Linnaeus, 1758 (which
is listed here under Pomacea subg. Effusa
Jousseaume, 1889), we follow Pilsbry (1927a:
251-252) in retaining poeyana Pilsbry, 1927,
as a valid species in Pomacea $. str.

pomatia

Ampullaria pomatia Martens, 1857: 194.
Brasilien. Syntypes: ZMHB 1366a (3 spms.),
1366b (3 spms.) (M. Glaubrecht, pers. comm.
to RHC, 1 March 2003). Distribution: Brasil.

pomum

Ampullaria pomum Philippi, 1851: 13, pl. 3,
figs. 3, 4 [1852b: 20]. [No locality given.] Type

NEW WORLD AMPULLARIIDAE 73

material: probably MNHNS. Distribution: un-
known.
porphyrostoma

Ampullaria porphyrostoma Reeve, 18565: pl.
6, fig. 30. [No locality given.] Syntypes:
BMNH 20020677 (3 spms.). Distribution: Ven-
ezuela (Baker, 1930: 5), New Granada [=
present-day Colombia and Panama from
1830 to 1903 and Colombia only from 1903
on] (Sowerby, 1909a: 353).

Remarks. Synonym of chemnitzii Philippi,
1852, teste Baker (1930: 5) and Pain (1956a:
74).

prasina
Ampullaria malleata var. Prasina Fischer &
Crosse, 1890: 235, pl. 48, fig. 4, 4a. Misantla,
provinciæ Vera Cruz. Type material: not
found by us in MNHN. Distribution: Mexico.

Remarks. Synonym of flagellata Say, 1829,
teste Pain (1964: 227).

producta
Ampullaria producta Reeve, 1856c: pl. 15,
fig. 68a, b. [No locality given.] Syntypes:
BMNH 20020678 (3 spms.). Distribution: “F.
Magdalen” [= Colombia] (Paetel, 1888: 481),
Amazon River (Sowerby, 1909a: 356), Neu-
Granada [= Colombia and/or Panama]
(Kobelt, 1913h: 205).

prourceus
Ротасеа (Pomacea) prourceus Boss &
Parodiz, 1977: 110, figs. 1-4. Chicocoa (a
single farmhouse on the east bank of the Río
Huallaga ...), east of Chasuta [Chazuta] (6°
352’ 5; 76° 112’ W), the Río Huallaga, Depart-
ment of San Martín, Peru. Holotype: MCZ
272899. Distribution: Peru.

Remarks. Tertiary fossil, possibly middle or

late Eocene (Boss 8 Parodiz, 1977: 110).

pulchra

Paludina pulchra Griffith 8 Pidgeon, 1834b:
599, pl. 1, fig. 6 [pl. 1 predated p. 599 and
was possibly published in 1833 (Cowan,
1969: 139)]. [No locality given.] Syntype:
BMNH 20020680. Distribution: South America
(Sowerby, 1909a: 356).

Remarks. Name attributed to Gray by
Griffith & Pidgeon (1834b: 599). Placed in
Pomacea [аз Ampullaria] by Sowerby
(1909a: 356).

puncticulata
Ampullaria puncticulata Swainson, 1823a: pl.
143, figs. 3, 4 [middle figs.]. [No locality
given.] Type material: possibly MMUE (Dean,
1936: 232; H. McGhie, pers. comm. to RHC,
28 July 2002), not found by us in BMNH (cf.

Dance, 1986: 227). Distribution: Brasil [error]
(Drouét, 1859: 81), Colombia, Guyana,
French Guiana (Pain, 1950b: 72); also Ven-
ezuela (Gaudion, 1879: 38). Spelled as
“punctulata” by Mousson (1873: 18), Paetel
(1888: 481) and Ihering (1919: 332).

Remarks. Synonym of guyanensis
Lamarck, 1822, teste Pain (1960: 426), and
hence of urceus Muller, 1774 (see Tillier,
1980: 27). N. Syn.

puntaplaya

Ampullaria puntaplaya Cousin, 1887: 278, pl.
4, fig. 2. Punta-Playa. Syntypes: MNHN (2
lots, 4 spms.). Distribution: Ecuador
(Sowerby, 1909a: 356).

+ purpurascens

Ampullaria purpurascens Guppy, 1864: 243.
Trinidad [in publication title]. Syntypes
(Guppy, 1864: 248): BMNH, not found by us.
Distribution: Trinidad (Sowerby, 1909a: 356).

Remarks. Treated as a variety of urceus
Müller, 1774, by Guppy (1866: 44).

+ ругит

Атри/апа ругит Philippi, 1851: 18, pl. 5, fig.
2 [1852b: 21]. Brasilien. Syntype: 2$М
20012060 (Е. Schwabe, pers. comm. to
RHC, 28 July 2002). Distribution: Brasil
(Gaudion, 1879: 38).

Remarks. Variety of hopetonensis Lea,
1834 (= paludosa Say, 1829), teste Sowerby
(1909a: 353), but note the skepticism of Pain
(1964: 225) regarding this. Either it is not a
variety (or synonym) of paludosa Say, 1829,
or the locality (Brasil) is incorrect.

quercina

Ampullaria quercina Spix, in Wagner, 1827: 2,
pl. 3, fig. 2. in fluminibus Amazonum,
Solimoès, Japurá alliisque in interiore
continente Brasiliae aequatorialis [as for
olivacea Spix, 1827]. Syntype: ZSM 20012061
(E. Schwabe, pers. comm. to RHC, 28 July
2002; cf. Fechter, 1983: 221). Distribution:
Amazon drainage (Pain, 1960: 428).

Remarks. Authorship is given here as “Spix,
in Wagner”, following Cowie et al. (in ргер.),
who also explain the publication history of this
work. Spix illustrated quercina as a full spe-
cies, but Wagner, in writing the description,
treated quercina “Spix” as a variety of
olivacea Spix, 1827. Retained as a variety by
Sowerby (1909a: 355), but treated here as a
distinct species, following Pain (1960: 428).
Berthold (1991: 23) placed quercina “Wagner
non Spix” in Pomacea subg. Effusa
Jousseaume, 1889.

74

quitensis
Ampullaria quitensis Busch, 1859: 168. Ecua-
dor. Type material: location not known to us.
Distribution: Ecuador (Miller, 1879: 149).
Remarks. Synonym of cumingii King &
Broderip, 1831, teste Sowerby (1909a: 348).
reflexa
Ampullaria reflexa Swainson, 1823b: 377
[1823a: pl. 172]. [No locality given.] Type ma-
terial: possibly MMUE (Dean, 1936: 232; H.
McGhie, pers. comm. to RHC, 28 July 2002),
not found by us in BMNH (cf. Dance, 1986:
227). Distribution: Cuba (Paetel, 1873: 65,
1888: 481; Sowerby, 1909a: 353; Henderson,
1916: 322) [error; Alderson, 1925: 34]; Co-
lombia (Alderson, 1925: 34; Pain, 1964: 224).
Remarks. Alderson (1925: 31, 34) dis-
cussed the confused history of
misidentification of reflexa Swainson, 1823,
confusion that apparently continues, as it
was considered a synonym of paludosa Say,
1829, by Yong & Perera (1984: 121). Consid-
ered either a variety of flagellata Say, 1829, or
a distinct species by Alderson (1925: 34).
Retained here as a distinct species, following
Pain (1964: 224).

retusa

Ampullaria retusa Philippi, 1851: 18, pl. 5, fig.
1 [1852b: 21]. Guyana, namentlich der Rio
Rupunin, und Brasilien [? error]. Syntype:
ZMHB 1339 (M. Glaubrecht, pers. comm. to
RHC, 1 March 2003); type material possibly
also in MNHNS. Distribution: Brasil, Guyana
(Martens, 1857: 188, 1899: 424; Gaudion,
1879: 39) [? error].

Remarks. Name attributed to Olfers by
Philippi (1851: 18; 1852b: 21). Synonym of
flagellata Say, 1829, teste Pain (1964: 227).
However, flagellata Say, 1829, is a Central
American species, extending southwards
only into northern Colombia (Pain, 1964:
228), suggesting either that the localities
given for retusa Philippi, 1851, are incorrect
or that Pain was incorrect in synonymizing
the two species.
reyrei
Ampullaria Reyrei Cousin, 1887: 279, pl. 4,
fig. 7. Маро. Probable syntype: MNHN;
topotype: MCZ 92312. Distribution: Ecuador
(Sowerby, 1909a: 357).
robusta
Ampullaria robusta Philippi, 1852a: 50, pl. 15,
figs. 4, 5 [1852b: 27]. [No locality given.] Type
material: probably MNHNS. Distribution: un-
known.

COWIE & THIENGO

Remarks. Synonym of columellaris Gould,
1848, teste Alderson (1925: 54).

rugosa

Ampullaria rugosa Lamarck, 1801: 93. [No
locality given; “Mississipi” [? error] (Lamarck,
1822а: 177)]. Syntypes: the specimens illus-
trated in the works cited by Lamarck (1801:
93); possible syntype: MHNG 1093/93 (Y.
Finet, pers. comm. to RHC, 22 August 2002;
see also Mermod, 1952: 85). Distribution:
unknown.

Remarks. Synonym of urceus Müller, 1774,
teste Valenciennes (1833: 258), Gaudion
(1879: 41), Paetel (1888: 481), Sowerby
(1909a: 358), Alderson (1925: 10), Prashad
(1925: 72) and Mermod (1952: 86).

+ sanjosensis

Pomacea cumingii sanjosensis Morrison,
1946: 6, pl. 1, fig. 1. three small streams (not
of contiguous drainage) on the west side of
San Jose Island. Holotype: USNM 542136;
paratypes: ANSP 190947 (5 spms.), 215480
(3 spms.), 386773 (4 spms.), BMNH
1951.11.1.6-9 (4 spms.), MNCN 15.05/23733
(2 spms.) (Villena et al., 1997: 76), UF (1 lot,
3 spms.), USNM 598924, ZSM 20012076 (3
spms.) (E. Schwabe, pers. comm. to RHC,
28 July 2002), MCZ, UMMZ. Distribution:
Panama.

scalaris

Ampullaria scalaris Orbigny, 1835a: 31. Rio
Parana (republica Argentina) ... Guarayos
(republica Boliviana) ... provincia Santa-Cruz
de la Sierra (republica Boliviana). Syntypes:
BMNH 1854.12.4.333-4 (9 spms.), ММНМ (4
lots, 10 spms.), MHNG 33488 (1 spm.) (Y.
Finet, pers. comm. to RHC, 26 August 2002).
Distribution: Bolivia, Paraguay, Argentina,
Brasil, Uruguay (Paraguay-Parana drainage)
(Pain, 1960: 425).

scholvieni

Ampullaria scholvieni Kobelt, 1914b: 223, pl.
77, figs. 6, 7 [1914e: 178]. Puerto Cabello.
Holotype: ZMUH 15880 [destroyed; В.
Hausdorf, pers. comm. to RHC, 3 May 2002].
Distribution: Venezuela (Baker, 1930: 5).

Remarks. Synonym of chemnitzii Philippi,
1852, teste Baker (1930: 5) and Pain (1956a:
74).

semitecta

Ampullaria semitecta Mousson, 1873: 18.
nördlichen Süd-Amerika [in publication title].
Type material: location not known to us, not in
ZMZ (T. Meier, pers. comm. to RHC, 15 Au-
gust 2002), not found by us in MNHN (cf.

NEW WORLD AMPULLARIIDAE 75

Dance, 1986: 220). Distribution: Colombia,
Venezuela (Pain, 1956a: 75).

semperi

Ampullaria (? figulina var.) semperi Kobelt,
1914b: 221, pl. 77, figs. 2, 3 [1914e: 176].
[No locality given. “Fundort nicht genau
bekannt, doch sicher in Brasilien” (Kobelt,
1914e: 176)]. Type material: possibly
SMFD, ZMHB (Dance, 1986: 215), but not
found in ZMHB (M. Glaubrecht, pers.
comm. to RHC, 1 March 2003); possible
syntype(s): ZMUH fall ZMUH dry material
destroyed in the second world war; B.
Hausdorf, pers. comm. to RHC, 3 May
2002]. Distribution: ? Brasil.

Remarks. In the group of lineata Spix, 1827,
teste Kobelt (1914b: 221).
simplex
Ampullaria simplex Reeve, 1856d: pl. 21, fig.
98a, b. [No locality given.] Syntype: BMNH
20020682. Distribution: unknown.

Remarks. Synonym of lineata Spix, 1827,
teste Pain (1960: 422).
sordida
Ampullaria sordida Swainson, 1823a: pl. 143,
figs. 1, 2 [top and bottom figs.]. [No locality
given.] Type material: possibly MMUE
EM265907 (1 spm.) (H. McGhie, pers.
comm. to RHC, 29 July 2002), not found by
us in BMNH (cf. Dance, 1986: 227). Distribu-
tion: “Brésil - Rio-Janeiro - Plata” (Gaudion,
1879: 40), French Guiana (possibly intro-
duced; Tillier, 1980: 24).
spirata
Ampultaria [sic] spirata Deville & Huppé,
1850: 643. [No locality given.] Type material:
location not known to us. Distribution: un-
known.

Remarks. Name attributed to Orbigny. By
comparing it with Ampullaria aulanieri Deville
& Huppé, 1850, sufficient description was
provided to make the name available. Junior
primary homonym of Ampullaria spirata
Lamarck, 1804, which is now placed in family
Naticidae (see also Lamarck, 1822b: 549).
Not listed by Sherborn (1922-1933) or Ruhoff
(1980: 504).
sprucei
Ampullaria Sprucei Reeve, 1856e: pl. 28, figs.
134a, b. Tarapoto, east side of the Andes.
Syntypes: BMNH 20020684 (2 spms.);
topotype: ANSP. Distribution: Peru (Paetel,
1888: 481).

Remarks. Synonym of columellaris Gould,
1848, teste Alderson (1925: 54).

strebeli

Ampullaria malleata var. Strebeli Fischer &
Crosse, 1890: 235. Misantla, provinciæ Vera
Cruz. Syntypes: ZMHB 23203 (1 spm.; ? =
Strebel, 1873, pl. 3a, fig. 13a) (M. Glaubrecht,
pers. comm. to RHC, 1 March 2003; F.
Köhler, pers. comm. to RHC, 6 March 2003),
location of the 5 other spms. listed by Strebel
(1873: 26) not known to us. Distribution: East
Mexico (Martens 1899: 415).

Remarks. Described by bibliographic refer-
ence to Strebel (1873: 25, pl. 3, fig. 13, pl. 3a,
fig. 13a, b). Synonym of flagellata Say, 1829,
teste Pain (1964: 227). Martens (1899: 415)
considered it a full species and gave more
detailed locality information.

superba

Ampullaria superba Marshall, 1926: 3, pl. 1,
fig. 9 [holotype]. Cienaga Totuma, Depart-
ment of Atlantico, United States of Columbia
[= Colombia]. Holotype: USNM 362863. Distri-
bution: Colombia (Pain, 1956a: 77).

swainsoni

Ampullaria Swainsoni Philippi, 1852a: 53, pl.
16, fig. 5. [No locality given; Brasil given by
Swainson (1831-1832, pl. 64)]. Holotype:
MMUE (Swainson, 1831-1832: pl. 64). Distri-
bution: Brasil, Guyana [error] (Baker, 1930: 3).

Remarks. Philippi (1852a: 53) explicitly
based his description on Swainson's (1831—
1832) figure of “Ampullaria fasciata var.” [not
fasciata Swainson, 1822; see swainsonii
Hupe, 1857], which he copied, and although
Swainson had given the locality as Brasil,
Philippi stated that the locality was unknown.
Baker (1930: 3) mistakenly gave the locality
as Demerara [Guyana], which Swainson
(1831-1832: pl. 64) had mentioned but in ref-
erence to other specimens. See also
Swainson (1822c: 12 [Appendix]). Synonym
of lineata Spix, 1827, teste Sowerby (1909a:
354), but treated here as a distinct species,
following Baker (1930: 3). See also
hanleyana Alderson, 1926.

swainsonii

Ampullaria swainsonii Hupe, 1857: 66. Brasil.
Holotype: the shell illustrated by Swainson
(1821-1822a: pl. 103, fig. 2), possibly MMUE
(H. McGhie, pers. comm. to RHC, 28 July
2002), not found by us in BMNH (cf. Dance,
1986: 227). Distribution: Brasil.

Remarks. Introduced as a new name for
fasciata Swainson, 1822, which Hupe con-
sidered preoccupied by Lamarck, 1816 [also
Roissy, 1805]. However, fasciata Swainson,

76

1822 (see Swainson, 1821-1822a: pl. 103) is
a misidentification of fasciata Roissy, 1805,
so Hupé's citation of Swainson’s figure con-
stitutes the original description of this spe-
cies. Junior primary homonym of swainsoni
Philippi, 1852.

tenuissima

Ampullaria tenuissima Jousseaume, 1894:
120, text fig. La Coca, province d'Orient
(Equateur) [Ecuador]. Type material: not
found by us in MNHN (cf. Dance, 1986: 215).
Distribution: Ecuador (Sowerby, 1909a: 358).
testudinea

Ampullaria testudinea Reeve, 1856e: pl. 24,
fig. 114. [No locality given.] Syntype: BMNH
1900.2.13.20. Distribution: “Amazons”
(Sowerby, 1909a: 358); Brasil (Baker, 1914:
660).

Remarks. Synonym of lineata Spix, 1827,
teste Pain (1960: 422).
tristrami
Ampullaria tristrami Crosse & Fischer, in
Fischer & Crosse, 1890: 245. New name for
columbiensis Reeve, 1856, non Philippi,
1851. Distribution: Columbia; also “Pérou [ег-
ror] - Guatemala” (Gaudion, 1879: 26); also ?
Panama.

Remarks. Martens (1899: 413) considered
that tristrami Crosse & Fischer, 1890, referred
to the shell given to Tristram by Salvin, which
Tristram (1864: 414) had misidentified as
columbiensis Reeve, 1856. Martens argued,
therefore, that tristrami Crosse & Fischer,
1890, should not be accepted as a replace-
ment name for columbiensis Reeve, 1856, but
should stand as a valid name for Tristram’s
shell. However, the misidentification notwith-
standing, the nomenclatural act of Crosse &
Fischer was valid, even despite there already
being a new name for columbiensis Reeve,
1856 (1.е., martensiana Nevill, 1884). Pain
(1964: 228), placed Tristram’s “columbiensis”
in the synonymy of livescens Reeve, 1856.
Because columbiensis Reeve, 1856, is
treated here as a synonym of flagellata Say,
1829, tristrami Crosse & Fischer, 1890, is
also a synonym of flagellata Say, 1829. N.
syn.

+ unicolor

Ampullaria gigas Var. unicolor Philippi, 1852a:
47, pl. 10, fig. 2. [No locality given.] Type ma-
terial: probably MNHNS. Distribution: un-
known.

+ urabaensis

Pomacea cumingi urabaensis Pain, 1956a:

COWIE & THIENGO

75, text fig. (holotype). Golfo de Uraba, north-
ern Antioquia, Colombia. Holotype and three
paratypes (listed with dimensions by Pain
(1956a: 75) but without giving their location):
NMW.Z.1981.118.00114 (Pain collection, 3
spms. only); additional paratypes: MCZ (1 lot,
2 spms.). Distribution: Colombia.

Remarks. None of the NMW specimens is
large enough to be the holotype (H. Wood,
pers. comm. to RHC, 30 October 2001), the
location of which is therefore unknown.

urceus

Nerita urceus Muller, 1774: 174. in insulis
Indiae. Syntypes: the specimen figured by
Lister, as cited by Muller, and the specimen(s)
“In Museo Moltkiano” (Muller, 1774: 175), loca-
tion not Known to us, not in the Copenhagen
Museum (O. S. Tendahl, pers. comm. to
RHC, 18 April 2002). Distribution: Brasil, Peru,
Ecuador, Colombia, Venezuela, Guyana,
French Guiana, Trinidad (Pain, 1960: 426), ?
Surinam (Vernhout, 1914: 30) [? error;
Geijskes & Pain, 1957: 46, Tillier, 1980: 29],
Mexico [error] (Paetel, 1873: 65).

venetus

Ampullaria venetus Reeve, 1856b: pl. 4, fig.
17. [No locality given.] Syntypes: BMNH
20020686 (2 spms.). Distribution: Guatemala
(Paetel, 1888: 482).

Remarks. Synonym of flagellata Say, 1829,
teste Pain (1964: 227).

vermiformis

Ampullaria vermiformis Reeve, 1856b: pl. 12,
fig. 54. Paraguay. Syntype: BMNH 20020687.
Distribution: Paraguay.

Remarks. Synonym of canaliculata
Lamarck, 1822, teste Martens (1857: 210).
Synonym of gigas Эрих, 1827, teste Ihering
(1898: 49). Synonym of insularum Orbigny,
1935, teste Sowerby (1909a: 353; 1909b:
363):

vexillum

Ampullaria vexillum Reeve, 1856a: pl. 4, fig.
20. [No locality given; locality unknown
(Baker, 1930: 7; Pain, 1950b: 72).] Syntypes:
BMNH 20020688 (2 spms.). Distribution: Ven-
ezuela (Baker, 1930: 7).

Remarks. Synonym of puncticulata
Swainson, 1823, teste Sowerby (1909a: 356)
and Kobelt (1913e: 180), but retained here as
a valid species, following Pain (1950b: 72).

vickeryi

Pomacea vickeryi Pain, 1949a: 257; pl. 13,
figs. 1, 2. marsh near Buenos Aires, La Plata.
Holotype: BMNH 1946.10.2.3; paratypes:

NEW WORLD AMPULLARIIDAE 77

NMW.Z.1981.118.00107 (Pain collection, 2
spms.). Distribution: Argentina.

Remarks. Synonym of insularum Orbigny,
1935, teste Scott (1958: 295). Neither ofthe
two NMW specimens is large enough to be
the paratype for which Pain (1949a: 257)
gave measurements; presumably they are
two others of the total of 10 that were col-
lected (H. Wood, pers. comm. to RHC, 30
October 2001).
violacea
Ampullaria violacea Valenciennes, 1833: 260.
in sylvis Americæ. (Nova Hispania.). Lecto-
type (Fischer & Crosse, 1888: [explanation
of] pl. 46, fig. 4, 4a): MNHN. Distribution:
Mexico (Martens, 1899: 415; Sowerby,
1909a: 358).

Remarks. Synonym of flagellata Say, 1829,
teste Pain (1964: 226).
welwitschiana
Ampullaria Welwitschiana Drouët, 1859: 82,
pl. 3, figs. 33, 34. la rivière du Diamant, les
environs de Cayenne. Type material: not
found by us in MNHN, not mentioned by Tillier
(1980: 27-29). Distribution: French Guiana.

Remarks. Synonym of urceus Múller, 1774,
teste Tillier (1980: 27).
woodwardi
Ampullaria Woodwardi Dohrn, 1858: 134.
Ceylon [in publication title; error]. Lectotype
(Prashad, 1931: 168): BMNH 20020689. Dis-
tribution: South America (Prashad, 1931:
168).

Remarks. “Probably an abnormal and
somewhat eroded shell of Pomacea (Marisa)
cyclostoma” (Prashad, 1931: 168). This
statement does not definitively synonymize
woodwardi Dohrn, 1858, so it is retained here
as a valid species pending further research.
Prashad (1925: 85) could only find one shell
in the BMNH (as could we) and (Prashad,
1931: 168) mentioned “the unique type’,
hence designating that specimen the lecto-
type.

+ yatesii

Ampullaria Yatesii Reeve, 1856b: pl. 6, fig. 28.
River Магайоп. Type material: not found by
us in BMNH. Distribution: Peru (Pain, 1960:
427).

Remarks. Subspecies of urceus Müller,
1774, teste Pain (1960: 427). Boss &
Parodiz (1977: 111) implied that they are
synonyms, without formally synonymizing
them. The single specimen labeled as
yatesii Reeve, 1856, in the BMNH type col-

lection does not fit the hardened glue on the
board in its box, nor does it look like urceus
Müller, 1774, nor does it match Reeve’s fig-
ure. This specimen is therefore not yatesii
Reeve, 1856, the type material of which
must be considered lost.

yucatanensis

Ampullaria yucatanensis Crosse & Fischer,
1890: 110 [see also Fischer & Crosse (1890:
240, pl. 48, fig. 3, 3a)]. San Geronimo,
provinciae Yucatan dictae, reipublicae
Mexicanae. Type material: not found by us in
MNHN. Distribution: Mexico.

Remarks. Synonym of flagellata Say, 1829,
teste Pain (1964: 227).

yzabalensis

Ampullaria yucatanensis var. yzabalensis
Martens, 1899: 420, pl. 24, fig. 9. E. Guate-
mala: Lake of Yzabal. Syntypes: ZMHB 47109
(2 spms.) (M. Glaubrecht, pers. comm. to
RHC, 3 March 2002); type material not found
by us in BMNH or MCZ (cf. Dance, 1986:
218). Distribution: Guatemala.

Remarks. Synonym of flagellata Say, 1829,
teste Pain (1964: 227).

zeteki

Pomacea zeteki Morrison, 1946: 8, pl. 1, fig.
3. Chagres River near Gatuncilla, Republic of
Panama. Holotype: USNM 542137;
paratypes: ANSP 190941 (5 spms.), BMNH
1957. 1124. 10-15: (6 spms.), HUJ 2151511
spm.) (H. Mienis, pers. comm. to RHC, 4 Au-
gust 2002), USNM 542138, ZSM 20012055 (4
spms.) (E. Schwabe, pers. comm. to RHC,
28 July 2002). Distribution: Panama (Pain,
1956a: 76).

zischkai

Pomacea zischkai Blume & Pain, 1952: 267,
pl. 7. Chapara Region, at 400 m. Bolivia
tropica. Holotype: ZSM 20012062 (E.
Schwabe, pers. comm. to RHC, 28 July
2002); paratypes: ANSP 212117 (2 spms.),
FMNH 35467 (1 spm.), 38001 (3 spms.),
MHNG 33486 (1 spm.) (Y. Finet, pers.
comm. to RHC, 26 August 2002),
NMW.Z.1981.118.00117 (Pain collection, 2
spms.) (H. Wood, pers. comm. to RHC, 30
October 2001), ZMHB 98762 (2 spms.) (М.
Glaubrecht, pers. comm. to RHC, 1 March
2003), ZSM 20012050 (24 spms.) 20012051
(2 эрме.) 20012052 (10Т ‘spms-);
20012053 (155 spms.), 20012057 (9 spms),
20012058 (2 spms.) (E. Schwabe, pers.
comm. to RHC, 28 July 2002). Distribution:
Bolivia (Pain, 1960: 428).

78 COWIE & THIENGO

zonata
Ampullaria zonata Spix, in Wagner, 1827: 1,
pl. 2, fig. 1. in rivulis ... Provinciae Bahiensis.
Syntype: ZSM 20012056 (E. Schwabe, pers.
comm. to RHC, 28 July 2002; see also
Fechter, 1983: 221). Distribution: Columbia,
Brasil (Gaudion, 1879: 43; Sowerby, 1909a:

359).
Remarks. Authorship is given here as “Зрих,
in Wagner”, following Cowie et al. (in prep.).

Genus POMELLA Gray, 1847

POMELLA Gray, 1847: 148. Type species:
Ampullaria neritoides Orbigny, 1835 [=
megastoma Sowerby, 1825], by original des-
ignation.

Treated as a full genus, with two subgenera
(Pomella s. str., Surinamia) following Berthold
(1991: 24, 250).

Subgenus POMELLA Gray, 1847
Details as for genus Pomella Gray, 1847.

americanista
Ampullaria americanista lhering, 1919: 330,
[unnumbered text figure (“Cotipo”)]. Río
Paraná (Encarnación e Iguazú). Syntype:
MACN 8776a (Ihering, 1919: 331). Distribu-
tion: Argentina, Brasil, Paraguay (lhering,
1919: 335; Hylton Scott, 1958: 316).

Remarks. Placed in Pomella following

Hylton Scott (1958: 316) and S. C. Thiengo
(unpublished).

megastoma
Ampullaria megastoma Sowerby, 1825: 44
[name], x [description]. [No locality given.]
Holotype (“The only specimen ... that we
have seen”): not found by us in BMNH. Distri-
bution: Uruguay (Sowerby, 1909a: 359), Ar-
gentina (Ihering, 1919: 333).

neritoides
Ampullaria neritoides Orbigny, 1835a: 31. Rio
Uruguay (republica Uruguayensi orientali).
Syntypes: BMNH 1854.12.4.306-7 (4 spms.),
MNHN (2 lots, 3 spms.). Distribution: Uru-
guay.

Remarks. Synonym of megastoma

Sowerby, 1825, teste Pilsbry (1933: 74) and
Hylton Scott (1958: 314).

Subgenus SURINAMIA Clench, 1933

SURINAMIA Clench, 1933: 71. Type species:
Asolene (Surinamia) fairchildi Clench, 1933

[= sinamarina Bruguiére, 1792], by original
designation.

fairchildi

Asolene (Surinamia) fairchildi Clench, 1933:
71, pl. 7, figs. 1, 2. in the cataract of the
Surinam River below Kedjo, Dutch Guiana
(100 miles up river from Paramaribo). Holo-
type: MCZ 80515; paratypes: MCZ 80516,
ANSP 161782 (1 spm.), UMMZ (Clench,
1933: 72) [not found by us]. Distribution:
Surinam.

Remarks. Synonym of sinamarina
Bruguière, 1792, teste Geijskes 8 Pain
(1957: 46) and Tillier (1980: 17).

schrammi

-Ampullaria Schrammi Crosse, 1876: 102. in
flumine Oyapock, Guyanae Gallicae. Lecto-
type (Fischer-Piette, 1850: 150, pl. 5, fig. 81):
MNHN. Distribution: French Guiana.

Remarks. Synonym of sinamarina

Bruguière, 1792, teste Tillier (1980: 17).

sinamarina

Bulimus Sinamarinus Bruguiére, 1792: 342,
pl. 18, figs. 2, 3. la riviere de Sinamari dans la
Guyane francaise. Type material: not found
by us, nor by Tillier (1980: 17), in ММНМ. Dis-
tribution: Guyana, Surinam, French Guiana
(Vernhout, 1914: 43; Pain, 1950b: 73, 1952:
31; Geijskes 8 Pain, 1957: 46).

Remarks. Placed in Surinamia Clench,
1933, by Pain (1952: 31) and Tillier (1980: 17).

Incertae sedis in family AMPULLARIIDAE
Gray, 1824

The following species are not well enough
known to place them in a particular genus or in
some cases to definitively include or exclude
them as South American.

bilineata

Ampullaria bilineata Reeve, 1856e: pl. 23, fig.
110a, b. [No locality given.] Type material: not
found by us in BMNH. Distribution: unknown.

Remarks. Gaudion (1879: 24) gave
“Manille” [= Manila, Philippines] as the locality.
Placed in Pila Róding, 1798, by Sowerby
(1910: 56-57) and, confusingly, considered to
be “based on young shells” of globosa
Swainson (which is now placed in Pila
Róding, 1798) by Prashad (1925: 72; see
also Nevill, 1877: 2) but “certainly the same”
as gracilis Lea (also now placed in Pila
Röding, 1798) by Prashad (1925: 81). How-
ever, Alderson (1925: 30) treated it as a syn-
onym of buxea Reeve, 1856, considering it

NEW WORLD AMPULLARIIDAE 79

“nothing more than a stunted specimen of this
variety [buxea]”. Pilsbry (1927a: 247) treated
buxea Reeve, 1856, as a synonym of
fasciata Roissy, 1805 [Pomacea] and con-
sidered bilineata Reeve 1856, as a possible
synonym of fasciata Roissy, 1805.
equestris

Pila equestris Róding, 1798: 145. [No locality
given.] Type material: possibly Art and Natu-
ral History Museum, Gotha (Stewart, 1930:
35; Dance, 1986: 206). Distribution: un-
known.

Remarks. Not listed by Gaudion (1879) ог
Sowerby (1916: 71).
fasciata
Ampullaria fasciata Lamarck, 1816: 12 [liste],
pl. 457, fig. За, b [1822a: 177]. [No locality
given. “les rivières de l'Inde, des Moluques et
des Antilles” (Lamarck, 1822a: 177)]. Lecto-
type [“die von Lamarck citirte Figur der
Encyclopádie” (Philippi, 1852a: 53)]: MHNG
(Mermod, 1952: 88) [not MHNG 1093/92, teste
Y. Finet (pers. comm. to RHC, 22 August
2002).

Remarks. Junior primary homonym of
fasciata Roissy, 1805. Its correct placement
is unclear (e.g., Alderson, 1925: viii, 60) and
depends on further study. Mermod (1952: 87)
considered it probably a synonym of
ampullacea Linnaeus, 1758, which is now
placed in Pila Róding, 1798. Misidentified by
Swainson (1821-1822a: pl. 103); see
swainsonii Hupé, 1857.
gibbosa
Ampullaria gibbosa Paetel, 1887: 478. Un-
available name; nom. nud.

Remarks. Attributed to “Sw.” [= Swainson],
with reference to “Ad. Gen.” [= H. Adams & А.
Adams, 1853-1854], by Paetel (1887: 478).
However, it is not listed by H. Adams & A.
Adams (1853-1854), Sowerby (1916: 71),
Sherborn (1922-1933) or Ruhoff (1980: 288)
and appears never to have been made avail-
able. Paetel (1887: 480) listed “pachystoma
Benson” as a synonym, suggesting that the
species is Asian, since Benson worked in In-
dia (Naggs, 1997).
hepataria
Ampullaria hepataria Reeve, 1856c: pl. 17,
fig. 77. [No locality given.] Syntype: BMNH
20020661. Distribution: unknown.

Remarks. Synonym of corrugata Swainson
(which is now placed in Pila Róding, 1798),
teste Nevill (1884: 2). Listed as a "Western
Hemisphere” species and possibly a form of
hopetonensis Lea, 1834, by Sowerby (1909a:
351). However, Alderson (1925: 46) excluded

it from the synonymy of hopetonensis Lea,
1834 [= paludosa Say, 1829]. Also note the
skepticism of Pain (1964: 225) regarding
Sowerby's synonymies.

ignota

Pila ignota Róding, 1798: 146. [No locality
given.] Type material: possibly Art and Natu-
ral History Museum, Gotha (Stewart, 1930:
35; Dance, 1986: 206). Distribution: un-
known.

Remarks. Not listed by Gaudion (1879) or
Sowerby (1916: 71).

imperforata

Ampullaria imperforata Swainson, 1823b:
377. [No locality given.] Type material: possi-
bly MMUE (Dean, 1936: 232; H. McGhie,
pers. comm. to RHC, 28 July 2002), not
found by us in BMNH (cf. Dance, 1986: 227).
Distribution: unknown.

Remarks. Sowerby (1916: 70) was unable
to identify this species. Swainson (1823b:
377) said “operculum horny ?”, which would
suggest a New World species.

nucleus

Ampullaria nucleus Philippi, 1852a: 25, pl. 7,
fig. 1 [1852b: 23]. [No locality given.] Syntypes:
ZMHB 1374 (2 spms.; larger spm. = original
illustration) (M. Glaubrecht, pers. comm. to
RHC, 1 March 2003). Distribution: unknown.

Remarks. Considered close to crasssa
Swainson, 1823, and with a horny operculum
(Philippi, 1852a: 25), so included here as a
New World species.

obtusa

Ampullaria obtusa Deshayes, 1850: 45, pl.
72, fig. 24. [No locality given.] Type material:
not found by us in MNHN or BMNH (cf.
Dance, 1986: 210). Distribution: unknown.

pachystoma

Ampullaria pachystoma Paetel, 1887: 480.
Unavailable name; nom. nud.

Remarks. Attributed to Benson, but not
listed by Sowerby (1916: 72), Sherborn
(1922-1933) or Ruhoff (1980: 415) and ap-
pears never to have been made available.
Material not in Cambridge (R. C. Preece,
pers. comm. to RHC, 27 August 2002), not
found by us in BMNH. May be pachystoma
Philippi [Pomacea] or possibly an Asian spe-
cies (see gibbosa Paetel, 1887).

planorboides

Ampullaria planorboides Cristofori & Jan,
1832: [Section lla, Pars la] 7. Unavailable
name; nom. nud.

Remarks. Name attributed to Ziegler by
Cristofori & Jan (1832: 7), who gave “Austr.
N. Holl.” as the locality. Martens (1857: 208)

80

was unsure of the locality, saying “Botanybay
(?) Quid [= where]?”. Gaudion (1879: 37)
gave the locality as “Nouv. Hollande”. Not
listed by Sowerby (1916: 72). Material for-
merly in MCSN [destroyed; A. Garassino,
pers. comm. to RHC, 5 September 2002].
May not be an ampullariid.

rufilineata

Ampullaria rufilineata Reeve, 1856a: pl. 2, fig.
7. [No locality given.] Syntypes: BMNH
20020681 (3 spms.). Distribution: uncertain.

Remarks. Sowerby (1916: 69) placed this
species in Pila, giving “Pegu” as the locality
(the locality on the BMNH label). Martens
(1857: 209) and Gaudion (1879: 39), how-
ever, gave Venezuela as the locality. Blume &
Pain (1952: 267) considered the location and
generic placement uncertain.
sepulta
Pila sepulta Róding, 1798: 146. [No locality
given.] Type material: possibly Art and Natu-
ral History Museum, Gotha (Stewart, 1930:
35; Dance, 1986: 206). Distribution: un-
known.

Remarks. Not listed by Gaudion (1879) or
Sowerby (1916: 72).
tristis
Ampullaria tristis Gaudion, 1879: 41. Unavail-
able name; nom. nud.

Remarks. Name attributed to Say, al-
though Say appears never to have pub-
lished it (Binney, 1858: [237]). “Amérique
Septentrionale” given as locality. Possibly
Bulimus tristis Jay, 1839 [now placed in the
genus Lanistes Montford, 1810, which is Afri-
can].

trochulus

Ampullaria trochulus Reeve, 1856c: pl. 14,
fig. 66. [No locality given.] Syntype: BMNH
20020685. Distribution: unknown.

Remarks. Listed as a New World species
by Sowerby (1909a: 358).

Non-American species in family
AMPULLARIIDAE Gray, 1824

To prevent confusion, we list those non-

COWIE & THIENGO

Remarks. Nevill (1884: 10) gave the locality
as “South America”. Confirmed as African
(and hence to be placed in Pila Roding, 1798)
by Alderson (1925: 86).

aperta

Ampullaria aperta Philippi, 1849: 18. [No lo-
cality given.] Type material: probably MNHNS.

Remarks. Gaudion (1879: 24) and Paetel
(1873: 64, 1887: 476) gave the locality as
Venezuela. Considered Indian and placed in
Turbinicola Annandale & Prashad, 1921 [=
Pila, teste Berthold, 1991: 247], by Prashad
(1925: 88).

bruguieri

Ampullaria Bruguieri Deshayes, 1830a: 32.
Cayenne [?]. Syntypes: MNHN (3 spms.).

Remarks. One of the syntypes has a calci-
fied operculum and the label says “= A.
kordofana Parreyss” suggesting that it is a
species of Pila from Africa (see also Tillier,
1980: 16).

exigua

Ampullaria exigua Philippi, 1852a: 46, pl. 13,
fig. 4 [1852b: 26]. [No locality given.] Type
material: probably MNHNS.

Remarks. Sowerby (1909a: 349), following
Philippi (1852b: 27), first considered that it
“may be a variety of A. crassa, Swainson”
but subsequently (Sowerby, 1910: 58) treated
it as a species of Pila from Egypt. Listed
from Egypt by Paetel (1887: 478).

+ pallens

Ampullaria pallens Philippi, 1849: 17. Indiae
orientalis. Type material: probably MNHNS.

Remarks. Gaudion (1879: 36) gave Mexico
as the locality. Philippi (1852a: 32) had previ-
ously said “wahrscheinlich Ostindien” and
Martens (1901: 644) stated that it was from
the Philippines. Variety of virens Lamarck,
1822 (which is now placed in Pila Réding,
1798), teste Sowerby (1910: 62).

paludinoides

Ampullaria paludinoides Cristofori & Jan,
1832: [Section Па, Pars la] 7, [Mantissa] 3.
Am. mer. Type material: formerly MCSN [de-
stroyed; A. Garassino, pers. comm. to RHC,
5 September 2002].

American ampullariids that have at some time
been considered as American or possibly
American. The list may not be comprehensive.

Remarks. The locality was reiterated as
“America meridionalis” by Philippi (1852b: 24)
and “Amer. m.” by Paetel (1873: 65). Martens
(1857: 213) considered it African. Paetel
(1887: 480) listed it from “Moulmein” and it
was considered Indian by Nevill (1877: 7-9),
though perhaps based on misidentifications
by the previous authors he cited. Placed in

adusta
Ampullaria adusta Reeve, 1856a: pl. 3, fig.
11. [No locality given.] Type material: BMNH
20020690 [labeled “Zanzibar’].

NEW WORLD AMPULLARIIDAE 81

Pila Róding, 1798, Бу Sowerby (1910: 57,
62).
prunella

Ampullaria prunella Hupé, 1857: 67, pl. 12.
fig. 4, 4a. les parties centrales de l'Amérique
du Sud, de Rio de Janeiro a Lima, et de Lima
au Para [in publication title]. Syntypes: MNHN
(7 spms.; see also Tillier, 1980: 16); possible
syntype: ММНМ (1 spm.).

Remarks. The syntypes are labeled as
from “Cayenne” but have calcified opercula,
indicating that this is not an American spe-
cies (Tillier, 1980: 16).

rotundata
Ampullaria rotundata Say, 1829b: 245. St.
John's River in Florida. Type material: “not
found” (Baker, 1964: 168).

Remarks. Sowerby (1909a: 357) consid-
ered it “most likely a form of Hopetonensis”
[= paludosa Say, 1829], but it was subse-
quently synonymized with globosa Swainson
(which is now placed in Pila Róding, 1798) by
Pilsbry (1953: 60; see also Walker, 1918:
124; Clench, 1955: 107; Clench & Turner,
1956: 120). Spelled by Binney (1858: 147) as
“Ampluria rotundata”.

Unpublished names in family
AMPULLARIIDAE Gray, 1824

The following names of ampullariids, some of
them perhaps referring to American species,
have been found by us on museum collection
labels. They appear never to have been pub-
lished and are not nomenclaturally available.

“adjusta”. No author. Treated as a synonym of
sordida Swainson, 1823 in ANSP.

“burmeisteri”. Attributed to Ihering. ZMHB
109518 (M. Glaubrecht, pers. comm. to RHC,
1 March 2003).

“gualteriana”. No author. “nov. Pernambuco A.
fasciata Sw. var.?” on label in MNHN.

“miquitensis”. Attributed to Spix in UMMZ.

“palmieri”. Attributed to Preston in UMMZ.

“tacarigua”. Attributed to Pilsbry. ANSP 161137,
labeled “Holotype”.

“undata”. No author. USNM.

“unicolor”. Attributed to Martens in UMMZ.

“venezullum”. No author. UMMZ.

Non-ampullariids described originally
in family AMPULLARIIDAE Gray, 1824

The following taxa were described originally
in Ampullaria or Pomacea but are not now con-

sidered to belong to the Ampullariidae. Some
may be nomenclaturally unavailable. The list is
not comprehensive.

Ampullaria acuminata Lamarck, 1804
Ampullaria acuta Lamarck, 1804
Pomacea annularis Perry, 1811
Ampullaria avellana Lamarck, 1822
Ampullaria buccinoidea Young & Bird, 1828
Ampullaria bulimoides Deshayes, 1842
Ampullaria canaliculata Lamarck, 1804
Ampullaria canalifera Lamarck, 1822
Pomacea bibliana Marshall & Bowles, 1932
Ampullaria borealis Valenciennes, 1833
Ampullaria conica Lamarck, 1804
Ampullaria crassa Deshayes, 1830
Ampullaria crassatina Lamarck, 1804
Ampullaria depressa Lamarck, 1804
Ampullaria elongata Bennett, 1831
Ampullaria excavata Lamarck, 1804
Ampularia faujasii Serres, 1829
Ampullaria fragilis Lamarck, 1822
Ampullaria galloprovincialis Matheson, 1843
Ampullaria hybrida Lamarck, 1804
Ampullaria laevigata Deshayes, 1842
Pomacea linearis Perry, 1811

Ampullaria media Bennett, 1831
Ampullaria patula Lamarck, 1804
Ampullaria perovata Conrad, 1846
Ampullaria ponderosa Deshayes, 1825
Ampullaria proboscidea Matheson, 1843
Ampullaria pygmaea Lamarck, 1804
Ampullaria rosea Эрих, 1827

Ampullaria scalariformis Deshayes, 1825
Ampullaria sigaretina Lamarck, 1804
Ampullaria spirata Lamarck, 1804
Ampullaria tasmaniae Guillou, 1842
Pomacea variegata Perry, 1811

ACKNOWLEDGEMENTS

For information from the collections of their
various institutions, for assistance with obtain-
ing and dating literature, and for facilitating vis-
its to the collections in their care we thank
Paula Mikkelsen and Julia Sigwart (АММН),
Paul Callomon, George Davis, Mark Kitson and
Gary Rosenberg (ANSP), Peter Mordan, Fred
Naggs, Kathie Way and Richard Williams
(BMNH), Tim Pearce (СММН), О. $. Tendahl
(Copenhagen Museum), Rüdiger Bieler and
Jochen Gerber (FMNH), Henk Mienis (HUJ),
Gina Douglas (Linnean Society), Alessandro
Garassino (MCSN), Ken Boss and Silvard Kool
(MCZ), Yves Finet (MHNG), Laurence Cook

82 COWIE & THIENGO

and Henry McGhie (MMUE), Philippe Bouchet,
Virginie Héros, Pierre Lozouet and Philippe
Maestrati (MNHN), Sergio Letelier (MNHNS),
Luiz De Simone (MZUSP), Mary Seddon and
Harriet Wood (NMW), Fred Thompson (UF),
Jack Burch, Doug Eernisse, Daniel Graf and
Diarmaid О Foighil (UMMZ), Bob Hershler and
Richard Houbrick (USNM), Alfredo Castro-
Vazquez (Universidad Nacional de Cuyo, Ar-
gentina), Nestor Cazzaniga (Universidad
Nacional del Sur, Argentina), Roberto Cipriani
(Universidad Simon Bolivar, Venezuela),
Matthias Glaubrecht and Frank Kôhler (ZMHB),
Bernhard Hausdorf (ZMUH), Trudi Meier (ZMZ),

Michael Schródl and Enrico Schwabe (ZSM).

We also thank Matthias Glaubrecht, Peter
Mordan and Dick Petit for detailed reviews of
the manuscript, Eugene Coan, Neal Evenhuis,
Jochen Gerber, Riccardo Giannuzzi-Savelli,
Matthias Glaubrecht, Alan Kabat, Alan Kohn,
Dick Petit, Andrew Rindsberg, Barry Roth,
Michael Schródl, Enrico Schwabe and Fred
Thompson for additional assistance and dis-
cussion, and Rudiger Bieler for help with Ger-
man translation.

The United States Department of Agriculture
Special Grants Program for Tropical and Sub-
tropical Agriculture provided partial funding.

LITERATURE CITED

We have seen all the references listed, thereby
ensuring accuracy of citation. Dates of publication
in some cases have been taken from Evenhuis &
Cowie (1995) and Cowie (1998); other dates de-
rive from subsequent research. Citation is given
verbatim, unless a publication represents a pre-
sentation made at a meeting, in which case it can-
not be cited verbatim and a paraphrased title is
provided and placed in square brackets. The date
of publication, as accurately as could be ascer-
tained from the publication itself and from outside
sources, is placed in square brackets at the end of
the citation. The dates recorded here are the ear-
liest found for each citation. If the year of publica-
tion was different from that printed in the
publication itself, the actual year of publication is
placed in square brackets. In many instances of
works published in parts (Lieferungen, livraisons,
etc.), the original wrappers have not been seen,
only the complete bound work. Dating has then
been obtained from other, secondary sources.
The dates that were printed on the original wrap-

pers have therefore not always been verified. The
year(s) of publication of the entire work, if differ-
ent from that which actually appeared in the work
(usually on the frontispiece) are therefore not
placed in square brackets, pending further re-
search on the original wrappers. If no date other
than year could be found, the publication date
must be treated as 31 December until such time
as evidence of earlier publication is discovered.
Sources for dates listed here are held by the first
author. When an author published more than
one paper in a year, the papers are listed chrono-
logically and the year given a letter suffix corre-
sponding to the citation in the catalog. Where
tabular collation is given for publications issued in
parts, the date letter for each part is given in the
“Date of publication” column. An author’s initials
are placed in square brackets if not given in the
publication. Publications of the International Com-
mission on Zoological Nomenclature are cited with
authorship as “ICZN” in the catalog but spelled out
in this bibliography.

ABBOTT, К. T., 1955, The Titian К. Peale shell collection. The Nautilus, 68(4): 123-126, pl. 4. [28 April]

ACOSTA, В. О. & Б. $. V. PULLIN, 1991, Environmental impact of the golden snail (Pomacea sp.) on
rice farming systems in the Philippines. Freshwater Aquaculture Center, Central Luzon State
University, Munoz, Nueva Ecija; ICLARM, Manila. vi + 34 pp.

ADAMS, Н. & A. ADAMS, 1853-1854, The genera of Recent Mollusca; arranged according to their

organization. Vol. I. J. Van Voorst, London.

Part Plates Pages Date of publication
1 1-4 1=32 January 1853a
2 5-8 33-64 February 1853b
3 9-12 65-96 June 1853c
4 13-16 97-128 August 1853d
5 17-20 129-160 September 1853е
6 21-24 161-192 October 1853f
7 25-28 193-224 November 185349
8 29-32 225-256 December 1853h
9 33-36 257-288 January 1854a
10 37-40 289-320 February 1854b
11 41-44 321-352 March 1854c
12 45-48 353-384 April 1854d
13 49-52 385-416 May 1854e
14 54-56 417-448 June 1854f
15 57-60 449-484 July 1854g

NEW WORLD AMPULLARIIDAE 83

ALDERSON, Е. G., 1925, Studies т Ampullaria. Heffer, Cambridge. хх + 102 pp., 19 pls.

ALDERSON, E. G., 1926, The Ampullaria swainsoni of Philippi, Hanley, and Reeve. Proceedings of the
Malacological Society of London, 17(1): 42-43. [30 April]

ANDERSON, B., 1993, The Philippine snail disaster. The Ecologist, 23: 70-72.

ANDERSON, Е. M., 1928, Notes on lower Tertiary deposits of Colombia and their molluscan and
foraminiferal fauna. Proceedings of the California Academy of Sciences, (4)17(1): 1-[29]. [22 June]

ANTON, H. E., 1838, Verzeichniss der Conchylien welche sich in der Sammlung von Hermann Eduard
Anton befinden. Eduard Anton, Halle. xvi + 110 pp.

“Im Druck beendigt den 9 October 1838” appears on p. 110, which we translate as “Printing completed ... *,
although “1839” appears on the title page. Cernohorsky (1978: 299) showed that this publication was indeed
distributed in 1838.

APPUN, С. F., 1871, Unter den Tropen. Wanderungen durch Venezuela, am Orinoco, durch Britisch
Guyana und am Amazonenstrome in den Jahren 1849-1868. Erster Band. Venezuela. Hermann
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ARIAS, S., 1952, Algunos moluscos de la region Baruta-El Hatillo. Memoria de la Sociedad de
Ciencias Naturales La Salle, 12: 47-65.

BAKER, F., 1914, The land and fresh-water mollusks of the Stanford Expedition to Brazil. Proceedings
of the Academy of Natural Sciences of Philadelphia, 65[1913]: 618-672, pls. 21-27. [28 January]

BAKER, H. B., 1922, The Mollusca collected by the University of Michigan — Walker expedition in
southern Vera Cruz, Mexico. |. Occasional Papers of the Museum of Zoology, University of
Michigan, 106: 1-95. [18 February]

BAKER, H. B., 1930, The Mollusca collected by the University of Michigan — Williamson expedition in
Venezuela. Part VI. Occasional Papers of the Museum of Zoology, University of Michigan, 210: 1—
95. [14 February]

BAKER, H. B., 1964, Type land snails in the Academy of Natural Sciences of Philadelphia. Part Ill.
Limnophile and thalassophile Pulmonata. Part IV. Land and fresh-water Prosobranchia. Proceedings
of the Academy of Natural Sciences of Philadelphia, 116(4): 149-193. [20 October]

BARBOSA RODRIGUES, J., 1892, Les reptiles de la vallée de l'Amazone. Vellosia, 2: 41-58.

BERTHOLD, T., 1989, Comparative conchology and functional morphology of the copulatory organ of
the Ampullariidae (Gastropoda, Monotocardia) and their bearing upon phylogeny and
palaeontology. Abhandlungen des Natunwissenschaftlichen Vereins in Hamburg, (NF) 28: 141-164.

BERTHOLD, T., 1991, Vergleichende Anatomie, Phylogenie und historische Biogeographie der
Ampullariidae (Mollusca, Gastropoda). Abhandlungen des Naturwissenschaftlichen Vereins т
Hamburg, (NF) 29: 1-256.

BIELER, R., 1992, Gastropod phylogeny and systematics. Annual Review of Ecology and
Systematics, 23: 311-338.

BIELER, R., 1993, Ampullariid phylogeny - Book review and cladistic re-analysis. The Veliger, 36:
291-299.

BINNEY, W. G., ed., 1858, The complete writings of Thomas Say, on the conchology of the United
States. H. Bailliere, New York. vi + 252 pp., 75 pls.

BINNEY, W. G., 1865, Land and fresh-water shells of North America. Part Ill. Ampullariidae, Valvatidae,
Viviparidae, fresh-water Rissoidae, Cyclophoridae, Truncatellidae, fresh-water Neritidae,
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BLUME, W., 1957, Eine bis heute unbekannte Interart von Pomacea bridgesi Rve. Opuscula
Zoologica, 1: 1-2. [15 Мау]

BLUME, W. 8 Т. РАМ, 1952, A new species of Pomacea from Bolivia. Journal of Conchology, 23(8):
267-268, pl. 7.

BOSS, К. J. & J. J. PARODIZ, 1977, Paleospecies of Neotropical ampullariids and notes on other fossil
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Invertebrate Zoology, American Museum of Natural History. V. Mollusca, part 2 (Class Gastropoda
[exclusive of Opisthobranchia and Pulmonata], with supplements to Gastropoda [Opisthobranchia],
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canaliculata. Топики Journal of Agricultural Research, 40: 81-89.

BRUGUIERE, J. G., 1792, Sur une nouvelle espèce de Вийте. Journal d'Histoire Naturelle, 1(9): 339-
343, pl: 18, figs. 2, 3.

BURKY, А. J., 1974, Growth and biomass production of an amphibious snail, Pomacea urceus (Müller),
from the Venezuelan savannah. Proceedings of the Malacological Society of London, 41: 127-143.

BUSCH, [G.] von dem, 1859, On some new freshwater shells from Ecuador and New Granada, in the
collection of Hugh Cuming, Esq. Proceedings of the Zoological Society of London, 1859(2): 167—
169. [between July and October]

CAZZANIGA, N. J., 1987, Pomacea canaliculata (Lamarck, 1801) en Catamarca (Argentina) y un
comentario sobre Ampullaria catamarcensis Sowerby, 1874 (Gastropoda, Ampullariidae). /heringía,
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84 COWIE & THIENGO

CAZZANIGA, N. J., 2002, Old species and new concepts in the taxonomy of Pomacea (Gastropoda:
Ampullariidae). Biocell, 26(1): 71-81.

CERNOHORSKY, W. O., 1978, The date of publication of Anton's “Verzeichniss der Conchylien”. The
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СНЕМС, E. Y., 1989, Control strategy for the introduced snail Pomacea lineata, in rice paddy. Pp. 69-
75, in: I. Е HENDERSON, ed., Slugs and Snails in World Agriculture, British Crop Protection Council
Monograph 41.

CLENCH, W. J., 1933, Surinamia, a new ampullariid from Dutch Guiana. The Nautilus, 47(2): 71-72.
[1 November]

CLENCH, W. J., 1955, Melania cancellata Say. The Nautilus, 68: 107. [11 February]

CLENCH, W. J. & R. D. TURNER, 1956, Freshwater mollusks of Alabama, Georgia, and Florida from
the Escambia to the Suwannee River. Bulletin of the Florida State Museum, Biological Sciences,

1(3): 99-239. [3 October]

COUSIN, A., 1887, Faune malacologique de la République а l’Equateur. Bulletin de la Société
Zoologique de France, 12: 187-287, pl. 4.

COWAN, С. F., 1969, Notes on Griffith's Animal Kingdom of Cuvier (1824-1835). Journal of the
Society for the Bibliography of Natural History, 5(2): 137-140. [April issue]

COWIE, R. H., 1995, Identity, distribution and impacts of introduced Ampullariidae and Viviparidae in
the Hawaiian Islands. Journal of Medical and Applied Malacology, 5[1993]: 61-67.

COWIE, R. H., 1997a, Pila Róding, 1798 and Pomacea Perry, 1810 (Mollusca, Gastropoda):
proposed placement on the Official List, and Ampullariidae Gray, 1824: proposed confirmation as
the nomenclaturally valid synonym of Pilidae Preston, 1915. Bulletin of Zoological Nomenclature,
54(2): 83-88. [March]

COWIE, R. H., 1997b, Catalog and bibliography of the nonindigenous nonmarine snails and slugs of
the Hawaiian Islands. Bishop Museum Occasional Papers, 50: 1-66. [25 February]

COWIE, R. H., 1998, Catalog of the nonmarine snails and slugs of the Samoan Islands. Bishop
Museum Bulletin in Zoology 3. Bishop Museum Press, Honolulu. viii + 122 pp. [13 January]

COWIE, R. H., 2002, Apple snails (Ampullariidae) as agricultural pests: their biology, impacts and
management. Pp. 145-192, in: С. М. BARKER, ed., Molluscs as Crop Pests. CABI Publishing,
Wallingford.

COWIE, R. H., М. J. CAZZANIGA 8 М. GLAUBRECHT, in prep., The South American Mollusca of
Johann Baptist Ritter von Spix and their publication by Johann Andreas Wagner.

COWIE, К. H., А. К. KABAT & М. L. EVENHUIS, 2001, Ampullaria canaliculata Lamarck, 1822 (currently
Pomacea canaliculata; Mollusca, Gastropoda): proposed conservation of the specific name. Bulletin
of Zoological Nomenclature, 58(1): 13-18. [31 March]

COX, |. R., 1942, Publication dates of Traité élémentaire de conchyliologie, by С.Р. Deshayes.
Proceedings of the Malacologial Society of London, 25(3): 94-95. [20 December]

CRISTOFORI, J. de & С. JAN, 1832, Catalogus т IV. sectiones divisus rerum naturalium in museo
exstantium Josephi de Cristofori et Georgii Jan plurium Acad. Scient. et Societ. Nat. Cur. Sodalium
complectens adumbrationem oryctognosiae et geognosiae atque prodromum fauna et florae Italiae
superioris. Sectio Il. Pars I. Carmignani, Parmae. iv + 8 + [1] + 4 + 4 pp. [April]

CROSSE, H., 1871, Des espèces terrestres et fluviatilies quellon a considérées, à tort, comme
appartenant á la faune malacologique de la Nouvelle-Calédonie. Journal de Conchyliologie, 19(3):
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CROSSE, H., 1876, Diagnosis Ampullariae novae, Guyanae Gallicae incolae. Journal de
Conchyliologie, 24(1): 102. [1 March]

CROSSE, H., 1891, Description d'un Ampullaria nouveau de l'Amazone. Journal de Conchyliologie,
39(2): 214- 216, pl. 4, fig. 2. [20 August]

CROSSE, H. 8 P. FISCHER, 1890, Diagnoses Ampullariarum novarum Guatemalae et reipublicae
Mexicanae incolarum. Journal de Conchyliologie, 38(2): 110-114. [23 September]

CUVIER, [G. |. С. Е. D.], [1816], Le règne animal distribué d'après son organisation, pour servir de
base a l'histoire naturelle des animaux et d'introduction a l'anatomie comparée. Avec figures,
desinées d'après nature. Tome Il, contenant les reptiles, les poissons, les mollusques et les
annelides. Deterville, Paris. xviii + 532 рр. [2 December]

DALL, W. Н., 1898, Description of a new Ampullaria from Florida. The Nautilus, 12(7): 75-76. [7 November]

DALL, W. H., 1904, Notes on the genus Ampullaria. Journal of Conchology, 11(2): 50-55.

DALL, W. H., 1919, A new form of Ampullaria. The Nautilus, 33(1): 10-11. [16 July]

DALL, W. H., 1921, Two new South American shells. The Nautilus, 34(4): 132-133. [5 May]

DANCE, S. P., 1967, Report on the Linnaean shell collection. Proceedings of the Linnean Society of
London, 178(1): 1-24, pls. 1-10. [January issue]

DANCE, $. P., 1986, A history of shell collecting. E.J. Brill/Dr. W. Backhuys, Leiden. xv + 265 pp., 32 pls.

DAUTZENBERG, P., [1902], Descriptions de coquilles nouvelles rapportées du Pérou par М. Baer.
Journal de Conchyliologie, 49(4): 306-313. [28 April]

DEAN, J. D., 1936, Conchological cabinets of the e century. Journal of Conchology, 20(8): 225-
2921 July]

NEW WORLD AMPULLARIIDAE 85

DeKAY, J. E., 1843, Zoology of New York, or the New-York fauna; comprising detailed descriptions of
all the animals hitherto observed within the State of New York; with brief notices of those occasion-
ally found near its borders: and accompanied by appropriate illustrations. Part V. Mollusca. Carroll
and Cook, Albany. [xiv] + 271 pp., 40 pls.

DESHAYES, G. P., 1830-1832, Encyclopédie méthodique. Histoire naturelle des vers. Tome second.
Agasse, Paris. vii + 256 + 594 pp.

Published in parts. Dates of publication as follows (from Evenhuis, 2003):

Part Livraison Pages Date of publication
1 101 i-vil, 1-256 1 February 1830a
2 101 1-144 1 February 1830b
2 102 [part] 145-594 29 September 1832

Livraison 102 also included volume 3, with pages 595-1152.

DESHAYES, С. P., 1838, Tome huitième. Mollusques. Pp. 1-660, in: С. Р. DESHAYES & H. MILNE
EDWARDS, Histoire naturelle des animaux sans vertébres, présentant les caracteres généraux et
particuliers de ces animaux, leur distribution, leurs classes, leurs familles, leurs genres, et la citation
des principales especes qui s'y rapportent; précédée d'une introduction offrant la détermination des
caracteres essentiels de l'animal, sa distinction du végétal et des autres corps naturels; enfin,
l'exposition des principes fondamentaux de la zoologie. Deuxième édition. Revue et augmentée de
notes présentant les faits nouveaux dont la science s'est enrichie jusqu'a ce jour. J.B. Baillière,
Paris. [June]

DESHAYES, G. P., 1839-1857, Traité élémentaire de conchyliologie avec les applications de cette
science a la Géologie. Explication des planches. Victor Masson, Paris. 80 + xi [Appendice] pp., 132

pls.
Published in parts. Dates of publication as follows (Cox, 1942: 95):

Pages Date of publication
1-24 1839
25-48, Appendice i-iv 1850
49-80 1853
Appendice v-xi 1857

DEVILLE, Е. & [L.] H. НУРРЕ, 1850, Description de quelques coquilles nouvelles provenant de
l'expedition de M. de Castelnau. Revue et Magasin de Zoologie Pure et Appliquée, (2) 2: 638-644.
[December issue]

DOHRN, H., 1858, Descriptions of new species of land and freshwater shells collected in Ceylon,
from the collection of H. Cuming, Esq. Proceedings of the Zoological Society of London, 26: 133-
135. [12 July]

DROUET, H., 1859, Essai sur les mollusques terrestres et fluviatile de la Guyane Française. J.-B.
Baillière, Paris. 116 pp., 4 pls.

DUNKER, W. [B. R. H.], 1845, Vorláufige Diagnosen mehrerer neuer Conchylien aus der
norddeutschen Liasbildung, die náchstens ausführlicher beschreiben und abgebildet erscheinen
werden. Zeitschrift für Malakozoologie, 1(December 1844 issue): 186-188. [January]

DUNKER, W. [В. К. H.], 1853, Ampullaria eximia. Zeitschrift für Malakozoologie, 10(6): 93-95.

EVENHUIS, М. L., 2003, Dating and publication of the Encyclopédie Méthodique (1782-1832), with
special reference to the parts of the Histoire Naturelle and details on the Histoire Naturelle des
Insectes. Zootaxa 166: 1-48.

EVENHUIS, М. L. & R. Н. COWIE, 1995, Bibliography. Pp. 205-35, in: В.Н. COWIE, NL. EVENHUIS & С.С.
CHRISTENSEN, Catalog of the native land and freshwater molluscs of the Hawaiian islands.
Backhuys Publishers, Leiden. [3 June]

FARACO, F., I. Е. VEITENHEIMER-MENDES 8 E. BORGES, 2002. Felipponea neritiniformis
(Gastropoda, Ampullariidae): concha, rádula, complexo peniano e comportamento reprodutivo.
Biociéncias 10(2): 65-78.

FARFANTE, I. P., 1942, Moluscos de la region de Сатоа y Somorrostro y sus condiciones de vida.
Memorias de la Sociedad Cubana de Historia Natural, 16(1): 45-56. [30 May]

FECHTER, R., 1983, Liste des Typenmaterials der von J. B. v. Spix in Brasilien gesammelten
Gastropoda. Spixiana, Supplement 9: 221-223. [15 December]

FERUSSAC, [A. E. J. P. J. F. d'A. de], 1827, Suite du catalogue des espèces de mollusques terrestres
et fluviatiles, recueillies par M. Rang, dans un voyage aux Grandes-Indes. Bulletin des Sciences
Naturelles et de Géologie 10: 408-413.

FISCHER, Р. & Н. CROSSE, [1870-1902], Recherches zoologiques pour servir à l'histoire de la faune
de l'Amérique Centrale et du Mexique, publiées sous la direction de М. Мите Edwards, membre de

86 COWIE & ТНЕМСО

l'Institut. Septième partie. Études sur les mollusques terrestres et fluviatiles du Mexique et du
Guatemala. Imprimerie Nationale, Paris. 702 + 731 pp., 72 pls.

Published in livraisons containing feuilles, as follows:

Volume Livraison Feuilles Pages Plates Date of publication

1 1 1-19 1-152 1-6 1870
2 20-38 153-304 7-12 1872
3 39-48 305-384 13-16 1873a
4 49-58 385-464 17-20 1873b
5 59-68 465-546 21-24 1875
6 69-78 457-624 25-28 1877
7 79-88 625-702 29-31 1878

2 8 1-10 1-80 32-36 1880
9 11-16 81-128 37-42 1886
10 17-22 129-176 43-46 1888
11 23-32 177-256 47-48 1890
12 33-39 257-312 49-52 1891
15 40-49 313-392 53-54 1892
14 50-61 393-488 55-58 1893
15 62-72 489-576 59-62 1894a
16 13-82 577-655 63-66 1894b
17 83-92 657-731 67-72 1902

FISCHER-PIETTE, E., 1950, Liste des types décrits dans le Journal de Conchyliologie et conservés
dans la collection de ce journal. Journal de Conchyliologie, 90: 8-23, 65-82, 149-180, pls. 1-5.
FUJIO, Y., Н. KURIHARA & Е. von BRAND, 1991, Differences metric traits among three strains of apple

snail, Pomacea canalicualta. Топики Journal of Agricultural Research, 41(3-4): 61-68.

GAUDION, H., 1879, Liste alphabétique des espèces du genre Ampullaria de Lamarck. Bulletin de la
Société d'Étude des Sciences Naturelles de Béziers, 4: 20—43.

GEIJSKES, D. C. 4 T. PAIN, 1957, Suriname freshwater snails of the genus Pomacea. Studies on the
fauna of Suriname and other Guyanas, 1(3): 41-48, pls. 9, 10.

GMELIN, J. F., 1791, Caroli a Linné. Systema naturae per regna ша пашгае secundum classes,
ordines, genera, species, cum caracteribus, differentiis, synonymis, locis. Editio decima tertia,
aucta, reformata. Tom. 1. Pars VI. Georg Emanuel Beer, Lipsiae [= Leipzig]. pp. 3021-[3910]. [14
May]

GOULD, [A. A.], 1848, [Descriptions of shells from the collection of the Exploring Expedition].
Proceedings of the Boston Society of Natural History, 3: 73-75. [November]

GRAY, J. E., 1824, Zoological notices. The Philosophical Magazine and Journal, 63(311): 274-277.
[30 April]

GRAY, J. E., 1847, Alist of the genera of Recent Mollusca, their synonyma and types. Proceedings of
the Zoological Society of London, 15: 129-219. [November]

GRAY, J. E., 1854, List of the shells of Cuba in the collection of the British Museum, collected by М.
Ramon de la Sagra. Described by Prof. Alcide d'Orbigny, in the “Histoire de l'Ile de Cuba.”. British
Museum, London. [i] + 48 pp. [9 December]

GRAY, J. E., 1855, List of the shells of South America in the collection of the British Museum. Collected
and described by М. Alcide d’Orbigny, in the “Voyage dans l'Amérique Méridionale.”. British
Museum, London. [i] + 89 pp. [13 January]

GRIFFITH, E. 8 E. PIDGEON, [1833]-1834, The Mollusca and Radiata. Arranged by the Baron
Cuvier, with supplementary additions to each order. Whittaker and Co., London. vii + 601 pp.

Published as volume 12 of Griffith and others’ translation of Cuvier's Le règne Animal ..., in three parts, tentatively
dated by Cowan (1969: 139) as follows:

Part Date of publication

38 December 1833

39 March 1834a

40 June 1834b

GUERRERO), L., 1991, The biology of golden snail in relation to Philippine conditions. Pp. 10-11, in:
B. O. ACOSTA 8 R. S. V. PULLIN, eds., Environmental impact of the golden snail (Pomacea sp.) on rice
farming systems in the Philippines. Freshwater Aquaculture Center, Central Luzon State University,
Munoz, Nueva Ecija; ICLARM, Manila.

GUILDING, L., 1828, Observations on the zoology of the Caribaean Islands. The Zoological Journal,
3: 527-544. [after April]

GUPPY, R. J. L., 1864, Descriptions of new species of fluviatile and terrestrial operculate Mollusca
from Trinidad. Annals and Magazine of Natural History, (3) 14(82): 243-248. [issue for October]

NEW WORLD AMPULLARIIDAE 87

GUPPY, R. J. L., 1866, On the terrestrial and fluviatile Mollusca of Trinidad. Annals and Magazine of
Natural History, (3) 17(97): 42-56. [issue for January]

HANLEY, $. [C. T.], 1854-1858, The Conchological Miscellany. Illustrative of Pandora, Amphidesma,
Ostrea, Melo, the Melaniadae, Ampullaria and Cyclostoma. Williams and Norgate, London and
Edinburgh. 12 pp., 40 pls.

The plates treating Ampullaria were published in November 1854.

HENDARSIH, S., $. SURIAPERMANA, А. FAGI & |. MANWAN, 1994, Potential of fish in rice-fish culture
as a biological control agent of rice pests. Pp. 32-33, in: С.В. DELA CRUZ, ed., Role of fish т
enhancing ricefield ecology and in integrated pest management. Agency for Agricultural Research
and Development, Bogor; ICLARM, Manila.

HENDERSON, J. B., 1916, A list of the land and fresh-water shells of the Isle of Pines. Annals of the
Carnegie Museum, 10(3-4): 315-324. [July]

HERRMANNSEN, А. N., 1846-1849, Indicis generum malacozoorum primordia. Nomina subgenerum,
generum, familiarum, tribuum, ordinum, classium; adjectis auctoribus, temporibus, locis systematicis
atque literariis, etymis, synonymis. Praetermittuntur cirripedia, Tunicata et Rhizopoda. Vol. |. Т.
Fischer, Cassellis [= Cassel]. xxvii + 637 pp.

Published in Lieferungen, as follows (data from the Supplementa et corrigenda associated with this work; R. E.
Petit, pers. comm. to RHC, 22 February 2003):

Lieferung Pages Date of publication
1 I-xxvil, 1-104 1 September 1846a
2 105-232 1 December 18465
3 233-360 1 March 1847a
4 261-488 18 April 1847b
5 489-616 25 May 1847c
6 (part) 617-637 17 July 1847d

HIDALGO, J. G., 1866, Description d'espèces nouvelles de la République de l'Equateur. Journal de
Conchyliologie, 14(4): 343-344. [7 October]

HIDALGO, J. G., 1871, Description d'un Ampullaria nouveau, provenant du fleuve des Amazones.
Journal de Conchyliologie, 19(3): 206-207. [27 September]

HIDALGO, J. G., 1872, Description d'espèces nouvelles. Journal de Conchyliologie, 20(2): 142-144,
pl. 7, figs. 1, 2. [6 May]

HINKLEY, А. A., 1920, Guatemala Mollusca. The Nautilus, 34(2): 37-55. [6 November]

Plate 4, with one figure (Fig. 5) associated with this article, was published in The Nautilus 34(3) on 11 January 1921.

HUPÉ, [L.] H., 1857, Animaux nouveaus ou rares recueillis pendant l'expédition dans les parties
centrales de l'Amérique du sud, de Rio de Janeiro a Lima, et de Lima au Para; exécutée par ordre
du gouvernement français pendent les années 1843 a 1847, sous la direction du comte Francis de
Castelnau. Mollusques. P. Bertrand, Paris. 96 pp., 20 pls. [There are two spellings of this author's
name; see Deville & Huppé (1850)]

HYLTON SCOTT, М. 1., 1948, Moluscos del noroeste Argentino. Acta Zoologica Lilloana, 6: 241-274,
1 pl. [before 29 December]

HYLTON SCOTT, М. 1., 1958, Estudio morfologico y taxonomico de los ampullaridos de la Republica
Argentina. Revista del Museo Argentino de Ciencias Naturales “Bernardino Rivadavia” e Instituto
Nacional de Investigacion de las Ciencias Naturales. Ciencias Zoológicas, 3(5): [2] pp. + 233-333,
23 pls. [17 April]

IHERING, H. von, 1898, As especies de Ampullaria da Republica Argentina. Annales del Museo
Nacional de Buenos Aires, 4: 47-52. [13 August]

IHERING, H. von, 1915, Annexo М. 5. Molluscos. Commissáo de Linhas Telegraphicas Estrategicas de
Matto-Grosso ao Amazonas, Rio de Janeiro. 14 pp., 3 pls.

IHERING, H. von, 1919, Las especies de Ampullaria de la Argentina y la historia del Río de la Plata.
Pp. 329-350, pls. 37-38, in: Primera reunion de la sociedad Argentina de ciencias naturales.
Tucumán 1916. Imprenta y Casa Editora “Coni”, Buenos Aires. [May]

INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE [ICZN], 1999a, Opinion 1913.
Pila Róding, 1798 and Pomacea Perry, 1810 (Mollusca, Gastropoda): placed on the Official List,
and AMPULLARIIDAE Gray, 1824: confirmed as the nomenclaturally valid synonym of PILIDAE
Preston, 1915. Bulletin of Zoological Nomenclature, 56(1): 74-76. [March]

INTERNATIONAL COMMISSION ÓN ZOOLOGICAL NOMENCLATURE [ICZN], 1999b, International
Code of Zoological Nomenclature. Fourth edition. International Trust for Zoological Nomenclature,
London. xxix + 306 pp.

INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE [ICZN], 2002, Opinion 1997
(Case 3175). Ampullaria canaliculata Lamarck, 1822 (currently Pomacea canaliculata; Mollusca,
o specific name conserved. Bulletin of Zoological Nomenclature, 59(2): 137-138. [28

une

88 COWIE & ТНЕМСО

JAY, J. C., 1836, A catalogue of Recent shells with descriptions of new or rare species in the collection
of John С. Jay, M.D. Second edition. [Publisher not indicated], New York. 79 + [1] + [4] pp., 4 pls.
JAY, J. C., 1839, A catalogue of the shells, arranged according to the Lamarckian system; together
with descriptions of new or rare species, contained in the collection of John C. Jay, M.D. Third edi-
tion. Wiley & Putnam, New York and London. 125 + [1] pp., 10 pls. [after April]

JAY, J. C., 1850, A catalogue of the shells, arranged according to the Lamarckian system, with their
authorities, synonymes, and references to works where figured or described, contained in the
collectin of John C. Jay, M.D. Fourth edition. [Publisher not indicated], New York. [1] + 459 + [1] pp.

Supplement (pp. [460]-479) published 1852.

JOHNSON, R. 1., 1964, The Recent Mollusca of Augustus Addison Gould. United States National
Museum Bulletin, 239, 182 pp., 45 pls. [28 July]

JONAS, J. H., 1844, Vorläufige Diagnosen neur Conchylien, welche ausführlicher beschreiben und
abgebildet náchstens erscheinen werden. Zeitschrift fúr Malakozoologie, 1(March issue) : 33-37.

JONAS, J. H., 1845, Neue Conchylien. Zeitschrift für Malakozoologie, 2(November issue): 168-173.

JONAS, J. H., 1846, Molluskologische Beitráge. Abhandlungen aus dem Gebiete der Naturwissen-
schaften herausgegeben von dem naturwissenschaftlichen Verein in Hamburg, 1: 99-130, pls. 7-11.

JOUSSEAUME, F., 1877, Mollusques nouveaux de la République а l'Equateur. Bulletin de la Société
Zoologique de France, 12: 165-186, pl. 3. |

JOUSSEAUME, F., 1889, Voyage de M. Eugène Simon au Venezuela (Décembre 1887-Avril 1888).
Mollusques. Mémoires de la Société Zoologique de France, 2: 232-259, pl. 9.

JOUSSEAUME, F., 1894, Description d'un mollusque nouveau. Le Naturaliste, (2) 8(173): 120-121. [15 May]

КАВАТ, А. R., 1991, The classification of the Naticidae (Mollusca: Gastropoda): review and analysis of
the supraspecific taxa. Bulletin of the Museum of Comparative Zoology, 152: 417-449. [23 September]

КАВАТ, А. Б. & К. J. BOSS, 1997, Кап Eduard von Martens (1831-1904): his life and works. Depart-
ment of Mollusks, Museum of Comparative Zoology, Harvard University, Cambridge. vii + 417 pp.

KEAWJAM, БК. 5$. & E. S. UPATHAM, 1990, Shell morphology, reproductive anatomy and genetic pat-
terns of three species of apple snails of the genus Pomacea in Thailand. Journal of Medical and
Applied Malacology, 2: 49-62.

KING, P. P. & W. J. BRODERIP, 1831, Description of the Cirrhipeda, Conchifera and Mollusca, in a
collection formed by the officers of H.M.S. Adventure and Beagle employed between the years 1826
and 1830 in surveying the southern coasts of South America, including the Straits of Magalhaens
and the coast of Tierra del Fuego. The Zoological Journal, 5: 332-349. [after September]

KOBELT, W., 1911-1914, Die Gattung Ampullaria. Neue Folge. In Abbildungen nach der Natur mit
Beschreibungen. Pp. 1-236, pls. 22-79, in: H.C. KUSTER, Systematisches Conchylien-Cabinet von
Martini und Chemnitz. Neue Folge. Ersten Bandes zwanzigste Abtheilung. Baur & Raspe, Nurnberg
[= Nuremburg].

Text published in sections, which are dated on the first page of each section. The sections seem to have been
combined into “parts”, for which later dates are given by some authors. The earlier dates are taken as the dates
of publication in this catalog. The most recent collation is that of Welter-Schultes (1999), who had some later dates
than those accepted here.

Section Pages Part Printed date Date in other lists
1 1-8 550 13 March 1911a 1911
2 9-16 550 19 March 1911b 1911
3 17-24 550 19 May 1911c 1911
4 25-32 556 24 Мау 1911d 1912
5 33-40 556 15 October 1911e 1912
6 41-48 556 2 November 1911f 1912
И 49-56 557 4 November 19119 1912
8 57-64 557 25 November 1911h 1912
9 65-72 557 30 November 19111 1912
10 73-80 560 2 December 1911) 1912
11 81-88 560 5 January 1912а 1912
12 89-96 560 12 January 1912b 1912
13 97-104 560 10 January 1912c 1912
14 105-112 563 30 June 1912d 1913
15 113-120 563 12 September 1912e 1913
16 121-128 563 12 September 1912f 1913
17 129-136 563 1 November 19129 1913
18 137-144 565 12 November 1912h October 1913
19 145-152 565 4 March 1913a October 1913
20 153-160 567 8 July 1913b November 1913
21 161-168 567 12 July 1913c November 1913
22 169-176 567 14 July 1913d November 1913 (Continues)

NEW WORLD AMPULLARIIDAE 89

(Continued)
Section Pages Part Printed date Date in other lists
23 177-184 570 4 August 1913e 1913
24 185-192 570 21 December 1913f 1913
25 193-200 570 23 December 1913g 1913
26 201-208 574 29 December 1913h 1914
27 209-216 574 12 January 1914а 1914
28 217-224 576 9 July 1914b 1915
29 225-232 576 15 July 1914c 1915
30 233-236 576 16 July 1914d 1915

KOBELT, W., 1914e, Drei neue Ampullarienformen. Nachrichtsblatt der Deutschen Malako-
zoologischen Gesellschaft, 46(4): 176-178. [October]

LACANILAO, F., 1990, Reproduction of the golden apple snail (Ampullaridae [sic]): egg mass, hatch-
ing, and incubation. Philippine Journal of Science, 119: 95-105.

LAMARCK, J. В. [P. A. de М. de], 1801, Système des animaux sans vertebres ... L'auteur, Deterville,
Paris. viii + 432 pp. [21 January]

LAMARCK, [J. В. P. A. de M. de], 1804, Suite des mémoires sur les fossiles des environs de Paris.
Annales du Muséum National d'Histoire Naturelle, 5(25): 28-36.

LAMARCK, [J. B. P. A. de M. de], 1816, Encyclopédie méthodique. Tableau Encyclopédique et
méthodique des trois règnes de la nature. Vingt-troisième partie. Liste des objets representés dans
les planches de cette livraison. V. Agasse, Paris. 16 pp., pls. 391—488. [14 December]

This is the 84th livraison, which contains plates and 16 pages of explanations of the plates in the “Liste des objets”.

LAMARCK, [J. В. Р.А. de M.] de, 1822a, Histoire naturelle des animaux sans vertèbres ... Tome
sixième. 2”° partie. L'auteur, Paris. 232 pp. [April]

LAMARCK, [J. В. Р.А. de M.] де, 1822b, Histoire naturelle des animaux sans vertèbres ... Tome
septième. L'auteur, Paris. 711 pp. [August]

LAUP, S., 1991, Golden apple snail and its eradication in Papua New Guinea. Pp. 55-62, in: R.
KUMAR, ed., Proceedings of a seminar on pests and diseases of food crops - urgent problems and
practical solutions. Department of Agriculture and Livestock, Konedobu.

LEA, I., 1834, Observations on the naïades; and descriptions of new species of that and other fami-
lies. Transactions of the American Philosophical Society, (new series) 5: 23-119, pls. 1-19. [August
or September]

LEA, |., 1838, Description of new freshwater and land shells. Transactions of the American Philosophi-
cal Society, (new series) 6: 1-154, pls. 1-24. [after 15 June]

LEA, I., 1856, Description of thirteen new species of exotic Peristomata. Proceedings of the Academy
of Natural Sciences of Philadelphia, 8(3): 109-111. [after 26 June, before 15 August]

LEA, |., 1866, New Unionidae, Melanidae, etc., chiefly of the United States. Journal of the Academy of
Natural Sciences of Philadelphia, (new series) 6(2): 113-187, pls. 22-24. [December]

LINNAEUS, C., 1758, Systema naturae per regna tria naturae, secundum classes, ordines, genera,
species, cum caracteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. L. Salvii,
Holmiae [= Stockholm]. [iv] + 824 pp. [1 January]

MARSHALL, W. B., 1926, New land and fresh-water mollusks from Central and South America. Pro-
ceedings of the United States National Museum, 69(12): 1-12, pls. 1-3. [6 November]

MARSHALL, W. B., 1930, New land and fresh-water mollusks from South America. Proceedings of the
United States National Museum, 77(2): 1-7, pls. 1, 2. [25 January]

MARTENS, [K.] E. von, 1857, Die Ampullarien des Berliner Museums. Malakozoologische Blatter, 4:
181-213.

MARTENS, [K.] E. von, 1873, Die Binnenmollusken Venezuela's. Pp. 157-225, pls. 1-2, in: К. В.
REICHERT, ed., Festschrift zur Feier des hundertjáhrigen Bestehens der Gesellschaft
naturforschende Freunde zu Berlin. Ferd. Dúmmler's Verlag, Berlin.

MARTENS, [K.] E. von, 1890-1901, Biologia Centrali-Americana. Land and freshwater Mollusca. R.H.
Porter, London. xxviii + 706 pp., 44 pls.

Published in parts. Dates of publication are from Kabat & Boss (1997: 82-84, 86-87, 90-94) as follows:

Pages Plates Date of publication
1-40 1 1890
41-96 2-5 1891
97-176 6-9 1892
177-248 10-12 1893
= 13-15 1894
249-288 16 1897
289-368 17-20 1898
369-472 21-28 1899
473-608 29-41 1900

609-706, i-xxviil 42-44 1901

90 COWIE & THIENGO

MATHEWS, С. M. & T. IREDALE, 1912, “Perry's Arcana” — an overlooked work. Victorian Naturalist,
29(1): 7-16. [9 May]

MATON, W. G., 1811, Description of seven new species of Testacea. The Transactions of the Linnean
Society of London, 10: 325-332, pl. 24.

Maton's contribution was read on 7 November 1809, the date frequently cited for Helix platae. However, the correct
dates of publication of volume 10 are as follows:

Pages Date of publication

1-228 8 March 1810
229-414 (with index and title page) 7 September 1811

MCKILLOP, W. В. 8 А. D. HARRISON, 1980, Hydrobiological studies of eastern Lesser Antillean is-
lands V. St. Lucia: freshwater habitats, water chemistry and distribution of freshwater molluscs.
Archiv für Hydrobiologie Supplementband, 57: 251-290. [April]

MERMOD, G., 1952, Les types de la collection Lamarck au Muséum de Genève. Mollusques vivants,
Ш. Revue Suisse de Zoologie, 59(2): 23-97. [March]

MICHELSON, E. H., 1961, On the generic limits in the family Pilidae (Prosobranchia: Mollusca).
Breviora, 133: 1-10. [27 February]

MILLER, K., 1879, Die Binnenmollusken von Ecuador (Schluss). Malakozoologische Blatter, (Neue
Folge) 1: 117-203, pls. 4-15.

MOCHIDA, O., 1991, Spread of freshwater Pomacea snails (Pilidae, Mollusca) from Argentina to Asia.
Micronesica, Supplement, 3: 51-62.

MORELET, A., 1849, Testacea novissima insulæ Cubanæ et Americee centralis. Part 1. J.-B. Baillière,
Paris. 31 pp.

MORELET, A., 1857, Testacea nova Australiæ. Bulletin de la Société d'Histoire Naturelle du
Département de la Moselle, 8: 26-33. [after 2 April]

MORICAND, S., 1836, Mémoire sur les coquilles terrestres et fluviatiles, envoyées de Bahia par М...
Blanchet. Memoires de la Société de Physique et d'Histoire Naturelle de Genève, 7(2): 415-446, pl. 2.

MORRISON, J. P. E., 1946, The nonmarine mollusks of San José Island, with notes on those of Pedro
Gonzáles Island, Pearl Islands, Panamá. Smithsonian Miscellaneous Collections, 106(6): 1-49, pls.
1-3. [12 September]

MORRISON, J. P. E., 1952, Correction of the type locality of Pomacea cumingii (King) 1834. The
Nautilus, 65(3): 105-106. [25 February]

MOUSSON, A., 1869, Notiz über einige von Herrn Gustav Wallis aus dem nördlichen Süd-Amerika
zurückgebrachte Mollusken. Malakozoologische Blätter, 16: 170-189.

MOUSSON, A., 1873, Zweite Notiz über einige von Herrn Gustav Wallis aus dem nördlichen Süd-
Amerika zurückgebrachte Mollusken. Malakozoologische Blätter, 21: 1-19.

MULLER, ©. F., 1774, Vermium terrestrum et fluviatilium, seu animalium infusoriorum, helminthicorum,
et testaceorum, non marinorum, succincta historia. Volumen alterum. Heineck & Faber, Havniae [=
Copenhagen] & Lipsiae [= Leipzig]. xxxv + 214 + [10] pp.

NAGGS, F., 1997, William Benson and the early study of land snails in British India and Ceylon. Ar-
chives of Natural History, 24(1): 37-88.

NARANJO-GARCIA, Е. & A. GARCIA-CUBAS, 1986, Algunas consideraciones sobre el genero
Pomacea (Gastropoda: Pilidae) en Mexico y Centroamerica. Anales del Instituto de Biologia,
Universidad Nacional Аиюпота de Mexico, Serie Zoologia, 56(2): 603-606. [20 November]

NEAVE, $. A., 1940, Nomenclator zoologicus. A list of the names of genera and subgenera in zoology
from the tenth edition of Linnaeus 1758 to the end of 1935. Vol. Ш. Zoological Society of London,
London. 1065 pp.

NEVILL, G., 1877, Catalogue of Mollusca in the Indian Museum, Calcutta. Fasciculus E. Indian Mu-
seum, Calcutta. [iv] + 42 pp.

NEVILL, G., 1884, Hand list of Mollusca in the Indian Museum, Calcutta. Part Il. Gastropoda.
Prosobranchia-Neurobranchia (contd.). Indian Museum, Calcutta. x + 306 pp.

МС, P. K. L., L. M. CHOU & T. J. LAM, 1993, The status and impact of introduced freshwater animals
in Singapore. Biological Conservation, 64: 19-24.

NORRIS, А. & $. P. DANCE, 2002, Sylvanus Charles Thorp Hanley (1819-1899) a nineteenth-century
dilettante of the shell world. Journal of Conchology, 37(4): 363-382.

ORBIGNY, А. [D.] d’, 1835a, Synopsis terrestrium et fluviatilium molluscorum, in suo per Americam
meridionalem itinere. Magasin de Zoologie, 5(61-62): 1-44. [after 1 July]

ORBIGNY, A. [D.] d’, 1835-1847, Voyage dans l'Amérique Méridionale (le Brésil, la république
orientale de ГИгидиау, la république Argentine, la Patagonie, la république du Chili, la république
de Bolivia, la république du Pérou), exécuté pendant les années 1826, 1827, 1828, 1829, 1839,
1831, 1832 et 1833. Tome cinquième. 3.° partie: mollusques. P. Bertrand, Paris; V.* Levrault,
Strasbourg. 758 pp., 82 pls.

Published in livraisons as follows:

Livraison

Pages

1-48
49-72
73-104
105-128
129-152
153—176

177—184

185-232
233-280
281-328
329-376

377-408
409-424
425-472
473-488
489-528
529-600
601-656
657-704
705-728
729-758

Plates

‚ 32, 34

‚ 42, 43

‚ 50, 51

‚ 65
‚ 60-63

‚ 69

„ 67, 71

‚ 74, 79
‚ 76, 80

‚ 85
‚ 81
‚ 82

NEW WORLD AMPULLARIIDAE

Wrapper date

1834
1835
1835
1835
1834
1835
1834
1834
1835
1835
1835
1835
1835
1834
1836
1836
1836
1836
1836
1836
1837
1837
1837
1837
1837
1837
1837
1837
1837
1837
1835
1834
1837
1836
1839
1939
1839
1839
1839
1940
1840
1840
1841
1841
1841
1846
1845
1846
1846
1846
1842
1847
1847

Date of publication

(Evenhuis & Cowie, 1995)

15 May 1835b

15 May 1835c
Before 31 August 1835d
31 August 1835e

14 September 1835f
23 November 18359
7 December 1835h
4 January 1836a

18 April 1836b

30 May 1836c

11 July 1836e

1 August 1836f

26 September 18369
3 October 1836h

7 November 18361
27 February 1837a
3 April 1837b

5 June 1837c

19 June 1837d

7 August 1837e

18 September 1837f
6 November 1837h
5 March 1838a

23 April 1838b

6 May 1838c

11 June 1838d

15 October 1838e
12 November 1838f
8 April 1839a

29 April 1839b

24 June 1839c

11 November 1839d
21 November 1839e
6 September 1841a
8 November 1841b
8 November 1841c
8 November 1841d
15 November 1841e
15 November 1841f
15 November 1841g
15 November 1841h
14 February 1842a

91

92 COWIE & ТНЕМСО
ORBIGNY, А. [D.] d’, [1842]-1853, Mollusques. Tome second. Pp. [i-iv], 1-380, 28 pls., in: В. DE LA
SAGRA, Histoire physique, politique et naturelle de l'ile de Cuba. Arthus Bertrand, Paris.

Published in livraisons. Few details are available; the following are from С. Rosenberg (pers. comm. to КНС,
October 2001):

Pages Date of publication
12112 1842c

113-128 8 November 1844
129-208 16 February 1848
209-380 1853

PAETEL, F., 1873, Catalog der Conchylien-Sammlung von Fr. Paetel. Nebst Uebersicht des
angewandten Systems. Gebrüder Paetel, Berlin. [iv] + 172 pp. [after April]

PAETEL, F., 1887-1888, Catalog der Conchylien-Sammlung von Fr. Paetel. Vierte Neubearbeitung.
Erste Abtheilung: die Cephalopoden, Pteropoden und Meeres-Gastropoden. Gebrúder Paetel,
Berlin. [1 + 16 + 639 pp.

Published in Lieferungen as follows:

Lieferung Pages Date of publication
159 1-480 after June 1887
7,8 481-639 before 22 October 1888

PAIN, T., [1946]a, Two new species of Pila (= Ampullaria) from South America. Proceedings of the
Malacological Society of London, 26: 180-181, pl. 6. [31 January]

PAIN, T., 1946b, On Pila canaliculata and its locality. Proceedings of the Malacological Society of
London, 27(2): 58-59. [5 September]

PAIN, T., 1949a, Three new species of Pomacea from South America. Proceedings of the
Malacological Society of London, 27(6): 257-258, pl. 13. [14 January]

PAIN, T., 1949b, On the types of three species of Pomacea described by С.В. Sowerby Ш. Proceedings
of the Malacological Society of London, 28(1): 39-40, pls. 1, 2.

PAIN, Т., 1950a, A new species of Pomacea (Limnopomus) from Venezuela. Journal of Conchology,
23(4): 109-111. [July]

PAIN, T., 1950b, Pomacea (Ampullariidae) of British Guiana. Proceedings of the Malacological Society
of London, 28: 63-74, pls. 6-8.

PAIN, T., 1951, Pomacea hanleyana (Alderson). Journal of Conchology, 23(5): 145-146. [5 March]

PAIN, T., 1952, Notes on the Pomacea of Surinam, with special reference to Ampullaria sowerbyi
Vernhout. Basteria, 16(1/2): 30-32.

PAIN, T., 1953, Pomacea ghiesbreghti (Reeve) in Guatemala. Proceedings of the Malacological
Society of London, 29(6): 222-223. [16 January]

PAIN, T., 1956a, On a collection of Pomacea from Colombia, with description of a new subspecies.
Journal of Conchology, 24(3): 73-79. [20 February]

PAIN, T., 1956b, Notes on the generic names Pomacea and Ampullarius. Journal of Conchology,
24(3): 79. [20 February]

PAIN, T., 1957, Pomacea of the Sierra de Merida, Venezuela. Journal of Conchology, 24(5): 175-176.
[4 January]

PAIN, T., 1960, Pomacea (Ampullariidae) of the Amazon River system. Journal of Conchology, 24(12):
421-432. [16 December]

PAIN, T., 1964, The Pomacea flagellata complex in Central America. Journal of Conchology, 25(6):
224-231, pl. 13.

PAIN, T., 1972, The Ampullariidae, an historical survey. Journal of Conchology, 27: 453-462.

PAIN, T. & S. ARIAS, 1958, Descripcion de una especie nueva de Pomacea de Venezuela
(Mesogastropoda, Architaenioglossa, Mollusca). Novedades Cientificas. Contribuciones
Ocasionales del Museo de Historia Natural La Salle, Serie Zoológica, 24: 5-11. [22 December]

PARODIZ, J. J., 1969, The Tertiary non-marine Mollusca of South America. Annals of Carnegie
Museum , 40: 1-242. [30 June]

PARODIZ, J. J. & J. J. TRIPP, 1988, Types of Mollusca in the collection of the Carnegie Museum of
Natural History. Part 1. Bivalvia and Gastropoda (Prosobranchia and Opisthobranchia). Annals of
Carnegie Museum, 57: 111-154. [20 May]

PATTERSON, B., 1936, Caiman latirostris from the Pleistocene of Argentina, and a summary of South
American Cenozoic Crocodilia. Herpetologica, 1(2): 43-54. [28 December]

PERERA, С. & J. G. WALLS, 1996, Apple Snails in the Aquarium. T.F.H. Publications, Inc., Neptune
City, New Jersey. 121 pp.

PERRY, G., 1810-1811, Arcana; or the museum of natural history. Stratford, London. 84 pls.
[unnumbered], associated text.

Issued in monthly parts of unnumbered plates and associated text. The plates were numbered by Mathews 8
Iredale (1912) and monthly dates are here given following those authors. Pomacea maculata appears on pl. 12.

NEW WORLD AMPULLARIIDAE 93

However, Соме (1997a: 84) indicated that P. maculata appeared on pl. 11, based on pencilled numbers in the
BMNH copy. Neave (1940: 866) indicated that it appeared in signature G5. A full collation is in preparation and
confirms that P maculata appeared on pl. 12 in signature G5 (К. E. Petit, pers. comm. to RHC, 16 October 2000).

Plates Date of publication
[1-4] 1 January 1810a
[5-8] February 1810b
[9-12] 1 March 1810c

[13-16] April 1810d
[17-20] May 1810e
[21-24] June 1810f
[25-28] July 18109
[29-32] August 1810h

[33-36] September 18101
[37-40] October 1810]
[41-44] November 1810k
[45-48] December 18101
[49-52] January 1811a
[53-56] February 1811b
[57-60] March 1811c
[61-64] April 1811d
[65-68] May 1811e
[69-72] June 1811f
[73-76] July 18119
[77-80] August 1811h
[81-84] September 18111

PETIT, R.E. & М. С. HARASEWYCH, 1990, Catalogue of the superfamily Cancellarioidea Forbes and
Hanley, 1851 (Gastropoda: Prosobranchia). The Nautilus, Supplement, 1: 1-69. [6 March]

PHILIPPI, В. A., 1849, Centuria tertia testaceorum novorum. (Contin.) Zeitschrift für Malakozoologie,
6(2): 17-26. [May]

PHILIPPI, R. A., 1851-[1852], Die Gattung Ampullaria. т Abbildungen nach der Natur mit
Beschreibungen. 74 pp., pls. A, 1-21, in: H.C. KUSTER, Systematisches Conchylien-Cabinet von
Martini und Chemnitz. Neu herausgegeben und vervollstandigt. Ersten Bandes zwanzigste
Abtheilung. Bauer & Raspe, Nurnberg [= Nuremberg].

Published in parts as follows:

Part Pages Plates Date of publication
104 1-24 А, 1-5 1851

107 25-48 6-11 1852а

110 49-74 12-17 1852а

113 18-21 1852a

PHILIPPI, В. A., 18526, Centuria quinta testaceorum novorum. (Contin.) Zeitschrift für
Malakozoologie, 9(2): 20-29. [25 March]

PILSBRY, H. A., 1891, Land and fresh-water mollusks collected in Yucatan and Mexico. Proceedings
of the Academy of Natural Sciences of Philadelphia, 43: 310-334, pls. 14, 15.

Published in two parts as follows:

Pages Date of publication

310-328 25 August 1891a
329-334 22 September 1891b

PILSBRY, H. A., 1893, Notes on a collection of shells from the state of Tabasco, Mexico. Proceedings
of the Academy of Natural Sciences of Philadelphia, 44[1892]: 338-341, pl. 14. [24 January]

PILSBRY, H. A., 1899, A new Ampullaria. Proceedings of the Academy of Natural Sciences of Philadel-
phia, 51: 365. [8 September]

PILSBRY, Н. А., 1927a, Revision of the Ampullariidae of Jamaica and Cuba. Proceedings of the Acad-
emy of Natural Sciences of Philadelphia, 79: 247-253, pls. 21-22. [10 September]

PILSBRY, Н. A., 1927b, On Pomacea Perry (Ampullariidae). The Nautilus, 41(2): 63-64. [27 October]

PILSBRY, H. A., 1933, Zoological results of the Matto Grosso expedition to Brazil in 1931, — Il. Mol-
lusca. Proceedings of the Academy of Natural Sciences of Philadelphia, 85: 67-76, pl. 2. [17 July]

PILSBRY, H. A., 1944, Molluscan fossils from the Rio Pachitea and vicinity in eastern Peru. Proceed-
ings of the Academy of Natural Sciences of Philadelphia, 96: 137-153, pls. 9-11. [11 August]

PILSBRY, Н. А., 1953, The case of Paludina multilineata Say. The Nautilus, 67(2): 58-61. [11 November]

94 COWIE & ТНЕМСО

PILSBRY, Н. А. & J. T. BEQUAERT, 1927, The aquatic mollusks of the Belgian Congo. With а geo-
graphical and ecological account of Congo malacology. Bulletin of the American Museum of Natural
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aux instructions de S. Exc. M. le Vicomte du Bouchage, Secrétaire d’ Etat au Département de la
Marine, exécuté sur les corvettes de $. М. l'Uranie et la Physicienne, pendant les années 1817,
1818, 1819 et 1820; publié sous les auspices de S. Е. M. le Comte Corbière, Secrétaire d'Etat de
l'Intérieur, pouv la partie historique et les sciences naturelles, et de S. E. M. le Marquis de
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Aine, Paris.

Published in livraisons as follows:

Livraison Pages Date of publication
1 1-40 26 June 1824a
2 41-88 31 July 1824b
3 89-128 28 August 1824c
4 129-184 18 September 1824d
5 185-232 9 October 1824e
6 233-280 20 November 1824f
7 281-328 18 December 1824g
8 329-376 29 January 1825a
9 377-424 26 March 1825b
10 425-464 7 May 1825c
11 465-496 18 June 1825d
12 497-536 6 August 1825e
13 537-576 1 October 1825f
14 577-616 17 December 18259
15 617-664 26 April 1826a
16 665-712 14 June 1826b

In the “Préface” to this work (unnumbered page 3), Quoy & Gaimard thank Férussac for the nomenclature of the
terrestrial mollusks. In the introduction to the terrestrial and freshwater mollusks, Quoy & Gaimard state (pp. 463-
464) that “Nous devons à M. de Férussac la description des espèces que nous avons rapportées, dont il a fait
figurer plusiers dans son magnifique ouvrage sur les mollusques terrestres et fluviatiles.” Thus, authorship of the
descriptions of the terrestrial and freshwater mollusks (pp. 465-496, including Ampullaria) is Férussac, in Quoy
& Gaimard, though Quoy & Gaimard are the authors of the introductory text to that chapter (pp. 462-464).

REEVE, L. [A.], 1856, Monograph of the genus Ampullaria. Pls. 1-28, in: L. [A] REEVE, Conchologia
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This is a lambda book (a book in which plates were published separately, along with unnumbered pages of
explanatory text, as the plates were ready). After all plates were completed, they were bound into volumes. Dates
of publication as given on the bottom of the explanatory text for each plate are as follows for Ampullaria.

Plates Date of publication

2-4 June 1856a
5-12 August 1856b
13-20 October 1856c
211: 22 November 1856d
1, 23-28 December 1856e

NEW WORLD AMPULLARIIDAE 95

RICHARDS, Н. G., 1933, А conchological expedition to Cuba. Proceedings ofthe Pennsylvania Acad-
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RÓDING, P. F., 1798, Museum Boltenianum. Pars Secunda. J. С. Trapp, Hamburg. viii + 199 pp. [Sep-
tember]

Authorship of this work determined by ICZN Direction 48.

ROISSY, F. de, 1805, Histoire naturelle, genérale et particuliere, des mollusques, animaux sans
vertèbres et a sang blanc. Tome cinquième. Е. Dufart, Paris. 448 pp.

RUHOFF, Е. A., 1980, Index to the species of Mollusca introduced from 1850 to 1870. Smithsonian
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SAULCY, E. de, 1854, Note sur l'ampullaire œil d Ammon, Ampullaria effusa (Lamarck). Bulletin de la
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SAY, T., 1829, Descriptions of some new terrestrial and fluviatile shells of North America. The Dissemi-
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Published in parts, as follows:

Pages Date of publication

229-230 29 July 1829a
244-246 12 August 1829b
259-261 26 August 1829c
275-277 9 September 1829d
291-293 23 September 1829e
308-310 7 October 1829f
323-325 21 October 1829g
339-341 4 November 1829h
355-356 18 November 18291

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NEW WORLD AMPULLARIIDAE 97

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Revised ms. accepted 12 March 2003

INDEX OF LISTED TAXA belizensis Crosse & Fischer 58
bilineata Reeve 78-79
This index includes all ampullariid names brasiliensis Paetel 58

listed in the main body of the catalog. Family- bridgesii Reeve 42, 58, 62

group names are in BOLDFACE capitalized brownii Jay 46

letters. Genus-group names are in all bruguieri Deshayes 80

CAPITALIZED letters. Unavailable names are in bulla Reeve 58

italics. Unpublished names are listed in quotes buxea Reeve 58, 78-79

in plain type. caliginosa Reeve 58

camena Pain 58, 69
acuta Paetel 56, 81 canaliculata Lamarck 42-43, 57-59, 62, 65-66,
“adjusta” no author 81 76, 81
adusta Reeve 80 cassidiformis Reeve 59
aldersoni Pain 56 castanea Deshayes 52
amazonica Reeve 43, 56 castelloi Sowerby 59, 72
americanista Ihering 78 castelnaudii Hupé 59
AMPULLARIIDAE Gray 41-81 catamarcensis Sowerby 59
AMPULLAROIDES Gray 46 catemacensis Baker 59
AMPULLOIDEA Orbigny 46 cerasum Hanley 57, 59
AMPULLOIDES Orbigny 46 CERATODES Guilding 49-51
angulata Jay 56-57 chamana Hinkley 59
angulata Dunker 56-57 chaquensis Scott 59
angulata Deshayes 56 chemnitzii Philippi 59, 72-74
angulata H. Adams 8 A. Adams 57 chiapasensis Fischer & Crosse 59
aperta Philippi 80 chiquitensis Orbigny 50
arata Fischer & Crosse 57 chlorostoma Sowerby 52
archimedes Эрих & Wagner 57 cincta Cristofori & Jan 60
armeniacum Hupé 57 cingulata Philippi 52
ASOLENA Herrmannsen 46 citreum Reeve 60
ASOLENE Orbigny 41-42, 46-50, 60, 78 columbensis Jay 60
aulanieri Deville 8 Huppé 57, 60, 68, 75 columbiensis Philippi 60
auriformis Reeve 57 columbiensis Reeve 60, 68, 76
aurostoma Lea 57 columellaris Gould 51,60, 74-75
australis Orbigny 57 commissionis Ihering 60
autumnalis Reeve 57 CONCHYLIUM Cuvier 51
avellana Sowerby 57-58, 81 conica Guppy 52, 81
baeri Dautzenberg 52 conoidea Martens 60
balteata Philippi 52 consolatrix Ihering 60

batabana Paetel 58 contamanoensis Preston 60

98 СО\МЕ & ТНЕМСО

contrarius Müller 50

cornuarietis Linnaeus 49-51
cornucopia Reeve 60
costaricana Martens 60

cousini Jousseaume 61

crassa Orbigny 47-48

crassa Swainson 46-49, 67, 80
crocostoma Philippi 52
crosseana Hidalgo 61

cubensis Morelet 61

cubensis Reeve 61, 72

cumingii King & Broderip 61, 69, 72, 74
cuprina Reeve 42, 53
cyclostoma Spix & Wagner 47-49, 63, 77
dacostæ Sowerby 61

decussata Moricand 60-61, 67
delattrei Fischer & Crosse 61, 67
depressa Say, 62, 71,81

diffusa Blume 58, 62

dilatata Orbigny 62

disseminata Kay 62

dolioides Reeve 62

dolium Philippi 62

dorbignyana Philippi 62

dorbignyi Philippi 62

dubia Guilding 53, 55

dysoni Hanley 62

EFFUSA Jousseaume 42, 51-55, 61, 72-73

effusa Müller 53-54, 56
electrina Reeve 63

elegans Orbigny 63, 72
elongata Dall 49

elongata Orbigny 49, 63
equestris Róding 79

erogata Crosse & Fischer 63
erronea Nevill 63

erythrostoma Reeve 63

eumicra Crosse 8 Fischer 63
exculpta Fischer & Crosse 63
exigua Philippi 80

eximia Dunker 59, 63

expansa Miller 53

exumbilicata Spix 8 Wagner 47
fairchildi Clench 78

falconensis Pain 8 Arias 63
fasciata Lamarck 64, 79
fasciata Reeve 64

fasciata Roissy 51, 58, 60, 63-64, 76, 79
fasciatus Guilding 50-51
fasciolata Spix & Wagner 47
FELIPPONEA Dall 41-42, 49-50
ferruginea Martens 64

figulina Spix 8 Wagner 64, 75
flagellata Say 43, 57-58, 60, 62-77
flatilis Reeve 64

flava Smith 64

fumata Reeve 64

gallardoi Ihering 47

garciae Richards 64

georgii Williams 64

geveana Philippi 53, 55

gevesensis Deshayes 53-54, 56

ghiesbreghti Reeve 64-65, 69

gibbosa Paetel 79

gigantea Barbosa Rodrigues 65

giganteus Tristram 65

gigas Spix & Wagner 42-43, 52, 65-66, 69, 76

glauca Linnaeus 51-56, 61, 72

gossei Reeve 65

granulosa Sowerby 47, 49

guadelupensis Martens 44, 54

guaduasensis Anderson 65

gualtieri Orbigny 65

“gualteriana” no author 81

guatemalensis Martens 65

guyanensis Lamarck 63, 65, 70-71, 73

haemastoma Reeve 65

hanleyana Alderson 66, 75

hanleyi Reeve 66

haustrum Reeve 43, 66

hepataria Reeve 79

hollingsworthi Pain 66

hondurasensis Reeve 66

hopetonensis Lea 57-58, 66, 73, 79, 81

ignota Róding 79

iheringi Pilsbry 49

immersa Reeve 43, 66

imperforata Swainson 79

impervia Philippi 47-48

innexa Crosse & Fischer 66

insularum Orbigny 42, 57, 64-66, 76-77

intermedia Ferussac 54

intermedia Gray 50-51

interrupta Sowerby 60, 66

intropicta Reeve 67

knorrii Philippi 50-51

labiosa Philippi 67

lamarckii Philippi 67

lattrei Reeve 59, 61-62, 67

lemniscata Crosse 4 Fischer 67

leucostoma Swainson 67

levior Sowerby 67

LIMNOPOMUS Dall 47, 51, 57, 59-60, 66, 68—69

lineata Spix & Wagner 42-43, 59, 62, 64, 67,
70, 72, 75-76

linnaei Philippi 67

livescens Reeve 60, 65, 67, 76

lutea Farfante 67

luteostoma Swainson 51-55

lymnaeaeformis Reeve 67

maculata Perry 41, 43, 51, 61, 65, 68

malleata Jonas 57, 59, 63, 68-70, 73, 75

manco Pilsbry 68

manetou Róding 68

NEW WORLD AMPULLARIIDAE 99

marginatra Jonas 68

MARISA Gray 41-42, 49-51, 77

martensiana Nevill 60, 68, 76

martinezi Hidalgo 68

megastoma Sowerby 78

melanocheila Reeve 68

melanostoma Philippi 69

meridaensis Pain 69

mermodi Sowerby 69

meta lhering 69

metcalfei Reeve 69

mexicana Martens 69

miamiensis Pilsbry 69

miltocheilus Reeve 69

miltochilus Fischer 8 Crosse 69

minor Nevill 69-70

minuscula Baker 54

“miquitensis” Spix 81

modesta Busch 70

monachus Crosse & Fischer 70

monstrosa Sowerby 70

monticola Vernhout 47

nais Pain 70

naticoides Orbigny 48

neritina Gmelin 52, 54, 56

neritiniformis Dall 49

neritoides Orbigny 78

nigrilabris Philippi 70

nobilis Reeve 70

notabilis Reeve 70

novaegranadae Busch 70

nubila Reeve 48, 70

nucleus Philippi 79

oajacensis Fischer & Crosse 70

oblonga Nevill 48

oblonga Swainson 48, 70

obtusa Deshayes 79

ocanensis Kobelt 70

occlusa Crosse & Fischer 71

ochracea Jay 71

oculuscommunis Gmelin 54

oligista Pilsbry & Olsson 55

olivacea Spix & Wagner 71, 73

olivieri Deshayes 48

orinoccensis Troschel 55

ormophora Morelet 48

oviformis Deshayes 71

pachystoma Paetel 79

pachystoma Philippi 55

pallens Philippi 80

palmeri Marshall 71

“palmieri” Preston 81

paludinoides Cristofori & Jan 80

paludosa Say 42, 57-58, 62-64, 66-67, 69-74,
79, 81

papyracea Spix 8 Wagner 71

pattersoni Boss & Parodiz 55

patula Reeve 59, 71

pealiana Lea 71

penesma Kay 72

periscelis Róding 72

peristomata Orbigny 72

pernambucensis Reeve 72

pertusa Sowerby 72

petiti Crosse 48

phaeostoma Philippi 72

philippiana Baker 55

physis Hupé 72

physoides Reeve 72

picta Reeve 72

PILIDAE Preston 41, 46

pinei Dall 72

planogyra Pilsbry 50-51

planorboides Cristofori & Jan 79

planorbula Philippi 55

platae Maton 46-49

poeyana Pilsbry 61, 72

POMACEA Perry 41-43, 47, 49-51, 54-56, 58
60, 62-64, 66, 68-70, 72-74, 76-77, 79, 81

pomatia Martens 72

POMELLA Gray 41-42, 50, 78

pomum Philippi 72

porphyrostoma Reeve 73

prasina Fischer 8 Crosse 73

producta Reeve 73

prourceus Boss & Parodiz 73

prunella Hupé 81

prunulum Reeve 55

pulchella Anton 47-48

pulchra Griffith & Pidgeon 66, 73

puncticulata Swainson 73, 76

puntaplaya Cousin 73

purpurascens Guppy 73

pyrum Philippi 73

quercina Spix 8 Wagner 73

quinindensis Miller 55

quitensis Busch 69, 74

reflexa Swainson 74

retusa Philippi 74

reyrei Cousin 74

rhodostoma Appun 55

robusta Philippi 74

roissii Orbigny 48

roissyi Orbigny 47-48

rotula Mousson 51

rotundata Say 81

rufilineata Reeve 80

rugosa Lamarck 74

sanjosensis Morrison 74

scalaris Orbigny 56-58, 74

scholvieni Kobelt 74

schrammi Crosse 78

semitecta Mousson 74

semperi Kobelt 75

100 COWIE & ТНЕМСО

sepulta Róding 80 tristrami Crosse 8 Fischer 60, 76
simplex Reeve 75 trochulus Reeve 80

sinamarina Вгидшеге 78 “undata” no author 81

sloanii Férussac 48 “unicolor” Martens 81

solida Busch 49 unicolor Philippi 76

sordida Swainson 54, 57, 69, 75, 81 urabaensis Pain 76

sowerbyi Vernhout 49 urceus Müller 51, 62-63, 65-68, 70-71, 73-74,
spirata Deville 8 Hupé 75 76-77

spixii Orbigny 49 venetus Reeve 76

sprucei Reeve 75 “venezullum” no author 81
storeria Jay 49 vermiformis Reeve 76

strebeli Fischer & Crosse 75 vexillum Reeve 69, 76

superba Marshall 75 vickeryi Pain 58, 76
suprafasciata Kobelt 55 villata Sowerby 56

SURINAMIA Clench 78 violacea Valenciennes 77
swainsoni Philippi 66, 69, 75-76 welwitschiana Drouét 77
swainsonii Нире 75-76, 79 woodwardi Dohrn 77

“tacarigua” Pilsbry 81 yatesii Reeve 77

tamsiana Philippi 56 yucatanensis Crosse & Fischer 77
tenuissima Jousseaume 76 yzabalensis Martens 77

teres Philippi 56, 61 zeteki Morrison 77

testudinea Reeve 76 zischkai Blume & Pain 77

tristis Gaudion 80 zonata Orbigny 49

tristis Guppy 56 zonata Spix & Wagner 49, 57, 68, 78

MALACOLOGIA, 2003, 45(1): 101-108

ASSESSMENT OF GENETIC НЕТЕКОСЕМЕПУ WITHIN LABORATORY-
MAINTAINED SCHISTOSOMA MANSONI-RESISTANT STOCKS OF
BIOMPHALARIA GLABRATA SNAILS BY RAPD-PCR

Wannaporn Ittiprasert', Christopher Rowe? Carolyn Patterson’, André Miller’,
Nithya Raghavan?, Susan Bandoni’, Fred Lewis? 8 Matty Knight”

ABSTRACT

Random amplified polymorphic DNA (RAPD)-PCR analysis was used to assess the extent of
genetic diversity within two laboratory-maintained Schistosoma mansoni-resistant stocks of
Biomphalaria glabrata (10-R2 and BS-90). Both stocks routinely serve as parents in crosses
with susceptible snails for studying the genetics of parasite resistance in the snail host.
Genomic DNA was isolated from individual adult 10-R2 and BS-90 snails. From RAPD-PCR
conducted with 16 anonymous primers, no polymorphisms were detected within the BS-90
stock, whereas 13 primers revealed considerable intrastrain variations showing different
sized bands among the 10-R2 snails. The polymorphisms in the 10-R2 stock allowed us to
identify three distinct groups (Types 1, 2 and 3) within these snails. Random screening of
individual 10-R2 snails revealed that, of the three distinct types, Types 1 and 2 snails were
found at similar frequencies (approximately 45%), whereas 10% fell into the third group (Type
3). The identification of genetic variants within the 10-R2 stock demonstrates the need for
careful assessment of the existence of diverse forms in this stock prior to conducting genetic
crosses with these snails.

Key words: genetic heterogeneity, Biomphalaria glabrata, Schistosoma mansoni, interme-

diate snail host, intrastrain variation, resistance, DNA polymorphisms, RAPD-PCR.

INTRODUCTION

Much of the research on the genetics of the
interrelationship between the parasitic helminth
Schistosoma mansoni and the snail host
Biomphalaria glabrata has been made possible
because of the availability of several genetically
defined B. glabrata stocks that breed true for a
variety of traits (Richards, 1970). From crosses
conducted between many different snail stocks,
these have been grouped into four categories
(Types I-IV) based on susceptibility to $.
mansoni (Richards 8 Shade, 1987, review).
Snails in the Type | category are resistant to
parasite infection at any age. Type Il snails are
susceptible as juveniles, but adult resistant.
Type Ш snails are susceptible at any age,
whereas Type IV snails are juvenile susceptible
and adult variable. Of all these categories,
snails from categories Types | and III have re-
ceived the most attention by investigators study-

ing the molecular basis of resistance and sus-
ceptibility to infection. Accordingly, many of these
studies have been done using either the proto-
type Type | resistant stocks 10-R2, 13-16-R1
orthe “Salvador” strain (also referred to as BS-
90), or the susceptible Type Ш M-line and NMRI
snail stocks.

It is known that genes of both the snail and
parasite affect the outcome of this host/para-
site relationship. Based on this, it was sug-
gested as early as the late 50s (Hubendick,
1958) that one method for reducing transmis-
sion may involve the use of parasite resistant
snails to replace susceptible ones in an en-
demic area. Whether or not this form of control
will become reality, studying the molecular bi-
ology of the snail and parasite has become the
focus of considerable research in recent years.
For the emerging field of molecular malacology,
the existence of genetically defined B. glabrata
snail stocks has therefore been invaluable.

‘Department of Microbiology and Immunology, Faculty of Tropical Medicine, Mahidol University, Bangkok, Thailand
“Biomedical Research Institute, 12111 Parklawn Drive, Rockville, Maryland 20852, U.S.A.
“Department of Biology, SUNY-Geneseo, Geneseo, New York, U.S.A.

*Corresponding author: mknight@afbr-bri.com

102 ITTIPRASERT ET AL.

Although recent success has been achieved
in defining molecular markers for resistance
in B. glabrata (Knight et al., 1999), there is evi-
dence that molecular heterogeneity may exist
between individual snails within either resis-
tant or susceptible laboratory-maintained
stocks, potentially complicating interpretation
of experimental results. In earlier studies,
while searching for RFLPs in the small ribo-
somal gene, we reported on a smaller scale
the genetic diversity within both resistant (10-
R2) and susceptible (M-line) snail stocks
(Knight et al., 1991). Later, Mulvey 8 Bandoni
(1994) reported differences in allozyme fre-
quencies among nine enzymes from M-line
snails obtained from several different labora-
tories.

For this study, we were interested in further
documenting the genetic variability among our
resistant snail stocks. Our goal was to assess
the extent of genetic diversity in the present
laboratory-maintained resistant stocks by
means other than RFLP analysis before us-
ing them to conduct crosses with the suscep-
tible M-line snails. Although the original 10-R2
stock was maintained in the laboratory for
many generations without loss or reduction of
the refractory trait (Richards, personal com-
munication), some stocks in other laborato-
ries occasionally show partial susceptibility,
especially when they are exposed as juveniles.

The RAPD-PCR method only detects domi-
nant alleles and has the potential to show in-
consistencies in results if assays have not
been optimized (Williams et al., 1990; Welsh
& McClelland, 1990). On the other hand, the
method has been used effectively in assess-
ing genetic diversity in both laboratory main-
tained and field populations of B. glabrata
(Larson et al., 1996; Vidigal et al., 1994). More-
over, in comparison to other DNA genotyping
tools available, for example RFLP analysis and
examination of polymorphic microsatellite
(simple sequence repeats) loci, RAPD-PCR
requires no prior sequencing or cloning steps,
uses negligible amounts of DNA, and requires
no radioactive isotopes in the assay.

Here, we compare the genotypes of indi-
vidual schistosome-resistant BS-90 and 10-
R2 snails using RAPD-PCR analysis. Of the
two stocks, no genetic diversity was found be-
tween snails of the BS-90 stock by this analy-
sis. Genetic heterogeneity however was found
in the 10-R2 stock, allowing us to identify three
distinct substocks (herein after referred to as
Types 1-3).

MATERIALS AND METHODS
Snails

The two different $. mansoni-resistant stocks
of B. glabrata used in this study were the 10-
R2 (Richards, 1975) and BS-90 (Paraense €
Correa, 1963) snails. Adult snails (10-14 mm
in diameter) were reared as individuals in self-
fertilizing lineages in beakers (400 ml) with petri
dish covers in aerated tap water, and fed ro-
maine lettuce. Cohorts of the snails used in this
study, after exposure to $. mansoni miracidia,
have always displayed the resistance phenotype.

Genomic DNA Extraction

Genomic DNA was prepared from the whole
body of individual snails (Knight et al., 1998). Snails
were cleaned with a Q-tip and maintained over-
night in sterile deionized water (DW) containing
100 ug/ml of ampicillin at room temperature. In
some cases, DNA was isolated from snail ten-
tacles. For this, tentacles were snipped off live
snails with a pair of fine tipped forceps under a
dissecting microscope, placed in siliconized tubes
and kept frozen at —70°C until required. The fro-
zen tentacles were thawed into 200! CTAB buffer
(2% w/v of cetyltrimethylammonium bromide, 1.4
М NaCl, 20 mM EDTA, 0.2% v/v of f-
mercaptoethanol, 100 mM Tris-HCl, pH8.0) con-
taining proteinase K (0.1mg/ml), homogenized
with a motorized pestle (Kimble, Illinois), and in-
cubated at 55°C for 1 h and processed further
as described previously (Knight et al., 1998).

DNA was recovered by centrifugation and
washed in cold 70% ethanol, air dried and re-
suspended in an appropriate volume of sterile
distilled water. The quality of DNA was deter-
mined by horizontal flat-bed gel electrophore-
sis (0.8% agarose) resolved in TBE buffer (89
mM Tris-base, 89 mM Boric acid and 2 mM
EDTA, pH 8.0). DNA concentration was mea-
sured under UV illumination (Eagle Eye,
Stratagene, California) from the intensities of
ethidium bromide staining of the extracted DNA
samples compared to that of known amounts
of standard DNA spotted (1.0 ul) onto agarose
plates incorporated with ethidium bromide.

Random Amplified Polymorphic DNA
(RAPD)-PCR

Genetic diversity between individual resistant
Snails was analyzed by RAPD-PCR as described
previously (Larson et al., 1996). The genotypes

GENETIC HETEROGENEITY OF BIOMPHALARIA GLABRATA 103

TABLE 1. The frequency of detecting variant
forms (%) of 10-R2 snails based on genotypying
DNA from individual snails assayed by RAPD-
PCR with random oligonucleotide decamer prim-
ers. Some primers produced no product (N/P)
and others detected no genetic diversity within

the snails (indicated by 0).

Variants (%)

Sequence 5'>3' Type 1 Type 2 Type 3

Primer

ОРМ-05 СССААССТСТ 45 45 10
ОРМ-07 СССТСАСТСА 45 45 10
OPZ-05 TCCCATGCTG 45 45 10
OPM-01 GTTGTTGGCT 45 45 МР
ОРМ-04 GGCGGTTGTC 45 45 МР
ОРМ-08 TCTGTTCCCC 45 45 МР
ОРМ-09 GTCTTGCGGA 45 45 МР
ОРМ-10 TCTGGCGCAC 45 45 МР
OPZ-01 TCTGTGCCAC 45 45 МР
OPZ-03 САССАССССА 45 45 МР
OPZ-06 GTGCCGTTCA 45 45 МР
OPZ-07 ССАССАССАС 45 45 МР
OPZ-10 СССАСАААСС 45 45 МР
ОРМ-06 CTGGGCAACT 0 0 0
ОРМ-11 СТССАСТСТС 0 0 0
OPZ-11 СТССАСТСТС 0 0 0

of individual 10-R2 and BS-90 snails were deter-
mined by anonymous 10-mer oligonucleotide
primers (listed in Table 1) obtained from Operon
Technologies (Alameda, California). The control
(no template DNA) was sterile distilled water. Af-
ter amplification, the reaction was mixed with 5
ul of loading buffer (40% sucrose, 0.25% bro-
mophenol blue and 0.25% xylene cyanol) and run
on a 1.2% agarose gel containing 0.5 ug/ml of
ethidium bromide in TBE buffer (voltage at 100V).
The amplified bands were visualized by UV illu-
mination and sizes estimated based on the mi-
gration of a 100-base pair (bp) ladder (Gibco BRL,
Gaithersburg, Maryland).

RESULTS AND DISCUSSION

RAPD analysis was performed using DNA
extracted from two different stocks of B.
glabrata. DNA samples from 24 individuals
of the 10-R2 stock and 20 individual BS-90
snails were compared using 16 random prim-
ers. The results revealed that with all of the ran-
dom primers tested, no polymorphisms were
found between individual BS-90 snails, whereas
13 of 16 primers revealed variations (different

sized bands) between the 10-R2 snails ana-
lyzed (Table 1). Thirteen of the primers revealed
that approximately 45% of the individuals ana-
lyzed could be grouped into one of two geno-
types (Types 1 or 2). Three primers (OPM-05,
OPM-07, OPZ-05) revealed a third type (Type
3) but at a lower frequency (10%). Only three
primers (OPM-06, OPM-11 and OPZ-11)
showed no polymorphisms between all indi-
vidual 10-R2 snails tested.

Results of specific bands obtained for the
three 10-R2 types, using the 13 primers that
produced amplified products, are summa-
rized in Table 2. Seven primers (OPM-04,
OPM-05, OPM-07, OPM-08, OPM-10, OPZ-03
and OPZ-10) amplified different sized prod-
ucts for Type 1 snails, but not for Type 2, thus
allowing differentiation between the two
types. Six primers produced bands specific
for Type 2 snails and three primers amplified
specific products for Type 3 snails.

Representative examples of the polymor-
phisms that enabled us to segregate the 10-R2
snails into three distinct groups using three dif-
ferent primers OPM-05, OPM-07 and OPZ-05,
can be seen in Figures 1A-C, respectively.
Primer OPM-05 (Fig. 1A), revealed the pres-
ence of high molecular weight bands (shown
by the arrows) in Type 1 (approximately,
1600bp), and Type 3 (1550bp) snails that were
absent in Type 2 snails. Amplification using
primer ОРМ-07 (Fig. 1B) showed (indicated by
arrows) two specific bands (1500bp and
700bp) in Type 1 snails that were absent in
Types 2 and 3 snails and the presence of Type
3 specific bands (1600bp, 1500bp, 900bp and
750bp). DNA from the 3 types amplified using
primer OPZ-05 (Fig. 1C) produced specific
markers for each (a band of 550bp for Type 1,
650bp and 800bp for Type 2,and a 1600bp frag-
ment and a doublet at approximately 1500bp/
1550bp for Type 3). Amplification with the same
three primers (OPM-05, OPM-07 and OPZ-05)
using DNA from individual BS-90 snails (Figs.
2A-C, respectively) showed that unlike the
10-R2 snails, intrastrain variation was not de-
tected within this stock. In some cases, results
showed differences in the intensities of bands
but because the same size bands were ampli-
fied from all individual BS-90 snails, these dif-
ferences in band intensities may be due to
minor inconsistencies in the amount of tem-
plate DNA utilized rather than inherent genetic
differences in this stock.

Using RAPD-PCR, we thus revealed consid-
erable molecular diversity between individual

104 ITTIPRASERT ET AL.

TABLE 2. The detection of Type-specific bands by RAPD-PCR analysis of individual
10-R2 snails. The sizes of the various substock specific bands were determined by
agarose gel electrophoresis as described in Materials and Methods. No amplified
products (N/P) were produced by ten of the primers.

Specific bands (bp)

Primer Type 1
OPM-05 1,550
OPM-07 1,400, 650
OPZ-05 550
OPM-01 750, 400
ОРМ-09 1,400
OPZ-07 1,550, 400
OPM-04 1,550, 1,500
OPM-08 1,550, 700
OPM-10 1,550
OPZ-03 650
OPZ-10 1,550, 550, 500, 400
OPZ-01
OPZ-06
OPM-06
OPM-11
OPZ-11

10-R2 snails, one of the most commonly used
resistant stocks in investigations of the molecu-
lar basis of the В. glabrata/S. mansoni relation-
ship. Similar genetic heterogeneity was not
detected between snails of another commonly
used resistant stock (BS-90). Because of cur-
rent research interest in these snails in the
search for genes that define the resistance phe-
notype, we felt that a thorough background of
their genetic diversity/stability was warranted in
order to avoid misscoring of genotypes in fu-
ture molecular genetic studies. Although our
focus is not population genetics, we also point
to findings that may be pertinent for furthering
our understanding of the population biology of
these organisms.

From our earlier crosses between the resis-
tant (BS-90) snails and the susceptible (M-line)
snails we were able to demonstrate that RAPD-
markers segregated with the adult resistant
phenotype (Knight et al., 1999). We hoped
that crosses generated with other resistant snail
stocks (e.g., 10-R2) would enable us to not only
confirm our earlier results but help identify other
DNA markers that may be associated with the
refractory phenotype.

Type 2 Type 3
1,500
800
650, 800 1,600, 1500 (db)
500 N/P
700,550, 500 N/P
350 N/P
N/P
N/P
N/P
N/P
N/P
900 (db), 1,100 N/P
600 (db) N/P

no polymorphisms detected

The level of variability observed in the 10-R2
snails was higher than expected, given that
these snails were laboratory selected and main-
tained for many years by selfing. However,
Mulvey & Vrijenhoek (1981) previously reported
polymorphisms for four of 16 allozyme loci stud-
ied in this stock. In addition, the 10-R2 snails
are derived in part from the M-line stock devel-
oped by Newton (1955) (Richards, 1973). M-
line snails have been shown, by allozyme
analysis, to display a high degree of genetic
heterogeneity (Mulvey 8 Vrijenhoek, 1981;
Mulvey 8 Bandoni, 1994). Earlier molecular
studies in our laboratory using the 10-R2 snails,
based on RFLP analysis of ribosomal RNA
genes, had also revealed a certain degree of
genetic heterogeneity within this stock (Knight
et al., 1991). Our earlier study was limited in
scope, however, and the fact that we can dis-
tinguish three distinct subgroups within the
stock with only a small subset of primers by
RAPD-PCR was unexpected.

Contamination might explain the variability
observed in the 10-R2 stock, and has previ-
ously been reported in the M-line stock of
Biomphalaria glabrata (Mulvey 8 Bandoni,

GENETIC HETEROGENEITY ОЕ BIOMPHALARIA GLABRATA 105

Туре 1 Туре 2 Туре 3 Control

ОРМ -05

ОРМ -07

OPZ-05

FIG. 1. Ethidium bromide stained agarose gels showing RAPD-PCR products amplified by anony-
mous primers (A) OPM-05, (B) OPM-07, and (C) OPZ-05 using DNA from individual 10-R2 snails that
segregated into three different substocks (Types 1-3). The different Type-specific bands identified
with these primers are indicated by the arrows. Control lanes represent amplifications done in the
absence of template DNA.

106 ITTIPRASERT ET AL.

A)

1500

800 ОРМ-05

600
400

В)

1500

800
600

400

200

OPZ-05

FIG. 2. Ethidium bromide stained agarose gels showing RAPD-PCR products using primers (A) OPM-
05, (B) OPM-07 and (C) OPZ-05 and DNA from individual BS-90 snails. Control lanes represent ampli-
fications done with the same primers without DNA template as control.

GENETIC HETEROGENEITY OF BIOMPHALARIA GLABRATA 107

1994). However, because the 10-R2 snails
were reared as individual selfing lineages, it is
very unlikely that the within-population varia-
tion seen in this stock is due to contamination.
The black-eye pigmentation of these snails
makes them readily distinguishable from al-
bino stocks maintained in the laboratory.

Several genetic mechanisms might explain
the presence of the three multilocus genotypes
observed. The occurrence of only three mul-
tiple locus genotypes would be consistent with
the history of selfing in the snails that we stud-
ied. It is possible that the three genotypes found
within these snails may reflect derivation from
separate self-fertilizing lineages. Further study
of these markers using progeny from selfing
and outcrossing individuals would be needed
to establish this. It is also possible that inbreed-
ing has produced distinctive combinations of
alleles at multiple loci that work best in concert,
without disruption. Again, further research is
needed in order to investigate this possibility.
Finally, spontaneous genetic mutations within
other susceptible stocks have also been re-
ported. For example, selection of mutants from
the NMRI stock gave rise to the LAC-line snails,
which display the non-susceptible phenotype,
in addition to other abnormalities (Cooper et al.,
1994: Cousin et al., 1995).

In contrast to our observations for the BS-90
snails, the laboratory-derived resistant 10-R2
stock, on which considerable research has
been reported, has proven to be morphologi-
cally more variable (Richards, personal com-
munication). This stock was selected for
juvenile resistance and maintained in self-fertil-
izing lineages. Frequent exposure of snails
from our 10-R2 stock, either as adults or juve-
niles, has revealed no susceptibility to the para-
site, and snails from the three types reported
here do not have obvious phenotypic differ-
ences. Several morphologic differences have,
however, been detected in this stock over the
years. These include deformed, everted
mouthparts, abnormal intestine, variations in
egg clutch size, abnormal position of the aorta,
unusual shell development, and abnormal ten-
tacles (Patterson & Richards, unpublished). It
is also not clear if any of these morphological
abnormalities may be related to inbreeding de-
pression in this stock, and warrants further in-
vestigation.

The mechanism(s) by which genome plas-
ticity occurs in 10-R2 snails remains un-
known. In unrelated, ongoing studies in our

laboratory, however, we have identified sev-
eral expressed sequence tags (ESTs) show-
ing a high degree of sequence identity to
genes normally associated with transposable
elements, such as transposase and reverse
transcriptase (Miller et al., 2001; Raghavan, in
preparation). Future studies will compare the
frequency (copy-number) and gene activity of
some of these retrotransposon-like se-
quences between 10-R2 and BS-90 snails.

The BS-90 snail stock, which has been
used for most of the resistance-related genet-
ics conducted in our laboratory, was derived
from snails that were isolated by Paraense &
Correa (1963) in the field (Salvador, Brazil).
Since its arrival in our laboratory 12 years ago,
this stock has remained robust and stable,
with no detectable changes either in morphol-
ogy or fecundity, regardless of whether they
are kept as pedigree selfing snails or in a
group.

The inability to detect polymorphisms within
the BS-90 stock was surprising, as allozyme
polymorphisms were detected in a previous
study (Bandoni et al., 1995). It is possible that
additional variation at the level of the DNA may
be revealed using a more sensitive tool, such
as the examination of polymorphic micro-
satellite loci. The analysis of variations within
microsatellite loci of B. glabrata as a means of
assessing diversity among snails is steadily
gaining significance (Jones et al., 1999;
Malvares et al., 2000). In recent years, ge-
nome sequencing projects for several organ-
isms have been initiated. It is hoped that, as is
being done for the mosquito vectors of ma-
laria, a genome project may be forthcoming
for В. glabrata. In view of the present study, we
hope that the inherent intra-strain diversity that
exists within these snails will be taken into
consideration before a particular snail stock is
chosen as representative of this organism.

ACKNOWLEGEMENTS

We would like to thank Ms. Frances Barnes
and Dr. Charles Richards for helpful discus-
sions in preparation of this manuscript. This
work was funded in part by NIH grant Al-27777.
Ms. Ittiprasert was supported by the Royal
Golden Jubilee Ph.D. Program, Thailand Re-
search Fund. Susan Bandoni was supported
by a Mid-Career Summer Fellowship from
SUNY Geneseo.

108 ITTIPRASERT ЕТ AL.

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MALACOLOGIA, 2003, 45(1): 109-120

DEPTH EFFECTS ON ZEBRA MUSSEL REPRODUCTION

Paride Mantecca', Giovanni Vailati', Letizia Garibaldi? & Renato Bacchetta™

ABSTRACT

Male and female Dreissena polymorpha from 2 т depth in Lake Iseo, northern Italy,
spawned synchronously, and their gametogenic cycle followed the annual pattern previ-
ously observed in other Italian and European populations. Water temperature of 12°C and
phytoplankton blooms triggered spawning, and seasonal variation in gametogenesis was
related to photoperiod. Some mussels at 25 m depth always had active gonads, and re-
production continued all year, with no seasonal gametogenic phases. Hypolimnetic envi-
ronmental conditions, such as slight variation in water temperature, darkness and low food
availability, may cause this reproductive strategy. No evidence of hermaphroditism or modi-
fied sex ratios were noted at either depth. Gametogenesis and spawning ability of zebra
mussels in the hypolimnion must be reconsidered.

Key words: Dreissena polymorpha, reproduction, depth, gametogenesis, histology.

INTRODUCTION

For the last two centuries, the zebra mussel,
Dreissena polymorpha, has been steadily
spreading over Europe (Stanczykowska,
1977). In Italy, D. polymorpha was first de-
tected at the end of the 1960s in Lake Garda
(Giusti & Oppi, 1972), and from then on, it
reached all the great subalpine lakes, the major
northern rivers, and, recently, Lake Trasimeno
in central Italy. By the 1980s, it had also in-
vaded the major river systems and numerous
inland lakes throughout the northeastern USA
(Ram 8 McMahon, 1996).

Among the factors that favor such a wide
geographic distribution, high fecundity is one of
the most important (Sprung, 1989). But the
mechanisms that regulate zebra mussel re-
productive behavior are not well understood.
Many authors have investigated its reproduc-
tion, both in Europe (Tourari et al., 1988;
Borcherding, 1991; Neumann et al., 1993;
Bacchetta et al., 2001) and North America
(Haag & Garton, 1992; Gist et al., 1997), and all
have found an annual cycle involving a gamete
development phase in winter and early spring,
spawning events in late spring and summer,
followed by a gonad resting stage.

Temperature is considered the main environ-
mental factor that regulates both gametogen-

esis and the start of spawning in D. poly-
morpha (Borcherding, 1991; Bacchetta et al.,
2001), whereas such other factors as food
availability and phytoplankton bloom are т-
volved in regulating the number of reproductive
events (Gist et al., 1997) and the onset of
spawning (Ram & Nichols, 1993). In laboratory
experiments, the gonadal cycle 1$ not closely
associated with photoperiod variation (Borcher-
ding, 1995).

In lakes, D. polymorpha forms a characteris-
tic belt around the shores, usually covering the
littoral and upper sublittoral zones. The area
occupied by this species varies greatly, de-
pending on the littoral zone width and slope. In
shallow lakes, it can occupy both inshore areas
and mid-lake zones, whereas in deep lakes it
has been found to 30 m (Lake Geneva) and
55 m (Lake Constance) (Stanczykowska,
1977). Although zebra mussels are most com-
mon between 2 and 8 m, they have been re-
ported from the wave zone to 110 m (Claxton &
Mackie, 1998).

In Lake Iseo, also known as Sebino (45°39-
49’N, 2°21—30'W), D. polymorpha forms dense
populations along the wave zones and occurs
to at least 50 m, although density progressively
decreases from 5 т and is very low at 30-40
m. Lake Iseo lies in the foothills of the Alps and
reaches a maximum depth of 251 m. Limno-

‘Dipartimento di Biologia, Universita degli Studi di Milano, Sezione di Zoologia e Citologia 26 Via Celoria, 1-20133

Milan, Italy

“Dipartimento di Scienze dell'Ambiente e Territorio, Universita degli Studi di Milano-Bicocca 15 Via Emanueli,

1-20126 Milan, Italy
“Corresponding author: renato.bacchetta@unimi.it

110 MANTECCAET AL.

logically, Lake Iseo is classified as warm
monomictic, and like other deep lakes, it is dis-
tinctively holo-oligomictic. Because of its geo-
graphic location in the temperate belt and its
morphology, complete overturn is uncommon,
only occurring in particularly cold, windy win-
ters (Ambrosetti et al., 1992). The lake is strati-
fied for a long period during summer-autumn
and is eutrophic (Garibaldi et al., 1997). These
attributes, and observations that depth may
control bivalve reproductive cycles (Mackie,
1984), led us to investigate possible differences
in reproductive behavior between shallow and
deep water mussels. To this end, and because
only one work (Claxton 8 Mackie, 1998) has
considered depth variation in gametogenesis
and spawning, we used qualitative histological
methods to follow the D. polymorpha repro-
ductive cycle from two depths and over two
spawning seasons in a Lake Iseo population.

MATERIALS AND METHODS
Sampling

To investigate the timing of gonadal develop-
ment, 20 samples of mussels were collected
from a densely settled site in Lake Iseo, near
the town of Tavernola Bergamasca, from March
1999 to September 2000. Scuba divers
brought up rocks covered with D. polymorpha
from about 2 and 25 m depths, above and
below the summer thermocline. On each sam-
pling occasion, about 60 specimens of D. poly-
morpha > 18 mm in length were detached from
the rocks and fixed in aqueous Bouin's for his-
tological analysis.

Histological Methods

After nearly one week in Bouin's fluid, about
30 mussels were randomly selected to deter-
mine gonadal condition. Specimens were
washed overnight in running tap water, and
then their visceral sacs were separated from
the remaining tissues, dehydrated in an as-
cending alcohol series, and embedded in Bio-
Plast tissue embedding medium (melting point
57°C). Using a rotary microtome, samples
were cut in 7 um transverse sections at the
proximal, central and distal levels of the gonad
in order to detect any heterogeneous develop-
ment within the ovaries. About ten serial sec-
tions from each portion were placed on
microscope slides and dried overnight at 37°C.

The slides were then stained with Mayer's
Haemalaun (Merck), counterstained with alco-
holic Eosin (Merck), mounted in Eukitt (Kindler
GmbH, Freiburg), and observed using a light
microscope with calibrated eyepiece. A total of
1,163 mussels were histologically examined.

Maturity Index and Sex Ratio

The stage of gametogenic development for
both males and females was described using
a four-step qualitative evaluation, as given by
Gist et al. (1997): stage 0 = gonad inactive,
stage 1 = developing, stage 2 = prespawn,
stage 3 = postspawn. The Maturity Index (MI)
was calculated for both sexes as the mean
gonadal stage for all the specimens examined
on each sampling occasion.

Mussels were sexed by microscopic exami-
nation of the histological slides and sex ratios
were estimated for all the samples.

Environmental Parameters

From February 1999 to September 2000, we
recorded water temperature, food availability,
photoperiod, and water transparency. Water
temperature was measured at 1, 10, 20 and
30 m. To evaluate food availability, chlorophyll-a
(Chl-a) concentration was determined in inte-
grated samples from six depths from 0 to 20 m
collected in a Van Dorn sampling bottle. After
collection, the samples were placed in polyeth-
ylene bottles, put in thermic bags in the dark,
and transported to the laboratory. Once in the
laboratory, each sample was filtered through a
GF/F glass microfibre filter (WHATMAN, pore
size 0.45 um) and then stored at -4°C until pro-
cessing. Chl-a concentrations were measured
with a standard spectrophotometric method
(Lorenzen, 1967) after 24 h extraction with
90% acetone. Daylength on each sampling oc-
casion was calculated from sunrise and sunset
times published by the Italian Airforce Weather
Bureau and reported as light-minutes per day.
A Secchi disk was used to estimate water
transparency. The epilimnion of Lake Iseo was
taken to be the upper 15 m, representing 10% of
the lake’s volume; the hypolimnion was the wa-
ter mass below 15 m (Garibaldi et al., 1997).

Statistical Analysis
Ninety-five percent confidence intervals were

calculated for the MI at each date and depth.
Thus, when the confidence intervals of two Mis

ОЕРТН EFFECTS ОМ ZEBRA MUSSEL REPRODUCTION

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112 MANTECCAET AL.

overlapped, we considered there to be no sig-
nificant difference between them. The relation-
ship between the Mls of the two depths was
tested using Spearman's Rank correlation
analysis (Snedecor & Cochran, 1980), and a
pairwise Pearson correlation matrix (Sokal &
Rohlf, 1981) was used to determine the corre-
lation between the environmental parameters
and the gonadal cycle. À Chi square goodness-
of-fit test (Snedecor & Cochran, 1980) was
used to test the hypothesis of a 1:1 sex ratio
and to determine the relationships between the
presence of oocyte degeneration in females
from the two depths. Differences were consid-
ered significant at the 5% level (p < 0.05).

RESULTS

The number of specimens at each gonadal
stage and the sex ratio on each sampling occa-
sion and at each depth are given in Table 1.
The histological appearance of the ovary and
testis at different maturity stages is illustrated in
Figures 1 and 2.

Reproductive Behavior at 2 m

The gonadal phases of development, matura-
tion, spawning, and inactivity were temporally
well defined (Fig. 3). Spawning events occurred
in spring/summer starting in May, when the first
postspawned specimens were seen, and end-
ing in July (1999) or August (2000). During
these periods, samples in pre- and post-
spawned stages were observed and showed а
continuous process of maturation and gamete
release. The onset of spawning was always
preceded by a period with gonads in the devel-
oping stage, followed by a quick gamete matu-
ration phase, highly synchronized among
individuals. At the end of the spawning season,
testes of all males became inactive until the
end of September; October signaled the restart
of gametogenesis (Fig. 3). The resting stage of
ovaries is shorter than that of testes and in-
volved only a few mussels. At the time of the
last spawning, some females were already de-
veloping gonads, and by late September all
were (Fig. 3). The MI trend confirmed these
observations. During the reproductive season,

AS
A

FIG. 1. Histology of the Dreissena polymorpha ovary. A. Developing stage: developing oocytes (=)
attached to wall of acini; B. Prespawn stage: acini filled with mature oocytes; C. Postspawned stage:
enlarged acini contain only few large oocytes in lumen (=); D. Mixed stage: acini filled with oocytes
at different maturity levels.

ОЕРТН EFFECTS ОМ ZEBRA MUSSEL REPRODUCTION 113

FIG. 2. Histology of the Dreissena polymorpha testis. А. Developing stage: detail of an acinus filled
with undifferentiated cells, arranged radially; В. Prespawn stage: villi-like processes with spermatozoa
(*), spermatocytes on periphery (=); С. Postspawned stage: villi detached from underlying generative
layers (=), spermatozoa free in lumen (x); D. Mixed stage: detail of an acinus with radially arranged

cells, together with mature spermatozoa (*).

both males and females showed values
around 2 or greater, but in the other months the
Mis were around 1, with lower values for males.
Only just before the spawning period were the
male and female Mls similar (Fig. 4). The
Spearman’s Rank correlation analysis showed
highly significant synchrony between the MI
trends of the two sexes (r = 0.91; p < 0.01).

Reproductive Behavior at 25 m

Postspawned females were present in all
samples, and none with inactive ovaries was
ever observed (Fig. 3). The frequency of the
postspawn stage was always high, except in
winter when the developing stage was domi-
nant. Prespawned ovaries were recorded
throughout 1999, whereas except for one
specimen in January they were not observed in
2000. This indicates that during 1999, female
mussels had spawned successfully following
massive oocyte maturation. This did not hap-

pen in 2000, suggesting a slackening in repro-
duction. In March 2000, a new gametogenic
stage was observed in which the ovary condi-
tion could not be assigned to any of the four
stages. We call these morphologies “mixed
stages”, because the ovary acini were filled
with oocytes at different maturity levels. Even
though no maturity stage predominated, we
underline the spawning activity initiated by 12
mussels, by classifying them as postspawned
(Fig. 3, Table 1).

Prespawned, postspawned, and inactive
male mussels were almost always present.
Synchrony among individuals was low. All four
stages of testis maturity occurred contempora-
neously at seven different times (Fig. 3). Thus,
we hypothesize a tendency towards continu-
ous male gamete production and release even
if, contrary to the females, individual mussels
often underwent a brief inactive period. The fre-
quency of inactive males increased in the fall
and was maximal in November 1999. By winter

114 MANTECCAET AL.

and until March, reproductive activity was re-
duced, with most males in an inactive or devel-
oping stage.

The tendency towards continuous reproduc-
tion was confirmed by the presence of “mixed
stages” in males in May and June 1999, and in
February 2000. Their testes had acini with radi-
ally arranged cells, typical of the developing
stage, together with mature sperm, typical of
the postspawned stage, but without the usual
detachment between generative layers (Fig. 2).
These, too, were classified as postspawned.

In both sexes, MI values were almost always
high, particularly in females, for which they ex-
ceeded 2 except in January and February 2000
(Fig. 4). The MI trends of the two sexes were
synchronous (Spearman г = 0.62; р < 0.01).

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# Stage 0 | Stag

Significantly, more mussels from 25 m depth
showed oocyte degeneration than those from 2
т (y? = 25.73, р < 0.01). Specimens with this
histopathological condition were frequently ob-
served in the 25 m mussels, while it was de-
tected only in May and June 1999 in the 2 m
mussels (Table 1). Degenerating oocytes pre-
sented a vacuolated apical portion that de-
tached from the lower cytoplasm and
accumulated in the lumen of the acini. At times,
the oocytes were completely disgregated.

Sex Ratio
Of the 562 mussels from the 2 m depth, 294

were females and 268 males, whereas of 601
mussels from 25 m, 322 were females and

-2m males

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Мау 07

FIG. 3. Gametogenic cycles of adult mussels from 2 (upper panels) and 25 т in depth (lower panels).
Histograms show relative frequency of the four gonadal stages for each sampling. Stage 0 = gonad
inactive; stage 1 = developing; stage 2 = prespawn; stage 3 = postspawn.

DEPTH EFFECTS ОМ ZEBRA MUSSEL REPRODUCTION 115

279 males (Table 1). At both depths, there was
no significant difference in the total number of
males and females (y? = 1.20, р > 0.05 for
mussels from 2 m; y? = 3.07, p > 0.05 for
mussels from 25 m). The monthly compari-
sons showed no statistical difference in male
and female numbers, except in March 2000 for
the 25 m deep mussels (x? = 3.90, р < 0.05).
No evidence of hermaphroditism was found.

————— =

N
|

Gonadal stage

Environmental Parameters

At 2 m, water temperature showed the same
trend in both years, with a rapid increase in
spring-summer, reaching the highest values in
August 1999 (22.6°C) and July 2000 (23.0°С),
with a clear decrease in autumn to a minimum
of 6.3°C in winter (Fig. 5). At the beginning of
May in both years, temperature rose above

Females

o +
Mar 28 — _ TE

Nov 14 MM _ _—_____

Jan 09 MM

Feb 20
Mar 25
Jun 04 |
Jul 01
Jul 29

Gonadal stage

4

Mar 28

Nov 14 mi z

Aug 19 == |
Sep 23 |

FIG. 4. Maturity Index values from the two depths. Data are presented as the mean gonadal stage
+ 95% c.i. Black bars = mussels from 2 т depth. Shaded bars = mussels from 25 т depth.

MANTECCAET AL.

116

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FIG. 5. Environmental parameters. Water temperature (upper panel), chlorophyll-a

concentration (mid panel), transparency (lower panel).

ОЕРТН EFFECTS ОМ ZEBRA MUSSEL REPRODUCTION 117.

12.0°C, which has been considered the tem-
perature that triggers spawning (Borcherding,
1991). At 20 m, the temperature stayed below
10.0°C for most of the year and varied mini-
mally, reaching 12.0°C only in November 1999
and September 2000. At 30 m, the temperature
curve flattened even more and ranged only be-
tween 6.4°C and 8.4°C.

Chl-a concentrations showed two peaks per
year, one in April before the onset of spawning
(especially high in 2000), and one in summer.
During the 2000 spring bloom, the highest Chl-
a concentration value recorded was 32 ug/L.
Minimum values occurred in winter, except for
the June 2000 sample, when the Chl-a was 2
ug/L (Fig. 5).

Transparency showed an opposite trend
from Chl-a, indicating that algae were the main
component of suspended matter. Minimum val-
ues were observed in August, concurrent with
the summer algal bloom, whereas the highest
values were in February. However, transpar-
ency never exceeded 11.2 m, indicating that
25 m mussels were always in the dark.

DISCUSSION

The reproductive cycle of zebra mussels in
northern Italian waters is annual, as in the rest
of Europe (Bacchetta et al., 2001). This was
confirmed for the 2 m mussels from Lake Iseo,
where the spawning period started in May and
ended in July, in both 1999 and 2000. This pe-
riod followed well-synchronized phases of ga-
mete development and maturation. Oogenesis
restarted soon after the end of the last spawn-
ing, while spermatogenesis began later. In gen-
eral, males and females showed similar
reproductive patterns, culminating in synchro-
nous spawning, except for the longer resting
gonad period in males than in females. In fact,
females had empty gonads only in a few
cases, indicating that they began gamete pro-
duction for the next season just before the end
of the late spawning events.

These results corroborate those from other
Italian water bodies (Bacchetta et al., 2001),
suggesting that in shallow waters a similar
course of reproductive events may be general-
ized for all subalpine populations of D. poly-
morpha.

Reproductive behavior at 25 m differed
greatly from that at 2 m. Intense spawning ac-
tivity occurred throughout the study period, ex-
cept during winter months, when reproduction

slackened. The constant presence of spawning
mussels indicates a much reduced annual re-
productive pattern for D. polymorpha at this
depth. The female MI trend confirmed this (Fig.
4), and MI values always exceeded 2, except in
January and February 2000. In these two
months, the MI was still over 1, which agreed
with the reduction in spawning activity, in con-
trast to clear reproductive inactivity.

The lack of prespawn females at 25 m in
2000 may indicate different reproductive be-
havior in the two years. By March 2000 females
may be unable to mature large numbers of oo-
cytes simultaneously, modifying the reproduc-
tive event from an “explosive” phenomenon to a
slow and continuous release of gametes. But
why such behavior? At 25 m, environmental
conditions may prevent a regular annual
course of gamete maturation and spawning.
Thus, reproductive events may only happen
once in two or more years. The high frequency
of mussels with developing ovaries in January
and February 2000, not followed by prespawn
specimens, support this assertion. Moreover,
the “mixed stages” observed in females after
these months strengthen this hypothesis.

Regarding males, their reproductive pattern
was similar in the two years and, as in the fe-
males, differed from that of those living near the
surface (Fig. 3). The presence of all four stages
of testis maturity in many samples indicated a
very low level of synchronization, but suggests
a tendency towards continuous reproduction.
Nevertheless, the course of the male gonadal
cycle at 25 m also differed from that of the fe-
males, mainly in the percentage of inactive go-
nads observed in males all year round. The
difference between sexes may be a result of the
major energy demand to mature oocytes,
which at 25 m, could be a limiting factor.

The loss of seasonality in deep water may
result from the environmental conditions in the
hypolimnion, including small variation in water
temperature, darkness, and low food availabil-
ity (perhaps because of the stable deep water
environment, contrasting with the shallow wa-
ter wave movement). In the epilimnion, many
environmental factors contribute to regulate
gametogenesis and spawning in bivalves. The
most important are usually considered to be
water temperature, food availability, photope-
riod, and depth (Giese 8 Pearse, 1979; Mackie,
1984). It is commonly held that for D. poly-
morpha water temperature is the main factor
involved in triggering spawning, whereas food
availability plays a role in determining the num-

118 MANTECCAET AL.

ber and intensity of reproductive events
(Borcherding, 1991; Ram et al., 1996). А|-
though photoperiod does not affect gametoge-
nesis in laboratory experiments (Borcherding,
1995), it has well-recognized regulatory prop-
erties in many other invertebrate species
(Bohlken 8 Joosse, 1982; Olive & Pillai, 1983;
Foster & Hodgson, 1995), including bivalves
(Giese, 1959). With regard to depth, the tem-
poral aspects of breeding strategy vary with
depth, although it is often difficult to separate
the effect of temperature and depth (Mackie,
1984; Claxton & Mackie, 1998).

In many European, Russian, and American
populations, breeding starts when water tem-
perature exceeds 12°C, even if other environ-
mental conditions differ markedly according to
the populations examined. Stanczykowska
(1977) indicated 15°C as the temperature
when spawning starts, whereas Sprung
(1987) stated that a range of 12-14°C is suit-
able for spawning, with an optimum around
18°C. Tourari et al. (1988) observed that ga-
mete spawning occurs when temperature
reaches 16-17°C, whereas Borcherding
(1991) and Neumann et al. (1993) reported
the onset when water temperature exceeds
12°C. In North America, Haag & Garton (1992)
found that spawning started between 22°C
and 23°C, even if in the same waterbody,
Lake Erie, disjunct populations start spawning
at different temperatures (Nichols, 1996).
McMahon (1996) indicated that spawning can
begin at 12°C, but is maximized around 17-
18°C, whereas Gist et al. (1997) reported the
onset of the breeding season when water tem-
perature is above 20°C.

Our results indicate that 2 т deep mussels
started spawning in May when water tempera-
ture reached 13.2°C (1999) and 12.1°C
(2000), thus triggering the event, confirming
the observations of Borcherding (1991) and
Neumann et al. (1993) in central Europe, and
of Bacchetta et al. (2001) in other north Italian
water bodies.

The lack of data about the relationship be-
tween photoperiod and the D. polymorpha re-
productive cycle does not permit speculation
about the role of this factor on the onset of
gamete deposition. However, previous data by
Bacchetta et al. (2001) show delayed
spawnings in a river population compared to a
lacustrine population, despite the same light
conditions. In fact, while in Lake Como ga-
mete release started at the beginning of May,
when water temperature reached 13°C, in the

Adda River this threshold was reached only
two weeks later.

Photoperiod played a major role in regulating
gametogenic phases, because in 2 m deep
mussels, this factor was correlated with Mls
trend (r = 0.75; p < 0.001 for females, r = 0.69;
p < 0.01 for males).

Even if Chl-a concentration was not signifi-
cantly related to the reproductive cycle, we
suggest that the two Chl-a peaks preceding
the spawning season may be involved in
spawning induction by signaling the trophically
advantageous conditions for success of the
subsequent larval developmental period. In D.
polymorpha, phytoplankton blooms, with their
associated chemicals, may be the first in-
ducer of gamete release, followed by further
chemical stimulus associated with gametes
that induces spawning in the opposite sex
(Ram & Nichols, 1993). In conclusion, the D.
polymorpha gametogenic events strictly fol-
lowed the seasonal variations of photoperiod,
while spawning events were mainly regulated
by food availability and water temperature.

At 25 m depth, below the thermocline, this
reproductive pattern vanished. Water tem-
perature never exceeded the threshold of 12-
13°C for the onset of spawning, but stayed for
most of the year below 10°C, the value that
has been considered the minimum spawning
and fertilization temperature for D. poly-
morpha (Sprung, 1987). We can say nothing
about fertilization and larval developmental
success at this depth, but both males and fe-
males matured and released their gametes,
as evidenced by the presence of spawning
specimens, even when temperature was 6-
7°C. Dreissena polymorpha also reproduces
at low temperatures in other regions: in Lake
Constance, at 4.5-5.5°C, and in Lake Grosser
Plóner at 2.5°С (Walz, 1978). Nichols (1996)
reported that zebra mussel larvae were col-
lected at temperatures below 5°C, and that D.
polymorpha is theoretically able to produce
larvae at temperatures below 10°C.

Another dreissenid, the quagga mussel,
Dreissena bugensis, has been reported to
spawn at low temperatures in deep waters
(Roe & Maclsaac, 1997). These authors ob-
served gonadal development and spawning at
4.8°C in Lake Erie, and Claxton & Mackie
(1998) found that D. bugensis and D.
bugensis “profunda” spawned in the same
lake at 9-10°C.

While it has now been shown that D.
bugensis spawns in the hypolimnion at tem-

ОЕРТН EFFECTS ОМ ZEBRA MUSSEL REPRODUCTION 119

peratures below the minimum dreissenid
spawning and fertilization temperature
(Claxton & Mackie, 1998), our findings suggest
that this minimum must also be reconsidered
for D. polymorpha, and that the zebra mussel
is able to reproduce in a hypolimnion environ-
ment. This contrasts with the findings of
Claxton & Mackie (1998), who found no game-
togenic development or spawning in hypo-
limnetic Lake Erie zebra mussels.

In our study, significantly more females with
degenerating oocytes were observed at 25 m
than at 2 т (y? = 25.73, р < 0.01). Oocyte de-
generation in D. polymorpha is triggered by
several cues, among them, low temperature
and low food availability (Borcherding, 1995).
The presence of degenerating oocytes in the
ovaries, together with detritus clearly visible in
the lumen of the acini, indicates intense recy-
cling activity by the gonad in disadvantageous
environmental conditions. This has been previ-
ously observed т D. polymorpha (Bielefeld,
1991), as well as in other mollusks, for ex-
ample, Mytilus edulis (Pipe, 1987) and
Mactra veneriformis (Chung & Ryou, 2000).
The hermaphroditic state in gonochoristic
bivalves is also determined by environmental
factors (Mackie, 1984), but even ifthe particu-
lar hypolimnetic conditions in Lake Iseo
caused oocyte degeneration and altered
spawning activity, they did not seem to influ-
ence sex determination. In fact, the sex ratio,
even in deep waters, was not significantly dif-
ferent from 1:1, and no hermaphrodites were
seen, contrary to Antheunisse (1963), who
found 4% hermaphroditic mussels in the
Amstel River.

In conclusion, the reproductive behavior of
25 m deep mussels differed significantly from
that of those in shallow water, where an annual
pattern was confirmed. In deep water, at the
low limit of the thermal discontinuity, reproduc-
tion continued all year, without seasonal
changes of gametogenic phases. Elsewhere,
reproduction can continue throughout the year,
depending mainly on the geographic locality of
a given water body and its thermal condition,
as in warmer reservoirs, where the reproduc-
tive period is longer (Stanczykowska, 1977).
Even in the hypolimnetic waters of Lake Iseo,
where temperatures were always low, zebra
mussels were observed to spawn continu-
ously, suggesting that variations in water tem-
perature are more important in regulating the
timing of reproduction than absolute tempera-
ture.

ACKNOWLEDGEMENTS

Our field samplings were assisted by Scuba
divers of the Federazione Italiana Attività
Subacquee. We thank Drs. Lorenzo Colombi
and Maddalena Pesenti for their help. We are
grateful to the reviewers who improved the
clarity of the manuscript.

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Revised ms. accepted 6 February 2003

MALACOLOGIA, 2003, 45(1): 121-124

THE EGG OF OXYCHILUS (DROUETIA) ATLANTICUS
(PULMONATA: ZONITIDAE):

SURFACE STRUCTURE AND CARBOHYDRATE COMPOSITION

Armindo S. Rodrigues", Regina T. Cunha! 8 Benjamin J. Gomez?

ABSTRACT

Oxychilus atlanticus is an oviparous species with eggs of the heavily calcified type. The
eggshell is formed of calcite crystals and is endowed with large amounts of neutral
polysaccharides, galactogen and glycogen, providing nutritive reserves for embryo
development. The egg is surrounded by mucopolysaccharides as it passes along the
spermoviduct. This component probably acts as a mechanical support element, as well as
a chelation promoter of calcium ions during eggshell construction.

Key words: oviparous snail; calcite crystals; galactogen; glycogen; mucopolysaccharides.

INTRODUCTION

Apart from the works of Bayne (1966, 1968),
Tompa (1974, 1976, 1979), Baur (1994), Baur
& Baur (1998), and Heller (2001), little is known
about the structure and composition of pulmo-
nate eggs, and no information exists about egg
formation inside the spermoviduct of oviparous
species. This could be due to the fact that, in
Oviparous species, such as the stylom-
matophoran land snails and slugs, the eggs are
laid as they are made inside the spermoviduct
and distal female genital ducts (Tompa, 1979).

The surface structure, composition and func-
tion of molluscan eggs have received little atten-
tion. In general, the egg is known to protect the
embryo against adverse environmental effects,
but it is also considered to act as a calcium re-
serve for itself and the newly hatched juvenile
(Fournié & Chétail, 1982a). Baur & Baur (1998)
consider that eggs are decisive for the survival
of the offspring in invertebrates, especially when
the species do not have post-laying egg care.

After studying a great number of eggs from
different gastropods, Hall & Taylor (1971) con-
cluded that the eggshell is generally made by
calcite or aragonite crystals, although they re-
fer to a new form of calcium carbonate crys-
tals, vaterite, in the eggshell of four species of
Ampullaria Lamarck, 1799, for the first time. Ac-
cording to Tompa (1976), 36 of 65 stylom-

matophoran families have eggshells made of
calcium carbonate, that is, in calcareous eggs,
commonly in the form of calcite. Tompa (1974)
suggests that the reason for the common pres-
ence of aragonite in the body shell is due to its
higher resistance to the abrasion by soil particles,
whereas calcite, frequently found in the eggshell,
involves less investment because it occupies
more space per mole of CaCO, secreted.

Tompa (1976) classified the eggs of the
Stylommatophora into three types — heavily cal-
cified (Cepaea nemoralis Linnaeus 1758), partly
calcified, and uncalcified (Limacidae), accord-
ing to the degree of calcification. In the same
work, the size of the egg and its surface ultra-
structure were considered to be of taxonomic
value, even between closely related species.

Concerning the resources of the eggs, Heller
(2001) considers that they are rich in energy
and nutrients, including proteins, mucopolysac-
charides and calcium.

This study describes the carbohydrate com-
position of the egg of Oxychilus (Drouetia)
atlanticus (Morelet & Drouét 1857) (Pulmonata
Zonitidae) during its formation inside the
spermoviduct, as well as the structure and com-
position of its surface after being laid.

The aim of this work is to contribute to the
knowledge of the reproductive biology of an
endemic species from Sao Miguel Island,
Асогез.

‘Departamento de Biologia, Universidade dos Açores, 9500 Ponta Delgada, Portugal
“Departamento de Zoologia y Dinamica Celular Animal, Facultad de Ciencias, Universidad del Pais Vasco, Apdo

644, 48080 Bilbao, Spain
“Corresponding author: rodrigues@notes.uac.pt

122 RODRIGUES ET AL.

MATERIALS AND METHODS

Specimens of O. atlanticus were collected at
Abelheira, 3 km north of Ponta Delgada, be-
tween June and September. The species
reaches maturity between June and November,
with a shell diameter of 7 mm (Rodrigues et
al., 1998). From over 200 specimens analysed,
only two adults had eggs inside the sperm-
oviduct, observable through the translucent
shell. Therefore, only these two specimens
were used for histochemical tests.

Genitalia with eggs were dissected and fixed
in Baker's formol (Culling, 1974), embedded in
paraffin, and sectioned at 7 ит thickness. Sec-
tions were routinely processed for light micros-
copy and stained using various histological and
histochemical methods.

The following histochemical methods were
carried out: The Periodic acid-Schiff (P.A.S.)
technique was used as a general method for
identifying neutral carbohydrates (Culling, 1974).
Acetylation, followed by saponification and dia-

FIG. 1. A. Cross section of a fractured egg of
Oxychilus (Drouetia) atlanticus. Arrowheads
indicate the outer surface; arrows show the inner
border (scale bar = 10 um). В. Surface view of
the egg showing the large calcite crystals (scale
bar = 10 um). C. Surface view of the egg showing
individual calcite crystals (scale bar = 1 pm).

stase treatment, were used as controls (Martoja
& Martoja-Pierson, 1970). Alcian blue staining
was used at pH = 0.5 to stain strongly sulphated
mucosubstances and at pH = 2.5 for carboxy-
lated and weakly sulphated mucosubstances
(Martoja & Martoja-Pierson, 1970). Best's car-
mine was used to identify glycogen deposits,
with previous digestion with diastase as a con-
trol. Best's carmine was also used to detect
the presence of galactogen, according to
Grainger & Shillitoe (1952).

In order to study the eggshell structure, indi-
viduals of O. atlanticus were collected from the
field and laid eggs under laboratory conditions
from November 1997 to January 1998. The di-
ameter of 40 eggs was measured under a ste-
reomicroscope with the aid of a camera lucida.
To perform scanning electron microscope
(SEM) observations, the shells of ten eggs were
broken and their fragments placed in a cham-
ber with silica gel for two days of dehydration.
The material was then mounted on specimen-
stubs, coated with carbon and gold-palladium
(60-40%) in a vacuum evaporator (JEE 400)
for observation with a JEOL SEM (JSM 5410)
at 15 kV or 25 kV.

RESULTS

The eggs of O. atlanticus are 1.5 + 0.05 mm
in diameter, with a hard and brittle surface, and
a thickness of 34.1 + 1.9 um (Fig. 1A). The egg
surface consists of a continuous layer of cal-
cium carbonate, which reacts with hydrochlo-
ric acid. The calcium carbonate develops as
geometric forms (Fig. 1B, C), with a symmetry
that fits into a hexagonal system, typical of cal-
cite (Almeida, personal communication). Sol-
ids of calcite show a perfect cleavage in three
directions that may originate rhombohedral hab-
its (Dana, 1969), as shown in Figure 1B and C.

The histochemical tests reveal the presence
of an organic layer, 5 um in thickness, over the
calcified shell, mainly composed of acid muco-
polysaccharides and some neutral polysaccha-
rides, which react strongly with alcian blue
solution (pH 2.5) and Best's carmine, respec-
tively. The inner shell membranes between the
calcified shell and the perivitelline fluid show a
high positivity for P.A.S. and a moderate posi-
tivity for Best’s carmine, revealing the presence
of neutral carbohydrates. The histochemical
composition of this layer is very similar to that
of the perivitelline fluid, the most important fea-
ture of which is a complete maintenance of the

THE ЕСС ОЕ OXYCHILUS ATLANTICUS

123

TABLE 1. Staining reactions of О. atlanticus eggs pulled from the spermoviduct. P.A.S. = Periodic
acid-Schiff; Acetyl./P.A.S. = control Acetylation/P.A.S.; Acetyl./S/P.A.S. = control Acetylation/saponifi-
cation/P.A.S.; gr. = granules; - = no reaction; + = weak positivity; + = moderate positivity; ++ = high

positivity ; +++ = very intense positivity.

Carbohydrates Outer Calcified Inner shell Perivitelline
Stain detected shell layer shell membranes fluid Vitellus
Alcian blue pH 0.5 Strongly sulphated + - - gr. + - -
mucosubstances
Alcian blue pH 2.5 Carboxylated and ++ - + - -
weakly sulphated
mucosubstances
Best's carmine Glycogen and + - + - +
galactogen
Control + - + + y
PAS. Neutral + = ++ gr. ++ gr. ++
carbohydrates
Acetyl./P.A.S. 2 E Е - -
Acetyl./S/P.A.S. + 2 + gr. ++ gr. ++
P.A.S.-without _ L = - я
oxidation

reaction intensity with Best's carmine test and
control, indicating the presence of galactogen.
In the vitellus, the observed polysaccharide is
glycogen because it shows a high positivity for
Schiffs reagent and Best's carmine, but it is
digested by diastase (Table 1).

DISCUSSION

Oxychilus atlanticus is an oviparous species
in which a sequential synthesis and release of
each egg occurs, with a clutch consisting of 5—
6 eggs (Rodrigues & Cunha, unpubl. data). The
difficulty in finding gravid individuals in O.
atlanticus may be related to its oviparity, as
stated by Tompa (1979) for other oviparous
species. He reports that, in thousands of speci-
mens analysed, not a single individual from the
oviparous species was found to be gravid, in
contrast with the ovoviviparous and viviparous
species.

The eggshells of O. atlanticus bear a com-
pact and brittle construction of calcite crystals,
which we classify as of the heavily calcified type
(Tompa, 1976; 1984). This strong structure may
function as a protection against predators and
desiccation, in addition to its role in the support
of the internal components (Bayne, 1966). Fur-

thermore, the eggshell may also function as a
calcium reserve to supply the developing em-
bryo with enough calcium to form its embry-
onic shell (Heller, 2001).

In the egg of O. atlanticus, mucopolysaccha-
rides prevail in the vicinity of the shell, whereas
neutral polysaccharides are the main elements
internally.

According to Grainger & Shillitoe (1952), dia-
stase digests the polysaccharide glycogen, but
it is unable to digest galactogen. Thus, this
methodology is suitable for detecting the pres-
ence of galactogen, indicating that the perivi-
telline fluid and vitellus mainly contain
galactogen and glycogen, respectively. Wijsman
& Wijck-Batenburg (1987) mentioned that
galactogen and proteins are the major compo-
nents of the egg of Lymnaea stagnalis
Linnaeus, 1758. According to Bayne (1968),
neutral polysaccharides, as well as proteins,
are abundant in the perivitelline fluid, whereas
mucopolysaccharides occur in the external lay-
ers of the egg; the neutral polysaccharides and
proteins are mainly involved with the nutrition
of the embryo, and the mucopolysaccharides
are more involved in mechanical support. We
suggest that the mucopolysaccharides found
between the eggshell and the spermoviduct
epithelium may also act in the chelation of the

124 RODRIGUES ET AL.

calcium ions (Chétail et al., 1982; Fournié &
Chétail, 1982b), promoting eggshell mineraliza-
tion.

ACKOWLEDGEMENTS

This work was supported by the project
Praxis/2/2.1/B1A/169/94-Biodiversidade, and
Centro de Investigaçäo em Recursos Naturais,
University of Azores. We thank Jorge Medeiros
for his support with scanning electron micros-
copy, Helena Almeida for her help on the deter-
mination of the structure of calcium carbonate
crystals, Robert Cameron for helpful discus-
sions, and Brian Morton for his help and com-
ments on the first drafts of this paper.

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Técnicas de histología animal, 1st ed.
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ТОМРА, А. $., 1974, The structure of calcareous
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TOMPA, А. S., 1976, A comparative study of the
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Revised ms. accepted 19 December 2002

MALACOLOGIA, 2003, 45(1): 125-132

INFLUENCE OF INORGANIC COMPOUNDS ON FOOD SELECTION
BY THE BROWN GARDEN SNAIL CORNU ASPERSUM (MÜLLER)
(GASTROPODA: PULMONATA)

Laure Chevalier, Martine Le Coz-Bouhnik? & Maryvonne Charrier™

ABSTRACT

Cornu aspersum's (зупопут: Helix aspersa) ingestion rates were determined for 19 wild
plant species and revealed strong feeding preferences. The plant species were classified
according to their palatability. The inorganic contents of 3 appetent and 3 inappetent
plants, analysed by ICP-MS, revealed significant differences, the appetent species being
poorer in zinc and richer in calcium than inappetent ones.

Trials facing isolated snails with a test solution against distilled water as control were
performed. Farm snails previously fed with a calcium-rich food did not react differently to-
wards two solutions of different calcium carbonate concentrations, whereas wild snails fed
with a calcium-lacking food significantly preferred a 1 mg x ml * calcium carbonate solution.
Farm snails were significantly repelled by a 13 mg x ml ‘zinc sulphate solution.

The detection of inorganic compounds by snails and its possible influence on feeding
regulation is discussed.

Key words: Cornu aspersum, Helix aspersa, food selection, inorganic compounds, cal-

cium, zinc

INTRODUCTION

Chemoreception via the tentacles, lips and
foot (Croll, 1983; Kohn, 1983; Chase, 2002)
enables a snail to direct itself towards the food
source (olfaction), to analyse it by touch and
taste, and to determine whether it is a suitable
food or not. This analysis requires the interven-
tion of a specialized olfactory neuronal system
(Chase, 2002) and learning capacities linked
with previous feeding history (Balaban, 1993;
Desbuquois & Daguzan, 1995; Ungless,
1998). Associative learning and post-ingestive
effects may also modify future food selection
(Gelperin, 1975; Chevalier et al., 2000).

Several factors may influence the feeding
choices of terrestrial molluscs. Secondary
metabolites produced by plants, such as terpe-
noids (Gouyon et al., 1983; Linhart & Thomp-
son, 1995), glucosinolates (Glen et al., 1990)
or alkaloids (Speiser et al., 1992; Chevalier et
al., 2000), were recognized as deterrent for
slugs and snails, and physical features of
plants such as height (Rathcke, 1985) or tex-

ture (Wadham & Wynn Parry, 1981) may also
constitute a barrier against feeding activity.
Some inorganic compounds, such as copper,
calcium and trace metals, are necessary nutri-
ents for molluscs (Johannsen & Solhoy, 2001),
whereas chrome, nickel or selenium are not
essential (Simkiss & Mason, 1983). Because
of seasonal needs for calcium, snails store
this mineral mainly as calcium carbonate and
possess high reallocation capacities (Tompa &
Wilbur, 1977; Fournié & Chétail, 1984). As cal-
cium phosphate, it constitutes the mineral rings
of the spherites located in the calcium (crypt)
cells of the digestive gland (Almendros &
Porcel, 1992) that have a heavy metal detoxify-
ing function (Beeby & Richmond, 1988).

Few studies have dealt with the role that inor-
ganic compounds might play in snail nutrition.
Williamson & Cameron (1976) and Wadham &
Wynn Parry (1981) hypothesized that the silica
content of grasses may be responsible for
their rejection by the snail Cepaea nemoralis
and the slug Deroceras reticulatum, respec-
tively. High levels of cadmium, copper and zinc

'U.M.R. C.N.R.S. 6553 "EcoBio", Université de Rennes 1, 263 Avenue du Général Leclerc, 35042 Rennes Cedex,

France

“Géosciences Rennes, UMR C.N.R.S. 6118, Université de Rennes 1, 263 Avenue du Général Leclerc, 35042 Rennes

Cedex, France

*Corresponding author: maryvonne.charrier@univ-rennes1.fr

126 CHEVALIER ET AL.

were found to reduce the food intake of snails
and be toxic (Laskowski & Hopkin, 1996;
Gomot-de Vaufleury, 2000). Iglesias &
Castillejo (1999) have suggested that the
appetence of Urtica dioica for Cornu
aspersum was linked with its richness in cal-
cium.

The aims of this study were to investigate
the plant preferences of С. aspersum and re-
late these preferences to the mineral compo-
sition of six selected food plants. Trials were
then performed to observe the snail's gusta-
tory sensitivity when faced with the most im-
portant minerals revealed by the analysis.

MATERIALS AND METHODS
Consumption of Plants

Samples of 19 plant species were collected
in two coastal sites where populations of C.
aspersum were well established. Three plant
species were common to the two sites.
Those sites were in the salt-pans of
Guérande, France, and in the polders of the
Mont-Saint-Michel bay, where we determined
previously the diet of C. aspersum (Chevalier
et al., 2001). Twenty-two groups of farmed
snails, each of 15 snails of equivalent sizes
(32.4 + 1.0 mm) and weight (11.0 + 1 g) were
used (ANOVA on sizes: df = 21, F= 0.218, р =
0.99). These snails were considered as naive
as they originated from a snail farm, had al-
ways been fed with a cereal-based flour and
had never experienced any fresh plant mate-
rial. They were starved two days before the
experiment.

Snails were kept individually in plastic boxes
and received a fresh sample of one plant spe-
cies weighing around 2 g. The relationships
between fresh and dry plant weights were es-
tablished with plant samples oven-dried to
constant mass. After 48 h, plant remains were
weighed, dried and the dry weight of plant in-
gested calculated. It was then divided by the
fresh weight of the snail to obtain ingestion
rates. Each snail was used once and each
plant was tested against 15 snails.

ICP-MS Analysis

Six of the 19 plant species studied were
chosen according to their strong appetence
or inappetence: three from Guérande: Picris
echioides and Carduus tenuifloris (Astera-

ceae) and Beta maritima (Salsolaceae), and
three from the Mont Saint-Michel polders:
Urtica dioica (Urticaceae), Brachythecium
rutabulum (Bryophyta Brachytheciaceae), and
Elytrigia repens (Poaceae).

Leaf samples of those six plants were col-
lected in the field, rapidly frozen in liquid nitro-
gen, and dried by lyophilisation. Each
powdered sample (nominally 100 mg) was di-
gested in а screw-top Teflon® bomb (Savillex®,
USA) on a hot-plate by three treatments using:
(1) nitric acid (HNO, 14N sub-boiling grade);
(2) hydrogen peroxide (Analytical reagent
30%); (3) nitric acid together with hydro-
xyfluoric acid (HF 29N sub-boiling grade). The
sample was dissolved in HNO, and evapo-
rated to dryness. The completely mineralised
samples were dissolved in 100ml of HNO,
0.37N (HNO, 14N diluted with ultrapure deion-
ized water Milli-Q® system).

Samples spiked with 100 ppb of indium (In-
ternal standard), were analyzed with an Induc-
tively Coupled Plasma Mass Spectrometer
(ICP-MS Agilent-Technologie® model HP4500)
(Table 1). Calibrations were determined using
synthetic multi-element solutions. To quantify
the accuracy of our ICP-MS analyses, we
used the NIST standard SRM 1573a (Standard
Reference Material of Tomato leaves — National
Institute of Standards and Technology, USA),
and a reagent blank.

Calcium Carbonate and Zinc Sulphate Experi-
ments

Six batches of 35 to 50 adult snails of
equivalent sizes (30.6 + 1.0 mm) and weights

TABLE 1: ICP-MS operating conditions

Instrument parameters:

Plasma gas 15L/min
Auxiliary gas 1.0L/min
Carrier gas 1.13L/min
Nebulizer Cross flow
Spray chamber Scott
T° spray chamber 24€
CeO'/Ce' 0.6%
Ce*'/Ce' 1%
Data acquisition parameters:

Quantitative analysis 3s/mass
Repetition 2

INORGANIC COMPOUNDS IN SNAIL FEEDING CHOICES

(8.9 + 1.0 g) (АМОМА, df = 59, Е = 0.01, р =
0.991) were used. These snails, originating
from different clutches were maintained at
20°C, 80% relative humidity under a 12 h Папе
12 h dark period. Two days before the experi-
ment, they were starved in order to enhance
their feeding motivation. 30 min before the test,
their locomotion was stimulated by a gentle
spray of tepid distilled water. At the beginning of
the experiment, carried out during the
scotophase, they were individually placed at
the bottom of a plastic box (11.5 x 8.5 x 4.5 cm)
between two nylon gauzes of 24 ст? each.
One gauze was impregnated with 2 ml of dis-
tilled water, the other with 2 ml of the test solu-
tion. Boxes were covered with a transparent
glass sheet. They were observed in dim-light
for 30 min and the time spent on the gauzes
was checked. Time was recorded when snails
remained active, not retracted into their shells,
head and lips in contact with the gauze.
Among the six batches, two comprised naïve
snails reared with a specific meal rich in cal-
cium carbonate (20% CaCO.) (“farm snails”).
Two other batches comprised wild snails from
Rennes, France, fed with lettuce for two
months before the experiment (“wild snails”).
Lettuce is well known as attractive to snails
and also poor in calcium, between 300 and 500
mg x kg * (Feinberg et al., 1991; Couplan,
1998). A batch of farm snails and a batch of
wild snails were tested with a 0.1 mg x ml?

127

CaCO, solution, while the two others were
tested with a 1 mg x ml" CaCO, solution.

One of the two remaining batches of farm
snails was tested with a 1 mg x ml * ZnSO,
solution, that roughly corresponds to the zinc
concentration of lettuce (Dallinger & Wieser,
1984) and the second with a 13 mg x ml"
ZnSO, solution, a value higher than the toxic
threshold of zinc determined by Gomot-de
Vaufleury (2000) in the food of C. aspersum.
This high value compensated for the short
time of exposure (30 min), the objective being
to demonstrate the perception of this metal by
the snail.

Statistical Analysis

Differences in ingestion rates among plant
species were evaluated using ANOVA followed
by a Fisher's PLSD test when a significant dif-
ference was detected. The normality of the
data was checked by the Wilk-Shapiro
method.

ICP-MS data from Guérande and Mont-Saint-
Michel, and data from preferred and rejected
plants, were compared by a Student t-test.

The trials data were analyzed using a
Wilcoxon, Mann 8 Whitney test for paired
samples. The total activities of the snails —
time spent on the test gauze + time spent on
the control — were compared between different
concentrations by a Student t-test.

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FIG. 1. Laboratory feeding trials showing the ingestion of 19 plant species by 15 adult Cornu
aspersum. Values are means + S.E. G = saltpans of Guérande, BM = polders of the Mont-Saint-
inappetent plants; ANOVA, р < 0.05.

Michel bay. A group = appetent plants; lA group =

128 CHEVALIER ET AL.

RESULTS
Consumption of Plants

The results, figured according to decreasing
ingestion rates were recorded for 19 species:
14 dicotyledonous, four monocotyledons
(Poaceae), and one Bryophyta (Fig. 1). Signifi-
cant differences between ingestion rates were
highlighted by ANOVA (df = 21, F = 12.205, p<
0.0001) but data were continuous so that we
could not separate all the plants species into
different groups according to their appetence.
From the ingestion rates, we could only con-
sider two groups: the A group of four appetent
plants that were significantly more ingested
than the IA group of 5 inappetent plants (Fig. 1).
Then, the ICP-MS analysis was performed on
three plants belonging to the A group (Urtica
sp., Carduus sp., Picris sp.) and the three oth-
ers belonging to the IA group (Beta sp., Elytrigia
sp., Brachythecium sp.).

Among the 11 most ingested plants, Poa-
ceae were represented by Festuca rubra,
Dactylis glomerata and Bromus hordeaceus,
whereas the tough Elytrigia repens figured
among the less eaten species. Festuca rubra
and Dactylis glomerata seemed to be less pal-

4

O Control
в CaCO,

2

Time spent on one gauze (min)

0.1mg/ml 1mg/ml

Farm snails

atable for C. aspersum when they came from
the Mont-Saint-Michel than from Guérande.

Inorganic Compounds

Forty-two elements were quantified in the
plant samples, but we show only the 12 ele-
ments having biological interest, that is, Na, Mg,
K, Ca, Mn, Ni, Cu, Zn, As, Sr, Cd, Pb (Table 2).

With 52 and 21 g x kg * respectively, B. mar-
itima and P. echioides from Guérande showed
high sodium contents compared to the other
species (Table 2). The lead content of the
moss B. rutabulum reached ten times the
level found in E. repens or U. dioica; B. mar-
itima was about eight times richer in cadmium
than E. repens or B. rutabulum.

Plants from Guérande were significantly
richer in sodium than Mont-Saint-Michel plants
(t-test, p = 0.049). No other significant differ-
ence, for any compound, was observed be-
tween the plants from the two sites (t-test, n =
6, p > 0.05). IA plants showed significantly
lower concentrations of calcium and strontium
than the A ones. A plants were roughly ten times
richer as the IA plants. On the other hand, IA
plants had a high zinc concentration (t-test,
n=6, р = 0.049).

в wll

0.1mg/ml 1mg/ml

Wild snails

FIG. 2. Time spent by adult snails on a gauze impregnated with CaCO, solutions compared to a
control gauze saturated with distilled water. Values are means + S.E.

129

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FIG. 3. Time spent by adult farm snails on a gauze
impregnated with ZnSO, solutions compared to a
control gauze saturated with distilled water. Val-
ues are means + S.E.

Calcium Carbonate and Zinc Sulphate Experi-
ments

The total activities — that is, time spent on
both gauzes — of the wild and the farm snails
for the four CaCO, trials were similar (t-test, n
= 140, p = 0.32). Wild snails significantly pre-
ferred the gauze impregnated with the 1mg x
mi? calcium carbonate concentration
(Wilcoxon, Mann & Whitney test, n = 35, p =
0.036). Wild snails subjected to 0.1 mg xml
CaCO, and farm snails for both concentrations
did not behave differently towards the two
gauzes (Fig. 2).

The total activities of the two batches of farm
snails exposed to two different concentrations
of zinc sulphate were not significantly different
(t-test, п = 100, р = 0.44). During the 1 mg x
ml” ZnSO, trials, snails showed a non-signifi-
cant tendency to avoid the zinc sulphate solu-
tion (Wilcoxon Mann & Whitney, п = 50, р =
0.16) (Fig. 3). This reaction became significant
with the 13 mg x ml" ZnSO, concentration
which was tasted half as often as the control
(Wilcoxon Mann & Whitney, п = 50, р = 0.04).

DISCUSSION

Amongst the plant species eaten by snails,
grasses were represented by Festuca rubra
and Dactylis glomerata, whereas the tough
grass Elytrigia repens was less appreciated.

CHEVALIER ETAL.

This result is consistent with our previous data
(Chevalier et al., 2001). Williamson & Cameron
(1976) and Waddham & Wynn-Parry (1981)
hypothesized that the tough texture of some
grasses may be responsible for their deter-
rence towards slugs and snails. Differences in
the ingestion rates of the same grass species
from the two different locations may be due to
the fact that grass species often present sub-
species (Metcalfe, 1960). The soil and climate
differences between the two sites could also
induce phenological and/or toughness differ-
ences.

The rejection of the bryophyte Brachythecium
rutabulum and the sea beet Beta maritima
could be linked to their richness in lead and
cadmium respectively. Bryophyta are known to
accumulate heavy metals but the much higher
concentrations known to be toxic to C.
aspersum — 146 mg x kg* for cadmium
(Russell et al., 1981), 12700 mg x kg" for lead
(Laskowski & Hopkin, 1996) — cannot support
this hypothesis.

The sodium content of В. maritima and P
echioides, representing nearly 50% and 30%
respectively of the total amount of inorganic
compounds recorded may be related to the
proximity of the saltpans. Some halophytes
were also met near our studied area: Suaeda
sp., Obione sp. or Salicornia sp. Considering
that free salt (NaCl) is often used as a barrier
for molluscs, we may hypothesize that a deter-
rence threshold for sodium could take place for
concentrations around 50 g x kg * found in Beta
maritima.

Although other chemical (secondary metabo-
lites) and physical (height, hairiness) character-
istics might account for differences т
consumption, it seems that inorganic com-
pounds and especially calcium, strontium and
zinc could play a role in C. aspersum's feeding
choices. Calcium and strontium values are of-
ten expressed as Sr/Ca ratios in the literature,
as they show similar variations (Klein et al.,
1996). In this study, “farm snails” were reared
with a flour supplemented with calcium carbon-
ate. Since snails have strong calcium storage
and reallocation capacities, especially from
their shells (Fournié 8 Chétail, 1984), it is diffi-
cult to assess precisely whether a snail lacks
calcium. This becomes possible with wild
snails fed with lettuce, 10-fold poorer in cal-
cium compared to industrial snail food. The
positive reaction towards calcium carbonate
shown by wild snails can therefore be related to
this lack. Snails may thus be able to detect the
CaCO, available in the soil or in the plants and

INORGANIC COMPOUNDS IN SNAIL FEEDING CHOICES 181

to balance their diet in order to optimize their
calcium intake. In term of optimal foraging, nu-
trients constraints sometimes drive feeding
decision (Boyer, 1997). Such an active balance
was observed т slugs towards amino-acids
(Cook et al., 2000) and was suggested in
snails for calcium by Iglesias & Castillejo
(1999). Recently, damages made by C.
aspersum have been observed on house
paints in different regions of Brittany. The work
carried out with 16 water-based house paints
proved that the more calcium they contained,
the more snails ingested them (Chevalier &
Charrier, 1999; Charrier, in Chesnais, 1999). А
comparison between two groups of snails liv-
ing in Brittany, one group originating from a nar-
row band of soil rich in calcium and the other
from a chalky poor soil, supported the hypoth-
esis that snails lacking this mineral ate more
calcium-rich paints. Many works and the
present study allow us to state that calcium is
one of the major mineral elements governing
the feeding strategy of С. aspersum, regard-
less of which factors are controlling its cellular
incorporation.

Like copper, zinc is an essential nutrient for
snails but over a certain threshold it has inhibi-
tory effects on the growth, development and fit-
ness of snails (Laskowski & Hopkin, 1996;
Gomot-de Vaufleury, 2000). Those effects are
often accompanied by a decrease in food in-
gestion (Russell et al., 1981; Simkiss 8
Watkins, 1991; Laskowski 8 Hopkin, 1996).
The 13 mg x ml * zinc concentration that de-
terred snails during our no-choice trials 1$
higher than those found in the three deterrent
plant species, but within 30 minutes, the dura-
tion of the trial, the response was a fast rejec-
tion after contact chemoreception. In contrast,
in nature, the snails are regularly exposed to
the metals in plants and soil. Therefore, post-
ingestive effects may occur after several days,
the compounds being stored and becoming
toxic by a cumulative pattern.

Our experiment accounts for oral chemore-
ception, but uptake of calcium (Fournié &
Chétail, 1984) and water (Prior et al., 1989) can
also be integumental, suggesting the presence
of Ca** channels and aquaporines in the snail
integument.

This study has shown that gastropods can
use inorganic compounds as clues for deciding
whether or not to feed. Nutritional quality of the
plants may influence the distribution of the
snails in their environment. Snails' pressure on
their food plants may in turn influence the bal-

ance of competition between plants, resulting
in modifications of their distribution. Such stud-
ies might contribute to the knowledge of eco-
logical niche occupancy in relation to food
resources.

Further investigations on inorganic com-
pounds are needed by using complete artificial
diets with different ratios in elements (Ca, Na,
Zn) concentrations and by testing these diets
on snails deprived specifically of such ele-
ments. This could help us to assess whether
snails have the capacity to respond to the inor-
ganic compounds contents of plants and to
regulate their intake.

ACKNOWLEDGEMENTS

We wish to thank Prof. S. E. R. Bailey
(Manchester University, United Kingdom) and
Dr. Annette Gomot-Berset de Vaufleury
(Université de Besançon, France) for their val-
ued comments on this manuscript and Valérie
Briand (Université de Rennes, France) for her
kind help.

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Revised ms. accepted 7 March 2003

MALACOLOGIA, 2003, 45(1): 133-140

SHELL-BAND COLOR POLYMORPHISM IN CEPAEA VINDOBONENSIS
AT THE NORTHERN LIMIT OF ITS RANGE

Alois Honek

Research Institute of Crop Production, Drnovska 507,
CZ-16106 Praha 6-Ruzyne, Czech Republic;
honek@hb. vurv.cz

ABSTRACT

The distribution of black-, brown- and faint-banded morphs in populations of Cepaea
vindobonensis (Férussac) was established at the northwestern edge of the species’
distribution. In the Czech Republic, the species was sampled at 132 localities in the Morava
and Elbe river basins, between 48°45’ and 50°15’N. Even in this narrow zone, there was а
significant trend for increasing frequency of the black-banded morph with increasing
geographic latitude and altitude. The variation paralleled the decreasing length of sunshine
(April-August), which is > 10% shorter at the northern than at the southern localities. The
habitats occupied by northern populations (grassy steppe-like stands) differed from those
preferred by southern populations (synanthropic weed stands, mostly stinging nettle).
Geographic trends and habitat differences in morph frequencies may be affected by climatic
selection. Other possible causes of variation are discussed, comparisons are made with
other species, and gaps in evidence are identified.

Key words: Cepaea, Helicidae, shell-band color, polymorphism, geographic distribution.
climatic selection, microclimate, vegetation.

INTRODUCTION

Cepaea vindobonensis (Férussac) is a me-
dium-sized helicid snail (shell diameter 20-25
mm) living in steppe or ruderal localities of
southeast Europe. In the northwest, its distri-
bution extends to north Carpathian, Sudete and
east Alp mountains, and only a few populations
pass this limit along the Danube and Elbe riv-
ers (Schilder & Schilder, 1953; Lozek, 1956;
Kerney & Cameron, 1999). Cepaea vindo-
bonensis has been studied less than its close
relatives, C. nemoralis (L.) and C. hortensis
(Muller), although this species is common in the
area of its distribution. Consequently, geo-
graphic variation and ecological significance of
shell banding and color polymorphisms are less
understood. Variation of these characters is,
however, simpler and less marked than in the
two congeners, which should make interpreta-
tion easier.

The shell of C. vindobonensis is whitish with
five dark bands. Two or more bands may fuse
together, but individuals with confluent bands
are rare. The polymorphism consists usually in
differences of band color, which varies between
black and pale yellow. As the band color be-

133

comes paler, the sharpness of band margins
decreases so that finally the color of the shell
becomes uniformly yellowish. The decolorized
“faint-banded” individuals dominate some popu-
lations of the Balkan Peninsula and southern
central Europe (Zimmermann, 1919; Lozek,
1956; Schilder & Schilder, 1957), whereas
northern populations consist of dark-banded in-
dividuals. Detailed studies of morph distribution
in the northern Balkan Peninsula (Jones, 1973,
1974; Jones et al., 1977) revealed that propor-
tion of morphs in local populations depends on
microclimate.

Cepaea vindobonensis is abundant in lowland
areas of the basin of the Morava River (Lisicky,
1991). This eastern part of the Czech Republic
(Moravia, approx. east of 16°E) is essentially a
valley opening to the south. In the western part
of the Czech Republic (Bohemia), C. vindo-
bonensis lives in north-central lowland parts,
mainly along the Elbe River. Earlier investiga-
tion (Zimmermann, 1919; Lozek, 1956) showed
that Moravian and Bohemian populations differ
in the proportion of faint-banded individuals.
However, the precise distribution of forms and
its environmental correlates were not known.
Assuming the effect of climatic selection, | ex-

134 НОМЕК

pected a north-south cline of increasing ргорог-
tion of faint-banded morph in local populations.
The existence of this variation was studied in
106 Moravian and 26 Bohemian populations of
C. vindobonensis that were sampled and for
which geographic and climatic conditions were
established.

MATERIALS AND METHODS
Color Morphs

Shell pattern and coloration of C. vindo-
bonensis has been illustrated (Jones, 1973;
Kerney & Cameron, 1999) and described
(Jones, 1973, 1975; Cook, 1998; Staikou, 1998,
1999; Kerney & Cameron, 1999) several times.
The descriptions require some precision with
respect to variation encountered in populations
о the Czech Republic. The shell has five bands
conventionally indicated band 1 (dorsal) to 5
(ventral). In populations of the Czech Republic,
the occurrence of shells with absent or con-
fluent bands was very rare. Considering this
variation was therefore immaterial for quantita-
tive description of the composition of popula-
tions, and only differences in band and shell
ground color were important. The categories

used here are based on bands 3-5 because
bands 1 and 2 are narrow and often paler. | dis-
tinguish three morphs with respect to color of
bands 3-5: (i) black-banded morph with black
or dark-violet-black bands (violet coloration of-
ten appears in worn living and dead weathered
shells), (ii) brown-banded morph, and (iii) faint-
banded morph with yellow bands. The ground
coloration in black-banded individuals is always
white. In band morphs (ii) and (iii) the band
margins may be blunted. Brown or yellow color
then becomes “diffuse” and “spreads” into ar-
eas of white coloration between bands so that
final shell coloration may become uniformly pale
brown or yellow.

Sampling and Habitat Description

The snails were sampled т 1990-2001 at lo-
calities (Appendix) of southern and central
Moravia and central Bohemia (Fig. 1). Search
for localities started in areas where species
occurrence was already established (Lozek,
1956; Lisicky, 1991) and progressed to neigh-
boring territories up to the edge of the species
distribution. The sampling was made in April-
June when snails are most active. At each lo-
cality, living animals and well-preserved dead
shells were collected at a plot of usually

FIG. 1. The presence of shell-band color morphs at local C. vindobonensis populations (Appendix).
A. Black-banded individuals only. B. Brown-banded individuals present. C. Faint-banded (and brown-
banded) individuals present. Dashed line: isotherm of 17°C mean June temperature. Dotted line:
isotherm of 15°С mean April-September temperature. Numbers 1-7 indicate the position of
meteorology stations (Table 2). The reaches of Elbe (E) and Morava (M) rivers are indicated.

BAND COLOR POLYMORPHISM IN СЕРАЕА VINDOBONENSIS

< 1000 т? size. The number of animals of each
morph was recorded together with geographic
coordinates, altitude (established by Global
Position System or read from 1:50,000 maps)
and vegetation of the locality. Vegetation stands
were divided into four categories: grassy closed
(grass sward with no or few small spots of bare
ground), grassy open (sparse grassy vegeta-
tion with large parts of bare ground), forb closed
(dense forb stands with no bare ground between
the plants) and forb open (forb stands not
closed, with large spots of bare ground). Cli-
matic data were taken from published sources.
The Atlas of the Climate of Czechoslovakia
(Anonymous, 1958) indicates the isotherms of
mean monthly temperatures or mean tempera-
tures accumulated over selected longer peri-
ods, calculated from data of 1901-1950. The
Statistical Yearbook of the Czech Republic
2001 (Anonymous, 2001) indicates average
monthly temperatures and sunshine hours for
selected meteorology stations, calculated from
data of 1961-1990.

Data Processing

The multiple linear regression of proportion
of the black-banded morph in local populations
(arcsin transformed) on geographic latitude and
altitude of the locality and the second order re-
gression (P = b,+b,S+b,S?, where P is arcsin
transformed proportion of black-banded morph
and S is sunshine hours) of this characteristic
on sunshine hours recorded at the nearest me-
teorology station were calculated. The differ-
ences in morph frequency on localities with
different types of plant cover were tested by G-
test and Fisher exact test. All calculations were
made using Statistica for Windows (StatSoft,
1994).

RESULTS

Geographic Distribution of Shell-Band Color
Forms

The distribution of populations containing
faint-banded and brown-banded morphs (Fig.
1) was limited to south Moravia. The faint-
banded animals were found at 26 localities situ-
ated below 49°20’М and 320 m altitude (aver-
age altitude 217 m a.s.l.). The brown-banded
morph was found at all these and further 44 lo-
calities, that 1$, at a total of 70 localities situ-
ated below 49°30’М and 340 т (average alti-
tude 230 m a.s.l.). The populations consisting

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FIG. 2. The regression of the proportion of black-
banded individuals (arcsin transformation) on the
geographic latitude and altitude of the locality of
collection. Data for populations of Moravia.

only of black-banded morph were found at 36
localities between 48°48’N and 49°46'N and 180
to 360 m (average altitude 260 m a.s.l.). The
occurrence of morphs was thus not geographi-
cally separated. Nevertheless, within Moravia,
the proportion of the black-banded morph in C.
vindobonensis local populations (Fig. 2) signifi-
cantly increased (R*= 0.3166, F,, 192) = 23.628,
р < 0.001) with geographic latitude (t,,,= 4.67,
р < 0.001) and altitude (t,,,= 2.31, р < 0.05) of
the locality. In Bohemia only populations con-
sisting of black-banded individuals were found
(Fig. 1).

Habitats

The vegetation cover of localities occupied
by C. vindobonensis populations changes with
latitude. In south Moravia (< 49°10’N), half of
the populations lived in open or closed forb
stands (Fig. 3). The latter were in most cases
the patches of stinging nettles (Urtica dioica |.)
growing on eutrophic soils surrounding human
settlements. As a consequence, the occurrence
of C. vindobonensis was largely synanthropic.
By contrast, in the north Moravia and Bohemia
about 80% of populations occupied closed or
open grassy stands. Thus, the preferred sites
changed from ruderal to steppe localities.

The relationship between plant cover and fre-
quency of shell color morphs was investigated
for 59 localities of south Moravia. The popula-
tions with faint-banded and brown-banded

136 НОМЕК

100%

Y
a
x)
x

SS

50%

Y

PLANT COVER

25%

—

Ш

0%

Могама $ Могама М Bohemia

FIG. 3. The proportion of C. vindobonensis
localities covered by different types of vegetation
in the southern Moravia (left), northern Moravia
(middle), and Bohemia (right). The localities of
Moravia were partitioned into south and north
by 49°0’М parallel (northern edge of a common
occurrence of the faint-banded morph). The
patterns filling the columns represent types of
vegetation cover: FO: forb open, FC: forb closed,
GO: grassy open, GC: grassy closed.

morphs were significantly more represented at
localities with grassy stands than in forb stands
(Table 1). By contrast, populations consisting
only of black-banded morphs were frequently
found at localities grown with forb stands.

Climatic Correlates of Morph Distribution

In Moravia, the geographic distribution of
shell- band color morphs was correlated with
climatic characteristics of the localities. The
faint-banded morph was present (with few ex-
ceptions) at localities situated south of the 17°С
isotherm of mean June temperature (Fig. 1).
The northern limit ofthe area where populations
of brown-banded morph were found was ap-
proximately the 15°C isotherm of mean April-
September temperature (Fig. 1). In Bohemia
there exist small areas where mean April-Sep-
tember temperatures also exceed 15°С (Fig.
1). However, the brown-banded shells were not
found at the Bohemian localities.

The increase of the frequency of black-
banded morph was parallel to the geographic
trend in the change of climate (Table 2). As one
moves from south to north average April-Au-
gust temperature decreased only slightly by
about 1°С. A consistent trend was found in the
decrease of sunshine hours with a total of

TABLE 1. The frequency of south Moravia (below
49°10'N) populations containing particular shell-
band color morphs in relation to plant cover at
the locality. Above: populations where faint-banded
morph was present vs. populations where only
brown and/or black-banded (not faint-banded)
morphs were present. Below: populations where
only black-banded morph was present vs.
populations where faint- and/or brown-banded
morphs were present. Expected frequencies for
each category are in brackets.

Plant cover

Forb Grass Total P
Not faint- 22 14 36
banded (UTE (18.3)
Faint-banded 7 16 23

CES (LI)
Total 29 30 59 <0.05а
Black-banded 11 2 13

(6.4) (6.6)
Brown- or 18 28 46
faint-banded (22.6) (23.4)
Total 29 30 59 <0.01b

a: G-test on 2x2 contingency table
b: Fisher exact test

TABLE 2. Mean temperature and mean monthly
sunshine duration in April-August at seven
meteorology stations situated inside the area of
C. vindobonensis distribution (Fig. 1), Velké
Pavlovice (48°50’N; 180 m a.s.l.), Kucharovice
(48°50’М; 300), Brno (49°10’М; 230), Olomouc
(49°40'N; 220), Praha (50°00’М; 260), Hradec
Králové (50°10’М; 240), Doksany (50°20’М; 160).
Both climatic characteristics were calculated as
means of 1961-1990 data.

Temperature Sunshine
(°C) duration (h)
1 Velké Pavlovice 16.1 223
2 Kucharovice Sl 219
3 Brno 15.3 212
4 Olomouc 15.4 208
5 Praha 157 202
6 Hradec Králové 14.9 203
7 Doksany 14.8 190

> 10% smaller in northern than southern areas.
There was a highly significant regression of
arcsin transformed percentage of black-banded
morph on average sunshine hours of the near-
est meteorology station (b,= 53.89, b, = 0.5959,
b,= 0.00155, R*= 0.3817, р < 0.001).

BAND COLOR POLYMORPHISM IN СЕРАЕА VINDOBONENSIS 137

DISCUSSION

This study demonstrates a latitudinal trend in
the frequency distribution of shell banding
morphs of С. vindobonensis near the edge of
its range. It confirms, in more detail, the earlier
finding that shells are all dark-banded at the
northwestern extreme of distribution (Schilder
8 Schilder, 1957; Honek, 1995b), whereas more
southerly populations contain proportions of
brown or pale-banded individuals. Where
polymorphism is usual (Moravia, south of
49°10'N), populations living in open, grassy
habitats are more likely to include pale morphs,
and to hold them at higher frequencies, than
those in dense stands of forbs. The frequency
of black-banded forms also increases with
altitude, although this is a much smaller trend,
and the range of altitude slight.

These trends all suggest the operation of cli-
matic selection in response to both macro- and
microclimatic differences. However, some prob-
lems should be discussed before such a con-
clusion is accepted. Over the whole range of
the species, the effects of local microclimatic
differences seem to be more consistent than
any response to regional macroclimates. Thus,
pale morphs are rare in northern Italy (Sacchi,
1984), but abundant in some populations of the
Balkan Peninsula. In Croatia, populations in
valley bottoms subject to temperature inver-
sions are black-banded, whereas those on
insolated slopes nearby have up to 50% of pale-
banded shells (Jones, 1973, 1974). A similar as-
sociation, reinforced by altitude, was found in
northern Greece (Staikou, 1999). Only in one
study in Romania was no association found
(Jones, 1975).

In the closely related C. nemoralis and C.
hortensis, there is also evidence for climatic
selection (Jones, 1973; Vicario et al., 1988;
Stine, 1989). In these species, however, sys-
tematic differences between regions of con-
trasting climate are clearer than associations
with local microclimates (Cook, 1998). There
exist large-scale geographic trends between
shell color (Jones, 1973; Jones et al., 1977) or
body color (Cowie 4 Jones, 1985) and local cli-
mate. Studies of shell color (e.g., Falniowski et
al., 1993; Gardner et al., 1995) or body color
(Cowie, 1990) of other species also demon-
strated association between climate and snail
color.

Local differences in morph frequencies as-
sociated with habitats are more usually ac-

counted for as a product of visual selection for
crypsis by predators (Cain & Sheppard, 1954;
Cook, 1998). In this context, there 1$, as yet, no
direct evidence on the selectiveness of preda-
tion on С. vindobonensis. It is worth noting,
however, that at the northwestern extremity of
its range, black-banded shells dominate in popu-
lations confined to open grassy habitats,
whereas further south, they are most frequent
in shadier places, and decline in the open habi-
tats. This is not consistent with strong selec-
tion for crypsis, but it is with climatic selection.

Genetic drift and the history of colonization
may also be significant, especially in small iso-
lated or marginal populations (Honek, 1995a;
Cameron et al., 1998). Monomorphy of marginal
populations (e.g., in Bohemia) could derive from
small founding populations.

Studies on the possible mechanisms of cli-
matic selection, however, strengthen the case
for its operation in this case. In other species,
variation in thermal equilibria and/or rate of heat-
ing in sunshine 1$ related to the degree of mela-
nism in the shell (Etter, 1988; Honek, 1993), and
such morphs may also differ in resistance to
desiccation (Arad et al., 1993a, b). There is
evidence that this applies to C. vindobonensis.
The thermal equilibrium of the black-banded
morph under sunshine has twice been shown
to be about 1°C higher than that of paler mor-
phs (Jones, 1973; Staikou, 1999), and in the
latter case, the rate of desiccation in the black-
banded morph was also higher. In Staikou's
study, however, differences between popula-
tions were greater than differences between
morphs. Other factors clearly affect the bal-
ance, and help to explain why local differences
in the proportion of morphs are more consis-
tent than regional ones.

The consequences of these differences need
further study. Snails are generally nocturnal
(Bailey, 1981; Blanc et al., 1989; Lorvelec et al.,
1991), asis C. vindobonensis, with activity usu-
ally ceasing in the morning, later in pale-banded
individuals than in those with dark bands
(Staikou, 1999). We have no evidence to dis-
tinguish between selection due to increased
activity of dark forms under cool conditions and
that due to superior survival of pale forms un-
der insolation (Jones, 1974). Resting snails are
sometimes exposed to direct sun when resting
on plants, rocks or walls. Despite these gaps
in our knowledge, the evidence overall indicates
that climate has a strong influence on visible
variation in this snail.

138 НОМЕК

ACKNOWLEDGEMENT

| thank R.A. D. Cameron for helpful comments,
particularly very important improvement of the
discussion, and perfection of the English.

LITERATURE CITED

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BAND COLOR POLYMORPHISM IN СЕРАЕА VINDOBONENSIS 139

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Revised ms. accepted 17 September 2002

APPENDIX

Proportion of shell-band color morphs at lo-

cal C. vindobonensis populations considered
in this study. For each locality the information
is provided as:
Nearest village (geographic latitude of the col-
lection site [degrees and minutes N], geo-
graphic longitude [degrees and minutes Е],
altitude [m above sea level], black-banded [no.
individuals]-brown-banded [no. individuals]-
faint-banded [no. individuals])

Localities in Moravia

Lanzhot (4843, 1658, 160, 4-3-1); Uvaly
(4844, 1642, 230, 6-10-12); Uvaly (4844,
1642, 220, 21-21-12); Hevlin (4846, 1626,
180, 7-10-2); Мабегайсе (4848, 1606, 230, 5-
0-0); Suchovské Mlyny (4848, 1735, 360, 7-0-
0); Zajeci (4853, 1647, 220, 5-3-0); Straznice
(4853, 1723, 180, 14-0-0); Cejkovice (4855,
1655, 270, 4-3-6); Troskotovice (4855, 1625,
190, 5-8-11); Hustopece (4855, 1644, 200, 9-
6-0); Kurdéjov (4856, 1646, 250, 8-7-3);
Horni Bojanovice (4857, 1647, 200, 1-8-1);
Vlasatice (4857, 1628, 180, 10-10-36):
Vlasatice (4857, 1628, 180, 20-41-55);
Hovorany (4857, 1700, 220, 15-1-0);
Hostéradice (4857, 1615, 220, 20-5-0);
Pouzdrany (4858, 1638, 180, 7-7-10); Mistrin
(4858, 1705, 200, 14-1-0); Bzenec (4858,
1717, 190, 8-1-0); Brumovice (4858, 1654,
200, 62-7-0); MoraSice (4858, 1613, 270, 26-
0-0); Diváky (4900, 1649, 220, 10-23-7);

Cvréovice (4900, 1632, 180, 47-37-8);
Zeravice (4900, 1715, 230, 25-4-0);
Olbramovice (4900, 1623, 220, 13-0-0); Hluk
(4900, 1731, 240, 7-0-0); Zidlochovice
(4902, 1636, 180, 28-13-3); Jezefany (4902,
1626, 220, 6-2-0); Lodénice (4902, 1628,
200, 4-3-0); Zeletice (4902, 1700, 190, 48-5-
0); Archlebov (4903, 1703, 200, 14-7-1);
Trbousany (4903, 1627, 210, 18-11-1);
Blucina (4903, 1640, 200, 6-12-2);
Hrusovany u Brna (4903, 1635, 220, 28-13-
3); Némcicky (4903, 1630, 190, 81-13-5);
Násedlovice (4903, 1658, 190, 25-2-0);
Moravsky Krumlov (4903, 1620, 260, 23-0-
0); Lovéice (4904, 1703, 240, 5-14-2);
Bohuslavice (4904, 1708, 210, 55-6-4);
Bohuslavice (4904, 1708, 210, 30-9-0);
Rokytná (4904, 1620, 270, 40-0-0); Otnice
(4905, 1650, 230, 15-14-3); Zdánice (4905,
1702, 230, 18-0-0); Mohelno (4906, 1611,
360, 14-0-0); RaSovice (4907, 1653, 240, 64-
5-0); Silúvky (4908, 1628, 280, 32-42-9);
Nesovice (4908, 1705, 240, 61-3-0);
Brankovice (4908, 1708, 260, 26-8-0);
Vícemilice (4908, 1702, 230, 31-25-0);
Krizanovice (4909, 1656, 230, 40-2-0);
Slavkov (4909, 1653, 220, 80-4-0); Hajany
(4909, 1634, 260, 11-6-0); Nesovice (4909,
1703, 260, 69-9-0); Bucovice (4909, 1659,
220, 126-15-0); Slavkov (4909, 1652, 220,
10-0-0); Milonice (4910, 1705, 260, 38-2-0);
Lísky (4910, 1714, 320, 63-0-0); Napajedla
(4910, 1731, 210, 18-0-0); Rostoutky (4911,
1703, 290, 51-1-0); Kozlany (4911, 1702,
310, 24-0-0); Strabenice (4912, 1714, 320,
42-8-3); Rostoutky (4912, 1703, 270, 19-3-
0); Holubice (4912, 1650, 280, 19-4-0);
Rousinov (4912, 1653, 250, 26-8-0);
Rousinov (4913, 1653, 270, 24-2-0);
Zdounky (4914, 1717, 230, 22-2-0);
Morkovice (4914, 1714, 280, 19-2-0);
Kvasice (4914, 1730, 200, 8-4-0); Skrzice
(4915, 1720, 210, 16-1-0); Netcice (4915,
1717, 240, 61-4-0); Rosténice (4915, 1658,
260, 5-8-0); Zborovice (4915, 1716, 260, 30-
0-0); VySkov (4917, 1701, 240, 49-0-0);
Heroltice (4918, 1704, 240, 18-2-0); TéSice
(4918, 1710, 210, 60-0-0); Pustimér (4919,
1702, 290, 44-7-1); Kfenovice (4919, 1715,
200, 48-2-0); Dfevnovice (4920, 1709, 240,
27-36-2); Nezamyslice (4920, 1800, 250, 64-
4-0); Zeleë (4921, 1704, 340, 121-15-0);
TiSnov (4922, 1623, 290, 44-22-0); Brodek u
Prostéjova (4921, 1705, 290, 64-0-0);
Brodek u Prostéjova (4922, 1705, 280, 22-0-
0). РА (4923, 1710; 280, 20-220);
Polkovice (4923, 1719, 200, 74-0-0);

140 НОМЕК

Vranovice (4924, 1706, 250, 11-0-0); Henclov
(4926, 1723, 200, 17-1-0); Prostéjov (4930,
1705, 240, 24-5-0); Kostelec na Hané (4930,
1704, 250, 30-7-0); Zdetin (4930, 1659, 350,
13-0-0); Ptení (4930, 1658, 340, 14-0-0);
Celechovice (4931, 1706, 280, 11-15-0);
Celechovice (4931, 1704, 280, 14-0-0);
Smrzice (4931, 1709, 250, 45-0-0); Blatec
(4932, 1715, 230, 57-0-0); Blatec (4932, 1715,
230, 51-0-0); Slatinice (4933, 1706, 250, 28-0-
0); Cechy pod Козйет (4933, 1701, 290, 18-
0-0); Pénéin (4934, 1701, 330, 11-0-0);
Olomouc (4935, 1717, 210, 17-0-0);
Drahanovice (4935, 1705, 250, 53-0-0);
Ludéfov (4935, 1703, 280, 25-0-0); Ludérov
(4935, 1703, 330, 25-0-0); Cholina (4940,
1702, 270, 26-0-0); Bila Lhota (4946, 1658,
290, 16-0-0)

Localities in Bohemia

Malá Leëice (4950, 1422, 280, 10-0-0);
Srbsko (4957, 1408, 260, 39-0-0); Karl&tejn
(4957, 1410, 260, 17-0-0); Vysoké Myto (4957,
1610, 270, 20-0-0); Kosténice (5000, 1554,
240, 20-0-0); Uhersko (5000, 1602, 240, 20-0-
0): Platenice (5001, 1557, 240, 20-0-0); Praha
(5002, 1420, 300, 40-0-0); Daëice (5002,
1556, 230, 20-0-0): Hradéany (5009, 1517,
250, 20-0-0); Podmorän (5010, 1420, 290, 24-
0-0): Bléany (5014, 1327, 280, 15-0-0):
Mécholupy (5016, 1333, 290, 15-0-0); Veltrusy
(5016, 1420, 180, 20-0-0); Kly (5018, 1433,
220, 60-0-0); Nová Ves (5020, 1418, 200, 20-
0-0); Roudnice (5025, 1417, 210, 9-0-0); Raná
(5025, 1347, 380, 40-0-0); Libochovice (5026,
1403, 170, 11-0-0); Charvatce (5026, 1348,
350, 25-0-0); Charvatce (5026, 1349, 330, 25-
0-0); Steti (5027, 1423, 170, 20-0-0); Vrbice
(5028, 1417, 160, 36-0-0); Chcebuz (5029,
1423, 230, 72-0-0); Vrutice (5030, 1417, 160,
20-0-0): Polepy (5031, 1419, 160, 20-0-0)

MALACOLOGIA, 2003, 45(1): 141-148

КАКУОТУРЕ$ OF EUROPEAN SPECIES OF RADIX
(GASTROPODA: PULMONATA: LYMNAEIDAE)

AND THEIR RELEVANCE TO SPECIES DISTINCTION IN THE GENUS

Alexandr V. Garbar' & Alexei V. Korniushin?”

ABSTRACT

Karyotypes of Radix auricularia (Linnaeus, 1758) and three disputable taxa considered by
different authors as distinct species or assigned as forms of Radix peregra (Múller, 1774),
sensu lato — R. labiata (Rossmássler, 1835), R. balthica (Linnaeus, 1758), and R. ampla
(Hartmann, 1821) — were studied with preparations obtained from gonad tissues by the air-
drying method. The studied taxa have the same diploid number (2n = 34), but are character-
ized by different morphology of some chromosome pairs. п particular, К. labiata (traditionally
identified as R. peregra, $. $.) and R. balthica (= К. ovata in traditional understanding) differ
in the number of subtelocentric chromosomes (1 and 5, respectively), species status of
these taxa being also supported by pronounced differences in centomeric indexes of chro-
mosome pairs 4 and 16. Species distinctness of К. ampla is supported by differences in
three chromosome pairs, and karyological similarity between this taxon and К. balthica 1$
also noted. FN values varied among the studied taxa from 56 in В. ampla to 66 in К. labiata.
The known karyological characters are traced on phylogenetic trees suggested by recent
molecular reconstructions. This study demonstrates that karyology can be an effective tool

for aiding taxonomic distinctions of historically problematic groups of molluscs.
Key words: Radix, karyotypes, taxonomy, species distinctions.

INTRODUCTION

The group of lymnaeid species bearing the
name Radix Montfort, 1810, is defined mainly
by its thin-walled fragile shell with a relatively
large aperture (Falkner, 1990; Gloer & Meier-
Brook, 1998; Jackiewicz, 1998; Glóer, 2002).
The distinctive karyological character of Radix,
namely its chromosome number (n = 17) devi-
ating from the other members of the family (typi-
cally n = 18, or in some taxa n = 16 or 19), has
also been known for a long time (Inaba, 1969;
Choudhary et al., 1992).

Despite intensive research by different meth-
ods (Hubendick, 1951; Inaba, 1969; Patterson
& Burch, 1978; Kruglov & Starobogatov, 1983,
1993; Remigio & Blair, 1997; Jackiewicz, 1998;
Bargues et al., 2001), many taxonomic prob-
lems of Radix remain unresolved. In particular,
the rank of the group is alternatively defined as
subgeneric within Lymnaea (Hubendick, 1951;
Kruglov & Starobogatov, 1983, 1993;
Jackiewicz, 1998; Kerney, 1999) or generic

(Patterson & Burch, 1978; Falkner, 1990; Glöer
& Meier-Brook, 1998; Bargues et al., 2001;
Falkner et al., 2002). A distinct subgenus
Peregriana was recognized in Lymnaea along-
side Radix by Kruglov & Starobogatov (1983).
Still uncertain also is the number of species
within this group. Stressing the lack of distinc-
tive anatomical characters and existence of in-
termediate shell morphotypes, British (Kerney,
1999) and Polish (Jackiewicz, 1998) authors
recognized only two European species, namely
Lymnaea (Radix) auricularia (Linnaeus, 1758)
and L. (R.) peregra (Muller, 1774), distinguish-
ing in the latter up to four ecological forms: L.
peregra $. $. (= f. typica, sensu Jackiewiecz,
1998), f. ovata (Draparnaud, 1805), f. lagotis
(Schrank, 1803), and f. атр/а (Hartmann,
1821). Evidence for species distinctness of
Radix ovata was provided by С бег & Meier-
Brook (1998), whereas Falkner (1990) also rec-
ognized R. ampla as a full species. Five
European taxa are supported by recent molecu-
lar studies (Bargues et al., 2001) and have been

‘Department of Natural Sciences, Zhitomyr Pedagogical Institute, |. Franko Str., Zhitomyr, Ukraine
21.1. Schmalhausen Institute of Zoology, The National Academy of Sciences, В. Khmelnitsky Str. 15, 01601 Kiev, Ukraine

*Corresponding author: root@iz.freenet.kiev.ua

GARBAR & KORNIUSHIN

142

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KARYOTYPES ОЕ EUROPEAN RADIX 143

included as species in the latest European
checklists (Falkner et al., 2002; Glôer, 2002)
with the following names: Radix auricularia, R.
labiata (Rossmässler, 1835) (substituting К.
peregra, $. $. of previous authors), К. balthica
(Linnaeus, 1758) (= R. peregra of Müller, = К.
ovata, auctt.), R. ampla, and R. lagotis. An even
more profound subdivision was suggested by
Kruglov & Starobogatov (1983, 1993) but not
supported by any ofthe later studies.

Until now, chromosome numbers were
mainly involved in discussions about taxonomy
and relationships among freshwater gastro-
pods. However, interspecific differences in chro-
mosome morphology were recently found in
Viviparus (Bar$iene et al., 2000). Furthermore,
preliminary investigations of Garbar (1998,
2000) on Radix and on Stagnicola (Garbar &
Korniushin, 2002) have shown that morphologi-
cal characters of chromosomes may be also
helpful by species distinction in lymnaeids. This
paper summarizes results of karyological in-
vestigation in four European taxa of Radix, des-
ignated in modern reviews as R. auricularia, R.
labiata, R. Баса and R. ampla.

FIG. 1. Shells of the studied species: А. Radix
auricularia (Linnaeus, 1758); В. R. labiata (Ross-
mássler, 1835); С. К. Баса (Linnaeus, 1758);
О. В. ampla (Hartmann, 1821). Scale bar, 5 mm.

MATERIALAND METHODS

Material was collected by the first author in
1997-2000 in western and central Ukraine
(Table 1). In order to minimize the influence of
local factors, each species was sampled in two
remote (at a distance of at least 150 km) locali-
ties; the sampled populations inhabit different
river systems (of the Danube, Yuzhny Bug, and
Dnieper drainages) and live under somewhat
different climatic conditions. Some populations
included in the earlier karyological studies
(Garbar, 1998, 2000) were re-sampled. Species
identification was based on traditional concho-
logical and anatomical characters (Glóer 8
Meier-Brook, 1998; Jackiewicz, 1998). The
studied group is treated in our work as a genus
and the studied forms as species, following the
latest systematic and phylogenetic works
(Remigio 8 Blair, 1997; Bargues et al., 2001;
Falkner et al., 2002). Nomenclature of the latest
European monographic review (Glöer, 2002) is
used herein; in order to avoid misunderstanding,
we provide the list of the studied taxa with syn-
onyms used in the cited publications (Table 1).

Pictures of shells are provided in Figure 1.
Voucher specimens have been deposited in the
mollusc collection of the Museum fúr Natur-
kunde, Humboldt Universitát zu Berlin, Ger-
many.

Chromosome preparations were obtained
from the gonad tissue according to the recom-
mendations of BarSiene et al. (1996) and Garbar
(1998). Molluscs were placed for 17 h ina
0.002% solution of colchicine. Pieces of gonad
were fixed in a mixture of ethanol and acetic
acid (3:1). The cell suspension was prepared
by maceration in a mixture of concentrated ace-
tic and 60% lactic acids (30:1) and dispersed
with a capillar pipette on microscopic slides
heated at 50°C. Dried preparations were
stained 10-15 min in 10% solution of azur-
eosine after Romanovski, prepared on 0.01M
phosphate buffer. Stained preparations were
placed for short time in xylol and embedded in
Canada Balsam. These preparations were stud-
ied under a Biolam-L-212 microscope with mag-
nification 10 x 90. The plates with a good
dispersion of chromosomes and moderate de-
gree of spiralization were selected for photo-
graphing and measuring. The relative length and
centromeric index were then calculated for
each chromosome. Chromosomes were clas-
sified according to Levan et al. (1964). The Fun-
damental Number (FN) was calculated as the
number of autosome arms in haploid comple-
ment, with a value of 4 given to metacentric and

144 GARBAR & KORNIUSHIN

submetacentric chromosomes, and a value of
2 to subtelocentric chromosomes (no telocen-
tric chromosomes were found in the studied
taxa). Quantitative data from the most numer-
ous samples were processed statistically us-
ing standard methods.

RESULTS
Descriptions of Karyotypes

Radix auricularia. 2n = 34. Chromosomes of
adjacent pairs similar in size, their relative length
varies between 9.21% and 4.15% (Table 2).
Karyotype includes 11 pairs of metacentric, four
pairs of submetacentric, and two pairs of

N FR

7. A

Je pel EA
a 4
EN,” LA, \ 2% x

q `
(MINE Os ıe
ny \ RADA AN 58 кА tn С
> ER ka ВА 13 ER ЕК Ев

AR ai an ЗА aa ИК ла ха D

FIG. 2. Chromosomes of Radix auricularia:
А. Mitotic metaphase of a specimen from Zhito-
туг; В. The same of a specimen from Vinnitsa;
С, D. Karyotypes of specimens from Zhitomyr and
Vinnitsa region, respectively; E. Ideogram (based
on data from Table 2). Scale bar, 10 um.

10 mkm

10
RL,%

subtelocentric chromosomes (Fig. 2, Table 2).
FN = 64.

Radix labiata. 2n = 34. Chromosomes of ad-
jacent pairs similar in size, with relative length
between 9.69% and 3.74% (Table 2). Karyotype
includes 12 pairs of metacentric, four pairs of
submetacentric, and one pair of subtelocentric
chromosomes (Fig. 3, Table 2). FN = 66.

Radix balthica. 2n = 34. Chromosomes of
adjacent pairs similar in size, with relative length
between 9.29% and 3.95% (Table 2). Karyotype
includes eight pairs of metacentric, four pairs
of submetacentric, and five pairs of subtelo-
centric chromosomes (Fig. 4, Table 2). FN = 58.

Radix ampla. 2n = 34. Chromosomes of ad-
jacent pairs similar in size, with relative length
of chromosomes between 9.32% and 4.05%
(Table 2). Karyotype includes eight pairs of
metacentric, three pairs of submetacentric, and
six pairs of subtelocentric chromosomes (Fig.
5, Table 2). ЕМ = 56.

«A wT oc. ee À 4
Se Swan
ve “+ mA Aye A Ча, а

96 83 34 Ex 45 вк XA хи ax

xx BA 4X SR BSA XX RB ха E

$$ BE EX ВЕ St GA NS 35563

KE 69 Ux ва аа za az ss О

10 mkm_

FIG. 3. Chromosomes of Radix labiata:
A, B. Mitotic metaphases of specimens from
Zhiomyr and Vorokhta, respectively; C, D. Karyo-
types of specimens from Zhitomyr and Vorokhta,
respectively; E. Ideogram (based on data from
Table 2). Scale bar, 10 um.

10
RL,%

KARYOTYPES OF EUROPEAN RADIX 145

Comparisons

All studied taxa are characterized by the same
chromosome number (2n = 34). Morphological
similarity is demonstrative also in some indi-
vidual chromosome pairs, that is, pairs 1, 3, 5,
7, 9, 11, 12 and 15, belonging to one and the
same type in all these taxa. On the other hand,
distinctive features of chromosome morphol-
ogy were noted not only for the doubtless spe-
cies Radix auricularia, but also for three taxa of
disputable status — R. labiata, R. balthica, and
R. ampla. The karyotype of R. labiata (= R.
peregra, auctt.) differs from that of R. balthica,
and R. ampla in morphological type in seven to
nine pairs. The higher rate of subtelocentric
chromosomes in two latter species (five to six
out of 17 pairs) is also reflected in the lower
values of FN (58 and 56, respectively), com-

CPUS BK A reas sun Kt

$) FS AM AB ай AB ча 54 С
{УС UR 56 3053 If аа xx

IR BR AR AS вв ви цв ла D

10 mkm
hu ———

FIG. 4. Chromosomes of Radix balthica:
À, B. Mitotic metaphases of specimens from
Olevsk and Kiev, respectively; C, D. Karyotypes
of specimens from Olevsk and Kiev, respectively;
E. Ideogram (based on data from Table 2). Scale
bar, 10 um.

10
RL,%

pared to FN of R. labiata (66). In some cases,
such as in the pairs 2, 6, and 13, mean values
of centromeric indexes in the compared spe-
cies were close (Table 2), and assignment of
chromosomes to different types might be influ-
enced by individual variation. However, that was
not the case in chromosome pairs 4 and 16,
for which interspecific differences were the most
pronounced. Taking into account that chromo-
somes adjacent to the mentioned pairs in the
ideograms (Figs. 2-5) were morphologically
similar among the studied taxa, we conclude
that observed differences could not be caused
by errors in identification of individual chromo-
somes. Therefore, they are further referred to
as taxonomic characters.

Similarity in chromosome morphology be-
tween К. labiata and R. auricularia is notewor-
thy: only two pairs (4 and 6) were assigned to

> wes,
> 4 ’ ae al
>. ERS te, 48
IGE LE AS
= А ES B

à АА № ЗАКИ ВИ АХ KE à

KA NA KA Им ЧА м» С

1) 9599000004 1) 05 28

BE IE LE LU AU te 8400 о

10 mkm
boon 4

FIG. 5. Chromosomes of Radix ampla:
À, B. Mitotic metaphases of specimens from
Zhitomyr and Kharkiv, respectively; C, D. Karyo-
types of specimens from Zhytomyr and Kharkiv,
respectively; Е. Ideogram (based on data from
Table 2). Scale bar, 10 um.

10
RL,%

GARBAR & KORNIUSHIN

146

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IS 58'0+7/`02 800+1/5% WS 25117015 УОбОЗУСУ W EBOFOFEr SO0F9Ly ш 821+9/'/6 600+9%7% 9
ш 09111585 900796 W HPLOFLLIF 700+9 W 8/0FS9SZb 900+95У ш BELFIGEr 900+89% SI
WS J60FEGPE 900+59% WS JLELFSGLSE SO00F/9p ш ZZLFOS6E 900+66У W pyyLFOL8E 900F6/ FL
\s 26`0 + |3'УС 70'0+06% IS ZLLF60EZ SOOFSSbh WS JO, FJZSZE SOOFSLh WS OYyLFLZ9OZ 800F00G EL
w 76`0 + 0985 900FZLG ш 86011507 $00+006 ш /90F/9€t SCOFIOS W OFIFrZIr LOOFSGZTS ZI
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WS HG, FZISE 70010595 W LOLFEOOF $00+6%6 W 890+90°57 POOFZSGS ш 6YLFELBE SOOFSSS OL
w LOL FEZBE vOOFZGS Ш 8/0+1007 700195 W /90+F+6c+ 700+5/5 W 9YLFZGLE 90'0+5/°9 6
1s O9LFPSOZ pOOFI8S IS LOLFS9ZZ POOFLES WS pyg0F8z9z SOOFZOO WS SOLFOEE VvOOFLOS 8
ш 91+ 08'66 900FS09 W 98011607 SOOFELY9 ш 7/0+985у7 POOFOTI W 680F06Er SOOFSIYI 2
1s GLL¥9S¢0Z 800FZE9 IS. ZLULFESELZ SOOFEVO WS 101+8896 SOOFZVO IS $7,+199р $0'0+0$°9 9
w GGL+F/0lt SOOFZS9 ш Z80F/06€ SOOF6S9 ш 69OFSEEY HOOFrE9 ш ItÿLl+F/tÿct SO0FSV9 G
JS GGLFEOEZ LLUOFZB9 IS 9LÉLFEZZZZ LOOFIOL ш 690+F1I/2+ 900+689 WS OLLFEYSC 8007599 y
ш 6LLFLG Lp 9001821 W pyZOFStHZH 900Fte/ W 080FO0O€€b LOOFESL W 6yLFELZL 8007889 €
ws 0914/0256 60O0FIZZ WS SELFSE/E 800F/08 ш 960F6S6€ 600F008 ш OGSLF/08E ILOFSEZ С
Ш Ср 0917 6101256 W ELOFEZIF ILOF66 ш 690F18 lb 800+696 W ILLFEZOY L70F126 |
adh] (as +) (as +) adhy (as +) (as +) adh, (dS =) (as +) adh, (as +) (as +) “ouJIeg
% 19 % Ty % 19 % Ty % 19 % Ty % 19 % Ty
ejdue y вошуед “Y eJeige) 4 euejnoune y

'A9/9]9| J3MY YÁLOYNYZ шо е/аше y pue ‘HSA8IO ‘чобэч
JÁLWOYUYZ шо 29/4I/8Q ‘у pue чАшоуцт шо eyeige/ "Y 'NSJ3]a] Janıy YÁLUOYYZ шо емепэипе XIpey JO (JWOSOWOAYI D11JUSDO|SIANS - JS ‘ощиээеэшап$
- WS ‘ощиаэеаш - LU) зэшозошолио JO иоцеэцизео pue (иоцемлэа PIEPUEIS - GS 'xapur энэшодщиеэ - 19 ‘чзбиз| элце|э - TY) Зиэшеллзеэи ‘с FIGVL

KARYOTYPES OF EUROPEAN RADIX 147

different types in these two taxa, and values of
FN were also close. The karyotype of R. ampla
is close to that of К. balthica, differing in mor-
phological type of three chromosome pairs;
presence of one more subtelocentric pair (16)
is the most characteristic feature of the former
taxon.

DISCUSSION

Diploid numbers of all studied species (2n =
34) agree well with the published literature on
the genus Radix (Inaba, 1969; Patterson &
Burch, 1978; Coudhary et al., 1992; BarSiene
et al., 1996; Garbar, 1998, 2000). No cases of
hypodiploidy or polyploidy were observed by our
study, this result being in contrast with data on
Spanish populations identified as R. peregra
(BarSiene et al., 1996). The presence of telo-
centric (t) chromosomes in the karyotype of R.
auricularia from the Zhitomyr population reported
by Garbar (1998) was also not confirmed by
this study. In all probability, this work dealt with
an artifact caused by very profound spiralization
of chromosomes. At the same time, our present
observations оп К. labiata and R. balthica
agree with the data of Garbar (2000) on these
species (for correspondence of nomenclature:
Table 1). The karyotype of К. ampla has been
studied here for the first time.

Differences in the chromosome morphology
(most pronounced in the pairs 4 and 16) sup-
port species status of R. balthica (= R. ovata)
and К. labiata (traditionally referred to as К.
peregra) — two taxa considered conspecific by
many taxonomists dealing with shell and ana-
tomical characters (Hubendick, 1951; Jackiewicz,
1998; Kerney, 1999). However, karyological dis-
tinction between R. labiata and R. balthica
shown by this study should be checked on the
representative material taken throughout their
distributions. Noteworthy, chromosome pair 4
is apparently of the same type in the karyotype
of Spanish R. peregra shown by BarSiene et al.
(1996) and Ukrainian specimens of R. /abiata
included in this study, but similarities/differences
in other chromosome pairs cannot be evalu-
ated, because centromeric indexes for the
Spanish specimens were not provided.

Species status of R. ampla is also supported
by this study, but the karyological differences
between this taxon and R. balthica were appar-
ently less pronounced than those reported for
R. balthica and R. labiata. Furthermore, the
karyotype of L. (Peregriana) fontinalis (Studer,

1820), as described by Garbar (2000), is similar
to that of R. ampla (especially in having
subtelocentric chromosome pair 16), with mod-
erate (about 6%) difference in mean values of
centromeric indexes of chromosome pair 2. This
result is surprising, because L. fontinalis (in the
understanding of Russian authors) corresponds
in its conchological and anatomical characters
(Kruglov & Starobogatov, 1983: fig. 2, 3; Garbar,
2000: fig. 1) to R. balthica of modern western
European reviewers (Glóer, 2002) and is appar-
ently different from К. ampla. Thus, correlation
between chromosome morphology and the other
characters in the R. balthica/R. ampla complex
should be checked by further studies.

The observed pattern of karyological differ-
ences in Radix is consistent with the phyloge-
netic trees based on ITS-2 sequences (Bargues
et al., 2001), supporting the following topology
in the clade of European Radix species (no-
menclature as used here): (R. /abiata, (R. au-
ricularia, (К. lagotis, К. ampla, R. balthica))). In
particular, the peculiar karyotype of К. labiata
and the karyological similarity between R. ampla
and R. balthica corroborate this phylogenetic
hypothesis. In particular, the overwhelming
prevalence of meta- and submetacentric chro-
mosomes characterizes the basal taxa R.
labiata and R. auricularia, whereas the high
number of subtelocentric chromosomes is a
common feature of R. balthica and R. ampla,
which belong to the terminal clade. Thus, the
state of the karyotype of the former may be in-
terpreted as plesiomorphic, and the latter as
apomorphic within the analysed group. The re-
sults of our investigation are also consistent with
the molecular analysis (Bargues et al., 2001)
in suggesting, that R. атр/а is a valid species
alongside R. balthica. Neither karyological nor
molecular characters support the subgenus
Peregriana of Kruglov & Starobogatov (1983,
1993) as including, among other species, R.
labiata, R. balthica, R. lagotis and R. ampla, but
not R. auricularia. Broad understanding of Lym-
naea peregra (Jackiewicz, 1998) also contra-
dicts both data sets.

The results of this work, as well as earlier
observations on other freshwater gastropods
(BarSiene et al., 2000; Garbar & Korniushin,
2002), show that the study of chromosome
morphology may provide additional characters
for species diagnosis and phylogenetic analy-
sis. Karyological study of Lymnaeidae should
be continued, given the parasitological impor-
tance of this group and remaining uncertainty
about its species-level taxonomy.

148 GARBAR & KORNIUSHIN

ACKNOWLEDGEMENTS

The authors are grateful to Dr. Valentina
Manilo (Kiev) for the help by processing and
interpreting chromosome preparations, and to
the anonymous reviewers for their helpful cor-
rections and suggestions on the manuscript.

LITERATURE CITED

BARGUES, M. D., M. VIGO, Р. HORAK, J.
DVORAK, R.A. PATZNER, J. P. POINTIER, M.
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COMA, 2001, European Lymnaeidae (Mol-
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BARSIENE, J., С. КВ! & D. BARSYTE, 2000,
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FALKNER, G., 1990, Binnenmollusken. Pp. 112-
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FALKNER, G., Т.Е. J. RIPKEN 8 M. FALKNER,
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GARBAR, А. V., 1998, A karyotype of Lymnaea
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Lymnaeidae) from central Polissya. Vestnik
Zoologii, 32 (5-6): 137-138 [in Russian, with
English summary].

GARBAR, A. V., 2000, Description of karyotype
of three species of genus Lymnaea (Gas-
tropoda, Pulmonata, Lymnaeidae) of the Fauna
of Ukraine. Vestnik Zoologii, Suppl. 14: 40—47
[in Russian, with English summary].

GARBAR, A. V. & A. V. KORNIUSHIN, 2002,
Karyotypes of two European species of the
genus Lymnaea with disputable taxonomic sta-

tus (Gastropoda: Pulmonata: Lymnaeidae).
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Museum für Tierkunde Dresden, 20 (1): 235-
246.

GLOER, P., 2002, Mollusca |. Süßwasser-
gastropoden Nord- und Mitteleuropas.
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breitung. ConchBooks, Hackenheim. 327 pp.

GLOER, P. & C. MEIER-BROOK, 1998,
Süsswassermollusken. Ein Bestimmungs-
schlüssel für die Bundesrepublik Deutschland.
Deutscher Jugendbund für Naturbeobachtung,
Hamburg. 136 pp.

HUBENDICK, B., 1951, Recent Lymnaeidae, their
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Vetenskapakademiens Handlingar, 3 (1):
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lymnaeid snails. Malacologia, 7 (2/3): 143-168.

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171-217, in: V. FRETTER 4 J. PEAKE, eds.,
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REMIGIO, E. A. 8 D. BLAIR, 1997, Molecular
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Revised ms. accepted 28 December 2002

MALACOLOGIA, 2003, 45(1): 149-166

USE OF MICROSATELLITE VARIATION AND RAPD-PCR TO ASSESS
GENETIC POLYMORPHISM IN BIOMPHALARIA GLABRATA SNAILS
FROM A SINGLE LOCALE INA SCHISTOSOMIASIS ENDEMIC AREA

Christopher Rowe*, Wannaporn Ittiprasert?, Carolyn Patterson', Claudia Eliff', Kristen
Page*, Susan Bandoni*, Thomas Wilke?, Dennis Minchellaÿ, Fred Lewis' & Matty Knight'*

ABSTRACT

Genetic variation was assessed in Biomphalaria glabrata snails using variations in
microsatellite loci and by RAPD-PCR analysis. Populations of snails examined were field-
collected isolates from a small pond in a schistosomiasis endemic region in Brazil, after
standard conditions were developed for analyzing snails from two laboratory-maintained
stocks. The analyses were performed using a total of 60 microsatellite primer sets and, for
RAPD-PCR, a total of 19 random primers. We show that genetic diversity can readily be
detected by both molecular methods among the field-collected snails from this small site.
In addition, RAPD-PCR bands that were found in another study to segregate with parasite
resistance were not detected in any of the field-collected snails analyzed.

Keywords: Biomphalaria glabrata, RAPD-PCR, microsatellites, genetic diversity, AMOVA,

schistosomiasis.

INTRODUCTION

Schistosomiasis is a parasitic disease of glo-
bal significance. Nearly every stage of the
schistosome!'s life cycle, which alternately in-
volves mammalian and snail hosts, has been
the target for intervention, but often these con-
trol efforts result in short-term success.
Although combined efforts to reduce transmis-
sion, for example by mollusciciding and mass
chemotherapy, are initially effective, rapid
recolonization by the snails and reinfection in
the human population make long-term control
difficult to achieve. Developing an effective vac-
cine against schistosomes is a challenging
task (Bergquist, 1998), making it even more
important to explore other control measures.
The strategy of replacing susceptible snails in
schistosomiasis-endemic areas with parasite-
resistant snails 15 a suggested form of
biological control (Hubendick, 1958). Implemen-
tation of these control methods and the need to
better understand the epidemiology of schisto-
somiasis has stimulated interest in better

characterizing the population genetic struc-
tures of both the snail hosts and parasites
(Bandoni et al., 1990; Johnston et al., 1993;
Hoffman et al., 1998; Langand et al., 1999;
Curtis & Minchella, 2000; Sire et al., 2001).
Some degree of genetic diversity in snail field
populations has been noted using a variety of
methods, including allozyme analysis, RAPD-
PCR, and variations in microsatellite loci
(Bandoni et al., 1990; Vidigal et al., 1994; Jones
et al., 1999; Bandoni et al., 2000; Mavares et
al., 2000; Charbonnel et al., 2000). Applying
RAPD-PCR and microsatellite analysis to
study snail population structure 1$ relatively
new, but both methods have found wide use for
examining the frequency of genotypes in rela-
tion to disease, genetics and ecology in nu-
merous species of medical and agricultural
importance. Over the last several years, how-
ever, less frequent use has been made of
multi-locus molecular markers, such as
RFLPs and RAPDs, than of microsatellites for
population genetic studies (review: Jarne «
Theron, 2001). This has occurred even though

‘Biomedical Research Institute, 12111 Parklawn Drive, Rockville, Maryland 20852, U.S.A.
“Department of Microbiology and Immunology, Faculty of Tropical Medicine, Mahidol University, Bangkok, Thailand

“Wheaton College, Wheaton, Illinois, U.S.A.

‘Department of Biology, SUNY-Geneseo, Geneseo, New York, U.S.A.
‘Department of Microbiology and Tropical Medicine, The George Washington University, Washington, D.C., U.S.A.
‘Department of Biological Sciences, Purdue University, West Lavayette, Indiana, U.S.A.

“Corresponding author: mknight@afbr-bri.com

150 ROWE ET AL.

RAPD-PCR has some obvious advantages,
notably the ability to identify genetic loci
(scored as the presence or absence of
bands) without prior cloning or sequencing
steps (Williams et al., 1993). Detecting micro-
satellite loci, on the other hand, depends on
initial time-consuming cloning and sequencing
steps to identify the simple sequence repeats
(SSR, or microsatellite loci), and the design of
primers flanking SSRs for amplification and
detection of polymorphic loci (Чате & Lagoda,
1996).

Most of the studies on snail hosts have ana-
lyzed the genetic structure of Biomphalaria
glabrata populations, the major intermediate
host for Schistosoma mansoni in South
America and several Caribbean islands. Re-
cently, this species was also found in Egypt,
where it could complicate the extensive con-
trol efforts that have been in place there for
the last several years (Yousif et al., 1996).
Vidigal et al. (1994) showed that snails in a
population in one geographic region were
more homogeneous (based on RAPD-PCR
analysis) than snails from different geographic
regions. However, for the African species B.
pfeifferi, Hoffman et al. (1998) found consider-
able heterogeneity in populations even within a
few hundred meters along a river system. As-
sessing genetic diversity in B. glabrata by tar-
geting polymorphic SSRs in the genome 1$
less established (Jones et al., 1999; Mavares
et al., 2000), but has been shown to be a pow-
erful tool for linkage analysis for certain traits
in other species. The purpose of this study
was to use RAPD-PCR and microsatellite
variation in assessing the genetic diversity
among В. glabrata snails collected in a single,
small field site. Our goal was to determine the
sensitivity of each method to detect genetic
diversity in a single population. We have be-
gun by examining genomic polymorphisms on
a small scale in order to better understand
patterns of variation where geographic dis-
tance is not a complicating factor. By begin-
ning on a small scale before applying these
methods to analyze snails from a broad geo-
graphic region, we will be better able to choose
markers and sampling protocols for larger-
scale analyses. Furthermore, the occurrence
of previously identified RAPD markers that
segregate with resistance to $. mansoni infec-
tion in laboratory maintained snails (Knight et
al., 1999) was assessed in field collected
snails for the first time in this study.

MATERIALS AND METHODS
Snails

Initial standardization of techniques was per-
formed on laboratory-maintained B. glabrata
snails from two well-established stocks: (1) the
BS-90 snails, a pigmented, parasite-resistant
stock that was isolated from Salvador, Brazil
(Paraense 8 Correa, 1963), and (2) the M-line
snails, an albino stock highly susceptible to S.
mansoni infection (Newton, 1953).

Field-collected B. glabrata snails were ob-
tained from a schistosomiasis endemic area in
Corrego de Melquiades, Governador Valadares
municipality, Minas Gerais, Brazil. The epide-
miology of schistosomiasis in this area has
been documented by others (Kloos et al., 2001;
Bethony et al., 2001). Snails (> 5 mm shell
dia.) were collected from a single fishpond
(approx. 40 т? surface area) fed by a single
small stream. Before being transferred to the
laboratory, the snails were maintained in plas-
tic jars containing water collected from their
respective sites and fed lettuce or decaying
aquatic vegetation. To detect trematode infec-
tion in field-collected snails, the snails were
placed individually in water in 24-well plates
and exposed to incandescent light for 1-2 hr at
room temperature. The water was then exam-
ined under a dissecting microscope for the
presence of cercariae. Snails exhibiting signs
of trematode infection were excluded from the
DNA analysis.

DNA Isolation

Snails were placed overnight in water con-
taining 0.1 mg/ml of ampicillin for preparation
for DNA extraction. The shells were removed
after gently crushing the snails between two
glass slides and, in the case of field-collected
snails, the tissues further examined micro-
scopically for evidence of trematode infections.
The headfoot of individual snails was severed
from the posterior region of the body with a
sharp scalpel blade, then the headfoot either
used immediately for DNA extraction, or frozen
in liquid nitrogen and stored at -70°C until re-
quired. DNA was extracted as described previ-
ously (Knight et al., 1998). Genotypes of
individual laboratory and field-collected snails
were initially established by RAPD-PCR using
the 10-mer primers OPM-04 and OPZ-11, as
described by Knight et al. (1999).

GENETIC VARIATION IN BIOMPHALARIA GLABRATA 151

Construction of Small Insert Genomic Library

Small insert genomic DNA libraries were con-
structed either by using restriction enzyme Sau
3Al to generate small fragments (0.5-2.0 kb), or
by multiple enzyme digestions, as described by
Ostrander et al. (1992). Restriction enzyme di-
gestions were performed using 10 ид genomic
DNA from an individual BS-90 snail according to
manufacturer’s instructions (Life Technologies).
DNA concentration was determined by spotting
samples (1 ul) on agarose plates impregnated
with ethidium bromide (0.2 pg/ml) and com-
pared with DNA samples of known concentra-
tion under UV light. The DNA fragments
generated by Sau ЗА! digestion were ligated into
Bam HI linearized pUC-18 vector (Amersham
Pharmacia Biotech) under standard conditions.
In the case of DNA fragments generated by
multiple enzyme digestions, blunt-ended frag-
ments, filled in by a standard method using the
Klenow fragment of DNA polymerase, were li-
gated into the Sma | site of pUC-18.

Transformation of heat-inactivated ligation re-
actions into E. coli strain DH5a was performed
according to manufacturer’s instructions (Life
Technologies). Libraries were plated on LB agar
(Invitrogen) containing ampicillin, X-gal, and
IPTG and stored with 15% glycerol at —70°C
until required. For screening, libraries were
thawed, diluted and plated at a density of 2,000
colonies/plate (180 mm dia.) onto LB agar con-
taining ampicillin, and colonies screened by lift-
ing onto nitrocellulose filters according to
manufacturer's instructions (Schleicher 8
Schull).

Baked filters were hybridized under moderate
to low stringency conditions (Benton & Davis,
1977) using y[*?P] labeled oligonucleotides con-
taining -[АТ] о, -[СТ]о, and -[AAT], repeats
(Pharmacia). Positive colonies were detected
by autoradiography on Kodax X-Omat film at —
70°C for 2 days using intensifying screens.
Plasmid DNA was isolated from positive clones
using the Wizard plasmid DNA kit (Promega)
from overnight cultures grown in LB medium
containing 0.1mg/ml amplicillin. Nucleotide se-
quences of recombinant plasmids were deter-
mined using M13 primers (forward and reverse)
by the dideoxy-chain termination Sequenase
Kit (Amersham). From this information, primers
flanking potential microsatellite sites were de-
signed and obtained commercially (Genosys).
University of Wisconsin Genetics Computer
Group (UWGCG) software (Deveraux et al.
1987) was used to mine existing sequences of

potential B. glabrata microsatellite loci from
EST sequences in GenBank (dbEST).

Detection of Microsatellite Variation and RAPD-
PCR Analysis

Genetic diversity between snails was as-
sessed by RAPD-PCR analysis, using 19 arbi-
trary primers, and by variations in microsatellite
loci. Variations in microsatellite loci were evalu-
ated by PCR using primer sets flanking either
15 previously described loci (Jones et al., 1999;
Mavares et al., 2000) or 45 new loci identified
for this study. Genomic DNA was amplified us-
ing ‘touchdown’ PCR (Don et al., 1991) with
oligonucletide primer pairs (17—19 mers) ob-
tained from Sigma-Genosys Ltd. The negative
control was sterile distilled water. PCR was per-
formed in a total volume of 5.0 ul containing 5
ng of DNA template, 0.5 ul of 10X PCR buffer
(same as in 10X RAPD buffer), 1.0 pl of 1mM
dNTPs, 0.25 ul a-*°S dATP (Specific Activity,
1000Ci/umol Amersham Pharmacia Biotech,
U.K.) and 0.002 units of Taq DNA polymerase
(Promega, Wisconsin). “Touchdown” PCR was
performed as follows: denaturation, 30 sec at
95°C; annealing, 30 sec (temperature was de-
creased by 1°C every two cycles from an initial
temperature of 10°C above optimal Ta as deter-
mined from the manufacturers for the new 45
primer pairs or as previously published by
Jones et al. (1999) and Mavares et al. (2000),
then held at optimal annealing temperature for
20 cycles; extensions were at 72°C for 30 sec.
Stop buffer (1.0 ul, 0.1% xylene cyanol 0.1%
bromophenol blue, 10mM EDTA in 95% deion-
ized formaldehyde) was then added to each re-
action. Amplified alleles were separated by
electrophoresis on 6% urea/polyacrylamide se-
quencing gels in conjunction with known stan-
dards: (1) a 25 bp DNA ladder (Invitrogen)
end-labelled with у-3?Р; and (2) a sequencing
ladder of pUC-19 generated using the universal
M13 (reverse) primer. Gels were fixed in 10% (v/
У) acetic acid and 10% (v/v) methanol for 15
minutes at room temperature prior to drying at
80°C. Dried gels were set up for autoradiogra-
phy on X-ray film (X-OMAT, Kodak) overnight at
room temperature without intensifying screens.
RAPD-PCR using 25 ng of genomic DNA was
performed as previously described (Larson et
al., 1996). The control was distilled water, and
amplified samples were resolved by gel electro-
phoresis on agarose gels (1.2% w/v). The
bands were stained by ethidium bromide and
visualized by UV transillumination.

152 ROWE ET AL.

AMOVA Analysis

Phenotypic variation of RAPD products
(primers OPX-6, OPM-19, OPAW-07) was in-
vestigated by analyses of molecular variance
(AMOVA; Excoffier et al., 1992) implemented in
the computer package ARLEQUIN Ver. 2.001
(Schneider et al., 2000). Only those bands that
could be unequivocally scored across all
samples were included in the analysis. Subse-
quent AMOVA analysis proceeded with 13
markers for three BS-90 snails, three M-line
snails and 36 field-collected snails. A matrix of
Euclidian square distances was computed us-
ing the pairwise difference method. This matrix
was used for the analysis of genetic structure
including partitioning of variation (A) within the
three populations, (B) among the three popula-
tions, and (C) among groups of populations
(i.e., lab strains versus field strain). Statistical
significance of variance components was as-
sessed with 10,000 random permutations.

RESULTS

Assessing Genetic Differences Between Labo-
ratory Snails from Two Different Stocks

Before embarking on the analysis of field
population snails, we tested our ability to detect
genetic differences, by variations in micro-
satellite loci and RAPD-PCR analysis, be-
tween laboratory-maintained snails. DNA
profiles of two well-established laboratory
stocks of B. glabrata were used as reference
samples to test all RAPD-PCR primers and
microsatellite primer sets for consistency in
amplification. These snails, from stocks BS-90
and M-line, are fully resistant or susceptible,
respectively, to the NMRI strain of S. mansoni
used in our laboratory. Previously, we have
shown considerable genetic homogeneity be-
tween individuals within each stock by RAPD-
PCR with numerous primers (Knight et al.,
1999; Ittiprasert et al., 2003). Here we deter-
mined the occurrence of polymorphisms be-
tween them by examining variations at 60
microsatellite loci and by RAPD-PCR using 19
random primers. The new loci were identified
either by screening a small insert library, or in
silico by mining for simple sequence repeats
(SSR) from B. glabrata sequences deposited
in GenBank.

Primer sets and annealing temperatures (Ta)
used for the present study for all 60 micro-

satellite loci are given in Table 1. Variations in
the microsatellite loci examined between the
two laboratory maintained snails, listing the
number of alleles and their observed sizes, are
shown in Table 2. Of the six loci previously de-
scribed by Jones et al. (1999), null alleles (N/A)
were observed for three of these (uBg1, Bgu10
and Bg 16) in both snails, and unscorable mul-
tiple products (MP) were detected in both
stocks with primer sets flanking locus Bgu8.
For the two other loci (uBg2 and Bgy 15) single
alleles that were polymorphic between the two
laboratory snails were identified. Allelic size
ranges observed were more consistent with
the size seen in the BS-90 stock compared to
M-line stock. For locus Bgy 15, the single allelic
band of 178 bp (previously reported) appeared
as two bands (176 bp and 178 bp) in the BS-
90 snail, and a single band of 161 bp in the M-
line snail.

For the nine microsatellite loci reported pre-
viously by Mavares et al. (2000), two (BgE2,
BgE3) were undetected (null alleles), and
primer sets corresponding to locus BgC8 pro-
duced multiple products in both snail stocks.
Of the remaining six loci, all except one (locus
BgE5) were detected in both stocks. The
primer set corresponding to the BgE5 locus
produced no detectable band in the M-line
snail, but produced two allelic bands of 204 bp
and 234 bp in the BS-90 snail, thus demon-
strating polymorphism in this locus between
the two laboratory strains. Similarly, genetic dif-
ferences between the two stocks were de-
tected with the primer sets corresponding to
loci BgC7, BgE1, BgE4 and BgE6. For these
sites, the allelic size range detected fell within
the expected size range as previously de-
scribed (Mavares et al., 2000), with the BS-90
snail showing better correspondence in size
expected than the M-line stock.

Data in Table 2 describe the presence of the
45 previously unreported, potential micro-
satellite loci (BGMSCA series, BGMSAT se-
ries, and BGMSGATA series). Included are the
number of alleles and observed sizes amplified
from the two laboratory stocks using primers
flanking these loci. Of these, variations were
detected in 20 of the sites between the two
laboratory snails. Null alleles were ob-
served with 14 primer sets (combined data
from BS-90 and M-line snails), and multiple
products were detected with primer sets corre-
sponding to four sites.

The results of the RAPD-PCR comparison,
using 19 random primers, are summarized in

153

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GENETIC VARIATION IN BIOMPHALARIA GLABRATA

Table 3 (column 3). All but one primer (OPV-20)
revealed polymorphisms between strains. In all
cases, the polymorphisms detected were ma-
jor and readily observed by ethidium bromide
staining of agarose gels. Representative ex-
amples of the DNA profiles from RAPD-PCR
using two random primers are shown in Figure
1A and B. Major differences could be detected
with both primers in the RAPD profiles of both
the BS-90 and M-line snails. For example,
primer OPAW-07 reveals a major band of 700
bp that is absent from the M-line snail. Using
primer OPZ-05 a band of 1.0 kb was detected
only in the M-line snail.

In summary, it was clear that both micro-
satellite variation and RAPD-PCR analysis
were adequate in assessing polymorphisms
between snails from the two laboratory stocks.
Of the microsatellite sites analyzed, 26 loci
were found to be polymorphic, while 8 sites
were monomorphic between the two stocks.
With RAPD-PCR, all but one primer revealed
polymorphisms between the two lab stocks.

Assessing Genetic Diversity in Field-Collected
Snails from a Single Locality

After conditions were optimized, DNA poly-
morphisms were evaluated following the estab-
lished procedure among field isolates of

157

B. glabrata. Results in Table 4 describe micro-
satellite variations (observed allelic sizes and
number of alleles) between 20 different indi-
vidual field collected snails for 26 primer sets.
The primer pairs used for field evaluation were
those that gave consistent results either when
amplifications were performed at different
times or by different investigators on the same
field-collected snail DNA template. Primer sets
that failed to show amplified products (null alle-
les) when laboratory snail stock DNA was used
as template were omitted from the field study.
In addition, 24 primer sets that produced mul-
tiple products when field collected samples
were analyzed have not been included in Table
4. In comparison to the laboratory snails, field-
collected snails showed significant variations
for several of the loci examined. For example,
for two loci previously identified by Mavares et
al. (2000) (BgE1 and BgE5), we detected more
intrastrain variation at these loci compared to
the laboratory stocks, where only one allele
was detected. Correspondence in the observed
size range to the expected size was consistent
between laboratory-maintained and field-col-
lected samples, although the number of alleles
varied between them for the majority of loci
analyzed. Invariant alleles (in size and number)
between the laboratory snails and field isolates
were found with only two loci (ВСМ$САО5 and

TABLE 2. Observed allelic sizes and number of alleles for 60 microsatellite loci in resistant
(BS-90) and susceptible (M-line) laboratory-maintained snails.

BS-90 (R) M-line (S)
Observed Number of Alleles Observed Number of Alleles

Locus Sizes (bp) Sizes (bp)

mBg1 N/A N/A N/A N/A
mBg2 249 1 259 1
Bgm8 MP MP MP MP
Bgm10 N/A N/A N/A N/A
Bgm15 176, 178 2 161 1
Bgm16 N/A N/A N/A N/A
BgC6 N/A N/A N/A N/A
BgC7 321 1 325 1
BgC8 MP MP MP MP
BgE1 1314137 2 103, 135 2
BgE2 N/A N/A N/A N/A
BgE3 N/A N/A N/A N/A
BgE4 205 1 177 1
BgE5 204, 234 2 N/A N/A
BgE6 539 1 289 1

(Continues)

158 ROWE ЕТ AL.

(Continued)
TIA A A A Е.
BS-90 (R) M-line (S)
Observed Number of Alleles Observed Number of Alleles
Locus Sizes (bp) Sizes (bp)
BGMSCAO1 126, 128 2 MP MP
BGMSCAO2 N/A N/A N/A N/A
ВСМ$САОЗ 158, 160 2 166, 168 2
BGMSCAO4 135, 164 2 164 1
BGMSCAO5 152 1 152 1
ВСМ$САОб 152,154 2 152,154 2
BGMSCAO7 149 1 153 1
BGMSCA08 N/A N/A N/A N/A
BGMSCA09 155 1 il 1
BGSMCA10 N/A N/A N/A N/A
BSMGCA11 MP MP MP MP
BGMSCA12 N/A N/A N/A N/A
BGMSCA13 N/A N/A N/A N/A
BGMSCA14 199 1 193, 195 2
BGMSCA15 224 1 190 1
BGMSCA16 334 1 334 1
BGMSCA17 N/A N/A N/A N/A
BGMSCA18 162 1 162 1
BGMSCA19 151 1 154 1
BGMSCA2O 162 1 164 1
BGMSCA21 203 1 N/A N/A
BGMSCA22 154 1 154 1
BGMSCA23 112 1 112 1
BGMSCA24 180, 182 2 185, 187 2
BGMSCA25 134 1 138 1
BGMSCA26 on 1 153 1
BGMSCA27 196 1 204, 212 2
BGMSATO1 N/A N/A N/A N/A
BGMSATO2 130 1 N/A N/A
BGMSAT03 N/A N/A N/A N/A
BGMSAT04 N/A N/A N/A N/A
BGMSATO5 140, 146 2 140, 146 2
BGMSATO6 MP MP MP MP
BGMSATO7 MP MP MP MP
BGMSAT08 220 230 2 224, 234 2
BGMSATO9 307 1 332 1
BGMSAT10 136 1 136 1
BGMSAT11 MP MP MP MP
BGMSAT12 335 1 N/A N/A
BGMSAT13 319,333 2 333 1
BGMSAT 14 N/A N/A N/A N/A
BGMSAT15 431 1 431 1
BGMSAT16 232 1 1851233 2
BGMSGATAO1 MP MP MP MP

BGMSGATAO2 372 1 374 1

oo оля ._————

СЕМЕТ!С VARIATION IN BIOMPHALARIA GLABRATA

TABLE 3. Detection of polymorphisms (+/-) by RAPD-PCR analysis among
laboratory stocks and among field-collected snails.

Lab Interstrain Field Intrastrain

A

OPAW-07

B

RAPD Primers Sequence 5’ to 3’ Variation (+/-) Variation (+/-)
OPAL-10 AAGGCCCCTG + -
OPAJ-17 ACCCCCTATG + -
OPAV-11 GACCCCGACA + -

OPT-11 TTCCCCGCGA + -
OPS-04 CACCCCCTTG + +
OPV-12 АССССССАСТ + т
ОРАО-09 АСТСССССТС + Ae
OPX-04 CCGCTACCGA + +
OPM-18 CACCATCCGT + -
OPAJ-08 GTGCTCCCTC + -
ОРАР-08 АСССССАСАС + -
OPAW-07 AGCCCCCAAG + +
OPV-20 CAGCATGGTC - -
OPM-19 CCTTCAGGCA + =
OPX-06 ACGCCAGAGG + +
OPV-15 CAGTGCCGGT E +
OPZ-20 ACTTTGGCGG ях -
OPZ-05 TCCCATGCTG + +
OPAH-04 СТСССАСАС + -

OPZ-05

FIGURE 1. RAPD-PCR amplification of BS-90 (B) and M-line (M) snail DNA
with random primers (A) OPAW-07 and (B) OPZ-05. For molecular weight mark-
ers (MW), a 100 bp ladder was used.

159

160 ROWE ET AL.

TABLE 4. Observed allelic sizes and number of alleles for 44 microsatellite loci
of field-collected В. glabrata snails.

Number of Observed Allelic Sizes

Locus Snails (bp) Number of Alleles

mBg2 19 215 1
1 259) 275 2

Bgm15 13 175 1
5 175.176 2
1 176 1
1 N/A

BgC7 4 326, 327 2
15 327 1
1 N/A

BgE1 2 99, 103 2
3 99, 107 2
1 99 1
т 103 1
7 N/A

BgE4 14 181 1
6 181, 209 2

BgE5 4 230 1
1 230, 234 2
1 230, 242 2
4 234 1
4 234, 242 2
6 N/A

BGMSCAO1 20 126, 128 2

ВСМ$САОЗ 19 166, 168 2
1 N/A

BGMSCA05 20 152 1

BGMSCA14 18 197 1
2 197, 198 2

BGMSCA18 20 162 1

BGMSCA19 19 160 1
1 N/A

BGMSCA20 20 N/A

BGMSCA22 20 152 1

BGMSCA23 20 N/A

BGMSCA25 20 144 1

BGMSCA26 20 N/A

BGMSCA27 17 200 1
3 N/A

BGMSATO5 14 130, 136 2
6 N/A

BGMSAT08 10 230 1
5 230,231 2
3 231 1
2 N/A

BGMSAT10 20 154 1

BGMSAT12 12 335 A
8 N/A

BGMSAT15 4 429 1
1 429, 437 2
2 437 1
13 N/A

BGMSAT16 11 184 1
6 184,211 2
3 N/A

BGMSGATAO2 19 520,521 2
1 N/A

GENETIC VARIATION IN BIOMPHALARIA GLABRATA 161

bp EMMA Bj ld isolates ити

FIGURE 2. Microsatellite variation of locus BGMSCA14. Sequencing ladder of pUC-19 was used as
standard. B and M represent lanes showing allelic bands amplified at this locus for the BS-90 and M-
line laboratory stocks, respectively. Amplification at the same locus for DNA from 29 field-isolated
snails, isolated from a single site, is shown. Note the difference in number and size of alleles between

the lab stocks and among the field-collected samples.

BGMSCA18), where single allelic size bands of
152 bp and 162 bp, respectively, were detected
with all DNA samples analyzed.

In several cases, multiple products were ob-
tained for the same loci in snails from both
laboratory and field, that is, Bgu8 and BgC®.
Although null alleles were infrequent for the
majority of loci, these were associated more
with the M-line snail than with either the BS-90
or field-collected snails. A representative ex-
ample of microsatellite variation in the locus
BGMSCA14 is shown in Figure 2. As can be
seen, polymorphisms in this locus were de-
tected between the laboratory-maintained
snails, manifested by differences in sizes of

MW

Field isolates

allelic bands in the BS-90 (199 bp) compared
to the M-line snail (193 bp and 195 bp). Diver-
sity within this locus for field-collected snails
was also clearly visible. In this case, most
samples examined from this population (29
snails) either displayed allelic size bands of
197 bp and 198 bp or, as seen in one snail
from this population, a single allelic band of 198
bp was detected.

A summary of the results obtained using
RAPD-PCR to assess genetic diversity among
the field-collected snails is shown in Table 3
(column 4). Of the 19 random primers utilized,
we observed intra-strain variation with nine of
them. Figure 3 shows a representative ex-

Field isolates

OPAW-07

OPZ-05

FIGURE 3. RAPD-PCR amplification of DNA from individual field isolates of В. glabrata snails, using
random primers (A) OPAW-07 and (B) OPZ-05. For molecular weight markers (MW), a 100 bp ladder
was used.

162 ROWE ET AL.

ample of RAPD-PCR analysis of field-collected
snails, using random primers OPAW-07 and
OPZ-05. Of the 19 DNA samples analyzed,
both primers revealed major intrastrain variation
among them. For primer OPAW-07, two bands
(doublet) of 700 bp and 750 bp were present in
most snails analyzed, but a single 750 bp band
was seen in five snails examined. Also, with this
primer, a major 500 bp band seen in most
snails was absent in two analyzed (lanes 1 and
2). Variations with primer OPZ-05 were clearly
visible in several bands. For example, polymor-
phisms were detected in bands at 550 bp and
750 bp in these field-collected snails.

Assessing Population Structure Based on
RAPD Data

The analysis of molecular variance (AMOVA)
showed that most of the variation resides within
populations (68%) and the permutation tests
indicated that the within-population differences
are highly significant (P < 0.00001). The varia-
tion among populations accounts for 54% of
the total variation (P = 0.0283). However, the
difference between the two lab-strain popula-
tions and the field-strain population (-22% of the
total variation) is not significant (Table 5). A
pairwise AMOVA differentiation test between
populations showed that all three comparisons
were significant (BS-90 snails vs. M-line snails
0.02712; BS-90 snails vs. field-collected snails
0.00157, and M-line snails vs. field-collected
snails 0.00161).

DISCUSSION

The genetics of both the snail host and the
parasite determine the outcome of the mollus-
can stage of parasite development. For this rea-

son, more is being done to understand how the
genetic structures of the parasite and snail host
populations affect transmission and epidemiol-
оду of this complex disease. In this study, we
have compared the relative sensitivities of two
DNA fingerprinting methods to assess genetic
heterogeneity among snails from a single, re-
stricted field site in a schistosomiasis-endemic
area. This analysis was done following estab-
lishment of the profiles of representative snails
from two well-established laboratory stocks that
are either resistant or susceptible to parasite
infection, and on which considerable molecular
profile data already exist (Knight et al., 1999;
Ittiprasert et al., 2003). The genome wide scan-
ning tools we used in this study (microsatellite
variation and RAPD-PCR) are particularly use-
ful for studying organisms in which only limited
molecular information is available. From our
results it was clear that both methods showed
significant polymorphisms between snails in
this restricted field site.

In previous work, we showed by RAPD-PCR
the segregation of two markers (1.2 kb and 1.0
kb) with the inheritance of adult resistance
(Knight et al., 1999), a known Mendelian single
gene {гай т В. glabrata (Richards, 1984). In-
cluded in our study here was the first analysis
to assess the potential frequency of these
markers in field-collected snails from a known
endemic area for schistosomiasis. We had
hoped that by extending these studies to the
field, the presence, if any, of resistant snails
and their role in the dynamics of transmission
can start to be evaluated. However, neither
marker was detected in any of the field-isolated
snails we analyzed. Whether these markers
universally segregate with resistance in all B.
glabrata populations, or whether resistant
snails were absent from the present population
studied is not known.

TABLE 5. Summary of AMOVA analysis. Statistics include: degrees of free-
dom (df), sum of squares (SSD), variance-component estimates (CV), and
percentages of the total variance (% Total) contributed by each component.

Source of Variation df
within populations 3
among populations 1
among groups 1

SSD CV % Total
14.500 0.39189 68.19
1.625 0.30828 53.61
1.730 -0.12511 21.70:

“not significant after 100172 permutations; the negative value reflects that this statistic
is actually a covariance where negative values can occur when the actual values are

close to zero (Excoffier et al., 1992).

GENETIC VARIATION IN BIOMPHALARIA GLABRATA 163

Several explanations may account for the
absence of the 1.2 kb and 1.0 kb markers in
our field population. One possible explanation is
that the markers do not universally segregate
with resistance in all B. glabrata populations.
Differences between the M-line, BS-90 and the
field population may also reflect intraspecific
diversity in B. glabrata as a whole. Previously,
Paraense (1959) found reduced infertility
among some populations of B. glabrata, sug-
gesting there may be considerable genetic het-
erogeneity in this species. A second
explanation that might account for the absence
of the 1.2 and 1.0 kb markers in the field popu-
lation would be evolution in the laboratory
stocks. It is possible that when these stocks
were founded that alleles that were rare in
natural populations became more common,
either by chance or because of the removal of
selection against them. Laboratory stocks can
be extremely useful for working out patterns
that are difficult to see in natural populations
because there may be more pronounced and
less variable responses, that is, a more favor-
able ratio of signal to noise. However, a poten-
tial pitfall of using laboratory stocks is that they
may not be reflective of what actually happens
in nature. Our findings therefore point to a criti-
cal need for further comparisons of laboratory
and field populations of B. glabrata. Our results
also show that not all microsatellite primer sets
that were developed using laboratory stock
DNA were useful when field collected snails
were analyzed, thus indicating that more work
needs to be done on field collected snails in
developing more representative markers.

Finally, it is possible that the markers of resis-
tance were simply absent from the sample we
examined. Experimental studies suggest that
resistance may be costly to snails (Minchella &
LoVerde, 1983). Mulvey & Vrijenhoek (1984)
have also proposed that genetic drift in popula-
tions of B. glabrata might produce patchy sus-
ceptibility and resistance over time and space.

For population genetic analysis of schisto-
some snail hosts by RAPD-PCR, evidence for
the existence of both inter- and intrapopulation
diversity has been reported. Vidigal et al. (1994),
using four primers, showed that diversity be-
tween snail populations can readily be as-
sessed by this method, but it was not as
revealing among snails from the same geo-
graphic region. With another Biomphalaria spe-
cies, however, Hoffman et al. (1998) reported
that this method could reveal a high frequency
of genetic diversity in B. pfeifferi populations

that resided only a short distance apart in the
same river. Likewise, studies examining
Bulinus snails involved in transmission of S.
haematobium have shown similar intrapopula-
tion genetic diversity by RAPD-PCR (Davies et
al., 1999). Our study clearly shows that RAPD-
PCR analysis can be a legitimate tool to reveal
polymorphisms between snails collected from a
single site, in which mating would presumably
be restricted, and to study population structure
of groups of populations.

As stated earlier, limitations to the RAPD-
PCR method for population studies exists, es-
pecially because it allows for the detection of
only dominant alleles. In earlier studies on labo-
ratory snail stocks, we detected reproducible
polymorphisms between schistosome-resis-
tant and -susceptible snails with 90% of the
primers utilized in the RAPD-assay. In the
present study, we found that by using RAPD-
PCR a somewhat higher frequency of diversity
is seen between our laboratory stocks, com-
pared to variation observed in the field isolated
snails — a not altogether unexpected finding.
This may reflect the fact that, compared to the
field snails, the two laboratory snails used here
originated from very different geographic lin-
eages; the albino M-line was derived by a
cross between Puerto Rican and Brazilian iso-
lates (Newton, 1953), whereas the wildtype
BS-90 snails are descendents of an original
field collection from Brazil (Paraense & Correa,
1963).

In addition to the B. glabrata microsatellite
loci previously described by others (Jones et
al., 1999; Mavares et al., 2000), we now show
results of genetic variation in 45 new
microsatellite loci. Population genetic studies
with microsatellites for other schistosome
transmitting species have been carried out,
especially for Bulinus sp., measuring param-
eters such as mating systems and geneflow
(Viard et al., 1996; Stothard et al., 2001). Invari-
ant loci detected were further evaluated by
SSCP to determine the unequivocal evidence
of homoplasy (Angers et al., 2000). Our study
shows that two loci (ВСМ$САО5 and
BGMSCA18) appear to be fixed between both
laboratory stocks and in field snails. Whether
the same size bands are identical by descent
remains to be seen, but nucleotide sequence
analysis of the same sized bands should re-
veal their relationship.

The value of data mining for SSRs in ex-
pressed sequence tags (ESTs) is evident from
the results of our study, allowing us to generate

164 ROWE ЕТ AL.

44 microsatellite primer sets. This data mining
strategy for SSRs has also been used with
success in plant molecular biology for identifi-
cation of potential microsatellite loci in barley,
maize, rice, sorghum and wheat (Kantety et
al., 2002). With current interest in a genome
project for B. glabrata, it is anticipated that the
availability of more sequence information from
this snail will increase the output of micro-
satellite markers, thus benefiting high density
mapping efforts. It is envisaged that high
throughput analysis of B. glabrata DNA will fa-
cilitate the development of other modern ge-
nome analysis tools, such as single nucleotide
polymorphisms (SNPs), another useful tool for
studying the diversity of complex genomes
(Wang et al., 1998).

It is not known what role SSRs play in MRNA
transcripts, because one would expect the
gene coding region of the transcript to remain
neutral. Polymorphisms, manifested by the ex-
pansion of tri-nucleotide repeats in the coding
region of MRNA, have been shown, however,
to play a role in certain human genetic disor-
ders, such as Huntington’s disease and mus-
cular dystrophy (Maat-Kievit et al., 2001;
Margolis et al., 2001). In humans, the most
abundant di-nucleotide repeat motif found is the
CA/GT repeat (Weber, 1990). In the new sites
described in this study, 15 repeat motifs were
CA/GT, but most were AT/TA rich. Compound
and interrupted repeats were also detected.

We found that null alleles were infrequent for
most microsatellite loci examined. In the case
of the loci described by Jones et al. (1999),
amplification of our laboratory snails showed no
product for three out of the six loci reported by
these investigators. We likewise observed null
alleles with several of the loci described by
Mavares et al. (2000) in BS-90 and M-line
snails. With the newly described loci we ob-
tained in silico, we were surprised to observe
null alleles for some of them, since these se-
quences reside in functional transcripts of the
snail. One explanation for this could be that
these sequences may not be readily detected
using the amplification conditions employed in
this analysis.

We found that several loci were non-
scorable, and amplification of these showed
multiple bands with sizes that deviated signifi-
cantly from the size expected. Because
microsatellites have been defined as highly
polymorphic single locus regions in the ge-
nome, we omitted primers that produced these
multiple products from our study. It is likely,

however, that these may constitute hyper-
variable sites in the B. glabrata genome. Nucle-
otide sequence analysis should help to clarify
the nature of these repetitive bands.

In summary, we found that RAPD-PCR is a
practical method for assessing population
structure of lab- and field-collected strains of
Biomphalaria glabrata. In addition, variations in
previously described and newly identified
microsatellite loci are useful for assessing ge-
netic differences between two laboratory main-
tained snails, and among snails collected from
a single field site. With the substantial variation
shown by microsatellite analysis in the field-
collected snails, however, it will be important to
select only those primer sets that do not dis-
play hypervariability for detailed population
studies. Before settling on primer sets to use
for studying snail population structure in other
geographic regions, it may be important to pre-
screen representative snails from that region.
The extensive variation in snails we showed
through microsatellite analysis may be magni-
fied by geographic distance, revealing even
more profound differences than what we dem-
onstrated even in this very restricted field col-
lection site.

ACKNOWLEDGMENTS

This work was supported in part by NIH
Grants Al-27777 and 42768. We thank Helmut
Kloos and Rodrigo and Andrea Correa-Oliviera
for coordination of snail collections in Brazil, as
well as Nithya Raghavan and André Miller for
their molecular help and critique of the manu-
script. We also gratefully acknowledge the lo-
gistical and administrative support from Jeff
Bethony and Phil LoVerde.

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Revised ms. accepted 27 March 2003

RESEARCH NOTES

MALACOLOGIA, 2003, 45(1): 169-174

А NEW PANAMIC SPECIES OF THE BIVALVE GENUS SEMELINA (SEMELIDAE)

Eugene V. Coan*

Department of Invertebrate Zoology and Geology
California Academy of Sciences, Golden Gate Park,
San Francisco, California 94118-4599, U.S.A.; gene.coan@sierraclub.org

ABSTRACT

A new species of Semelina, S. campbellorum, is described from the Panamic province,
differing from S. subquadrata (Carpenter, 1857) in having a longer, more tapered posterior
end, orthogyrate rather than opisthogyrate beaks, more lamellar commarginal ribs with fine
commarginal threads between the larger lamellae, and a longer, more confluent pallial sinus
that reaches the anterior adductor muscle scar. Semelina subquadrata is very similar to the
western Atlantic S. nuculoides (Conrad, 1841). An internal ligament has probably evolved
more than once within the Tellinoidea, and the Semelidae is probably polyphyletic. The
genus Semelina is one of several genera that are of somewhat uncertain position.

Key words: Semelina, Semelidae, Panamic province.

INTRODUCTION

In identifying material in connection with
preparation of a manual on the Panamic Bivalvia,
it was realized that there are two tropical eastern
Pacific species of the genus Semelina. Study of
Carpenter's type material of “?Montacuta”
subquadrata was necessary to be certain which
of the two species he described.

As first suggested by Maxwell (1991) in a talk
and accompanying abstract, the Semelidae as
presently constituted is probably polyphyletic.
He cited several pairs of genera with similar
external shell morphology, differing chiefly in
the presence of only an external ligament
(Tellinidae) or also having an internal resilifer
(Semelidae). Whereas some examples that he
cited may actually represent convergence in
general shell morphology, it is likely that an in-
ternal ligament has evolved more than once
among taxa now allocated to the Semelidae.
Indeed, Kamenev 8 Nadtochy (1999) demon-
strated that juvenile Macoma have a small in-
ternal ligament and are not very different from
species allocated to Abrina in the Semelidae.
Gustav Paulay (personal communication, 19
December 2002) has pointed out that some
IndoPacific species, mostly as yet unde-
scribed, now allocated to the tellinid genera
Exotica and Semelangulus actually have both

an external and an internal ligament, and some
of these are similar to the New World genus
Semelina, although only the three taxa dis-
cussed in this paper have been allocated to
Semelina in the literature. Additional studies are
clearly much needed to sort out the clades
within the Tellinoidea.

The following institutional abbreviations are
used here: BMNH, British Museum of Natural
History collection, The Natural History Mu-
seum, London, England; CAS, California Acad-
emy of Sciences, San Francisco, California,
U.S.A.; LACM, Natural History Museum of Los
Angeles County, Los Angeles, California,
U.S.A.; USNM, United States National Museum
collection, National Museum of Natural History,
Smithsonian Institution, Washington, D.C., U.S.A.

SYSTEMATIC TREATMENT
Semelina Dall, 1900: 986, 994

Type species (original designation): Amphi-
desma nuculoides Conrad, in Hodge, 1841:
347; Conrad, 1845: 73, pl. 41, fig. 7. Natural
Well, Duplin County, North Carolina; Duplin
Formation (sensu stricto), 3.2 Ma, late
Pliocene. Synonyms: Semele nuculoidea,
auctt., пот. null.; “Semele?” virginiana

*Mailing address: 891 San Jude Avenue, Palo Alto, California, 94306-2640, U.S.A.; also Research Associate, Santa
Barbara Museum of Natural History and the Natural History Museum of Los Angeles County.

170 СОАМ

Meyer, 1888: 143; $. nuculoidea lirulata Dall,
1900: 994; $. sirulata Dall, auctt., nom. null.

Diagnosis

Small, longer anteriorly; internal ligament in a
short to elongate resilifer, not produced ven-
trally beyond the hinge plate; external ligament
in a narrow groove; right valve with elongate
anterior and posterior lateral teeth, the left
valve fitting into the grooves between them and
the dorsal margin; lunule and escutcheon
present; right valve with a narrow, inconspicu-
ous anterior cardinal and a large posterior car-
dinal; left valve with a large anterior cardinal
and a narrow posterior cardinal that defines the
posterior edge of the resilifer; pallial sinus
large, deep. Sculpture of fine, dense
commarginal ribs.

This genus first appears in the early Miocene
Chipola Formation of Calhoun County, Florida,
with Semelina cythereoidea Dall, 1900 (p. 994,
pl. 44, fig. 5). The eastern U.S./western Atlantic
Semelina nuculoides is treated by Campbell
(1993: 42, pl. 17, fig. 156), Díaz M. 8 Puyana H.
(1994: 97, pl. 28, fig. 268), Gardner (1944: 102—
103, pl. 17, figs. 18-21), Lamy (1915), Redfern
(2001: 231-232, pl. 99, fig. 948), and Rios
(1994: 275, pl. 94, fig. 1352). It now occurs
from North Carolina to the West Indies and
Brazil, and is recorded as early as the early
Pliocene (3.8 Ma) in the southeastern U.S.A.

Semelina subquadrata (Carpenter, 1857)
Figures 1-4

“?Montacuta” subquadrata Carpenter, 1857a:
248, nom. nud.; Carpenter, 1857b: 113;
Brann, 1966: 35, pl. 10, fig. 162

“?Mysella” subquadrata (Carpenter) — Dall,
1899: 881

Rochefortia subquadrata (Carpenter) — Hertlein
8 Strong, 1947: 135 [in part; their specimen
from Bahía Santa Inez, Baja California Sur, 1$
a Mysella — CAS 162714; Loc. 17746]

Semelina nuculoides Conrad, non Conrad, in
Hodge, 1841 — Hoffstetter, 1952: 41

Mysella subquadrata (Carpenter) — Keen,
1958: 107

Semelina subquadrata (Carpenter) — Olsson,
1961: 375, pl. 66, fig. 11; Keen, 1968: 395, fig.
11, 400; Keen, 1971: 259, 260, fig. 661

Description

Ovate-elongate, evenly inflated; beaks al-
most at posterior end, opisthogyrous; posterior

end subtruncate; surface with fine, even,
rounded commarginal ribs, some of which be-
come lamellar on posterior slope, while others
die out before reaching the posterior slope; pal-
lial sinus ending well short of anterior adductor
muscle scar and confluent with the pallial line
for only about a third of its length; shell white,
sometimes with a pinkish flush. Length to
6.6 mm.

Type Material & Locality

BMNH 1857.6.4.503/1, lectotype herein, the
larger specimen, a right valve, length = approxi-
mately 3.2 mm (Fig. 4); BMNH 1857.6.4.503/2,
paralectotype, a smaller left valve measuring
approximately 1.2 mm. Both specimens re-
main glued to Carpenter's original glass slide.
Mazatlán, Sinaloa, México (32.2°N); Frederick
Reigen. A lectotype is designated because the
small glue-covered left valve is too small to be
reliable identified.

Distribution

Bahía Magdalena, Pacific coast of Baja Cali-
fornia Sur (24.5°М) [LACM 49-234.1], into the
Golfo de California as far north as Isla
Danzante, Baja California Sur (25.8°N)
(Skoglund Collection), and Estero Soldado,
Sonora (27.9°N) [LACM 73-5.45], México, to
Manglaralto, Guayas, Ecuador (1.9°S) [CAS
162257]; probably as far south as Punta Santa
Elena, Guayas, Ecuador (2.2°S), where it has
been recorded as a subfossil (Hoffstetter, 1952:
41); Isla Marchena, Islas Galapagos, Ecuador
(0.3°N) [LACM 34-285.5]; from the intertidal
zone to 220 т (mean = 43.1 т; п = 17); no
bottom types noted on labels. | have seen 20
lots; Carol Skoglund provided data for 4 addi-
tional lots.

Referred Material

Stations in México: LACM 49-234.1 — Bahia
Magdalena, Baja California Sur — 33 m;
Skoglund Collection — Isla Danzante, Baja
California Sur — 30-45 т; Skoglund Collec-
tion — Los Frailes, Baja California Sur — 50—
66 m; LACM 73-5.45 — Estero Soldado,
Sonora — intertidal zone; Skoglund Collection
— Bahía San Carlos, Sonora — 15-30 т;
BMNH 1857.6.4.503 — Mazatlán, Jalisco —
type lot; CAS 161375 — Mazatlán, Jalisco —
“dredged”; Skoglund Collection -
Cuastecomate, Jalisco — 12-30 m; LACM
38-263.1, CAS 164336 — Black Rocks,

ANEW PANAMIC SEMELINA 171

FIGS. 1-4. Semelina subquadrata. FIG. 1. External view о left valve; length, 4.6 mm. SBMNH 348120;
Balboa, Panamá; commercial dredgings; ex Skoglund Collection. FIG. 2. Same specimen: close-up of
sculpture on border between central and posterior slopes; width of ribs approximately 60 um. FIG. 3.
Sketch showing hinge, adductor muscle scars, pallial sinus, pallial line, and cruciform muscle scars
of right valve; CAS 161375; Mazatlán, Sinaloa, México; length, 5.1 mm. FIG. 4. Lectotype of
“?Montacuta” subquadrata Carpenter; BMNH 1857.6.4.503/1, right valve approximately 3.2 mm long.
FIG. 5. Semelina nuculoides. External view of left valve; length, 4.5 mm. SBMNH 348121: 40 miles SE
of Charleston, South Carolina; 60 m; ex Campbell Collection.

FIGS. 6-7. Semelina campbellorum. FIG. 6. Holotype; external views of left and right valves; SBMNH
348119; Bahía de Santiago, Colima, México; length, 4.4 mm. FIG. 7. Paratype; SBMNH 348119: external
view of left valve; same station; length, 3.7 mm.

FIGS. 8, 9. Semelina campbellorum. FIG. 8. Same specimen as Fig. 7; close-up of sculpture оп
border between central and posterior slopes; distance between higher ribs approximately 100 um.
FIG. 9. Sketch showing hinge, adductor muscle scars, pallial sinus, pallial line, and cruciform muscle
scars of right valve; LACM 37-218.2; Bahía San Ignacio, Sinaloa, México; length, 5.0 mm.

Jalisco — 37 т; LACM 65-16.18 — Bahia
Banderas, Jalisco; LACM 33-136.1 -
Petatlan, Guerrero — 11 т; LACM 34-241.1 —
Petatlan, Guerrero — 183-256 т; LACM
38.9.12 — Bahia Guatulco, Oaxaxa — 73-128 т

Stations in Nicaragua: CAS 164337 — Corinto,
Chinandega — no depth recorded; CAS
164338 — Corinto, Chinandega — no depth
recorded

Stations in Costa Rica: LACM 72-19.39 — Bahia
de Salinas, Guanacaste - 1.5-11 т; LACM
80-60.17 — Cabo Santa Elena, Guanacaste —
intertidal zone; LACM 86-26.33 — Playa
Nancite, Guanacaste — beach drift; Skoglund
Collection — Playa Tamarindo, Guanacaste —
6-15 m; LACM 84-152.5 — Bahia Ballena,
Puntarenas — 15-21 т.

Stations in Panama: LACM 39-259.2 — Isla
Ladrones, Chiriquí — 99 т, Skoglund Collec-
tion — Balboa, Panama

Stations in Ecuador: CAS 162257 -
Manglaralto, Guayas — no depth recorded;
LACM 34-285.5 — Isla Marchena, Islas
Galápagos — 37 т

Discussion

As pointed out by Olsson (1961: 375), this
species is very similar to the western Atlantic
Semelina nuculoides (Conrad, in Hodge, 1841),
and the two may be indistinguishable. Resolution
of this question would require more abundant
material of Semelina nuculoides from its type lo-
cality in the Pliocene of North Carolina than was
at my disposal. (Conrad's type material was not
located in the Academy of Natural Sciences of

Philadelphia by Moore, 1962: 80.) Eastern Pa-
cific material of S. subquadrata differs from Re-
cent specimens from North Carolina identified as
S. nuculoides (Fig. 5) in having higher, more
pointed, more posteriorly placed beaks.

Semelina campbellorum Coan, 2002,
new species
Figures 6-9

Description

Subtrigonal; anterior end more inflated; beaks
about two-thirds of way to posterior end,
orthogyrous; posterior end tapered, slightly
sinuous, subtruncate ventrally; surface with
lamellar commarginal ribs, becoming broader
ventrally, and with fine commarginal threads
between them; major lamellar ribs more higher
near and on posterior slope, whereas some of
them become thread-like and end anterior to
posterior slope; pallial sinus just touching ante-
rior adductor muscle scar, and confluent with
the pallial line for most of its length; internal liga-
ment in a short to elongate resilifer, not pro-
duced ventrally beyond the hinge plate;
external ligament in a narrow groove; right
valve with elongate anterior and posterior lat-
eral teeth, the left valve fitting into the grooves
between them and the dorsal margin; lunule
and escutcheon present; right valve with a nar-
row, inconspicuous anterior cardinal and a
large posterior cardinal; left valve with a large
anterior cardinal and a narrow posterior cardi-
nal that defines the posterior edge of the
resilifer. Length to 7.0 mm.

ANEW РАМАМ!С SEMELINA 178

Type Material & Locality

SBMNH 348118, holotype; length, 4.4 mm;
height, 3.3 mm; width, 2.0 mm (Fig. 6); SBMNH
348119, paratypes, 7 pairs, including the two fig-
ured herein (Figs. 7, 8); USNM 1008293; para-
type, 1 pair. Skoglund Coll., paratypes, 5 pairs.
Off Punta de Juluapan, Bahia de Santiago, Coli-
ma, México (19°5’N, 104°23’W); 30-60 m; Paul
& Carol Skoglund; December 1975 and later.

Distribution

In the СоЮ de California as far north as Bahia
de los Angeles, Baja California (29.1°М) [LACM
86-195.5], and Bahia San Ignacio, Sinaloa
(25.4°N) [LACM 37-218.2], Mexico, to Islas Lo-
bos de Afuera, Lambayeque, Perü (6.9°S)
[LACM 35-161.1]; Isla Socorro, Islas Revillagige-
dos, México [LACM 34-246.3, 34-247 .5]; Isla
Marchena, Islas Galäpagos, Ecuador (0.3°N)
[LACM 34-285.6]; 5-100 m (mean = 44 т; п =
29). The only bottom type noted on labels is
sand. | have seen 28 lots, and Carol Skoglund
provided data for an additional lot.

Referred Material

Stations in Mexico: LACM 86-195.5 — Bahia de
Los Angeles, Baja California — 5 m; Skoglund
Collection — Bahía Concepción, Baja Califor-
па Sur — 8-15 т; LACM 39-99.10 — Bahia
Coyote, Baja California Sur — 4-5 т; LACM
37-185.5 — Isla Ildefonso, Baja California Sur —
91 т; LACM 49-238.1 — Bahía San Francisco,
Baja California — 46 m; LACM 78-120.20 — Isla
Danzante, Baja California — 43-55 т; LACM
36-144.3 — Isla San Francisco, Baja California
— 42 т; LACM 49-237.1 — Bahía Frailes, Baja
California Sur — 91 т; LACM 34-247.5 — Isla
Socorro, Islas Revillagigedos — 7-18 m; LACM
34-246.3 — Isla Socorro, Islas Revillagigedos —
37 т; LACM 37-218.2 — Isla San Ignacio,
Sinaloa — 42 m; SBMNH 348118, 348119,
USNM 1008293, Skoglund Collection — Punta
de Juluapan, Bahía de Santiago, Colima — 30—
60 т - Type lot; LACM 38-265.4 — Bahia
Chacahua, Oaxaca - 75m

Stations in Costa Rica: LACM 72-13.30 — Bahía
Juanilla, Guanacaste — 37 т; LACM 72-7.31
— Bahía Santa Elena, Guanacaste — 1.2-11
m; LACM 72-30.26 — Punta Santa Elena,
Guanacaste — 12-15 т; LACM 72-57.37 —
Punta Quepos, Puntarenas — 21 т; LACM
72-66.38 — Isla del Caño, Puntarenas — 56 m.

Stations in Panamá: LACM 39-259.1 — Isla Lad-
rones, Chiriquí — 99 m; LACM 38-184.2 — Islas

Secas, Chiriquí — 22 т; LACM 34-251.5 — Islas
Secas, Chiriquí — 34-146 т; LACM 34-252,9 —
Bahía Honda, Veraguas — 55—64 т; LACM 34-
114.17 - Isla Jicarita, Veraguas — 44 m

Stations in Colombia: LACM 35-179.5 — Bahía
Octavia, Choco — 82 т; LACM 38-224.2 — Isla
Gorgona, Nariño — 18-37 m

Stations in Ecuador: LACM 33-180.1 — Bahía
Santa Elena, Guayas — 46 т; LACM 34-307.4
— Isla Santa Clara, Guayas — 64 m; LACM 34-
285.6 — Isla Marchena, Islas Galápagos — 37 т

Station in Perú: LACM 35-161.1 — Bahía Norte,
Islas Lobos de Afuera, Lambayque — 22 т

Discussion

This species differs from S. subquadrata
(Carpenter, 1857) in having a longer, more ta-
pered posterior end, orthogyrate rather than
opisthogyrate beaks, more lamellar commarginal
ribs with fine commarginal threads between the
larger ribs, and a more elongate pallial sinus. It
differs from small specimens of Semele, such
as $. jamesi Coan, 1988 (pp. 33-35, figs. 62,
63), which never attains a large size, in having
more prominent lateral teeth, especially the ante-
rior lateral, in the right valve and in having a less
conspicuous posterior cardinal in the left valve.

Etymology

This species is named for the Campbell clade
of Spartanburg, South Carolina, U.S.A., all of
whom have studied and published on the Mol-
lusca — Lyle D. Campbell, Sarah C. Campbell,
David C. Campbell, Matthew R. Campbell, and
Andrew C. Campbell.

ACKNOWLEDMENTS

Lyle D. Campbell kindly provided Pliocene and
Recent specimens of Semelina nuculoides for
examination. Kathie Way of The Natural History
Museum, London, loaned Carpenter's type ma-
terial of Semelina subquadrata; Lindsey Groves
of the Natural History Museum of Los Angeles
County, California, and Elizabeth Kools of the
California Academy of Sciences, San Francisco,
California, loaned material from those collec-
tions. Carol Skoglund made material and data
from her collection available to me, including the
specimens that became the type lot of S.
campbellorum. Gustav Paulay provided informa-
tion about systematic relations in the Tellinoidea.
Yolanda Camacho took the SEM photographs,
and Daniel L. Geiger prepared the plates.

174 СОАМ

LITERATURE CITED

BRANN, D. C., 1966, Illustrations to “Catalogue
of the collection of Mazatlan shells” by Philip
P. Carpenter. Ithaca, New York (Paleontologi-
cal Research Institution). 111 pp., 60 pls.

CAMPBELL, L. C., 1993, Pliocene molluscs from
the Yorktown and Chowan River formations
in Virginia. Virginia Division of Mineral
Resources Publication, 27: vii + 259 pp., incl.
43 pls.

CARPENTER, P. P., 1857a, Report on the
present state of our knowledge with regard to
the Mollusca of the west coast of North
America. Report of the British Association for
the Advancement of Science, 26[for 1856]:
159-368 + 4, pls. 6-9.

CARPENTER, P. P., 1857b, Catalogue of the
collection of Mazatlan shells, in the British
Museum: collected by Frederick Reigen. Lon-
don (British Museum). xii + 552 pp. [some as
i-iv + ix-xvi] [also published simultaneously as
Catalogue of the Reigen collection of
Mazatlan Mollusca, in the British Museum.
Warrington (Oberlin Press). viii + xii + 552 pp.]
[reprinted by Paleontological Research Insti-
tution, 1967].

COAN, E. V., 1988, Recent eastern Pacific spe-
cies of the bivalve genus Semele. The Ve-
liger 31(1/2): 1-42.

CONRAD, T. A., 1841. See Hodge 8 Conrad
(1841).

CONRAD, T. A., 1845, Fossils of the (medial
Tertiary or) Miocene formation of the United
States, no. 3. Philadelphia (Dobson). Pp. 57-
80, pls. 30-32.

DALL, W. H., 1899, Synopsis of the Recent and
Tertiary Leptonacea of North America and the
West Indies. Proceedings of the United States
National Museum, 21(1177): 873-897, pls. 87,
88.

DALL, W. H., 1900, Contributions to the Ter-
tiary fauna of Florida, with especial reference
to the silex beds of Tampa and the Pliocene
beds of the Caloosahatchie River, including
in many cases a complete revision of the ge-
neric groups treated of and their American
Tertiary species. Part V. Teleodesmacea:
Solen to Diplodonta. Transactions of the
Wagner Free Institute of Science of Philadel-
рта, 3(5): 949-1218, pls. 36-47.

DIAZ M., J. M. & М. PUYANA H., 1994, Moluscos
del Caribe Colombiano. Un catálogo ilustrado.
Santafe de Bogota (Colciencias & Fundación
Natura Colombia). 291 pp., [12] + 78 pls.

GARDNER, J., 1944, Mollusca from the Miocene
and lower Pliocene of Virginia and South Caro-
lina. Part |. Pelecypoda. United States Geo-
logical Survey Professional Paper, 199A: iv +
178 рр., 23 pls:

HERTLEIN, L. С. 8 А. М. STRONG, 1947, East-
ern Pacific expeditions of the New York Zoo-
logical Society. XXXVI. Mollusks from the west
coast of Mexico and Central America. Part V.

New York Zoological Society. Zoologica, 31(4):
129-150, pl. 1.

HODGE, J. T., with an appendix by T. A.
CONRAD, 1841, Observations on the Second-
ary and Tertiary formations of the southern
Atlantic states. American Journal of Science
and the Arts, 41(2): 332-348, pl. 2 [Conrad
appendix: pp. 344-348, pl. 2].

HOFFSTETTER, R., 1952, Moluscos subfösi-
les de los estanques de sal de Salinas (Pen.
De Santa Elena, Ecuador). Comparación con
la fauna actual del Ecuador. Boletín del
Instituto de Ciencias Naturales, 1(1): 3-79.

KAMENEV, G. M. 8 V. A. NADTOCHY, 1999, Spe-
cies of Macoma (Bivalvia: Tellinidae) from the
Pacific coast of Russia, previously described
as Abrina (Bivalvia: Semelidae). Malacologia
41(1): 209-230.

KEEN, A. М., 1958, Sea shells of tropical west
America; marine mollusks from Lower Califor-
nia to Colombia, 1st ed. Stanford, California
(Stanford University Press). xii + 624 pp., 10
pls. [repr.: 1960].

KEEN, A. M., 1968, West American mollusk types
at the British Museum (Natural History), IV.
Carpenter's Mazatlan collection. The Veliger,
10(4): 389-439, pls. 55-59.

KEEN, A. М., 1971, Sea shells of tropical west
America; marine mollusks from Baja Califor-
nia to Peru, 2nd ed. Stanford, California
(Stanford University). xiv + 1064 pp., 22 pls.
[repr., 1984 with only 12 pls.].

LAMY, E., 1915, Note sur le Semele nuculoides
Conrad. Bulletin du Muséum d'Histoire
Naturelle 21(1): 17-18.

MAXWELL, P.A., 1991, Clades vs. grades in bi-
valve classification — some examples from the
Tellinacea. American Malacological Union,
Program and Abstracts, 1991: 42.

MEYER, O., 1888, On Miocene invertebrates
from Virginia. Transactions of the American
Philosophical Society 25(127): 135-144, 1 pl.

MOORE, Е. J., 1962, Conrad's Cenozoic fossil
marine mollusk type specimens at the Acad-
emy of Natural Sciences of Philadelphia. Pro-
ceedings of the Academy of Natural Sciences
of Philadelphia, 114(2): 23-120, 2 pls.

OLSSON, A. A., 1961, Mollusks of the tropical
eastern Pacific particularly from the southern
half of the Panamic-Pacific faunal province
(Panama to Peru). Panamic-Pacific
Pelecypoda. Ithaca, New York (Paleontologi-
cal Research Institution). 574 pp., 86 pls.

REDFERN, C., 2001, Bahamian seashells: a
thousand species from Abaco, Bahamas. Boca
Raton, Florida (Bahamianseashells.com). x +
280 pp., 124 pls.

RIOS, E. de C., with the collaboration of, M.
HAIMOVICI, J. A. ALVARES PERES & К.
AGUIAR DOS SANTOS, 1994, Seashells of
Brazil, 2nd ed. Rio Grande (Universidade do
Rio Grande). 368 pp., 113 pls.

Revised ms. accepted 13 January 2002

MALACOLOGIA, 2003, 45(1): 175-178

AFIELD STUDY OF THE LIFE HISTORY
OF AN ENDEMIC HAWAIIAN SUCCINEID LAND SNAIL

Susan G. Brown, B. Kalani Spain & Karen Crowell

Social Sciences Division, University of Hawaii at Hilo,
200 W. Kawili Street, Hilo Hawaii 96720-4091, USA; susanb@hawaii.edu

The Hawaiian land snail fauna is noted for its
diversity and high level of endemism (Cowie,
1995). Almost nothing, however, is known of the
ecology of any of the species, with the excep-
tion of the life histories of a few achatinelline tree
snail species (Hadfield et al., 1993). The Hawai-
ian Succineidae comprise about 42 species
found in diverse habitats from arid coastal dune-
land to rainforest (Cowie et al., 1995), but the
only published work on their ecology is a labora-
tory study of growth and reproduction in Succinea
thaanumi Ancey, 1899, and Catinella rotundata
(Gould, 1846) (Rundell & Cowie, in press).

Here, we report a field study of the life history
of S. thaanumi. Unlike many Hawaiian land snail
species, S. thaanumi remains relatively com-
mon. It occurs on the eastern side of the island
of Hawaii, the largest in the Hawaiian chain. The
study was conducted in the Pu'u Maka’ala Natu-
ral Area Reserve in two areas (10 m? and 2 m?
about 12 m apart) at an elevation of 1,067 m.
The under-story consisted of native plants of
the following genera: Cyanea, Broussaisia,
Pipturus, Straussia, Freycinetia, and Cibotium.
The over-story consisted primarily of the Ha-
waiian endemic tree Metrosideros polymorpha.
Snails and their egg clutches were found on all
plants but mostly on Broussaisia, one of the
most common under-story plants in the study
site. Data were collected, usually twice weekly,
from 22 February 2000 to 4 June 2001, a total
of 104 occasions. In addition, rainfall was ге-
corded on 88 occasions with a rain gauge
placed in a clearing on the perimeter of the study
site, temperature on 68 occasions, and humid-
ity on 37 occasions.

Each plant in the study site was examined for
the presence of snails and egg clutches. Egg
clutches consisted of transparent, viscous
material containing developing embryos and
were laid primarily attached to leaf tips but also
occasionally on the stems and other parts of
the plants. This contrasts with other land snails,
which lay their eggs primarily in moist soil
(Tompa, 1984), and with Succinea putris

175

Linnaeus, 1758, which lays its eggs among the
roots of rushes (Rigby, 1965). Maximum shell
length of each snail was measured with cali-
pers, with a minimum amount of contact and
without moving the snail. The number of em-
bryos in each clutch was recorded. In addition,
the location of the snail on a plant (e.g., on the
top, bottom, or petiole of a leaf, or on the stem)
and whether the snail's body was retracted into
the shell or not were recorded.

Average snail size (F (11 92) = 19.6; р < 0.0001)
and the number of new egg clutches (F(44,44) =
2.8; p < 0.008) varied through the year. In Feb-
ruary and March of 2000 and 2001, the size-
frequency distribution was unimodal, with the
highest numbers in the 5—6 mm range (Fig. 1).
Few egg clutches were observed in February,
but by March, the number of clutches had in-
creased (Fig. 2). By April of both years, mean
snail size had increased but the size-frequency
distribution remained unimodal. In May, how-
ever, the distribution became bimodal, with the
appearance of large numbers of very small
(1 mm) snails, which we interpret as the off-
spring of the snails in the larger size-class (Fig.
1). Numbers of egg clutches also increased in
April (Fig. 2). The distributions remained bimo-
dal through August, but by this time fewer large
snails were observed, presumably because
they were dying off. The numbers of egg
clutches increased from April to August (Fig. 2).
From September through November, the size
distribution became skewed by the increased
numbers of newly emerged snails. These snails
appeared to be growing about 1 mm per month
because the modal size of this size-class in-
creased from 1 mm in September to 3 mm in
November. Growth continued through Decem-
ber and January (Fig. 1). Numbers of egg
clutches decreased from September through
January (Fig. 2). The number of embryos found
in a clutch ranged from 1 to over 16. In most
months, the average was 6-8, except for Feb-
ruary and March when it averaged 1-2. This
difference is significant (F 41.33) = 6.3; р < 0.001).

176 BROWN ЕТ AL.

N

8 9 10 11121314 123456

1
Ва. |. all.
| Ih. ho
alll k

7 8 91011121314

October 00 February 01

dl

ember 00

Marc

в. _ | alu ar

ember 00 Apni 01

ha. lla

May 01

January 9

alm. | №

6 7 8 9 1011121314

FIG. 1. Monthly size-frequency distributions of snail size. The Y-axis is the number of snails of a
particular size, averaged over data collecting occasions. The X-axis is snail size.

In 2000, copulations were first observed on
13 June and in 2001 on 30 May. Most copula-
tions were between pairs of snails (succineids
are hermaphrodites) with the top snail acting
as the male, mating by “shell mounting” (Asami
et al., 1998). In one case, we observed three
snails with the middle one acting as both a male
and a female. We observed one snail laying an
egg clutch. The viscous material containing the
eggs was exuded from the snail's genital pore.
The completed clutch was about three times
larger than the snail, which may be explained if
the viscous material absorbed moisture from
the air, as reported for Ovachlamys fulgens
(Gude, 1900) (Barrientos, 1998). We interpret
these patterns as representing an annual,
semelparous life-cycle, which agrees well with
the results of the laboratory study of S. thaanumi

of Rundell & Cowie (in press). Reproduction is
primarily in the May—November period, but
snails of all sizes and some egg clutches were
observed throughout the year. Therefore, at least
some Snails were growing and reproducing out
of synchrony with the overall population.

Snail behavior was related to the microclimate
of the study area. Numbers of snails recorded
were negatively related to temperature (r =
-0.44; Г 65) = 15.8; р < 0.0001), suggesting
that the snails moved into a different part о the
habitat as the temperature increased. However,
temperature did not vary significantly through
the year (F(11 55, = 0.86; р = 0.59), possibly be-
cause the Sud site was in the interior of an
upland rainforest. The number of new clutches
dropped to essentially zero when the humidity
was below 80% (Fig. 3). Regression analyses

LIFE HISTORY OF A SUCCINEID SNAIL

30-

20-

10-

0 < u |,

JFMAMJJASOND

FIG. 2. Frequency distribution of the number of
egg clutches observed during a month, averaged
over the two years of the study. The letters on
the X-axis are the first letters of the month. Bars
are standard errors of the means.

Number of Clutches

60 70

177

showed that the percent of snails observed on
the tops of leaves (F ¡3 зо) = 9.4; р < 0.0001) and
the bottoms of leaves (F3 зо) = 33.6; р < 0.0001),
and the percent of snails observed with their
bodies extended out of their shells (F(3 зо) = 32.5;
p < 0.0001) were significantly related to tem-
perature and humidity. In low humidity, the snails
tended to be on the bottoms of leaves; as hu-
midity increased, especially ifitwas raining, they
tended to be on the tops. When humidity
reached 80%, nearly all of the snails' bodies
were extended out of their shells. Snails on the
bottoms of leaves were usually retracted into
their shells, whereas snails on the tops of leaves
were usually extended. Presumably this behav-
ior is related to one of the major problems faced
by terrestrial snails, desiccation (Riddle, 1983).

ACKNOWLEDGMENTS

We thank Judy Spain and Johnnie Murphy for
help in collecting data, and Judy Spain for help in
reviewing the manuscript. We also thank the re-
viewers for their thoughtful and helpful comments.

100

80 90
Humidity

FIG. 3. Scatterplot of the relationship between number of new clutches re-
corded on an observation occasion and humidity.

178 BROWN ET AL.

LITERATURE CITED

ASAMI, Т., К. Н. COWIE & К. OHBAYASHI, 1998,
Evolution of mirror images by sexual asymmet-
ric mating behavior in hermaphroditic snails.
The American Naturalist, 152: 225-236.

BARRIENTOS, Z., 1998, Life history of the ter-
restrial snail Ovachlamys fulgens (Stylom-
matophora: Helicarionidae) under laboratory
conditions. Revista de Biologia Tropica, 46:
369-384.

COWIE, Б. Н., 1995, Variation in species diver-
sity and shell shape in Hawaiian land snails: in
situ speciation and ecological relationships.
Evolution, 49: 1191-1202.

COWIE, В. Н., М. Е. EVENHUIS &:C. С.
CHRISTENSEN, 1995, Catalog of the native
land and freshwater molluscs of the Hawaiian
Islands. Leiden, Backhuys Publishers, vi +
248 pp.

HADFIELD, MAG; S. Es MILLERSS AMEL
CARWILE, 1993, The decimation of endemic
Hawal'ian [sic] tree snails by alien predators.
American Zoologist, 33: 610-622.

RIDDLE, W. A., 1983, Physiological ecology of
land snails and slugs. Pp. 431-461, in: w. D.
RUSSELL-HUNTER, ed., The Mollusca, Volume
6. Ecology. New York, Academic Press.

RIGBY, J., 1965, Succinea putris: a terristrial
opisthobranch mollusc. Proceedings of the
Zoological Society of London, 144: 445-486.

RUNDELL, R. J. 8 К. Н. COWIE, in press, Growth
and reproduction in Hawaiian succineid land
snails. Journal of Molluscan Studies.

TOMPA, A. S., 1984, Land snails (Stylomma-
tophora). Pp. 47-140, in: A. S. TOMPA, ed., The
Mollusca Volume 7. Reproduction. New York:
Academic Press.

Revised ms. accepted 16 January 2003

MALACOLOGIA, 2003, 45(1): 179-184

DRY SEASON SURVIVAL IN A FLORIDA APPLE SNAIL
(POMACEA PALUDOSA SAY) POPULATION

Philip С. Darby, Patricia L. Valentine-Darby & H. Franklin Percival

Department of Biology, University of West Florida, 11000 University Parkway,
Pensacola, Florida, USA - 32514; pdarby@uwf.edu

ABSTRACT

Previous reports concluded that Florida apple snails (Pomacea paludosa Say) have
little tolerance to dry conditions, which stands in contrast to other Ampullariidae studied to
date. Given that inconsistency, and the fact that we do find snails in wetlands that periodi-
cally dry down, we were interested in elucidating dry season survival patterns in this spe-
cies. We conducted a field study in 1995 in which snails with miniature radio transmitters
were monitored weekly in flooded and dry marsh. Snails from this same wetland were
collected during the 1996 dry season and monitored during a laboratory study designed
to simulate a marsh in dry down conditions. We found that apple snails die at a rate of
approximately 10-15% per week in May and June regardless of water levels. Dry condi-
tions exacerbated snail mortality in the laboratory study (y* = 6.53, df = 1, Р = 0.011), but
not in the field (y? = 0.48, df = 1, P = 0.49). The mean size of snails still alive at the end of
the laboratory study were significantly smaller than those that had died during the study
(T = 2.25, 9 df, P = 0.025), indicating they were young of the year. Our data support
previously unsubstantiated reports that Pomacea paludosa is essentially an annual spe-
cies that experiences a post-reproductive die-off near the end of the dry season. lt ap-
pears previously reported apple snail deaths attributed to dry conditions were confounded
by an annual post reproductive die-off. Florida apple snails may be well equipped to sur-
vive in these fluctuating wetland environments, but how drying events affect their popula-

tion demography remains largely unanswered.
Key words: Pomacea paludosa, apple snail, survival, water level, wetland, Florida.

INTRODUCTION

Estimates of survival are critical to under-
standing population dynamics and the relation-
ship between population density, environmental
gradients, and the impacts of environmental
fluctuations. Freshwater snails of the genera
Pila and Pomacea (Caenogastropoda: Am-
pullariidae) commonly referred to as apple
snails, inhabit tropical and subtropical wet-
lands, some of which are subject to periodic
drying events (i.e., the water table falls below
ground level). Pila and Pomacea species stud-
ied to date can survive dry conditions for 3 to
25 months (Cowie, 2002), although direct com-
parisons of survival under wet versus dry con-
ditions have not been made for any apple snail
species. Apple snails have drawn attention be-
cause of their critical role as prey items for
wetland vertebrate fauna (e.g., Snyder 8

179

Snyder, 1969; Donnay & Beisinger, 1993) and
because they are considered pests in rice and
taro agriculture (Cowie, 2002). Information link-
ing survival with hydrology is fundamental to
understanding apple snail autecology and the
potential impacts of water management prac-
tices on snail populations.

The research described here was prompted
by several reports about the inability of Florida
apple snails, Pomacea paludosa Say, to toler-
ate drying events, in contrast to other Pila and
Pomacea species. Florida wetlands experi-
ence a dry season that generally extends from
November through May or June (Chen 8
Gerber, 1990), and in some years culminates
in a dry down (Kushlan, 1990; Duever et al.,
1994). Despite the persistence of snail popula-
tions in these wetlands, one field study
(Kushlan, 1975) and two lab studies (Little,
1968; Turner, 1994) concluded that Florida

180 DARBY ET AL.

apple snails were intolerant to dry downs, even
those less than one month in duration. How-
ever, there is also indirect evidence (accumula-
tions of empty shells in the field) of an annual
spring adult die-off that would typically coincide
with the May-June drying events (Hanning,
1979). If Florida apple snails were an annual
species, previous reports of dry down intoler-
ance may have been confounded by a coinci-
dental adult die-off. The objective of this study
was to compare survival of P. paludosa in wet
versus dry marsh in order to address the con-
tradictory available information on dry down tol-
erance in this species.

METHODS
Field Study

We conducted a field study in 1995 in the
easternmost portion of the Blue Cypress Water
Management Area (BCWMA), a wetland unit
that is part of the Upper St. Johns River Basin,
Indian River County, Florida. Darby et al. (2002)
described the study site, which had the highest
ground elevation of the basin wetlands and was
therefore most likely to dry out. Areas that even-
tually went dry were adjacent to areas that re-
mained inundated (Darby et al., 2002).

We studied the same snails from BCWMA
for which movements were monitored (Darby
et al., 2002). Snails were located weekly via
miniature radio transmitters affixed to their
shells. Maximum transmitter battery life was
60 d. We documented weekly survival for four
months by releasing transmitters in a stag-
gered fashion (six in March, four in April, 26 in
May, nine in June). We skewed transmitter re-
lease in May to maximize the probability of the
snails encountering a dry down. We also moni-
tored six snails that were initially found in May
via hand searches in dry marsh (the 45 snails
described above were found in flooded marsh).
Snails in dry marsh do not move (Darby et al.,
2002), so we could monitor them without trans-
mitters by flagging their location. Snail survival
was assessed first by tapping the shell to look
for the behavioral response of retracting the
operculum and, if there was no response, gen-
tly prying one corner of the operculum away
from the aperture to inspect the soft body tis-
sue. If an empty shell was found, evidence of
predation was noted as described by Snyder 8
Snyder (1969). We measured water depth and
temperature throughout the study, as described
by Darby et al. (2002).

Laboratory Study

In 1996, we assessed survival for stranded
snails in tanks designed to simulate a dry
down. A laboratory setting permitted control
over moisture conditions and eliminated factors
(e.g., precipitation, predation) that confounded
interpretation of field data.

Snails were collected from eastern BCWMA
via wire traps (Darby et al. 2001) from 29 April
through 10 May 1996 (n =232). Snail shell
widths were 22-42 mm (mean + SD = 34.1 +
3.2 mm). Based on size/maturation relation-
ships reported by Hanning (1979) and our own
observations of snails mating and laying eggs,
snails > 30 mm were considered adults. Ap-
proximately 10% of our lab study population
were juveniles.

On 12 May, a total of 18-24 snails were
placed in each of twelve 120 cm L x 61 cm W x
46 cm H polyethylene tanks. All tanks started
with 15 cm (above the substratum) of filtered,
aerated well water. The substratum consisted
of a 5 cm layer of stone in a size gradation (di-
ameter) from 1.9 cm to 0.6 cm (upper layer)
and topped by a 13 cm layer of either sand (n =
6 tanks) or unprocessed commercial peat (n =
6). Two substrata were included in order to see
if peat, with its greater moisture holding capac-
ity, would enhance survival for stranded snails
relative to sand. Tank locations were random-
ized for substratum type and water regime.

Water was replaced every 3 to 7 days during
the experiment. Snails were fed Utricularia sp.
in excess of demand. Uneaten food and waste
were removed every 3 days. The tanks were
outdoors and therefore subject to ambient tem-
peratures, but covered to prevent rain from en-
tering. Well water was allowed to reach
ambient temperature prior to tank distribution.

For each substratum (peat and sand), we
had three control tanks with continuous 15 cm
water depths and three dry down tanks
wherein water was dropped from 15 cm to 0
cm over 28 days. The water withdrawal rate
approximated the 28 d period prior to dry down
conditions in BCWMA in 1995. Water depths
reached 0 cm on 10 June. On 11 June all the
water was drained and the substratum began
drying.

We measured water and substratum tem-
peratures three times weekly. Substratum
moisture levels were measured three times
weekly in three locations per tank as percent
saturation using a moisture meter (Lincoln In-
dustries, NE) inserted 5 cm below the surface.
Snail survival was checked weekly as de-

DRY SEASON SURVIVAL IN À FLORIDA APPLE SNAIL POPULATION 181

scribed for the telemetry study. All tanks con-
taining water were inspected daily to remove
dead snails.

Statistical Analyses

Cumulative survival for the laboratory snails
was estimated at weekly intervals using the
Kaplan 8 Meier procedure (1958). A variation of
this procedure to accommodate staggered
transmitter release and failed transmitters was
used to estimate survival in the telemetry study
(Pollock et al. 1989). We used the approach
described by Bennetts et al. (1999) to accom-
modate situations in which animals previously
found in one condition (e.g., flooded marsh)
were subsequently found in another (e.g., dry
marsh). The hypothesis that survival for snails
in dry down conditions (19 May-7 July) was
lower than in flooded conditions over the same
period was tested using a log-rank test (Pollock
et al., 1989). We tested for sex differences and
tank effects using the same approach. In all
cases we report the most conservative у? val-
ues of the three log-rank tests described in
Pollock et al. (1989), but for all tests the con-
clusions were the same regardless of how y?
was Calculated.

RESULTS

Snail survival in both the field and the labora-
tory studies declined to less than 10% (Fig. 1),
regardless of water levels. Survival in the field
study remained at 100% for the initial six-week
period through mid-April before declining at an
average rate of 14% per week through June.
Water levels declined steadily (Darby et al.
2002) and water temperatures rose from 23°C
in March to a peak of 38°C (afternoon mea-
surements) in late May. Nine of these snails
were eaten by predators (snail kites and
limpkins). [There may have been more preyed
upon, but for several dead snails the evidence
for predation was not clear]. Survival for males
(n = 24) did not differ from females (n = 27)
(y? = 0.048, df = 1, = 0.83). Survival of the
twelve snails stranded in dry down was not dif-
ferent from snails remaining in flooded marsh
over the same period (y? = 0.48, df = 1, P =
0.49). The mean survival time in dry down con-
ditions was 3.9 + 2.2 weeks.

Substratum type had no effect on survival
(x? = 0.121, df = 1, = 0.73 and y? = 0.08,
df = 1, P = 0.78, for controls and dry down

tanks, respectively), despite the fact that mois-
ture levels in the peat substratum remained 3-
5 times higher than the sand substratum (data
not shown). The peat and sand tank survival
data were therefore pooled to examine the
overall effects of drying. Survival for all labora-
tory snails declined approximately 10-15% per
week by late May, prior to the 11 June dry
down. The simulated dry down did, however,
appear to exacerbate death rates (y? = 6.53,
df = 1, P = 0.011). Tank water temperatures
ranged from 23°C to 30°C over the study pe-
riod.

At the end of the lab study, it appeared that
surviving snails were smaller than those that
had died. We used a T-test for groups with un-
equal variance (F-test: F = 0.27, P = 0.0002) to
test for size differences between surviving and
dead snails. The mean (+ SD) size of the ten
snails living at the end of the lab study was sig-
nificantly smaller (30.1 + 3.1 mm) than that of
those that had died (34.3 + 5.9 mm) (T = 2.25,
df = 9, P = 0.025). Sizes were similar for the
47 dead (35.9 + 2.4 mm) and four living snails
(34.9 + 2.5 mm) at the end of the field study
(T = 0.70, df = 3, = 0.27). At the end of both
studies, aestivating snails were removed from
their dry conditions and placed in water. They
became active within 2 to 24 h.

DISCUSSION

Florida apple snails, regardless of hydrologic
conditions, exhibit a steady decline in survival
late in the dry season (May-June). Dry down
conditions exacerbated mortality in these
snails, but dry down intolerance clearly was
not the primary cause of mortality for snails
observed in the lab or the field. Our data from
both the field and lab studies support previously
unsubstantiated estimates of a 1-1.5 year life
span for P. paludosa (Hanning, 1979; Ferrer et
al., 1990). It appears that the life cycle of apple
snails terminates in a post-reproductive die-off.
Egg cluster surveys by Hanning (1979), Odum
(1957) and Darby et al. (1999) consistently
show an April-May peak in egg production. In
our study, steepest declines in survival oc-
curred in May and June. The telemetry study
was initiated early in the breeding season, so
this would explain the six-week period of high
survival prior to the May—June die-off. Snails
alive at the end of the lab study were likely
young of the year that had reached sufficient
size to be captured in the wire traps.

182 DARBY ET AL.

a)
1.00
0.80
©
2 060
>
>
=
[7]
0.40
—— Snails in flooded marsh
020 —=— Snails in dry marsh
0.00 -
3/3 3/17 3/31 4/14 4/28 BAZ 5/26 6/9 6/23 117 7/21
b)
1.00
0.80
<
> 0.60
=
D
-e- Sand Control
eet —= Sand Dry Down
== Peat Control
0.20 += Peat Dry Down
0.00 TE т

-

3/3 3/17 3/31 4/14 4/28 5/12 5/26 6/9 6/23 TT T2

FIG. 1. Cumulative survival for apple snails monitored in wet and dry conditions in a) the 1995 field

study and b) the 1996 laboratory study. Dashed vertical lines indicate the date on which snails first
became stranded in dry conditions.

DRY SEASON SURVIVAL IN A FLORIDA APPLE SNAIL POPULATION 183

Bearing transmitters could influence survival.
However, the staggered entry design meant
that snails in any given week had carried trans-
mitters for a range of different times. For ex-
ample, in the first week of June, eight snails
that died had worn transmitters from 1 to 8
weeks and 11 snails still alive had worn trans-
mitters 1 to 9 weeks. Four of the snails were
initially found via hand searches in dry marsh,
demonstrating that snails do get stranded with-
out bearing transmitters.

We also considered that high water tempera-
tures (33-38°С) we measured in the field in
May 1995 might have contributed to the steep
decline in survival. In the lab study, however,
we found that a similar survival pattern
emerged even when water temperatures
stayed < 30°C. Lethal temperatures of 35°C to
45°C have been reported for other apple snail
species (Cowie, 2002).

lt appears previously reported apple snail
deaths attributed to dry conditions (Little, 1968;
Turner, 1994) were confounded by an annual
post reproductive die-off. Apple snails may be
well equipped to survive in these fluctuating
wetland environments, but how drying events
affect Florida apple snail population demogra-
phy remains largely unanswered.

ACKNOWLEDGEMENTS

This work was funded under a joint contract
by the South Florida Water Management Dis-
trict (WMD) (Contract No. C-E6609) and the St.
Johns River WMD (Contract No. 950159) and
conducted under the auspices of the Florida
Cooperative Fish and Wildlife Research Unit —
Florida Fish and Wildlife Conservation Com-
mission (FFWCC), University of Florida, Ц. 5.
Geological Survey, and Wildlife Management
Institute, cooperating — through cooperative
agreement #1434HQ97RU01544. This is con-
tribution No. R-08977 of the Florida Agricultural
Experiment Station Journal Series.

We thank Patrick Dean (FF WCC) and family
and Steve Darby for help during the lab studies.
We also thank Rob Bennetts (USGS-BRD)
and two anonymous reviewers for suggestions
on draft manuscripts.

LITERATURE CITED

BENNETTS, К. Е., V. J. DREITZ, W. М. KITCH-
ENS, J. E. HINES & J. D. NICHOLS, 1999, An-

nual survival of snail kites in Florida: radio te-
lemetry versus capture-resighting data. The
Auk, 116: 435-447.

CALOW, P., 1978, The evolution of life-cycle
strategies in fresh-water gastropods. Mala-
cologia, 17: 351-364.

CHEN, E. & J. Е. GERBER, 1990, Climate. Pp.
11-34, in: В. L. MEYERS 4 J. J. EWEL, eds., Eco-
systems of Florida. University of Central
Florida Press, Orlando, Florida.

COLES, С. C., 1968, The termination of aestiva-
tion in the large fresh-water snail Pila ovata
(Ampullariidae)-I. changes in oxygen uptake.
Comparative Biochemistry and Physiology, 25:
517-522.

COWIE, R. H., 2002, Apple snails (Ampullariidae)
as agricultural pests: their biology, impacts
and management. Pp. 145-192, in: G M.
BARKER, ed., Molluscs as crop pests. CABI
Publishing, Wallingford, New Zealand.

DARBY, P. C., В. E. BENNETTS, J. D. CROOP,
P. L. VALENTINE-DARBY & W. М. KITCHENS,
1999, A comparison of sampling techniques
for quantifying abundance of the Florida apple
snail (Pomacea paludosa Say). Journal of
Molluscan Studies, 65: 195-208.

DARBY, P. C., P. L. VALENTINE-DARBY, H. F.
PERCIVAL & W. M. KITCHENS, 2001, Collect-
ing Florida applesnails (Pomacea paludosa)
from wetland habitats using funnel traps. Wet-
lands, 21: 308-311.

DARBY, P. С., К. Е. BENNETTS, $. J. MILLER &
Н. F. PERCIVAL, 2002, Movements of Florida
apple snails in relation to water levels and dry-
ing events. Wetlands, 22: 489-498.

DONNAY, Т. J. & $. К. BEISINGER, 1993, Apple
snail (Pomacea doliodes) and freshwater crab
(Dilocarcinus dentatus) population fluctua-
tions in the llanos of Venezuela. Biotropica, 25:
206-214.

DUEVER, М. J, J. Е. MEEDER, L. С. MEEDER 8
J. М. McCOLLOM, 1994, The climate of south
Florida and its role in shaping the Everglades
ecosystem. Pp. 225-248, in: $. М. DAVIS & J. С.
OGDEN, eds., Everglades: the ecosystem and
its restoration, St. Lucie Press, Delray Beach,
Florida.

FERRER, J. R., С. PERERA & М. YONG, 1990,
Life tables of Pomacea paludosa (Say) in
natural conditions. Florida Scientist, Supple-
ment 53: 15 pp.

HANNING, G. W., 1979, Aspects of reproduction
п Pomacea paludosa (Mesogastropoda:
Pilidae). Master’s Thesis, Florida State Univer-
sity, Tallahassee, Florida, USA.

KAPLAN, Е. L. 8 P. MEIER, 1958, Nonparametric
estimation of incomplete observations. Journal
of the American Statistical Association, 53:
457-481.

KUSHLAN, J. A., 1975, Population changes of
the apple snail (Pomacea paludosa) in the
southern Everglades. Nautilus, 89(1): 21-23.

KUSHLAN, J. A., 1990, Freshwater marshes. Pp.
324-363, in: R. L. MEYERS, 8 J. J. EWEL, eds.,
Ecosystems of Florida, University of Central
Florida Press, Orlando, Florida.

184 DARBY ET AL.

LITTLE, C., 1968, Aestivation and ionic regula-
tion in two species of Pomacea (Gastropoda,
Prosobranchia). Journal of Experimental Biol-
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ductivity of Silver Springs, Florida. Ecological
Monographs, 27: 55-112.

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BUNCK & P. D. CURTIS, 1989, Survival analy-
sis in telemetry studies: the staggered entry
design. Journal of Wildlife Management, 53:
7-15.

SCIENCE SUBGROUP, 1996, South Florida
ecosystem restoration: scientific information
needs. Report to the Working Group of the
South Florida Ecosystem Restoration Task
Force. http://everglades.fiu.edu/taskforce/
scineeds/index.html.

SNYDER, М. Е. 8 H. A. SNYDER, 1969, A com-
parative study of mollusk predation by
limpkins, everglade kites, and boat-tailed
grackles. Living Bird, 8: 177-223.

TURNER, R. L., 1994, The effects of hydrology
on the population dynamics of the Florida
apple snail (Pomacea paludosa). Florida Insti-
tute of Technology. Final Report for the St.
Johns Water Management District.

USFWS, 1999, South Florida multi-species re-
covery plan. US Fish and Wildlife Service, De-
partment of the Interior, Atlanta, Georgia, USA;
http://verobeach.fws.gov/Programs/Recovery/
vbms5.html.

Revised ms. accepted 7 March 2003

MALACOLOGIA, 2003, 45(1): 185-187

LUNTIA INSIGNIS SMITH, 1898, IS А SYNONYM OF
STREPTOSTELE (TOMOSTELE) MUSAECOLA (MORELET, 1860)
(GASTROPODA: STREPTAXIDAE) - AN AFRICAN TRAMP
AND ITS DISTRIBUTION IN AMERICA

Bernhard Hausdorf! & Clara Ines Medina Bermudez?

Smith (1898) described Luntia insignis as a
new genus and new species of the family
Stenogyridae (= Subulinidae) from Trinidad.
This species has been recorded as probably
introduced in Aruba (Hummelink, 1940a, b),
Guyana (Morrison, 1943), Suriname (van
Regteren Altena, 1960, 1964, 1975), Nicaragua
(Löpez & Pérez, 1996), and in a greenhouse in
the Netherlands (Meeuse & Hubert, 1949, as
Varicella clappi [non Pilsbry, 1907]; see van
Regteren Altena, 1964). It has also been re-
corded from Saba (Haas, 1962) and Barbados
(Chase & Robinson, 2001).

Thiele (1931) included Luntia Smith, 1898, as
a subgenus in Leptinaria Beck, 1837 (Subulini-
dae), whereas Baker (in van Regteren Altena,
1975), who examined the radula of the species,
transferred it to Varicella L. Pfeiffer, 1856
(Oleacinidae).

Pilsbry (1930) was the first to record the Afri-
can Streptostele (Tomostele) musaecola
(Morelet, 1860) (Streptaxidae) as an introduced
species in Panama. The native range of
Streptostele musaecola extends in western Af-
rica from Guinea to the Congo (Pilsbry, 1919).
Pilsbry (1930) recognized that this might be a
“tramp” species carried around on bananas, on
which it was originally discovered (Morelet,
1860). Later, Streptostele musaecola has been
recorded as introduced in Bermuda (Bieler &
Slapcinsky, 2000), Vanuatu (= New Hebrides)
(Solem, 1989), American Samoa (Solem,
1989; Cowie, 1998, Cowie & Rundell, 2002;
Cowie et al., 2003), and the Society Islands
(Solem, 1989).

We have examined the holotype of Luntia
insignis Smith (The Natural History Museum,
London, NHM 1898.12.5.18; Fig. 1) and three
specimens labelled as syntypes of Achatina
musaecola Morelet in The Natural History Mu-
seum (МНМ 1893.2.4.276-8; Fig. 2). As locality
of the putative syntypes of A. musaecola,
“Gabon” is given in Morelet’s handwriting,

whereas Morelet (1860: 190) gives Guinea as
type locality. However, Ancey (1885) also said
that A. musaecola is from Gabon. It is unclear
whether this was an error or whether Ancey
knew that the type locality actually was Gabon
and not Guinea.

A comparison of these type specimens, new
material from Colombia (Zoologisches Mu-
seum der Universität Hamburg, ZMH 2918; Fig.
3), and figures of shells identified as Luntia
insignis (Meeuse & Hubert, 1949; Haas, 1962)
and Streptostele musaecola (Pilsbry, 1919,
1930; Solem, 1989; Bieler & Slapcinsky, 2000),

FIGS. 1-3. Streptostele (Tomostele) musaecola
(Morelet). FIG. 1. Trinidad: Port of Spain (holo-
type of Luntia insignis Smith, NHM 1898.12.5.18).
FIG. 2. Gabon (syntype? of Achatina musaecola
Morelet, NHM 1893.2.4.276-8). FIG. 3. Colombia:
Finca Torreblanca near Silvania (ZMH 2918).
Scale bar = 1 mm.

'Zoologisches Museum der Universitat Hamburg, Martin-Luther-King-Platz 3, D-20146 Hamburg, Germany;

hausdorf@zoologie.uni-hamburg.de

2Universidad Militar Nueva Granada, Facultad de Ciencias: Biología Aplicada, Cra. 11 No 101 - 80, Bogotá, Colombia

186 HAUSDORF 8 MEDINA BERMÚDEZ

FIG. 4. Distribution of Streptostele (Tomostele) musaecola (Morelet) in America
(1°-grid).

show that Luntia insignis Smith, 1898 is a syn-
onym of Streptostele (Tomostele) musaecola
(Morelet, 1860). Consequently, Luntia Smith,
1898, is asynonym of Tomostele Ancey, 1885,
of which Achatina musaecola Morelet, 1860, is
the type species.

Streptostele musaecola is known from the
following American localities (Fig. 4): Bermuda:
Hamilton, 32°18’М, 64°47’W (Bieler &
Slapcinsky, 2000). Nicaragua: Masatepe, Е!
Arenal, El Pochote, 455 т altitude, 11°55’N,
86”08'46"W (López & Pérez, 1996); Masatepe,
El Arenal, El Mango, 455 т altitude, 11°55’N,
86°08'46"W (López 8 Pérez, 1996). Costa
Rica: La Lola, 28.3 miles W of Puerto Limón,
10°N, 83°W (Florida Museum of Natural His-
tory FLMNH 211842). Panama: Mount Hope,
9°20’М, 79°54’W (Pilsbry, 1930); Colon,
922'N, 79°54’W (FLMNH 211843). Colombia,
Departamento Cundinamarca: Finca Torre-
blanca near Silvania, 1560 m altitude, forest,
4°24'14"N, 74°23°12"W (ZMH 2918). Aruba:
Fontein, 12°30'N, 69°54’W (Hummelink, 1940a).
Saba: Road to Bottom, 17°38’N, 63°15’W
(Haas, 1962). Saint Lucia: Grande Anse,
14°00’М, 60%54'W (FLMNH 281110). Barbados:
Holetown, Porter's House, 2-4 m altitude,
13°11'44”М, 59°38'18"W (Chase & Robinson,
2001); Holetown, hill NE of Royal West-
moreland Landscape Garden Centre, 10 m

altitude, 13°12’01”М, 59*38'04”"W (Chase &
Robinson, 2001); Bathsheba, Andromeda
Botanic Gardens, 60-90 т altitude,
13°12’25”М, 59°31’04”W (Chase 8 Robinson,
2001); 200 m S of Harrison Point Lighthouse,
30 т altitude, 13°18’23”М, 59°38'58"W (Chase
8 Robinson, 2001). Trinidad: Port of Spain,
10°39’М, 61°31’W (holotype of Luntia insignis
Smith, NHM 1898.12.5.18). Guyana: Kyk over
Al Island, 6°23’N, 58°41’W (Morrison, 1943).
Suriname: Jodensavanne, 5°25’N, 54*59'W
(van Regteren Altena, 1964, 1975); Albina,
5°30’N, 54°03’W (van Regteren Altena, 1975);
Paramaribo, 5°50’N, 55°10’W (van Regteren
Altena, 1964, 1975); Tambaredjo, 5°50’N,
55°33’W (van Regteren Altena, 1960).

The newly discovered occurrence near
Silvania in Colombia represents probably the
highest locality from which Streptostele
musaecola has been recorded. However, we
do not know whether a stable population has
been established there or whether there was
only an unsuccessful introduction.

Streptostele musaecola is widespread
throughout tropical America. Its impact on the
native fauna should be monitored, especially
because it is a carnivorous species. Actually, it
might have been implicated in the extinction of
a native species in American Samoa (Miller,
unpublished report, quoted in Cowie, 1998).

STREPTOSTELE MUSAECOLA IN AMERICA 187

ACKNOWLEDGEMENTS

We thank Dr. Peter Mordan (NHM) for lending
as the relevant type material, Dr. F. Thompson
(FLMNH) for communicating additional records
and Dr. R. H. Cowie and two anonymous re-
viewers for helpful comments on the manu-
script.

LITERATURE CITED

ANCEY, С. F., 1885, Nouvelles contributions
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BIELER, R. 4 J. SLAPCINSKY, 2000, A case
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Nemouria, 44: 1-99.

CHASE, К. & 0. С. ROBINSON, 2001, The un-
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, 1998, Catalog of nonmarine
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HAAS, F., 1962, Caribbean land molluscs:
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HUMMELINK, P. W., 1940a, A survey ofthe mam-
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HUMMELINK, P. W., 1940b, Zoogeographical re-
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LOPEZ, A. & A. М. PEREZ, 1996, Nuevos
registros de caracoles terrestres “ad-
venedizos” en Nicaragua: Leptinaria insignis,
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MEEUSE, А. О. J. 8 В. HUBERT, 1949, The mol-
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MORELET, A., 1860, Description de nouvelles
especes de l'Afrique occidentale, rapportées
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MORRISON, J. P. E., 1943, A new type of fresh-
water clam from British Guiana. Nautilus, 57:
46-52, pl. 8.

PILSBRY, H. A., 1907, Manual of Conchology.
Second Series: Pulmonata. 19 (74): 65-128,
pls. 11-20. Academy of Natural Sciences,
Philadelphia.

PILSBRY, H.A., 1919, A review of the land mol-
lusks of Belgian Congo chiefly based on the
collections of the American Museum Congo
Expedition, 1909-1915. Bulletin of the Ameri-
can Museum of Natural History, 40: 1-370,

pls. 1-23.
PILSBRY, H. A., 1930, Results of the Pinchot
South Sea Expedition — Il. Land mollusks of

the Canal Zone, The Republic of Panama, and
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339-354, pls. 28-30.

SMITH, E. A., 1898, On some land shells from
Trinidad. Journal of Conchology, 9: 27-29.
SOLEM, A., 1989, Non-camaenid land snails of
the Kimberley and Northern Territory, Australia.
|. Systematics, affinities and ranges. Inverte-

brate Taxonomy, 2: 455-604.

THIELE, J., 1929-1931, Handbuch der syste-
matischen Weichtierkunde. G. Fischer, Jena,
1: 1-376 [1929]; 377-778 [1931].

VAN REGTEREN ALTENA, С. O., 1960, On a
small collection of land Mollusca from Surinam
(Dutch Guyana). Basteria, 24: 48-51.

VAN REGTEREN ALTENA, С. O., 1964, Notes
on some Surinam land snails. Zoologische
Mededelingen, 40: 139-141.

VAN REGTEREN ALTENA, С. O., 1975, Land
Gastropoda of Suriname, with description of a
new species of Nesopupa. Basteria, 39: 29-
50.

Revised ms. accepted 24 March 2003

MALACOLOGIA

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Vol. 45(1) 2003

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VOL. 45, МО. 1 MALACOLOGIA 2003

CONTENTS

SUSAN G. BROWN, B. KALANI SPAIN, 8 KAREN CROWELL
A Field Study of the Life History of an Endemic Hawaiian Succineid Land
A NR NN 175
LAURE CHEVALIER, MARTINE LE COZ-BOUHNIK, & MARYVONNE CHARRIER
Influence of Inorganic Compounds on Food Selection by the Brown Garden

Snail Cornu aspersum (Muller) (Gastropoda: Pulmonata) .............. 125
EUGENE V. COAN
A New Panamic Species of the Bivalve Genus Semelina (Semelidae) .... 169

ROBERT H. COWIE & SILVANA C. THIENGO
The Apple Snails of the Americas (Mollusca: Gastropoda: Ampullariidae:
Asolene, Felipponea, Marisa, Pomacea, Pomella): A Nomenclatural and
MVOCKGARGIOG. cp teu tahoe ne be ue eter a ams Mer sale 41
PHILIP С. DARBY, PATRICIA L. VALENTINE-DARBY, & H. FRANKLIN PERCIVAL
Dry Season Survival in a Florida Apple Snail (Pomacea paludosa Say)
SO CAO ue ae 179
ALEXANDR V. GARBER & ALEXEI V. KORNIUSHIN
Karyotypes of European Species of Radix (Gastropoda: Pulmonata:
Lymnaeidae) and Their Relevance to Species Distinction in the Genus ... 141

MATTHIAS GLAUBRECHT, THOMAS VON RINTELEN, 4
ALEXEI V. KORNIUSHIN
Toward a Systematic Revision of Brooding Freshwater Corbiculidae т
Southeast Asia (Bivalvia, Veneroida): On Shell Morphology, Anatomy and
Molecular Phylogenetics of Endemic Таха from Islands in Indonesia ..... 1
BERNHARD HAUSDORF & CLARA INÉS MEDINA BERMÚDEZ
Luntia insignis Smith, 1898, Is а Synonym of Streptostele (Tomostele)
musaecola (Morelet, 1860) (Gastropoda: Streptaxidae)—An African Tramp

and its: DISIMDUTIONAMAMONICA occasion 185
ALOIS HONEK

Shell-Band Color Polymorphism in Cepaea vindobonensis at the Northern

Mo A reese ays A ores eon eet Scone Gs 133

WANNAPORN ITTIPRASERT, CHRISTOPHER ROWE, CAROLYN PATTERSON,

ANDRE MILLER, NITHYA RAGHAVAN, SUSAN BANDONI, FRED LEWIS, &

MATTY KNIGHT
Assessment of Genetic Heterogeneity within Laboratory-Maintained
Schistosoma mansoni-Resistant Stocks of Biomphalaria glabrata Snails by
O A ee 101

PARIDE MANTECCA, GIOVANNI VAILATI, LETIZIA GARIBALDI, &
RENATO BACCHETTA
Depth Effects on Zebra Mussel Reproduction ....................... 109
ARMINDO S. RODRIGUES, REGINA T. CUNHA, 8 BENJAMIN J. GÓMEZ
The Egg of Oxychilus (Drouetia) atlanticus (Pulmonata: Zonitidae): Surface
Structure and Carbohydrate Composition .......................... 121
CHRISTOPHER ROWE, WANNAPORN ITTIPRASERT, CAROLYN PATTERSON,
CLAUDIA ELIFF, KRISTEN PAGE, SUSAN BANDONI, THOMAS WILKE,
DENNIS MINCHELLA, FRED LEWIS, & MATTY KNIGHT
Use of Microsatellite Variation and RAPD-PCR to Assess Genetic Poly-
morphism in Biomphalaria glabrata Snails from a Single Locale in a
Schistosomiasis Endemic. Alea ен de meee $d Sew ete ees 149

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CONTENTS

MATTHIAS GLAUBRECHT, THOMAS VON RINTELEN, 4

ALEXEI V. KORNIUSHIN
Toward a Systematic Revision of Brooding Freshwater Corbiculidae in
Southeast Asia (Bivalvia, Veneroida): On Shell Morphology, Anatomy and
Molecular Phylogenetics of Endemic Taxa from Islands in Indonesia .....

ROBERT H. COWIE 8 SILVANA С. THIENGO
The Apple Snails of the Americas (Mollusca: Gastropoda: Ampullariidae:
Asolene, Felipponea, Marisa, Pomacea, Pomella): A Nomenclatural and
Туве Catalog. ео

WANNAPORN ITTIPRASERT, CHRISTOPHER ROWE, CAROLYN PATTERSON,
ANDRE MILLER, NITHYA RAGHAVAN, SUSAN BANDONI, FRED LEWIS, &
MATTY KNIGHT

Assessment of Genetic Heterogeneity within Laboratory-Maintained

Schistosoma mansoni-Resistant Stocks of Biomphalaria glabrata Snails by

RAPDEPER u.a Be a Ho PER AN
PARIDE MANTECCA, GIOVANNI VAILATI, LETIZIA GARIBALDI, &
RENATO BACCHETTA

Depth Effects on Zebra Mussel Reproduction .......................
ARMINDO $. RODRIGUES, REGINA T. CUNHA, & BENJAMIN J. GOMEZ

The Egg of Oxychilus (Drouetia) atlanticus (Pulmonata: Zonitidae): Surface

Structure and Carbohydrate Composition ..... еее eee
LAURE CHEVALIER, MARTINE LE COZ-BOUHNIK, & MARYVONNE CHARRIER

Influence of Inorganic Compounds on Food Selection by the Brown Garden

Snail Cornu aspersum (Muller) (Gastropoda: Pulmonata) ..............
ALOIS HONEK

Shell-Band Color Polymorphism in Cepaea vindobonensis at the Northern

Limit OF iS Range . 20.000220 das nat Os cle pes RE
ALEXANDR V. GARBER & ALEXEI V. KORNIUSHIN

Karyotypes of European Species of Radix (Gastropoda: Pulmonata:

Lymnaeidae) and Their Relevance to Species Distinction in the Genus ...
CHRISTOPHER ROWE, WANNAPORN ITTIPRASERT, CAROLYN PATTERSON,
CLAUDIA ELIFF, KRISTEN PAGE, SUSAN BANDONI, THOMAS WILKE,
DENNIS MINCHELLA, FRED LEWIS, & MATTY KNIGHT

Use of Microsatellite Variation and RAPD-PCR to Assess Genetic Poly-

morphism in Biomphalaria glabrata Snails from a Single Locale in a

schistosomiasis Endemic Area „осо eee

RESEARCH NOTES

EUGENE V. COAN
A New Panamic Species of the Bivalve Genus Semelina (Semelidae) ....
SUSAN G. BROWN, B. KALANI SPAIN, & KAREN CROWELL
A Field Study of the Life History of an Endemic Hawaiian Succineid Land
SMA... 2.5 ca AA Me ee eee a .
PHILIP C. DARBY, PATRICIA L. VALENTINE-DARBY, & H. FRANKLIN PERCIVAL
Dry Season Survival in a Florida Apple Snail (Pomacea paludosa Say)
РОВНОМ ооо ое A о RE
BERNHARD HAUSDORF & CLARA INÉS MEDINA BERMÚDEZ
Luntia insignis Smith, 1898, Is a Synonym of Streptostele (Tomostele)
musaecola (Morelet, 1860) (Gastropoda: Streptaxidae)—An African Tramp
and is Distribution in Amenea -:..... onan sas ce eae o о

2003

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MALACOLOGIA, 2004, 45(2): 195-440

CLASSIFICATION OF THE HELICINIDAE: REVIEW OF MORPHOLOGICAL
CHARACTERISTICS BASED ON A REVISION OF THE COSTA RICAN SPECIES
AND APPLICATION TO THE ARRANGEMENT OF THE CENTRAL AMERICAN
MAINLAND TAXA (MOLLUSCA: GASTROPODA: NERITOPSINA)

га Richling

Zoologisches Institut, Christian-Albrechts-Universität zu Kiel
Olshausenstrake 40, 24098 Kiel, Germany; ira@richling.de

ABSTRACT

The present study combines a taxonomical revision of the poorly known Costa Rican
Helicinidae, with a detailed investigation of certain morphological structures with respect
to their relevance for systematics, culminating in a discussion of the arrangement of the
Central American mainland species.

The revision of the Costa Rican species is based on the examination of nearly all type
material, coupled with extensive field work and investigations of the collections of the
Instituto Nacional de Biodiversidad de Costa Rica and the Florida Museum of Natural His-
tory, Gainesville, along with perusal of additional historical material. With minor exceptions,
all these species were investigated with respect to the features of shell, operculum, surface
sculpture of embryonic shell and teleoconch, internal shell structures, radula, and female
reproductive system. In addition, analyses of morphometry and sexual dimorphism were
carried out. Faced with a limited amount of material, it became necessary to develop a new
preparation method to separate the soft body from the shell without damaging either.

For the higher classification and comparative analysis of the different morphological char-
acteristics, similar examinations emphasizing formerly poorly studied or neglected charac-
teristics, such as embryonic shell and female reproductive system, were carried out for 17
additional species representing the most important related Central American supraspecific
taxa using their type species when available. For taxa with inaccessible material, data from
the available literature were critically incorporated.

For Costa Rica, 15 species were recognized, among them seven new species, partially
published in Richling (2001) — Helicina echandiensis, H. talamancensis, H. monteverdensis,
H. chiquitica, H. escondida, Alcadia (Microalcadia) hojarasca, and A. (M.) boeckeleri — and
two new subspecies - H. punctisulcata cuericiensis, and H. beatrix riopejensis. Other pre-
viously subspecifically separated taxa (H. funcki costaricensis Wagner, 1905; H. tenuis
pittieri Wagner, 1910) were shown to fall within the range of intraspecific variability. Records
of the Guatemalan and Mexican species Helicina oweniana L. Pfeiffer, 1849, and subspe-
cies, H. amoena L. Pfeiffer, 1849, as well as those of H. fragilis Morelet, 1851, were proven
to be based on faulty identifications and were therefore excluded from the Costa Rican
fauna. This fact, together with the recognition of the several new species, shows that the
faunal composition of Costa Rica is much more distinct from that of the northern areas than
previously assumed. The transitional zone of Nicaragua, however, still remains widely
uninvestigated. Only Helicina tenuis L. Pfeiffer, 1849, being ecologically very tolerant,
Lucidella lirata (L. Pfeiffer, 1847), and Pyrgodomus microdinus (Morelet, 1851) are wide-
spread, extending from Mexico to Costa Rica, perhaps even farther south. The distribution
of the typical Costa Rican species follows the topographical subdivision created by the
Central Cordilleras, along with its corresponding effects on the climate.

Contrary to former assumptions, certain features of the female reproductive system
proved very useful for the classification of the Helicinidae. For the first time, monaulic con-
ditions have been recognized for Helicina and Eutrochatella, necessitating the correction
of previous descriptions in this respect. Furthermore, the monaulic or diaulic state is char-
acteristic of the genera and 1$ paralleled by consistent changes in the embryonic shell struc-

195

196

RICHLING

ture. Because primitive members of the Helicinidae possess a diaulic system, the monaulic
condition is regarded as the derived state. The Central American genera Helicina, Alcadia,
Eutrochatella, Lucidella and Schasicheila were properly distinguished and described by this,
as well as by other differences in the female reproductive system. The anatomies of the type
species of Helicina and Alcadia were examined for the first time, and earlier descriptions
of Eutrochatella and Lucidella were corrected in major points. On the basis of this new
evidence, the assignment of traditional subgeneric units of Helicina and Alcadia, previously
based mainly on vague radula and shell characteristics, was especially reassessed. The
subgenera Sericea and Analcadia were transferred to Helicina, as well as the mainland land
species summarized under the preoccupied taxon “Gemma”. Tristramia, Oxyrhombus,
Pseudoligyra, Oligyra, Succincta, “Cinctella” [also preoccupied] and Punctisulcata were
confirmed in their association with Helicina. Due to its monaulic condition, the former ge-
nus Ceochasma is reduced to a subgenus of Helicina. In addition, exemplary non-type
Antillean species were studied, including Helicina jamaicensis Sowerby, 1841, which had
to be shifted to Alcadia $.1., and Alcadia (Analcadia) platychila (von Mühlfeldt, 1816), which
is now assigned to Helicina s.s. On one hand, the new arrangement excludes Alcadia as
previously known from the Central American mainland, but, on the other hand, examination
of the newly discovered Costa Rican species Helicina hojarasca and H. boeckeleri required
the establishment of a new subgenus of Alcadia, Microalcadia n. subgen. on the mainland,
based mainly on the features of the female reproductive system and embryonic shell struc-
ture. The occurrence of A/cadia with only a few diminutive species on the mainland of Cen-
tral America corresponds to the distribution of the genera Eutrochatella/Pyrgodomus and
Lucidella.

The Central American mainland species of Helicina seem to show a closer relationship
among each other than to the northern South American subgenera Analcadia and Sericea.
The Brazilian taxon Angulata, previously a subgroup of Helicina, deviates remarkably in
embryonic shell structure and shows differences in anatomy that still require final
confirmation, and it thus deserves recognition as a separate genus.

Contrary to the well-supported differentiation at the generic level, the attempt to
characterize subgroups of the Central American mainland species of Helicina has been only
partially successful. Certain similarities in teleoconch surface structure, relative development
of the accessory structures of the female reproductive system, and the degree of sexual
dimorphism become obvious and are discussed to some extent, but intermediate
characteristics complicate a satisfactory solution. Besides Ceochasma, three Central
American mainland subgenera are recognized: Oligyra, Tristramia most closely resembling
Helicina s.s., and “Gemma”. The latter preoccupied name is tentatively retained, because
the proposal of a new name seems inappropriate at this stage.

Investigation of the morphological features other than the embryonic shell sculpture and
the female reproductive system revealed the following additional results, mainly based on
the Costa Rican species of Helicina:

Characteristics of teleoconch, operculum, and radula, previously regarded as substantial
for classification, were repeatedly demonstrated to be subject to convergent development,
thus limiting their value for systematics. Different examples are given, such as the T-shaped
lateral of the radula or periostracal hairs, and further evidence is provided by the necessary
re-arrangements outlined above. Nevertheless, these features play a supplementary or
supporting role.

The mantle pigmentation of arboreal Helicinidae is closely related to the transparency of
the shell and functionally replaces shell color in thin shells. The physiological possibility of
an obviously adaptable mantle pigmentation could provide the opportunity for survival with
thin, transparent shells as adaptation to the limited availability of calcium carbonate.
Whereas varying and patterned mantle color are characteristic for arboreal thin-shelled
species, the color of the head and foot is seldom species specific.

Size differences of the embryonic shell have not previously been studied for Helicinidae.
Embryonic shell size is shown to increase with the shell size within a group of related
species and also altitude within different populations of a species. Furthermore, it may show

CLASSIFICATION OF HELICINIDAE

a certain species specificity. Preliminary data on Lucidella and Eutrochatella/Pyrgodomus
suggest a consistently smaller embryonic shell size than in Helicina or Alcadia.

Internal shell structures — axial cleft and muscle attachments - seem characteristic for
certain systematic units, for example, Lucidella and Schasicheila. The length of the axial
сей is confirmed to be constant within a species, but, contrary to former assumption, it 1$
not related to the whorl count.

The data on sexual dimorphism given in this study represent the most comprehensive
approach to date to analyze this phenomenon for Helicinidae. The sexual dimorphism may
manifest itself in differences in volume, a male's size being only about 62-70% of that of
the female's, but formerly assumed deviations in shape could not be proved to be of
significance for species of Helicina. A certain value for the degree of differences in
uncovering systematic affinities is indicated.

Keywords: Helicinidae, Costa Rica, Central America, classification, reproductive system,
radula, embryonic shell, new species.

RESUMEN

El presente estudio combina una revisión de los poco conocidos helicinidos de Costa
Rica con un análisis detallado de varias estructuras morfológicas y su utilización para
resolver preguntas sistemáticas. Con base en esto se discute a profundidad la clasificación
de las especies continentales de Centroamérica.

La revisión de las especies costarricenses se basa en un exhaustivo trabajo de campo,
en el análisis de casi todo el material tipo, de las colecciones del Instituto Nacional de
Biodiversidad de Costa Rica y del Museo de Historia Natural de Gainesville, asi como de
material histórico. Con pocas excepciones se estudiaron para todas las espécies los
carácteres de la concha, del opérculo, de la estructura superficial de la concha embriónica,
asi como de la teleoconcha, estructuras internas de la concha, la rádula, y el tracto
reproductor femenino. También se efecturon estudios sobre morfometría y dimorfismo
sexúal. Considerando la escacéz de material, para efectuar un estudio a gran envergadura,
fue necesario desarollar una metodología de disección para separar el cuerpo blando de
la concha sin ningún detrimento.

Para efectuar una clasificación más amplia y una comparación de las diferentes
estructuras morfológicas se tomaron los mismos datos de otras 17 especies, que
representan los taxa supraespecificos emparentados más importantes de Centroamérica.

Énfasis se puso en las estructuras poco estudiadas hasta ahora como la concha
embriónica y el tracto reproductor femenino. Hasta donde se pudo se trabajo con material
de especies que corresponden a los tipo. En donde no se pudo obtener material anatómico
para estudiar se interpretaron cuidadosamente los datos de la literatura.

Para Costa Rica se determinaron 15 especies, entre las cuales siete son nuevas, y en
parte publicadas en Richling (2001) - Helicina echandiensis, Н. talamancensis, Н.
monteverdensis, H. chiquitica, H. escondida, Alcadia (Microalcadia) hojarasca y A. (М.)
boeckeleri, además dos nuevas subespécies H. punctisulcata cuericiensis y H. beatrix
riopejensis. Los taxa subespecíficos, hasta ahora separados, H. funcki costaricensis
Wagner, 1905, y H. tenuis pittieri Wagner, 1910, caen dentro de la variación intraespecífica.
La presencia de las especies mejicanas y guatemaltécas Helicina oweniana L. Pfeiffer,
1849 con sus subespécies, H. amoena L. Pfeiffer, 1849 y H. fragilis Morelet, 1851 no fue
confirmada ya que el material estaba mal identificado y por esto se las elimina del listado
faunístico de Costa Rica. Debido a este hecho y al descubrimiento de algunas nuevas
espécies se puede distinguir la fauna de Costa Rica más claramente de otras regiones más
al norte de lo que se suponia hasta ahora. Nicaragua que es el territorio de transición está
casi inexplorado. Solamente las espécies Helicina tenuis L. Pfeiffer, 1849, que presenta
una gran tolerancia ecológica, asi como Lucidella lirata (L. Pfeiffer, 1847) y Prygodomus
microdinus (Morelet, 1851) se distribuyen desde México hasta Costa Rica y también más
hacia el sur. La distribución de las espécies típicas costarricenses sigue la subdivisión
topográfica de las cordilleras centrales y sus efectos correspondientes al clima.

197

198

RICHLING

Contrario a supociciones anteriores, se pudo demostrar que los carácteres del tracto
reproductor femenino son muy útiles en la clasificación de los helicinidos. Por primera vez
se pudieron reconocer condiciones monáulicas en Helicina y Eutrochatella por lo que
descripciones previas se deben corregir a este respecto. Además la condición monáulica
o diáulica son característicos para cada género y paralelamente hay cambios consistentes
el la estructura de la concha embriónica. Ya que los miembros primitivos de los helicínidos
poseen un sistema diáulico, la condición monáulica es considerada como derivada. Los
géneros centroamericanos Helicina, Alcadia, Eutrochatella, Lucidella y Schasicheila se
distinguen claramente y son descritos por estos y otros carácteres de la genitalia femenina.
La anatomía de las espécies tipo de Helicina y Alcadia se estudiaron por primera vez y las
descripciones anteriores de Eutrochatella y Lucidella se debieron corregir en varios puntos
importantes. Sobre esta nueva base, especialmente la asignación de las tradicionales
unidades subgenéricas de Helicina y Alcadia, que estaban previamente basadas en
carácteres vagos de la rádula y la concha, fueron reordenaron. Los subgéneros Sericea
y Analcadia se transfirieron a Helicina asi como las especies continentales comprendidas
bajo el taxón preocupado “Gemma”. La pertenencia a Helicina de Tristamia, Oxyrhombus,
Pseudoligyra, Oligyra, Succincta, “Cinctella” (también preocupada) y Punctisulcata se
confirma. Debido al tracto genital monáulico, el género Ceochasma se ordena como
subgénero de Helicina. Espécies de la Antillas solo se estudiaron ejemplarmente, Helicina
jamaicensis Sowerby, 1841, se incluyó dentro de Alcadia s.l. y Alcadia (Analcadia)
platychila (von Mühlfeldt, 1816) se le asigna a Helicina s.s. Por una parte estos datos
excluyen al género Alcadia del continente centroaméricano, por otra parte el análisis de las
nuevas espécies costarricenses encontradas de Helicina hojarasca y H. boeckeleri
requirieron la instauración del subgénero Alcadia, Microalcadia п. subgen., para el
continente basandose mayormente en los carácteres del tracto reproductor femenino y de
la concha embriónica. La presencia de A/cadia con solo unas cuantas pequeñas especies
en el continente corresponde a la distribución de Eutrochatella/ Pyrgodomus y Lucidella.

Las espécies continentales centroaméricanas de Helicina parecen estar más
emparentadas entre si, qué con los subgéneros Analcadia y Sericea del norte de
Suramérica. El taxón brasilero Angulata, subordinado a Helicina, posee una estructura de
la concha embriónica claramente distinta y diferencias anatómicas todavia por corroborar,
por esto se le considera como un género aparte.

Contraria a la clara diferenciación a nivel genérico, el intento de agrupar las espécies
continentales de Helicina ha sido solo en parte exitoso. Algunas similitúdes en la estructura
superficial de la teleoconcha, el desarróllo relativo de las estructuras accesórias del aparato
reproductor femenino, y el grado de dimorfismo sexual son obios y se discuten en parte,
pero estadios intermedios complican la solución satisfactoria de este problema. Además
de Ceochasma, se reconocen tres subgéneros centroaméricanos continentales: Oligyra,
Tristramia muy parecido a Helicina s.s. y “Gemma”. Este último está preocupado pero se
le retiene tentativamente, ya que proponer un núevo nombre a este nivel по se considera
apropiado.

El estudio de otros carácteres morfológicos diferentes a la estructura de la concha
embriónica y del aparato reproductivo femenino revelan los siguientes resultados
adicionales, basados especialmente en las espécies costarricenses de Helicina:

Se demostró repetidamente que las características de la teleoconcha, opérculo y rádula,
previamente considerados fundamentales para la clasificación, son objeto de desarrollos
convergentes, limitando asi su valor sistemático. Diferentes ejemplos son dados como el
diente lateral en forma T de la rádula o los filamentos del perióstraco. Evidencia adicional
es dada en la reorganización requerida mencionada anteriormente. Sin embargo estos
carácteres juegan un papel suplementario o de soporte.

La pigmentación del manto de los helicínidos arbóreos está fuertemente relacionada con
la transparencia de la concha y reemplaza funcionalmente la coloración de la concha en
conchas delgadas. La posibilidad fisiológica de la pigmantación del manto se considera
como una adaptación obia que permite la supervivencia con conchas delgadas, en
ambientes con poco calcio. Mientras que los patrónes de coloración del manto son

CLASSIFICATION ОЕ HELICINIDAE

característicos para las espécies arbóreas con conchas delgadas, la coloración de la
cabeza y el pié son raramente característicos a nivel de espécie.

Las diferencias del tamaño de la protococnha no se habian estudiado previamente en los
helicínidos. Se demuestra que el tamaño de la concha embriónica aumenta con el tamaño
de la concha en un grupo de espécies relacionadas y con la altitúd de la localidad de
differentes poblaciones de una espécie. En algunos casos el tamaño de la concha
embriónica puede ser característico para una espécie. Primeros datos de Lucidella y
Eutrochatella/Pyrgodomus muestran constantemente una concha embriónica más pequeña
que en Helicina o Alcadia.

Las estructuras internas de la concha - apertura axial e inserción de los músculos —
parecen ser característicos para algunas unidades sistemáticas, e. j. Lucidella y
Schasicheila. La longitud de la apertura axíal es constante dentro de las espécies, pero,
contrariamente a lo que se suponia, no está relacionada con el número de vueltas.

Los datos sobre dimorfismo sexúal dados en este trabajo representa la aproximación más
amplia hasta la fecha para analizar este fenómeno en los helicinidos. Este estudio muestra
diferencias significativas en el volúmen, en donde los machos en casos extremos
solamente alcanzan aproximadamente entre el 62 y el 70% del tamaño de las hembras.
Contrariamente a las suposiciónes anteriores no se pudieron comprobar diferentes formas
en el género Helicina. El grado de dimorfismo sexúal parece tener también valor al

199

determinar las relaciones de parentesco.

INTRODUCTION

The Helicinidae and a few related families
belonging to the Neritopsina represent the
earliest branch of gastropods evolved to ter-
restrial existence from as-yet unknown
diotocardian marine ancestors. Their recent
distribution encompasses two main regions —
the subtropical and tropical zones of North
and South America and the Indopacific and
Pacific islands and small areas of the Asian
and Australian continents. А particular high
diversity has developed on the Caribbean Is-
lands and оп the Philippines. The family 1$
comprised of approximately 550 species, of
which a little more than half occur in the New
World. Most species are small, with only the
largest representatives reaching nearly 3 cm.

Early classifications of the Helicinidae were
based on shell characters only (e.g., L.
Pfeiffer, 1850-1853). Later, Wagner (1907-
1911) provided the still most extensive, but
much criticized (e.g., Fulton, 1915; Solem,
1959: 166-167) monograph on the family
worldwide, incorporating features of the oper-
culum for his systematic arrangement. At
about the same time, a very detailed, compre-
hensive anatomical investigation of several
New and Old World species of different gen-
era, including histology, was published by
Bourne (1911). His study demonstrated a con-
siderable uniformity of the morphological
structures within the Helicinidae, indicating
their very limited value for revealing system-

atic affinities, especially with respect to the
reproductive system. In conclusion, Bourne
(1911) favored radular characteristics as the
safest feature for a classification. Baker
(1922a, 1923) followed this concept to clarify
the relationships of the American mainland
taxa (United States to northern South
America) and nomenclaturally corrected,
modified and consolidated the system of
Wagner (1907-1911) through radula charac-
teristics that were believed to provide system-
atically relevant information. Subsequent
anatomical studies on the same group of spe-
cies with emphasis on the reproductive sys-
tem (Baker, 1926, 1928) did not allow similar
conclusions to be drawn due to the uniformity
of the structures and the limited material. Later
authors interpreted the results in the sense of
Bourne (1911) and stated that the “... general
uniformity of the genitalia of the Helicinidae
makes them useless for diagnostic purposes”
(Boss & Jacobson, 1974: 6). The radula char-
acteristics were partly accepted, but other
authors questioned their value for certain taxo-
nomical units (Rehder, 1966; Boss &
Jacobson, 1973). The most recent contribu-
tion to systematic issues of Helicinidae by
Thompson (1982) highlights the conservative
character of embryonic shell sculptures as a
criterion for determining relationships, but its
further application was beyond the scope of
his study on a species-group from the West
Indies. The few publications dealing with the
classification of the Australasian and Pacific

200 RICHLING

species are based on shell structures or
vague differences in the radula, respectively.
In conclusion, it can be stated that the system-
atics within the Helicinidae still remain contro-
versial, and due to the fact that the studies of
the different structures were mostly based on
different taxa, they are not comparable and
any interpretation is, at the very least, partially
questionable.

Faced with the absence of a detailed inves-
tigation of the applicability of different features
to reveal affinities within the Helicinidae and a
comparison of all these characteristics for one
and the same group of species, this study tries
to bridge this gap. Furthermore, preliminary
studies on the female reproductive system of
a Costa Rican species showed deviations
from the previous results, rendering these or-
gans more informative than described above.
Therefore, the present study intends to inves-
tigate several morphological characters for
their value in determining relationships on the
species level (rather highly adaptable) and in
higher systematics (rather conservative). The
chosen characteristics will encompass widely
applied aspects, such as shell in general,
operculum and radula, but will also focus on
less investigated or neglected structures, such
as teleoconch surface structure, embryonic
shell, internal shell structures, female repro-
ductive system, and the phenomenon of
sexual dimorphism.

Because single structures can only partially
be assessed in their possible adaptability by
their complexity and functionality alone, they
also have to be discussed within the context of
the best possible, well-founded synthesis of all
possible characteristics, that is, the proposed
classification. Therefore, the analysis of struc-
tures is based on revision of one group of
closely related species (species level) and
study of other related supraspecific taxa
(higher systematics, e.g., type species of re-
spective genera and subgenera). This will re-
sult in a new proposal for the classification of
the taxa studied, which is compared with pos-
sibly deviating previous concepts.

This study will be based on the Costa Rican
representatives of Helicinidae, which encom-
pass a reasonable number of species for de-
tailed analysis. According to Wagner
(1907-1911) and Baker (1922a, 1926), most
of the species belong to one or two genera,
Helicina or Helicina and Alcadia. Single spe-
cies of Lucidella and Pyrgodomus represent
relatives of genera with otherwise Antillean
distributions. Two newly discovered species

(Richling, 2001) still await proper classifica-
tion. Thus, а fairly wide scope of systematic
units is included and, with respect to the Costa
Rican species, part of the Central American
mainland fauna has been chosen for which
the most data for comparison, mainly from the
works of Baker, are available.

The focus on the Costa Rican species pro-
vided the opportunity to carry out a revision of
the Helicinidae of a poorly investigated area
as well. Because von Martens (1901: xii) still
has characterized the molluscan fauna as
“one of the best known within Central
America”, a few scattered publications in the
1930s (e.g., Pilsbry) remained in complete
neglect until recently, when the growing inter-
est in tropical biodiversity, spearheaded by the
foundation and work of the Instituto Nacional
de Biodiversidad de Costa Rica (INBio), re-
sulted in a new approach. The cooperation
with the Zaidett Barrientos of the Malacology
Section of INBio in providing access to the
comprehensive collection of national molluscs
greatly ameliorated the disadvantage of the
unfavorable geological conditions of Costa
Rica for collecting terrestrial snails which re-
sult in extremely low abundances and there-
fore present practical difficulties for obtaining
sufficiently large numbers of specimens for
certain aspects of the study.

MATERIALS AND METHODS
Area of Investigation

Costa Rica is situated in southern Central
America adjacent to Nicaragua to the north
and Panama to the south (about 8° to
11°15’N). Located between the Pacific Ocean
and the Caribbean Sea, small area of just
51,100 square kilometers rises up to 3,820 m
above sea level. The central mountain chain,
northwest to southeast in orientation, sepa-
rates a larger Caribbean from a hilly Pacific
plain. The mountains are subdivided into the
northern Cordillera de Guanacaste, the Cordil-
lera de Tilaran, and the Cordillera Central, a
chain of volcanoes, some of them still active,
and the southern Cordillera de Talamanca
which has been uplifted as a result of the sub-
duction of the Cocos Ridge (Fig. 1).

The climate is characterized by a dry anda
rainy season, with the dry season lasting from
about December to May. Whereas the north-
western and central parts of the country really
experience a dry period, the southern Pacific

CLASSIFICATION OF HELICINIDAE

side as well as the Caribbean side always
have humid conditions. This is reflected in the
variation of the vegetation, the tropical dry for-
est only being found in the northwestern area
in the transition to the Peninsula de Nicoya.
The vegetation of the remaining part of the
country is classified as moist, wet or rain for-
est (Tosi, 1969), with the humidity mainly in-
creasing With the altitude (Fig. 2). The distribution
of the annual precipitation is given in Fig. 3.

Materials

Fieldwork:

Costa Rica. The field work was carried out
on five visits of about 4 to 9 weeks each to

201

Costa Rica between 1998 and 2001. With
one exception during the rainy season of July
to September, the field trips were carried out
during the dry season in February and March.
Several localities scattered around the coun-
try were investigated for distributional data.
Selected areas were visited several times in
order to gather sufficient material of certain
populations for comparative studies, because
their abundance in the tropical rain forests is
very low. Due to the arboreal life-style of most
of the Costa Rican species, manual searches
had to be conducted. The detailed material
and localities are listed under each species.
Main collecting sites are shown on the gen-
eral map (Fig. 1).

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FIG. 1. Map of Costa Rica, including the most important collecting sites, the central mountain chains,

and the provinces.

202

RICHLING

| rain forest

moist forest
dry forest

> 6500 mm

5500-6500 mm

| 4500-5500 mm

| 3500-4500 mm

| 3000-3500 mm
2500-3000 mm

| | 2000-2500 mm
1500-2000 mm

<1500 mm

FIG. 3. Annual precipitation in Costa Rica [mm/year] (based on
Ministerio de Agricultura y Ganaderia & Instituto Meteorologico
Nacional, 1985).

CLASSIFICATION OF HELICINIDAE 203

JAMAICA/UNITED States. Because the type
species of the most important Central Ameri-
can genera (Helicina, Alcadia, Lucidella,
Eutrochatella) occur in Jamaica, and pre-
served material was not available in collec-
tions, supplementary research was carried
out during two weeks in May/June 2001.
Specimens of Helicina orbiculata (Say,
1818) were collected in Gainesville, Florida,
in May 2001.

Museum Collections: Material of the following
institutions has been studied, subsequently
only the abbreviations will be used:

ANSP Academy of Natural Sciences of
Philadelphia, Philadelphia, USA
(Dr. Gary Rosenberg, Dr. Igor
Muratov)

APHIS- Malacological Collection of United

PPQ States Department of Agriculture,
Philadelphia, USA (Dr. David G.
Robinson)
The Natural History Museum, Lon-
don (formerly British Museum,
Natural History), Great Britain (Dr.
Fred Naggs, Richard Williams)
HNC Haus der Natur Cismar, Germany
(Dr. Vollrath Wiese)
INBio Instituto Nacional de Biodiversidad
de Costa Rica, Santo Domingo,
Costa Rica (Dr. Zaidett Barrientos)
IR Material collected by Ira Richling,
partially deposited as vouchers at
INBio, otherwise accessible
through the collection of the НМС;
a few of the numbers refer to field
observations only
Musée d'Histoire Naturelle,
Neuchâtel, Switzerland (Dr. Jean-
Paul Haenni)
MIZ Museum and Institute of Zoology
of the Polish Academy of Sci-
ences in Warszawa, Poland
(Prof. Dr. Adolf Riedel)
Naturhistorisches Museum Bern,
Bern, Swiss (Dr. Margret Gosteli)
Nationaal Natuurhistorisch Mu-
seum, Leiden (formerly
Rijksmuseum van Natuurlijke
Historie), The Netherlands (Wim
Maassen)
SMF Naturmuseum und Forschungs-
institut Senckenberg, Frankfurt a.M.,
Germany (Dr. Ronald Janssen)

MHNN

NMBE

RMNH

UF Florida Museum of Natural His-

tory, Gainesville, USA (Dr. Fred

G. Thompson, John Slapcinsky)

United States National Museum,

Washington, D.C., USA (Dr. Rob-

ert Hershler)

ZMB Museum für Naturkunde, Humboldt-
Universität, Berlin, Germany (for-
merly Zoological Museum Berlin)
(Dr. Matthias Glaubrecht)

ZMH Zoologisches Museum, Universität
Hamburg, Hamburg, Germany (Dr.
Bernhard Hausdorf)

USNM

INBio: Within a context of considerable re-
cent efforts towards an inventory of the
biodiversity of Costa Rica, the institute
houses a very extensive collection of mol-
luscs. All available specimens of Helicinidae
from this material were studied, partially dur-
ing personal visits in Costa Rica, partially by
loans to Germany.

UF: This institution houses probably one of
the most comprehensive collections of Cen-
tral American terrestrial molluscs. During a
two-week visit, about 1,100 lots of
Helicinidae were studied with the emphasis
on the mainland species yielding consider-
able distributional data.

ZMB/MHNN: The only important historical
collections in Costa Rica were made by the
Swiss naturalists Biolley and Pittier at the
end of 19" century. Their material ended up
in different collections, parts of it in the ZMB
and MHNN respectively, other parts re-
mained in the Museo Nacional in San Jose,
Costa Rica (see under Helicina pitalensis).
The ZMB collection was visited personally,
whereas material in the MHNN was
searched for by J.-P. Haenni, Neuchätel,
and kindly loaned to the author. According to
J.-P. Haenni, an up-to-date catalogue of the
mollusc collection does not exist and the
materials of Pittier and Biolley are scattered
throughout the collection, which has never
actually been catalogued and which was
moved in the past, and it is possible that
some of the material has not yet been found.
A detailed list of material studied is given
under each species.

Locations/Maps: During the field work, coordi-

nates of the localities were registered using
the Global Positioning System (Magellan
GPS 3000) whenever possible, otherwise

204

they were taken from maps in 1:50,000
scale produced by the Instituto Geografico
Nacional, San José, Costa Rica, in different
editions, but all based on data from between
1961 and 1966. The staff of INBio uses the
same maps. All Costa Rican records from
literature or other sources without exact data
were localized as accurately as possible and
coordinates were estimated based on the
map: Los Parques Nacionales y otras areas
protegidas de Costa Rica. — Fundacion
Neotrópica, San José, 1993, | Reimpressión
1995. Information on some historical collect-
ing sites was provided by Zaidett Barrientos
and Maribel Zuñiga, INBio. All further expla-
nations that were subsequently added are
given in brackets.

The map of Costa Rica used throughout this
study is based on: Costa Rica. Mapa fisico-
politico 1:500.000 — Instituto Geográfico
Nacional, San José, edition 1987.

Methods

Measurements: The following linear measure-
ments (Fig. 4) were used, when measure-
ments of single species given, the following
sequence is given, separated by */” (unless
otherwise stated):

height

major diameter
greatest diameter
minor diameter
expansion of outer lip

of outer lip

N height of
А last whorl
height
= A | expansion

heig

columellar axis

RICHLING

height of last whorl
height of columellar axis

Because some helicinid species display
variation in the development of the outer lip
which mainly influences the measure of the
greater diameter, measurement of “major
diameter” has been introduced. This mea-
surement was taken just behind the reflec-
tion of the outer lip (Fig. 4). For the height,
this modification was not applied, because it
is not affected as much and 1$ furthermore
not uniformly practicable.

The greatest diameter is usually included
only in measurements given for single speci-
mens to comply with traditional measure-
ments.

Measurements were taken with a microme-
ter gauged on 0.01 mm scale. In view of the
deviation shown below that were minimized
by personal experience, values given to
characterize single specimens were
rounded to a 0.1 mm scale. Deviations are
mainly due to effects of an imperfect perpen-
dicular orientation of the shell with respect to
the measuring axis, a problem that can be
minimized with experience if the same per-
son carries out the measurements. How-
ever, errors probably cannot be excluded in
globular shells, but their range 1$ tolerable.
To check the average deviations, three
shells of different shapes were measured
the different times and the mean value, the
standard deviation and absolute deviations
were analyzed (Table 1).

major diameter

minor
diameter

ht of

FIG. 4. Measurements and counting of postembryonic whorls.

205

CLASSIFICATION OF HELICINIDAE

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206

In addition to the linear measurements, the
weight and volume of empty shells were
analyzed. Sartorius scales (scale 0.001 g)
were used. The volume was measured as
the difference of weight of the shell filled with
distilled water and the weight of the empty
shell. To obtain comparable data, shells
were always filled until the water showed a
plain surface in the aperture. In the weight
measurements, the hole caused by the
preparation procedure did not influence the
results, because the wall at the beginning of
the last whorl is thin and the amount of ma-
terial removed was below the scale of reso-
lution.

Except for the specimens studied with the
SEM, the diameter of the embryonic shell
was measured under a stereomicroscope
(scale 20 um). Otherwise, measurements
were taken from photographs, which are
much more exact. Whorls were counted ac-
cording to Fig. 4.

Fixation: The preservation of the live collected
material was carried out in two ways:

(1) Collections until 1999: Specimens were
relaxed in water for several hours and sub-
sequently transferred to isopropanol (about
80%).

This method has disadvantages: lt is difficult
to find the right time to stop the relaxation
process, because it depends on so many
factors, such as specimen size, water vol-
ume and temperature. Under the conditions
of field work and travel by bus, it is difficult to
carry out lengthy procedures. Furthermore,
the shocks received during transport also
influence timing. As a result, the specimens
may be badly preserved or contracted. In
case they close their opercula again, there
remains the risk that the alcohol will not pen-
etrate into the shell.

Beginning in 2000, | developed the new
method to remove the body from its shell
described below, which allowed another
preservation method mitigating these disad-
vantages.

(2) Collections after 1999: Specimens were
removed from the shell alive and immedi-
ately dropped in isopropanol.

It has the advantage that specimens can be
preserved immediately and with a constant
result. The problem of the closure of the
operculum becomes irrelevant. If a relax-
ation is required for subsequent investiga-
tions, the body can still be dropped in water
or other solutions and will be anaesthetized

RICHLING

much faster due to the greater unprotected
surface for medium exchange. For the
present study and due to the need of a fast
working method, a relaxation process has
usually not been applied, because retrac-
tions are limited to the foot and the two por-
tions of the retractor muscle and do not
greatly affect other organs.

Preparation and Storage of Material: Against

the background of the low abundance of
Helicinidae in Costa Rica and the various
aims of the study (e.g., aspects of anatomy,
sexual dimorphism), two requirements had
to be met at the same time: the shell and the
animal had to be separated and they had to
be kept as intact as possible.

When normally pulling a more or less re-
laxed animal out of its shell, in most cases
the head-foot and the anterior pallial portion
will be released, but the remaining part will
be torn off within the shell. This is due to the
fact that in the Helicinidae, contrary to most
other gastropods, by the dissolution of the
inner whorls of the shell, the visceral mass
forms one large complex, which has a
greater diameter than the remaining part of
the last whorl or aperture respectively,
through which it has to pass. Furthermore,
air cannot penetrate to allow the body to be
released. Besides the obvious disadvan-
tages, the resulting rupture of the body di-
rectly divides the pallial gonoduct at an
important section and often makes its study
impossible.

In a newly developed method, a small hole
is made on the periphery within about the
second quarter of the last whorl (Fig. 5, ar-
row) with a nail file or insect needle of differ-
ent size, depending on the shell thickness.
This can be performed without injury to the
animal when applying the method to live in-
dividuals. Subsequently, a needle, curved if
necessary, is carefully inserted between
shell wall and body and the two retractor
muscles are detached. Afterwards, the ani-

FIG. 5. Hole for removal of the body.

CLASSIFICATION OF HELICINIDAE 207

mal can easily be removed by pulling the
operculum (live animals) or by a needle in-
serted in the foot (preserved animals). One
must be careful to allow air to enter the hole.
Live animals can then be fixed. In preserved
specimens, it is usually more complicated
during the final removal to avoid the damage
described above, because the visceral mass
is no longer very flexible or may suffer from
poor preservation. By the aid of the needle
(through the hole), the visceral mass then
has to be squeezed through the remaining
part of the last whorl. The success in pre-
served specimens greatly depends on the
shell shape (relation of shape and volume of
visceral mass to the diameter of the aper-
ture) and the prior fixation. During the
present study, the method seldom failed.

In my own material, shells were separated
from the bodies in all adult and live collected
specimens. They were individually stored,
enumerated and labeled.

Sex Determination: The determination of the
sex was done by external inspection of the
soft body. According to Baker (1926) and
personal experience, in most cases and
many species a dissection is not necessary.
Females are recognized by the compara-
tively small lobes of the ovary, widely
spaced, regular constrictions of the pallial
gonoduct (not in all species), and the dark
color of a distinct portion of the distal pallial
gonoduct. Males are characterized by the
comparable larger lobes of the testis, the
absence of the distinct dark color, a very
densely lobed apical, and smooth distal part
of the pallial gonoduct. In some cases, the
shiny white vas deferens may shimmer
through the visceral mass. Normally not all
these features are visible in one and the
same specimen, but each one may be un-

digestive /

y 1 Jen
gland =
Г п i 1

ovary ten) ey

intestine

Vee N IN A
ja ‚+77 hypobranchial gland

pallial reproductive system 4 417

LT DS
--/ / NON
/
< \
/ =
/ x м
1
\
)
4 1
es! 1
y \
/ \
\ /

equivocal. { mainly depends on the body
pigmentation, the species and the individual
development. In ambiguous cases, the
specimen was dissected.

Reproductive System: Dissections were made

in 70% isopropanol or ethanol. For the in-
vestigation of the reproductive system, the
mantle cavity was opened along the left side
of the intestine, with the latter remaining
along the pallial gonoduct. А second cut was
made between the pallial gonoduct and the
right retractor muscle, along or through the
hypobranchial gland up to the apical part of
the pallial portion of the reproductive system
(Е9.6):

Histology: The separated female reproductive

system was dehydrated through a series of
ethanol, transferred to paraffin via acetone
(100%) and embedded in paraffin. Serial
sectioning was done at 5-7 pm with a sliding
microtome. The tissue was subsequently
stained with a sequence of paraldehyde
fuchsin solution, nuclear fast red and orange
G/ light-green.

Preparation of Shells for SEM: In order to re-

duce lasting effects to the shell by gold coat-
ing, the specimens were usually mounted on
aluminum specimen stubs using adhesive
conductive tape. Subsequently, they were
tightly covered with laboratory film (Parafilm
“М”®), which adhered to the remaining sur-
face of the adhesive conductive tape. Fi-
nally, the embryonic shell or other areas of
interest were uncovered and coated. After
the SEM investigation the laboratory film can
easily be removed and the shell extracted.

Preparation of the Radula: The radula was re-

moved from the buccal mass. It was cleaned

stomach

kidney
heart and pericard
digestive gland

y D, Y A Up N
right retractor a / / 5 mantle cavity
muscle fey /

( (ex ih /

WE J
operculum NYY A

A

FIG. 6. General anatomy of the Helicinidae.

ovary

\ =9— left retractor
Y ANY muscle

operculum

208 RICHLING

from remaining tissue in NaOH-solution (1
N) for about 24 h at 50°C. Subsequently the
radula ribbon was washed in distilled water
several times and dehydrated through a se-
ries of ethanol (70%, 80%, 96%, 100%). Af-
ter the pure alcohol, it was dried and
arranged to the final mounting position with
the marginal teeth turned up by using prepa-
ration needles. Finally, the radula was
mounted with conductive carbon cement on
the aluminum specimen stubs for SEM ex-
amination. Only a few radulae were studied
with the light microscope. For light micros-
copy, the radulae were transferred to the
slides directly after removal.

SEM Investigation of Shells and Radulae:
Samples were sputtered with gold for 140
sec by using a BALTEC SCD 050 Sputter
Coater. Investigations were carried out with
a LEO 420 scanning electron microscope
(LEO V 02.04). Radulae had to be studied
under low voltage conditions (about 2.5 kV),
because the structure of the rhipidogloss
radula causes extremely high charge distri-
butions, rendering adequate studies and
exposures under high voltage conditions
impossible.

Figures: Unless otherwise stated, all draw-
ings, maps and photographs in the study
were made by the author. Drawings were
made at a LEICA MZ 8 stereomicroscope by
the aid of a camera lucida. Except for Fig-
ures 140, 228, 249, 257, and all live animals
photographed with a 35 mm SLR camera, all
shells were digitized with a Sony Digital Still
Camera DSC-F505V.

Addditional Abbreviations:
ad./ads. — adult/s
coll. — collection
juv./juvs. — juvenile/s
SEM - scanning electron microscope

RESULTS

The results are presented into two parts: (1)
Revision of all Costa Rican species of
Helicinidae including the investigation of the
shell — general aspects, internal structure,
surface structure, embryonic shell, morphom-
etry and sexual dimorphism — the radula, the
soft body color, the female reproductive sys-
tem, and data on the habitat and distribution.

(2) The morphological characters of the
supraspecific taxa relevant for the classifica-
tion of the Central American mainland
Helicinidae.

GENERAL ASPECTS

The discussion under each species will fo-
cus on the species-relevant data. Aspects of
the morphological characteristics will be dis-
cussed in context with the classification sub-
sequent to the Results, as will some general
results for the Costa Rican fauna and the clas-
sification of the Helicinidae. The account for
each species has the following outline, in
which | have here included an overview of the
morphological characters.

Literature Records (without heading): All litera-
ture records of the respective species are
listed. In some cases of questionable deter-
minations, attempts to re-examine the origi-
nal material were made. Some citations
nevertheless remained uncertain, those are
marked by a “?”.

Synonymy: For clarity, the synonyms are
reitterated from the Literature Records. This
includes only synonyms that were proved
and accepted during this study.

Original Description: Complete citation of the
Original description.

Type Material: This exclusively includes the
type material of the respective species.

Type Locality: Only the type locality of the re-
spective species is given under this heading.

Type Material of Synonymous Taxa or Similar
Species: If necessary for comparison, infor-
mation on the type material of synonymous
taxa or similar species is also provided, be-
cause for many Central American taxa ad-
equate figures cannot be found in the
literature. For those species, the type local-
ity is given here.

Examined Material: For a better finding of the
data of the lots, the material is arranged ac-
cording to the collections (leg. |. Richling,
collection INBio, other sources) and, only
secondarily, according to localities (Costa
Rica: different provinces; other countries).

CLASSIFICATION OF HELICINIDAE 209

FIG. 7. Changes in shell surface structure exemplary shown for Helicina gemma. А. |: Embryonic shell;
|: Transitional structure; Ш: Oblique diverging grooves; IV: Smooth surface with fine growth lines.
В. Enlarged view of the transitional structure (section II). С. Pattern of oblique diverging grooves with
transformation of section Il. D. Enlarged view of the smooth surface with fine growth lines (section IV)
scale bars 500 um (A); 100 um (B-D).

210

To shorten the descriptions of the localities
and to facilitate the search for the complete
locality data mentioned in the text, the fol-
lowing typological convention is used for the
Richling and INBio material. In the case of a
single locality for lot(s), only a shortened
name of the locality is set in italics. In the
case of several lots from sublocalities, the
entire general description is in italics, fol-
lowed by a colon; the colon 1$ then followed
by further specifications in italics applying to
the lots following the second colon. In some
cases, there 1$ a further subdivision, such as
altitude, given in the same format. Locality
data in roman type refer only to the subse-
quent lot.

Description:

Shell: General description of the species.
Internal Shell Structures: Contrary to most
other gastropods, Helicinidae dissolve the
inner parts of their shells so completely that
only a septum of a certain length subdivides
the shell internally. This septum extends
from the remains of the columella to the su-
ture of the last whorl. The length of the sep-
tum or, referring to the soft body, of the axial
cleft is figured here. Additionally, the posi-
tions of the attachments of the retractor
muscles are shown. In Helicinidae, the col-
umellar muscle in separated in two portions,
one attaching somewhere in the umbilical
area, the other in the upper part of the shell,
often close to the beginning of the axial cleft.
Teleoconch Surface Structure: During
growth, the Helicinidae produce different
shell surface structures. A possible se-
quence of different patterns is shown for
Helicina gemma (Fig. 7) covering the varia-
tions among in Costa Rican helicinids.

The embryonic shell (Fig. 7A: 1) is sharply
distinguished from the teleoconch by a dis-
tinct pattern and a more or less clear growth
mark. The subsequent part exhibits an ir-
regular, coarse and wrinkled surface (“tran-
sitional structure”) (Figs. 7A: Il, В). It
changes continuously with pits elongating to
grooves to a pattern consisting of groups of
parallel grooves that diverge acutely with
other obliquely orientated groups of grooves
(Figs. 7A: Ш, С). The grooves follow two
main orientations (this structure will subse-
quently be referred to as “pattern of oblique
diverging grooves”). Finally, this pattern 1$
predominated by fine growth lines forming
an otherwise smooth, shiny surface (Figs.

RICHLING

ТА: IV, D). Only a vestige of the oblique
grooves may still be visible. This surface
structure is maintained to the aperture.

This general scheme 1$ not completely real-
ized in all Costa Rican Helicinidae, certain
sections may be absent, for example, the
pattern of oblique diverging grooves contin-
ues for the rest of the postembryonic shell
and the smooth surface is absent. Apart
from differences in detail, the pattern of the
major part of the postembryonic shell does
not change again and starts at the latest at
the beginning of the second whorl (Fig. 7A,
III-IV). Therefore, a section of this whorl is

preferably described for species compari-

sons.

Embryonic Shell: If available, at least three
specimens of each species or subspecies
respectively were investigated for embryonic
shell structures. Individuals were chosen
randomly and depending on the preserva-
tion. Especially in cloud forest areas, the
embryonic shell seems to erode very
quickly.

Unless otherwise stated, relative descrip-
tions refer to the structures of Helicina
funcki.

Operculum: The operculum of most species
of the Helicinidae is concentric and consists
of two plates, an inner horny plate (attached
to the foot) and an outer calcareous plate.
The horny plate projects beyond the margins
of the calcareous plate. In all Costa Rican
species except for Pyrgodomus, the calcar-
eous plate is thin and becomes thickened
only towards the columellar edge, determin-
ing the shape of this margin, whereas the
palatal margin is shaped by the further ex-
tending horny layer; the calcareous layer
becomes indistinguishable and normally
does not reach this margin.

Animal: Expecting species-specific differ-
ences in the mantle color of certain species,
as many specimens as possible were docu-
mented as to their color, but due to consid-
erable variation, especially among different
populations, the comparison did not reveal
many species-specific differences. A gener-
alized description will be given for each spe-
cies.

Radula: The helicinid radula consists of
three groups of teeth: the centrals, the later-
als and the marginals (Fig. 8). The central
field is composed of an unpaired central or
rhachidian tooth (R), which is flanked by
three paired teeth, called A-, B-, and C-cen-

CLASSIFICATION ОЕ HELICINIDAE 211

FIG. 8. Part of radula ribbon (shown in Helicina
funcki); “А”-”С”: respective central teeth, ap:
accessory plate, cl: comb-lateral, В: rhachidian
tooth; scale bar 100 pm.

tral with the A-central aside to the rhachidian
tooth (some authors such as Keen, 1960;
Thompson, 1980, but not 1982; Stanisic,
1997, include the three paired teeth into the
laterals). The laterals, also called the
capituliform complex, are formed by two par-
tially fused teeth, the inner comb-lateral (cl)
and the accessory plate (ap). Within the
Helicinidae, the comb-lateral is developed in
two main types: (1) the true “comb”-lateral: a
broad tooth with numerous cusps at the cut-
ting edge (Fig. 8) or (2) a very strong tooth
T- or mushroom shaped (also called T-lat-
eral) without cusps (Fig. 246B). The
marginals encompass numerous long, slen-
der teeth in oblique rows that bear a varying
number of accuminate cusps. The terminol-
ogy follows Baker (1922a).

The radulae of the Costa Rican species of
Helicina do not show many differences
among the individual species, but within
populations of the species themselves, there
is some variation, especially regarding the
number of cusps on the central teeth. A cer-
tain number of cusps is usually not ex-
ceeded, but the cusps are often vestigial or
absent, forming a crenulate margin at the
cutting edge. Throughout each radula, the
different teeth are very uniformly developed
with a very constant numbers of cusps.
The rhachidian tooth is triangular to trap-
ezoid shaped and lacks cusps. The A- and
B-centrals project laterally, with broad faces
forming an oblique cutting edge. The C-cen-
tral narrows towards its face and represents
the outer tip of the central cutting edge.

The two teeth of the capituliform complex
were always observed to be fused, and, un-
der the conditions and the magnifications
studied with the SEM, the demarcation line
between the teeth was not visible. The
cusps on the comb-lateral only show in-
traspecific fluctuations of one or two, but
aberrant developments do occasionally oc-
cur (e.g., many more cusps or lacking any at
all). The relative size of the cusps appears to
be constant. In most species, the cusps
slightly decrease in length towards both
ends of the edge, with the inner a little
longer. The accessory plate 1$ usually
slightly smaller than the comb-lateral and
projects laterally.

With the occasional exception of the inner-
most tooth, the marginals increase in num-
ber of cusps outwards starting with 2-3 to
more than 10. Two tendencies were recog-
nized: (1) slowly and (2) rapidly increasing
number of cusps; in the first, there are re-
markably more teeth with 2, 3 and 4 cusps,
that is, also more teeth with pronounced ter-
minal cusps, whereas in teeth with more
cusps the latter tend to arrange themselves
laterally along the tip which 1$ therefore
turned sidewards to bring the cutting edge
into action.

In the account on the radula for the species,
only the distinguishing features are outlined
in addition to the figures.

Unless otherwise stated, the radulae of at
least three specimens of each species and
in some cases also of different populations
were investigated. For Helicina funcki and H.
beatrix riopejensis n. subsp. eight speci-
mens were studied to check for intraspecific
variability.

Female Reproductive System: Parts of the
reproductive system of the Helicinidae show
several peculiarities for which authors have
introduced special terminology (Bourne,
1911; Baker, 1925 & 1926). Because terms
were exchanged and confused, a summary
is given, and the present use is indicated
(Table 2, Fig. 9). The terminology implies
certain functional aspects, but the function
has been controversially discussed for differ-
ent taxa (e.g., Bourne, 1911) and still re-
mains partially doubtful, especially with
respect to the structures for sperm storage.
The terms used in this study follow the tradi-
tional usage and strike a balance between
possible confusions, but will not be modified
for functional correctness to avoid any fur-

oes |ешбело.а

хщепаоэ esinq

ula] эшоэа$ ou

SIUILUSS
uunjnoejdeces
oes
wiads 1105539998

jonpino |ееа

RICHLING

oes |ешбело.а

esinq Álojendoo
/xigyejndoo esinq

oes

weds Ало$$аээе
oes

wueds Ало$$аээе

JONpIAO feed

oes jeulbeaoid
esinq |едиэл
(ejlayaıseyaS JO}) Des

uyods Ало$$аээе Alepu09as

oes uads А10$$э99эе

saunjon1]s A10SS299\/

snjon

oes jeulBeaojd

xI1Je¡ndoo
esing Jo eoayjeuuads

oes wiads А10$$э99эе

snJajn Jo
yonpouob Aiepuoses

oes 1ешбело.а

adÁjoo
jo шпоэеэ

SIUILUSS
un¡noejdaoas

adÁjoo

yonp jeuıßenoud Jo saquieyo
и014э99/ U}IM pajeioosse ‘jesioq

Jaqueyo
uondaoal UM pajerdosse чедиэл

Jsqweyo
uondasas ou! Адоэлр биизиа oes

¡aoipad
элоае jsnf qua, Buipuaosap чо

ueBo-A yo do} чо

unjoal
0} ja¡¡esed 'yonpino jo реа jejsıq

_ эре}9э9э/ зиеб.о A1oss299e
Jaqueyo иоцаэээ/ ¡gunas Jaquueyo иоцаэээ/ Jaquieyo чоцетицАэ; - ¡esjanas BUIAIS981 JONPIAO JO UDILOG
Jaquieyo иоцаэээл
0} dn иоцц}зи09 разешбели
jsoıpad _I891ped ¡aoipad - - uo ueBo-A Jo uoiyod jeseg
qui, Buipusosap _[eoipad qui] ya] - qui] Buipuaose ¡aoipad oyuı Buipee] иебло-л Jo qui]
yonpiao Алешиа
qui, Buipusose qui, Buipusose qui зчбы ци 38114 qui, Buipuaosap Japuaj¡s jo чоцепициоэ j9aJ1g
JONPIAO JO зиеа uonIsod jeolwojyeuy
Ápn]s juasaJd (0861) NOSdWOHL (9261) v3vg (6061) d3nvg (1161) anunog

212

‘эершионэн Ul WEISÄS anonpoldas ajeway ay} jo ÁBojoua] ‘г FIGVL

CLASSIFICATION OF HELICINIDAE 219

V-organ
enlarged:
primary
oviduct

/__ hypobranchial
orifice

pallial
oviduct

opening near
mantle edge

intestine

A

descending

reception 7
chamber pallial $

\/-огдап - ascending limb

/ receptaculum seminis
$ pedicel of V-organ
| bursa copulatrix

provaginal
sac

E

oviduct |

provaginal duct
and vagina

FIG. 9. Female reproductive system in Helicinidae (Helicina orbiculata), ovary to slender portion of the
oviduct omitted, ventral view. A. Organs in natural position. B. Apical complex enlarged and artificially
arranged to show the different organs and their connections (modified after Baker, 1926).

ther confusion until these aspects have been
finally clarified. Furthermore, the term “pal-
lial” only refers to the topographical position
and not to ontogenetic origin.

The reproductive system of female
Helicinidae consists of a folliculous ovary
that discharges into a thin-walled spherical
structure, which continues as the slender
primary oviduct. This oviduct is curved
anteriad before it enters the V-organ. The V-
organ is subdivided in an ascending limb, a
descending limb, and a pedicel and leads
into the reception chamber. The oviduct con-
tinues as an elongated pallial part parallel to
the intestine and opens near the mantle

may receive the duct of a receptaculum
seminis, or is associated with sac-like struc-
tures (accessory sperm sac) at its very be-
ginning (e.g., Lucidella). Besides the oviduct
(pedicel), the reception chamber is con-
nected with a ventral bursa, a dorsal
provaginal sac, and a provaginal duct that
opens into the mantle cavity. Because of the
two openings, the female system is called
diaulic.

Because the general structure is similar in
Costa Rican Helicina, it is described as fol-
lows, and only specific deviations are added
under each species.

The ascending limb of the V-organ is straight

edge. The descending limb of the V-organ and a little longer than descending limb and

(Continued from opposite page)

‘ The caecum of the ootype and the provaginal sac sensu Bourne (1911) were differently interpreted by Baker. In 1925,
he assigned the organs correctly as given in my Table, whereas in 1926, he exchanged this assignment of the terms,
regarding his former interpretation as wrong. He pointed out that in Bourne's figure (1911: pl. XXXV, fig. 25) the caecum
would clearly be located dorsally (as only is the provaginal sac) with respect to the oviduct. Actually, he disregarded the
accurate description of the position of the organs. He was probably misled by the fact that in Alcadia palliata (С. В. Adams,
1849) the general appearance of this apical complex with both ventral bursa and provaginal sac elongated and without
lobes differs somewhat from the mainland species that he had dissected.

© Thompson misinterpreted the pedicel (term introduced by Baker, 1926) as the whole descending limb of the V-organ.
In Helicinidae, it is demarcated by an invaginated constriction and subsequent distal swelling before entering into the
reception chamber, also histologically differentiated. In the Ceresidae and Proserpinidae, which Thompson studied, the
descending limb and a pedicel are not externally demarcated, but since histological data are lacking, the identification of
the “descending” limb with the pedicel only (and absence of the “non-pedicel” part) is not verified for the two families.
Thompson intended to comply with other prosobranch terminology, but because both Latin and English terms were used
(e.g., bursa copulatrix and copulatory bursa) for the same organ, “seminal receptacle” would be synonymous with
“receptaculum seminis”, a term being already in use for an accessory structure. Furthermore, as far as it is known just the
reception chamber is not a place for sperm storage, i.e. a receptaculum seminis.

214

pedicel together, and т natural position it
approximately reaches the transition of the
reception chamber to the pallial oviduct.
Situated between the limbs of the V-organ, a
comparatively small, always simple sac-
shaped receptaculum seminis enters the
descending limb with a slender дис. The
ventral bursa copulatrix is always lobed, but
to a different extent. The provaginal sac 1$
well developed, but rather simple shaped,
and possesses a fairly long stalk as connec-
tion to the reception chamber. Contrary to
the general scheme given above and the
descriptions of other Central American spe-
cies of Helicina by Baker (1926), the
provaginal duct or vagina does not exist and
the system is monaulic. The thick walls of
the pallial oviduct are always variously
folded, which 1$ reflected in surface constric-
tions. A short, distinct portion just before the
distal opening is dark brownish, whereas the
remaining part of the reproductive system 1$
whitish-opaque if not otherwise stated.

If material was sufficient the reproductive
system of at least three females of each
species or population were dissected. п
addition, serial sections were studied for
Нейста funcki, H. tenuis, H. beatrix
confusa, H. beatrix riopejensis n. subsp., H.
gemma, Alcadia hojarasca, and Lucidella
lirata to confirm the results of the dissec-
tions.

In the drawings, the reproductive system is
normally shown from the ventral side and
the accessory organs of the apical part were
artificially separated to allow an adequate
presentation of this complex structure (Fig.
9B). If not otherwise stated, relative descrip-
tions refer to the structures of Helicina
funcki.

Morphometry and Sexual Dimorphism: Due to
the paucity of material, the number of speci-
mens of each population/species studied
could not be standardized, but, as far as
possible, maximized; the number of speci-
mens is indicated in each case. The follow-
ing measurements were analyzed: height,
minor diameter, height of last whorl and col-
umellar axis, extension of outer lip, volume
and weight, if available. The major diameter
is given only for comparison, but it is not in-
cluded in diagrams, because the shells in-
crease regularly in size, and it is therefore
correlated with the minor diameter, which
can be measured more exactly.

RICHLING

For the comparison with populations of un-
known sex (е.д., INBio material, type mate-
rial) the sex-independent mean value 1$
always indicated in the diagrams by shad-
ing, it is given as the average of the mean
values of both sexes. In this way, it more
closely approaches the theoretical 1:1 distri-
bution of females and males than the mean
value of the total population.
When relations of the shell size of the differ-
ent populations of one species to other pa-
rameters will be analyzed, the minor
diameter is preferred over the shell height,
because it is better correlated to the volume
- (shown for Helicina funcki, Fig. 30). The lat-
ter would display the size best, but the vol-
ume is normally not available for all
populations.

Habitat: The description of the habitat is nearly
exclusively based on the author’s own field
observations.

Distribution: In addition to the description, for
the distribution within Costa Rica a detailed
map is provided based on all records criti-
cally revised and the material studied. The
sources of the localities will be indicated di-
vided into recent collections (IR, INBio), and
literature records and the other material ex-
amined.

Discussion: Here, mainly the taxonomical
problems of each species will be discussed.
For broader aspects, see the general Dis-
cussion.

REVISION OF THE COSTA RICAN
HELICINIDAE

The following species are recognized for
Costa Rica:

Helicina (Tristramia) funcki L. Pfeiffer, 1849

Helicina (Tristramia) pitalensis Wagner, 1910

Helicina (Tristramia) tenuis L. Pfeiffer, 1849

Helicina (Tristramia) echandiensis n. sp.

Helicina (Tristramia) punctisulcata
cuericiensis n. subsp.

Helicina (“Gemma”) beatrix beatrix Angas,
1879

Helicina (“Gemma”) beatrix confusa (Wagner,
1908)

Helicina (“Gemma”) beatrix riopejensis п.
subsp.

CLASSIFICATION ОЕ HELICINIDAE 215

Helicina (“Gemma”) talamancensis (Richling,
2001)

Helicina (“Gemma”) gemma Preston, 1903

Helicina (“Gemma”) monteverdensis n. sp.

Helicina (“Gemma”) escondida n. sp.

Helicina (“Gemma”) chiquitica (Richling, 2001)

Pyrgodomus microdinus (Morelet, 1851)

Alcadia (Microalcadia) hojarasca (Richling,
2001)

Alcadia (Microalcadia) boeckeleri (Richling,
2001)

Lucidella (Perenna) lirata (L. Pfeiffer, 1847)

Questionable:

Helicina (Oligyra) flavida Menke, 1828

Helicina (Tristramia) funcki
L. Pfeiffer, 1849

Helicina funcki L. Pfeiffer, 1849: 121 (not fig-
ured)

Helicina funcki — L. Pfeiffer, 1850: 33, pl. 9,
figs. 1,2

Helicina funcki — L. Pfeiffer, 1852a: 361

Helicina tuncki [sic] — L. Pfeiffer, 1852b: 261-
262

Helicina funckii [sic] — Sowerby, 1866: 288, pl.
273, fig. 271

Helicina funcki — Bland, 1866: 9

Helicina funcki — Reeve, 1874: pl. 17, fig. 152

Helicina funki [sic] — Angas, 1879: 484, pl. XL,
fig. 7 (living animal): Costa Rica: Talamanca,
all the coast region, and to the lower hills
(Gabb)

Helicina funcki — von Martens, 1890: 33:
Costa Rica: Talamanca, all the coast region,
and to the lower hills (Gabb); Cache [Cachi?
09°50’°N, 83°48 W, Cartago Province]
(Rogers)

Helicina funcki var. a, b — Biolley, 1897: 4—5:
Costa Rica: San Miguel, Sarapiqui, 200 m
[about 10%19'N, 84°11'30"W, Alajuela Prov-
ince], Тиз, 600 т [about 09°51’N, 83°35’W,
Cartago Province]

Helicina funcki var. с, а — Biolley, 1897: 4-5:
Costa Rica: Azahar de Cartago, 1,500 m,
Tarbaca, 1,600 m [09°49’25"М, 84°06’39"W,
San José Province]

Helicina funcki — Ancey, 1897: 87: E-Nicara-
gua: Greytown, N-Panama: Monkey Hill,
near Colon (leg. Aillaud)

Helicina funcki — von Martens, 1900: 603-604:
E-Nicaragua: Greytown; NE-Costa Rica:
San Miguel, valley of the Sarapiqui, 200 m
[about 10°19’N, 84°11’30"W, Alajuela Prov-
ince]; Puerto Viejo [about 10°28'N,
84°00'30"W, Heredia Province] (Biolley), on

the borders of the Río San Juan [along bor-
derline to Nicaragua in Alajuela, Heredia,
Limón provinces, cannot be specified]
(Pittier), E-Costa Rica: Tuis, 600 m [about
09°51'N, 83°35’W, Cartago Province]
(Biolley, Pittier); Turrialba, 750 m [about
09°54’30"N, 83°41’W, Cartago Province]
(Biolley), central Costa Rica: Azahar de
Cartago [not clear, if referring to the town
Cartago, ?about 09°52’N, 83°55’W, Cartago
Province] and Tarbaca, 1,500-1,600 m
[09°49'25"N, 84°06’39"W, San José Prov-
ince], only the smaller varieties (Biolley); N-
Panama: Monkey Hill, near Colon [in part]

Helicina (Retorquata) funcki — Wagner, 1905:
232-233

Helicina (Retorquata) funcki costaricensis
Wagner, 1905: 233, pl. XIII, fig. 12 а-с:
Costa Rica (“von San José [14 km NW of
Upala, about 10%58'N, 85°08'W, Alajuela
Province] in Costarica besitze ich
Exemplare dieser Form, welche größer und
einfarbig weiß sind, ferner % bis % Umgang
mehr aufweisen”)

Нейста fucki [sic] - Wagner, 1910a: 306307,
pl. 61, figs. 11-15: Neu Granada (obviously
only in part of Panama), Costa Rica:
Azachar Centajo, Tarbaca

Нейста funcki costaricensis — Wagner,
1910a: 307, pl. 61, fig. 16: Costa Rica: St.
Jose [see above] and Sta. Clara [7.5 km NW
of Upala, about 10°56’М, 85°05’W, Alajuela
Province]; “eine ähnliche Form, jedoch mit
deutlicher Kante am letzten Umgang und
höherem Gewinde liegt in meiner Sammlung
mit der Fundartsangabe Ylalag in Mexico”

Helicina funcki — Pilsbry, 1910: 503: Panama:
Canal Zone: Tabernillo (Brown)

Helicina funcki — Pilsbry, 1920a: 3: Costa Rica:
Guapiles, 980 ft. [about 10°14’N, 83°47’W,
Limön Province] (Calvert)

Helicina deppeana parvidens Pilsbry, 1920a: 3
(not figured): Costa Rica: Juan Viñas, farther
waterfall, 3300 ft., also on the road to Rio
Reventazon, 3000 ft. [about 09°54’N,
83°44'30"W, Cartago Province] (Calvert)

Helicina (Tristramia) funcki funcki — Baker,
1922a: 51

Helicina (Tristramia) funcki parvidens — Baker,
1922а: 51

Helicina (Tristramia) funcki costaricensis —
Baker, 1922a: 51

Helicina funcki — Pilsbry, 1926a: 59, 69, 71,
fig. 3C: Panama: Escobal on Gatun Lake
(Chapin), Bocas del Toro Province: Mono
Creek (Olsson), Gatun (Harrower), Canal

216 RICHLING

Zone: Barro Colorado Island and near
Darien (Zetek)

Helicina funcki — Pilsbry, 1926b: 127: Costa
Rica: Talamanca Valley, < 100 ft. [approxi-
mately 09°34'N, 83°W, not specified, Limon
Province] (Olsson)

Helicina (Tristramia) funcki — Baker, 1926: 42:
Panama: Gatun, Canal Zone (Harrower), pl.
V fig. 8, pl. VI, fig. 9 (female and male repro-
ductive system)

Helicina funcki — Pérez, 1994: 746: Costa
Rica: La Selva [about 10°26’N, 84°W,
Heredia Province]

Helicina funki [sic] — Monge-Najera, 1997:
113: Costa Rica

Helicina funcki — Robinson, 1999: 434: USA:
sometimes mistakenly imported

Synonymy

Helicina funcki costaricensis Wagner, 1905
Helicina deppeana parvidens Pilsbry, 1920

Original Description

“Hel. testa conico-subglobosa, tenuiuscula,
sub lente tenuissime oblique striatula, vix
nitidula, flavida, roseo-nebulosa; spira
conoidea, obtusiuscula; anfractibus 5,5
planiusculis, ultimo utrinque convexiore, obso-
lete angulato; apertura obliqua, semiovali;
columella subarcuata, linea impressa verticali
notata, basi subnodosa, in callum sensim
tenuiorem retrorsum abiente; peristomate late
expanso, margine supero subrepando.
Diam. 13,5, altit. 9 mill.

From San Yago, New Granada (Funck)

”

Type Material

BMNH 20010497.1-4: Santiago, New
Granada, Funk, H. Cuming collection

The type lot contains four similar specimens.
The shell that 15 slightly larger than the other
three is herein selected as lectotype of
Helicina funcki (Fig. 10). It shows the traces of
some lead pencil painting which could have
been applied to the specimen as a drawing
aid, probably reflected in the dark shading vis-
ible in the figure in L. Pfeiffer (1850: pl. 9, figs.
1, 2). Furthermore, it is the only specimen that
attains 13.5 mm in its greatest extension (not
perpendicular to the shell axis). The height
given in the original description cannot be at-
tributed to a conventional adjustment of the
shell. The specimen is yellowish, and the red-
dish tinge is barely visible, whereas it is well
developed in the three paralectotypes in the
second half of the body whorl between suture
and the periphery.
Dimensions:
Lectotype BMNH 20010497.1:
10.6/11.9/13.2/10.7/7.9/8.8/8.2 mm
Paralectotypes BMNH 20010497 .2—4:
10.1/11.5/12.6/10.2/7.3/8.3/7.7 mm
10.0/11.0/12.4/10.0/7.1/8.2/7.6 mm
10.0/11.1/12.4/10.1/7.4/8.2/7.9 mm

Type Locality

“San Yago, New Granada’, this most prob-
ably refers to Santiago, which today belongs
to Panama, Veraguas Province.

Type Material of Synonymous Taxa or Similar
Species

Helicina funcki costaricensis Wagner, 1905

Type Material: MIZ 8989: Costa Rica, Sta.
Clara, 250 m alt., Biolley legit
In the original description Wagner gives Costa
Rica as the origin of the new subspecies and
mentions additional specimens from San

FIG. 10. Нейста funcki, lectotype, BMNH 20010497.1, height 10.6 mm; scale bar 5 mm.

CLASSIFICATION ОЕ HELICINIDAE 217

José, which are said to be of greater size and
of uniformly white color, the description being
obviously based on further unspecified mate-
rial. In his collection stored in the MIZ, there
are two lots: the typical one with the locality
mentioned later (Wagner, 1910a), as given
above and the one from San José. The
syntypes MIZ 8989 consist of two specimens,
a yellowish-greenish one and a reddish tinged
one. In comparing the figures in Wagner
(1905, 1910a), it is obvious that different speci-
mens were illustrated: the later figure shows a
yellowish-greenish specimen somewhat more
elevated and with a more strongly developed
denticle at the transition of the outer lip into the
columella. Thus, the reddish specimen was
first to be depicted and has therefore been
selected here as lectotype (Fig. 11). It dis-
plays a slight crack in the last whorl which,
however, did not result in any deformation or
damage in the shell. The specimen was dead
collected, whereas the paralectotype was col-
lected alive, complete with its operculum.
Dimensions:

Lectotype MIZ 8989a:
12.0/13.2/14.6/11.8/8.7/9.7/9.3 mm
Paralectotype М! 8989b:
12.3/13.1/14.6/11.9/8.9/10.1/9.5 mm

Type Locality: “Costa Rica”; restricted by type
selection to Sta. Clara, 250 m a.s.l. [7.5 km
NW of Upala, about 10°56’N, 85°05'W,
Alajuela Province]

“Santa Clara” is a name used for various lo-
calities in Costa Rica. Biolley mentions it
several times as a collecting site, also
“Delicias near Santa Clara” and a “San
José” (see “Discussion” for Helicina funcki

costaricensis) that is definitively not the capi-
tal. This combination suggests the identifica-
tion with the village of Santa Clara near
Upala, because Las Delicias and San José
are nearby. The exact altitude of Santa Clara
is 40 m, but it is known that in former times
(personal communication with Zaidett
Barrientos) the whole region was called
“Llanuras de Santa Clara” [plains of ...].
Therefore, in case it was not just an inaccu-
rate measurement of the altitude, it is likely
that the specimens were collected a little to
the southeast of the village approaching the
Cordillera de Guanacaste.

Helicina deppeana parvidens Pilsbry, 1920

Type Material: Holotype ANSP 105286 (Fig. 12),
Paratype ANSP 105252 (original designation)

Type Locality: Costa Rica: “Juan Viñas, farther
waterfall, 3300 ft.” [about 09°54’N,
83*44'30"W, Cartago Province]

Examined Material

Lee. |. RICHLING

Guanacaste: N Santa Elena: Reserva Sta.
Elena, Sendero Rio Negro, about
10°20’31"М, 84%47'53"W, 1,550 т a.s.l.:
14.08.1999: (IR 924); Sendero at Mirador
Gerardo, 10°22’19"М, 84*48'25"W, 1,450 т
a.s.l.: 14.08.1999: (IR 928); 19.02.2000: (IR
1230)
N of Nuevo Arenal: area of primary rain for-
est, 10°33’32"N, 84*51'40"W, 800 m a.s.l.:
05.03.1999: (IR 737); “Las Pavas” (private
reserve in preparation), secondary rain for-

FIGS. 11-12. Helicina spp. FIG. 11. Helicina funcki costaricensis, lectotype, MIZ 8989a, height 12.0
mm; scale bar 5 mm. FIG. 12. Helicina deppeana parvidens, holotype, ANSP 105286, height 10.0 mm;
scale bar 5 mm (photograph: D. Robinson).

218

est, about 10°33’30"М, 84*51'53"W, 800 m
a.s.l., to 10°33’26"М, 84°51'57"W, 760 т
a.s./.: 05.03.1999: (IR 742); 17.08.1999: (IR
952): 24.02.2000: (IR 1273); (В 1331);
03.2001: (IR 1637)

NW Nuevo Arenal, “Eco Lodge”, Sendero
Cabana, about 10°34’37"N, 84°55’35"W, 750
m a.s.l., 18.08.1999: (IR 955)

Parque Nacional Rincón de Vieja: trail from
Aquas calientes to Las Pailas, about
10°46'00"N, 85°19'13"W, 800 m a.s.l.:
20.08.1999: (IR 972); E Casona Sta. Maria,
trail to Canal, 10%45'57"N, 85°17’06"W, 750
т a.s.l.: 21.08.1999: (IR 979)

Alajuela: Near Volcán Arenal, trail along vol-
cano in rainforest, about 10°29’07"М,
84°42’55"W, 720 т a.s.l.: 24.02.1998: (IR
390); 01.08.1999: (IR 884); on Heliconiaceae,
25.02.2000: (IR 1286)

Heredia: S Puerto Viejo de Sarapiqui, Zona
Protectora La Selva, near OTS-Station, about
10%25'53"N, 84°00'18"W, 60 т а.5.1.,
05.09.1999: (IR 1061); 06.09.1999: (IR
1062); 13.02.2000: (IR 1182); (IR 1184)
Limón: Parque Nacional Cahuita, trail from
Cahuita to Puerto Vargas, coastal forest with
coco palms and swampy areas: 09°44’01"N,
82°49'48"W, 1-5 т a.s.l.: 11.03.1997: (IR
107); (IR 106); (IR 108); about 09°43'27"N,
82°50'28"W, 4 m a.s.l.: 10.03.1999: (IR 757);
07.08.1999: (IR 897); 08.08.1999: (IR 898);
04.03.2000: (IR 1312); 14.03.2001: (IR
1555), (IR 1630), (IR 1639); 15.03.2001: (IR
1557), (IR 1648); 09°43’13"М, 82%50'39"W, 4
т a.s.l.: 13.09.1999: (IR 1095); near Puerto
Vargas, 09°42’49"N, 82°49’20"W, 1 т a.s.l.:
08.08.1999: (IR 907)

Refugio Nacional de Fauna Silvestre
Gandoca-Manzanillo, S Manzanillo, trail
along coast line to S, coastal forest, about
09”38'06"N, 82°38’26"М/, 50 m а.5.1.,
14.9.1999: (IR 1096); 5.3.2000: (IR 1320); (IR
1322); (IR 1642)

Near Cruce Penshurt, mouth delta of Rio
Estrella, Aviarios del Caribe, about
09°48’30"М, 82°54W, 20 m asll.,
09.08.1999: (IR 912)

About 9 km W of Matina, road Limón to
Siquirres, a little stream up Río Barbilla, small
banana plantation, about 10°03’29"М,
83°22'24"W, 70 m a.s.l., 12.03.2001: (IR 1545)
S Siquirres, road Limón to Siquirres, along
footpath stream up Rio Pacuarito,
10 05.38 83228 11:W. Мои аз.
18.03.2001: (IR 1612); (IR 1613)

Siquirres, along footpath stream up Rio
Siquirres and along a southern tributary,

RICHLING

10°05'37°N;183/30 92 W100: Mas:
11.03.2001: (IR 1623); 19.03.2001: (IR 1617)
W Guayacän, abandoned banana plantation,
10*01:53'N,, 83°32"4"W,. 520) ть as”
03:09:1999:, (IR 1079); ав: 1090).
12.09.1999: (IR 1090): 17.03.2001: (IR 1608)
И/ Liverpool, Mexico: near Rio Blanco,
0956:37"N, 83°09'41"М 35 та!
13.03.1997: (IR 150); near Rio Blanco,
abandoned area with bananas and some old
trees, 09°58’32"N, 83°08’32"W, 35 m a.s.l.:
14.02.1998: (IR 274): 21.02.1999: (IR 627);
footpath along small creek and through
bush, 09°59’04"М, 83%08'04"W, 40 m a.s.l.:
16.02.2000: (IR 1191): 22.02200E2 IR
1406); (IR 1408)

S Liverpool: near Rio Rene, swampy area
and forest, 09°57’33"М, 83°08'15"W, 20 т
a.s.l.: 13.03.1997: (IR 148); (IR 140); along
Rio Victoria, 09°56'01"N, 83°10’24"W, 80 т
a.S./.: 05.03.1998: (IR 465)

SW Liverpool: Rio Quito, at bridge,
09°57’11"N, 83°1037"W 40 т a.s.l.:
04.03.1998: (IR 448); E of Río Peje, forest
on little hill, 09°57’46"N, 83°13’26"W, 270 т
a.S./.: 12.03.1997: (IR 131); Río Peje and
small tributary, 09*56'35"N, 83*14'01"W, 110
m a.s.l.: 12.03.1997: (IR 126); (IR 127): Во
Peje, bordering forest, 09°56’23"М,
83°14’06"W, 160 m a.s.l.: 09.03.1999: (IR
753); along Rio Peje, bordering forest with
palms, 09°55’46"N,83°13'15"W, 135 m a.s.l.:
04.03.1998: (IR 441); 09.03.1999: (IR 751);
03.03.2000: (IR 1300); (IR 1502);
13.03.2001: (IR.1552)

N Shiroles: along Quebrada Кто,
09°35’38"N, 82°57’20"W 100 m asl:
12.03.1999: (IR 763); 09.08.1999: (15390:
16.03.2001: (IR 1596); (IR 1644); Сето
Mirador, along trail, 09°36’37"М, 82°57’43"W,
430 m a.s.l.: 16.03.2001: (IR 1599)

W Bribri, road to Uatsi, about 09°38’11"М,
82°51 48"W, 30 т a.s.l.: abandoned field
with Heliconiaceae and Eucalyptus:
12.03.1999: (IR 766); 15.09.1999: (IR 1114);
wooded valley within banana plantation, 50
m:a.s.l.: 15.3.2001: (IR 1572); at crossing
with Río Carbón, 30 m a.s.l.: 17.3.1997: (IR
183);

И/ Uatsi, along Кю Ча, 09°37’30"М,
82°53’30"W, 60) т а.$1., 15.03.2008
1632)

Zona Protectora Tortuguero, near
Tortuguero, about 10°34’N, 83°31 W, 10 т
a.s.l.: Sendero Ranita: 10.3.2000: (IR 1348);
N of village: 16.03.2001: (IR 1620);
21.03.2001: (IR 1653)

CLASSIFICATION OF HELICINIDAE 219

Cartago: W Turrialba, near Catie, forest along
road Turrialba to Siquirres, 09°53’01"М,
83"39'17"W, 610 m a.s:l.. 15:03.2000: (IR
1350)

Puntarenas: Near Monteverde, about
10°17’24"М, 84°48'04"W: small piece of for-
est along road to reserve, 1,330 m a.s.l.:
27.02.1997: (IR 22); 1 km before entrance
on road to reserve, 1,500 т a.s.l.:
26.07.1999: (IR 826); 1999: (IR 1391);
13.8.1999: (IR 927); Bosque de los Niños,
10%17'59"N, 84°48'44"W, 1,380 т a.s.l.:
29.07.1999: (IR 860)

Zona Protectora Arenal-Monteverde:
Reserva Biológica Bosque Nuboso
Monteverde (about 10°18'08"N, 84°47’41"W,
1,500-1,650 m a.s.l.): 27.07.1999: (IR 843);
18.02.2000: (IR 1194); (IR 1199); (IR 1627):
Sendero Bosque Nuboso: 25.02.1997: (IR
14); 24.02.1999: (IR 628); Sendero Roble:
18.02.1998: (IR 301); Sendero Chomogo:
25.02.2001: (IR 1435)

Monteverde, Cerro Plano, Finca Ecologica,
Sendero Mirador, 10°18'47"N, 84°49’30"W,
1,330 Im а 25.02.1999: (В 651);
28.07.1999: (IR 859); 15.08.1999: (IR 946);
20.02.2000: (IR 1246)

About 4 km N Santa Elena, Skywalk,
10°18'33"N, 84°4942"W, 1,330 m a.s.l.,
20.02.1998: (IR 332)

INBio COLLECTION

Guanacaste: Zona Protectora Arenal-
Monteverde: Santa Elena, sendero Encantado,
10%21'57"N, 84°47'27"W, 1,200 т a.s.l., leg.
Kattia Martinez, 21.06.1996: 8 ads. (INBio
1498638)

Zona Protectora Tenorio: Río San Lorenzo,
Tierras Morenas, 10°36’38"М, 84°59'42"W,
1,050 m a.s.l., leg. Gladys Rodriguez,
28.10.1995: 1 ad. (INBio 1485411); Tenorio,
Alrededores de la estación, 10°36’51"М,
85”00'07"W, 900 m a.s.l., leg. Gladys
Rodriguez, 18.09.1996: 1 ad. (INBio
1498593)

Parque Nacional Rincón de la Vieja: Sector
Las Pailas: 4.5 km SW del Volcán Rincón de
la Vieja, 10%46'36"N, 8521'07"W, 800 т
a.s.l., leg. malacological staff of INBio,
09.12.1992: 1 ad. (INBio 1466644); sendero
Pailas, 10°46'36"N, 85°21'07"W, 800 m
a.s.l., leg. Karla Taylor, 23.08.1995: 3 ads.
1 s.ad. (INBio 1498739); Sector Santa
Maria: 10°45’58"М, 85°18'19"W, 800 m
a.s.l., leg. Dunia Garcia, 14.10.1995: 1 ad.
(INBio 1487945); sendero Bosque
Encantado, 10°46’36"N, 85°21'07"W, 800 т

a.s.l., leg. Karla Taylor, 23.08.1995: 1 ad.
(INBio 1498744)

Parque Nacional Guanacaste: Estación
Góngora, 10°53’22"М, 85°28’33"М/, 580 т
a.s.l.: leg. Zaidett Barrientos, 11.02.1994: 2
ads. (INBio 1480300); leg. Dunia Garcia,
20.10.1994: 1 juv. (INBio 1478682); 2 juvs.
(INBio 1478739); 2 ads. (INBio 1483409);
leg. Dunia Garcia, 28.10.1994: 1 ad. (INBio
1480475); leg. Dunia Garcia, 08.03.1995: 1
ad. (INBio 1488083); leg. Dunia Garcia,
28.06.1995: 1 ad., 3 juvs. (INBio 1484993);
Sector Góngora [rio Góngora], 10°53’22"М,
85°28'33"W, 580 m a.s.l.: leg. Kattia
Martinez, 26.05.1995: 1 juv. (INBio 1498514)
Parque Nacional Guanacaste: Sector Orosi
(antes: Maritza), sendero Casa Fram,
10%57'40"N, 8529'45"W, 600 m a.s.l., leg.
Zaidett Barrientos, 15.07.1996: 3 ads. (INBio
1494681); Río Tempisquito, 10%57'45"N,
85”29'05"W, 600 m a.s.l., leg. Dunia Garcia,
08.03.1996: 1 juv. (INBio 1488078)
Parque Nacional Guanacaste: La Cruz, 9
km S de Santa Cecilia, Estación Pitilla:
10°59’25"М, 85°25’38"W, 700 т a.s.l.: leg.
Petrona Rios, 09.12.1994: 1 ad. (INBio
1480289); Lado S del Río Orosí, leg. Calixto
Moraga, 16.08.1994: 3 ads. (INBio
1480319); leg. Calixto Moraga, 23.08.1994:
1 аа. (INBio 1480318); 10°59’33"М,
85°25'46"W, 700 т a.s.l.: leg. malacological
staff of INBio, 08.01.1993: 1 ad. (INBio
1463787); leg. Calixto Moraga, 10.07.1993:
1 juv. (INBio 1467560); Sendero Nacho,
10°59'33"N, 85°25’46"W, 700 m a.s.l.: leg.
malacological staff of INBio, 13.10.1993: 1
ad. (INBio 1463946); Sendero Mena, 400 m
W de la Estación Pitilla, 10°59’25"М,
85°25'51"W, 700 m a.s.l.: leg. Calixto
Moraga, 09.01.1994: 1 ad. (INBio 1480043);
Fila Orosilito, 10°59'02"N, 85°26 01"W, 900
т a.s.l.: leg. Calixto Moraga, 20.04.1994: 1
ad. (INBio 1480329); Finca del Estado:
Casa de Roberto, 11°00’09"N, 85*25'33"W,
600 m a.s!l: leg. Calixto Moraga,
22.08.1994: 1 s.ad. (INBio 1480342)

La Esperanza, 6 km E de Santa Cecilia de la
Cruz, 11°00’42"N, 85°22’45"W, 400 m a.s.l.,
leg. Calixto Moraga, 09.01.1994: 1 ad.
(INBio 1480050)

Alajuela: Reserva Biológica Los Angeles, 7

km NE de los Angeles Norte de San Ramón,
10°12’12"М, 84°29'10"W, 1,100 m a.s.l., leg.
Zaidett Barrientos, 06.11.1995: 1 ad., 2
s.ads. (INBio 1482570)

Reserva Biológica San Ramón, 10°13’30"N,
84°35'17"W, 800 т a.s.l.: leg. malacological

220

staff of INBio, 15.02.1994: 1 аа. (INBio
1477816); leg. Gerardo Carballo,
10.07.1994: 1 ad. (INBio 1476150); Sendero
Liz, leg. Gerardo Carballo, 08.08.1994: 1 ad.
(INBio 1476214)

Sector Colonia Palmareña, 10°14’09"N,
84°33'15"W, 700 m a.s.l., leg. Еда Fletes,
13.04.1995: 1 ad. (INBio 1485385)

Zona Protectora Arenal-Monteverde: Sector
Alemán, Finca dos Ases, 10°17’56"N,
84°46'08"W, 1,140 т a.s.l.: leg. Zaidett
Barrientos, 13.10.1994: 1 ad. (INBio
1468276); leg. Kattia Martinez, 04.12.1995: 2
ads. (INBio 1485227); Sendero Alemán,
10°17'59"N, 84°45'38"W, 1,080 т a.s.l.: leg.
Kattia Martinez, 18.08.1994: 1 ad. (INBio
1480101); 19.08.1994: 1 ad. (INBio
1478523); Sector Peñas Blancas, Estación
Alemán, 10°18'09"N, 84°44’52"W, 900 m
a.s.l.: leg. Каша Martinez, 11.10.1994: 2
ads. (INBio 1498802); 11.12.1994: 1 ad.
(INBio 1480605)

Parque Nacional Guanacaste-Rincön de la
Vieja, Estación San Cristóbal, 10°52’55"М,
85°23'26"W, 600 m a.s.l.: leg. Dunia Garcia,
08.01.1995: 5 ads., 2 s.ads. (INBio 1488065);
leg. malacological staff of INBio, 18.08.1995:
8 ads., 5 s.ads. (INBio 1498494)

Sector las Cubas, Bosque Urbina,
10%53'41"N, 84°47"20"W, 40 m a.s.l., leg.
Kattia Martinez, 25.04.1994: 1 ad. (INBio
1466940)

Caño Negro: Veracruz, 10°50’22"М,
84°52’52"W, 35 m a.s.l.: leg. Каша Flores,
14.02.1997: 1 juv. (INBio 1487125); Finca
Delicias, 10°54’01"N, 84°47’20"W, 35 т
a.s.l.: leg. Каша Flores, 14.12.1996: 1 ad.
(INBio 1487043); 01.11.1997: 1 ad. (INBio
1487611); en el Pueblo, 10°53’38"N,
84*47'20"W, 35 т a.s.l.: leg. Каша Flores,
09.10.1994: 1 ad. (INBio 1480029);
07.04.1995: 5 ads. (INBio 1501040)
Refugio Nacional de Vida Silvestre Caño
Negro: Caño Negro, San Antonio, Finca
Juan Cubano 2, 10°54’50"N, 84%45'12"W,
35 m a.s.l., leg. Каша Flores, 16.11.1996: 1
ad. (INBio 1487878)

Monte Cele, sendero La Tepezcuintle,
10°57'27"N, 85°24’20"\М, 700 m a.s.l., leg.
Dunia Garcia, 09.09.1995: 4 ads. (INBio
1488042)

Estación Playuelas, 50 m del Río Frío,
10°57'29"N, 84°44’55"W, 40 m a.s.l., leg.
Kattia Martinez, 08.01.1994: 4 ads. (INBio
1479506)

Sector Playuelas, 10°57’29"N, 84°45’15"W,
35 т a.s.l.: leg. Каша Martinez, 21.08.1996:

RICHLING

2 ads. (INBio 1498571); leg. Kattia Flores,
08.11.1996: 1 ad. (INBio 1487809)

Heredia: Frente al bosque de la hoja,

10°04'13"N, 84°05’40"W, 1,800 m a.s.l., leg.
Zaidett Barrientos, 14.05.2000: 1 ad. (INBio
3562231)

Limón: Reserva Indígena Talamanca: Sector

Amubri, 09*30'53"N, 82*57'19"W, 70 m
a.s.l.: 14.06.1994: 1 ad. (INBio 1477585);
15.06.1994: 1 ad. (INBio 1477569);
26.09.1994: 1 ad. (INBio 1483302); 4 s.ads.
(INBio 1483303); 1 juv. (INBio 1483376); 1
ad. (INBio 1483382); 1 s.ad. (INBio
1483388); 2 ads. (INBio 1483392); 1 ad.
(INBio 1483407); 2 s.ads. (INBio 1483408);
27.09.1994: 1 ad. (INBio 1483381); 1 ad.
(INBio 1483389); 2 s.ads. (INBio 1483402);
29.09.1994: 2 s.ads. (INBio 1483403); 1 ad.
(INBio 1483394): 30.09.1994: 2 s.ads.
(INBio 1483395); 2 s.ads. (INBio 1483401);
18.10.1994: 1 ad. (INBio 1483390); 1 ad.
(INBio 1483386); 19.10.1994: 1 s.ad. (INBio
1483383); 1 ad. (INBio 1483387); 1 s.ad.
(INBio 1483396); 27.11.1994: 1 ad. (INBio
1483398); 28.11.1994: 1 s.ad. (INBio
1483378); 1 s.ad. (INBio 1483385);
29.11.1994: 1 sad. (INBio 1483397);
30.11.1994: 1 ad. (INBio 1483400) (all leg.
Gerardina Gallardo); Amubri, Sendero Soki,
09°30'53"N, 82°57'19"W, 70 m.a.s.l.: "leg!
Angela Mora Maroto, 17.04.1995: 1 ad.
(INBio 1484735); leg. Gerardina Gallardo,
17.05.1994: 4 ads. (INBio 1467294); 4 ads.
(INBio 3395382); leg. Angela Mora Maroto,
22.04.1995: 3 ads. (INBio 1485382); leg.
Angela Mora Maroto, 04.08.1995: 1 ad.
(INBio 1485365); leg. Gerardina Gallardo,
27.11.1996: 3 ads., 1 s.ad. (INBio 1493444)
Reserva Indígena Talamanca: Cerca Río
Lan, 09*32'57"N, 82°58’25"W, 80 mars
leg. Gerardo Carballo, 17.06.1994: 1 ad.
(INBio 1476073); Suirí, orillas del Rio Telire,
09°33’56"М, 82*55'50"W, 30 m a.s.l.: leg.
Gerardina Gallardo, 25.11.1996: 1 ad., 2
s.ads. (INBio 1487336)

Reserva Biológica Hitoy Cerere: Sector
Miramar: 09°38’ОЗ"М, 83°00’45"W, 300 т
a.s.l.: leg. Zaidett Barrientos, 08.10.1994: 1
ad. (INBio 1475720); 1 ad. (INBio 1475725);
Senderos а Rio Мот, 09°37’44"N,
83°00’32"W, 150 m a.s.l.: leg. Zaidett
Barrientos, 08.11.1994: 2 juvs. (INBio
1475228); 1 ad. (INBio 1475234); Hitoy
Cerere, 09°37’50"М, 83°00’52"W, 300 т
a.s.l.: leg. Gerardo Carballo, 12.05.1994: 3
ads., 1 s.ad. (INBio 1476376); leg. Gerardo
Carballo, 13.06.1994: 4 ads. (INBio

CLASSIFICATION ОЕ HELICINIDAE 221

1476490); leg. Gerardo Carballo,
04.07.1994: 2 ads. (INBio 1475694); leg.
Marianella Segura, 07.12.1994: 1 ad. (INBio
1480272); Sendero Moin, 09°37'50"N,
83°00'52"W, 300 m a.s.l.: 14.01.1994: 3 ads.
(INBio 1475930); 27.02.1994: 1 ad. (INBio
1476687); 1 ad. (INBio 1476688) (all leg.
Gerardo Carballo)

Reserva Biológica Hitoy Cerere: Cruce entre
Sendero Revienta Pechos y Sendero
Espavel, 09°39’12"М, 83%00'58"W, 600 т
a.s.l.: leg. Alexander Alvarado Mendez,
24.04.1999: 1 ad. (INBio 1497851); Sector
Hitoy Cerere, Sendero Catarata,
09°40'18"N, 83°01’45"W, 100 т a.s.l.: leg.
Gerardo Carballo, 22.02.1994: 1 ad. (INBio
1476262); Sendero Tepezcuintle,
09°40’22"М, 83%01'40"W, 140 m a.s.l.:
25.04.1999: 2 ads. (INBio 1497862); 2 ads.
(INBio 3090624); 05.07.1999: 1 ad. (all leg.
Alexander Alvarado Mendez) (INBio
1497905); Sendero Bobócara, 09°40’31"N,
83°00'31"W, 200 m a.s.l.: leg. malacological
staff of INBio, 10.01.1993: 1 ad. (INBio
1466444); Sendero Toma de Agua,
09°40’31"М, 83°01 36"W, 100 m a.s.l.:
20.04.1994, leg. Zaidett Barrientos: 2 ads.
(INBio 1473832); leg. Gerardo Carballo: 1
ad. (INBio 1476246); leg. Zaidett Barrientos,
08.09.1994: 1 ad. (INBio 1475438); Estación
Hitoy Cerere, 09*40'35"N, 83°01' 36"W, 100
т a.s.l.: leg. malacological staff of INBio,
15.11.1993: 3 ads. (INBio 1463392); 400 т
NE de la Estación de Hitoy Cerere, Sendero
la “Finca”, 09°40'35"N, 83%01'26"W, 110 т
a.s.l.: 03.06.2000: 1 ad. (INBio 3098418);
20.07.1999: 1 $.аа. (INBio 1497844);
27.09.2000: 2 ads. (INBio 3091789) (all leg.
Alexander Alvarado Mendez); Sendero
Chato: 09*40'41"N, 83°01’26"W, 100 т
а.$.1., leg. Marianella Segura, 14.07.1994: 1
s.ad. (INBio 1478197)

Refugio Nacional de Vida Silvestre Gandoca-
Manzanillo: Sector Gandoca, Camino a
Gandoca, 09°38'04"N, 82*38'37"W, 10 т
a.s./.: 28.04.1999: 5 juvs. (INBio 3097941);
Sector Manzanillo: 1 km S de la escuela,
99237531"М;. 92°39:36\№ 4 т аз,
02.02.2000: 2 ads. (INBio 3097906):
Camino a Gandoca, 09°38’13"М,
82°38'40"W, 100 т a.s.l., 28.01.2000: 2 ads.
(INBio 3097895); Sendero a Gandoca,
09°38'04"N, 82°38'43"W, 8 m as,
04.02.2000: 2 s.ads. (INBio 3097899) (all
leg. Alexander Alvarado Mendez)

1 km $ de Рита Cocles, 09°38’17"М,
82"43'25"W, 40 m a.s.l., leg. Zaidett

Barrientos, 20.08.1996: 1 ad., 1 juv. (INBio
1487850)

Parque Nacional Cahuita: Sector Cahuita:
800 т E de la Casetilla, 09°44’00"N,
82°49'57"W, 10 m a.s.l., 05.11.1999: 1 s.ad.,
1 juv. (INBio 3096430); Sector Puerto
Vargas: Sendero a Cahuita, 09°43’43"N,
82°49'11"W, 0 та.5.1., 01.09.1999: 3 ads.
(INBio 3095846); 600 m E de la Casetilla,
09°42’54"М, 82°48’58"W, 8 т asi.
27.09.2000: 1 juv. (INBio 3091796) (all leg.
Alexander Alvarado Mendez)

Isla Uvita, frente al muelle de Limón,
09°59'45"М, 83*00'50"W, 5 m a.s.l., leg.
Alexander Alvarado Mendez, 11.10.2000: 2
ads., 1 juv. (INBio 3315386)

Zona Protectora Río Pacuare: 1.3 km NW
de la Estación Barbilla, 09°59’25"М,
83°28'04"W, 500 m a.s.l., leg. Alexander
Alvarado Mendez, 02.11.2000: 1 ad. (INBio

3315302)
Reserva Indígena Barbilla-Dantas: Sector
Colonia Puriscaleña, 10°00'17"N,

83°23'02"W, 300 m a.s.l., leg. Alexander
Alvarado Mendez, 03.03.2000: 2 juvs.
(INBio 3098016)

Sector Guápiles: 10°11'51"N, 83°51'22"W,
300 m a.s.l., leg. Alexander Alvarado
Mendez, 08.03.2000: 2 ads., 1 s.ad. (INBio
3097950)

Orillas del rio Aguas Frias, 10°24’05"М,
83°35'60"W, 10 т a.s.l.: leg. Elias Rojas,
29.11.1996: 3 ads. (INBio 1487980)

Finca Montaña Grande, 10°31’39"М,
83°43'33"W, 10 т a.s.l.: 400 т N de la
estación, a orillas de la quebrada:
13.09.1993: 3 ads. (INBio 1498610); 300 m
N de la estación: 21.09.1996: 1 ad. (INBio
1501097); 600 т N de la estación Cedrales:
13.11,1996: 2 juvs: (INBio 1501055);
14.12.1996: 1 juv. (INBio 1498623) (all leg.
Elias Rojas)

Finca Toty Castro, 1.7 km S de la estación
Cedrales, 10%31'39"N, 83°43’33"W, 10 m
a.s.l., leg. Elias Rojas, 16.10.1996: 1 ad.
(INBio 1501098)

Refugio Nacional de Vida Silvestre Barra del
Colorado: Pococí, Colorado, Sector Cerro
Cocorí, 30 km N de Cariari, 10°35’39"N,
83°42’59"W, 160 m a.s.l.: leg. malacological
staff of INBio, 10.12.1993: 6 ads. (INBio
1465446); leg. malacological staff of INBio,
04.10.1994: 3 ads. (INBio 1478061); leg.
Elias Rojas, 10.05.1994: 5 ads. (INBio
1483360); leg. Elias Rojas, 24.08.1994: 1
ad., 1 s.ad. (INBio 1480255); leg. Elias
Rojas, 10.09.1994: 1 ad. (INBio 1483208); 1

222

juv. (INBio 1483209); leg. Elias Rojas,
13.09.1994: 2 ads. (INBio 1480261); 1 juv.
(INBio 1480281); leg. Elias Rojas,
10.10.1994: 1 ad. (INBio 1483017); leg. Elias
Rojas, 05.12.1994: 1 ad. (INBio 1467174)
Refugio Nacional de Vida Silvestre Barra del
Colorado: Barra del Colorado, Estación
Sardinas: 10°38’52"М, 83°43’52"W, 50 m
a.s.l.: 05.01.1994: 1 ad. (INBio 1478283);
10.02.1994: 4 ads. (INBio 1484010);
12.05.1994: 1 ad., 1 s.ad. (INBio 1484585);
5 ads., 1 s.ads. (INBio 1484587); 6 ads.
(INBio 1484589); 24.05.1994: 1 s.ad. (INBio
1478305); 11.07.1994: 4 s.ads., 2 juv.
(INBio 1484432); 25.07.1994: 1 ad. (INBio
1478294); 28.08.1994: 1 ad. (INBio
1480051); 12.10.1994: 3 juv. (INBio
1484372); 3 ads., 1 s.ad., 2 juv. (INBio
1484372); 12.10.1994: 3 ads., 1 s.ad., 2
juv. (INBio 1484374); 16.10.1994: 3 ads.
(INBio 1484013); 22.10.1994: 2 ads. (INBio
1484991); 7 ads., 1 s.ad. (INBio 1485284);
5 ads., 1 s.ad., 2 juvs. (INBio 1485285); 7
ads., 1 s.ad., 2 juvs. (INBio 1485289); 2 juvs.
(INBio 1485290); 09.11.1994: 2 s.ads.
(INBio 1480044); 09.12.1994: 1 ad. (INBio
1480041); 01.02.1995: 1 ad., 1 s.ad. (INBio
1485145); 02.06.1995: 1 ad. (INBio
1484748); 4 ads. (INBio 1484749) (all leg.
Flor Araya); 10°39’11"N, 83°44’21"W, 15 т
a.s.l.: leg. malacological staff of INBio:
13.01.1994: 1 ad. (INBio 1478017);
16.04.1994: 1 ad. (INBio 1477915); 800 т М
de la Estación Sardinas, Sendero Tono,
10°39’05"М, 83°44’31"W, 50 т a.s.l.: leg.
malacological staff of INBio, 21.11.1993: 1
juv. (INBio 1465699); 1 ad. (INBio 1465700)
Cartago: Parque Nacional Tapantí-Macizo de
La Muerte: Sendero Oropéndola,
09°45'09"N, 83°47’08"W, 1,260 т a.s.l.: leg.
Rosa Guzman, 03.10.1997: 1 ad. (INBio
1488194); Estación Quebrada Segundo,
09°45'45"М, 83*47'18"W, 1,360 т a.s.l.: leg.
Roberto Delgado, 18.10.1994: 1 ad. (INBio
1479646); leg. Roberto Delgado,
03.07.1995: 1 ad. (INBio 1487842)
Monumento Nacional Guayabo: Turrialba,
Santa Teresita, 09°58’26"М, 83°41’42"W,
1,000 m a.s.l., leg. Zaidett Barrientos,
16.12.1994: 1 ad. (INBio 1476052)
Puntarenas: Quebrada Chanchera, 800 m W
de la Playa, 08°37’26"М, 83°26’39"W, 1 т
a.s.l., leg. Socorro Avila, 08.12.1996: 1 ad.
(INBio 1486976)

San Luis, Finca Buen Amigo, 10°16’36"М,
84°47'48"W, 1,100 т a.s.l., leg. Zobeida
Fuentes, 26.06.1995: 1 ad. (INBio 1484382)

RICHLING

Zona Protectora Arenal-Monteverde:
Reserva Biológica Bosque Nuboso
Monteverde: Sendero Brillante, 10°17’59"М,
84*47'10"W, 1,520 m a.s.l.: leg. Каша
Martinez, 17.06.1994: 2 ads. (INBio
1466835); Sendero Bosque Nuboso,
10%17'59"N, 84°47’36"W, 1,600 т a.s.l.:
24.05.1994: 5 ads. (INBio 1466884); 3
ads. (INBio 1466954); 2 ads. (INBio
1467003); 25.05.1994: 1 ad. (INBio
1466842); 1 ad. (INBio 1466870); 2 ads.
(INBio 1466891); 1 ad. (INBio 1466905); 1
ad. (INBio 1467024); 14.06.1994: 3 ads., 2
sads. (INBio 1467031); 15.07.1994: 2 ads.
(INBio 1479528); 16.07.1994: 2 ads.
(INBio 1479539); 25.09.1995: 1 ad. (INBio
1498806); 28.10.1995: 1 ad. (INBio
1498590); 20.10.1996: 4 ads. (INBio
1498828) (all leg. Kattia Martinez); Sendero
Bosque Nuboso, 10°17’59"М, 84°47’36"W,
1,520 m a.s.l.: leg. Zaidett Barrientos,
14.10.1994: 1 s.ad., 1 juv. (INBio 1468141);
1 ad. (INBio 1468211); 1 ad. (INBio
1468212); Sendero el Camino, 10°18'03"N,
84°47'15"W, 1,560 m a.s.l.: 23.05.1994: 5
ads., 1 s.ad. (INBio 1466912); 1 s.ad. (INBio
1466947); 23.05.1994: 1 s.ad. (INBio
1466996); 2 ads. (INBio 1467010);
25.05.1994: 1 ad. (INBio 1466863); 1 s.ad.
(INBio 1466968); 10.06.1994: 7 ads. (INBio
1480426); 14.07.1994: 2 ads., 1 s.ad. (INBio
1480126); 2 ads. (INBio 1480128); 5 ads.
(INBio 1480129); 1 ad. (INBio 1480149);
08.08.1994: 3 ads. (INBio 1479517); 2 ads.
(INBio 1479550); 1 juv. (INBio 1479838);
16.09.1994: 1 ad. (INBio 1480098);
10.10.1994: 1 ad. (INBio 1485422);
26.09.1995: 1 ad. (INBio 1498807) (all leg.
Kattia Martinez); Sendero el Roble,
10°18'16"N, 84*47'27"W, 1,600 т a.s.l.: leg.
Kattia Martinez, 08.11.1994: 1 ad. (INBio
1480132); Sendero Chomogo, 10°18’22"М,
84°47'23"W, 1,640 m a.s.l.: 13.08.1994: 1
ad. (INBio 1480152); 10.10.1994: 1 ad.
(INBio 1485426); 08.12.1994: 1 ad. (INBio
1477521); 15.12.1994: 1 ad. (INBio
1484687); 06.03.1995: 1 ad. (INBio
1485441) (all leg. Каша Martinez); Sendero
Bosque Eterno, 10°18’22"М, 84°47’40"W,
1,600 т a.s.l.: 09.06.1994: 1 ad. (INBio
1480119); 06.08.1994: 1 sad. (INBio
1466793); 28.10.1995: 1 ad. (INBio 1498581)
(all leg. Kattia Martinez); Sendero el Rio,
10°18’29"N, 84°47'37"W, 1,600 т a.s.l.:
15.07.1994: 1 juv. (INBio 1479353);
08.12.1994: 3 ads. (INBio 1480127); 1 ad.
(INBio 1480130); 1 ad. (INBio 1480131);

CLASSIFICATION OF HELICINIDAE 223

04.07.1995: 1 s.ad. (INBio 1485234); 1 ad.
(INBio 1485235) (all leg. Kattia Martinez);
Estación la Casona, 10°18"11"N,
84°47’50"W, 1,520 т a.s.l.: 08.09.1994: 1
ad. (INBio 1479451); 22.09.1995: 3 ads.
(INBio 1498804); 28.10.1998: 3 ads. (INBio
1498632) (all leg. Kattia Martinez);
10°18’15"М, 84°47’46"W, 1,520 m a.s.l.,
leg. malacological staff of INBio,
28.07.1994: 5 ads. (INBio 1477749)
Finca tomas, por Casa Boby, 10°18'12"N,
84%48'22"W, 1,520 m a.s.l., leg. Kattia
Martinez, 24.10.1995: 1 s.ad. (INBio
1498808)

Cerro Plano, 10°18'58"N, 84°49’09"W, 1,300
m a.s.l., leg. Каша Martinez, 02.09.1996: 1
ad. (INBio 1498652)

Limón: Los Diamantas Farm, 11.08.1971: 1
ad. (UF 69846); Los Diamantes Farm, 12 mi
SE Guapiles [about 10%11'N, 83°37’W], leg.
R.W. McDiarmid, 13.08.1971: 1 ad. (UF
214167)

Moin, hill #1 [about 10°М, 83°04’W}, leg. С.
Little, 29.09.1967: 1 ad. (UF 214158)
Cueva Castil, near Limon [about 10%N,
83°02’W], leg. Colin Little, 30.08.1967: 5
ads. (UF 214165)

Puerto Limon, football field adjacent to Stan-
dard Fruit Box Factory [about 10°N,
83°02’W], leg О.С. Robinson (TU-954),
19.05.1984: 1 ad. (UF 155820)

Along road cut, along south side of Rio
Banano, opposite La Bomba, 09°54'49.7"М,
83°03'56.4"W, leg. О.С. Robinson & J.M.
Montoya, 21.09.1998 (APHIS PPQ USDA)

OTHER SOURCES
COSTA RICA

Pandora [about 09°43’М, 82°58'W], leg. Jay
Savage, 01.05.1964: 1 ad. (UF 214156); leg.

Guanacaste: Tilaran [about 10°28’30"М,
84”58'30"WJ], leg. Univ. Alabama, М. Smith
coll.: 6 ads. (UF 95283)

1.7 mi S Tilaran on road to Quebrado
Grande [about 10°27’N, 84°58’W], leg. R.W.
McDiarmid, 28.08.1971: 2 ads., 1 s.ad. (UF
214166)

10 mi W Tilaran [about 10°26 N, 85”06'W],
leg. Ronald Heyer, 06.08.1964: 1 ad. (UF
214163)

Monte Verde [about 10°18’М, 84°47’W], leg.
Savage & Scott, 13-16.05.1964: 6 ads., 3
s.ads. (UF 214170)

Alajuela: San Carlos [about 10°20’М,
84°26’W], leg. McGinty coll., ex Preston 8
Tomlin: 3 ads. (UF 160150)

Cariblanca [about 10%17'N, 84°12’W)],
Sarapiqui, 600 m a.s.l., P. Biolley (#267): 5
ads. (MHNN)

Chemin de Sarapiqui, S. Miguel [about
10°19’N, 84°11’30"W], leg. P. Biolley: 11
ads. (MHNN)

Тезайа [Tetsalia?, about 10°21’N, 84°24'W],
leg. R. W. McDiarmid, 18-20.07.1971: 1 ad.
(UF 214164)

Heredia: Puerto Viejo [de Sarapiqui, about
10°28’N, 84*00'30"W], leg. P. Biolley: 2 ads.
(ZMB 103242)

Río Frio, Standard Fruit Co., 10°20'N,
83°53’W, 300 ft., leg. Michael J. Corn,
21.11.1969: 1 ad. (UF 214160); 22.11.1969:
2 s.ads. (UF 214172)

[not: “Alajuela”], Río Frio [about 10°20'N,
83°53’W], leg. Michael J. Corn, 05.05.1970:
rad (УЕ 214161); 15.05. 1970: 1:ad. (UF
214171)

San José:

F. G. Thompson (FGT-100), 05.08.1964: 3
ads. (UF 214157)

3.2 кт N Pandora [about 09°45'N,
82°58'W], leg. Е.С. Thompson (FGT-98),
04.08.1964: 8 ads, 1 s.ad. (UF 214155)

1 km NW of Cahuita, 09°44.5’N, 82°50.9’W”,
leg. FG. Thompson (FGT-5616),
25.02.1996: lad. (UF 258427)

Trib[utary] to Rio Moin [Valle de
Talamanca!], 572 500 Е, 397 600 $, 430 m
a.s.l. [09°37’45"N, 83°00'18"W], leg. E.L.
Raiser (ELR-082), 10.08.1994: 1 ad. and in
alcohol (UF 41438) (UF 41437); leg. Е.
Alvando (ELR-087), 11.08.1994: 2 ads. (UF
41442)

Amubre [about 09°32’N, 82°57 30"W], leg.
Norman Scott, 16.03.1964: 1 ad. (UF
214168)

Tarbaca [about 09°49’25"М,
84°06’39"\М, leg. P. Biolley: 2 ads. (ZMB
103246)

2 lots mixed: Cartago: 1. Azahar de Cartago
[not clear, if referring to the town Cartago,
?about 09°52’М, 83°55’W], San Jose: 2.
Tarbaca [about 09°49’25"М, 84°06'39"W],
leg. P. Biolley: 10 ads. (MHNN)

Cartago: 2 lots mixed: Cartago: 1. Azahar de

Cartago [not clear, if referring to the town
Cartago, ?about 09°52’N, 83°55’W], San
José: 2. Tarbaca [about 09°49'25"N,
84°06’39"W], leg. Р. Biolley: 10 ads. (MHNN)
Tapanti, 4300 ft. [about 09°47’М, 83°48’W],
leg. F. G. Thompson (FGT-23), 26.06.1963:
1 ad. (UF 214169)

Turrialba [about 09°54’30"N, 83°41’W], ex
coll. S.G.A. Jaeckel: 2 ads. (HNC 39842);

224 RICHLING

coll. Bosch, ex Rolle, ex Wagner: 6 ads.
(SMF 180790/6); Turrialba, versant de
l'Atlantique, 750 т [about 09°54’30"М,
83°41'W], leg. P. Biolley (#146), 07.1893: 4
ads. (MHNN)

Valleé de Tuis [about 09°51’N, 83°35’W], H.
Pittier, 9.1893 ex coll. Wiegmann: 1 ad.
(ZMB 70633)

Cartago?: Cache [Cachí?, about 09°50’N,
83°48'W], leg. Roger, ex Godwin & Salvin: 1
ad. (ZMB 40836)

Puntarenas: 1.5 mi NE Monte Verde [about
10°19'N, 84°47'W], leg. R.W. McDiarmid
(RWM-11), 17.02.1966: 6 ads. (UF 214162)

Costa Rica, without locality further specified:
leg. Beal-Maltbie coll., ex W. F. Webb coll.: 4
spec. (UF 237539); leg. H. G. Lee, ex G.D.
Robinson, W.F. Webb: 1 ad. (UF 166943);
leg. Univ. Alabama, T.H. Aldrich coll. (THA-
8213), ex Webb: 1 ad. (UF 95254); 1 ad.
(UF 214110); leg. P. Biolley: 4 ads. (MHNN);
leg. Carmiol: 2 ads. (ZMB 103244); ex
Fulton: 3 ads. (ZMB 64488); 1 ad. (ZMB
103245)

NICARAGUA
Not further specified: Sumichrast: 2 ads. (UF
214108)

PANAMA

Bocas Del Toro: Colon Island, leg. McGinty
coll.: 2 ads. (UF 185608); Isla Colon, ca. 12
km NNW of Bocas del Toro, 09°25’00”М,
82°1623"W, leg. Е.С. Thompson (FGT-
4726), 19.09.1990: 1 ad. (UF 167537); Isla
Colon, limestone knoll along E coast, 5 km
NNE of Bocas del Toro, 09%23'05"N,
82”14'09"W, leg. Е.С. Thompson (FGT-
4727), 20.09.1990: 1 ad. (UF 167538)

М end of Isla San Cristobal, 09°17’28”М,
82°15'51"W, leg. Е.С. Thompson (FGT-
4730), 21.09.1990: 1 ad. (UF 167541)

Isla Bastimentos, 0.5 km NE of Bastimentos
Town оп trail to Wizard Beach, 09°20’59”М,
82°12'15”W, 60 т a.s.l., leg. Е.С. Thompson
(FGT-4731), 22.09.1990: 1 ad. (UF 167544)
Ojo de Agua, Filo Almirante, 09°17’32”М,
82”27'43"W, 300 m a.s.l., leg. Е.С. Thomp-
son (FGT-4733), 24.09.1990: 6 ads. (UF
167551)

Colón, Canal Zone: 0.5 mi SE Achiote, $5. В.
Telford, 12.1969, 1 ad. (UF 214173); 4.8 km
SE Achiote, leg. F.G. Thompson (FGT-1130),
27.04.1969: 20 ads. (UF 214154)

0.8 km SW Madden Dam, leg. F.G. Thomp-
son (FGT-1131), 02:05:1969: 1. аа. (UF
214159)

N bank Chagres River, 6 km NNE Gamboa,
leg. S.R. Telford, 22.04.1969: 1 ad. (UF
214174)

Canal Zone, not further specified: leg. Univ.
Alabama, M. Smith coll., ex Clark 5 ads., 1
s.ad. (UF 95284); leg. Univ. Alabama, M.
Smith coll.: 11 ads. (UF 95285)

Panama, without locality further specified: leg.
Beal-Maltbie coll., ex W. Webb coll. (UF
237401)

Description

Shell (Fig. 335A-C): Conical-subglobose,
solid, relatively large sized and dull to
slightly shiny. Color: basic color yellowish to
whitish-opaque, towards apex and on upper
half of whorls often a more or less intensive
tinge ranging from reddish-brown to flesh
colored, in some specimens involving the
whole shell with exception of outer lip. The
color is slightly overlapped with fine light to
transparent patches and lines giving the
shell a special ornamentation. Surface tex-
tured with fine growth lines and oblique
grooves of different individual orientations
but of same general direction (Fig. 14),
causing the dull appearance. Embryonic
shell of about 1 whorl; 4—4°/, (lectotype: 472)
subsequent whorls nearly straight and only
very slightly convex; last whorl with a touch
of angulation on the periphery; whorls
equally extending in size, forming a very
regular, blunt spire. Suture very slightly im-
pressed. Aperture oblique and nearly
straight, last whorl only very slightly de-
scending, inserting exactly at periphery or
just below it. Outer lip independent of color
of whorls, always yellowish-whitish, remark-
ably thickened and broadly expanded, upper
palatal region slightly sinuate. Reflection
nearly rectangular to the whorl; transition to
columella with a more or less pronounced
denticle. Columella slightly curved, its tran-

FIG. 13. Axial cleft and muscle attachments of
Helicina funcki, IR 757; scale bar 5 mm.

CLASSIFICATION ОЕ HELICINIDAE 225

FIG. 14. Teleoconch surface structure of Helicina funcki. A. On 1* whorl. В. On 2™ whorl. С.
On 3% whorl. D. On 4" whorl (inset same magnification as in С); scale bar 100 um.

sition to body whorl marked with a perpen-
dicularly impressed line or even a groove.
Basal callus weakly developed and nearly
completely smooth or very slightly granu-
lated.

Juvenile specimens exhibit a roundly cari-
nated periphery sometimes bearing
periostracal spiral lines.

Internal Shell Structures: (Fig. 13)

Teleoconch Surface Structure: In Helicina
funcki, the transitional structure is followed
by a pattern of oblique diverging grooves,
which is maintained in all whorls (Fig. 14B-
D). The grooves only increase in length and
become more widely spaced.

226

Embryonic Shell: The surface is structured
with pits arranged in concentric lines (Fig.
15А). The diameter of these pits is approxi-
mately equal to the interspacial distance
between the pits in a line as well as between
the lines of pits themselves, although the
arrangement is somewhat irregular. This is

RICHLING

the typical structure for Helicina funcki, but
deviations also occur (Fig. 15B). The “com-
pressed” pattern obviously results from ir-
regularities experienced during growth (in
the present case, during the younger part)
which caused a different form of the embry-
onic shell and more closely spaced growth

lines resulting in a reduction of the diameter
(Fig. 15А: 1040 pm, Fig. 15B: 930 pm).
Concerning the size of the embryonic shell,
the type material falls within the range of
Costa Rican lowland populations (e.g.,
Cahuita), whereas specimens from higher
altitudes of Monteverde consistently develop
a much larger embryonic shell.

Diameter: 954 um (+ 41) (870-1,040) (п =
16) (IR 1630, IR 1639, IR 1642, IR 1648,
Cahuita); 1,160 um (+ 43) (1,060-1,240) (п
= 20) (IR 843, Monteverde); 980 um (+ 40)
(940—1,040) (п = 4) (BMNH 20010497.1-4,
type lot, lectotype: 1,000 um); 970 um (+ 50)
(9201,020) (n = 2) (MIZ 8989, type lot of
Нейста funcki costaricensis, lectotype:
1020 pm).

SES AE
La. y

Operculum (Fig. 16): Only slightly calcified,
calcareous plate not fully extended over
horny plate, leaving a free margin, thickened
towards columellar side. Color dark reddish-
brown to nearly black, only the margin trans-
parent. Columellar side slightly S-shaped,
upper end acute and pointed or nearly rect-
angular, lower end rounded, but slightly trun-
cated towards the columella.

Animal (Fig. 337A, B): The color of the animal
does not show any great variation, either at
different sites or within the populations. Sole

FIG. 15. Embryonic shell of Helicina funcki, A.
Typical. B. Irregularities during growth; scale bar
100 um.

FIG. 16. Operculum of Helicina funcki, IR 757;
scale bar 2.5 mm.

CLASSIFICATION OF HELICINIDAE 227

and sides of the foot are whitish-yellowish
changing gradually to dark brown-greyish
towards the upper side and head. The ten-
tacles are also dark brown-greyish with a
light tip. The mantle has a whitish-greenish
pigmentation shining through the shell, thus
providing the live specimens with a some-
what greenish appearance. The mantle only
bears a dark color in juvenile specimens
(Fig. 337B), being sometimes spotted with
yellow, thus causing those juvenile individu-
als to appear darker.

Radula (Fig. 17): The cusps on the A- and C-
central are vestigial, only the B-central with
4-5 more or less-well developed cusps.
Comb-lateral with 7-9 cusps, cusps on
marginals slowly increasing in number.
Radula with about 70-105 rows of teeth.

Female Reproductive System (Figs. 18, 19):
The receptaculum seminis enters at the
middle of the inner side of the descending
limb of the V-organ. It is a cylindrical, slender

FIG. 17. Radula of Helicina funcki. A. Centrals. B. FIG. 18. Female reproductive system of Helicina
Comb-lateral. C. Marginals; scale bar 50 um (A, funcki, IR 1312; scale bar 2 mm.
В), 100 pm (С).

228

JS _
Yo, > )
( Bl = 4 — Е
и IDS}
N ps
| ( \ { cE
| | \
| — sy )
| | € | | /
Ç — \ \ — À | ( \ >
= LS LA
\ \ | р \ = y \ Ÿ
\ < ES N He
\ = N <4
T2 SO TES ES
KO) NÖ) —

NS 9 Ya

FIG. 19. Variability ofthe female reproductive sys-
tem of Helicina funcki, IR 1312; scale bar 2.5 mm.

sac. The bursa copulatrix consists of numer-
ous remarkably elongated lobes, some of
which are always further subdivided. The
provaginal sac is irregularly shaped, dors-
oventrally flattened and bears lobe-struc-
tures at its distal side. The stalk is
comparatively long and deeply curved ante-
riorly, as is the adjacent part of the reception
chamber. The pallial oviduct is relatively long

0.4 T T T T

RICHLING

and shows mainly transversal constrictions.
In Figure 19, the right drawing shows the
genital for a slightly immature specimen. The
main difference consists in the much less
thickened pallial oviduct, in which, except for
a slight enlargement, the final shape of the
accessory structure is already developed.

Morphometry and Sexual Dimorphism
(Tables 3-4, Figs. 21-28)

From my own material, all adult specimens
of known sex and populations with at least a
few specimens of each sex were compared. A
few populations with scanty material were in-
cluded because of their otherwise undocu-
mented origin.

The measurement of the weight is especially
difficult in Helicina funcki, because a consider-
able part of the weight results from the
strongly developed, broadly expanded outer
lip. In Fig. 20, the increase of weight during
growth is illustrated for the population from
Cahuita (juveniles were studied from lot IR
1312). An additional non-mature shell from
Rio Peje is included as an example for heavy-
shelled specimens to demonstrate that the
increase of weight during juvenile growth pe-
riod continues at about the same rate. Shells

RE T T =
| Cahuita adult +
weight | Cahuita juvenile + |
r . O
[9] Río Peje adult © a
O
2] je]
0.3 Е О 2 = =]
O
oo
po El
0.25 + ce 4
7 О Au О
D a
ee ©
0:2) = + + + =
++ + и +
| + O ++ + ae
ES
0.15 Ш. 4 -
+ о +
ы + a E en +4 + N E
0.1 P ++ + + a + ES 52
4 + +4 + ©
nos o ES 2e juvenile
| o
o ©
o OU
o
y Lee 1 1 L fu u 1 —Η
0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 volume [ml] 0.9

FIG. 20. Increase of weight during growth of Helicina funcki from Cahuita
(juveniles IR 1312), compared also with adults from Río Peje (one juvenile
included); arrows indicate “thin-lipped” adults; juvenile = shells without
expanded lip.

CLASSIFICATION OF HELICINIDAE

229

TABLE 3. Measurements of different populations of Helicina funcki from material collected by the
author, given as mean value with standard deviation, minimum and maximum value (min, max), and
number of specimens (тт./тах. diam. = minor/major diameter, col. axis = columellar axis); linear
measurements [mm], weight [g], volume [ml].

Height
Height
Maj. diam.
Maj. diam.
Min. diam.
Min. diam.
Outer lip
Outer lip
Last whorl
Last whorl
Col. axis
Col. axis
Weight
Weight
Volume
Volume

Height
Height
Maj. diam.
Maj. diam.
Min. diam.
Min. diam.
Outer lip
Outer lip
Last whorl
Last whorl
Col. axis
Col. axis
Weight
Weight
Volume
Volume

Sex

= ió = a Gl Bra ee ii

Sex

= a = a = ni de Fu a

“Rincón de la Vieja” (altitude 800 m)

Mean
value

11.14
10.60
12.02
11.56
10.84
10.39
195
7.83
9.08
8.86
8.73
8.14
0.078
0.093
0.471
0.409

lots IR 972, IR 979

Deviation

0:33
0.32
0.31
0.69
0.23
0.56
0.20
0:52
0.17
0.27
0.17
0:13
0.008
0.032
0.031
0.053

Min
10.80
10:22
141.57
10.66
10.53

9.58

7.56

7.13

8.78

8.43

8.50

7.96
0.066
0.032
0.435
0.342

Max Number

11.52
11.04
12.63
1285
11.22
11.03
8.11
8.76
9.30
9.30
8.97
8.29
0.094
0.132
0.515
0.471

4

AAAAAHARAAAAAAAAAA

“Monteverde - Finca Ecológica”
(altitude 1330 m)

lots IR 859, IR 946, IR 1246

Mean
value

10.96
10.03
11.91
11.09
11.16
9.94
8.04
7.65
9.02
8.32
8.55
7.88
0.156
0.126
0.444
0.345

Deviation

0.22
0.22
0.43
0.44
0.36
0.33
0.21
0.30
0.25
0.30
0.19
0.27
0.037
0.046
0.028
0.020

Min

10.65
9.70
11.33
10.43
10.45
9.45
1.19
7.34
8.56
7.88
8.16
1.98
0.105
0.057
0.407
0.315

Max Number

11,31
10.32
12.54
11.70
11:72
10.34
8.30
8.09
9.30
9:55
8.86
8.28
0.198
04177.
0.490
0.365

WAWAWAWAWAWAWAHOWH

“Mirador Gerardo” (altitude 1450 m)

Mean

value Deviation

11.05
10.17
12.42
11.90
11.19
10.48
817.
7.83
9.15
8.61
8.62
7.89
00137
0.142
0.491
0.400

0.48
0.11
0.36
0.12
CS
0.08
0.14
0.26
0.27
0.06
0:32
0.11
0.025
0.015
0.031
0.005

Min
10.55
10.04
11.96
11.67
10.78
10.36

7.92

7.49

8.82

8.49

8.08

7.67
0.117
0.125
0.452
0.390

lots IR 924, IR 928, IR 1230

Max Number

11.74 4
10.34
12.96
12.07
11.67
10.65
8.36
8.25
9.48
8.68
9.10
8.09
0.187
0.162
0.553
0.408

AAHRAAAAAAAAAAAA

“Monteverde” (altitude 1500 т)
lots IR 843, IR 927, IR 1194, IR 1435,

Mean
value

1.31
10.29
12.73
12.10
11.45
10.77
8.16
1.89
9.39
8.71
8.82
7.98
0.174
0.132
0.521
0.420

IR 1627

Deviation Min

0.33
0.38
0.36
0.34
0.34
0.37
0.20
0.23
0.25
0:32
027
0.26
0.018
0.024
0.039
0.035

10.24
9.71
11.70
11.54
10:25
10:19
7.40
7.15
8.57
8.26
8.14
1-02
0137
0.075
0.385
0.368

Max Number

12517 18
11:55 15
13.38 18
12.98 15
12.35 18
11:99 15
8.65 18
8.54 15
10.09 18
9.74 18
9.45 18
8.81 15
0.235 18
0.188 15
0.653 18
0.518 15

(Continues)

230

(Continues)

Height
Height
Maj. diam.
Maj. diam.
Min. diam.
Min. diam.
Outer lip
Outer lip
Last whorl
Last whorl
Col. axis
Col. axis
Weight
Weight
Volume
Volume

Height
Height
Maj. diam.
Maj. diam.
Min. diam.
Min. diam.
Outer lip
Outer lip
Last whorl
Last whorl
Col. axis
Col. axis
Weight
Weight
Volume
Volume

Sex

= PS ав

“Las Pavas” (altitude 800 т)
lots IR 952, IR 955, IR 1273, IR 1637

Mean

value Deviation

12.12 022
1111 0.48
13.38 082
12.30 0.35
12.11 0.18
11.03 0.34
8.74 0.23
8.40 0.28
10.14 0.10
9.31 0.32
933 013
8.49 0.40
0.152 0.035
0.151 0.020
0.636 0.035
0.473 0.042

“La Selva”

lots IR 1061,

Mean

value Deviation

12,65
11.02
14.06
13.37
12.88
12.00
9.96
9.08
10.58
9.86
9.77
8.93
0.236
0.219
0.729
0.580

0.29
0.36
0.21
0.23
0.14
0.18
0.24
0.29
0.19
027
0.17
0.21
0.036
0.044
0.037
0.041

Min

11.81
10-18
12.93
11.85
11.61
10.60
8.32
7.39
10.00
8.76
9.17
У
0.103
0.128
0.593
0.404

(altitude 60 т)
IR 1062, IR 1182

Min

12:16
10.92
13.56
12:78
12:51
11.51
9.03
8.30
10.21
8.98
9.38
8.37
0.134
0.141
0.652
0.505

RICHLING

Max Number

12.75
12.08
14.08
13.00
12.44
11.60
9.23
8.75
10.40
9.96
9.18
9.25
0.253
0.197
0.703
0.549

Max Number

13.06
12:26
14.37
13.82
13.09
12:32
10.18

9.47
10.95
10.22
10.00

9.35
0277
0.278
0.783
0.638

юю<юююююююююююююю CO

Où Y OY O1 Y an O1 on own on

“Tortuguero” (altitude 0-10 т)
lots IR 1348, IR 1620, IR 1653

Mean
value Deviation
12.22 1.13
11273 0.21
14.08 0.78
19:27 0.09
12.66 0.77
11.78 0.16
9.40 0.85
8.81 0.01
10.36 0.86
9.69 0.03
9.24 0.48
872 0.08
0.147 0.081
0.177 0.062
0.731 0.089
0.568 0.003
“Guayacán”
lots IR 1079,
Mean
value Deviation
12.14 0.34
11:50 0.31
13.18 0.36
1270 0.17
11.97 0.27
11.41 0.10
8.73 0.11
8.52 0.04
10.14 0.29
9.61 0.07
9.34 0.24
8.76 0.11
0.191 0.032
0.175 0.004
0.615 0.040
0.534 0.001

Min

11.09
11:52
13.31
13.17
11.84
11.62
8.55
8.80
9.50
9.66
8.02
8.64
0.061
0.115
0.602
0.565

Max

13.34
11.94
15:25
13.36
13.82
11.93
10.25

8.82
1122

9.72

9.81

8.80
0.268
0.239
0.865
0.570

Number

NOWNWNWNNNNN WN WN DY

(altitude 520 m)
IR 1090, IR 1608

Min
11.47
11.19
12.69
12.45
11.65
11227

8.48

8.48

9.66

9.54

8.88

8.63
0.138
0.171
0.533
0.533

Мах

12:55
11.80
13.62
12.89
12.42
11:56
8.88
8:55
10.43
9.68
9:56
8:92
0.233
0:179
0.669
0.534

Number

N O1 N O1 © UN UN Y Y UY UN Y

(Continues)

CLASSIFICATION OF HELICINIDAE 231

(Continues)

“México” (altitude 40 m) “Río Peje” (altitude 160 т)
lots IR 274, IR1191, IR 1406 lots IR 751, IR 1300, IR 1552

Mean Mean
Sex value Deviation Min Max Number value Deviation Min Max Number

Height f @ 11.22 0.23 10.92 11.68 4 12.54 0.33 11.56 18:64 17
Height m 10.40 0.16 10.15 10.62 6 11.68 0.40 10.88 12.55 20
Ма]. Чат. f 13.21 0.30: 12.82 13.64 4 13.91 0.32 13.17 14.65 17
Ма]. dam. m 12.11 0.37 1132 12.55 6 13.13 0.31 1261 13:97 20
Min. diam. f 11.98 0.29 11.59 12.40 4 12.74 0.29 11.93 13.27 {TA
Min. diam. т 10.90 0.28 10.36 11.29 6 11.80 0.24 11.34 12.44 20
Outer lip f 8.68 0.15 8.49 8.98 4 9.45 0.28 8.75 9.98 17
Outer lip m 8.01 0.26 7.41 8.41 6 8.98 0.28 842 9.87 20
Last whorl f 9.70 0.24 9.43 10.08 4 10.61 0.28 9.84 11.47 17
Last whorl т 8.90 0.13 8.65 9.09 6 9.97 0.25 9.54 10.66 20
Col. axis f 8.67 0.16 849 8.99 4 9.59 0.32 886 10.66 17
Col. axis m 8.05 0.09 793 8.17 6 8.95 0.29 813 9.78 20
Weight f 0.200 0.030 0.175 0.260 4 0.259 0.036 0.142 0.344 17.
Weight m 0.200 0.034 0.130 0.238 6 0.249 0.040 0.131 0.331 20
Volume Е 0.574 0.030 0.542 0.624 4 0.703 0.050 0.548 0.820 16
Volume т 0.432 0.036 0.371 0.494 6 0.562 0.038 0.452 0.686 19
“Río Barbilla” (altitude 70 т) “Uatsi’ (altitude 30 m)
lot IR 1545 lots IR 766, IR 1114, IR 1632
Mean Mean
Sex value Deviation Min Max Number value Deviation Min Max Number
Height | 1222 0100 12.21. 12.22 2 12.61 0.18 12.28 12.79 6
Height m: 141232 050" 10.75 12212 5 11.94 0.18 11.59 12.15 5
Mai. dam. f 13.93 0.00 13.93 13.93 2 14.04 0.15 13.78 14.33 6
Ма]. Чат. m 12.73 0.47 11.97 13.61 5 13.36 0.60 12.64 14.35 5
Min. diam. f 12.60 0.02 12.58 12.62 2 12.68 0.19 12.45 12.97 6
Min. dam. тм 11.37 0.43 10.73 12.09 5 12.05 0.44 11.52 12.98 5
Outer lip Е 9.29 0.02 9.27 9.30 2 9.50 0.19 916 9.95 6
Outer lip m 8.57 0.29 8.04 8.95 5 9.19 0.35 8.32 9.68 5
Last whorl f 10.31 0.10 10.21 10.41 2 10:51 0.20 10.04 10.81 6
Last whorl m 9.60 0.455 8.78 10.30 5 9.98 0.30 9.56 10.57 5
Col. axis f 9.41 0.25. 9 16 9.66 2 9.53 0.12 9.36 9.81 6
Col. axis т 8.62 03545 11932 5 8.89 0.16871 911 5
Weight f 0.240 0.012 0.228 0.251 2 0.263 0.036 0.219 0.335 6
Weight т 0.157 0.041 0.089 0.219 5 0.148 0.049 0.087 0.226 5
Volume f 0.688 0.019 0.669 0.707 2 0.708 0.021 0.681 0.744 6
Volume т 0.547 0.073 0.442 0.695 5 0.631 0.074 0.546 0.764 5

(Continues)

232 RICHLING
(Continues)
“Cahuita” (altitude 0-10 т)
lots IR 107, IR 757, IR 897, IR 898, IR
“Shiroles” (altitude 100 т) 907, IR 1095, IR 1312, IR 1555, IR
lots IR 911, IR 1596, IR 1599, IR 1644 1557, IR 1630, IR 1639, IR 1648
Mean Mean
Sex value Deviation Min Max Number value Deviation Min Max Number
Height f 12.54 0.57 1197 13.10 2 1.31 0.551021 12.73 63
Height m 12.04 0.30 11.67 12.50 4 10.66 0.54 9.20 12.08 51
Maj. diam. f 14.16 0.63 13.53 14.79 2 12.76 0.52 11:59 14.55 63
Maj. diam. m 13.49 0.25, 13.14 13.82 4 12.26 0.53 10.85 14.43 Si
Min. diam. f 12.81 0.29’ 12.52 13.10 2 11:67 0.46 10.62 13.26 68
Min. dam. m 12.21 0.19 11.82 12.41 4 11.04 0.45 9.88 12.67 51
Outer lip f 9.24 0.06 918 9.30 2 8.63 0.41 7.76 9.89 63
Outer lip m 9.16 0.21 8.76 9.48 4 8.37 0.41 1.397 9775 51
Last whorl f 10.63 0.40 10.22 11.03 2 9.56 0.47 844 10.71 63
Last whorl m 10.27 0.13 10.10 10.45 4 9.05 0.44 8.17 10.19 51
Col. axis f 9:53 0.30 9.23 9.82 2 8.68 0.43 7.93 9.91 63
Col. axis m 9.24 0.15 9.07 9.49 4 8.12 0.41 7.11 9.18 57
Weight f 0.298 0.030 0.268 0.328 2 0.134 0.026 0.077 0.224 63
Weight т 0.292 0.014 0.274 0.306 4 0.141 0.039 0.057 0.282 Si
Volume f 0.718 0.060 0.658 0.777 2 0.564 0.070 0.437 0.817 60
Volume m 0.622 0.025 0.572 0.656 4 0.471 0.058 0.329 0.662 51
“Manzanillo” (altitude 0-10 т)
lots IR 1096, IR 1320, IR 1642
Mean
Sex value Deviation Min Max Number
Height f 11.81 0.43 10.43 12.45 10
Height Mii” 0.34 1044 12.45 19
Maj. diam. + 13.43 0.43 1239 14.43 10
Maj. diam. m 12.79 0.45 12:01 13.70 19
Min. diam. + 12514 0.40 11.25 12.96 10
Min. dam. m 11.48 0.33 10.99 12.25 19
Outer lip f 9.11 O47 197 9:76 10
Outer lip m 8.66 0.28 8:03 9.23 19
Last whorl f 9.97 0.35: 9.28: 10:55 10
Last whorl т 9.31 0.24 8.79 10.03 19
Col. axis f 8.93 0.34 7.91 9.57 10
Col. axis m 8.56 0.27 7.96 9.29 19
Weight Е 0.180 00180 13210217 10
Weight m..0:168. 70.031 0:28 20257 19
Volume f 0615 0.053 0.486 0.719 10
Volume m 0.522 0.043 0.452 0.630 19

still lacking their thickened ощег lip but other-
wise being fully grown weigh from approx. 50
to 100 mg depending on the size, with adults
ranging up to 344 mg. Thus, the weight is
mainly influenced by the thickness of the outer
lip, which not only depends on the age of the
individual but also on size. A recently devel-

oped outer lip is probably thinner than the one
of a truly fully grown individual, a factor that
cannot be differentiated in field collections, al-
though the attempt was made to at least ex-
clude non fully grown adult shells in
populations with sufficient material. The ar-
rows in the figure exemplary indicate such

CLASSIFICATION ОЕ HELICINIDAE 233

“thin-lipped” adults. In fact, this may have re-
sulted in too low measurements, thereby re-
ducing the mean value.

In the diagrams, the populations are roughly
grouped according to their locations: Rincon
de la Vieja to Monteverde (NW-Costa Rica:
mountain chains of Guanacaste and Tilaran),
Las Pavas to La Selva (western and northern
Caribbean plain), Guayacan to Barbilla
(middle Caribbean plain), and Uatsi to
Manzanillo (Southern Caribbean plain).

Because the sex of the additional popula-
tions from the INBio material could not be de-
termined, they have to be compared to the
average value of both sexes.

Rincon de la Vieja (n=4/4)

Mirador Gerardo (n=4/4)

—
Monteverde - Finca Ecológica get)
—

Monteverde (n=18/15)

Las Pavas (n=7/5)

Tortuguero (n=2/2)

La Selva (n=9/9)

ja (n=5/2)

Morphometry: The comparison of the popula-
tions showed that they differ in all character-
istics. Except for weight, these differences
between the populations exhibit a similar
pattern in each characteristic (Figs. 21-28);
that is, “Monteverde — Finca Ecológica” al-
ways has the smallest dimensions, sug-
gesting that relations between the
measurements at each locality are constant.
In fact, several relations were tested and,
except for the size, no significant differences
were found at the different locations. The
individual data can diverge remarkably from
the mean value and as may be expected,
due to the fact that as more specimens are

Rincón de la Vieja (n=4/4 9
Sy AS = ——— м

Mirador Gerardo (n=4/4)

ST - Finca (n=4/3)

Monteverde (n=18/15)

Las Pavas (n=7/5)

=

Tortuguero (n=2/2)

La Selva (n=9/9)

SIE (n=5/2)

Mexico (n=4/6)
——
Rio Peje (n=17/20)

Barbilla (n=2/5)

Mexico (n=4/6)

Rio Peje (n=17/20)

Barbilla (n=2/5)

Uatsi (n=6/5)

SU
—

Shiroles (n=2/4)

Cahuita (n=63/51)

Manzanillo (n=10/19)

= 1 = 1 1
0 8 9 10 11 12

13 [mm] 14

FIG. 21. Shell height of different populations of
Helicina funcki in Costa Rica according to Table 3;
on each line: mean value, standard deviation,
absolute range; number of individuals given as “n
= females/males”; upper line: females, lower line:
males; in between and shaded: average of both
for comparison with populations of unknown sex.

Uatsi (n=6/5)

iy SES zz
Shiroles (n=2/4)

Cahuita (n=63/51)

Manzanillo (n=10/19)

1 1 1 L 1 1 L
ás 6 11 [mm] 12

FIG. 22. Expansion of outer lip of different
populations of Helicina funcki in Costa Rica
according to Table 3; for explanations see Fig.
21.

234

included in the investigation, the wider the
range of the data can become.

Regarding these size differences, the vol-
ume appears to illustrate them best, espe-
cially because it is the only measurement
directly reflecting the actual living conditions
of the animal. The extrema: the biggest indi-
vidual from Tortuguero (0.865 ml) had a shell
with 2.75 times the volume of the smallest
from Monteverde — Finca Ecológica (0.315
ml); the mean value of the population of
Shiroles is 1.7 times higher than that of
Monteverde - Finca Ecológica. At the upper
four locations in Fig. 27 (mountain chains of
Guanacaste and Tilarán), representing the
highest altitudes, consistently smaller-
shelled populations are found, as well as at
two far distant localities in the Caribbean
plain (México and Cahuita). Shells from Las

Rincón de la Vieja (n=4/4)

RICHLING

Pavas, Guayacán and Manzanillo are of in-
termediate size (Fig. 29).

Additional populations from the collection of
INBio were subsequently compared only in
the minor diameter, because the volume
could not be measured and - as previously
demonstrated — other characteristics varied
in the same way. The minor diameter was
chosen instead of the height, because it can
be measured more exactly and it is better
correlated with the volume (Fig. 30, Cahuita
population). The populations show similar
differences in size at different sites (Figs. 24,
29). The two individuals of Isla Uvita repre-
sent the smallest Helicina funcki measured
in this study. The few corresponding locali-
ties (Monteverde, Manzanillo, Rincón de la
Vieja, Mirador Gerardo — Santa Elena,
Shiroles close to Hitoy Cerere) agree sur-

Santiago (type lot BMNH) (n=4)

Mirador Gerardo (n=4/4)

a

Monteverde - Finca Ecológica (n=4/3)

Monteverde (n=18/15)

Santa Clara (type lot H. funcki costaricensis (n=2)
E] Á SET

Estación Góngora (n=6)
Es) fe — ee

Estación San Cristóbal (n=12)
== |

Rincón de la Vieja (n=6)
JE D -———————

Shiroles (n=2/4)

y & = == Tenorio (n=2)
iz ES on
Las Pavas (n=7/5) EU Elena (n=8) ,
mr EE SS
— Peñas Blancas (n=3)
Tortuguero (n=3/2) Sur de GSD)
un onteverde (n=
a — sy
La Selva (n=9/9) a, (n=8)
AAA e
_ Monte Cele (n=4)

Guayacan (n=5/3) =) А er

ER } pe Estación Playuelas (n=7)
aja

México (n=4/6) Caño Negro (n=8)
E I] i | Barra del Colorado (n=59)

Rio Peje (n=17/20)

> RE Certo Cocorí (n=19)
Barbilla (n=2/5) Finca Montaña Grande (n=4)
A Río Aguas frías (n=3)
Uatsi (n=6/5) E]
= Isla Uvita (n=2)
py) jaa >

Fey >

Cahuita (n=63/51)

Manzanillo (n=10/19)

Hitoy Cerere (n=13)
y fs Sn A u

Hitoy Cerere - Miramar (n=18)
ss]

Amubri (n=23)
|

Manzanillo (n=4)
У fa

—_—

1H 1 1 1 1 1 1 1
8 9 10 11 12 13 [mm] 14

FIG. 23. Minor diameter of shell of different
populations of Helicina funcki in Costa Rica
according to Table 3; for explanations see Fig. 21.

0 8 9 10 11 12 13 [mm] 14
FIG. 24. Minor diameter of shell of different
populations of Helicina funcki in Costa Rica,
INBio collection, according to Table 4, and type
material of Helicina funcki and Helicina funcki
costaricensis; for explanations see Fig. 21.

CLASSIFICATION OF HELICINIDAE 235

prisingly well in their mean values, although
this is often not supported by very extensive
data in my own material or INBio's, respec-
tively. This even agrees with the results of
the analysis of “non-statistical” numbers of
specimens.

Following up the fact that the highest locali-
ties always had the small-shelled popula-
tions, the average minor diameter of the 34
populations (Isla Uvita excluded) was plotted
against the elevation (Fig. 31). A constant
decline of the maximum values with increas-
ing elevation 1$ clearly visible, suggesting an
influence by elevation. Furthermore, all the
data scattered below this decline in maxi-
mum size indicate that altitude is not the
only important parameter. The range of pos-
sible influences is too wide, and Helicina
funcki, although widely distributed, occurs at

Rincón de la Vieja (n=4/4)

Mirador Gerardo (n=4/4)

El =

very scattered locations so that very detailed
and local studies of environmental condi-
tions would be required to trace any further
correlation.

Sexual Dimorphism: Besides weight, females

in all measurements and in all populations
are clearly bigger than males, although the
range of measurements overlaps widely, as
exemplified for Cahuita by the minor diam-
eter-height-relation (Fig. 32). Even with a
small sample size of sometimes as few as
two individuals, this result is always con-
firmed. Furthermore, the range of the differ-
ences is often about the same as in the
more extensively supported data of the
populations from Cahuita, Rio Peje and
Monteverde. The males have a volume of
about 81% of females.

Rincon de la Vieja (n=4/4)

+0

= o
Mirador Gerardo (n=4/4)

—

Monteverde - Finca Ecológica (n=4/3)

Monteverde (n=18/15)

Las Pavas (n=7/5)

Tortuguero (n=3/2)

La Selva (n=9/9)

E E A

Guayacán (n=5/3)
y a

México (n=4/6)

Monteverde - Finca Ecológica (n=4/3)

Monteverde (n=18/15)

a
Las Pavas (n=7/5)
SEE
Tortuguero (n=2/2)
de

jas (n=9/9)

AS (n=5/2)

México (n=4/6)

Río Peje (n=17/20)

Barbilla (n=2/5)

Río Peje (n=17/20)

Barbilla (n=2/5)

=
Uatsi (n=6/5) Uatsi (n=6/5) = —
Shiroles (n=2/4) Shiroles (n=2/4)
Fr (n=63/51) Cahuita (n=63/51)
zz
Manzanillo (n=10/19) Manzanillo (n=10/19)
0 7 6 7 8 9 10 11 [mm] 12 ù т 7 8 y 10 11 [mm] 12

FIG. 25. Height of last whorl of different
populations of Helicina funcki in Costa Rica
according to Table 3; for explanations see
Eig: 21.

FIG. 26. Height of columellar axis of different
populations of Helicina funcki in Costa Rica
according to Table 3; for explanations see
Е 21:

Rincôn de la Vieja (п=4/4)

Q, +0

Mirador Gerardo =

236 RICHLING
Rincón de la Vieja (n=4/4) 9
м
| Mirador Gerardo (n=4/4)
>
| Monteverde - Finca Ecolögica (n=4/3)
|
—

Monteverde - Finca Ecológica (n=4/3)

Monteverde (n=18/15)

Monteverde (n=18/15)

Las Pavas (n=7/5)

Las Pavas (n=7/5)

Tortuguero (n=3/2)

Tortuguero (n=3/2)

ER

La Selva (n=9/9)

La Selva (n=9/9)

| yp (n=5/2) Guayacan (n=5/2)
|
México (n=4/6) México (n=4/6)
SN SES
Río Peje (n=16/20) Río Peje (n=17/20)
Barbilla (n=2/5) Barbilla (n=2/5)
| = —
Uatsi (n=6/5) Uatsi (n=6/5)
—— nn

Shiroles (n=2/4)

Cahuita (n=60/51)

Shiroles (n=2/4)

a

Cahuita (n=63/51)

Manzanillo (n=10/19)

1 alt 1 1 1 1 1 1
0.5 0.6 0.7 0.8 [ml] 0.9

FIG. 27. Shell volume of different populations of
Helicina funcki in Costa Rica according to Table
3; for explanations see Fig. 21.

Manzanillo (n=10/19)

ES

1 1 4 1

0.25

0 0.05 9 035
FIG. 28. Shell weight of different populations of
Helicina funcki in Costa Rica according to Table

3; for explanations see Fig. 21.

TABLE 4. Minor diameter measurements [тт] of different populations of Helicina funcki from the
INBio collection and type material, given as mean value with standard deviation, minimum and
maximum value (min, max), and number of specimens.

Mean Devi-
Locality value ation Min Max
Santiago (type lot 10.24 0.22 9.95 10.67
ВММН)
Santa Clara (type lot Н. 11.84 0.02 11.82 11.86
funcki costaricensis)
Estación Góngora 11.42 0.34 10.80 11.85
Estación San 11.09 0.38 10:26 11.68
Cristóbal
Rincón de la Vieja 10.95 0.47 10.34 12.27
Tenorio 11.37 2017220 NA
Santa Elena 10:81 0.30 110.18: 11:48

Number Lots
4 BMNH 20010497.1-4
2 MIZ 8989
6 INBio 1480300, 1480475, 1483409,
1484993, 1488083
12 INBio 1488065, 1498494
6 INBio 1466644, 1487945, 1498739,
1498744
2 INBio 1485411, 1498593
8 INBio 1498638

(Continues)

CLASSIFICATION OF HELICINIDAE 237

(Continued)

Mean Devi-
Locality value ation Min Max Number Lots

Peñas Blancas 10.82 0.44 10.45 11.48 3 INBio 1480605, 1498802

Monteverde 11.13. -0:45 989 12.03. 92 INBio 1466835, 1466842, 1466863,
1466870, 1466884, 1466891,
1466905, 1466912, 1466954,
1467003, 1467010, 1467031,
1468211, 1477521, 1477749,
1479517, 1479528, 1479539,
1479550, 1480098, 1480119,
1480126, 1480127, 1480128,
1480129, 1480130, 1480131,
1480132, 1480149, 1480152,
1480426, 1484687, 1485422,
1485426, 1485441, 1498581,
1498590, 1498632, 1498804,
1498806, 1498807, 1498828

Pitilla 11.94 049 11.22 13.08 8 INBio 1463787, 1463946, 1480043,
1480289, 1480318, 1480319

Monte Cele 10:91 0.22 10.58 11.26 = INBio 1488042

Estacion Playuelas 10.49 044 953 11.32 7 INBio 1479506, 1487809, 149857 1

Caño Negro 12.20 0.47 11.18 12.78 8 INBio 1466940, 1480029, 1487043,
1487611, 1487878, 1501040

Barra del Colorado (239% 2035 1109" 1397 53 INBio 1465700, 1477915, 1478017,
1478283, 1478294, 1480041,
1480051, 1484010, 1484013,
1484372, 1484374, 1484585,
1484587, 1484589, 1484748,
1484749, 1484991, 1485145,
1485284, 1485285, 1485289

Cerro Cocori 12.67 0.34 11.98 13.25 19 INBio 1465446, 1467174, 1478061,
1480255, 1480261, 1483017,
1483208, 1483360

Finca Montana 1308 033 233 14.15 4 INBio 1498610, 1501098
Grande
Rio Aguas frias 13.16 0.50 12.41 13.66 3 INBio 1487980
Isla Uvita 895 0.09 8.86 9.03 2 INBio 3315386
Hitoy Cerere 12.72 .-0.44. 1182 1355 13 INBio 1463392, 1466444, 1473832,

1475438, 1476246, 1476262,
1497862, 1497905, 3091789

Hitoy Cerere - 12.13 0.39 11291 13:81 1% INBio 1475234, 1475694, 1475720,
Miramar 1475725, 1475930, 1476376,
1476490, 1476687, 1476688, 1480272
Amubri 11.192 0.28 1032 1181 24 INBio 1467294, 1477569, 1477585,

1483302, 1483381, 1483382,
1483386, 1483387, 1483389,
1483390, 1483392, 1483394,
1483398, 1483400, 1483407,
1485365, 1485382, 1493444
Manzanillo 196: 0:39) 11:08; 1233 4 INBio 3097895, 3097906

238

RICHLING

Tortuguero

La Selva

Monteverde - Finca Ecolégica
Monteverde

E =
Rio . .
Guayacan, aes à +.
= México\ ~
és

Cahuita
Uatsi ~\Manzanillo

Shiroles

Г

FIG. 29. Size variations in Costa Rican populations of Не/ста funcki: shell
height in figures reflects the mean value of the respective females; each shell
originates from the respective locality and is randomly chosen according to
the approximation of the mean value.

height >
minor diameter +

0.3 0.4 0.5 0.6 0.7 volume [mi] 0.9

FIG. 30. Relation of shell height and minor diamater respectively to the volume
in Helicina funcki exemplary for the females of the population from Cahuita.

CLASSIFICATION OF HELICINIDAE 239

T T
material leg. RICHLING o
min. material INBio +

er:

o SE
ne + o
o —
1 1 - le ——
0 200 400 600 800 1000 elevation [m] 1600

FIG. 31. Relation of minor diameter of shell to elevation of locality of different
populations of Helicina funcki in Costa Rica; sex-independent mean values
were used.

female +
male +

9 10 11 12 height [тт] 14

FIG. 32. Range of measurements in females and males exemplary for height
and minor diameter in the population from Cahuita.

240 RICHLING

0.4 ==="

female >

weight | Cahuita male +
[9]

0.3 r
+

0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 volume [ml] 0.9

FIG. 33. Relation of weight to volume in females and males of the population
of Helicina funcki from Cahuita (material according to Table 3).

female >
weight Manzanillo male + |
[9]
0.3 4

0.25

0.2

0.15

0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 volume [ml] 0.9

FIG. 34. Relation of weight to volume in females and males of the population
of Helicina funcki from Manzanillo (material according to Table 3).

CLASSIFICATION OF HELICINIDAE 241

0.4 T T T Ii T T
female >
weight Río Peje male +
[9] A j
o
E o
0.3 : a ® |
E
© ©
+t + > 5%
0.25 F pa Se -
+ o o $
+ + N |
A
0.2 +
г
0.15 + -
©
ri
0.1 Al
0.05 4
0 1 1 E AAA AE
0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 volume [ml] 0.9

FIG. 35. Relation of weight to volume in females and males of the population
of Helicina funcki from Rio Peje (material according to Table 3).

т T ann ГИС |
female >
weight La Selva male + |
[9]
0.3 + -

0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 volume [ml] 0.9

FIG. 36. Relation of weight to volume in females and males of the population
of Helicina funcki from La Selva (material according to Table 3).

242 RICHLING

0.4
female >

weight Monteverde male +
[9]

0.3

0.25

0.2. |-

0.15 F

0.1

0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 volume [ml] 0.9

FIG. 37. Relation of weight to volume in females and males of the population
of Helicina funcki from Monteverde (material according to Table 3).

0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 volume [ml] 0.9

FIG. 38. Relation of weight to volume in adults of Helicina funcki in Costa Rica
(all material listed in Table 3 included).

CLASSIFICATION OF HELICINIDAE 243

Considering the problems connected with
the weight measurements, the data (Fig.
28) indeed show contrary results even in
otherwise rather similar populations (e.g.,
upper rows: Rincon de la Vieja to
Monteverde). When only looking at the bet-
ter-supported data — namely Monteverde,
La Selva, Rio Peje, Cahuita, Manzanillo —
the difference between males and females
is greater in the population at Monteverde
than in La Selva, Rio Peje and Manzanillo,
whereas at Cahuita this relationship is even
reversed. Taking into account the greater
size of all females, it seems that in the low-
land populations the males invest more
material in their shells than females of the
same volume do. To test this assumption,
the relation of weight to volume is plotted
for these populations (Figs. 33-37). Be-
cause the mathematical relation between
weight and volume is unknown and males
and females fall into a different range, the
data of all specimens of Helicina funcki
used in the morphometric analysis were
plotted as an adjustment for comparison
(Fig. 38). As expected, the latter measure-
ments are widely scattered, but a linear
approximation or a function of greater de-
gree would better match the data. A higher
volume should result in a higher weight.
The Monteverde population clearly demon-
strates this relation for both sexes (Fig. 37),
whereas the males of the lowland popula-
tions on average weigh the same as fe-
males with a higher volume (Figs. 33-36).
For other localities, the deviations ex-
plained above seem to be interposed with
the actual results.

Habitat

Biolley (1897) found the species on the
trunks of trees, the stems of plantains (Musa)
and also on the ground. Except for the last
habitat, these observations could be con-
firmed during the field work for this study.
Moreover, Helicina funcki often crawls and
aestivates on the underside or more seldom
on the upper side of different kind of leaves.
The recognized plants belong not only to
Musaceae, Heliconiaceae, and palms, but
also to various herbs of the undergrowth.
Helicina.funcki may even be found on climbing
species such as the Araceae Monstera spec.
Probably because of the relatively large size
of the species, it is found on plant species with

large leaves. But it lives on trunks, branches
and twigs of trees, bushes and tree ferns as
well. Helicina funcki not only crawls on live leaf
surfaces, it was also found in the dead, dried
and curled-up leaves, especially those of ba-
nanas. In areas of human influence speci-
mens were observed on concrete walls of
buildings or wooden fences. Thus, H. funcki is
a typical arboreal species, having been ob-
served up to 7 т or more above the ground.
With regards to alimentation, it was definitely
found feeding on the surface of trunks and on
living and dead leaves.

Distribution

Helicina funcki is confined to southern Central
America. Although for Nicaragua it has thus far
only been recorded by Ancey (1897) from
Greytown at the mouth of the Rio San Juan,
with a further unspecified lot in the collection of
the UF and a site somewhere along the south-
ern border of the Rio San Juan (von Martens,
1901), the distribution range extends at least
from southern Nicaragua to the Canal Zone in
Panama. It most probably occurs in the eastern
Caribbean lowlands further north in Nicaragua
as well, because the habitats do not change
greatly. Furthermore, the wide distribution in
northern Costa Rica and the morphometric
data suggest that in this area H. funcki has not
come close to its distribution limit.

Due to the lack of literature records for the
better investigated countries, such as Hondu-
ras, Guatemala and Belize, and due to the
absence of H. funcki in the extensive Central
American collection of the UF (checked per-
sonally) any occurrences north of Nicaragua
can be excluded. The relatively large size fur-
thermore renders the species unlikely to be
overlooked. The records from Ylalag (Mexico:
Oaxaca) by Wagner (1910a) therefore seems
very questionable.

In Costa Rica, the species is fairly widely
distributed throughout the Caribbean plain
and on the mountain slopes (Fig. 39). The dis-
tribution is mainly influenced by the central
mountain chains subdividing the country.
Helicina funcki crosses the northern volcanic
mountains (Cordillera de Guanacaste and
Tilaran, Cordillera Central), where the upper
Pacific slopes are connected to the Caribbean
side by various valleys between the separate
volcanoes. According to the present data, the
species is known to occur up to 1,800 т. А
limitation by altitude is furthermore supported

244 RICHLING

by the decline of the shell size with increasing
elevation of the localities. In fact, the southern
Cordillera de Talamanca, highly elevated as a
continuous mountain chain (approximately
3,000 m), forms a clear barrier in the distribu-
tion of H. funcki. The exact occurrence on the
Caribbean slope of this Cordillera is known
only fragmentarily because the area is difficult
to reach and has not been investigated. Con-
{пита downhill towards the northern Pacific
and in the Valle Central the climate becomes
drier (Figs. 2, 3), therefore appearing to be the
most important factor limiting the distribution.
Except for the most southern Caribbean plain,
H. funcki does not occur in areas of less than
2,000 mm annual precipitation. On the more
humid southern Pacific plains and slopes, H.
funcki is replaced by H. pitalensis.

The single record of H. funcki on the
Península de Osa (INBio 1486976) seems to
contradict the otherwise continuous distribu-
tion. Upon request, the data were confirmed
by INBio. The specimen is small (9.4/12.1/9.5

mm). There 1$ no reason to question the find-
ing, despite the fact that several collecting ef-
forts of INBio up to now have yielded only one
specimen, because species of Helicinidae are
extremely rare on Península de Osa.

Biolley in 1897 reports the species as the
most common land snail of the country. Nowa-
days due to the extreme change of the land
use (e.g., deforestation in large areas), it prob-
ably will be shown that synanthropic snails like
Subulina octona (Bruguière, 1789), the intro-
duced Ovachlamys fulgens (Gude, 1900)
(Barrientos, 2000) and Succinea costaricana
von Martens, 1898, the latter known as pest
species in agriculture (Villalobos et al., 1995),
are now much more common.

Discussion

The differences of Helicina funcki
costaricensis to the nominal species men-
tioned by Wagner (1910a) can be summarized
as differences in size and in a more strongly

T

1

3500 - 4000 m

3000 - 3500 m
2500 - 3000 m
2000 - 2500 m
1500 - 2000 m
1000 - 1500 m

500 - 1000 т
100 - 500 m
0-100m |

|
1

e coll. IR
| e coll. INBio |
| e others

FIG. 39. Records of Helicina funcki in Costa Rica.

CLASSIFICATION OF HELICINIDAE 245

developed outer lip. The original description of
the subspecies does not include any compari-
son With H. funcki. Astonishingly, Wagner
(1910a) gives higher dimensions (12-15/15-
18/12-14 mm) than in his publication in 1905
(11.0/13.3/11.3 mm), which on one hand
clearly are exaggerated, on the other hand the
indeed higher values from specimens from
San José were most likely included in his
measurements. The morphometric investiga-
tions of different populations of H. funcki sug-
gest that the size depends on environmental
factors and 1$ not suitable for the separation of
a subspecies in absence of other differentiat-
ing characters. The type lots of both the nomi-
nal form and the subspecies fall in the range
of the Costa Rican specimens (Fig. 24).
Helicina funcki reaches maximum sizes in
lowlands to which the type locality of H. funcki
costaricensis belongs. The nominal species is
described from close to the southern limit of its
distribution (Fig. 39), which makes it likely that
environmental conditions of this area are less
favorable for the species, such as perhaps at
high elevations, which may result in smaller
shells. Therefore, H. funcki costaricensis 1$
regarded as a synonym of H. funcki. Regard-
ing the comparably large shells mentioned by
Wagner (1905, 1910a) (checked: MIZ 8990: 2
ads.: 13.4/16.0/13.2 mm; 13.9/15.7/12.9 mm)
from San José, the locality given without com-
ment is misleading, because one would imme-
diately think of San José, capital of Costa Rica
(in the historical times of Wagner a possible
locality). But considering the relation of shell
size to elevation of the locality (Fig. 31), the
site appears to be in contradiction to the shell
size because San José 15 located at about an
altitude of 1,160 m. Biolley (1897) reported a
small form of H. funcki from Cartago (close to
San José, at a similar elevation). A closer ex-
amination of the map of Costa Rica reveals a
second San José in the Alajuela Province,
close to Santa Clara, which is here suggested
to be the locality “San José”. It also supports
the localization of “Santa Clara”. Under H.
funcki costaricensis, Wagner (1910a) men-
tions a somewhat dubious form from Ylalag,
Mexico in his collection, which is said to be
more elevated and remarkably angulated at
the periphery. It could not be checked and
therefore cannot be discussed any further,
especially because of the outstanding locality
for Helicina funcki.

The description of Helicina deppeana
parvidens has to be discussed in the context

of material in the ZMB. Helicina deppeana von
Martens, 1863, was described from Mexico
(locality unknown) and was figured later (von
Martens, 1865, 1890) together with a variety
from Yalalag (State of Oaxaca, Mexico). The
study of the original material of the figures
stored in the ZMB revealed the following: the
typical H. deppeana (syntypes ZMB 4571) are
not conspecific with H. funcki, for example,
they do not have the typical ornamentation of
lighter patches, and are more solid and
unicolored. The specimens of the variety from
Yalalag (ZMB 1743) look exactly like speci-
mens of H. funcki and thus are specifically dif-
ferent from H. deppeana, an observation
already remarked on the label by Wagner:
“nach meiner Ansicht stellen die vorliegenden
Exemplare nur Helicina funcki dar” [= in my
opinion the specimens only represent H.
funcki]. Interestingly, Wagner (1910a) сот-
pletely avoids any comment on this in his
monograph, although it is certain that he had
seen the collection prior to his publication,
because various types of newly described
species (e.g., H. pitalensis, H. tenuis pittieri)
are in the ZMB collection. The singular Mexi-
can locality of H. funcki is discussed in the
paragraph “Distribution”. Returning to H.
deppeana parvidens, it is very likely that when
Pilsbry (1920a) published his work on Costa
Rican land molluscs, he used the Biologia
Centrali-Americana (von Martens, 1890-
1901), representing the only comprehensive
contribution for the area even today. It there-
fore appears probable that Pilsbry was misled
by this figure and classified part of his Costa
Rican material of H. funcki as a new subspe-
cies of the Mexican H. deppeana.

Helicina (Tristramia) pitalensis
Wagner, 1910

Helicina funcki — von Martens, 1900: 603-604:
Costa Rica: SW-Costa Rica: Bay of Terraba
[mouth of Rio Terraba, about 09°00’М,
83°36’W, Puntarenas Province], Tocori in the
valley of the Rio Paquita [NE of Quepos,
canton Aguirre, 09°29'43"N, 84*04'52"W, 10
m a.s.l., Puntarenas Province], middle part
of the Rio Saveque [now: Rio Savegre,
about 09°29’N, 83°56’W, San José Prov-
ince] and lower part of the Rio Pacuare [now
Rio Pacuar south of San Isidro de El Gen-
eral [not Rio Pacuare on Atlantic slope!],
about 09°16’N, 83°39’W, San José Prov-
ince] (Pittier); El Pital, in the valley of the Rio

246 RICHLING

Naranjo [near Londres? (about 09°27’М,
84°05’W, Puntarenas Province], some
specimens banded and others more el-
evated (Pittier) [in part] [поп L. Pfeiffer, 1849]

Helicina pitalensis Wagner, 1910a: 308, pl. 61,
figs. 17-19

Helicina amoena — Monge-Nájera, 1997: 113:
Costa Rica [non L. Pfeiffer, 1849]

Original Description

“Gehause kegelfórmig mit gewólbter Basis,
festschalig, leicht glánzend, zitrongelb mit
undeutlichen weissen Flecken und Punkten,
sowie einer schmalen rotbraunen Binde úber
der Naht und dem Kiel. Die Skulptur besteht
aus feinen, etwas ungleichmássigen Zu-
wachsstreifen, auch erscheint die Epidermis
unter der Lupe sehr fein gerunzelt. Das
regelmässig spitzkegelförmige Gewinde
besteht aus 5-5'/, leicht gewölbten, langsam
zunehmenden Umgängen, welche durch eine
hell berandete, schwach eingedrückte Naht
geschieden werden; der letzte ist beiderseits
gleichmässig gewölbt, an der Peripherie
deutlich kantig bis stumpf gekielt und steigt
vorne nicht herab (unmittelbar vor der
Mündung ein wenig hinauf). Die abgerundet
dreieckige Mündung ist schief, innen gelb mit
durchscheinender Binde. Der leicht ver-
dickte, gelbliche Mundsaum erweitert; der
Oberrand schmal und an der Insertion
vorgezogen, der Aussen- und Basalrand breit
umgeschlagen. Die kurze, abgerundete
Spindel ist senkrecht oder leicht nach links
gebogen; am Uebergange derselben in den
Basalrand der Mündung eine zahnartig
vorspringende Ecke. Der sehr dünne, fein-
gekörnelte Basalkallus nur im Umkreise der

Spindel deutlich. Das Grübchen
Nabelgegend undeutlich.

D = 14, d = 11,5, H = 13,5 mm.
Deckel birnförmig mit seitlich gekrümter [sic]
Spitze schwarzbraun bis pechschwarz mit
lichterem Streifen entlang der Sigmakante; die
dünne, feingekörnelte Kalkplatte nur am
Spindelrande etwas leistenartig verdickt; in
den übrigen Verhältnissen typisch.

Fundort: El Pital im Tale des Rio Naranjo im
südwestlichen Costarica. Da abgebildete Ex-
emplar im k. Museum zu Berlin.

Von der ähnlichen Helicina funcki Pfeiffer
unterscheidet sich vorstehende neue Art
durch die lebhafte Färbung mit deutlicher
Binde, die glänzende Oberfläche mit deut-
licheren Zuwachsstreifen, das höhere
Gewinde mit deutlich gewölbten langsam und
regelmässig zunehmenden Umgängen, den
weniger erweiterten, aber deutlich kantigen
bis stumpfgekielten letzten Umgang, sowie
besonders die abweichenden Verhältnisse der
Mündung und des Mundsaumes.”

in der

Type Material

ZMB 103240 “El Pital, 200 m, 111.1893,
Vallée du Rio Naranjo, leg. Madame Pittier de
Fahega” (the lot contains one specimen)
Because the original description refers to one
specimen in the ZMB which matches the fig-
ure, it is the holotype (Fig. 40).

Dimensions:
Holotype: 13.0/12.5/14.0/11.2/9.1/10.6/10.0 mm

Type Locality

“El Pital im Tale des Rio Naranjo im
südwestlichen Costarica”; El Pital could not be

FIG. 40. Helicina pitalensis, holotype, ZMB 103240, height 13.0 mm; scale bar 5 mm.

CLASSIFICATION OF HELICINIDAE 247

localized on recent detailed maps. The Rio
Naranjo leads into the Pacific Ocean at the
southern border of the Parque Nacional de
Manuel Antonio, a little south of Quepos. By
the elevation given in the data remaining with
the original material, the type locality can be
assumed to be located near Londres [about
09°27’N, 84°05’W], Puntarenas Province.

Examined Material

Lee. |. RICHLING

Puntarenas: S San Vito, Wilson Botanical
Garden, Las Cruces, sendero a Rio Jaba,
08°46’57"N, 82%57'40"W, 1,160 т a.s.l.,
27.08.1999: (IR 1013); 28.08.1999: (IR
1016)
N Neily, road from Ciudad Neily to San Vito,
open area with a few trees, 08°40’23"М,
82°56 44"W, 180 m a.s.. М Neily,
23.03.1997: (IR 209)
Fila de Cal, road from Ciudad Neily to San
Vito, S Campo Dos, burned area,
08°41'00"N, 82°56 29"W, 630 m a.s.l.,
23.03.1997: (IR 191)
Fila Costeña, north of Bajo Bonito (locally
called Llano Bonito), N of Río Claro, rain for-
est, 08°44'41"М, 83°02’09"W, 980 m a.s.l.,
24.03.1997: (IR 221); 15.02.1999: (IR 579);
29.08.1999: (IR 1028); 06.03.2001: (IR
1485)

INBio COLLECTION
Puntarenas: Parque Nacional Corcovado:
Estación Sirena, 08°28'52"N, 83*35'32"W,

5 m as!l: leg. Mario Chinchilla,
23.03.1995: 1 juv. (INBio 1485050);
Sendero los Espaveles, Sirena,

08°28'49"N, 83°35’42"W, O т a.s.l.: leg.
Annia Picado, 25.03.1995: 1 ad. (INBio
1482837); 1 ad. (INBio 1482842); Sendero
Espaveles, 08°29'05"N, 83°35’29"W, 0 т
a.s.l.: leg. Socorro Avila, 23.03.1995: 1 ad.
(INBio 1482627); Sendero Espaveles,
08°29’22"N, 83°35'14"W, 0 m a.s.l.: leg.
Billen Gamboa R., 03.12.1995: 1 ad. (INBio
1485173); Estación Sirena, Sendero Las
Ollas, 08*28'47"N, 83*35'40"W, 5 m a.s.l.:
leg. Alejandro Azofeifa, 25.03.1995: 1 juv.
(INBio 1484670); Estación Sirena, Sendero
Las Ollas, 08*28'57"N, 83°35’20"W, 20 т
a.s.l.: leg. Francisco Alvarado, 25.03.1995:
1 s.ad. (INBio 1484221); Sendero los
Patos, 3.5 km al N. de la Estación Sirena,
08°30’46"М, 83°35’56"W, 0 т a.s.l.: leg.
Ramon Angulo, 26.08.1994: 1 juv. (INBio

1480506); Rio Pavo, 08°30'51"N,
83°35'44"W, 20 т a.s.l.: leg. M. Madrigal,
03.04.1996: 2 ads. (INBio 3542542)
Reserva Forestal Golfo Dulce: Cerro La
Torre, Finca La Purruja, 08°32’04"М,
83°25'53"W, 400 m a.s.l: leg. Javier
Quesada, 05.05.1994: 1 s.ad. (INBio
1477485); Agujas, alrededores de la
estación, 08*32'13"N, 83°25’33"W, 300 m
a.s.l.: leg. A. Berrocal, 01.11.1998: 1 ad.
(INBio 3397130)

Fila Cal: 24 km de San Vito hacia Ciudad
Neilly, 08%41'36"N, 82°56’36"W, 780 т
a.s.l.: 29.08.1995: 1 ad. (INBio 1485456);
29.08.1995: 1 s.ad., 1 juv. (INBio 3121204)
(all leg. Marianella Segura); 24.5 km S en
la carretera de San Vito hacia Ciudad
Neilly, 08*40'55"N, 82°56’23"W, 600 т
а.5.1.: leg. Zaidett Barrientos, 21.11.1995: 1
juv. (INBio 1485120)

45 km NW de Ciudad Neily, Camino
Paralelo al Río Caño Seco, Colectado en
hojarasca en helechos, 08°40’50"М,
82°57'25"W, 180 т a.s.l.: leg. М. Chinchilla,
22.11.1995: 1 ad. (INBio 3542525)

Linda Vista, Río Claro: 3 km NE de la
Escuela de Llano Bonito, 08°44’54"М,
83°02'04"W: 920 m a.s.l., leg. Socorro
Avila, 24.03.1997: 1 s.ad., 1 juv. (INBio
1494393); 950 m a.s.l., leg. Alexander
Alvarado Mendez, 15.02.1999: 1 s.ad., 1
juv. (INBio 3091134)

OTHER Sources

COSTA RICA

Alajuela: La Paz, Chemin du rivière Sarapiqui
[not localized, near Isla Bonita?, about
10%15'30"N, 84°11’W], Biolley, ex Godet,
12.1892 received: 1 ad. (ZMB 45501)

Description

Shell (Fig. 335D-E): Conical-subglobose,
solid, relatively large, slightly shiny to dull.
Color: basic color lemon yellow, sometimes
less bright, with slender reddish-brown band
between sutures or suture and the periphery
respectively, in some specimens very light or
obsolete. On account of this band, the upper
whorls may appear darker. The periphery is
always lighter. As in Helicina funcki, the color
is overlapped by fine white patches and lines
giving the shell a special ornamentation.
Surface textured with fine, irregular growth
lines and oblique grooves of different indi-
vidual orientation but of same general direc-

248 RICHLING

FIG. 41. Axial cleft and muscle attachments of

Helicina pitalensis, IR 579; scale bar 5 mm.

FIG. 42. Teleoconch surface structure of
Helicina pitalensis. A. On 2" whorl. В. On
4" whorl; scale bar 100 um.

FIG. 43. Embryonic shell of Helicina pitalensis;
scale bar 100 pm.

tion (Fig. 42), causing the dull appearance.
Embryonic shell with about 1 whorl; 4'/,-4°7,
subsequent whorls slightly convex; periph-
ery remarkably angulated; whorls equally
extending in size and slightly descending,
only towards aperture slightly ascending;
spire very regular. Suture slightly impressed
and marginally lighter in color. Aperture ob-
lique and nearly straight, inserting a little
above periphery. Outer lip yellowish-whitish,
very thickened, broadly expanded, only in
the upper palatal part a little less strongly
developed. Reflection nearly rectangular to

FIG. 44. Operculum of Helicina pitalensis, IR
579; scale bar 2.5 mm.

CLASSIFICATION OF HELICINIDAE 249

the whorl; transition to columella with a ге- Juveniles are roundly angulated, and in
markably protruding denticle. Columella some cases a few rows of periostracal hairs
short, slightly curved, umbilical area without are present at the periphery.

any groove or impressed line. Basal callus

only close to columella present, thin, slightly Internal Shell Structures: (Fig. 41)

granulated.

Teleoconch Surface Structure (Fig. 42): The
transitional structure is developed, but as in
Helicina funcki the pattern of oblique diverg-
ing grooves continues up to the aperture. In
the shell illustrated, the grooves remain finer
than in Я. funcki (see 4" whorl), but this as-
pect is subject to individual variation.

Embryonic Shell (Fig. 43): The embryonic
shell of Helicina pitalensis is very similar to
that of Helicina funcki. The specimens mea-
sured came from altitudes of nearly 1,000 m.
In comparison with the H. funcki-popula-
tions, the intermediate size therefore sug-
gests equal dimensions, assuming a similar
dependence on altitude.

Diameter: 1,089 um (+ 23) (1,040-1,150)
(n = 10) (IR 579, IR 1013, IR 1028, IR 1485).

Operculum (Fig. 44): Slightly calcified, calcar-
eous plate leaving a free margin, thickened
towards columellar side. Color dark reddish-
brown to even black, only non-calcified mar-
gin transparent. Columellar side regularly
S-shaped, upper end acute and pointed,
lower end well rounded.

Animal (Fig. 337C): Only specimens that were
very similarly colored from Bajo Bonito were
studied. The foot is whitish-yellowish

€ NE | Ñ
ES we NS
ia \ / Ss LN
£ > BR )
у \ a À } )
Es 7 N
17 NUN \ ir)
MS» | \ = a
E] N SIA
| E | N LR Я \
EA = U
CT
a <Q a a
м SET
<<)
FIG. 45. Radula of Helicina pitalensis. А. FIG. 46. Female reproductive system of Helicina
Centrals. В. Comb-lateral. С. Marginals; pitalensis, apical complex enlarged, IR 579;

scale bars 50 pm (A, B), 100 pm (C). scale bars 2 mm (left), 1 mm (right).

250 RICHLING

Bajo Bonito (n=2/3) 9
— ©”
Rio Naranjo (type)
ff :
San Vito (n=3)
Я = —=_ о
Fila de Cal (n=4)
ЧЕ nn
Beh ACE all (n=8)
———
Peninsula de Osa (n=6)
JE mm

L | A 1 1 1 1
0 8 9 10 11 12

13 [mm] 14

FIG. 47. Shell height of different populations of
Helicina pitalensis in Costa Rica, according to
Table 5; on each line: mean value, standard
deviation, absolute range; number of individuals
given as “п = females/males or total”; upper line:
females, lower line: males if separate; in
between and shaded: average of both for
comparison with populations of unknown sex.

throughout, the head region, especially
around the black eyes and occasionally the
upper part of the snout, is distinctly white.
The tentacles become gradually darker to-
wards their tips. The mantle has a whitish
pigmentation.

Radula (Fig. 45): Only two specimens were
investigated. The cusps on the A- and C-
central are vestigial, only the B-central bears
5 to 9 cusps. Comb-lateral with 6-8 cusps,
cusps on marginals slowly increasing in
number. Radula with about 95-99 rows of
teeth.

Female Reproductive System (Fig. 46): Only
three female specimens were available for
dissection. The pallial reproductive system
closely resembles that of Helicina funcki.

Bajo Bonito (n=2/3)

rt ?
+ o
Río Naranjo (type)
ТЕ I
San Vito (n=3)
Fila de Cal (n=4)
Bajo Bonito, all (n=8)
Peninsula de Osa (n=6)
a] faz —
1H 4 1 1 L 1 1 1
0 8 9 10 11 12 13 [тт] 14

FIG. 48. Minor diameter of shell of different
populations of Helicina pitalensis in Costa Rica
according to Table 5; for explanations see Fig. 47.

The bursa copulatrix differs in that it is as
elongated as the whole organ, and the lobes
project from a central axis and are much
shorter, similar to those in the other species,
although they are occasionally further subdi-
vided. The provaginal sac seems to be
smaller than in Helicina funcki.

Morphometry and Sexual Dimorphism

Despite the number of lots of Helicina

pitalensis, the material for morphometric
analysis is scant because of a high proportion
of juvenile shells.
- Individuals from the lowlands of the
Península de Osa differed from the typical
specimens in having a more prominent aper-
ture. The few individual records summarized
as “Península de Osa” originate from the
same region near Estación Sirena and were
compared to populations from “San Vito”,
“Bajo Bonito” and “Fila de Cal”, which are lo-
cated close to each other in the mountainous
country on the southern Pacific side (Fig. 52).
The only specimens whose sex was deter-
mined belong to the population of Bajo Bonito.
Any comparisons in this species can only hint
at possible tendencies because of the scanty
data.

Morphometry: The different populations show
remarkably little differences in their minor
diameter (Table 5, Fig. 48), which therefore
provides a good reference for the pattern of
differences among the populations for other
measurements (Figs. 47, 49-51). The popu-
lation “Península de Osa” displays the high-
est deviations; the shells are relatively less
elevated in every respect (height, height of
last whorl and columellar axis), but the aper-
ture and outer lip is much more expanded.
This confirms the observations noted above
(Fig. 52). In general, the “Bajo Bonito” popu-
lation best matches the type in proportions
and size. This excludes a correlation of the
differing shell shape of the “Península de
Osa” specimens to the altitude, because the
type lot also originates from lowlands (200
m) whereas the other sites are located at
700 to 1,160 m (San Vito). Thus, the special
shell shape of the population “Península de
Osa” seems to be a local peculiarity. Fur-
thermore, current data do not support a cor-
relation of shell size and altitude as for
Helicina funcki.

CLASSIFICATION OF HELICINIDAE

251

TABLE 5. Measurements of different populations of Helicina pitalensis given as mean value with
standard deviation, minimum and maximum value (min, max), and number of specimens; only
population in last column separated in females and males, these individuals are also included in “Bajo
Bonito, all” (min./max. diam. = minor/major diameter, col. axis = columellar axis); linear measurements
[mm], weight [9], volume [ml].

Height
Maj. diam.
Min. diam.
Outer lip
Last whorl
Col. axis

Height
Maj. diam.
Min. diam.
Outer lip
Last whorl
Col. axis

Mean
value

12.48
12.34
11.04
8.46
10.02
9.45

“Bajo Bonito, all” (altitude 920-980 т)
lots IR 221, IR 579, IR 1028, IR 1485

Mean

value Deviation Min

12.64
12.49
11.13
8.51
10.21
9:50

“San Vito” (altitude 1160 m)

“Fila de Cal” (altitude 600-780 m)
lots IR 191, 209, INBio 1485456, INBio

lots IR 1016

Mean

Deviation Min Max Number value
0.47 11.78 12.89 3 1242
0.87 1179 12.65 3 12.31
0.34 10.53 11.31 3 Mate
0.08 839 8.58 3 8.48
0.31 9.56 10.33 3 9.84
040 885 9.79 3 9.39

0.45
0.33
0.27
0.27
0.29
0.33

Height
Height

Maj. diam.
Maj. diam.
Min. diam.
Min. diam.
Outer lip
Outer lip
Last whorl
Last whorl
Col. axis
Col. axis
Weight
Weight
Volume
Volume

Max Number

3542525

Deviation

0.30
0.19
0.30
0.27
0.25
0.48

Min
11273
11.96
10.81

7.93

9.36

8.91

Max Number

12.72 4
12.58
11.60
8.82
10.14
9.88

A ap

“Peninsula de Оза” (altitude 0-20 т)
lots INBio 3542542, INBio 1482627, INBio
1482837, INBio 1482842, INBio 1485173

Mean
value

11.69
11.95
10.93
8.78
DUT
8.89

Deviation

0.50
0.43
0.40
0.42
0.39
0.52

“Bajo Bonito” (altitude 980 т)
lots IR 579, IR 1028, IR 1485

Min

12.82
12.29

Мах

13.70
12.65

Number

11.96 13.70 8
12.07 13:12 8
10.33 11.66 8
7.85 9.08 8
9.68 10.78 8
8.75 10.48 8
Mean
Sex value Deviation
f 13.26 0.44
m 12.41 0.16
Е 13.09 0.04
т 12.34 0.19
f 11.64 0.02
m 11.08 0.07
f 8.81 0.28
m 8.61 0.18
f 10.67 0.12
m 10.21 0.15
f 9.95 053
m 9.43 0.10
f 0.074 0.000
m “ONZA 0.024
f 0.603 0.000
m 0.507 0.027

13.05
12217
11:02
10.98

8:58

8.35
10.53
10.07

9.42

9.27
0.074
0.091
0.603
0.476

13.12
12.63
11.66
11.18

9.08

8.88
10.78
10.43
10.48

9.54
0.074
0.156
0.603
0.547

0 =0_0N0N0ayYN © ND WN W DY

Min
10.78
11.30
10835

8.25

9.27
91

Max Number

12.44
12162
11.54
925
10.30
JET

000000

252 RICHLING

Bajo Bonito (n=2/3)

Bajo Bonito (n=2/3)

Río Naranjo (type)
ТЕ

Rio Naranjo (type)
==] f=

FIG. 49. Expansion of outer lip of different
populations of Helicina pitalensis in Costa Rica
according to Table 5; for explanations see Fig. 47.

Sexual Dimorphism: Although not well sup-
ported, the data for only two females and
three males (Table 5, Figs. 47-51, upper
row) suggest that females are bigger. The
clear differences between both sexes for
height, minor diameter, and height of last
whorl may be only a coincidence.

Habitat

My live material came from two localities,
near Bajo Bonito and near San Vito. They are
characterized by steep mountain forests, prob-
ably primary rain forests, the first bordered by
secondary growth and small manually tended
agricultural areas. Helicina pitalensis lives in
arboreal environments mainly on the underside
of leaves of palms and Heliconiaceae. It was
also found in the dried and curled-up leaves of
abandoned banana trees. It thus has a very
similar habitat as Н. funcki.

Distribution (Fig. 53)
According to the relatively few records,

Helicina pitalensis is confined to the southern

Bajo Bonito (n=2/3)

a Naranjo (type)

Е zt
San Vito (n=3)
TE pe —
Fila de Cal (n=4)
TE =
Bajo Bonito, all (n=8)
IE —
Península de Osa (n=6)
E 1 L —
LH 4 1 1 1 1 a 4
0 6 7 8 9 10 [mm] 11

FIG. 50. Height of last whorl of different
populations of Helicina pitalensis in Costa Rica
according to Table 5; for explanations see Fig. 47.

I a
San Vito (n=3) San Vito (n=3)
or. E ==
Fila de Cal (n=4) Fila de Cal (n=4)
ЧЕ Ес 1t I
Bajo Bonito, all (n=8) Bajo Bonito, all (n=8)
TE ————— Y — _——_—
un de Osa (n=6) Peninsula de Osa (n=6)
E A A e ———
Y 1 1 L ap 1 1 1 11 1 1 | Й i: 1 1
0 5 6 7 8 9 10 [mm] 11 0 5 6 T4 8 9 10 [mm] 11

FIG. 51. Height of columellar axis of different
populations of Helicina pitalensis in Costa Rica
according to Table 5; for explanations see Fig. 47.

Pacific slopes and coastal lowlands in Costa
Rica. On the Península de Osa and in the Fila
de Cruces it is found at various localities.
From the area around the type locality and the
connecting area to the southern localities
there are no recent records. This may be ex-
plained by lack of investigations in these areas
and the relatively low abundance of the spe-
cies on one hand, and the fact that the Pacific
plains were transformed into agricultural plan-
tations to a large extent starting in the 1950s.
The records of Pittier date back to the end of
the 19" century when the areas where largely
unexplored and under closed forest cover.
Assuming that the interpretation of the records
listed in von Martens (1900) 1$ correct, H.
pitalensis at least was well distributed over the
area of the southern Pacific plain, replacing H.
funcki in this region. In the Fila Cruces area
and on Península de Osa, H. pitalensis 1$
found sympatrically with H. talamancensis.
A typical specimen (ZMB 45501) of H.
pitalensis is labeled as originating from La
Paz, a location at the Río Sarapiqui north of
the Cordillera Central on the Caribbean slope.
This location seems to contradict the assumed
distribution. Pending better knowledge, it 1$
considered here to be erroneous.

Discussion

The species most resembles Helicina funcki,
which is of about equal size and shows the
same shell ornamentation. Helicina pitalensis
is relatively higher and has more convex
whorls. All specimens of H. funcki studied
show neither banding nor the distinct angula-
tion of the periphery. The strong denticle at the
transition of the basal outer lip to the columella
is characteristic for H. pitalensis, whereas it
lacks the groove or angulation in the transition
from the columella into the body whorl. Fur-

CLASSIFICATION OF HELICINIDAE 253

Peninsula de Osa
Ч

FIG. 52. Variations м Costa Rican Не/ста pitalensis: shell height in figures reflects
the mean value (enlarged), each shell originates from the respective locality.

3500 - 4000 m
3000 - 3500 m
2500 - 3000 m
2000 - 2500 m
1500 - 2000 m
1000 - 1500 т
500 - 1000 m
100 - 500 m
0- 100 m

e coll. INBio
| © others

FIG. 53. Records of Не/ста pitalensis т Costa Rica.

254 RICHLING

thermore, the soft body color differs and is
constantly lighter in H. pitalensis.

The interpretation of the additional records
of H. funcki listed in von Martens (1900) in
SW-Costa Rica is difficult, because one of
these locations became the type locality of H.
pitalensis with description of this species by
Wagner. Von Martens (1900) remarked:
“some specimens banded and others more
elevated”, obviously for the specimens from El
Pital. As here, Wagner saw only the holotype
from the ZMB. If the remark of von Martens
(1900) also referred to the specimens from the
other locations, it would render their identifica-
tion as H. pitalensis very likely. Although other
material of Pittier is kept in the ZMB or the
МНММ respectively, these lots could, unfortu-
nately, not be found in either of the collections
and thus could not be verified. According to
personal information of Zaidett Barrientos
(INBio), those parts of the historical collections
in Costa Rica in the Museo Nacional in San
José could not yet be found when she
searched for this material. The museum had
passed through several crises and the where-
abouts of the material is uncertain. Regarding
the fact that, according to the present data, H.
funcki and H. pitalensis do not occur sympat-
rically and the latter species has not been re-
corded from the southern Pacific plain (except
the one doubtful record from the Península de
Osa, see: under H. funcki) and the question-
able interpretation of von Martens' remark, it
seems more appropriate, until better knowl-
edge comes along, to refer the records from
“Bay of Terraba, Tocori in the valley of the Río
Paquita, middle part of the Río Saveque and
lower part of the Río Pacuare (Pittier)” also to
H. pitalensis.

The record of Helicina amoena L. Pfeiffer,
1849, for Costa Rica by Monge-Nájera (1997)
was based on the material in the INBio collec-
tion. The subsequent revision of the material
revealed one lot of H. amoena (INBio 1485173)
collected and determined before 1997, on the
basis of which the publication must have been
based. This specimen can clearly be referred to
H. pitalensis in its typical form from the
Península de Osa. Helicina amoena is distin-
guished from H. pitalensis by its less elevated
shell, which is more strongly angulated at the
periphery and marked with distinct spiral stria-
tions. The color is also different. Except for one
doubtful record from Panama by von Martens
(1890) (“Helicina amoena var. b” from Cham-

pion) H. amoena has not yet been reported
south of the Mosquito Coast of Nicaragua
(Fluck, 1906: Mosquito Coast: NW Kukallaya,
Wounta River; Jacobson, 1968: Bonanza). The
original material of this doubtful record was
assumed to be in the collection of the ZMB, as
is other material of Champion, but it remains
lost, although it was also searched for under
other possible designations. It was later cited
by Pilsbry (1910, 1926a), but who claimed not
to have seen the specimen.

All recent records of H. pitalensis refer more
to the south than the type locality. The speci-
mens from the area of the Fila Cruces (Linda
Vista, Fila Cal, San Vito) at higher altitudes
(about 6001,000 m) are very similar to the
type material. Only the basic lemon-yellow
color is sometimes replaced by light orange-
brownish. Specimens from the Península de
Osa sometimes lack the band and show cer-
tain deviations in shape and a stronger infla-
tion of the whorls immediately below the
suture. But because other characteristics do
not differ (e.g., color, development of columel-
lar region, protruding denticle at the transition
of outer lip to columella, roundly angulated pe-
riphery) and adult material from other loca-
tions on the Península de Osa is not available,
a separation of this form is not yet warranted.

Helicina (Tristramia) tenuis
L. Pfeiffer, 1849

Helicina tenuis L. Pfeiffer, 1849: 124-125 (not

figured)

Helicina vernalis Morelet, 1849: 20 (not fig-
ured)

Helicina tenuis — L. Pfeiffer, 1850: 40, pl. 7,
figs. 33, 34

Helicina tenuis — L. Pfeiffer, 1852a: 372

Helicina vernalis — L. Pfeiffer, 1852a: 372

Helicina tenuis — L. Pfeiffer, 1852b: 269

Helicina vernalis — L. Pfeiffer, 1852b: 269-270

Helicina vernalis — L. Pfeiffer, 1853: 71, pl. 10,
figs. 12-14

Helicina chiapensis L. Pfeiffer, 1856: 237 (not
figured); 1857: 380 (not figured)

?Не/ста тает! — Tristram, 1862: 5: Guate-
mala: neighbourhood of Dueñas [according
to Tristram, 1864] (Salvin) [non L. Pfeiffer,
1849]

?Helicina lindeni — Sowerby, 1866: 288, pl.
272, figs. 258260 [non L. Pfeiffer, 1849]

Helicina chiapensis — Sowerby, 1866: 288, pl.
272, figs. 255-257

CLASSIFICATION OF HELICINIDAE 255

Helicina vernalis — Sowerby, 1866: 288, pl.
273, fig: 273

Helicina tenuis — Bland, 1866: 9

Helicina vernalis — Bland, 1866: 9

Helicina chiapensis — Bland, 1866: 9

Helicina vernalis — Reeve, 1874: pl. 18, fig.
156

Helicina chiappensis [sic] — Reeve, 1874: pl.
13, fig. 110

Helicina vernalis — von Martens, 1875: 649:
Guatemala: Coban, Vera Paz

Helicina vernalis — von Martens, 1876: 259:
Guatemala: Coban

Helicina lindeni — Angas, 1879: 484: Costa
Rica [non L. Pfeiffer, 1849]

?Helicina lindeni var. minor — Ancey, 1886:
258-259: Honduras, Atlantic coast (smaller
specimens)

Helicina tenuis — von Martens, 1890: 34-35:
Central Mexico: Sayula in Jalisco, Irapuato
near Guantajuoto; E-Mexico: Soledad, be-
tween Cordova and Orizaba; SE-Mexico:
Chiapas; Teapa and San Juan Bautista in
Tabasco, Tapinapa; Yucatan; N-Guatamala:
Peten Province; Cubilguitz, valley of the
River de la Pasion; Coban; San Geronimo
and the neighbouring mountains in Vera
Paz; Panzos; Chacoj; San Juan (all in the
valley of the Polochic River); Purula; S-Gua-
temala: Totonicapam mountains 8,500 to
10,500 feet (small variety); El Reposo 800
feet; Las Mercedes 3,000 feet; Cerro Zunil
4,000 feet; San Isidro 1,600 feet, all on Pa-
cific slope; Zapote, on the slope of the
Volcan de Fuego; Nicaragua: Toro Rapids?;
Costa Rica

Helicina tenuis var. chiapensis — Pilsbry, 1892:
339: Mexico: Tabasco: Poana (Rovirosa)

Helicina (Oligyra) lindeni — Fischer & Crosse,
1893: 416-420, pl. LVI, figs. 1-3: same data
as von Martens (1890) [non L. Pfeiffer, 1849]

Helicina tenuis — Biolley, 1897: 5: Costa Rica:
Turrubares, 200 m [Зап Pablo de
Turrubares, about 09°55’N, 84°27'W, San
José Province] and La Paz, 900 m, en el
camino del Sarapiqui [along the River
Sarapiqui] [not exactly localized, near Isla
Bonita?, about 10°15’30"М, 84°11'W,
Alajuela Province]

Helicina tenuis — von Martens, 1900: 604: SE-
Mexico: Poana, Tabasco; Honduras: East
Coast — smaller spec.; NE-Costa Rica: La
Paz , on the road to the Rio Sarapiqui
Sarapiqui [not localized, near Isla Bonita?,
about 10°15’30"М, 84*11'W, Alajuela Prov-
ince] (Biolley); Central Costa Rica: Alajuela,

900-1,000 т [town or province?, town about
10%01'30"N, 84°13’W] (Orosco), SW-Costa
Rica: Turubares, 200 m [San Pablo de
Turrubares, about 09°55’М, 84°27'W, San
Jose Province] (Biolley); along the Rio de
los Platanales and the Golfo Dulce [correct:
Rio de los Platanares, S of Puerto Jiménez,
Peninsula de Osa, about 08°31’30"М,
83°18’W, Puntarenas Province] (Pittier)

Helicina vernalis — Wagner, 1905: 233-234,
pl. XIII, fig. 13a-c: Guatemala: Petén;
Verapaz: Rio Polochic

Helicina vernalis verapazensis Wagner, 1905:
234, pl. ХИ, fig. 14: Guatemala: Verapaz

Helicina tenuis pittieri Wagner, 1910a: 303-
304, pl. 60, fig. 24

Helicina tenuis — Wagner, 1910a: 302-303, pl.
60, figs. 15-23, 25: S-Mexico to Panama:
Mexico: Tabasco and Chiapas; Guatemala:
Coban, Totonicapan, St. Isidoro, Río
Polochic, Mercedes and Vera Paz; Costa
Rica: Turrubares [San Pablo de Turrubares,
about 09°55’М, 84°27' W, San José Prov-
ince] and Alajuela [town or province?, town
about 10%01'30"N, 84°13’W]

Helicina tenuis маг. lindeni — Hinkley, 1920: 49,
52: Guatemala: Jocolo plantation on north
side of Lake Isabal; Alta Verapaz: Chama
between Río Tsalbha and Río Negro [non L.
Pfeiffer, 1849]

Helicina (Tristramia) tenuis — Baker, 1922a:
50, pl. Ш fig. 7, pl. М, fig. 14 (radula)

Helicina (Tenuis) tenuis — Baker, 1922b: 35-
36: Mexico: S Vera Cruz, near Hacienda de
Cuatolapam (Río San Juan - Arroyo
Hueyapam, canton of Acayacan (Michigan-
Walker-Expedition)

Helicina tenuis — Pilsbry, 1926a: 59, 71:
Panama: Los Santos Province: Tonosi
(Olsson)

?Helicina tenuis var. — Pilsbry, 1930: 339:
Panama: Barro Colorado Island (Pinchot-
Expedition)

Helicina (Tristramia) lindeni — Bequaert &
Clench, 1933: 543: not found again in
Yucatan [non L. Pfeiffer, 1849]

Helicina tenuis — Goodrich & van der Schalie,
1937: 12, 15, 32: Guatemala: Petén: region
of headwater of Río San Pedro de Mártir,
lower Río de la Pasión; Alta Verapaz: upper
part of Río de la Pasión

Helicina tenuis — van der Schalie, 1940: 6, 9,
10: Guatemala: Alta Verapaz: Pacala and
Chama, 290 m a.s.l., Samac, 1,300 т a.s.l.
[W of Coban], Panzamala, 1,250 т a.s.l. [$
of Lanquín] (Stuart)

256 RICHLING

Helicina (Helicina) tenuis — Haas, 1949: 137-
138: Guatemala: Chimaltenango: Yepocapa,
4800 ft.; Zacapa: Santa Clara, valley in the
interior of the Sierra de las Minas, N of
Cabañas, 5500 ft. (Wenzel &Mitchell)

Нейста tenuis — Bequaert, 1957: 207:
Chiapas: Selva Lacandona: Monte Líbano,
600 m, El Real, 600 m

Helicina tenuis tenuis — Thompson, 1967:
228-229: Mexico: Campeche: 10.2 mi E
Escárcega, rare (1 spec., dead), Chiapas:
15.8 mi NW Ocozocoautla

Helicina tennuis [sic] — Pérez 8 Lopez, 1993:
27: Nicaragua

Helicina oweniana — Monge-Nájera, 1997: 113:
Costa Rica [in part] [non L. Pfeiffer, 1849]

Synonymy

Helicina vernalis Morelet, 1849

Helicina chiapensis L. Pfeiffer, 1856

Нейста vernalis verapazensis Wagner,
1905

Helicina tenuis pittieri Wagner, 1910

Original Description

“Hel. testa, turbinata, tenuissima, vix
striatula, pellucida, corneo-albida, rubro obso-
lete trifasciata; spira conica, acuta; anfractibus
6 vix convexiusculis, ultimo basi planiusculo;
apertura fere verticali, triangulari-semiovali;
columella brevi, basi retrorsum subdentata,
superne in callum nitidum, circumscriptum,
dilatata; peristomate tenui, angulatim
expanso, margine basali cum columellae basi
angulum formante.

Diam. 11, altit. 8'/, mill.
From Yucatan.”

Type Material

BMNH 20010496.1-7 “Yucatan &

Barbadoes, coll. Hugh Cuming”
The type lot contains seven specimens, la-
beled as originating from Yucatan and
Barbadoes. The latter locality is not given in
the original description. In fact, the lot is a mix-
ture of two species, and only five specimens
agree with the description of Helicina tenuis.
The other two exhibit a less elevated spire,
less convex whorls and the first whorls in-
crease more rapidly in size. Furthermore, the
shells, lacking the spiral color bands, are col-
ored uniformly whitish, except for a broken
nearly transparent thin spiral line above the
periphery. Finally, the characteristic denticle of
H. tenuis at the basal outer lip is less strongly
developed. Thus, these specimens (BMNH
20010496.6-7) are excluded from the syntype
lot of Helicina tenuis, because they do not
agree with the original description and the
later given figure. It is very likely that the lot
was mixed subsequently to the studies of L.
Pfeiffer.

The largest specimen is here selected as
lectotype (Fig. 54), because it best agrees
with the figure in L. Pfeiffer (1850). It is the
only specimen in the lot with banding, without
operculum and about the size given in the
description. In comparison with the figure the
bands are faded, but it may be an exaggera-
tion in the drawing since they are described as
“rubro obsolete trifasciata”.

Dimensions:
Lectotype BMNH 20010496.1:

9.8/9.8/10.8/8.9/6.5/7.7/7.6 mm
Paralectotypes BMNH 20010496.2-5:

9.6/9.1/10.4/8.4/6.4/7.5/7.2 mm

FIG. 54. Helicina tenuis, lectotype, BMNH 2001496.1, height 9.8 mm; scale bar 2.5 mm.

CLASSIFICATION OF HELICINIDAE Zor

8.9/8.7/9.4/8.1/5.6/6.8/7.0 mm

9.8/9.0/10.0/8.3/6.0/7.2/7.5 mm

8.7/7.9/8.7/7.3/5.4/6.4/6.7 mm

It is remarkable that even the type lot shows
a comparatively great variation in size and
shape (e.g., lectotype and _ smallest
paralectotype).

Type Locality

“Yucatan” [Not clear, whether it refers to the
Mexican State of Yucatan or to the whole
Yucatan Peninsula, shared by Mexico, Guate-
mala, and Belize. The present data of distribu-
tion suggest its origin rather in the Mexican
State of Campeche or the Guatemalan Petén
Department.]

Type Material of Synonymous Taxa or Similar
Species

Helicina vernalis Morelet, 1849

Type Material: BMNH 1893.2.4.1991-1993:
Morelet coll., purchased from H. Fulton
The Morelet collection was bought by H.
Fulton and later purchased by the BMNH.
Fischer & Crosse (1893) studied the origi-
nal material in the Morelet collection and
figured a shell that can be identified by the
mark of a “x” and the clear similarity to the
figure. This shell is here selected as lecto-
type of Helicina vernalis (BMNH
1893.2.4.1991) (Fig. 55), because it is un-
certain whether Fischer & Crosse’s com-
ment in the figure caption (pl. LVI, fig. 1, 1a,
1b: “premier type de Г Helicina vernalis”)
can be regarded as a type selection. The
lectotype is colored uniformly whitish and

still possesses its operculum, whereas the
paralectotypes are whitish below the pe-
riphery and above tinged reddish-brownish
or yellowish with two reddish-brownish
bands respectively.
Dimensions:
Lectotype BMNH 1893.2.4.1991:
9.9/9.9/10.8/8.9/6.5/7.7/7.5 mm
Paralectotypes BMNH 1893.2.4.1992-
1993:
9.4/9.4/10.3/8.3/6.3/7.5/7.2 mm
9.2/9.2/10.0/8.2/6.2/7.3/7.0 mm

Туре Locality: “Petenensis sylvas” [Guate-
mala, Petén Department]

Helicina chiapensis L. Pfeiffer, 1856

Type Material: Syntype ZMB 65624: leg.
Ghiesbreght, ex coll. L. Pfeiffer (Fig. 56)
The description of Helicina chiapensis was
published in two journals. ш the earlier pub-
lication (December 1856), L. Pfeiffer stated
that he had received specimens from Hugh
Cuming, leg. Ghiesbreght, which he prob-
ably kept in his collection. The second pub-
lication (May 1857) refers to material in the
collection Hugh Cuming, leg. Ghiesbreght.
Thus additional syntypes are possibly in the
collection of the BMNH housing the main
collection of Hugh Cuming, although they
have not yet been found in the type collec-
tion.

Dimensions (height/greatest diameter/minor
diameter):
Syntype: 10.2/11.4/9.4 mm

Type Locality: “Mexico, Chiapa” [Mexico, State
of Chiapas]

FIG. 55. Helicina vernalis, lectotype, BMNH 1893.2.4.1991, height 9.9 mm; scale bar 2.5 mm.

258 RICHLING

Helicina lindeni L. Pfeiffer, 1849

Helicina lindeni L. Pfeiffer, 1849: 123 (not fig-
ured)

Helicina lindeni — L. Pfeiffer, 1850: 52, pl. 8,
figs. 25, 26

Material Studied: Helicina lindeni var. - BMNH

20010757: Mexico, Hugh Cuming coll., three
specimens
The type material could not be located in the
collection of the BMNH, although it was listed
in the catalogue of the BMNH collection by L.
Pfeiffer (1852b: 282) as coming from the type
locality Tapinapa, Mexico (leg. Linden).
The specimens in BMNH 20020757 defi-
nitely do not belong to Helicina tenuis, but
rather agree well with the original description
of Helicina lindeni, especially, because in
contrast to Н. tenuis it is slightly angulated at
the periphery and less elevated. None of the
shells shows a trace of spiral color bands.
The outer lip is more reflexed.

Helicina tenuis pittieri Wagner, 1910

Type Material: Holotype ZMB 103241: leg.
Pittier
Because the original description refers to one
specimen in the ZMB which also matches the
figure it is the holotype (Fig. 57).
Dimensions:
Holotype: 9.2/8.6/9.4/8.0/6.0/7.4/7.1 mm

Type Locality: “Costa Rica, Río de los
Plutunales, Golfo Dolce” [correct: Río de los
Platanares, S of Puerto Jiménez, Península
de Osa, about 08°31’30"М, 83°18’W,
Puntarenas Province]

Examined Material

Lee. |. RICHLING

Guanacaste: 3 km Е Nuevo Arenal,
10%31'53"N,84*52'50"W, 640 m a.s.l.: prop-
erty of pension Villa Decary, rain forest:
03.03.1997: (IR 52); 01.03.1999: (1R715):
02.03.1999: (IR 722); 31.07.1999: (IR 880);
23.02.2000: (IR 1266); along small creek E
of Villa Decary: 04.03.1999: (IR 730)

Heredia: S Puerto Viejo de Sarapiqui, Zona
Protectora La Selva, near OTS-Station,
about 10%25'53"N, 84°00'18"W, 60 m a.s.l.,
05.09.1999: (IR 1057); (IR. 71058);
12.02.2000: (IR 1181)

Puntarenas: Reserva Natural Absoluta Cabo
Blanco, 09°35’16"N, 85%05'45"W, 30 т
a.s.l.: Sendero Danes and trail from en-
trance: 25.08.1999: (IR 1001); (IR 1002);
27.02.2000: (IR 1289); (IR 1291): Sendero
Sueco: 02.03.2001: (IR 1481)

INBio COLLECTION

Guanacaste: 500 m E de la Estación
Almendros, 11*02'04"N, 85°31'10"W, 280 т
a.s.l., leg. Elba Lopez, 01.08.1994: 1 ad.
(INBio 1477167)
Parque Nacional Barra Honda, Los
Mesones: 10°10’12"М, 85°21’03"W, 300 т
a.s.l., 29.05.1993: 2 ads. (INBio 1463476);
10°10"12"N, 85°20/50"W,; 100: пота,
31.05.1993: 4 ads., 1 s.ad. (INBio 1463452)
(all leg. malacological staff of INBio)
Refugio Nacional de Vida Silvestre Bosque
рта, Sector Diriá: Sendero Espavel,
10°10’19"М, 85°35’44"W, 220 m a.s.l.: leg
Alexander Alvarado Mendez, 13.05.1999: 6
ads., 4 juvs. (INBio 3096450); 200 m a.s.l.:
leg. A. Berrocal, 22.11.1998: 2 ads. (INBio

FIG. 56. Helicina chiapensis, syntype, ZMB 65624, height 10.2 mm; scale bar 2.5 mm.

CLASSIFICATION OF HELICINIDAE 299

3435759); Camino a Esperanza, 10°10’32"М,
85°35'11"W, 260 т a.s.l.: 14.05.1999: 1ad., 1
s.ad., 2 juvs. (INBio 1498286); 2 ads. (INBio
1498287) (all leg Alexander Alvarado Mendez)
Puntarenas: Parque Nacional Carara:
Quebrada Bonita, 09*46'29"N, 84°36’34"W:
50 m a.s.l., 03.06.2000: 4 s.ads. (INBio
3324332); 100 m a.s.l., 02.07.2000: 3 ads.
(INBio 3129469); Carara, sendero Laguna
Meandrica, 09*48'20"N, 84*35'02"W, 100 т
a.s.l.: 15.07.2000: 1 s.ad. (INBio 3395010)
(all leg. malacological staff of INBio)
Parque Nacional Corcovado: Río Sirena,
08°30’25"М, 83°29'23"W, 545 m a.s.l.: leg.
Enia Navarro, 24.05.1995: 1 ad. (INBio
1484663); 2 km SW del Mirador, 08*32'30"N,
83°30'57"W, 200 m a.s.l.: leg. Socorro Avila,
22.05.1997: 1 ad. (INBio 1487810)
Reserva Forestal Golfo Dulce: Fila Casa
Loma, 1,600 m S de la Escuela de Rincón,
08°41’33"М, 83°29'17"W, 170 т a.s.l.: leg.
Socorro Avila, 10.10.1996: 2 ads. (INBio
1487328); Península de Osa, Instalaciones
de IDA, 08°41’38"М, 83°29'07"W, 60 т
a.s.l.: leg. Ramon Angulo, 07.06.1994: 1 ad.
(INBio 1480502)
Reserva Natural Absoluta Cabo Blanco:
Sector Balsitas, Sendero Central,
093502 N, 85°0726°W, 120 т; a.s.l.:
18.05.1994: 6 juv. (INBio 1473990); 2 ads.
(INBio 1475801); 1 ad. (INBio 1475805) (all
leg. Zaidett Barrientos); Sector Cabuya,
Sendero Sueco, Río Ariolo, 09°35’16"М,
85°05'41"W, 20 m as: leg. Ulises
Chavarría, 08.11.1994: 1 ad. (INBio
1480012); Sector San Miguel, Sendero Ma-
ven, 09°35’09"М, 85°08’12"W, 200 т a.s.l.:
leg. Zaidett Barrientos, 17.05.1994: 1 ad.
(INBio 1474149)

Cóbano, Estación Cabo Blanco, 09°35’30"М,
85°05'45"W, 15 m a.s.l., leg. malacological
staff of INBio, 09.01.1993: 4 ads., 5 s.ads.,
3 juvs. (INBio 1465481)
Sendero Camino Maven, orilla de quebrada
San Miguel, 09°35'18"N, 85°08'12"W, 100 т
a.s.l., leg. Alexander Alvarado Mendez,
21.01.1999: 2 ads. (INBio 1498272); 2 ads.
(INBio 1498276)
Quebrada San Miguel, 09°35’15"М,
85°08'15"W, 100 m a.s.l., leg. Socorro Avila,
05.10.1995: 1 ad. (INBio 1484853)
Alajuela: San Ramón, 10°05’19"М, 84°29'18"W,
1,060 m a.s.l., leg. malacological staff of
INBio, 16.09.1993: 1 ad. (INBio 1464319)
Estación Playuelas, 50 m del Río Frío,
10%57'29"N, 84%44'55"W, 40 m a.s.l., leg.
Kattia Martinez, 08.01.1994: 1 ad. (INBio
1479297)

OTHER SOURCES

COSTA RICA

Guanacaste: Las Cascadas, Quebrada San
Diego, 10°10'59.5"N, 85°20'18.5"W, leg.
D.G. Robinson & J.M. Montoya, 20.09.1998
(APHIS PPQ USDA)
Karst exposure, Cerro Barra Honda, approx.
10°10'10"N, 85°22'10"W, leg. О.С. Robinson
& J.M. Montoya, 19.09.1998 (APHIS PPQ
USDA)
Nicoya [about 10°08’30"N, 85°27'30"\М, leg.
H.G. Lee, ex G.D. Robinson, W.F. Webb: 1
ad. (UF 166944)
Pederal de Nicoya [about 10°08'N,
85°26’W], leg. Univ. Alabama, M. Smith coll.
(MS-15277): 12 ads. (UF 95336)
2.2 mi SE Nicoya [about 10°07’30"М,
85°26’W], 500 ft. leg. Е.С. Thompson (FGT-
106), 10.08.1964: 1 ad. (UF 214333)

FIG. 57. Helicina tenuis pittieri, holotype, ZMB 103241, height 9.2 тт; scale bar 2.5 mm.

260 RICHLING

3.8 mi S Nicoya [about 10°05’N, 85°28’W],
leg. Е.С. Thompson (FGT-111), 11.08.1964:
1 ад. (ЧЕ 35511)

1.2 mi E Caimital [about 10°04’N, 85°27’W],
leg. F.G. Thompson (FGT-109), 11.08.1964:
1 ad. (UF 214332)

Monte Alto conservation area, near
Pilangosta, Canton Hojancha, 10°00'48.1"N,
85°24'08.1"W, leg. О.С. Robinson & J.M.
Montoya, 19.09.1998 (APHIS PPQ USDA)

Alajuela: “Alajuela” [city or province?, town
about 10°01’30"М, 84°13’W], Orosco (ZMB
103247)

La Paz (Chem. du Sarapiqui) [not exactly
localized, near Isla Bonita?, about
10°15'30"М, 84*11'W], leg. P. Biolley (#90):
2 ads. (MHNN)

“San José Prov.:” San José [really San José,
capital of Costa Rica?, or province later
added and originally referring to San José in
Alajuela Province, here preferred: 14 km
NW of Upala, about 10%58'N, 85°08’W,
Alajuela Province], leg. McGinty coll., ex
Preston & Tomlin: 1 ad. (UF 160158)

San José: Turubares, Versant du Pacifique
[San Pablo de Turrubares, about 09°55’N,
84°27'W], 500 m a.s.l., leg. P. Biolley (#140),
06.1893: 21 ads., 1 s.ad. (MHNN)

Cartago: Turrialba [about 09°54’30"М,
83°41'W], coll. С. Bosch, ex coll. Rolle, ex
Wagner: 4 ads. (SMF 180786/4); coll. Rolle:
12 ads. (ZMB 103802)

Puntarenas: Golfito [about 08°39’N, 83°10’W],
leg. F.G. Thompson et al., 14.06.1964: 1 ad.
(UF 35510)

Costa Rica, without locality further specified:
leg: McGinty coll.: 1 ad. (UF 263576); 1 ad.
(UF 214331)

GUATEMALA

El Peten: S of Sayaxche, beyond L
Petexbatun, leg. J. Polisar, 31.08.1994: 4
ads. (UF 234127)

Huehuetenango: Cave below Finca Chiblac,
ca. 5 km W of San Ramon, 15°52’45"N,
91°14’34"W, 700 m a.s.l., leg. Е.С. Thomp-
son et al. (FGT-4828), 05.03.1991: 5 ads.
(UF 190327); (UF 190329: 1 of 6 spec.)

Alta Verapaz: 2 km WNW of Lanquin,
15°34’38"М, 89°59'19"W, 300 m a.s.l., leg.
S.P. Christman (FGT-4791), 21.02.1991: 2
ads. (UF 190068)

4 km W of Lanquin, 15°34’37"N, 90%01'06"W,
330 m a.s.l., leg. Е.С. Thompson (FGT-4793),
21.02.1991 (UF 190093: 1 of 5 spec.)

9 кт \М of Lanquin, 1573503М,
90°03'20"W, 690 т a.s.l., leg. Е.С. Thomp-

son et al. (FGT-4787), 20.02.1991: 1 ad. (UF
190036); (UF 190045: 2 of 4 spec.)

11 km W of Lanquin, 15*33'29'N:
90%04'02"W, 1,000 т a.s.l., leg. Е.С. Thomp-
son et al. (FGT-4801), 22.02.1991: 2 ads.
(UF 190142)

6.5 km SE of Lanquin, 15°32’52"N,
89°57'22"W, 400 m a.s.l., leg. Е.С. Thomp-
son (FGT-4796), 21.02.1991: 2 ads. (UF
190108)

8 km SE of Lanquin, 1532433
89°56'49"W, 350 m a.s.l., leg. Е.С. Thomp-
son et al. (FGT-4797), 21.02.1991: 1 ad. (UF
190116)

2 km ESE Cojaj, 15°33'25"М, 90°0656"W;
1,250 m a.s.l., leg. Е.С. Thompson (FGT-
4783), 20.02.1991: 3 ads. (UF 190006)

8 km by road N of Coban, 15°31’30"N,
90°23'11"W, 1,340 m a.s.l., leg. Е.С. Thomp-
son et al. (FGT-4776), 18.02.1991: 3 ads.
(UF 189950)

4 km E of Coban, 1,260 т a.s.l., leg. F.G.
Thompson et al. (FGT-4803), 23.02.1991
(UF 190156: 1 of 2 spec.)

Coban, Sumichrast: 2 ads. (UF 214336); leg.
Univ. Alabama, T.H. Aldrich coll. (THA-8198),
ex Mohr coll.: 2 ads. (UF 095334)
Limestone knoll 11 km S of Coban,
15°24'57"М, 90°24’09"W, 1,350:mra:sil;,
leg. F. G. Thompson et al. (FGT-4805),
04.02.1991: 1 ad. (UF 190163)

2.5 km by road NE of Puente Chixoy,
1521'32"N, 90°39'10"W, 810 m a.s.l., leg.
Е.С. Thompson (FGT-4781), 19.02.1991 (UF
189988)

Limestone knoll 17.5 km NW of Tactic,
15°21’29"М, 90%25'25"W, 1,330 m а.$1., leg.
Е.С. Thompson et а. (FGT-4764),
16.02.1991: 3 ads. (UF 189840)

10.5. km. SE of ‘El Tactic, MSMIGIS ONF
90°18'11"W, 1,460 m a.s.., leg. S.P.
Christman (FGT-4810), 26.02.1991: 2 ads.
(UF 190204)

E of Finca el Volcan, leg. J. Schuster,
22.07.1984: 1 ad. (UF 114090)

Izabal: Río Tameja, 12.9 km SSW Livingston,
leg. F.G. Thompson (FGT-54), 04.07.1964: 1
ad. (UF 214330)

Zacapa: La Union, Cerro Mona (N), 1,350-
1,500 т a.s.l., leg. Е.М. Smith, 20.06.1994:
1 ad. (UF 244447)

Retalhuleu: Retalhuleu, leg. Univ. Alabama, T.
H. Aldrich coll. (THA-8197) ex Mohr coll.: 2
ads. (UF 95335)

Guatemala, without locality further specified:
La Paz [localization?: perhaps Verapaz or in
Honduras?], ex coll. S. С. A. Jaeckel: 3 ads.

CLASSIFICATION ОЕ HELICINIDAE 261

(HNC 39843); leg. Beal-Maltbie coll., ex W.
Webb coll.: 1 ad. (UF 237376); leg. Beal-
Maltbie coll., ex W. Webb coll.: 1 ad. (UF
237377)

EL SALVADOR

Ahuachapän: 6 km W of Atiquizaya, on road to
Ahuachapän, leg. А. Zilch, 21.09.1951: 4
ads. (SMF)

HONDURAS

Colôn: Limestone ridge, 2.6 km SW of La
Brea, 15°45'39"N, 86°00’08"W, 100 m a.s.l.,
leg. FE: Thompson (FGT-5253),
22.10.1993: 1 ad. (UF 212023)

Olancho: Vicinity of Magua Cave, ca. 15 km
SSW of Gualaco, 14°56.5'N, 86°07.5'W, 940
m a.s.l., leg. Е.С. Thompson et al. (FGT-
5216), 11.03.1993: 4 ads. (UF 194339)

MEXICO

Guerrero: 1 km E Petaquillas, 1158 m a.s.l.,
leg Е.С. Thompson (FGT-1584), 03.11.1970:
9 ads. (UF 217551)

2.2 mi ММЕ of Mazatlan, 4800 ft., leg. Е.С.
Thompson (FGT-672), 14.06.1966: 1 ad.
(UF 77607)

Limestone hill, 1 km NW of Naranjito,
18°05’03"М, 101°50’45"W, 675 т a.s.l., leg.
F.G. Thompson (FGT-5087), 04.11.1992: 1
ad. (UF 200647)

Oaxaca: Lagunas, 259 т a.s.l., leg. Е.С.

Thompson, 18.07.1966: 2 ads. (UF 214337)
Limestone ridge, 4 km W of Cuauht, moc,
17°05'56"М, 94°54'25"W, 100 m a.s.l., leg. Е.
G. Thompson et al. (FGT-5271), 02.08.1993:
тс аа. (UF 211326)
Limestone knoll, 13 km ENE of Sarabia,
17°05'54"М, 94°56 34"W, 125 m a.s.l., leg. F.
С. Thompson et al. (FGT-5269), 02.08.1993:
3 ads. (UF 211316); leg. F. G. Thompson
(FGT-5280), 03.08.1993: 2 ads. (UF
211427)

Veracruz: 5 km ЕМЕ of Cuauht, moc, Оахаса,
17°06’59"М, 94°51 10"W, 75 m a.s.l., leg.
Е.С. Thompson et al. (FGT-5273),
03.08.1993: 1 ad. (UF 211337)

7 Кт $, 7 km Е of Catamaco, 350 m a.s.l.,
leg. Е.С. Thompson et al. (ЕСТ-4608),
03.01.1990: 4 ads. (UF 159375)

Laguna Encontada, 20.08.1962: 1 ad. (UF
214338)

Limestone knoll, 2 km SW of Plan Arroyo,
17°14’15"М, 94°37’36"W, 100 m a.s.l., leg.
Е.С. Thompson et al. (FGT-5278),
03.08.1993: 2 ads. (UF 211398)

Tabasco: 3 km N of Vicente Guerrero,
17°31'09"N, 92°56’00"W, 160 m a.s.l., leg.
F.G. Thompson (FGT-4873), 03.04.1991: 3
ads. (UF 190725)

6.8 km W Теара, leg. Е.С. Thompson (ЕСТ-
427), 08.07.1965: 2 ads. (UF 214344)

Campeche: 16.4 km Е Escárcega, leg. Е.С.
Thompson (FGT-406), 19.06.1965: 1 ad.
(UF 19296)

Chiapas: 15.1 km W San Cristobal, 2469 m
a.s.l., leg. Е.С. Thompson (FGT-446),
15.07.1965: 1 Juv; (UP 214335)4 ad. (УЕ
214340)

18.3 km М Tuxtla Gutierrez, 1372 m a.s.l.,
leg. F.G. Thompson (FGT-465), 22.07.1965:
1 juv. (UF 214341)

12.9 km М Tuxtla Gutierrez, 1158 m a.s.l.,
leg. F.G. Thompson (FGT-459), 19.07.1965:
3 ads. (UF 214343)

4.8 km SSE Tuxtla Gutierrez, 823 т a.s.l.,
leg. F.G. Thompson (FGT-763), 25.07.1966:
2 ads. (UF 214345)

7.5 km NNE Huixtla, 183 m a.s.l., leg. Е.С.
Thompson (FGT-757), 23.07.1966: 1 ad.
(UF 214346)

21.3 mi NW Huixtla, 300 ft., leg. D.R.
Paulson et al., 31.07.1965: 2 ads. (UF
214339)

Stream, 44.4 km NW Ocozocoautla, 610 m
a.s.l., leg. Е.С. Thompson (FGT-464),
21.07.1965: 1 ad. (UF 214342)

25.4 km NW Ocozocoautla, 823 т a.s.l., leg.
F.G. Thompson (FGT-462): 20.07.1965: 1
ad. (UF 19295)

34.1 km E, 16.4 km S Comitan, 1524 m
a.s.l., leg. Е.С. Thompson (FGT-441),
14.07.1965: 1 ad. (UF 214145)

Ruins of Palenque, leg. H.W. Campbell,
04.05.1970: 3 ads. (UF 214334)

Mexico, without locality further specified: leg.
Univ. Alabama, T.H. Aldrich coll. (THA-8195):
2 ads. (UF 95291)

Description

Shell (Figs. 58, 335F-1): Conical-globose,
semi-fragile to thin, sometimes semitrans-
parent, medium sized and only slightly shiny
to dull. Color: basic color yellowish to whit-
ish-opaque to horncolored, with up to three
indistinct reddish bands on body whorl: one
between suture and periphery and one or
two below the periphery. The lower band
only very weakly developed or obsolete.
Surface textured with fine irregular growth
lines and oblique grooves of different indi-

RICHLING

FIG. 58. Helicina tenuis. A-C. Cabo Blanco, IR 1001. A. Height 8.3 тт. В. Height 7.3 тт. С. Height
8.5 mm. D. La Selva, IR 1057, height 8.7 mm; scale bar 2.5 mm.

vidual orientation but of the same general
direction (Fig. 60), causing the dull appear-
ance. Embryonic shell with about 1 whorl,
4%-5 (lectotype: 4%) subsequent whorls
well inflated, remarkably convex, the last
whorl regularly rounded or sometimes with a
slight angulation at the periphery, under the
suture slightly shouldered; whorls equally
extending in size, forming a very regular
conical, pointed spire. Suture deeply im-
pressed. Aperture oblique and nearly
straight, last whorl regularly descending and
inserting exactly at the periphery. Outer lip
always yellowish-white, slightly thickened
and broadly expanded. Reflection nearly
rectangular to the whorl; transition to col-
umella with a remarkably protruding den-
ticle. Columella short. Basal callus weakly
developed and nearly completely smooth or
very little granulated, umbilical area without
groove.

Internal Shell Structures: (Fig. 59)

Teleoconch Surface Structure (Fig. 60): The
transitional structure extends about half a

whorl, the subsequent pattern of oblique di-
verging grooves continues up to the aper-
ture.

Embryonic Shell (Fig. 61): The structure re-
sembles that of Helicina funcki, occasionally
the pits are somewhat smaller. The embry-
onic shell size of the Costa Rican specimens
agrees fairly well with the larger shells (see
“Morphometry”) of the type lot of H. tenuis
which came from the Peninsula de Yucatan.
Diameter: 838 um (+ 28) (780-900) (п = 25)
(IR 1001, IR 1002); 834 um (+ 27) (800-860)

FIG. 59. Axial cleft and muscle attachments of
Helicina tenuis, |R 1001; scale bar 5 mm.

CLASSIFICATION ОЕ HELICINIDAE 263

FIG. 62. Operculum of Helicina tenuis, IR 1001;
scale bar 2 mm.

(п = 5) (BMNH 20010496.1-5, type lot, lec-
totype: 860 um); 813 pm (+ 9) (800-820) (п
= 3) (BMNH 1893.2.4.1991-1993, type lot of
Helicina vernalis, lectotype: 820 ит).

Operculum (Fig. 62): Very slightly calcified,
calcareous plate leaving a free margin,
thickened towards the columellar side. Color
reddish horny-amber, only the central area
yellowish-transparent. Columellar side
nearly regular S-shaped, upper end acute
and pointed, lower end continuously chang-
ing into outer margin.

FIG. 60. Teleoconch surface structure of Helicina
tenuis оп 2" whorl; scale bar 100 pm.

Animal (Figs. 337D, E): Foot and head are
greyish and become darker towards the
dorsal side; tentacles are greyish too. The
mantle pigmentation shows a high variabil-
ity: seldom unicolored light or dark, often
basic color light yellowish with two (or sel-
dom one) brown distinct but irregular bands
on the last whorl above and below the pe-
riphery and more or less irregularly brown-
ish spotted throughout the mantle. The few
specimens from Arenal were only brownish
spotted with small dots. The pattern 1$ al-
most always clearly visible through the
shell.

Radula (Fig. 63): A-central without well-de-
fined cusps, B-central in most cases with 3—
4, C-central only occasionally with up to 6
small cusps. Comb-lateral with 8-9 cusps,
cusps on marginals slowly increasing in
number. Radula with about 66-86 rows of
teeth. Description agrees with Baker (1922a:
pl. Ш, fig. 7, pl. IV, fig. 14).

Female Reproductive System (Figs. 64, 65):
FIG. 61. Embryonic shell of Helicina tenuis; scale The receptaculum seminis is a small, simple
bar 100 um. drop-shaped sac, the bursa copulatrix pos-

264 RICHLING

FIG. 64. Female reproductive system of Helicina
tenuis, IR 1001; scale bar 1 mm.

sesses few rather large simple lobes and is
of moderate size. The provaginal sac is ob-
long and well inflated, its distal end bears a
few small processes. lt has a slightly grey-
ish-brownish pigmentation. The stalk is
shorter than in Helicina funcki and rather
stout.

FIG. 63. Radula of Helicina tenuis. A. Centrals. FIG. 65. Variability of the female reproductive
B. Comb-lateral. C. Marginals; scale bar 50 um system of Helicina tenuis, IR 1002; scale bar 2.5
(A, B), 100 um (C). mm.

CLASSIFICATION ОЕ HELICINIDAE 265

TABLE 6. Measurements of different populations of Helicina tenuis given as mean value with standard
deviation, minimum and maximum value (min, max), and number of specimens; only population from
“Cabo Blanco” and “La Selva” were determined for the sex (min./max. diam. = minor/major diameter,
col. axis = columellar axis); linear measurements [mm], weight [g], volume [ml].

“Cabo Blanco” (altitude 30 m) “La Selva” (altitude 60 m)
lots IR 1001, IR 1002, IR 1289 lots IR 1057, IR 1181

Mean Mean
Sex value Deviation Min Max Number value Deviation Min Max Number

Height f 8.12 0.36 7.28 8.98 54 = = E = -
Height m 7.34 025 6:71 792 30 8.23 0.27 7.83 8.67 4
Maj. diam. f 7.93 829 7.18 8.69 54 - - - - -
Maj. diam. т 7.33 0.19 6.90 7.72 30 8.06 0.21 7.15 8.36 4
Min. diam. f 7.31 0.27 667 3803 54 - - - - -
Min. diam. т 6.68 018 6:33 100 30 1438 0.14 715 755 4
Outer lip f 5.39 0.24 481 6.05 54 - - - - -
Outer lip m 5.03 0.16 449 599 30 5:53 O16 5:37 5.70 4
Last whorl f 6.43 0.29 564 7.24 54 - - - - -
Last whorl т 5.83 021 547 62 30 6.39 0.23, 6.15 “6.67 4
Col. axis f 6.37 O26M5 65712054 - - - - -
Col. axis m 5.74 0.19. 501 6.23 30 6.50 0.187 6.13 . 6.73 4
Weight f 0.030 0.009 0.015 0.092 54 - - - - -
Weight m 0.027 0.007 0.013 0.048 30 0.053 0.011 0.037 0.065 3
Volume Е - 0.159 0.018 0.119. 0.207 54 - - - - -
Volume m 0.117 0010 "0100 0136 30 0.162 0.011 0.145 0.172 3
“Она” (altitude 220-260 m) “Barra Honda” (altitude 100-300 т)

lots INBio 1498286, 1498287, 3096450 lots INBio 1463452, 1463476

Mean Mean

value Deviation Min Max Number value Deviation Min Max Number
Height 7.89 0.307 7.15 843 8 8.48 0.36; 767 902 6
Maj. diam. 7.91 022076114832 9 8.11 0.29 7.59 8.68 6
Min. diam. 7.14 0.20 6.82 7.48 9 1655 0.26 - 6:98 7.98 6
Outer lip 5.19 0.18 5.00 585 8 5.50 021025235743 6
Last whorl 5.99 0.19 5.66 6.36 8 6.42 0:41 5:60 6.93 6
Col. axis 6:27 0.20 5:73 "16:68 9 6.88 0.36 6.03 7.34 6

“Cabo Blanco, INBio” (altitude 15-120 т)

lots INBio 1465481, 1475801, 1475805,

1480012, 1484853, 1498272, 1498276, “Turrubares” (altitude 500 т)

IR 1481 lot MHNN

Mean Mean

value Deviation Min Max Number value Deviation Min Max Number
Height 7.94 0:39 6.92 8168 22 7.88 050% 16:9]. 900 21
Maj. diam. 7.95 0.35 7.007 860 22 7.90 0.418 720 8:80 %21
Min. diam. 7.22 0:33 6.32 77321 eal 036 649 797 21
Outer lip 5.30 026 46565 585 22 5:22 OMAN 45385 = 5.805 21
Last whorl 6.23 0.32 530 6178 22 6.05 085 5:40 6.84 21

Col. axis 6.25 0:34: 5.29’ 6.90: 22 6.09 0.41 5.30 И 21

(Continues)

266
(Continued)
“Сагага” (altitude 100 т)
lot INBio 3129469

Mean

value Deviation Min Max
Height 8.08 0.52 17231 8.68
Maj. diam. 7.60 0:29 718 07:93
Min. diam. 7.10 039 6.51 7.60
Outer lip 5.28 039 2707 957
Last whorl 6.21 0.27 5.81 6.60
Col. axis 6.22 0.46 5.53 6.88

Morphometry and Sexual Dimorphism

Although Helicina tenuis is widely distributed,
only data from Costa Rica and the type lot or
those of type lots of synonyms, respectively,
were included, because the lots studied from
other areas consisted of only a very few speci-
mens. Except for “La Selva”, the Costa Rican
populations originated from the Pacific side.
The only southern specimens are those sum-
marized as “Peninsula de Osa” and the holo-
type of H. tenuis pittieri (Table 6, Figs. 66-70).

BE Blanco (n=54/30)

2

o
La Selva (n=0/4)
——
et (type lot) (n=5)
—
Malle (non tenuis) (п=2)
—— —

Petén (type lot H. vernalis) (n=3)
men) fi

Osa (type H. tenuis pittieri)
Es) fo

Ома (n=8)
E E

Barra Honda (n=6)
7H

Cabo Blanco (INBio) (n=22)
==

Turrubares (п=21)
TE

Carara (n=3)
= ja

Peninsula de Osa (n=4)

RICHLING

“Оза” (altitude 60-545 m)
lots INBio 1480502, 1484663, 1487328,

1487810
Mean
Number value Deviation Min Max Number
3 7.55 0.83 6.70 8:68 4
3 6.98 0:58 635 806 5
S 6.54 0.57 588 735 4
3 4.95 0.38 451 5.74 5
3 5.99 0.58 5.44 6.86 5
3 6.00 0.69 524 6.85 4

Morphometry: Regarding the type lot of
Helicina tenuis, the non-conspecificity of all
specimens is confirmed in the measure-
ments, especially in the height-diameter-re-
lation. Helicina tenuis pittieri closely
approaches the mean value of the type lot in
all characteristics. The same is true for the
type lot of H. vernalis, which is larger, but
otherwise shows similar relations between
the different measurements, additionally
supporting the status as a synonym.
Except for “Peninsula de Osa”, the Pacific
populations are remarkably similar to each
other in all characteristics. The specimens
from Barra Honda have a bigger shell, which
is more highly elevated (height, columellar

Cabo Blanco (n=54/30) 9
o”

La Selva (n=0/4)

Yucatán (type lot) (n=5)
pj

Yucatán? (non tenuis) (n=2)
pp [A

Petén (type lot H. vernalis) (n=3)
pj À

Osa (type H. tenuis pittieri)
EJ

Diriá (n=9)
Barra Honda (n=6)
De |

1 | 1 1 1 1 L 1 TE
0 4 5 6 7 8 9 [mm] 10

FIG. 66. Shell height of different populations of
Helicina tenuis in Costa Rica according to Table
6; on each line: mean value, standard deviation,
absolute range; number of individuals given as
“п = females/males or total”; upper line: females,
lower line: males if separate; in between and
shaded: average of both for comparison with
populations of unknown sex.

Cabo Blanco (INBio) (n=21)
==

Turrubares (n=21)

Carara (n=3)

zen elle de Osa (n=4)

— | 1
9 [mm] 10

FIG. 67. Minor diameter of shell of different
populations of Helicina tenuis in Costa Rica
according to Table 6; for explanations see Fig. 66.

CLASSIFICATION OF HELICINIDAE 267

Cabo Blanco (n=54/30) 9

о

La Selva (n=0/4)

Cabo Blanco (п=54/30)

La Selva (n=0/4)

— —-
Yucatán (type lot) (n=5) Yucatán (type lot) (n=5)
pa] {i= I
Yucatän? (non tenuis) (n=2) Yucatán? (non tenuis) (n=2)
= —/ mm
Petén (type lot H. vernalis) (n=3) Petén (type lot H. vernalis) (n=3)
Е == Е ——
Osa (type H. tenuis pittieri) Osa (type H. tenuis pittieri)
pee] А I =p: E
Diriá (n=8) Diriá (n=9)
=] = Е ——_
Barra Honda (n=6) Barra Honda (n=6)
—— =f —_—=> о
Cabo Blanco (INBio) (n=22) Cabo Blanco (INBio) (n=22)
===>] у = —_
Turrubares (n=21) Turrubares (n=21)
рев
Сагага (n=3) Сагага (n=3)
E==/ | — ee
Peninsula de Osa (n=5) Peninsula de Osa (n=4)
pay ay
| 1 1 1 1 | 1 1 il 1 1 1 1
EU 3 4 5 6 7 [mm] 8 o 3 4 5 6 7 [mm] 8

FIG. 68. Expansion of outer lip of different
populations of Helicina tenuis in Costa Rica
according to Table 6; for explanations see Fig. 66.

axis). Besides the small sample size, the
comparatively high deviations among the
specimens from Peninsula de Osa presum-
ably reflect the fact that they originate from
different sites on the Peninsula and cannot
be considered as a real population the same
as the others. Contrary to their small size the
type of H. tenuis pittieri, collected about 100

Cabo Blanco (n=54/30)

La Selva (n=0/4)

Yucatan (type lot) (n=5)
=| se

Yucatän? (non tenuis) (n=2)
f szecaeare

Bay (type lot H. vernalis) (n=3)

Osa (type H. tenuis pittieri)
=f

Diria (n=8)

Barra Honda (n=6)
1

Cabo Blanco (INBio) (n=22)
pee]

Turrubares (n=21)
Be

Carara (n=3)
=>

Península de Osa (n=5)
E

FIG. 69. Height of last whorl of different
populations of Helicina tenuis in Costa Rica
according to Table 6; for explanations see Fig. 66.

FIG. 70. Height of columellar axis of different
populations of Helicina tenuis in Costa Rica
according to Table 6; for explanations see Fig. 66.

years ago on the same peninsula, is excep-
tional big for the Pacific populations. It sug-
gests that H. tenuis displays greater size
variation in this area, but the scanty material
does not allow further conclusions.
Considering the sexual dimorphism, the av-
erage shell height of Caribbean specimens
from “La Selva” can be estimated approxi-
mately 8.6 mm, thus being bigger than the
Pacific populations. This may be caused by
the drier climate on the northern Pacific side
as compared to the Caribbean plain.

In general, the average of the type lot of H.
tenuis appears to be typical for the Mexican
and Guatemalan areas, since many single
specimens from this region were measured
and approach a similar size. Nevertheless,
smaller specimens were also present, as
was the case for the type lot that also may
have consisted of specimens from various
localities in Yucatan. Goodrich & van der

| Cabo Blanco (n=54/30) 9
| y — nn — 4

La Selva (n=0/3)

1 1

| ul L 1 1
0 0.05 0.075 0.1 0.125 0.15 [ml] 0.2

FIG. 71. Shell volume of different populations of
Helicina tenuis in Costa Rica according to Table
6; for explanations see Fig. 66.

268 RICHLING

|
| Cabo Blanco (n=54/30)

La Selva (n=0/3)

— o
1 1 1 1 L 1 1

0 0.025 0.05 0.075 0.1 0.125 [9] 0.175

FIG. 72. Shell weight of different populations of
Helicina tenuis in Costa Rica according to Table
6; for explanations see Fig. 66.

Schalie (1937) state for Péten and North Alta
Verapaz, northern Guatemala, that shells
from the southern region are a little smaller,
unfortunately without giving any measure-
ments. In conclusion, it can be assumed that
the shell size of H. tenuis varies throughout
the whole range of distribution, obviously
depending on environmental factors. Near
its southern limit of distribution in Costa
Rica, the average size 1$ smaller.

Sexual Dimorphism: The 84 specimens of the
Cabo Blanco population show clear differ-
ences with the average size of females
larger than males. The measurements over-
lap, as shown for height and minor diameter
in Fig. 73, but to a smaller degree than in H.

funcki (Fig. 32). As may be expected, the
volume (Fig. 71) best reflects the differ-
ences, the average volume of male amounts
only 73.6% of the females. The shell weight
of both sexes 15 nearly equal (Fig. 72), there-
fore males possess relatively heavier shells
(Fig. 74).

Habitat

During this study, Helicina tenuis was only
found in comparatively high abundance during
the rainy season in the Cabo Blanco reserve.
During the daily rains, the snails were seen
crawling on and under living and dead leaves
of bushes and palms and on stems. None
were collected on the ground. In the same
place, H. tenuis was nearly “absent” during the
dry season, except for very few specimens
that were aestivating in folded palm leaves.
Two of these seven specimens found were
visibly parasitized by larvae of trematodes,
whereas no other helicinids ever were found
to be infected in this obvious way. It is not
clear where the majority of specimens retreat
to during the dry period. In addition to the ar-
boreal habitats, searches were conducted in
the leaf litter and around the stems of bushes

Re =
min
diam. +
[mm]
8 I
7.5 +
7 |
&
ar
nah
6.5 | В
a

6 6.5 7 7.5

8 8.5 9

female +
male +

height [mm] 10

FIG. 73. Range of measurements in females and males exemplary for height
and minor diameter in the population from Cabo Blanco.

CLASSIFICATION OF HELICINIDAE 269

0.1

| female >
weight

L male +
[9] juvenile «

0.08

0.07

0.06

0.05

0.04

0.03

0.02

0 0.05 0.1 0.15 volume [ml] 0.25

FIG. 74. Relation of weight to volume in females and males of the populations
from Cabo Blanco.

86° | 85° 84° | ‚83°

am

3500 - 4000 m |
3000 - 3500 | Mts... 1 | —
2500 - 3000 m | 08
2000 - 2500 m |
1500 - 2000 m |
1000 a 1500 m Er sn | = = à
500 - 1000 m
100 - 500 m |

e coll. IR À
om e coll. INBio |
o others | |

FIG. 75. Records of Не/ста tenuis in Costa Rica.

270 RICHLING

and palms on the ground but without success.
In other areas without such contrasting sea-
sonal changes (e.g., near Nuevo Arenal, La
Selva), H. tenuis was only found occasionally
during the dry as well during the rainy season.
There it was additionally found on the under-
side of leaves of Musaceae or Heliconiaceae,
respectively, which are absent in the drier
Cabo Blanco area.

These habitats correspond to those ob-
served by Biolley (1897) for Costa Rica and
Baker (1922b) from southern Veracruz in
Mexico, who additionally found the species
“on the ground and on leaves of shrubs and
cacti in the savannah forests”. Van der Schalie
(1940) reported H. tenuis from Alta Verapaz in
Guatemala as common and as being found
near Panzamala “moving about on the vegeta-
tion at night”.

With its occurrence on the Península de
Nicoya, H. tenuis tolerates the highest level of
dryness among the Costa Rican Helicinidae
and is the only species that can withstand the
regular extended dry period during the year.
As cited above, it also inhabits the Savannah
in association with cacti. This comparatively
high ecological tolerance of Helicina tenuis
among the Helicinidae provides a possible
explanation for its remarkably wide distribu-
tion.

Distribution

The species reaches its northern limit in
southern Mexico (states of Guerrero, Oaxaca,
Veracruz) and occurs throughout Guatemala,
Honduras, El Salvador, Nicaragua, Costa Rica
to western Panama. Even more northern sites
in Central Mexico (states of Jalisco and
Guantajuato) were listed by von Martens
(1890-1901), but the specimens have not
been re-examined. The most southeastern
record comes from the Tonosi, Los Santos
Province, Panama (Pilsbry, 1926a). The
specimens from Isla Barro Colorado in the
Canal Zone of Panama (Pilsbry, 1930) seem
to belong to another species (see “Discus-
sion”). Helicina tenuis is found on the Carib-
bean as well as on the Pacific side of the
central mountain chains. Except for Lucidella
lirata, it is thus probably the most widely dis-
tributed species of Helicinidae of the Central
American mainland.

In Costa Rica, the species is not common,
but was nevertheless found at several distinct
localities (Fig. 75). According to the collec-

tions, it seems to occur in relatively greater
numbers in the Pacific plain, where it also ap-
pears to be more widely distributed. Com-
pared with areas investigated and inhabited by
other helicinids (e.g., see H. funcki), the ap-
parent lack of H. tenuis on the Caribbean side
at many localities is remarkable, because the
species is found in a similar habitat and 1$
comparatively large. In fact, it is completely
absent throughout the large province of Limón
stretching along the entire Caribbean coast of
Costa Rica. La Selva and Turrialba represent
the most southeastern localities.

Discussion

The nomenclatural discussion of Helicina
tenuis is complicated because several confu-
sions have arisen and been maintained in lit-
erature.

First, it is important to note that Helicina
tenuis is not preoccupied by Helicina tenuis C.
В. Adams, 1849 (now Stoastomops adamsi
Baker, 1934) from Jamaica, because the latter
name was published in September 1849
(Baker, 1934a) and not as stated by von Mar-
tens (1890) or Bequaert 8 Clench (1933) in
1840.

Traditionally, Helicina tenuis and H. lindeni,
both described by L. Pfeiffer in the same pa-
per, the second one page before the other, are
regarded as synonyms or varieties of one spe-
cies. Sowerby (1866) only mentions H. lindeni,
and his drawing probably represents H. tenuis,
but both figure and the very short paragraph
on the species do not provide sufficient infor-
mation to assess the status. Von Martens
(1890: 34-35) proposed the synonymy with-
out further explanation, except for a statement
about the figure of H. lindeni in L. Pfeiffer
(1850) “not good”, and he used H. tenuis as
the valid name, which also would have estab-
lished priority because von Martens was the
first revising author. Fischer 8 Crosse (1893)
agreed upon the conspecificity, but claimed
that H. lindeni had page priority and H. tenuis
became a variety. None of these authors men-
tions an investigation of the original material
(Fischer 8 Crosse did so for H. vernalis), nor
did they give reasons for their opinion. Inter-
estingly enough, von Martens (1900: 604) re-
plied in his supplemental part to the French
authors, regarding tenuis as the most appli-
cable name and remarked on the rather great
distinctness L. Pfeiffer attributes to these spe-
cies (see below). Because both publications

CLASSIFICATION OF HELICINIDAE 271

are standard contributions on terrestrial mol-
luscs for Central America, subsequent authors
used the one or other name, but commonly
adopted the synonymy.

L. Pfeiffer (e.g., 1852a: 372, 388) assigned
his two species to different higher groups (“8.
8. Есаппаае” [H. tenuis] and “8. 10.
Subcarinatae” [H. lindeni]). The descriptions
and the subsequently published figures (L.
Pfeiffer, 1850) (reprinted here in Fig. 76) are in
fact not similar enough to support the syn-
onymy. Obvious differences can be summa-
rized in a less elevated shell in H. lindeni
(“globosa-conica” instead of “turbinata”; “spira
acutiuscula” instead of “acuta” and the mea-
surements). Furthermore, H. lindeni is slightly
angulated and does not bear color bands, the
outer lip is “breviter expanso, reflexiusculo”
instead of “tenui, angulatim expanso”. L.
Pfeiffer's descriptions are short, but very pre-
cise in certain details. Regarding the literature,
the conclusions of von Martens and Fischer &
Crosse can thus not be understood, especially
the “not good” figure of H. lindeni, because it
perfectly matches the written description.
Since the type material of H. lindeni is still
unavailable, possible deviations of the original
material (perhaps seen by other authors) from
the description that could have explained
those conclusions, remain subject to specula-
tion. А variety of H. lindeni from the Cuming
collection (BMNH 20010757) fits well to the
description and figure of H. lindeni. In conciu-
sion, H. tenuis, for which a lectotype could be
chosen in full agreement with the description
and the current interpretation, is regarded as
specifically distinct from H. lindeni. The Costa
Rican specimens clearly belong to H. tenuis.
According to comments and figure H. lindeni
sensu Fischer & Crosse (1893) is synony-
mous with H. tenuis.

The type material of Helicina vernalis and H.
chiapensis was investigated and the species
are confirmed as synonyms of H. tenuis. The
taxon H. vernalis verapazensis proposed by
Wagner (1905) was included into the syn-
onymy of H. tenuis by himself.

FIG. 76. Reproduction of the figures from L. Pfeiffer
(1850) of A. Helicina tenuis. B. Helicina lindeni.

The present status of Helicina tenuis pittieri
is doubtful, because comparable material from
the Peninsula de Osa is very scarce and the
few specimens available show a high variation
in size, are always not only smaller, but also
belong to different sites. A common feature is
the whitish band at the periphery, which is
lacking in other Costa Rican populations. Con-
sidering the high variation of the widespread
H. tenuis and the lack of further distinguishing
characteristics, H. tenuis pittieri is tentatively
regarded as a synonym. Wagner (1910a:
303), judging H. tenuis as variable and even
not constant in local forms, presents only the
new subspecies at the southern limit of the
distribution as a “auffallender unterschiedene
und anscheinend konstante Form” [strikingly
different and apparently constant form]. Ac-
cording to the original description and as far
as it could be traced in collections (ZMB, SMF,
MIZ [Wagner coll.]), it seems very likely that
he only knew a single specimen, the holotype
of the so-called “constant form”. More north-
ern records in Costa Rica Wagner included in
the nominal form.

Several records of H. tenuis from high eleva-
tions (e.g., Cerro Zunil) given by von Martens
(1890-1901) likely refer to H. punctisulcata
zunilensis, in one example cited it is very likely
that the record exactly originates from the
specimen on which Wagner (1910a) based his
new subspecies. The record from the Canal
island of Panama (Pilsbry, 1930) seems to be
based on another species, because the size of
the specimen (5/5.4 mm) is clearly beyond the
range of H. tenuis.

The record of H. oweniana for Costa Rica by
Monge-Najera (1997) was checked in the
INBio collection. The lots INBio 1463452 and
1464319 clearly determined before 1997,
have to refer to H. tenuis. H. oweniana is fi-
nally distinguished from H. tenuis by its or-
ange colored outer lip, a more solid shell, a
less impressed suture with its lower margin
whitish. Helicina oweniana lacks the typical
denticle at the transition from the outer lip to

the columella.

Helicina (Tristramia) echandiensis
Richling, n. sp.

Type Material

Holotype: INBio 3542520, female (leg.
Alexander Alvarado Mendez, 14.11.2001)

272 RICHLING

FIGS. 77, 78. Helicina echandiensis n. sp. FIG. 77. Holotype, INBio 3542520, height 7.2 mm. FIG. 78.
Paratype 1, INBio 3542521, height 6.5 mm; scale bar 2.5 mm.

Paratype: INBio 3542521, male (same data as
holotype)

Additional paratypes: INBio 3428246: 6 ads.,
1 s.ads., 9 juvs., INBio 3574064: 1 s.ad.
(same data as holotype)

Dimensions:

Holotype: 7.2/6.8/7.3/6.2/4.4/5.3/5.4 mm

Paratype 1: 6.5/6.1/6.5/5.7/3.9/4.8/5.1 mm

Type Locality

S-Costa Rica, Puntarenas Province, Parque
Nacional La Amistad, Sector Las Alturas,
Southern Cordillera de Talamanca, S of Cerro
Echandi, campamento Echandi, 09*01'33"N,
82*49'12"W, 2,840 т a.s.l.
Etymology

The name refers to the origin of the species,
the Cerro Echandi.

Examined Material

INBio COLLECTION

Puntarenas: Zona Protectora Las Tablas, sec-
tor Las Alturas, campamento de los
nacientes del Río Vella Vista, 08°59'39"N,
82°49'18"W, 2,100 m a.s.l.: leg. E. Alfaro,
13.11.2001: 1 ad. (INBio 3505804)

Description

Shell (Figs. 77, 78, 335J-K): conical, thin and
fragile, medium to small sized, only slightly
shiny to dull. Color: basic color light orange-
brownish; apex and upper whorl unicolored,
only lighter towards the suture, about the 1.5
last whorls above periphery with a pattern of
irregular, mostly parallel distinct white stripes
in the same orientation as growth lines,
about as wide as interspaces; stripes start-
ing at suture and all ending at the same level
a little above periphery. Surface textured
with irregular growth lines and oblique
grooves of different individual orientation but
of same general direction (Fig. 80), causing
the rather dull appearance. Embryonic shell
with about 1 whorl; 37/, (372-4) subsequent
whorls very straight; last whorl also straight

CLASSIFICATION OF HELICINIDAE 2783

FIG. 79. Axial cleft and muscle attachments of
Helicina echandiensis n. sp., INBio 3542520;
scale bar 2.5 mm.

above and round at periphery and below;
whorls equally extending in size, forming a
very regular, pointed spire. Suture very
slightly impressed. Aperture slightly oblique
and straight, last whorl very slightly ascend-
ing towards the aperture and inserting just
below the periphery. Outer lip of a bright or-
ange, thickened, moderately and equally
expanded. Reflection nearly rectangular to
the whorl; transition to columella forming a
blunt edge with a very small denticle. Col-
umella oblique and rather straight, transition
to the body whorl smooth. Basal callus
weakly developed, at the base more pro-
nounced and granulated.

Internal Shell Structures: (Fig. 79)

Teleoconch Surface Structure: Helicina
echandiensis n. sp. seems to lack the tran-
sitional pattern (Fig. 80A), the whole
teleoconch exhibits a structure of oblique
diverging grooves (Fig. 80B). About the up-
per half of the beginning of the 1° whorl is
occasionally sculptured with fine wrinkles
parallel to the growth lines. Overlapping
equally spaced periostracal spiral ridges
also begin immediately at the teleoconch
(Fig. 80A, arrow).

Embryonic Shell (Fig. 81): Faced with the pau-
city of material for Helicina echandiensis n.
sp., only one specimen was studied under
the SEM (INBio 3574064). The younger part
ofthe embryonic shell appears compressed,
as if it developed in slower growth, but this is
not likely to be a typical feature. It is prob-
ably not related to living conditions at high
altitudes where the species is found, because
the embryonic shell of H. punctisulcata

FIG. 80. Teleoconch surface structure of Helicina
echandiensis n. sp. A. Embryonic shell and
begin of 1% and 2" whorl, arrow indicates
exemplarly an early spiral ridge. В. 3“ whorl;
scale bars 500 um (A), 100 um (B).

274 RICHLING

FIG. 81. Embryonic shell of Helicina echandiensis
п. sp.; scale bar 100 um.

cuericiensis n. subsp. is normally developed.
The arrangement of the pits is less regular
than in the previous species, and the pits are
relatively smaller. The embryonic shell size
is much larger than in H. escondida n. sp. of
equal shell size and even exceeds that of
specimens of H. funcki from the lowlands.
Diameter: 1,026 um (+ 36) (960-1,120) (n =
9) (INBio 3428246, INBio 3542520, INBio
3542521).

FIG. 83. Radula of Helicina echandiensis n. sp.
FIG. 82. Operculum of Helicina echandiensis n. A. Centrals. B. Comb-lateral. C. Marginals; scale
sp., INBio 3542520; scale bar 1 mm. bars 50 um (A, B), 100 um (C).

CLASSIFICATION OF HELICINIDAE 279

Operculum (Fig. 82): very slightly calcified,
calcareous plate covering only part of outer
surface. Color horny-amber to orange, only
near the columella whitish or transparent.
Columellar side slightly irregular S-shaped,
upper end acute and pointed, lower end con-
tinuously changing into outer margin.

Animal: In the preserved specimens, the soft
body is greyish-blackish throughout. Only
towards the sides and underside of the foot
does the color become lighter. The sides of
the foot and parts of the mantle are occa-
sionally only spotted greyish.

Radula (Fig. 83): Due to the lack of material,
the radula of only one specimen was inves-
tigated. Cutting edges in centrals rather
crenulate than bearing cusps, comb-lateral
with 10-11 cusps, cusps on marginals slowly
increasing in number. Radula with 72 rows
of teeth.

Female Reproductive System (Fig. 84): The
receptaculum seminis is long and slender
and joins the descending limb of the V-organ
at the middle of its inner side. The bursa

=
7

FIG. 84. Female reproductive system of Helicina
echandiensis п. sp., INBio 3542520; scale bar
1 mm.

copulatrix is moderately lobed, the flattened
provaginal sac is of about equal size. It is
clearly demarcated from its short and stout
stalk, the distal side is irregularly subdivided.
The pallial oviduct is mainly transversally
constricted.

Morphometry and Sexual Dimorphism (Table
7, ЕЮ. 85)

The material available is very limited, but
because the sex of all these eight adult speci-
mens could be determined (two by removal
from the shell, the rest by external inspection
enabled by the transparency of the shells), it
seems worthwhile including them in the data.

The measurements show a range of devia-
tions that is higher than in the populations of
the similarly sized Helicina escondida n. sp.
for which a comparable number of specimens
was analyzed. À sexual dimorphism is indi-
cated with the females being bigger than the
males, but the data overlapping slightly. The
differences for height and minor diameter in
females and males amount less than in such
species as H. gemma and H. beatrix and re-
semble those of H. escondida п. sp. In inter-
polation from the minor diameter, males have
a volume of about 75% that of females.

Habitat
The type locality is located in an area charac-
terized by montane rain forest. The field notes

of Alexander Alvarado Mendez state that the
specimens were found in very humid, primary

height (n=5/3)

minor diameter (n=5/3)

outer lip (n=5/3)

—
last whorl (n=5/3)
—
columellar axis (n=5/3)
1 1 1 1 1 1 1
0 1 2 3 4 5 6 [mm] 7
FIG. 85. Measurements of Helicina

echandiensis n. sp. according to Table 7; on
each line: mean value, standard deviation,
absolute range; number of individuals given as
“п = females/males “; upper line: females,
lower line: males; in between and shaded:
average of both.

276

RICHLING

TABLE 7. Measurements of Helicina echandiensis n. sp.
given as mean value with standard deviation, minimum
and maximum value (min, max), and number of
specimens (min./max. diam. = minor/major diameter, col.
axis = columellar axis); linear measurements [mm].

“Cerro Echandi” (altitude 2840 m)
lots INBio 3428246, 3542520, 3542521

Mean
Sex value Deviation Min Max Number
Height f 6.75 034 6.28 718 5
Height m 6.24 0.22. 592 6.53 3
Mai. diam. f 6.40 0:23 6:02 681 5
Maj. diam. m 5.78 0.21 5:50 6.10 3
Min. diam. f 5.87 0.22 559 76.15 5
Min. diam. m 5.34 023 508 568 3
Outer lip f 427 0.19 4.00 4.55 5
Outer lip m 3.19 016 5352 392 3
Last whorl + 5.09 0.25, 478 545 5
Last whorl m 4.62 0.14 4.41 4.79 3
Col. axis fl 55 0.22 486 5.44 5
Col. axis m 4.79 0.237 24507 513 3
[86° \ / N 85 u [84° 1 — В И |
u ae En
11°
| |
| = их =
| | 3 |
3500 - 4000 m
3000 - 3500 т a ee =
2500 - 3000 т |
2000 - 2500 m |
1500 - 2000 m |
1000 - 1500т |. у |
500-1000m | |
100 - 500 m И. |
@ coll.
aa: 0199 m e coll. INBio
| | © others |

FIG. 86. Records of Helicina echandiensis п. sp. т Costa Rica.

CLASSIFICATION OF HELICINIDAE 2TH

forest on black soil. The undergrowth mainly
consisted of Heliconiaceae. Considering the
shell color and the habitats of comparable spe-
cies, it seems likely that Helicina echandiensis
n. sp. was also found on these plants.

Distribution (Fig. 86)

Helicina echandiensis n. sp. is Known only
from the southern slopes of Cerro Echandi a
little below the summit, from altitudes of 2,100
to 2,840 m. The area is part of the central
mountain chain of the Cordillera de Talamanca.

Discussion

Helicina echandiensis n. sp. is unique in its
combination of characteristics. It can be distin-
guished from the other species with a bright
reddish-orange outer lip — H. gemma, H.
beatrix riopejensis n. subsp. — by the straight
and uncurved form of the latter and the surface
structure of oblique diverging grooves. Among
helicinids of this shape and shell surface tex-
ture it is comparable in size only to H.
escondida n. sp., which has a light yellowish
outer lip, a less pronounced surface structure,
and more convex whorls. Furthermore Н.
escondida n. sp. lacks the characteristic diagonal
white stripes and seems to be restricted to the
Caribbean side of the central mountain chains.

Helicina (Tristramia) punctisulcata
cuericiensis
Richling, n. subsp.

Type Material

Holotype: INBio 3542622 (leg. A. Picado,
19.01.1996)

Paratype: INBio 3542541, female (09°33'19"N,
83°40'13"W, 2,600 m as: colectado

mediante sombrereta [collected by beating
vegetation], leg. В. Gamboa, 29.10.1995)
Dimensions:
Holotype: 5.9/6.5/6.8/5.8/4.1/4.4/4.8 mm
Paratype: 7.9/7.3/7.7/6.8/4.5/5.6/6.5 mm

Type Locality

Central Costa Rica, San José Province,
Cordillera de Talamanca, Estación Cuerici, 4.5
km E de Villa Mills, Sendero el Mirador,
09°33’28"М, 83°40'13"W, 2,700 m a.s.l.

Type Material of Relevant Taxa
Helicina punctisulcata von Martens, 1890

Helicina punctisulcata von Martens, 1890: 36-
37, pl. 1, fig. 10

Type Material: Lectotype ZMB 103326a: leg.
H. H. Smith, additional paralectotypes ZMB
103326b, ZMB 103326c, ZMB 103325
Von Martens based the description on mate-
rial collected by H. H. Smith, which 1$ in the
collection of the ZMB. Four specimens from
ZMB 103326 were marked to be figured by
von Martens, of which only one matches the
measurements given in the original descrip-
tion, the other being much smaller (about 1.3
to 1.8 mm smaller in the greater diameter).
Furthermore, it best fits his upper right basal
view, with a minute groove in the columellar
region. This specimen 1$ herein selected as
lectotype (Fig. 87).

Dimensions (height/greatest diameter/minor
diameter):
Lectotype: 7.2/8.9/7.8 mm

Type Locality: “W Mexico: Omilteme, 8000 ft.
on the Sierra Madre del Sur, State of
Guerrero, Pacific side of the main cordillera”

FIG. 87. Helicina punctisulcata, lectotype, ZMB 103326a, height 7.2 тт; scale bar 2.5 mm.

278 RICHLING

FIG. 88. Helicina punctisulcata zunilensis, holotype, ZMB 103324, height 9.2 mm; scale bar 2.5 mm.

Helicina punctisulcata zunilensis Wagner, single specimen matching the figure is the
1910 holotype (Fig. 88).
Dimensions (height/greatest diameter/minor
Helicina punctisulcata zunilensis Wagner, diameter):
1910a: 295, pl. 59, fig. 9 Holotype: 9.2/10.2/8.7 mm
Type Material: Holotype ZMB 103324 Type Locality: “Vulkan Zunil in Guatemala”
Because the original description refers to [Guatemala, at border of Quezaltenango and
one specimen in the museum in Berlin, the Solola departments, volcano Volcán Zunil]

FIGS. 89, 90. Helicina punctisulcata cuericiensis n. subsp. FIG. 89. Holotype, INBio 3542622, height
5.9 mm. FIG. 90. Paratype, INBio 3542541, height 7.9 mm; scale bar 2.5 mm.

CLASSIFICATION ОЕ HELICINIDAE 279

“A

|
|

FIG. 91. Axial cleft and muscle attachments of
Helicina punctisulcata cuericiensis n. subsp.,
INBio 3542622; scale bar 2.5 mm.

Etymology

The name refers to the origin of the species,
the Cerros Cuerici.

Examined Material

INBio COLLECTION

San José: Estación Cuerici: Sendero el
Mirador, 4.5 km E de Villa Mills, 09°33’28"N,
83°40'13"W, 2,700 m a.s.l.: leg. A. Picado,
19.01.1996 (INBio 3542622); 19.01.1996
(INBio 3542527); colectado en una planta
[collected on a plant] 26.06.1996, leg. B.
Gamboa (INBio 3544828); 2,750 m a.s.l.:
leg. A. J. Mora, 27.11.1995 (INBio 3542528);
09°33’19"N, 83%40'13"W, 2,600 m a.s.l.:
colectado mediante sombrereta [collected
by beating vegetation], leg. B. Gamboa,
29.10.1995 (INBio 3542541); recolectado en
una rubiaceae caminando [collected crawl-
ing on a Rubiaceae], leg. A. Picado,
26.08.1995 (INBio 3542539)

Description

Shell (Figs. 89, 90, 335L—M): Conical, solid,
medium to small sized and only slightly
shiny to dull. Color: apex and upper whorl
dark yellow, becoming lighter with growth
and increasingly whitish spotted, towards
the aperture changing to whitish with small
yellowish spots. Surface textured with ir-
regular growth lines and oblique grooves of
different individual orientation but of the

same general direction (Fig. 92), causing the
dull appearance; last two whorls with 3-4
equally spaced spiral grooves. Embryonic

FIG. 92. Teleoconch surface structure of Helicina

punctisulcata cuericiensis п. subsp. A. 2" whorl,

раму eroded. В. 4" whorl; scale bar 100 um.

280 RICHLING

FIG. 93. Embryonic shell of Helicina punctisulcata
cuericiensis п. subsp.; scale bar 100 um.

shell with about 1 whorl; 3°/, (3°/.—4%) sub-
sequent whorls straight, the last whorl very
slightly angulated at the periphery and
rounded below; whorls equally extending in
size, forming a very regular, pointed spire.
Suture moderately impressed. Aperture ob-
lique and straight, last whorl slightly de-
scending towards aperture and inserting
below the periphery. Outer lip yellowish, re-

REA

FIG. 94. Operculum of Helicina punctisulcata FIG. 95. Radula of Helicina punctisulcata
cuericiensis n. subsp., INBio 3542622; scale bar cuericiensis n. subsp. A. Centrals. B. Comb-
1 mm. lateral. C. Marginals; scale bar 50 um.

CLASSIFICATION OF HELICINIDAE 281

markably thickened and equally expanded,
edge appearing rounded. Transition to col-
umella protruding, forming a blunt edge with
a denticle. Columella very short and curved,
transition to the body whorl with sharply im-
pressed line. Basal callus well developed,
very pronounced in umbilical area and finely
granulated.

Internal Shell Structures: (Fig. 91)

Teleoconch Surface Structure (Fig. 92): In all
of the few available specimens, the begin-
ning of the teleoconch is eroded. On the
second whorl, the surface is sculptured with
oblique diverging grooves continuing
throughout the whole teleoconch. This pat-
tern is interposed with distinct, irregular
growth lines (Fig. 92B) and spiral grooves,
which are characteristic for Helicina
punctisulcata cuericiensis п. subsp.

Embryonic Shell (Fig. 93): Only a single speci-
men could be studied. The structure 1$ simi-
lar to that of Helicina funcki. As in H.
echandiensis п. sp. the diameter 15 relatively
very large.

Diameter: 1,038 um (+ 15) (1,000-1,060) (п =
5) (INBio 3544828, 3542541, 3542539,
3542528, 3542622).

FIG. 96. Female reproductive system of Helicina
punctisulcata cuericiensis п. subsp., INBio
3542528; scale bar 1 mm.

Operculum (Fig. 94): Very slightly calcified,
calcareous plate covering only part of the
outer surface. Color horny-amber, at
nucleus nearly transparent. Columellar side
slightly regularly S-shaped, upper end acute
and pointed, lower end rounded, but slightly
truncated.

Animal: In preserved specimens, the soft body
15 greyish-black throughout, only towards the
sides and underside of the foot the color
becomes lighter to whitish-yellowish. The
dark color of the mantle gives the shells of
live specimens a greenish tinge.

Radula (Fig. 95): Only two specimens were
investigated. The B-central bears 8 well de-
fined cusps, whereas A- and C-central may
be a little crenulated. Comb-lateral with 8-11
cusps, cusps on marginals slowly increasing
in number. Radula with about 60 rows of
teeth.

Female Reproductive System (Fig. 96): The
V-organ is comparatively slender, the oblong
receptaculum seminis joins its descending
limb about the middle of the inner side. The
bursa copulatrix is relatively large and com-
pact, К is subdivided in a few simple lobes.
The provaginal sac is oblong and finely ir-
regularly lobed at its distal side, a stout,
short duct continues to the reception cham-
ber. The sac is blackish pigmented.

Morphometry and Sexual Dimorphism

The amount of material is too limited to be
analyzed. The shape of the holotype 1$ герге-
sentative for all other specimens except for
the paratype, which is higher elevated and
more evenly rounded at the periphery. The
holotype is the smallest specimen, whereas
the paratype 1$ the largest.

Two specimens dissected for anatomical
studies are females, of which one is the
paratype. The other specimen represents the
smallest of the four live-collected individuals.

Habitat

The field notes from the collectors of INBio
indicate that the species climbs on vegetation,
where it was found “en una rubiaceae
caminando [crawling on a Rubiaceae]” or by
beating vegetation. The type locality is situ-
ated in a transitional zone of montane rain for-
est to paramó vegetation.

282 RICHLING

Distribution (Fig. 97)

Helicina punctisulcata cuericiensis n. subsp.
is only recorded from the main ridge of the
northern Cordillera de Talamanca west of the
Cerros Cuerici.

Discussion

The specimens were tentatively classified as
a new subspecies of Helicina punctisulcata
because of the resemblance to this species
and its subspecies H. p. zunilensis in the shell
surface structure, color, shell thickness, and
development of the outer lip, with a protruding
denticle and the impressed line near the col-
umella. In fact, differences are only shell
shape, size, and color detail. The compara-
tively widely spaced spiral grooves are the
most remarkable common feature that also
distinguishes the “punctisulcata”-group from
other species of Helicinidae of similar size.
The only exception is H. raresulcata L. Pfeiffer,
1861, differing in a more globose, slightly

=" |

[841 о 83

angulated and shouldered shape of the shell
(rather similar to H. merdigera see under H.
monteverdensis п. sp.), which furthermore
occurs on the Caribbean side of Mexico in
Veracruz, whereas the subspecies of the
“punctisulcata”-group all originate from high
altitudes in the Pacific or Central mountains.
Besides the types of H. punctisulcata, similar
spirally grooved specimens come from El Sal-
vador (Laguna de las Ranas, 1,730 m a.s.l.,
leg. A. Zilch, 16.7.1951, SMF), Guatemala,
Huehuetenango Department (5 km W of
Aguacatan, 15°20'44"N, 91°23'03"W, 1,910 т
а.$.1., leg. Е. С. Thompson et al.UF 190472,
UF 190225), and Honduras, Santa Barbara
Department (Cerro Santa Barbara, ridge
above El Cedral, 14°54’55"N, 88°07’30"\М,
2,800 m a.s.l., leg. J. Polisar, UF 242644). As
can be seen, all these specimens also come
from high elevations.

Helicina p. cuericiensis n. subsp. is smaller
and has a more intense yellow color. Contrary
to H. р. zunilensis, the spiral grooves are re-
stricted to the upper half of the whorls. The

3500 - 4000 m

3000 - 3500 m pe EE
2500 - 3000 m
2000 - 2500 m
1500 - 2000 т
1000 - 1500 т
500 - 1000 т
100 - 500 т
0- 100 m

09°

Le coll. IR
| e coll. INBio
| © others

FIG. 97. Records of Helicina punctisulcata cuericiensis n. subsp. in Costa Rica.

CLASSIFICATION OF HELICINIDAE 283

nominal subspecies appears to be consis-
tently less elevated (broader than high), and
the lower margin of the ощег lip is less рго-
truding. The few Costa Rican specimens vary
remarkably in the relation of height and diam-
eter. But lacking more material for either of the
subspecies, the extent of variations, the distri-
bution and ecological data remain only
fragmentarily known, which is why the sub-
specific classification is maintained, although
H. p. punctisulcata diverges more strongly
from the two southern subspecies. Assuming
a restriction to higher altitudes, the three sub-
species are separated by the low elevations at
the Isthmo de Tehuantepec and the Nicara-
guan depression respectively.

An additional single specimen from Costa
Rica (San José Province, Parque Nacional
Chirripó, Fila Cementerio de la Máquina, 4 km
E de San Gerardo, 09°27'49"N, 8333'40"W,
2,200 т a.s.l., leg. Alexander Alvarado
Mendez, 08.10.2001, INBio 3428245) 1$ larger
than H. punctisulcata cuericiensis n. subsp.
and is similar to H. punctisulcata zunilensis,
but, due to the lack of sufficient material, its
proper determination must await further com-
parative data.

Helicina (“Gemma”) beatrix beatrix
Angas, 1879

Helicina beatrix Angas, 1879: 484, pl. XL, fig. 13

Helicina beatrix — Pilsbry, 1891: 332

Helicina flavida var. — von Martens, 1890: 39

Helicina beatrix — Fischer & Crosse, 1893: 435

Helicina flavida var. beatrix — von Martens,
1900: 606: E-Costa Rica: Talamanca (Pittier);
Valley of Туз [about 09°51’М, 83°35 W]

(Pittier & Biolley); Santa Clara, 200 m [7.5
km NW of Upala, about 10°56’N, 85°05'W,
Alajuela Province] (Biolley); Valley of Alta
Coca, near Talamanca, 1,000 m (Pittier)
[probably referring to Alto Coén, recently
called San Jose Cabécar, about 09°30'35"N,
83°08'22"W, 500 m a.s.l., Limón Province];
between Uiskur and Mokri [not localized],
Alta Talamanca, further in Alta Uren [Alto
Uren: 09°23’50"N, 82°59'02"W, 900 m а.5.1.,
Limon Province], and between Ukatschka
and Bruschik, in Alta Taruria [Alto Tararia,
about 09°14’30"N, 83°00'30"W, 2,500 m a.s.!.
or downstream, Limon Province] (Pittier)

Alcadia (Lelalcadia) beatrix — Wagner, 1908:
83-84, pl. 14, figs. 19-22

Oligyra (Succincta) beatrix beatrix — Baker,
1922a: 45

Helicina (Oligyra) beatrix — Pilsbry, 1926a: 59,
69, fig. 3A, 71: Panama: Bocas del Toro:
Mono Creek (Olsson)

?Helicina beatrix — Pilsbry, 1926b: 127: Costa
Rica: La Emilia, < 100 ft. [not localized] (Rehn)

Helicina beatrix — Monge-Nájera, 1997: 113:
Costa Rica [in part]

Original Description

“Shell conical, solid, shining; as seen through
the lens, very finely transversely striated;
whorls 6, very slightly convex, the four upper-
most chestnut, the fifth dark red, with an
opaque whitish band below the suture, the last
pale olive-green, with a similar opaque band at
the suture; outer lip thickened, a little expanded
and reflexed; aperture quadrately semilunate.
Var. Smaller and straw-coloured thoughout.
Diam. 4'/,, alt. 5 lin.

FIG. 98. Helicina beatrix, lectotype, BMNH 1879.7.22.29, height 10.1 mm; scale bar 2.5 mm.

284 RICHLING

Very few specimens. “Found only on the hills
up to an elevation of 2,500 feet. Animal dark
grey above, sides and foot white” (Gabb). Ap-
proaches H. heloisae, Sallé, but larger and
much more conical.”

Type Material

BMNH 1879.7.22.29-31 (leg. Gabb)
Angas (1879: 475) stated that his type mate-
rial would be placed in the collection of the
British Museum, the lot is labeled with “type”.
Of the three specimens, one represents the
dark red opaque whitish banded typical form,
the other two the straw colored variety sepa-
rated by the author. Therefore, the latter two
specimens are not regarded as syntypes. The
remaining specimen (BMNH 1879.7.22.29),
also perfectly matching the figure in the origi-
nal description, is here selected as lectotype
of Helicina beatrix (Fig. 98). It still bears its
operculum and was probably collected live.
The two other specimens (one with operculum
inside) are much smaller and show a whitish
to slightly yellowish color (perhaps faded since
the description of Angas), and in one a slight
whitish subsutural banding is visible. They are
regarded here as H. beatrix confusa.
Dimensions:

Lectotype BMNH 1879.7.22.29:

10:1/8.3/8.9/7 .7/5.3/7.1/8.2 mm;

BMNH 1879.7.22.30-31 Helicina beatrix var.

sensu Angas, now referred to H. beatrix

confusa:

7.3/6.9/7.2/6.3/4.3/5.5/5.7 mm

7.0/6.3/6.6/5.9/4.1/5.4/5.6 mm

Type Locality

“Costa Rica, only on the hills up to an eleva-
tion of 2,500 feet”.

Type Material of Synonymous Taxa or Similar
Species

Helicina beatrix nicaraguae (Wagner, 1908)

Alcadia (Leialcadia) beatrix nicaraguae
Wagner, 1908: 84, pl. 14, figs. 23-24

Type Material: MIZ 8408: “Nicaragua”
Wagner did not refer to any type material,
but his collection contains only one lot with
two specimens. It is labeled to be figured
and the larger shell perfectly matches the
drawing. It is here selected as lectotype
(Fig. 99). The paralectotype (MIZ 8408b) is
not fully grown.

Dimensions:
Lectotype MIZ 8408a:
10.2/8.7/9.1/8.1/5.3/7.3/8.1 mm

Type Locality: Nicaragua
Unfortunately, the locality Nicaragua is not
further specified on Wagner's rewritten label,
and an original label is lacking, therefore, it
cannot be traced further.

Examined Material
Lee. |. RICHLING

Limon: W Guayacan, abandoned banana
plantation, 10%01'53"N, 83°32'14"W, 520 т

FIG. 99. Helicina beatrix nicaraguae, lectotype, MIZ 8408a, height 10.2 mm; scale bar 2.5 mm.

CLASSIFICATION OF HELICINIDAE 285

a.s.l., 03.09.1999: (IR 1078); (IR 1081);
12.09.1999: (IR 1087); 15.03.2000: (IR
1360); 17.03.2001: (IR 1606)

INBio COLLECTION

Limón: Suerre de Jiménez, 10%11'31'N,
83"44'49"W, 330 т a.s.l., leg. Richard
Helling, 26.02.1994: 1 ad. (INBio 1467201)
Reserva Biológica Hitoy Cerere, Sendero
Bobocara, 09°40’53"М, 83°04’09"W, 798 т
a.s.l., leg. Alexander Alvarado Mendez,
17.06.1999: 1 ad. (INBio 3542522)

Cartago: (determination uncertain) ?Parque
Nacional Barbilla, bosque secundario,
09°57’52"N, 83°26°59"W, 400 m a.s.l., leg.
malacological staff of INBio, 12.01.2001: 1
ad. (INBio 3324279)
?Zona Protectora Rio Pacuare, Sector de la
Estaciön de Barbilla, 09°58’50"М,
83°27 08"W, 500 т a.s.l., leg. Alexander
Alvarado Mendez, 05.09.2000: 1 s.ad.
(INBio 3542905)
Zona Protectora Río Pacuare, Las Brisas de
Pacuarito, 10°02’00"N, 83°28’00"W, 400 т
a.s.l., leg. malacological staff of INBio,
29.04.2001: 1 ad. (INBio 3418572)

OTHER SOURCES

COSTA RICA

Limon: Finca Los Diamantes, 1,000 ft. [about
10°11'N, 83°37’W], leg. А. Starrett,
22.08.1963: 1 ad. (UF 243509)
Entre Ukatschka et Brushik, Haut Tararia
[about 09°14’30"N, 83°00’30"W, 2,500 т
a.s.l. or downstream, Limón Province], leg.
H. Pittier, 1X.98 (ZMB 103251)

San José: Carillo [?about 10°09’N, 8357'W],

coll. Е.К. Sykes (ВММН)

Cartago: Turrialba [about 09°54’30"N,

83°41'W], coll. H. Jaeckel: 1 ad. (SMF
209575/1); coll. H. Rolle, coll. С. Bosch: 3
ads., 1 s.ad. (SMF 180668/4); coll. Rolle: 3
ads. (ZMB 103812); Plattino, Turrialba
[about 09°54’30"М, 83°41’W], leg: University
of Alabama, M. Smith coll. (MS-15183): 4
ads. (UF 95337)

Тиз [about 09°51’N, 83°35’W], leg. H.
Pittier: 1 ad. (ZMB 103252)

Costa Rica, without locality further specified:

coll. Wagner (MIZ 8407); leg. P. Biolley: 3
ads., 1 s.ad. (МНММ)

Description

Shell (Figs. 100, 336A): Conical-globose,

solid, medium sized and shiny. Color: upper
whorls chestnut to reddish-brown, getting
darker from apex down, towards last whorl
changing to pale olive-green-greyish, to-
wards aperture even opaque, in 2.5 last
whorls an opaque whitish band directly be-
low suture. Periostracum very thin, shiny
and smooth, except for very fine growth
lines. Embryonic shell with about 1 whorl;
4°/.—5 (lectotype: 5) subsequent whorls very
slightly convex; last whorl equally rounded at
periphery; upper whorls more rapidly ex-
tending in size; whorls rapidly descending,
forming a high spire. In the area of the band
surface more inflated. Suture slightly im-
pressed. Aperture oblique and in its middle
part remarkably curved backwards. Outer lip

FIG. 100. Helicina beatrix beatrix, Guayacán, IR 1087, height 9.4 тт; scale bar 2.5 mm.

286 RICHLING

FIG. 101. Axial cleft and muscle attachments of
Helicina beatrix beatrix, IR 1087; scale bar 5 mm.

always whitish-opaque, similar to the band,
thickened and very narrowly reflexed; tran-
sition into columella continuous without any
notch or only a very small one. Basal callus
weakly developed and nearly completely
smooth or very little granulated.

Internal Shell Structures: (Fig. 101)

Teleoconch Surface Structure (Fig. 102): The
section of the transitional structure encom-
passes about the first half whorl. A very short
zone structured with oblique diverging

FIG. 102. Teleoconch surface structure of Helicina
beatrix beatrix, 2" whorl; scale bar 500 um.

FIG. 103. Embryonic shell of Helicina beatrix
beatrix; scale bar 100 um.

grooves a replaced by fine growth lines con-
tinuing up to the aperture.

Embryonic Shell (Fig. 103): The spirally ar-
ranged pits are consistently much smaller
than in Helicina funcki, and the interspacial
distance exceeds the diameter of the pits.
The pattern appears much finer, the smooth
surface 1$ more prominent.

FIG. 104. Operculum of Helicina beatrix beatrix,
IR 1087; scale bar 1 mm.

CLASSIFICATION OF HELICINIDAE 287

Diameter: 963 рт (+ 33) (900-1010) (п =
15) (IR 1078, IR 1081, IR 1087, IR 1360, IR
1606); 1,040 um (BMNH 1879.7.22.29, lec-
totype).

Operculum (Fig. 104): Very slightly calcified,
calcareous plate covering only part of the
outer surface, thickened towards the col-
umellar side. Color horny-amber, only near
the columella whitish, but still somewhat
transparent. Columellar side slightly S-
shaped, both ends acute, upper end
pointed, lower slightly rounded.

Animal (Fig. 338A): The soft body 1$
unicolored, whitish yellow throughout, only
the tentacles may show a tinge of grey, the
mantle is whitish pigmented. There is no
trace of any dark spots.

Radula (Fig. 105): Because the radulae of the
different subspecies are very similar, they
are treated under Helicina beatrix beatrix.
Central A to С may occasionally bear a few
cusps, the B-central most frequently. Comb-
lateral with 6-8 denticles, only two aberrant
forms with a plain edge or 13 cusps respec-
tively. Cusps on marginals rapidly increasing

FIG. 105. Radula of Helicina beatrix beatrix. A. FIG. 106. Female reproductive system of Helicina
Centrals. B. Comb-lateral. C. Marginals; scale beatrix beatrix, IR 1087; scale bar 1 mm.
bars 50 um (А, В), 100 um (С).

288 RICHLING

FIG. 107. Variability of the female reproductive
system of Helicina beatrix beatrix, IR 1087; scale
bar 2.5 mm.

CE (n=7/5) H. beatrix DEAN 9

Rio Peje (n=92/61) H. beatrix riopejensis

Uatsi (n=11/13) H. beatrix confusa

Shiroles (n=18/10)
yy Li ii -_

Hitoy Cerere (n=10/6)

Hitoy Cerere - Miramar (n=2/1)

Guayacan (n=7/5) H. beatrix beatrix
E u + =
o
Río Peje (n=92/61) H. beatrix riopejensis
Uatsi (n=11/13) H. beatrix confusa
——

Shiroles (n=18/10)
|
$$
Hitoy Cerere (n=10/6)

Hitoy Cerere - Miramar (n=2/1)

7
Turrialba (n=10) H. beatrix beatrix
Е

Lectotype (H. beatrix)

Es EEE
| BMNH 1879.7.22.30-31 (n=2) Н. beatrix confusa
py] fm a

Brushik (type H. b. confusa)

mr —

1H 1 1 1 = 1 == L

0 4 5 6 7 8 9 [mm] 10

FIG. 109. Minor diameter of shell of different
populations of Helicina beatrix beatrix and
subspecies in Costa Rica according to Table 8;
for explanations see Fig. 108.

in number, only in nominal subspecies does
a change to more denticles a little more out-
wards take place, perhaps caused by of the
larger size of this form. The same 1$ true for
the number of rows of teeth: about 91-138
in HA. b. beatrix, in the other two subspecies
only about 66-79.

CE (n=7/5) H. beatrix beatrix 9

|
Rio Peje (n=92/61) H. beatrix riopejensis

Turrialba (n=10) H. beatrix beatrix :

: Uatsi (n=10/13) H. beatrix confusa
Lectotype (H. beatrix)
BMNH 1879.7.22.30-31 (n=2) H. beatrix confusa Shiroles (n=18/10)
Brushik (type H. b. confusa) +!

Be Hitoy Cerere (n=10/6)
1} 1 == L 1 Vo 1 ЕЕ
0 5 6 7 8 9 [тт] 10 Hitoy Сегеге - Miramar (n=2/1)

FIG. 108. Shell height of different populations
and subspecies of Helicina beatrix in Costa Rica
according to Table 8; on each line: mean value,
standard deviation, absolute range; number of
individuals given as “n = females/males or total”;
upper line: females, lower line: males if separate;
in between and shaded: average of both for
comparison with populations of unknown sex;
sex of individuals from Hitoy Cerere and Hitoy
Cerere - Miramar not determined anatomically
(see text).

Turrialba (n=10) H. beatrix beatrix
df E al
Lectotype (H. beatrix)
=f fa ET
BMNH 1879.7.22.30-31 (n=2) H. beatrix confusa
ES e
Brushik (type H. b. confusa)
Easy [sr
1 L 1 st 1 1 1
0 dE 2 4 5 6 7 [mm] 8

FIG. 110. Expansion of outer lip of different
populations and subspecies of Helicina beatrix in
Costa Rica according to Table 8; for explanations
see Fig. 108.

CLASSIFICATION OF HELICINIDAE 289

Guayacán (n=7/5) H. beatrix beatrix 9
[ a SR 7

Rio Peje (n=92/61) H. beatrix riopejensis

Uatsi (n=11/13) H. beatrix confusa

Shiroles (n=18/10)

Hitoy Cerere (n=10/6)
SK
—

>

Hitoy Cerere - Miramar (n=2/1)

H. beatrix beatrix

Turrialba (n=10)
= Es

one (H. beatrix)

SAUS —
H. beatrix confusa

BMNH 1879.7.22.30-31 (n=2)

au (type H. b. confusa)
I

/ L 1 JE L 1 |
0 2 3 4 5 6

7 [mm] 8

FIG. 111. Height of last whorl of different
populations and subspecies of Helicina beatrix
in Costa Rica according to Table 8; for
explanations see Fig. 108.

Female Reproductive System (Figs. 106,
107): The ascending limb of the V-organ is
relatively short and stout. The receptaculum
seminis is rather large. The bursa copulatrix
consists of a few irregularly shaped lobes;
the provaginal sac is somewhat inflated and
shows a simple outline, the stalk is short.
The pallial oviduct is strongly constricted.

Guayacán (n=7/5) H. beatrix beatrix
SES —
— o

Río Peje (n=70/49) H Beaux riopejensis

H —" confusa

EE I

Hitoy Cerere (n=10/6)

Uatsi (n=11/13)

Shiroles (n=17/9)

Hitoy Cerere - Miramar (n=2/1)

H. beatrix beatrix

Turrialba (n=10)
= =

Lectotype (H. beatrix)

——
H. beatrix confusa

BMNH 1879.7.22.30-31 (n=2)
Ey =

Brushik (type H. b. confusa)
y Er

1 | И 1 1 1 1
0 2 3 4 5 6

7 [mm] 8

FIG. 112. Height of columellar axis of different
populations and subspecies of Helicina beatrix in
Costa Rica according to Table 8; for explanations
see Fig. 108.

Guayacan (n=7/5 H. beatrix beatrix
y ) р 9
—— ©
Ко Peje (n=91/61) H. beatrix riopejensis
—
Uatsi (n=11/13) H. beatrix confusa
en 505
Shiroles (n=18/10)
—-
iL „te 1 = a | L 1
0 0.04 0.08 0.12 0.16 0.2 [mi] 0.28

FIG. 113. Shell volume of different populations and
subspecies of Helicina beatrix in Costa Rica
according to Table 8; for explanations see Fig. 108.

Morphometry and Sexual Dimorphism (Table
8, Figs. 108-114)

For comparison, the different subspecies are
all discussed in conjunction with the nominal
subspecies. The material available for
Helicina beatrix beatrix remains very scanty,
although during the field work several efforts
were made to find the species in greater abun-
dance and at different localities. The only
specimens studied anatomically are those
from Guayacán. Because the “Turrialba”
population is united from lots of three different
collections, they may originate from different
localities around Turrialba.

The specimens of H. beatrix confusa included
from the INBio collection (Hitoy Cerere, Hitoy
Cerere — Miramar) could not be analyzed for
their sex by dissection. To make them never-
theless available for morphometric comparison,
the degree of sexual dimorphism found in the
dissected populations was used to attribute the
probable sex to the specimens in reverse con-
clusion (see below). These data cannot prima-
rily be used to investigate sexual dimorphism.

Guayacán (n=7/5) H. beatrix beatrix

A

$
o

Rio Peje (n=91/61) H. beatrix riopejensis

Uatsi (n=11/13) H. beatrix confusa

Shiroles (n=18/10)

1 1

1 1 4 1 y
0 0.02 0.04 0.06 0.08 0.1 [9] 0.14

FIG. 114. Shell weight of different populations
and subspecies of Helicina beatrix in Costa
Rica according to Table 8; for explanations see
Fig. 108.

290 RICHLING

TABLE 8. Measurements of different populations and subspecies of Helicina beatrix given as mean
value with standard deviation, minimum and maximum value (min, max), and number of specimens;
sex of individuals from Hitoy Cerere and Hitoy Cerere — Miramar not determined anatomically (see
text) (min./max. diam. = minor/major diameter, col. axis = columellar axis); linear measurements [mm],
weight [g], volume [ml].

Helicina beatrix beatrix Helicina beatrix riopejensis п. subsp.
“Guayacán” (altitude 520 m) “Río Peje” (altitude 135 m)
lots IR 1078, IR 1087, IR 1081 lots IR 440, IR 752, IR 1303, IR 1550
Mean Mean

Sex value Deviation Min Max Number value Deviation Min Max Number

Height Е 9.59 0.16 931 9.88 7 1.202 0:26 695 85372092
Height т 8.39 0.13 8.26 8.71 5 6.51 0.20 6410!) 7-9 6H
Maj. Чат. + 8.67 0.22 8.33 9.08 7 6.89 0.19 6.00 7.93 92
Mai. Чат. т 7.75 0.08 7.64 7.88 5 5.97 015 546.6 3361
Min. diam. f 8:12 0.21 779 857 mA 6.39 0.18 5.5 700 92
Min. dam. m 7.15 0.10 702 7.32 5 5:50 0.14 5.20 597 61
Outer lip f 5.61 012 531 57 7 4.45 012 3.757 290 92
Outer lip m 5.13 007 503 522 5 3.98 0.13 3.68 4.40 61
Last whorl f 1:45 0.13 7.28) 7565 7 5.83 0.18 525 64292
Last whorl m 6.58 0.06 6.49 6.69 5 5.03 0.19 4.53 5.47 61
Col. axis f 7.83 0.13 756 8:01 7 6.28 022 559 6988170
Col. axis m 6.88 0.15 6.75 7.24 5 5.24 0.19 4:85 518 29
Weight f 0.069 0.014 0.046 0.092 7 0.035 0.005 0.018 0051 91
Weight т 0.053 0.018 0.034 0.082 5 0.027 0.003 0.018 0.044 61
Volume г 0232 9.014 0212 0.253 7 0.115 0.009 0.084 0.147 91
Volume m 0.156 0.006 0.145 0.169 5 0.071 0.005 0.058 0.089 61
Helicina beatrix confusa
Helicina beatrix confusa “Shiroles” (altitude 120 m)
“Ца! (altitude 30 т) lots IR 910, IR1327, IR 1594, IR 1600,
lots IR 1112, IR 1113, IR 1567 IR 1646
Mean Mean
Sex value Deviation Min Max Number value Deviation Min Max Number
Height f 8.17 028 159 3850 AM 116 0.17 140 93718
Height m 710 Оу 6721750 218 6.35 017 607 667 110
Ма]. Чат. + 7:38 07 7.201759 11 7.08 016 67’ ASIS
Mai. diam. m 6.63 0.16 630 6.94 13 6.03 0.13. 580 6.22880
Min. diam. f 6.93 010 1675676 11 6.63 017 635 696588
Min. diam. m 6.14 Ont, 593 637 13 5.61 O12 5.43 5688110
Outer lip f 4.81 0.08 461 5.03 10 4.67 0.12 447 500 als
Outer lip m 4.42 0.12 425 453 13 4.04 010 387 24250
Last whorl f 6.35 016 610 5656 11 6.09 0.14 572 648883
Last whorl m 5.64 0.09 542 584 13 5:05 0.18 ‘472 5A)
Col. axis f 6.68 025 630 108 Zi 6.32 0.16 590 696 217
Со! ах п 5.77 O15 5:47 6.06 13 5.07 0.17 2479 550 9
Weight f 0.050 0.009 0.040 0.067 11 0.047 0.008 0.031 0.064 18
Weight m 0.038 0.004 0.024 0.048 13 0.027 0.004 0.019 0.034 10
Volume Г 01243 0007 0.126 10.156 11 0.124 0.010 0.095 0.150’ 168
Volume т 0.097 0.006 0.087 0.110 13 0.073 0.004 0.063 0.079 10

(Continues)

Morphometry: The typical Helicina beatrix
beatrix clearly possesses the largest shells
among the Costa Rican subspecies of H.
beatrix. Its shells have a similar size at all
three localities. The lectotype is more highly
elevated, reflected mainly in height and
height of the columellar axis, whereas the
specimens from Turrialba have relatively the
largest minor diameter. Single specimens of
H. beatrix beatrix not included in the dia-
grams fall within the same size range.

The populations of the subspecies H. beatrix
confusa and H. beatrix riopejensis n. subsp.
show a very constant pattern of differences
between the populations for the different
measurements, displaying the same rela-
tions of the measurements. The specimens

CLASSIFICATION OF HELICINIDAE 291
(Continued)
Helicina beatrix confusa Helicina beatrix confusa
“Hitoy Cerere” (altitude 100-798 m) “Hitoy Cerere - Miramar”
lots INBio 1473618, 1473833, 1473837, (altitude 150-300 m)
1475069, 1498277, 1543340, 3096421 lots INBio 1475230, 1475695, 1476494
Mean Mean
Sex value Deviation Min Max Number value Deviation Min Max Number
Height f 7.68 0.34 7.09 843 10 7.97 0:26 77| 8.23 2
Height m 6.56 0.21 6.10 6.78 6 6.73 0.00 6.73 6.73 1
Mai. diam. f 6.93 020 647 755 10 7.01 0.04 6.96 7.05 2
Ма]. diam. m 6.04 0:25 550 646 6 6.04 0.00 6.04 6.04 1
Min. diam. f 6.56 018.632 723.10 6.68 0.08 6.60 6.76 2
Min. diam. т 5:69 OST 5.94 6 5.70 0.00 5.70 5.70 1
Outer lip Г 4.57 0.17 422 490 10 4.64 0.04 460 4.67 2
Outer lip m 3.96 0.21 369 4.30 6 4.09 0.00 4.09 4.09 1
Last whorl f 5.95 0.23 5.18 6.68 10 6.07 0.11 5.96 6.18 2
Last whorl т 5.04 0.21 4.74 5.31 6 552 0:00 7552 "5:52 1
Col. axis f 6.30 0.25. 564 7.17 10 6.54 0.33 6.21 6.86 2
Col. axis m 5.26 0.12. 75:01 5.46 6 5.59 0.00 5.59 5.59 1
Helicina beatrix beatrix
“Turrialba”
lots SMF 209575/1, SMF 180668/4, UF
95337, ZMB 103812
Mean
value Deviation Min Max Number
Height 8.81 0.47 806 9.70 10
Maj. diam. 7.96 0.42 7.43 862 11
Min. Чат. 7.39 0.38 6.68 8.00 11
Outer lip 5.24 0.11 494 5.41 9
Last whorl 6.75 0.27 6.19 7.08 10
Col. axis 7411 0.55 5.82 7.90 7

of Uatsi have the biggest shells. But they all
exhibit a smaller size than the nominal sub-
species. The relative constancy within the
populations and the nearly equal size of the
individuals from Shiroles and Hitoy Cerere
(Hitoy Cerere — Miramar) suggest a relation
to the distribution, because these localities
are closer to each other than Uatsi (Fig. 1).
Whereas in measurements the two speci-
mens of H. beatrix var. sensu Angas match
the two subspecies well, the lectotype of H.
beatrix confusa is much smaller. For the
subspecies, the lack of material from addi-
tional localities still prevents any investiga-
tion of a possible correlation of the size to
the altitude which could help to relate the
small size of the lectotype (from a much

292 RICHLING

9 Ta Se T ER a A aren T el Fee]
we ee o |
diam. + Males x
[mm] 2
SS
8 |- o |
o o
7.5 | -
ip
| |
A
7+ ae
6.5 |
6
5.5
5
6 6.5 7 7.5 8 8.5 9 height [mm] 10

FIG. 115. Range of measurements in females and males of Helicina beatrix
beatrix exemplary for height and minor diameter in the population from
Guayacan.

9
female +

min. | nalen

6 6.5 7 7.5 8 8.5 9 height [mm] 1о

FIG. 116. Range of measurements in females and males of Helicina beatrix
confusa exemplary for height and minor diameter in the population from
Shiroles.

CLASSIFICATION ОЕ HELICINIDAE

9 T T OO Eee |
Er female +
|
an male +
lam. + E
[mm]
8 + |
7.5 +
7 E o o 4
o
а
6.5 | Ñ EN -
5 o DE Po
IS % © & ©
о LES 2 И
6 o o a
o oof
. +
ны aes So
о
er te He |
4 A = y
++ ++
5 1 I A A > 1 - 1
6 6.5 7 7.5 8 8.5 9 height [mm] 10

FIG. 117. Range of measurements in females and males of Helicina beatrix
riopejensis n. subsp. exemplary for height and minor diameter in the
population from Rio Peje.

Q.

D

3
T

VEA = +

6 6.5 7 7.5 8 8.5 9 height [mm] 10

FIG. 118. Plot of measurements for height and minor diameter for individuals
of Helicina beatrix confusa of unknown sex, exemplary for the population of
Hitoy Cerere and the separation proposed.

293

294 RICHLING

higher altitude) to the recently collected
material. For H. beatrix beatrix, the present
data do not corroborate a correlation of size
with altitude. Within a range from elevations
of 330 т (Suerre de Jimenez) to 800-1,000
т (Turrialba) or even up to 2,500? т (Alto
Tararia), the size remains nearly constant.

Sexual Dimorphism: All populations and even
all measurements show a very clear differ-
ence between females and males, in many
cases not only within the range of the stan-
dard deviation but also for the whole range.
This is shown for a population of each sub-
species (Figs. 115-117) in data for height
and minor diameter, which best separates
the sexes. In populations with many indi-
viduals (e.g., Río Peje), high extrema are
more likely and result in a little overlap. In
volume, the males are only about ‘/, of that
(61% to 68%) of females.

The clear difference between both sexes
can be used to plot measurements (e.g.,
minor diameter to height) of specimens of
unknown sex (Fig. 118, Hitoy Cerere) in or-
der to attribute them to their most likely sex.
But the differences between the populations
also demonstrate that this method will only
work for specimens from one and the same
locality, and the lot from “Turrialba” could not
be separated (Fig. 119) because it does not
represent a single population.

The differences of the specimens of H.
beatrix var. sensu Angas to the lectotype of
H. beatrix are out of the range of sexual di-
morphism, supporting their exclusion from
typical H. beatrix.

Habitat

Helicina beatrix beatrix was found by the au-
thor at only one locality at Guayacán. There it
inhabits a small abandoned banana field on a
steep hillside surrounded by secondary growth
and partly swampy meadows for cattle. Speci-
mens were aestivating or on the underside of
green banana leaves or crawling in curled,
dried leaves. Originally the area was covered
by rain forest, and it seems to be a relic occur-
rence of the species. All negative records and
the few specimens in collections suggest that
H. beatrix beatrix is a rare subspecies.

Distribution (Fig. 120)

Although records are scarce, the occurrence
shows a remarkable pattern. As already ob-

served by Gabb and cited in the original de-
scription, the species is said to occur only on
hills up to an elevation of 2,500 feet. In fact
Helicina beatrix inhabits the Caribbean moun-
tain slopes of the Cordillera Central and the
Cordillera de Talamanca. The localities can be
attributed to the slopes of three regions: the
valley between the volcanoes Barva and
Irazú/Turrialba, the Caribbean side of the Valle
Central along the Rio Reventazon between
the Volcan Irazü and northern Cordillera de
Talamanca, and the Valle de Talamanca. The
verified range of altitude is from about 330 m
to 1,000 m or even up to 2,500 m depending
оп the exact location of “Alto Тагапа”. The
most northern record from Santa Clara near
the frontier to Nicaragua is uncertain.

Discussion

Helicina beatrix beatrix is understood as the
large whitish-opaque form with reddish-brown
upper whorls. The determination of two of the
three lots from the Barbilla/Rio Pacuare area
(INBio 3324279, INBio 3542905) is uncertain;
the size of the adult specimen is similar to the
nominal subspecies; the color approaches that
of H. b. confusa. A common feature of H.
beatrix beatrix and the other subspecies is the
general shape as described above and the
subsutural opaque band. Furthermore, the
outer lip is typically strongly curved backwards,
especially in females. Against the background
that intermediate forms of the subspecies are
lacking and a sympatrical occurrence implying
a specific separation is uncertain, the status of
subspecies is tentatively maintained or sug-
gested for the population from Rio Peje. But
certain hints for a sympatrical existence of H.
beatrix beatrix and H. b. confusa have to be
mentioned. The type locality of H. beatrix
confusa (Brushik, Alto Tararia) is probably close
to “between Ukatschka and Brushik, Alto
Tararia” recorded for H. beatrix beatrix, al-
though the exact location is uncertain (see be-
low). Furthermore, a recently collected lot in the
collection of INBio from the Sendero [= trail]
Bobócara in the reserve Hitoy Cerere contains
both subspecies, and the very top of the moun-
tain Cerro Bobócara is given as locality, but is
most likely not the source of all specimens. In
the extremely mountainous and steep terrain, a
short distance of the trail probably already en-
compasses different habitats at different alti-
tudes. Therefore, the data do not contribute to
an assessment of the status of H. beatrix and
subspecies.

CLASSIFICATION OF HELICINIDAE

9 = — fi = Е т MAS as
[ il
min.
diam. | -
[mm]
8 | ’ © al
7.5 | 2 y 4
7 + 6 o o -
6.5 Г -
6 | 1
5.5 | |
re июн aye = ie y y Ze
6 6.5 7 7.5 8 8.5 9 height [mm] 10

FIG. 119. Plot of measurements for height and minor diameter for individuals
of Helicina beatrix beatrix of unknown sex, exemplary for the specimens from
“Turrialba” (probably not single populations) for which a separation is not
possible.

86° \ HS о 84 À | 183°

| 3500 - 4000 т
3000 - 3500 m
2500 - 3000 m
2000 - 2500 m
1500 - 2000 m
1000 - 1500 m |

500 - 1000 m |

100 - 500 т
e coll. IR
0- 100m e coll. INBio
© others

FIG. 120. Records of Helicina beatrix beatrix in Costa Rica.

295

296 RICHLING

The type lot of H. beatrix is of unknown ori-
gin and most likely it is also composed of
specimens from different localities, because
the smaller two belong to a different subspe-
cies. The Knowledge of well-localized records
of H. beatrix is very limited, but among those,
the lot ZMB 103251 from high up in the moun-
tains from the Valle de Talamanca (southern
Caribbean side) most closely resembles the
lectotype in having a high, elevated shell. Ac-
cording to recent and previous findings, popu-
lations at Guayacän, Turrialba and Tuis are
characterized by more globular shells, thus
suggesting that the lectotype probably was not
collected in the area of the Rio Reventazôn
and its tributaries between Turrialba and
Siquirres, which was comparatively easily ac-
cessible at the end of the 19" century as the
most direct connection between San José and
Puerto Limón on the Caribbean.

The record from Santa Clara remains doubt-
ful, because it is far out of the verified distribu-
tion and the original material has not been
found. А confusion with H. gemma which oc-
curs in this region, can be excluded for three
reasons: (1) Biolley (1897) and von Martens
(1900) also reported this species for the area
as Н. ометапа anozona and von Martens
(1900) as H. oweniana coccinostoma, (2) von
Martens (1900) characterized the specimens
as no less than 10 mm in diameter and 9 in
height, whereas H. gemma displays a remark-
ably constant size of about 5 to 7 mm in
height, and (3) the orange ощег lip of H.
gemma would rather suggest an identification
as H. ометапа than H. beatrix, exactly as von
Martens obviously treated H. gemma in his
publication. Except for the doubtful record of
H. beatrix nicaraguae, the species has not
been reported from the adjacent Nicaragua. In
the face of absence of the original material
and the limited knowledge on Nicaraguan
Helicinidae, it still remains doubtful.

In Helicina beatrix nicaraguae, the whorls
very evenly increase in size, forming a regu-
lar spire, which is less inflated than in H.
beatrix beatrix. The whorls are more convex,
therefore the suture is more deeply im-
pressed. The whitish band under the suture
is less distinct and more slender. The basic
color is yellowish, with a tendency to green-
ish, towards the aperture lighter and chang-
ing to opaque.

On account of the poorly investigated Nica-
raguan terrestrial molluscan fauna, it is impos-
sible to render any judgement about the
possible origin or distribution. According to the

Costa Rican records, H. beatrix and subspe-
cies are confined to the southern Caribbean
slope and coastal plain with its most northern
record verified at about Guapiles (10°13’М), or
with uncertainty near the Nicaraguan border at
Santa Clara. The only record for Nicaragua 1$
that of Wagner (1908) of his new subspecies.
Supposing that the record from Santa Clara 1$
attributed to another species and considering
that H. beatrix and subspecies seem to be
absent from the very lowlands, that 1$, the
southern Nicaragua, there appears to be a
gap in the distribution towards Nicaragua.
Otherwise, specimens from Isla Colón (Isla
Colon, Las Gratas, 5 km NNW of Bocas del
Toro, 09°23’25"N, 82°16'15"W, 70 m a.s.l.,
leg. F.G. Thompson (FGT-4724), 17.09.1990:
2 ads. (UF 167532); interior of Colon Island,
leg. McGinty coll., 28.03.1953: 1 s.ad. (UF
185607) (Fig. 121), Bocas del Toro Province,
Panama, adjacent to the Costa Rica distribu-
tion, show a surprising similarity in shape and
color. With a height of 9.6 mm, the largest
shell nearly reaches the size of the lectotype.
An investigation of the Nicaraguan malaco-
fauna would be required to prove whether the
type locality of H. beatrix nicaraguae 1$ in this
country, or whether the lectotype in fact came
from Panama.

Von Martens (1890-1901) misinterpreted H.
beatrix as a variety of H. flavida because he
had not seen original specimens. Subsequent
authors (Pilsbry, 1891; Fischer 8 Crosse,
1893) stressed the distinctness of H. beatrix,
and von Martens agreed with this in his
supplement.

FIG. 121. Helicina beatrix nicaraguae,
Panama, Isla de Colón, UF 167532,
height 8.9 mm; scale bar 2.5 mm.

CLASSIFICATION ОЕ HELICINIDAE 297

Pilsbry (1926a) twice mentions H. beatrix
from Bocas del Toro Province, Panama: speci-
mens agreeing in size with the nominal sub-
species from a certain locality and additional
individuals not further specified in their origin
that are remarkably smaller, thus resembling
H. beatrix confusa in size. Furthermore, the
identification and localization of the Costa
Rican record by Pilsbry (1926b) remains
doubtful, because altitudes of less than 100
feet would be exceptional for the nominal sub-
species.

Helicina (“Gemma”) beatrix confusa
(Wagner, 1908)

Helicina beatrix var. — Angas, 1879: 484, pl. XL,
fig. 13 [non Helicina beatrix Angas, 1879]
Alcadia (Leialcadia) beatrix confusa Wagner,
1908: 84, pl. 14, fig. 25

Oligyra (Succincta) beatrix confusa - Baker,
1922a: 45

Helicina beatrix — Monge-Najera, 1997: 113:
Costa Rica [in part] [non Angas, 1879]

Original Description

“Gehäuse ме! kleiner [als Alcadia beatrix],
dúnnschaliger, gelbgrün mit rótlichem
Gewinde; das niedrigere, konvexe Gewinde
besteht nur aus 4'/, deutlicher gewólbten
Umgángen, der letzte ist unten weniger
abgeflacht.

FIG. 122. Helicina beatrix confusa, lectotype,
МЕ 8409, height 5.6 mm; scale Баг 2.5 mm.

D=7.3,H=7.2
Deckel wie bei der typischen Form.
Fundort: Costarica.”

Type Material

MIZ 8409: “Brusik [sic] Ht. Тагапа”
Wagner did nat refer to any type material, but
his collection contains only one specimen he
singled out as type material. It is labeled “to be
depicted” and agrees very well with the illus-
tration. The specimen is therefore here se-
lected as lectotype (Fig. 122).
Dimensions:
Lectotype: 5.6/5.3/5.7/4.9/3.4/4.2/4.4 mm

Type Locality

“Costa Rica” (figure caption erroneously
Nicaragua); restricted by the type selection to
Brushik, Alto Tararia [about 09°14’30"N,
83°00'30"W, 2,500 m a.s.l. or downstream,
Limon Province]

Unfortunately, Wagner used to rewrite most
of the labels and only occasionally retained
the original, thereby not always preserving all
information (Riedel, 2000). In the present
case, it can be referred to “Brushik, Haut
Тагапа” [Spanish: Alto Tararia], a source also
named by von Martens (1900) for material
collected by Pittier. At that time, Pittier was the
only one intensively studying the region of the
Valle de Talamanca and its adjacent mountain
slopes. Despite an intensive search for the
locality, it is difficult to rediscover it. It is known
that Pittier maintained good relations with the
indigenous Cabécar and Bribri, which inhabit
the Valle de Talamanca and settle along the
four main rivers — Rio Telire, Rio Coén, Rio
Lari and Rio Uren — and their tributaries high
up in the mountains. Throughout this region,
neither Brushik nor Alto Tararia or related
names could be found on detailed maps.
Much further south a Cerro Tararia [“cerro” =
“mountain”, 09°09’03"М, 82°58'27"W, 2,690 m
a.s.l.] exists, which most likely is not the local-
ity mentioned, because it forms a part of the
very central mountain chain and lacks any
access route. Furthermore, “Alto” followed by
a name of a river usually refers to a main
settlement along the river in the mountains, for
example, Rio Lari — Alto Lari. Therefore, Alto
Tararia may mean the upper part of the Rio
Tararia [about 09°14’30"N, 83°00’30"W, 2,500
т a.s.l. or downstream], which really exists
southeast of Cerro Kamuk. Again, it is difficult

298 RICHLING

to gain access to the region, settlements or
trails are not shown on maps. From Valle de
Talamanca, it means following the river Rio
Lari or Rio Urén to their headwaters and to
cross the Cerro Kamuk to reach the high re-
gion of Rio Tararia draining towards Panama.
Because other Pittier localities are very reli-
able, this appears to be the best interpretation,
because the material also may have been
given to Pittier by indigenous people and prob-
ably not all indigenous names and trails were
incorporated in maps, and some may even
have been forgotten or lost.

Examined Material

Lec. |. RICHLING

Limon: Southern road from Bribri to Shiroles,
small banana plantation near creek,
09735'17'№ "82°52 46 W,.. 50 "mr ais;
15.03.1997: (IR 170)
W Bribri, road to Uatsi, about 09°38'11"N,
82°51'48"W, 30 т a.s.l.: abandoned field
with Heliconiaceae and Eucalyptus:
17.03.1997: (IR 182); 12.03.1999: (IR 765);
15.09.1999: (В 112) (В 13
15.03.2001: (IR 1567); wooded valley within
banana plantation, 50 т a.s.l.: 15.03.2001:

(IR 1585)

N Shiroles: along Quebrada Kirio,
09°35'38"N, 82°57 20°W: 120° m as:
15:03. 19972" (RY 161) 100m a5:

12.03.1999: (IR 764); 09.08.1999: (IR 910);
06.03.2000: (IR: 1327; (IR. 1329);
16.03.2001: (IR 1594); (IR 1646); Сето
Mirador, along trail, 09°36’37"М, 82°57’43"W,
430 m a.s.l.: 16.03.2001: (IR 1600)

INBio COLLECTION

Limón: Parque Nacional La Amistad,
Quebrada Cachabri (toma de agua),
09°29’29"М, 82°59’37"W, 360 m a.s.l., leg.
Gerardina Gallardo, 26.11.1996: 1 ad.
(INBio 1488199)
Reserva Biológica Hitoy Cerere: Sendero
Toma de Agua, 09*40'31"N, 83°01'36"W, 100
m a.s.l.: 20.04.1994: 5 ads. (INBio 1473837);
1 ad. (INBio 1473618); 2 ads., 1 juv. (INBio
1473833); 13.08.1994: 3 ads., 2 s.ads., 13
juvs. (INBio 1475069) (all leg. Zaidett
Barrientos); Sendero Toma de Agua,
09°40’22"М, 83%01'35"W, 160 т a.s.l.: leg.
Marianella Segura, 14.07.1994: 1 s.ad.
(INBio 1478208); 1 juv. (INBio 1478209);
Sector Miramar, Hitoy Cerere, 09°37’50"N,
83*00'52"W, 300 m a.s.l.: 13.06.1994: 1 ad.
(INBio 1476494); 04.07.1994: 1 ad. (INBio
1475695) (all leg. Gerardo Carballo); Sector
Miramar, 09*38'03"N, 83°00’45"W, 300 m
a.s.l.: leg. Zaidett Barrientos, 08.10.1994: 1
ad. (INBio 1475716); Sendero a Captación
de Agua, 09*39'59"N, 83°01'31"W, 200 т
a.s.l.: leg. Alexander Alvarado Mendez,
28.04.1999: 3 ads., 2 s.ads. (INBio 1498277);
Sendero Tepezcintle, 09°40’18"М,
83°01'43"W, 140 m a.s.l.: leg. Alexander
Alvarado Mendez, 28.04.1999: 1 s.ad. (INBio
1496288); Sendero Bobócara, 09°40'53"N,
83°04'09"W, 798 т a.s.l.: leg. Alexander
Alvarado Mendez, 17.06.1999: 1 ad., 1 s.ad.
(INBio 1543340)
Reserva Indígena Talamanca, Sector
Amubri, 09°30’53"М, 82%57'19"W, 70 т
a.s./.: leg. Gerardina Gallardo, 14.06.1994: 1
ad. (INBio 1477505); 1 ad. (INBio 1477553);

FIG. 123. Helicina beatrix confusa, Shiroles, IR 910, height 7.8 тт; scale bar 2.5 mm.

CLASSIFICATION OF HELICINIDAE 299

Sector Miramar, Senderos а Rio Мот,
09°37’44"N, 83°00’32"W, 150 m a.s.l.: leg.
Zaidett Barrientos, 08.11.1994: 1 ad., 1
s.ad., 3 juvs. (INBio 1475230)

Reserva Indígena Tayni, Sendero
Tepezcuintle, 09°40'22"N, 83°01 46"W, 180
m a.s.l., leg. Alexander Alvarado Mendez,
22.04.1999: 1 ad. (INBio 3096421)

OTHER SOURCES

COSTA RICA

Limén: N of Rio Moin [Valle de Talamanca!],
012 000. Е, 397: 60005 110937 45"N,
8300'38"W], 220 m a.s.l., leg. E.L. Raiser
(ELR-086), 11.08.1994: 1 ad., 2 s.ads. (UF
41440)

Costa Rica, without locality further specified:
leg. Gabb: 2 spec. (BMNH 1879.7.22.30-
31)

Description

Shell (Figs. 123, 336B, C): Conical-globose,
rather thin, small sized, shiny. Color: upper
whorls yellowish-red, horny changing con-
tinuously to yellow at the beginning of body
whorl and getting nearly white towards aper-
ture. The opaque whitish band directly below
suture very slender. Shell surface shiny and
smooth, except very fine growth lines. Em-
bryonic shell with about 1 whorl; 4-5 (lecto-
type: 3%) subsequent whorls very slightly
convex; last whorl equally rounded at the
periphery; upper whorls slightly more rapidly
extending in size; whorls regularly descend-
ing, forming a nearly blunt spire. Suture
slightly impressed. Aperture oblique and in
its middle part remarkably curved back-
wards. Outer lip whitish-opaque similar to
the band, slightly thickened and very nar-
rowly reflexed; transition into columella con-

tinuous without a very little notch. Basal cal-
lus weakly developed and nearly completely
smooth or very little granulated, umbilical
area whitish.

FIG. 124. Axial cleft and muscle attachments of FIG. 125. Teleoconch surface structure of Helicina
Helicina beatrix confusa, IR 1113; scale bar beatrix confusa. À. Changes in the apical part. B.
2.5 mm! 2™ whorl; scale bars 500 um (A), 100 um (В).

300 RICHLING

Internal Shell Structures: (Fig. 124)

Teleoconch Surface Structure (Fig. 125): Simi-
lar to Helicina b. beatrix, but the relation be-
tween the transitional structure and the
pattern of oblique diverging grooves is re-
versed, the former nearly disappearing.

Embryonic Shell (Fig. 126): Among the speci-

mens investigated, the spiral lines are less
numerous than in the nominal subspecies.
Otherwise, the embryonic shell structure is
similar. The diameter is smaller.
Diameter: 889 um (+ 32) (800-950) (п = 21)
(IR 1113, IR 1567); 840 um (М 8409, lecto-
type); 900 pm (n = 2) (BMNH 1879.7.22.30-
31, Helicina beatrix confusa).

Operculum (Fig. 127): Very slightly calcified,
calcareous plate covering only part of the
outer surface. Color horny-amber, only near
the columella whitish, but still somewhat
transparent. Columellar side slightly S-
shaped, both ends acute, upper end
pointed, lower slightly rounded.

FIG. 126. Embryonic shell of Helicina beatrix
confusa; scale bar 100 um.

ay
4

FIG. 127. Operculum of Helicina beatrix confusa,
IR 1113; scale bar 1 mm.

Animal (Figs. 337F, G): The body color is simi-
lar to Helicina beatrix beatrix, especially
within the yellow-shelled population from
Uatsi. In some specimens from Shiroles
(more frequently in individuals with orange-
brownish tinged shells), the dorsal part of
the head region including eyes and tentacles
and the foot is more of less grey-blackish
and the mantle is greyish pigmented as well.

Radula: See Helicina beatrix beatrix.

Female Reproductive System (Figs. 128, 129):
The structures are similar to the nominal sub-
species; the bursa copulatrix bears even
fewer lobes.

/ a,
\ +
NS ZEN
=)
\

< <
=, ) < VA
Xt /
4

Y

FIG. 128. Female reproductive system of Helicina
beatrix confusa, IR 1113; scale bar 1 mm.

CLASSIFICATION OF HELICINIDAE

ee LY)
N NE
Е TS) ( )
N ==,
в ES
) uk \
= \ Ze AE
ae \ Ad
X \ = \ №.
4 N = _ /
O а y ee
GS | Ти, N
TS : AVENT)
—— \ > yoo
Sue > A

FIG. 129. Variability of the female reproductive
system of Helicina beatrix confusa, IR 1113;
scale bar 2.5 mm.
Morphometry and Sexual Dimorphism

See Helicina beatrix beatrix.

Habitat

The population “Uatsi” inhabits an appar-
ently abandoned agricultural area surrounded

301

by small banana fields. The vegetation con-
sists mainly of Heliconiaceae and some bam-
boo (Poaceae). Helicina beatrix confusa was
found on the underside of the leaves of
Heliconiaceae and occasionally on Monstera
spec. (Araceae) climbing the few big trees left
of the previous forest. By way of contrast, the
main site at Shiroles is a small creek in what
seemed to be primary forest. Snails were
crawling and aestivating on the leaves of vari-
ous small-leafed plants of the undergrowth
along the creek. Two specimens were discov-
ered in the leaf litter. Additionally specimens
were found on leaves of lower branches of big
trees within the forest. Near Shiroles, H.
beatrix confusa lives sympatrically with H.
funcki and H. escondida n. sp.

Distribution (Fig. 130)

Helicina beatrix confusa is confined to the
southern Caribbean mountain slopes of the
northern Cordillera de Talamanca and adja-
cent hilly areas, namely in the Valle de
Talamanca and Valle de Estrella. Like the
other subspecies, it seems to be absent near
the coast. A more northern occurrence 1$
questionable, a record from the Río Pacuare-

2000
1500

10

3500 -
3000 -
2500 -

1000 -
500 -

4000 m
3500 m
3000 m
- 2500 m
- 2000 m
1500 m
1000 m
0 - 500 т
0-100m

e coll. IR
| ® coll. IN
o others

Bio

FIG. 130. Records of Helicina beatrix confusa in Costa Rica.

302 RICHLING

Barbilla area at altitudes of 400 to 500 m has
tentatively been attributed to the nominal sub-
species. The foothills of the Talamanca have
otherwise only been poorly investigated, not
only because they are difficult to access, but
also because the very local and patchy distri-
bution renders the snails difficult to find. There
is evidence for a more northerly absence, at
least at lower altitudes, because several less
elevated sites at Río Siquirres, Río Pacuarito,
Río Barbilla, near the road between Siquirres
and Limón, were checked with negative re-
sults although other Helicinidae at other
places inhabiting the same habitats — H.
funcki, H. escondida п. sp., H. chiquitica, H.
gemma — were found.

Discussion

The description given above applies to the
lectotype. The specimen is adult, but the outer
lip is still not fully developed. The less el-
evated shell renders is likely to be a male, al-
though it cannot be concluded with certainty
due to the lack of comparative material from
the same locality. If this assumption is correct,

the average size of Helicina beatrix confusa
would be bigger than indicated by the lecto-
type. Nevertheless, all specimens studied are
clearly bigger. But because the specimens
morphometrically studied originate from lower
altitudes of sites not far from each other, al-
though comparatively far away from the type
locality at presumably higher altitudes, and the
shape and the mode of color is similar, they
are attributed to this subspecies. The lecto-
type looks like a reduced form. The distribu-
tion within the Valle de Talamanca and
adjacent slopes additionally supports this clas-
sification.

In the specimens recently collected, the
whitish band is like in the nominal subspecies
broader, but occasionally it can also be as
slender as in the lectotype. The color of the
whorls varies within the populations and
among them, but the outer lip and umbilical
area are constantly whitish-opaque. The indi-
viduals from Uatsi possess shells with brown-
ish whorls at the apex that during growth
change more or less quickly to a pale or more
often bright yellow color above the periphery.
At least the beginning of the last whorl is yellow

FIGS. 131, 132. Helicina beatrix riopejensis n. subsp., Rio Peje. FIG. 131. Holotype, INBio 3542625,
height 7.8 mm. FIG. 132. Paratype 1, INBio 3542626, height 6.6 mm; scale bar 2.5 mm.

CLASSIFICATION OF HELICINIDAE 303

below the subsutural whitish band. Towards the
aperture the color fades to pale yellowish-whit-
ish. Specimens from Shiroles may display a
similar color, but in many the brownish-reddish-
orange of the upper whorls does not change up
to the aperture. Similarly the color becomes
lighter below the periphery. In general, the yel-
low form seems to be more frequent.

Helicina (“Gemma”) beatrix riopejensis
Richling, n. subsp.

Type Material

Holotype: INBio 3542625, female (leg. 1.
Richling, 09.03.1999, ex IR 752)

Paratype 1: INBio 3542626, male (same data
as holotype)

Paratype 2: ZMB 103882, female (same data
as holotype)

Paratype 3: ZMB 103883, male (same data as
holotype)

Dimensions:

Holotype: 7.8/7.0/7.4/6.5/5.9/4.6/6.4 mm

Paratype 1: 6.6/6.0/6.5/5.6/5.0/4.1/5.3 mm

Paratype 2: 8.1/6.8/7.3/6.5/6.1/4.6/6.7 mm

Paratype 3: 6.3/5.9/6.5/5.5/5.2/4.1/5.1 mm

Type Locality

SE-Costa Rica, Limón Province, SW of
Liverpool (about 24 km W of Puerto Limón)
along Río Peje, 09°55'46"М, 83°13'15"W, 135
m a.s.l.

Etymology

The subspecies is named after its origin, the
Río Peje.

Examined Material

Lee. |. RICHLING

Limon: SW Liverpool: Rio Peje and small
tributary, 09°56’35"N, 83°14’01"W, 110 т
a.s:l.: 12.03.1997: (IR 125); along Rio Peje,
bordering forest with palms, 09°55’46"N,
83°13'15"W, 135 m a.s.l.: 04.03.1998: (IR
440); 09.03.1999: (IR 752); 03.03.2000: (IR
1303); (IR 1305); (IR 1306); 13.03.2001: (IR
1550)

Description

Shell (Figs. 131, 132, 336D): Conical-globose,
rather solid, medium sized, shiny. Color: up-

per whorls light yellowish-horny-amber, be-
coming darker from apex down, especially in
the course of the last whorl changing to bright
orange. Asmall but very distinct opaque whit-
ish band directly below the suture, color of
whorl most intensive towards the band. Shell
surface shiny and smooth, only structured
with very fine growth lines. Embryonic shell
with about 1 whorl; 4°/, (374-472) subsequent
whorls very slightly convex; last whorl equally
rounded at the periphery; upper whorls
slightly more rapidly extending in size; whorls
rapidly descending, forming a high spire. Su-
ture slightly impressed. Aperture oblique and
remarkably curved backwards, last whorl
regularly descending and inserting exactly at
periphery. Outer lip always bright orange in
continuation of last whorl, thickened and very
narrowly reflexed; transition into columella
continuous with a little notch. Basal callus
very weakly developed and nearly completely
smooth or very little granulated.

Internal Shell Structures: (Fig. 133)

Teleoconch Surface Structure (Fig. 134): Simi-
lar to Helicina b. beatrix, but the zone of ob-
lique diverging grooves is more pronounced.

Embryonic Shell (Fig. 135): The structure is
similar to that of Helicina beatrix confusa.
Figure 135B shows a common phenomenon
also seen in other species: a few spiral lines
of pits become indistinguishable.

La SS
/ A
№
а

Lar |

FIG. 133. Axial cleft and muscle attachments of
Helicina beatrix riopejensis n. subsp., INBio
3542625, 3542626; scale bar 2.5 mm.

304 RICHLING

FIG. 134. Teleoconch surface structure of Helicina beatrix riopejensis n. subsp. À. Embryonic shell to
2" whorl. В. 1* whorl, zone of transitional pattern and begin of transformation to next structure. С. 1*
whorl, pattern of oblique diverging grooves. D. 2% whorl, smooth surface with growth lines; scale bars

500 um (A), 100 um (B-D).

CLASSIFICATION ОЕ HELICINIDAE 305

FIG. 136. Operculum of Helicina beatrix riopejensis
n. subsp., INBio 3542625; scale bar 1 mm.

Diameter: 878 um (+ 31) (800-940) (n = 22)
(IR 1303, IR 1550).

Operculum (Fig. 136): Very slightly calcified,
calcareous plate covering only part of the
outer surface, thickened towards the col-
umellar side. Color orange to dark red, only
at columellar side and in the area of the
nucleus yellowish-transparent. Columellar
side slightly S-shaped, both ends acute,
upper end pointed, lower slightly rounded.

Animal (Fig. 337Н): As with other subspecies,
Helicina beatrix riopejensis n. subsp. lacks
any spotted pattern on the mantle. But the
dorsal and upper lateral sides of head and a
median stripe on the posterior foot are black.
Only on the middle of the head there 1$ a
lighter area. The tentacles are black as well.
А greyish-blackish mantle pigmentation
gives the semitransparent shell a greenish-
brownish appearance, making the white
band even more prominent.

Radula: See Helicina beatrix beatrix.

Female Reproductive System (Figs. 137,
138): The structures are similar to the nomi-
nal subspecies, except for the bursa
copulatrix, which is more regularly and
deeply lobed; the receptaculum seminis ap-
pears consistently smaller.

Morphometry and Sexual Dimorphism

FIG. 135. Embryonic shell of Helicina beatrix
riopejensis п. subsp.; scale bar 100 рт. See Helicina beatrix beatrix.

306 RICHLING

FIG. 137. Female reproductive system of Helicina beatrix
riopejensis п. subsp., apical complex in natural position, dorsal
and ventral view, IR 752; scale bars 1 mm (left), 0.5 mm (right).

Habitat coniaceae may have resulted from the much
larger surface of the leaves and the easier
At the type locality, this subspecies 1$ rela- search. When aestivating, the specimens

tively abundant. The undergrowth of the were found mainly on the underside of the
vegetation of the banks of the creek 1$ leaves close to the middle rib. On palms,

mainly composed of Heliconiaceae, different individuals were observed up to about 5-6
palm species, and Araceae. During wet m above the ground. Along the Río Peje, the
weather, Helicina beatrix riopejensis п. subspecies lives sympatrically with H.
subsp. was found crawling nearly every- funcki, but it was not discovered at several
where on the leaves with no obvious prefer- other localities in the area along Río
ence for any particular plants. A higher Victoria, Río Blanco, Río René or Río Quito,
abundance on leaves of palms and Heli- where H. funcki also occurs.
CS. = O) La (QA
PA Y TX ee NIE
AG - soir а м
Y \)7 = wee 3 2 44 bs y SA sy =
( ee ( en & a r if { / ]
д er al INS
En JA EA A is ) = y =
( = en > h N ) (SS
qe (35 ae =a
LA ee PS ESA
N VA Nazis) |
NS a ре © ae \ X=
\ ae NS Se wen
Tr ec Y EZ wa
5 — AY YD) a —=—_
A о

FIG. 138. Variability of the female reproductive system of Не/ста beatrix riopejensis п. subsp., IR 752;
scale bar 2.5 mm.

CLASSIFICATION ОЕ HELICINIDAE

Distribution (Fig. 139)

Up until now, the subspecies has only been
found along the upper part of the Rio Peje and
a few small tributaries around the type locality.
The site belongs to the hilly Caribbean low-
lands close to the northeastern foothills of the
Cordillera de Talamanca.

Discussion

Few specimens exhibit a dark red spot at the
apex. The tinge of orange towards the aper-
ture may be paler or even bright red. Other-
wise, the color is very constant within the
population investigated.

Helicina beatrix riopejensis п. subsp. differs
from the nominal species and other subspe-
cies in the color of the outer lip, which in the
other subspecies is consistently whitish, inde-
pendently of the varying color of the whorls.
Furthermore, in H. beatrix beatrix, H. b. con-
fusa, and H. b. nicaraguae the color of the
whorls becomes lighter towards the aperture
(normally whitish at least in the umbilical

307

area), whereas in H. b. riopejensis п. subsp. it
turns darker, even in the umbilical area. In
general, the whitish band 1$ more slender in H.
b. riopejensis n. subsp.

With respect to the orange outer lip, the sub-
species closely resembles H. gemma, but the
latter consistently lacks the distinct whitish
band under the suture. lts suture seems
somewhat more strongly impressed, and the
whorls appear to be more convex. Whereas
H. beatrix beatrix 1$ very clearly distinguished
from H. gemma, H. beatrix riopejensis n.
subsp. seems to represent a somewhat inter-
mediate form showing several similarities to
both species. Besides the aspects of the shell
color, the length of the axial cleft equals the
conditions of H. gemma and deviates from the
other subspecies of H. beatrix. On the other
hand, H. beatrix riopejensis n. subsp. com-
pletely lacks the spotted mantle pigmentation
of H. gemma and, with respect to the mor-
phometry, it would clearly represent the popu-
lation with the largest specimens. According to
all investigated populations of H. gemma, the
species is much more constant in size than

3500 -
3000 -

4000 m
3500 m a

2500 -
2000 -
1500 -

3000 m
2500 m
2000 m
1000 - 1500 m

500 - 1000 m
100 - 500 m
0-100m

e coll. IR
| e coll. INBio
| © others

FIGS 139.

Records of Helicina beatrix riopejensis п. subsp. in Costa Rica.

308 RICHLING

the other species. At the same locality at the
Rio Ре, it reaches its largest sizes, but in
Tortuguero, where H. gemma occurs sympatri-
cally, the population has a smaller shell size
than H. beatrix riopejensis n. subsp., making a
comparison with the conditions of other species
contradictory. With the present state of knowl-
edge, “riopejensis” is tentatively referred to H.
beatrix, but further data may reveal closer affini-
ties to H. gemma. Nevertheless, the differences
between “riopejensis” and both species justify
a recognition at subspecific level.

Helicina (“Gemma”) talamancensis
(Richling, 2001)

Helicina oweniana — Monge-Najera, 1997: 113:
Costa Rica [in part] [non L. Pfeiffer, 1849]
Helicina beatrix - Monge-Najera, 1997: 113:

Costa Rica [in part] [поп Angas, 1879]
Oligyra talamancensis Richling, 2001: 3-5
(text figure)

Original Description
See “Description”.
Type Material

Holotype: INBio 3404978, female (leg. |.
Richling, 24.3.1997) (Fig. 140)

Paratype 1: INBio 1494509, female
(Puntarenas, 3 km NE de la Escuela de LI-
ano Bonito, 08°44’54"N, 83°02'04"W, 920
m a.s.l., leg. Socorro Avila, 24.03.1997)

Paratype 2: INBio 3389580 (same data as
paratype 1)

Paratype 3: ZMB 103368, male (same data as
holotype)

Paratype 4: ZMB 103385, probably female,
empty shell (from type locality, leg. 1.
Richling, 29.8.1999)

Paratypes 5-12: INBio 1494642: 7 adults, 1
juvenile (same data as paratype 1)

Paratypes 13-14: INBio 1487761: 2 juveniles
(Puntarenas, 3.5 km de la Escuela de Llano
Bonito Carretera a San Vito, 08°44'37"N,
83”02'04"W, 840 т a.s.l., leg. Socorro Avila,
24.03.1997)

Dimensions (height/greatest diameter):

Holotype: 9.2/9.2 mm

Paratype 1: 9.1/8.7 mm

Paratype 2: 9.2/8.8 mm

Paratype 3: 8.2/8.3 mm

Type Locality

SW-Costa Rica, Puntarenas Province, Fila
Costeña, north of Bajo Bonito (locally called
Llano Bonito), north of Río Claro, 8°44'41"М,
83°02’09"W, 980 m a.s.l., probably primary
rain forest bordered by secondary growth.

Type Material of Synonymous Taxa or Similar
Species

Helicina terryae Rehder, 1940
Helicina terryae Rehder, 1940: 350, fig. 16

Type Material: USNM 536026 (not USNM
539026 as given in Rehder, 1940): holotype
(Fig. 141)

Dimensions (given in original description,
height/greatest diameter):
Holotype: 8.2/9.8 mm

Type Locality: Panama, Chiriquí Province.

FIG. 140. Helicina talamancensis, holotype, INBio 3404978, height 9.2 mm; scale bar 2.5 mm.

CLASSIFICATION ОЕ HELICINIDAE 309

Examined Material

Lec. |. RICHLING

Puntarenas: S San Vito, forest opposite the
Wilson Botanical Garden, Las Cruces,
08°46’57"М, 82°57'40"W, 1,160 m a.s.l.,
29.08.1999: (IR 1018)
N Nelly, road from Ciudad Neily to San Vito,
open area with a few trees, 08°40’23"М,
82°56 44"W, 180 m а.5.1., N Neily,
23.03.1997: (IR 210)
Fila Costeña, north of Bajo Bonito (locally
called Llano Bonito), М of Rio Claro, rain for-
est, 08°44'41"М, 83°02’09"W, 980 m a.s.l.,
24.03.1997: (IR 222); 15.02.1999: (IR 580);
29.08.1999: (IR 1029), (IR 1661); (IR 1030);
06.03.2001: (IR 1487); (IR 1489)
Refugio Nacional de Fauna Silvestre Со,
rain forest, 08°39’26"М, 83*10'50"W, 100 т
a.s.l., 14.02.1999: (IR 567); 10.02.2000: (IR
1166)

INBio COLLECTION

San José: San Isidro, Area de Conservación la
Amistad, Parque Nacional Chirripó, Estación
Santa Elena, Finca del Gringo, 09*23'31"N,
83°35'42"W, 1,300 m a.s.l.: leg. А. М. Maroto,
29.09.1995: 1 ad. (INBio 3542536)

Puntarenas: Reserva Forestal Golfo Dulce:
Cerro de Oro, 08°33’46"N, 83*29'24"W,
150 m a.s.l.: leg. Eida Fletes, 30.10.1995:
1 ad. (INBio 1498769), 1 ad. (INBio
1498766); Cerro de Oro, Quebrada
Terranosa, 08°34’11"М, 83*30'15"W, 140 т
a.S./.: leg. Ronald Villalobos, 05.10.1995: 1
ad. (INBio 1485176); Rancho Quemado,
08*40'35"N, 83*34'33"W, 250 т a.s.l.: leg.
Zaidett Barrientos, 18.03.1994: 1 juv.
(INBio 1475394)

Playa Blanca, 08°38'18"N, 83*26'16"W, О m
a.s.l., leg. Gillermo Mena, 04.09.1995: 1 juv.
(INBio 1479918)

Fila Cal: 24 km de San Vito hacia Ciudad
Neilly, 08*41'36"N, 82°56’36"W, 780 т
а.$.1.: 14.01.1995: leg. Luis Angulo, 2 ads.,
1 juv. (INBio 1480714); leg. Angela Mora
Maroto: 2 ads. (INBio 1481246); leg.
Socorro Avila: 4 ads., 2 juvs. (INBio
1481353); leg. Marcos Moraga: 1 ad., 1 juv.
(INBio 1481564); leg. Alejandro Azofeifa: 1
ad. (INBio 1482605); leg. Francisco
Alvarado: 3 s.ads. (INBio 1495690);
29.08.1995: leg. Marianella Segura, 3 ads.,
2 s.ads. (INBio 3121201); 740 m a.s.l.: leg.
Ronald Villalobos: 4 juvs. (INBio 1481514);
24.5 km S en la carretera de San Vito hacia
Ciudad Neilly, 08°40'55"N, 82°56’23"W,
600 m a.s.l.: leg. Zaidett Barrientos,
21.11.1995: 4 ads., 1 juv. (INBio 1485120);
leg. A. Picado, 21.11.1995: 2 ads., 2 s.ads.,
1 juv. (INBio 3542530); leg. M. Segura,
21.11.1995: 2 ad., 1 s.ad. (INBio 3542545)
4.5 km NW de Ciudad Neily, Camino
Paralelo al Río Caño Seco, Colectado en
hojarasca en helechos, 08°40’50"N,
82°57'25"W, 180 т a.s.l.: leg. М. Chinchilla,
22.11.1995: 4 ads., 1 s.ad. (INBio 3542526)
Jardín Botánico Wilson, Sendero a Río
Jaba, 08°47'13"N, 82°58 04"W, 1,160 m
a.s.l., leg. Zaidett Barrientos, 10.03.1995: 1
ad. (INBio 1485093)

Estación Pittier. 09%01'32"N, 82°57'46"W,
1,660 m a.s.l.: leg. Angela Mora Maroto,
15.01.1995: 1 ad., 2 juvs. (INBio 1481397);
Sendero Pittier, 09%01'11"N, 82%57'54"W,
1,540 m a.s.l.: leg. malacological staff of
INBio, 06.11.1995: 4 ads., 1 juv. (INBio
1488141); Sendero Río Gemelo, 09°01’36"М,

FIG. 141. Helicina terryae, holotype, USNM 536026, height 8.2 тт; scale bar 2.5 mm (photograph:
R. Hershler).

310 RICHLING

82°57'26"W, 1,640 m a.s.l.: leg. Annia
Picado, 13.01.1995: 1 juv. (INBio 1481168)
Parque Nacional La Amistad, Coto Brus,
sendero a Cerro Pittier, 600 m NW de la
Estación, 09°01’44"N, 82°57’54"W, 1,750 m
a.s.l., leg. Marcos Moraga, 06.11.1995: 1 ad.
(INBio 1484619)

Parque Nacional La Amistad, Estación
Pittier. Sendero a Cerro Pittier. 09%02'05"N,
82°57’39"W, 1,800 m a.s.l.: leg. Luis
Angulo, 06.10.1995: 1 juv. (INBio
1485495); 09°01’43"N, 82°57'54"W, 1,750
т a.s.l.: leg. М. Moraga, 19.06.1996: 1 ad.
(INBio 3542538); Sendero а Altamira, 900
т NW de la estación, 09%01'52"N,
82°58 05"W, 1,760 т a.s.l.: leg. Evelio
Alfaro, 15.01.1995: 1 ad. (INBio 1480719),
1 ad. (INBio 1480725); leg. Angela Mora
Maroto: 2 ads. (INBio 1481219), 1 ad.
(INBio 1481236); 1 ad. (INBio 3542544);
Sendero а Rio Canasta, 09°01’51"М,
82°58 05"W, 1,740 m a.s.l.: leg. М. Moraga,
14.06.1996: 1 ad. (INBio 3542540)
Parque Nacional La Amistad, Estaciôn
Altamira, Sendero a Estación Biolley:
09°01’59"N, 83%00'39"W, 1,340 m a.s.l.: leg.
Marianella Segura, 13.10.1994: 1 ad. (INBio
1485516); 09%01'47"N, 83°01’07"W, 1,300 т
a.S./.: leg. Alexander Alvarado Mendez,
10.09.2001: 1 ad. (INBio 3394313)

Parque Nacional La Amistad, Cerro Biolley,
09°02'25"N, 83*00'39"W, 1766 т a.s.l., leg.
Roberto Delgado, 17.06.1994: 1 ad. (INBio
1467066)

Parque Nacional La Amistad, Cabagra,
Puesto Altamira, Sendero a Cerro Biolley,
09°02'12"N, 83°00'39"W, 1,600 m a.s.l.,
13.06.2001: 1 s.ad. (INBio 3318186); 3 ads.
(INBio 3318194) (all leg. Alexander Alvarado
Mendez)

Puntarenas, Parque Nacional La Amistad,
Pittier, Puesto Altamira, sendero Casa Coca,

FIG. 142. Axial cleft and muscle attachments of
Helicina talamancensis, IR 1030; scale bar 5 mm.

09°02'25"N, 82°59'24"W, 1,800 m a.s.l., leg.
Alexander Alvarado Mendez, 12.05.2001: 1
ad., 1 s.ad. (INBio 3317088)

OTHER SOURCES

COSTA RICA

San José: determination uncertain? 4.3 mi
SW of San Isidro del General on Road to

Dominical [about 09°20’М, 83°44’W],
01.08.1971: 2 ads. (UF 69848)
Puntarenas: Rincón [about 08°42’30"М,

83°29'30"W], В. Casebeer, 28.06.1963: 1
ad. (UF 243510)

Etymology

The species is named after the southern
central mountain chain in Costa Rica, the Cor-
dillera de Talamanca, which forms the greatest
remaining undisturbed area of primary forest
in the country.

Description

Shell (Figs. 140, 336E): conical-globose, rather
solid, medium sized, shiny. Color: yellowish to

FIG. 143. Teleoconch surface structure of Helicina
talamancensis, 2" whorl; scale bar 100 um.

CLASSIFICATION OF HELICINIDAE otf

whitish-opaque (holotype); in some speci-
mens the last whorl yellowish-white and the
upper whorls with a more or less strong ten-
dency to a рае orangish-red color.
Periostracum very thin, shiny and smooth,
except very fine growth lines. Embryonic shell
with about 1 whorl; 4'/,-4°/, subsequent
whorls slightly convex; last whorl equally
rounded at periphery; upper whorls more rap-
idly extending in size, so that shell (especially
the female’s) appears somewhat rounded
and less pointed in apical part. Suture slightly
impressed. Aperture oblique and in its middle
part remarkably curved backwards. Outer lip
always whitish, thickened, very narrowly re-
flexed, appearing somewhat rounded at
edge; transition into columella continuous,
with a slight notch. Basal callus weakly de-
veloped and nearly completely smooth or
very little granulated.

Internal Shell Structures: (Fig. 142)

Teleoconch Surface Structure (Fig. 143): The
transitional pattern covers only about % of a
whorl; the structure is weakly developed.
The smooth zone with just the fine growth
lines follows directly.

uba
y be h y , + a À ©
ee es si

FIG. 144. Embryonic shell of Helicina
talamancensis; scale bar 100 um.

Embryonic Shell (Fig. 144): In comparison
with Helicina beatrix, the pitted pattern 1$
even less prominent in H. talamancensis.
The embryonic shell appears nearly smooth.
Diameter: 933 um (+ 40) (840-1,000) (n =
18) (IR 222, IR 1028, IR 1030).

Operculum (Fig. 145): Very slightly calcified,
calcareous plate covering only part of the
outer surface. Color horny-amber, only near
the columella whitish, but still somewhat
transparent. Columellar side S-shaped, both
ends acute, upper end pointed.

Animal (Fig. 338B): The appearance of living
Helicina talamancensis is striking: the body
is whitish-yellow throughout, the mantle pig-
mentation is also whitish; only the tentacles
are deep black. This characteristic color 1$
present in all live and preserved specimens
studied.

Radula (Fig. 146): Cutting edge of the centrals
smooth or crenulated. Comb-lateral with 8-9
cusps, cusps on marginals rather rapidly in-
creasing in number, but with a similar effect
as in Helicina beatrix beatrix. Radula with
about 60-75 rows of teeth.

Female Reproductive System (Figs. 147,
148): Compared to Helicina beatrix the as-
cending limb of the V-organ is elongated, the
receptaculum 1$ drop-shaped. The bursa
copulatrix 1$ very irregularly lobed, the elon-
gated provaginal sac shows a simple out-
line. Its stalk is short and stout. The pallial
oviduct is marked by a longitudinal furrow
and various transversal constrictions.

FIG. 145. Operculum of Helicina talamancensis,
holotype, INBio 3404978; scale bar 2 mm.

312 RICHLING

FIG. 146. Radula of Helicina talamancensis. A.
Centrals. В. Comb-lateral. С. Marginals; scale
bar 50 pm.

( с
I
Sa a N
re, 7 /
| NA \ | y
|
\
à
es à
я pao \
ЕО
y
А
+

FIG. 147. Female reproductive system of Helicina
talamancensis, IR 1030; scale bar 1 mm.

Morphometry and Sexual Dimorphism (Table
9, Figs. 149-155)

Helicina talamancensis could not be found in
high numbers, the only specimens studied
anatomically are those | collected at Bajo Bo-
nito. Populations included from the collection
of INBio with sufficient individuals (Fila de Cal,
Neily, Amistad, Bajo Bonito) that could not be
analyzed for sex were separated as in H.
beatrix to avoid artificial high deviations of
measurements with mixed sexes.

q 7 | > Wer
Be EEE
| SÓ) ve Or RE |
Hs CAN ee
LUZ —A Sy, we;
(ES / no / © )
vi { lex —% АС
PAU cou fre
“ef 2 a $ у
+) es: LIES
A e 7 <
fo | у А A
Cy =i
= а
= À > | а
И — une
SEEN LO | Ne

FIG. 148. Variability of the female reproductive
system of Helicina talamancensis, IR 1030;
scale bar 2.5 mm.

CLASSIFICATION OF HELICINIDAE 313

TABLE 9. Measurements of different populations of Helicina talamancensis given as mean value with
standard deviation, minimum and maximum value (min, max), and number of specimens; sex of
individuals from Fila de Cal, Neily, Amistad and Bajo Bonito INBio not determined anatomically (see
text) (min./max. diam. = minor/major diameter, col. axis = columellar axis); linear measurements [mm],
weight [g], volume [ml].

“Bajo Bonito” (altitude 980 m)
lots IR 1018, IR 1029, IR 1030, IR 1487

Mean
Sex value Deviation Min Max Number
Height f 9.03 0.33 8.48 9.61 8
Height m 7.81 0.187 750 6.22 8
Maj. diam. f 8.54 0.27 8.02 8.92 8
Maj. diam. m 7.46 0.13° 7.16 . 7.72 8
Min. diam. f 7.95 0.20 7.60 8.31 8
Min. diam. т 6.97 0.11 6.75 7.20 8
Outer lip f 5.61 0.15 -5.46 5.91 8
Outer lip m 5.12 0.17 480 5.41 8
Last whorl f 7:05 0.15. 6.58 747 8
Last whorl т 6.17 0.18 5.88 6.46 8
Col. axis f 7.34 0.26 6.77 7.87 8
Col. axis m 6.29 0.19 6.03 6.68 7
Weight Е 0.087 0.010 0.056 0.102 8
Weight т 0.066 0.007 0.055 0.085 8
Volume Е 0199 0.017 0.172 0.224 8
Volume т 0.130 0.008 0.115 0.145 8
“Fila de Cal” (altitude 600-780 т)
lots INBio 1480714, 1481246, 1481353,
1481564, 1482605, 1485120, 3121201, “Neily” (altitude 180 т)
3542530, 3542545 lot INBio 3542526
Mean Mean
Sex value Deviation Min Max Number value Deviation Min Max Number
Height f 1.13 0.17 135 198 10 TES 0.33 7.42 8.08 2
Height m 6.80 O21 5:82 HAN A 6.43 0.04 6.38 6.47 2
Maj. diam. f 7.14 0.14 6.95 7.42 10 7.42 0.097 77.332 7.50 2
Maj. dam. m 6.69 OMS 645 695 1 6.60 0.06 6:54 6.65 2
Min. diam. f 6.75 011 6.55 6.98 10 6.90 0:17 673 1.06 2
Min. diam. т 6.23 O42 6:08 6.50 11 5.95 0.107 585 16:04; 2
Outer lip f 4.71 0.16 441 487 10 4.78 0.02 4.75 4.80 2
Outer lip т 4.46 0.08 4.26 462 11 4.38 011. 427 2.48 2
Last whorl f 6.01 0.16 570 6:38 10 6.02 0:15 5897 61 2
Last whorl т 5.50 0.12 510 583 1 5.15 0:20 484595555135 2
Col. axis f 6.37 0.29; 15:95 7.31 10 6:23 0.23 6.00 6:45 2
Col. axis m 5:65 0.20» 5.29 726:27 11 SA 0106 511 5.23 2

(Continues)

314 RICHLING

(Continued)

“Amistad” (altitude 1340-1800 т)
lots INBio 1467066, 1480719, 1480725,
1481219, 1481236, 1481397, 1484619,

1485516, 1488141, 3317088, 3318194, “Bajo Bonito INBio” (altitude 920 т)
3542538, 3542544, 3542540 lots INBio 1494509, 1494642, 3389580
Mean Mean
Sex value Deviation Min Max Number value Deviation Min Max Number
Height f 8.50 0033 806 918 11 9.11 0.05 900 915 4
Height m 7.26 у 6887567 9 8.10 0.13 7.88 8.28 5
Maj. Чат. + 8.16 0.26 7.78 8.68 11 8.47 0.10 8.33 8.62 4
Maj. diam. т 7.20 0.21 638 7.50 9 7.67 017 725 1388 5
Min. diam. f 1.53 023 720 19511 7.88 0.09 7.77 8.02 4
Min. diam. m 6.58 0.15 5.94 6.74 9 7.08 012 (67 122 5
Outer lip f 537 0.19 5.08 5.81 11 5.62 0.107 542 519 4
Outer lip m 4.76 0.20 410 5.15 8 5.14 0.14 488 5.40 5
Last whorl f 6.55 0.19 645 692 14 1.12 0.07 7.02 7.20 4
Last whorl m 5.80 0.16 5.49 6.07 9 6.25 0.10 6.09 6.48 5
Col. axis f 6.94 0.29 6.36 7.63 11 7.29 0.13 712 155 4
Col. axis m 5.93 0.15 567 620 9 6:51 0.08 6.41 6.64 5
“Península de Osa” (altitude 140-150 т) “Chirripó” (altitude 1300 т)
lots INBio 1485176, 1498766, 1498769 lot INBio 3542536
Mean Mean

Sex value Deviation Min Max Number value Deviation Min Max Number

Height f 8.06 0.61 7.15 8.82 3 8.16 - - - 1
Maj. diam. f 8.04 O65. 707. 8:77 3 8.21 - - - 1
Min. diam. f 7.42 0.62 6.50 7.96 3 7.64 - - - 1
Outer lip f 5:26 0.41 465 5.66 3 5:27 - - - 1
Last whorl f 6.36 045 569 6.77 3 6.34 - - - 1
Col. axis f 6.61 0.56: 25.77. 7.15 E 6.66 - - - 1
Morphometry: The populations show remark- tion is smaller, but the weight is significantly
able differences in size with the individuals higher. It confirms the impression of more
from Fila de Cal and Neily being smallest. For solid shells in H. talamancensis.

each characteristic, the differences between
the populations are similar, implying that the Sexual Dimorphism: Аз in H. beatrix the sexes

relations are about the same. Although from clearly diverge in all measurements, in most
different altitudes and from localities relatively cases even without an overlap of the ex-
close to each other, the specimens from Neily trema (Fig. 156). The females are much big-
and the Fila de Cal are of about the same ger than the males, in volume the males are
size, which suggests rather a relation to the only */, of the females. This divergence al-
sites than to altitude. The few specimens from lows the separation of individuals of un-
the lowlands of Península de Osa approach known sex (Fig. 157), as explained under H.
the shells from “Amistad” at much higher al- beatrix beatrix. Although containing only
titudes more closely in size than those from three specimens, the lot from Península de
Neily or Fila de Cal. Osa probably consists of two females and
In comparison with Helicina beatrix beatrix, one male, because the average is shifted to

the shell volume of the Bajo Bonito popula- the higher value.

CLASSIFICATION OF HELICINIDAE 315

Bajo Bonito (n=8/8) 9
—— o
Fila de Cal (n=10/11)
—
SSS
Neily (n=2/2)
|
+
Amistad (n=11/9)
nn
SEN
Bajo Bonito INBio (n=4/5) +
po
Península de Osa (n=3)
EJ = SE
Chirripó
E=/ T
À | il 1 1 L L 1 1:
0 5 6 7 8 9 [mm] 10

FIG. 149. Shell height of different populations of
Helicina talamancensis in Costa Rica according
to Table 9; on each line: mean value, standard
deviation, absolute range; number of individuals
given as “п = females/males or total”; upper line:
females, lower line: males if separate; in
between and shaded: average of both for
comparison with populations of unknown sex;
sex of individuals from Fila de Cal, Neily,
Amistad, and Bajo Bonito INBio not determined
anatomically (see text).

Contrary to most of the populations of H.
funcki and H. tenuis, the smaller males also
weigh less than the females.

Habitat

The species is arboreal, at the type locality
specimens were found aestivating on the

Bajo Bonito (n=8/8)
ETES g
—— oO
Fila de Cal (n=10/11)
Neily (n=2/2
У ( ) =.
>
Amistad (n=11/9)
—
Bajo Bonito INBio (n=4/5)
SE +
Península de Osa (n=3)
Bf
Chirripó
Be
— | 1 1 | L 1 1 1
0 5 6 7 8 9 [mm] 10

FIG. 150. Minor diameter of shell of different
populations of Helicina talamancensis in Costa
Rica according to Table 9; for explanations see
Fig. 149.

Bajo Bonito (n=8/8
) ( ) 9
— O
Fila de Cal (n=10/11)
Neily (n=2/2) 4
Amistad (n=11/9)
Sl
Bajo Bonito INBio (n=4/5)
Peninsula de Osa (n=3)
== PP
Chirripó
=f f AA
1H 1 1 1 - 1 1 1
0 3 4 6 7 [mm] 8

FIG. 151. Expansion of outer lip of different
populations of Helicina talamancensis in Costa
Rica according to Table 9; for explanations see
Fig. 149.

lower side of leaves in the undergrowth,
mainly on Heliconiaceae and different species
of palms. They were more abundant during
the rainy season in August than they were in
February and March during the dry period. In
rainy weather, individuals were also observed
crawling on the upper side of leaves at night
near the Wilson Botanical Garden. At several
localities, Helicina talamancensis occurs sym-
patrically with H. pitalensis.

Distribution (Fig. 158)

Helicina talamancensis is only known from
the Pacific slopes in the southern parts of

Bajo Bonito (n=8/8) 9
== 0)
Fila de Cal (n=10/11)
Neily (n=2/2
ily (n=2/2) =
—
Amistad (n=11/9)
—
Bajo Bonito INBio (n=4/5) р
] —
Peninsula de Osa (n=3)
es |
Chirripó
Eee | р
1 L 1 4 _L 1 1
0 fe 3 4 5 6 7 [mm] 8

FIG. 152. Height of last whorl of different
populations of Helicina talamancensis in Costa
Rica according to Table 9; for explanations see
Fig. 149.

316 RICHLING

Bajo Bonito (n=8/7) 9
a
Fila de Cal (n=10/11)
| Мейу (n=2/2)
ily ie)
+
| Amistad (n=11/9)
| —
| Bajo Bonito INBio (n=4/5)
A SE
| Peninsula de Osa (n=3)
E —
| Chirripo
ЕЁ RSS
| 1H 1 1 1 1 1 1 1
0 2 3 4 5 6 7 [mm] 8

FIG. 153. Height of columellar axis of different
populations of Helicina talamancensis in Costa
Rica according to Table 9; for explanations see
Fig. 149.

Costa Rica. It occurs in the lowland rain for-
est on the Peninsula de Osa and near
Golfito close to the coast at elevations of ap-
proximately 100-250 т. In the steep moun-
tains of Fila Cruces and Fila de Cal and on
the slopes of the southern Cordillera de
Talamanca, H. talamancensis is found on
elevations of up to 1,800 m in the cloud for-
est area. Although corresponding in its distri-
bution to the southern records for H.
pitalensis, there is no indication and no his-
torical evidence for a more northerly occur-
rence of this species than the area of San
Isidro. With respect to records of Helicinidae,
the adjacent area of Chiriqui Province,
Panama, is virtually unexplored.

Discussion

The material of Helicina talamancensis con-
tains some specimens that show a completely
pale orange color, whereas yellowish speci-
mens are much more common.

| Bajo Bonito (n=8/8)

— a о
| En =
0

1 1 1 SL: 1
0.04 008 0.12 01 02 [mi] 0.28

FIG. 154. Shell volume of Helicina talamancensis
in Costa Rica according to Table 9; for
explanations see Fig. 149.

Bajo Bonito (n=8/8) 9
LLL @6U
1 1 1

| 1 1 1
0 0.02 0.04 0.06 0.08 0.1 la] 0.14

FIG. 155. Shell weight of Helicina talamancensis
in Costa Rica according to Table 9; for
explanations see Fig. 149.

The species most closely resembles H.
beatrix beatrix, but differs in the form of the
peristome. In H. beatrix beatrix, the outer lip is
less reflexed and thinner, and the aperture is
more strongly curved backwards so that the
upper part appears somewhat depressed in
frontal view. Furthermore, H. talamancensis
lacks the characteristic white band directly
under the suture. The color of the soft body of
H. talamancensis is unique among Costa
Rican Helicinidae.

Helicina terryae is of a nearly similar color
and size. Unfortunately, this species was de-
scribed from the holotype only, with the vague
locality of Chiriqui Province, Panama, and
there are no other similar specimens in the
USNM collection (pers. comm. Dr. R. Hershler,
USMN). Examination of photographs of the
type (USNM 536026, not USNM 539026, as
given in Rehder, 1940; pers. comm. Dr. R.
Hershler) revealed that H. terryae displays a
different outline of the last whorl in having the
curvature of the periphery more towards the
base. The outer lip is less reflexed, and the
spire is lower. Furthermore, the surface of the
shell appears to be similar to that of H. funcki,
for example, with “irregular, oblique, and
subspiral grooves”, rather than to the smooth
and shiny one of H. talamancensis.

The material of H. beatrix and H. oweniana
sensu Monge-Nájera (1997) was checked in
the INBio collection and was found to partially
belong to H. talamancensis. The differences to
H. beatrix are given above. Helicina oweniana
differs most obviously in the distinct orange
color of the outer lip and the less impressed
suture.

Rehder (1940) mentioned H. tenuis as the
most closely resembling species of H. terryae.
From H. talamancensis, H. tenuis can easily
be distinguished by its shell surface structure,
rectangularly expanded outer lip, occasionally
developed bands and color of shell and soft
body.

CLASSIFICATION OF HELICINIDAE

9 T T De
female >

mn male +
diam. |
[mm] e

o o |

8 & o
8
©
1.5 +

о

6 6.5 7 7.5 8 8.5 9 height [mm] 1

FIG. 156. Range of measurements in females and males of Helicina
talamancensis exemplary for height and minor diameter in the population
from Bajo Bonito.

min.
diam. |
[mm]
8 |- A Ñ -
©
© o
7.5 o 4
o
- females

6.5.7

5.5

6 6.5 7 7.5 8 8.5 9 height [mm] 10

FIG. 157. Plot of measurements for height and minor diameter for individuals
of Helicina talamancensis of unknown sex, exemplary for the population of
Amistad and the separation proposed.

317

318 RICHLING

3500 - 4000 m

3000 - 3500 m

2500 - 3000 т
2000 - 2500 m
1500 - 2000 m
1000 - 1500 m

500 - 1000 m
| 100 - 500 m
| 0- 100 т

e coll. IR
# coll. INBio |
© others

FIG. 158. Records of Helicina talamancensis in Costa Rica.

Helicina (“Gemma”) gemma
Preston, 1903

Helicina oweniana var. anozona — Biolley,
1897: 5: Costa Rica: Tuis, 600 m [about
09%51'N, 83°35’W, Cartago Province] and las
Delicias (Santa Clara), 400 т [10%57'37"N,
8502 W, 40 m a.s.|., Alajuela Province] [поп
von Martens, 1876]

Helicina oweniana var. coccinostoma — von
Martens, 1900: 605-606: E-Costa Rica: Las
Delicias, near Santa Clara, 400 m
[10%57'37"N, 85°02'W, 40 m а.$.1., Alajuela
Province] (Biolley) [non Morelet, 1849]

Helicina oweniana var. anozona — von Martens,
1900: 605-606: E-Costa Rica: Las Delicias,
near Santa Clara, 400 m [10%57'37"N,
85°02’;W, 40 m a.s.l., Alajuela Province]
(Biolley), Тиз, 600 m [about 09%51'N,
83°35’W, Cartago Province] (Pittier, Biolley)
[non von Martens, 1876]

Helicina gemma Preston, 1903: 4 (with text
figure)

Alcadia (Leialcadia) gemma — Wagner, 1908:
83, pl. 14, figs. 17-18

Oligyra (Succincta) gemma - Baker, 1922а: 45

Helicina ометапа — Monge-Nájera, 1997: 113:
Costa Rica [in part] [non L. Pfeiffer, 1849]
Helicina beatrix — Monge-Nájera, 1997: 113:

Costa Rica [in part] [non Angas, 1879]

Original Description

“Shell conical, elevated, bright yellow, apical
whorls crimson, last whorl tinged with orange-
scarlet for some distance from the mouth, the
outer lip being also of a vivid orange-scarlet
colour. Whorls 5, convex, very finely striated
with lines of growth. Peristome expanded and
slightly reflexed. Aperture rather high and nar-
row. Operculum reddish-brown, normal.
Diam. maj. 6, alt. 7 millim. Aperture (inside
measurement) diam. 2.5, alt. 3 millim.

Hab. — Costa Rica.

A very beautiful and striking shell, whose near-
est ally appears to be H. oweniana, Pfr., from
Mexico; from this, however, it differs in being
more globular, in the greater convexity of the
whorls and in having one less, in the narrower
aperture, and in the color of the outer lip (other-
wise similar in both species) extending further up
the body whorl than it does in H. oweniana.”

CLASSIFICATION OF HELICINIDAE 319

FIG. 159. Helicina gemma, lectotype, BMNH 1903.5.4.2, height 7.0 mm; scale bar 2.5 mm.

Type Material

Lectotype BMNH 1903.5.4.2 “San Carlos,
purchased from Мг. Н.В. Preston”; 1
paralectotype ZMB 53814 “San Carlos, ex
Preston”, 1 paralectotype ZMB 59238 (ex
Preston); 1 paralectotype ANSP 098181
“Costa Rica, San Carlos River, purchased from
Preston as cotype” (Robertson et al., 1986)

According to Dance (1986), most Preston
types are in the BMNH collection. Following the
advice for type selection of the International
Code for Zoological Nomenclature, the speci-
men ВММН 1903.5.4.2 is here selected as
lectotype of Helicina gemma (Fig. 159). Fur-
thermore, it was labeled by Preston as “type”
and is much better preserved than the ZMB
specimens suffering from Byne's disease. The
specimen still possesses its operculum. | did
not study the ANSP specimen.

Dimensions:
Lectotype BMNH 1903.5.4.2:

7.0/6.1/6.5/5.7/3.9/5.4/5.6 mm
Paralectotypes:

ZMB 53814: 6.0/5.4/5.8/5.0/3.5/4.5/4.7 mm

ZMB 59238: 6.7/6.0/6.2/5.5/3.7/5.0/5.4 mm

Type Locality

“Costa Rica”, by type selection restricted to
San Carlos [on one hand San Carlos is the old
name for Ciudad Quesada, about 10°20'N,
84°26’W in Alajuela Province; on the other
hand and following the data from the ANSP
specimen, a river in northern Costa Rica 1$
called Río San Carlos, bearing the name from
the confluence of Río Arenal and Río Peñas
Blancas near Boca de Arenal, about 10°33’N,

84°29'W, until it becomes a tributary of the Rio
San Juan at Boca San Carlos at the Nicara-
guan border, about 10°46’30"N, 84°12’30"W
in Alajuela Province, thus referring to a more
northern area. However, both possible loca-
tions share the trait that they are situated on
the Caribbean plain in the eastern part of
Alajuela Province].

Examined Material

Lec. |. RICHLING

Guanacaste: N of Nuevo Arenal: area of primary

rain forest, 10°33’32"N, 84*51'40"W, 800 т
а.$.!.: 05.03.1999: (IR 740); “Las Pavas” (pri-
vate reserve in preparation), secondary rain
forest, about 10°33’30"N, 84*51'53"W, 800 m
a.s.l., to 10°33’26"М, 84°51'57"W, 760 m a.s.l.:
05.03.1999: (IR 741); 17.08.1999: (IR 947);
(IR 948): 24.02.2000: (IR 1275); (IR. 1277);
27.02.2001: (IR 1460); (IR 1464); 01.03.2001:
(IR 1462); 03.03.2001: (IR 1463)
Parque Nacional Guanacaste, Volcan Ca-
cao, at southern slope, Estacion Cacao, W-
Sendero near station, forest, 10°55’35"М,
85 28 06 W, 1,110 miazsil., 18:03-1999; (IR
786); 09.03.2000: (IR 1333); (IR 1335)

Alajuela: Near Volcan Arenal, trail along vol-
cano т rainforest: about 10°29’07"N,
84°42'55"W, 720 m a.s.l.: 24.02.1998: (IR
387); 03.03.1999: (IR 734); 01.08.1999: (IR
885); 25.02.2000: (IR 1284); about
10°33-23-Ne (84-5101 W, 800 т ais:
05.03.1997: (IR 77); (IR 76)

Limon: Siquirres, along footpath stream up
Rio Siquirres and along a southern tributary,
100537 № 283-308 2W + 100 м аъ.
11.03.2001: (В 1536): 19.05.2007: “IR
1618); (IR 1635); (IR 1650; (IR 1652)

320

Zona Protectora Tortuguero, near Tortuguero,
N of village, about 10°34’N, 83°31 W, 10 т
a.s.l., 16.03.2001: (IR 1621); (IR 1654)

INBio COLLECTION

Guanacaste: Parque Nacional Guanacaste,
Estación Cacao: 10%55'29"N, 85°28'17"W,
1,100 m a.s.., leg. Dunia Garcia,
01.12.1995: 1 ad. (INBio 1484977);
10%56'05"N, 85°28'14"W, 1,100 m a.s.l., leg.
Dunia Garcia, 13.12.1995: 1 ad., 2 s.ads.
(INBio 1488058); 10%55'43"N, 85°28'20"W,
1,000 m a.s.l., leg. malacological staff of
INBio, 09.01.1995: 5 ads., 2 s.ads. (INBio
1539438); Sendero Los Naranjos,
10%55'38"N, 85”28'30"W, 1,100 т а.$.1., leg.
Dunia Garcia, 13.09.1995: 9 ads., 3 s.ads.
(INBio 1487886); Sendero Los Naranjos,
10%55'38"N, 85°28'30"W, 1,020 т a.s.l., leg.
malacological staff of INBio, 14.09.1995: 8
ads. (INBio 1539463)

Parque Nacional Guanacaste, La Cruz, 9 km
S de Santa Cecilia, Estación РИ:
10%59'25"N, 85°25’38"W, 700 m a.s.l.: leg.
Petrona Rios, 22.08.1994: 1 juv. (INBio
1480267); 1 ad. (INBio 1480284); 10°59'33"N,
85°25'46"W, 700 т a.s.l.: leg. malacological
staff of INBio, 10.09.1993: 1 ad. (INBio
1463737); Lado $ del Río Orosi, 10°59’25"N,
85°25'38"W, 700 т а.$.1.: leg. Calixto Moraga,
23.08.1994: 1 ad., 2 s.ads. (INBio 1480341);
leg. Marcos Moraga, 04.04.1995: 1 ad. (INBio
1484672); Sendero Mena, 400 m W de la
Estación РИ, 10°59’25"М, 85*25'51"W, 700
т a.s.l.: leg. Calixto Moraga, 09.01.1994: 1
ad. (INBio 1480045); Sendero a la Fila de
Orosilito, 10*59'24"N, 85*25'38"W, 700 т
a.s.l.: leg. Petrona Rios, 09.01.1994: 1 ad.
(INBio 1480270)

Parque Nacional Guanacaste, Sector Orosi
(antes: Maritza ); sendero Casa Fram,
10%57'40"N, 85°29'45"W, 600 т a.s.l., leg.
Zaidett Barrientos, 15.07.1996: 7 ads., 1
s.ad. (INBio 1487835)

Parque Nacional Rincón de la Vieja, Sector
Santa Мапа, 10°45'58"М, 85°18'19"W, 800
m a.s.l., leg. Dunia Garcia, 07.11.1996: 1 ad.
(INBio 1488039)

Alajuela: Reserva Biológica San Ramón,
10°13’30"N, 84°35'17"W, 800 т a.s.l., leg.
Gerardo Carballo, 14.12.1994: 1 ad. (INBio
1485501)

Sector Colonia Palmareña, San Ramón,
10°13’56"N, 84°33'12"W, 760 m a.s.l., leg.
Gerardo Carballo, 04.11.1995: 1 ad. (INBio
1484803)

Limón: Estación Cedrales: 800 т W de la
Estación Cedrales, 10°31'39"N, 83%43'33"W,

RICHLING

10 m a.s.!.: leg. Elias Rojas, 22.11.1996: 1 ad.
(INBio 1498586); Finca Leiva, 1 km W de la
estación Cedrales, 10°31’35"М, 83°43’33"W,
10 m a.s./.: leg. Elias Rojas, 17.10.1996: 1 juv.
(INBio 1501467); 3 ads. (INBio 3398104);
Finca Montaña Grande, 10°31'39"N,
83°43'33"W, 10 m a.s.l.: 600 m N de la
estación Cedrales, leg. Elias Rojas,
18.10.1996: 3 ads., 1 s.ad. (INBio 1501218);
500 т М de la estación Cedrales, leg. Elias
Rojas, 18.11.1996: 1 ad. (INBio 1498585)

Orillas del río Aguas Frías, 10°24’05"N,
83”36'00"W, 10 m a.s.l., leg. Elias Rojas,

29.11.1996: 20 ads., 15 s.ads,, 8 м5.

(INBio 1487942); 1 ad. (INBio 1488002)
Sector Guápiles, 10°11'51"N, 83°51’22"\М,
300 т a.s.l., leg. Alexander Alvarado Mendez,
08.03.2000: 1 ad. (INBio 3097951)

Refugio Nacional de Vida Silvestre Barra del
Colorado, Barra del Colorado, Estación
Sardinas: 10°38’52"N, 83°43’52"W, 50 т
a.s.l.: 10.02.1994: 1 ad., 1 s.ad. (INBio
1484009); 12.10.1994: 1 ad. (INBio 1484371);
16.10.1994: 1 ad. (INBio 1484012);
22.10.1994: 6 ads. (INBio 1485286) (all leg.
Flor Araya); 10%39'11"N, 83°44'21"W, 15 т
a.s./.: 13.01.1994: 2 ads., 1 s.ad. (INBio
1478019); 16.04.1994: 1 ad. (INBio 1477917)
(all leg. malacological staff of INBio)

Refugio Nacional de Vida Silvestre Barra del
Colorado, Pococí, Colorado, Sector Cerro
Cocori, 30 km N de Cariari, 10°35’39"N,
83*42'59"W. 160 m a.s.l.: leg. malacological
staff of INBio, 10.12.1993: 2 ads. (INBio
1465444); 2 ads. (INBio 1465444); 150 m
a.s.l.: leg. malacological staff of INBio,
04.10.1994: 3 ads. (INBio 1478057); 3 ads.
(INBio 1478057); 3 ads. (INBio 1478057)

Cartago: Parque Nacional Barbilla: Orilla de

río Dantas, cerca de la estación principal,
09°58’23"М, 83°27’03"W, 300 m a.s.l., leg:
Alexander Alvarado Mendez, 26.10.2000: 1
ad. (INBio 3316138); Sector de la Estaciön
de Barbilla, 0957'58"N, 83°27'41"W, 480 m
a.s.l., leg. Alexander Alvarado Mendez,
06.09.2000: 1 ad. (INBio 3100215)

Puntarenas: Parque Nacional Corcovado,

Estación Sirena, 08”28'52"N, 83*35'32"W, 5
m a.s.l., leg. Mario Chinchilla, 23.03.1995: 1
ad. (INBio 1485052)

OTHER SOURCES
COSTA RICA
Alajuela: Carriblanco [Cariblanco, about

10%17'N, 84°12’W], С.Н. Lancester (BMNH
1905.3.31.4)

Heredia: Rio Frio, Standard Fruit Co.,

10°20’N, 83°53’W, leg. Michael J. Corn,

CLASSIFICATION OF HELICINIDAE 321

05.12.1969: 1 ad. (UF 217595); Rio Frio Description
[about 10°20’М, 83°53’W], leg. Michael J.

Corn, 20.02.1970: 1 ad. (UF 217596); Shell (Figs. 160-162, 336F-H): Conical, thin
Costa Rica, without locality further specified: and fragile, medium to small sized, semi-
ex Sowerby 8 Fulton: 2 ads. (UF 243507: 2 transparent and shiny. Color: basic color
of 3 spec.) unicolored, more or less intensively yellow,
apical whorls sometimes crimson; last whorl

NICARAGUA: tinged with orange-brownish some distance
Zelaya Norte: Cerro Saslaya, Bosawas, leg. from aperture, sometimes also at the umbili-
Zamira Guevara M., 04.1999 (IR 3137) cal area. Surface textured with fine and

FIGS. 160-162. Helicina gemma. FIG. 160. Cacao, IR 1333, height 6.6 mm. FIG. 161. Las Pavas, IR
1460, height 7.3 mm. FIG. 162. Siquirres, IR 1536, height 7.0 mm; scale bar 2.5 mm.

322 RICHLING

FIG. 163. Axial cleft and muscle attachments of
Helicina gemma, IR 947; scale bar 2.5 mm.

regular growth lines (Fig. 164), causing the
glossy appearance. Embryonic shell with
about 1 whorl; 3°/,—4°/, (lectotype: 4) sub-
sequent whorls convex; last whorl very
evenly rounded at the periphery; whorls
equally extending in size, forming a very
regular, pointed spire. Suture moderately
impressed. Aperture oblique and curved
backwards, last whorl regularly descending
towards the aperture and inserting a little
below the periphery. Outer lip in continuation
of the whorl of a bright orange color, slightly

FIG. 164. Teleoconch surface structure of
Helicina gemma, 2"* whorl; scale bar 100 um.

thickened and very narrowly expanded and
reflexed. Transition to columella continuous
with a slight notch. Columella slightly
curved; transition to the body whorl without
any groove. Basal callus very weakly devel-
oped and slightly granulated.

Internal Shell Structures: (Fig. 163)

Teleoconch Surface Structure (Fig. 164): The

surface structure of Helicina gemma 1$ de-
scribed above representing the general
scheme: about Y whorl exhibits transitional

structure and subsequently oblique diverg-

ing grooves; the rest of the shell is smooth
with only fine growth lines.

Embryonic Shell: The embryonic shell of

Helicina gemma approaches the structure of
H. funcki with larger pits and similarly sized
interspacings. The pattern is relatively con-
stant within a population as well as at differ-
ent localities (investigated: Cacao, Las
Pavas, Tortuguero, Siquirres) (Fig. 165). The
diameter (range and mean value) increases
with the altitude of the locality.

Diameter: 925 рт (+ 23) (900-960) (п = 10)
(IR 786, IR 1333, Cerro Cacao); 845 pm (+
28) (760-900) (п = 24) (IR 1275, Las Pavas);
808 um (+ 25) (740-860) (n = 32) (IR 1635,
Siquirres); 800 um (BMNH 1903.5.4.2, lecto-
type).

Operculum (Fig. 166): Very slightly calcified,

calcareous plate covering only part of the
outer surface. Color yellowish to horny-am-
ber-reddish, only near the columella whitish
or transparent. Columellar side slightly regu-
lar S-shaped, upper end acute and pointed,
lower end continuously changing into outer
margin.

Animal (Figs. 338С-Е): The color of foot and

head is constant in all populations, whereas
the mantle pattern is subject to variation.
The sole is whitish yellow; the dorsal part
and upper side of the head region, including
tentacles and the dorsal portion of the foot,
are greyish to black. In all specimens from
the Cerro Cacao and near Volcán Arenal,
the mantle is unicolored and pale, whereas
in the populations from Las Pavas,
Tortuguero and Siquirres such forms are
very rare. Instead, the mantle pigmentation
displays a special pattern: a greyish-blackish
basic color mottled with whitish dots. In most

CLASSIFICATION OF HELICINIDAE 323

< a TR
PR < А р
ni wem...

rer, à
TIE

FIG. 165. Embryonic shell of Helicina gemma. A. Cerro Cacao, IR 786. B. Las Pavas, IR 1460. C.
Tortuguero, IR 1654. D. Siquirres, IR 1618; scale bar 100 рт.

324 RICHLING

FIG. 166. Operculum of Helicina gemma, IR 947:
scale bar 1 mm.

cases, these dots are so numerous that only
a network of thin dark lines is visible. The
mantle pigmentation is clearly visible
through the thin shell.

Radula (Figs. 167, 168): Sometimes B- and C-
central with 3-7 ог about 3 cusps respec-
tively, Figures 167A and 168 show
exemplary such variations. Comb-lateral
with (7—) 8-9 cusps, cusps on marginals rap-
idly increasing in number. Radula with about
62-85 rows of teeth.

Female Reproductive System (Figs. 169-171 E
The receptaculum seminis is quite large and
drop-shaped. The prominently developed
bursa copulatrix consists of a few relatively
large, simple and equal-sized lobes, some-
times further subdivided. They are only occa-
sionally smaller and more numerous. The
provaginal sac is simple, and its stout stalk is
shorter than in Helicina funcki. In comparison
to the apical complex, the pallial oviduct is
short. It is transversally constricted and often
exhibits an additionally longitudinal furrow.
Specimens from Las Pavas, Tortuguero and
Siquirres are similar; the single female dis-
sected from the Cerro Cacao has a smaller
bursa copulatrix (Fig. 170A).

Morphometry and Sexual Dimorphism (Table
10, Figs. 172-176)

The available material of Helicina gemma is
comparatively comprehensive, and four popu-
lations for anatomical investigations from dis-
tant sites were collected.

FIG. 167. Radula of Helicina gemma. A. Centrals.
B. Comb-lateral. C. Marginals; scale bar 50 um.

CLASSIFICATION OF HELICINIDAE 325

Whereas the three upper localities of the
collection IR and the collection INBio respec-
tively (Figs. 172-176) belong to the northwest-
ern Caribbean slopes, the others come from
the eastern plain. The type locality is situated
between these sites.

All populations included from the INBio col-
lection (Figs. 172-176, below thick line) that
could not be analyzed for their sex were sepa-
rated, as in H. beatrix, to avoid artificially high
deviations of measurements with mixed sexes.

Morphometry: The variations among the differ-
ent populations for the different measurements
are quite constant. The size differences corre-
spond to the origin of the specimens. “Cacao”
(IR and INBio), “Las Pavas”, “Volcan Arenal”
and “Orosi” are very similar to each other; only
the shell size of the population “Pitilla” is some-
what smaller. Except for Tortuguero which is
very close to the sea, Helicina gemma be-
comes bigger in the northeastern Caribbean
lowlands. More to the south, near southern
limit of Known distribution, the size declines
(Siquirres). The lectotype, higher than the av-
erage shell of the populations compared, has
an intermediate size, suggesting that it is a
female. As in some samples of H. funcki, the
corresponding mean values of the two
samples from “Cacao” treated separately sup-
port the reliability of results gained with small
samples sizes.

Following the suggestion that the shells be-
come larger in the lowlands, the average
minor diameter was plotted against the alti-
tude of the sites (Fig. 178). The values indi-
cate a slight decline of the size with

4 =
- :

FIG. 168. Radula of Helicina gemma, centrals;
scale bar 50 ит.

increasing altitude. The difference amounts
about 10% of the shell size of the population
with the largest individuals, which is only 5%
less than in H. funcki but which is yet found
up to 1,500 m.

Sexual Dimorphism: All measurements clearly

show a different range for both sexes, with
the females being bigger. Only in populations
with a high sample size (Siquirres, Las
Pavas) do the extrema overlap a little, this
being illustrated for the original set of data of
height and minor diameter for the populations
“Las Pavas” and “Siquirres” (Figs. 179, 180).
The volume of the males is only about */, of
that of the females. The differences displayed
for the populations are very constant for each
measurement. As explained in H. beatrix, the
well-developed sexual dimorphism allows a
separation of sets of mixed data. (illustrated
for Río Aguas Frías, Fig. 181)
The lectotype is assumed to be female, be-
cause it seems very unlikely that the type
(type lot) is extraordinarily big in a species of
low variation. Furthermore, the paralecto-
types are smaller.

Habitat

Helicina gemma is an arboreal species,
mainly climbing and aestivating on the lower

FIG. 169. Female reproductive system of
Helicina gemma, IR 1275; scale bar 1 mm.

326 RICHLING

A B -

FIG. 170. Variability of the female reproductive system of Helicina gemma, populations from A. Cerro
Cacao, IR 786. B. Las Pavas, IR 947, IR 1275; scale bar 2 mm.

and sometimes upper side of leaves. As ob-
served for H. beatrix, a special preference of
certain plant species could not be observed.
The predominant presence appears to de-
pend on the species composition of the under-
growth on whose leaves the species were
found. On the Cerro Cacao several small-
leafed plants provide a crawling and foraging
surface for the snails, whereas in Las Pavas
Heliconiaceae and different palms represent a
good place to search for H. gemma. In
Siquirres, specimens were often seen aestivat-
ing also on the upper surface of nearly every
kind of plant composing the secondary growth
near a small creek. On one occasion, the spe-
cies was found in dead decomposing Cecro-
pia-leaves on the ground in the rain forest.

A | B

Distribution

The species is limited to southern Central
America. The most northern record comes
from the Cerro Saslaya in Nicaragua, adjacent
to the Costa de Miskitos stretching along the
Caribbean coast. The southern limit is
reached around Siquirres and Parque
Nacional de Barbilla, the northeastern foothills
of the Cordillera de Talamanca. Helicina
gemma possibly occurs a little further the
south, but the data suggest that it is finally
absent in the Valle de Talamanca or even in
the Valle de Estrella, because the lower re-
gions which are normally inhabited by the spe-
cies have been fairly well investigated, and H.
gemma has not been reported.

FIG. 171. Variability of the female reproductive system of Helicina gemma, populations from A.
Tortuguero, IR 1654. B. Siquirres, IR 1652; scale bar 2 mm.

CLASSIFICATION OF HELICINIDAE 327

ST = ши

Orosi (n=3/4)

Cacao (n=6/2 Cacao (n=6/2)

( ) 9 ( Es Q

at o ms o
Las Pavas (n=42/24) | Las Pavas (n=43/25)
Vol A | (n=3/3 Volcan Arenal (n=3/3
ölcan Arenal (п ) ne В ( ) sa rs
Tort =6/3 Tortuguero (n=6/3
ortuguero (n ) Ä g ( )
— ni
Siquirres (n=45/78) Siquirres (n=45/78)
rn

nm
Orosi (n=3/4)

FIG. 172. Shell height of different populations of
Helicina gemma in Costa Rica according to
Table 10; on each line: mean value, standard
deviation, absolute range; number of individuals
given as “п = females/males”; upper line:
females, lower line: males; in between and
shaded: average of both for comparison with
populations of unknown sex; sex of individuals
from Orosi, Cacao INBio, Pitilla, Barra del
Colorado, Cerro Cocori, Finca Montafia Grande,
and Rio Aguas Frias not determined

anatomically (see text).

In Costa Rica, H. gemma is confined to the
Caribbean plain and the adjacent mountain
slopes (Fig. 182). The highest altitude is reached
in the northern Cordillera de Guanacaste on the
Cerro Cacao at about 1,100 m, where it just
crosses the chain of volcanoes. In this region,
the higher elevated areas still provide a suitable
climate on the otherwise drier Pacific slopes.
The absence in the intensively searched area
around Monteverde and its presence in the re-
gion of the Volcán Arenal and San Ramón at
lower altitudes provides evidence that H.
gemma does not occur much above 1,200 m. In
the Caribbean lowlands it is or was probably
fairly well distributed although not directly visible
on the map. On one hand, vast areas have been
deforested and used for agriculture, most prob-
ably causing a massive habitat loss, because
the species has thus far only been found in pri-

En Nr
Cacao INBio (n=10/14) Cacao INBio (n=10/14)
SEN —
ill =3/3 Pitill =5/3
Pitilla (n=3/3) Е )
— —
Barra del Colorado (п=8/2 Barra del Colorado (n=8/2
( ) 4 ( )
=—— —
Сегго Cocori (n=4/1 Cerro Cocori (n=4/1
( ) ( )
y +
Finca Montaña Grande (n=3/0) р Finca Montaña Grande (n=3/0) +
Rio Aguas Frias (n=9/11 Rio Aguas Frias (n=9/11
io Ag ( ) 9 ( )
— —
OS (holotype) San Carlos (holotype)
7 т т
1 1 1 1 ht 1 1 1 L 1 vf a 1 1 1
0 3 4 5 6 7 [mm] 8 0 1 2 3 4 5 6 [mm] 7

FIG. 173. Minor diameter of shell of different popu-
lations of Helicina gemma in Costa Rica according
to Table 10; for explanations see Fig. 172.

Cacao (n=6/2)

a, ю

Las Pavas (n=42/24)

E

Vólcan Arenal (n=3/3)

B
Tortuguero (n=6/3)

Siquirres (n=45/78)

Orosi (n=3/4)
ES
Cacao INBio (n=10/14)

Pitilla (n=3/3)

Barra del Colorado (n=8/2)

Cerro Cocorí (n=4/1)

Finca Montaña Grande (n=3/0)

Río Aguas Frías (n=9/11)

San Carlos (holotype)
=

ИВ

y 1
0 1 2 3 5 6 [mm] 7

FIG. 174. Expansion of outer lip of different popu-
lations of Helicina gemma in Costa Rica according
to Table 10; for explanations see Fig. 172.

328 RICHLING

| Cacao (n=6/2)

Las Pavas (n=42/24) ОВ

| Volcan Arenal (n=3/3)

=—
= = —
| Tortuguero (n=6/3)
= =
>
Siquirres (п=45/78
iqui ( )
—
Orosi (n=3/4)
+
Cacao INBio (n=10/14)
Pitilla (n=3/3)
—
Barra del Colorado (n=8/2)
ES
Cerro Cocori (n=4/1)
+ 1

Finca Montaña Grande (n=3/0)

Río Aguas Frías (n=9/11)

e o ня

I

San Carlos (holotype)

1
6 [mm] 7

FIG. 175. Height of last whorl of different popula-
tions of Helicina gemma in Costa Rica according
to Table 10; for explanations see Fig. 172.

Cacao (n=6/2)
er 2
+ o
Las Pavas (n=43/25)
Vólcan Arenal (n=3/3)
—
_ ne
Tortuguero (n=6/3)
u
+

Siquirres (n=45/78)

A >>
Orosi (n=3/4)

>
—
Cacao INBio (n=10/14)
ee
Pitilla (n=3/3)
—
+
Barra del Colorado (n=8/2)
=
Cerro Cocori (n=4/1)
+
Finca Montana Grande (n=3/0) +
Rio Aguas Frias (п=9/11)
—— |
San Carlos (holotype)
‘8
1 1 1 1 1: RES 1
0 1 2 3 4 5 6 [mm] 7

FIG. 176. Height of columellar axis of different
populations of Helicina gemma in Costa Rica ac-
cording to Table 10; for explanations see Fig. 172.

Cacao (n=6/2)

|

4
Las Pavas (n=33/11)

LS Pa ENT —

Tortuguero (n=6/3)
Sa —_— |
Siquirres (n=45/78)

l 1 1 1 e L 1
0 0.02 0.04 0.06 0.08 0.1 [mi] 0.14

FIG. 177. Shell volume of different populations of
Helicina gemma in Costa Rica according to
Table 10; for explanations see Fig. 172.

mary or secondary forest or forest-like habitats.
On the other hand, investigations (especially
INBio's) focus on protected areas compassing
mainly mountainous terrain (watersheds and
volcanoes). The only exception for the lowlands
is the zone of Colorado — Tortuguero on the NE
coast. The pattern of distribution is very similar
to that of H. funcki, except for the fact that the
latter has a wider range and obviously a higher
ecological tolerance.

Discussion

The color of the shells varies among the dif-
ferent populations in respect to the basic whorl
color. Specimens from the Arenal area (vol-
cano and Las Pavas) are pale yellow to even
whitish-transparent, whereas the populations
of the Caribbean lowlands (Siquirres,
Tortuguero, Cerro Cocori, Barra del Colorado)
and Pitilla are bright yellow. Only specimens
from the Cacao exhibit a strong tendency to-
wards brownish shells, which is otherwise only
very exceptionally observed in the Siquirres
population (about 3 out of 120 specimens).
The orange aperture is a common and con-
stant character of all specimens, as is the
crimson apex of a part of each population.

Preston (1903) compared Helicina gemma to
H. oweniana and recognized the greater con-
vexity (deeper impressed sutures) and fewer
whorls. He also stated that the orange color of
the outer lip extends further up the last whorl.
This is verified by the study of syntypes of H.
oweniana. Furthermore, Helicina oweniana is
more solid and larger, the outer lip is very
straight and perpendicularly expanded,
whereas in H. gemma it is narrowly reflexed
and curved backwards.

At the time of von Martens’ (1890-1901) and
Biolley’s (1897) reports on the Costa Rican
land molluscs, Helicina oweniana and subspe-

CLASSIFICATION OF HELICINIDAE 329

58 | -
56 [Es $ o o o |
©
o
5.4 -
o
5.2 4
5 E 1 - 1
0 200 400 600 800 altitude [т] 1200

FIG. 178. Relation of minor shell diameter (females used) to altitude of the
locality of the different populations of Helicina gemma in Costa Rica.

T
female >
ani male +
diam. +
[mm]
© o
6 1
$ o
8 Vo
¿9
o © © о
5.5 |- oo 08 8°
o ©
+ 9 ee
o 90
т o
i o
a о + + |
+ в
Е
+ ++
4.5 +
4 =e a ee
4 4.5 5 5.5 6 6.5 7 height [mm] 8

FIG. 179. Range of measurements in females and males of Helicina gemma
exemplary for height and minor diameter in the population from Las Pavas.

330 RICHLING

7 T —T T 7 Pen T
female >
male +
min
diam. + J
[mm]
©
6 FP o о © El
o
5 Yo o
œ
$$ PI
L% 2060
o 8
5.5 + we
o 4 o
o

+ +ou° ++

4 4.5 5 5.5 6 6.5 7 height [mm] 8

FIG. 180. Range of measurements in females and males of Helicina gemma
exemplary for height and minor diameter in the population from Siquirres.

4 4.5 5 5.5 6 6.5 7 height[mm] 8

FIG. 181. Plot of measurements for height and minor diameter for individuals
of Helicina gemma of unknown sex, exemplary for the population of Rio
Aguas Frias and the separation proposed.

CLASSIFICATION ОЕ HELICINIDAE 331

3500 - 4000 m
3000 - 3500 m
2500 - 3000 m
2000 - 2500 m
1500 - 2000 m
| 1000 - 1500 т
500 - 1000 m
| 100 - 500 m ae
eo co
| Е e coll. INBio
| © others |

FIG. 182. Records of Helicina детта in Costa Rica.

TABLE 10. Measurements of different populations of Helicina gemma given as mean value with
standard deviation, minimum and maximum value (min, max), and number of зреситепз; sex of
individuals of all populations included from the collection of INBio (both lower rows and Orosi) not
determined anatomically (see text) (min./max. diam. = ттог/та]ог diameter, col. axis = columellar
axis); linear measurements [mm], weight [9], volume [ml].

“Siquirres” (altitude 100 m)
“Tortuguero” (altitude 10 m) lots IR 1536, IR 1618, IR 1635, IR 1650,
lots IR 1621, IR 1654 IR 1652

Mean Mean
Sex value Deviation Min Max Number value Deviation Min Max Number

Weight
Volume
Volume

0.019 0.004 0.011 0.031 78
0.076 0.006 0.048 0099 45
0.053 0.003 0.045 0.068 78

0.015. 0.001 0:014 0.017
0:079 0:007 0:069 70.092
0057 0.001 0.056 70.058

Height f 6.67 0.25 7633 7.13 6 6.58 (0.24 ‘5:87 733 45
Height m 6.00 0.09 5.87 6.08 3 5.82 0:14 530 632 78
Maj. diam. f 6.05 0.15 590 6.28 6 6.06 0.18 538 655 “45
Maj. Чат. т 5.50 0.05 543 5.56 3 5.49 0:12 5.20 583 78
Min. diam. f 5.63 0.13 5.45 5:88 6 5:63 0:16 502 6:12 45
Min. diam. m 512 0.06. 5.02 518 3 5.05 0.10 4.74 538 78
Outer lip Е 3.80 0:12 357 3:98 6 3.81 0.12 3437 413 45
Outer lip m 3.55 0:05 350 3.62 3 3.54 0.08 332 3.79 78
Last whorl f 4.98 0:12 4:82 522 6 4.96 0.16; 448 552 45
Last whorl m 4.34 0.09 4.22. 4147 3 4.42 0.10 4.13 481 18
Col. axis f 531 CU 5:03 5.57 6 5.25 0:19 2174. 584 (45
Col. axis m 4.74 0.09 4.60 4.82 3 4.59 0.12 419 5094 78
Weight [0.015 0.002 00120018 6 (01022 0.003 100122003227 45

т 3

f 6

m 3

(Continues)

332 RICHLING
(Continued)

“Las Pavas” (altitude 760-800 m)

“Cacao” (altitude 1110 m) lots IR 947, IR 948, IR 1275, IR 1460,
lots IR 786, IR 1333 IR 1462, IR 1463
Mean Mean
Sex value Deviation Min Max Number value Deviation Min Max Number
Height f 6.41 0.24 590 6/72 6 6.36 0.2572 550, (729 242
Height m 5.48 017 531 5.64 2 5.53 0.15 514 60324
Maj. diam. f 5.92 0.17 566 6.16 6 5.98 O19" 1542 76571, 243
Ма). diam. т 5.27 0.10 15 5.37 2 5.31 0.16 5.03 5.90: 125
Min. diam. f 5.50 0.1677 5:21 5.70 6 5.53 0.17 493 612 43
Min. diam. т 4.68 0.22 4.46 4.90 De 4.85 0.13 4.52 553 125
Outer lip Г 3.85 012. 35651 4.03 6 SAT 0.13 3.38 4:28 42
Outer lip m 3.37 0.00 3.37 3.397 2 3.37 0.12 3.11 3.86 24
Last whorl f 5.10 0.14 484 5.33 6 4.90 0.20 410 555 42
Last whorl m 4.47 0.01 446 4.48 2 4.26 0.17 403 475 724
Col. axis f 525 0.14 “5:02 5.57 5 5.05 0.20 434 577 43
Col. axis m 4.39 0.10 4.29 448 2 4.34 012 408 472 25
Weight f 0.017 0.004 0.011 0.024 6 0.016 0.002 0.011 0.021 42
Weight m 0.015 0.03 0.012 0.017 2 0.014 0.003 0.007 0.025 21
Volume f 0.074 0.007 0.063 0.083 6 0.073 0.007 0.050 0.103 42
Volume m 0.047 0.000 0.047 0.047 1 0.047 0.003 0.040 0.061 21
“Vólcan Arenal” (altitude 720 m) “Orosi” (altitude 600 т)
lots IR 387, IR 740, IR 885, IR 1284 lot INBio 1487835
Mean Mean
Sex value Deviation Min Max Number value Deviation Min Max Number

Height f 6.46 O16 6.32 6.70 3 6.47 0.20 6.17 6.69 3
Height m 5.54 015 535—577 3 5.63 006 5.56 574 4
Maj. diam. + 6.08 0.10 5.92 6.16 3 6.17 024 581 5653 3
Maj. diam. m 5.39 0:04 532543 8 5.49 0.15 5.20 “5:75 4
Min. diam. f 5.60 0.12 5.50 578 3 5.62 0.16 5:38 57 3
Min. diam. m 4.92 0.06 4.84 5.01 3 5.04 0.10 489 5.24 4
Outer lip f 3.90 0.05 385 3.97 3 3.89 0.09 3.75 3.98 3
Outer lip m 3.61 006 3:54) 5.3.70 3 3103 0.11 3.48 3.76 4
Last whorl f 5.07 0.16 4.90 531 3 4.98 0.20 469 5.25 3
Last whorl m 4.32 0.10 4.22 4.48 3 4.48 0.07 4.40 4.56 4
Col. axis f 5.28 (01915.13 5:43 3 5.40 0.14 5.19 5.54 3
Col. axis т 4.50 0.12 436 4.68 3 4.68 0.15 456 490 9
Weight f 0.022 0.000 0.022 0022 1 - - - - -
Weight m - - - - - - - - - -

cies were the only known Central American
orange-lipped Helicinidae, which is why they
most likely assigned their Costa Rican or-
ange-lipped Helicinidae to the subspecies H.
oweniana coccinostoma or H. o. anozona.
Both probably synonymous subspecies are
more globular, with whorls a little more convex

(Continues)

than the nominal subspecies, thus rather re-
sembling H. gemma. But the differences to H.
gemma mentioned above refer to the subspe-
cies as well. Unfortunately, the original mate-
rial of these records has not yet been
rediscovered to check this assumption, but it
seems plausible.

(Continued)

“Cacao INBio” (altitude 1000-1100 т)
lots INBio 1484977, 1487886, 1488058,

“Barra del Colorado” (altitude 15-50 m)
lots INBio 1477917, 1484009, 1484012,

“Finca Montaña Grande” (altitude 10 т)

Mean

Sex value

Height f 6.38
Height My 552
Maj. diam. + 5.99
Maj. diam. m 5.24
Min. diam. f 5.61
Min. diam. m 4.80
Outer lip f 3:09
Outer lip m 3.43
Last whorl f 5.00
Last whorl т 4.27
Col. axis f 5.19
Col. axis m 4.54
Mean

Sex value

Height f 7.10
Height m 6.03
Maj. diam. f 6.44
Ма]. diam. m 5.55
Min. diam. f 6.01
Min. diam. т 5.16
Outer lip Г 4.12
Outer lip m 3.68
Last whorl f 5.50
Last whorl т 4.73
Col. axis f 5.80
Col. axis m 4.93
Mean

Sex value

Height f 7.20
Height m -
Maj. diam. f 6.63
Maj. diam. m -
Min. diam. + 6.15
Min. diam. т -
Outer lip Г 4.17
Outer lip m -
Last whorl f 5.48
Last whorl т -
Col. axis f 5.82
Col. axis m -

CLASSIFICATION ОЕ HELICINIDAE

1539438, 1539463

Deviation Min Max
023 5.74 6,84
015 516 517
OD 235.525 6:22
0.08 5.03 5.46
0.16 529 592
011 4.60 5.04
0.10 3.65 4.04
0.08 328 3:61
0.18 467 5.35
0.14 4.05 4.53
023 435 555
0.14 410 4.77

1484371, 1485286

Deviation Min Max
0.22 6.81 7.75
0.29 5.74 6.32
O15 607 650
017 538 512
0.14 5.69 6.27
0.13. 5.03 5.29
0.07 4.02 4.26
0.28 3.40 3.95
0417 531 5.95
0.29 444 5.02
0.18 5.49 6.23
0.13 4.80 5.05

lot INBio 1501218

Deviation Min Max
0:07 77.1072 7,30
0.07 6.53 6.74
0.06 6.12 6.24
0.06 4.08 4.26
0.04 5.42 553
0.06 5.72 5.88

Number

10
14
10

Number

N © N © N © N AON AN 00

Number

3

MIDI DEIN о ©:

“Pitilla” (altitude 700 m)
lots INBio 1463737, 1480045, 1480270,

333

1480284, 1480341, 1484672

Mean
value

6.33
9.92
9.19
4.85
5.34
4.56
3.64
3.30
4.80
4.22
5.36
4.30

Deviation Min Max
0.28 591 6.59
0.24 5.02 5.68
0.23 5.44 6.10
0.10 471 4.96
0.20 5.08 5.69
0.09 4.42 4.66
011 350 3.81
0.10 3.17 3.44
0.24 4.44 4.98
0.16 3.98 4.37
0.18 509 557
0:06 421 4.35

Number

CO Y CO CO © UY Y UU OW WwW

“Cerro Cocorí” (altitude 150 т)
lots INBio 1465444, 1478057

Mean
value

7.16
6.21
6.57
5.88
6.05
5.33
4.18
3.02
5.45
4.73
9.77
5.03

Deviation Min Max
0.26 6.83 7.55
0.00 6.21 6.21
0.20 6.32 6.88
000 5.88 5.88
0.16 583 6.32
000 533 5.33
0.17 4.03 4.52
0.00 3.82 3.82
028 502 5.91
0.00 4.73 4.73
0:28 536 612
0:00. 503 5503

Number

NAN = D ES

“Río Aguas Frías” (altitude 10 т)

Mean
value

7.14
9.98
6.56
5.64
6.06
5:19
4.11
3.69
9:92
4.50
5.81
4.85

lot INBio 1487942

Deviation Min Max
0.29 6.67 7.95
0.14 5.49 6.21
0-10: 6.22 6.78
0.10.1520 25:91
0.207 75:67 6539
0.09 4.88 5.38
0.13 38 4:39
0109 348385
02] ‘2.90 115.72
O12 4:26 4:84
035 488 6.49
Osi 74587 35:08

Number

9
11
g
11
9
11
9
ila
9
11
9
11

334 RICHLING

The records for H. oweniana and H. beatrix
by Monge-Nájera (1997) were checked in the
INBio-collection and partially belong to H.
gemma. For differences to Н. beatrix
riopejensis п. subsp. and H. monteverdensis
n. sp. see under these taxa.

Helicina (“Gemma”) monteverdensis
Richling, n. sp.

Helicina ометапа — Monge-Nájera, 1997:
113: Costa Rica [in part] [non L. Pfeiffer,
1849]

Helicina beatrix — Monge-Nájera, 1997: 113:
Costa Rica [in part] [non Angas, 1879]

Type Material

Holotype: INBio 3542627, female (leg. 1.
Richling, 24.02.1999, ex IR 634)

Paratype 1: INBio 3542628, male (same data
as holotype, 27.07.1999, ex IR 844)

Paratype 2: ZMB 103884, female (same data
as holotype)

Paratype 3: ZMB 103885, male (same data as
paratype 1)

Dimensions:

Holotype: 6.6/6.2/6.5/5.8/3.6/4.8/5.2 mm

Paratype 1: 5.9/5.5/5.8/5.1/3.6/4.6/4.8 mm

Paratype 2: 6.8/6.5/6.8/5.9/4.0/5.0/5.4 mm

Paratype 3: 5.6/5.7/6.1/5.1/3.6/4.6/4.5 mm

Type Locality

NW-Costa Rica, Puntarenas Province, Cor-
dillera de Tilarán, near Monteverde, Zona
Protectora Arenal-Monteverde, Reserva
Biológica Bosque Nuboso Monteverde,
Sendero Bosque Nuboso, about 10°18’08"М,
84°47’41"W, 1,550 т a.s.l., cloud forest.

Type Material of Synonymous Taxa or Similar
Species

Helicina merdigera L. Pfeiffer, 1855

Helicina merdigera L. Pfeiffer, 1855: 102:
Mexico: Vera Cruz (leg. Sallé, coll. Hugh
Cuming)

Type Material: BMNH 20010752: leg. Sallé,
coll. Hugh Cuming
The lot contains three specimens, of which
one is completely fragmented except for the
aperture with the operculum still inside. The

two remaining shells are very similar to each
other. The species was neither figured by
the author nor does the description give any
useful hints for the identification of the type.
The slightly larger shell is here selected as
lectotype (Fig. 183), the denticle at the tran-
sition of the basal outer lip into the columella
is stronger developed. Whereas the
paralectotype (Fig. 184) is whitish, it shows
a tinge of yellowish-brown.
Dimensions:
Lectotype 20010752.1:
5.0/5.1/5.4/4.7/2.9/3.7/4.0 mm
Paralectotype 20010752.2-3 (latter frag-
mented): 4.5/5.0/5.3/4.5/3.0/3.4/3.7 mm

Type Locality: “Vera Cruz, Mexico”.
Helicina fragilis Morelet, 1851
Helicina fragilis Morelet, 1851: 17 (not figured)

Type Material: BMNH 1893.2.4.809-12, coll.

Morelet, purchased from H. Fulton

The Morelet collection was bought by H.

Fulton and subsequently purchased by the

BMNH. The “type” among the four syntypes

(Figs. 185, 186) is marked with an “х” on the

shell. This specimen 15 here selected as lec-

totype (Fig. 185). It matches best the dimen-
sions given by Morelet. It is neither the largest
nor the smallest specimen of the type lot.

Dimensions:

Lectotype 1893.2.4.809:
6.0/5.5/5.7/5.1/3.4/4.2/4.9 mm; 5 teleo-
conch whorls

Paralectotypes 1893.2.4.810-12:
7.2/6.6/7.0/6.1/4.1/5.2/5.9 mm; 47/, teleo-
conch whorls
5.9/5.6/6.0/5.2/3.5/4.3/4.7 mm; 4*/, teleo-
conch whorls
5.2/5.2/5.6/4.8/3.2/3.9/4.1 mm; 4% teleo-
conch whorls

Type Locality: “sylvas Petenenses” [Guate-
mala: Petén Department].

Helicina mohriana L. Pfeiffer, 1861
Helicina mohriana L. Pfeiffer, 1861: 172-173
Type Material: Not located, probably lost.

Type Locality: Mexico, Orizaba (leg. Mohr)
[State of Vera Cruz]

CLASSIFICATION OF HELICINIDAE 339

FIGS. 183-188. Helicina spp. FIG. 183. Helicina merdigera, BMNH 20010752.1, lectotype, height 5.0
mm. FIG. 184. Helicina merdigera, BMNH 20010752.2, paralectotype, height 4.5 mm. FIG. 185. Helicina
fragilis, BMNH 1893.2.4.809, lectotype, height 6.0 mm. FIG. 186. Helicina fragilis, ВММН 1893.2.4.810-
812, paralectotypes, height 7.2, 5.9, 5.2 mm. FIG. 187. Helicina elata, syntype, NHMB 15269, height 4.0
mm. FIG. 188. Helicina diaphana, probable syntype, BMNH 196282, height 3.7 mm; scale bar 2.5 mm.

336 RICHLING

Helicina elata Shuttleworth, 1852
Helicina elata Shuttleworth, 1852: 304

Type Material: Syntype NHMB 15269: leg.
Jacot-Guillarmod (Fig. 187)
Shuttleworth stated “specimena pauca vidi”,
but his collection in the NHMB contains only
a single specimen. He probably exchanged
the others.
Dimensions:
Syntype: 4.0/4.3/4.3/3.8/2.4/2.9/3.1 mm

Type Locality: Mexico: Vera Cruz: Cordova
[Cordoba].

Helicina diaphana L. Pfeiffer, 1852

Helicina diaphana L. Pfeiffer, 1852: 98: Hondu-
ras (leg. Mr. Dyson, coll. Hugh Cuming)

Type Material: Probable syntype BMNH
196282: Honduras (It was stated at the time
of registration that it was possible that some
original labeling from the back of the speci-
men board was not kept. i.e. MC initials and
Dyson as collector.)

The assumption of the type status is sup-
ported by the fact that Rehder (1966) refers
to a photograph of the syntype in the BMNH,
probably the specimen was registered when
he requested the photograph; the time pe-
riod would make this seem likely. The speci-
men (Fig. 188) shows various details of the
original description. The operculum is still
inside the shell.

Dimensions:

Syntype?: 3.7/4.8/4.9/4.2/2.7/3.0/3.0 mm

Type Locality: “Honduras”
Examined Material

LEG. |. RICHLING

Guanacaste: N Santa Elena, Sendero at
Mirador Gerardo, 10°22'19"N, 84°48’25"\М,
1,450 m a.s.l., 28.07.1999: (IR 855); (IR
856); 14.08.1999: (IR 929); (IR 930);
19.02.2000: (IR 1226); (IR 1228); (IR 1229);
24.02.2001: (IR 1416); (IR 1418); (IR 1419);
(IR 1420)

Puntarenas: Near Monteverde, about
10°17'24"N, 84%48'04"W, 1 km before
entrance on road to reserve, 1,500 m a.s.l.:
26.07.1999: (IR 825); 13.08.1999: (IR 920);
(IR 921)

Zona Protectora Arenal-Monteverde:
Reserva Biológica Bosque Nuboso
Monteverde (about 10*18'08"N, 84°47'41"W,
1,500-1,650 m a.s.l.): 27.07.1999: (IR 844);
(IR 845); 18.02.2000: (IR 1196); (IR 1197);
25.02.2001: (IR 1436); Sendero Bosque
Nuboso: 24.02.1999: (IR 634); (IR 636);
Sendero Roble: 18.02.1998: (IR 302);
Sendero Chomogo: 18.02.1998: (IR 296)
About 4 km N Santa Elena, Skywalk,
10°18’33"М, 84°49'42"W, 1,330 т a.s.l.,
2702.1999: (IR 681)

INBio COLLECTION
Guanacaste: Zona Protectora Arenal-

Monteverde, Santa Elena: Sendero
Encantado, 10°21'57"N, 84°47'27"W, 1,400
m a.s.l.: leg. Kattia Martinez, 13.07.1994: 4
ads., 6 s.ads., 7 juvs. (INBio 1479270); 3
ads., 6 juvs. (INBio 1479273); Sendero
Encantado, 10°21'57"N, 84°47’27"W, 1,200
т a.s.l.: leg. Каша Martinez, 21.06.1996: 3
ads., 3 s.ads., 2 juvs. (INBio 1487482); 2 ads.
(INBio 1498545); Sendero Rancho Alegre,
10°21'24"N, 84°47’47"W, 1,440 m a.s.l.: leg.
Kattia Martinez, 13.11.1994: 2 ads. (INBio
1485425); 1 ad., 1 s.ad. (INBio 1485431)
Zona Protectora Arenal-Monteverde: 1km
NE de la casa de información Reserva
Santa Elena, 10%21'05"N, 84°47'40"W,
1,550 т a.s.l.: leg. Alexander Alvarado
Mendez, 12.01.2000: 3 ads., 2 s.ads. (INBio
3098462); Sendero Tabacón, 10°22'55"N,
84°47'40"W, 900 т a.s.l.: leg. Alexander
Alvarado Mendez, 03.09.1999: 1 ad. (INBio
1501135)

Alajuela: Zona Protectora Arenal-Monteverde:

Camino a El Valle, 10°19’44"N, 84°45’55"W,
1,580 m a.s.l.: leg. Кафа Martinez,
20.06.1996: 1 juv. (INBio 1498646);
Estación Alemán, 10%18'11"N, 84*44'49"W,
940 m a.s.l.: leg. Dunia Garcia, 10.11.1994:
1 s.ad. (INBio 1475971); Refugio El Valle,
10°19’04"М, 84°46’41"W, 1,800 m a.s.l.: leg.
Kattia Martinez, 11.01.1995: 2 ads., 1 s.ad.,
2 juvs. (INBio 1498728)

Zona Protectora Arenal-Monteverde, Sector
Peñas Blancas, Estación Aleman,
10°18’09"М, 84*44'52"W: 900 m a.s.l.: leg.
Kattia Martinez, 18.08.1994: 1 ad. (INBio
1480178); 1,140 m a.s.l.: leg. Zaidett
Barrientos, 10.11.1994: 2 ads., 1 juv. (INBio
1473354); 1 ad. (INBio 1473359)

Zona Protectora Arenal-Monteverde,
Reserva Biológica Bosque Nuboso
Monteverde: Sendero Pantanoso, 10°18'19"N,

CLASSIFICATION OF HELICINIDAE 087

84°47'10"W, 1,620 m a.s.l.: 08.11.1994: 1
s.ad., 1 juv. (INBio 1479473); 15.01.1995: 1
ad., 1 s.ad. (INBio 1498821); 04.04.1995: 1
ad. (INBio 1484024) (all leg. Kattia Martinez);
leg. Alejandro Azofeifa, 04.04.1995: 2 juvs.
(INBio 1484832)

Cartago: ?Reserva Indígena Chirripó, Zona de
captación Rio Humo, 09°42'47"N,
83°25'52"W, 1,550 m a.s.l., leg. Zaidett
Barrientos, 27.06.1996: 1 ad., 4 s.ad.s, 1 juv.
(INBio 1498787)

Puntarenas: Zona Protectora Arenal-
Monteverde, Reserva Biológica Bosque
Nuboso Monteverde: Estación la Casona,
10°18'11"N, 84*47'50"W, 1,600 т a.s.l.: leg.
Kattia Martinez, 30.10.1996: 2 ads., 2 juvs.
(INBio 1498680); 1 ad., 2 s.ads., 2 juvs.
(INBio 1498684); Sendero Bosque Eterno,
10°18’22"М, 84*47'40"W, 1,600 m a.s.l.:
20.01.1995: 1 ad., 1 s.ad. (INBio 1483833);
04.04.1995: 4 ads., 4 s.ads., 1 juv. (INBio
1485231); 04.07.1995: 3 ads. (INBio
1485229); 19.06.1996: 1 ad. (INBio
1498693) (all leg. Kattia Martinez); Sendero

Bosque Nuboso, 10°17'59"N, 84*47'36"W,
1,600 т a.s.l.:, 14.06.1994: 2 ads. (INBio
1466785); 16.07.1994: 1 ad., 1 s.ad. (INBio
1479239); 1 s.ad. (INBio 1479860);
16.09.1994: 2 ads., 1 s.ad. (INBio 1480117);
14.01.1995: 3 ads., 1 s.ad. (INBio 1498583);
22.06.1996: 2 ads., 1 juv. (INBio 1498519)
(all leg. Kattia Martinez); Sendero Bosque
Nuboso, 10°17’59"М, 84°47’36"W, 1,520 т
a.s.l.: leg. Zaidett Barrientos, 14.10.1994: 1
ad., 1 juv. (INBio 1468138), 1 ad. (INBio
1468209); Sendero Bosque Nuboso,
10°17’59"N, 84*47'43"W, 1,520 m a.s.l.: leg.
Alexander Alvarado Mendez, 03.02.1999: 1
ad. (INBio 1501428); Sendero Chomogo,
10°18’22"М, 84*47'23"W, 1,690 т a.s.!.: leg.
Kattia Martinez, 13.08.1994: 1 ad. (INBio
1480143); 1 juv. (INBio 1480156); Sendero
Chomogo, 10°18’22"М, 84*47'23"W, 1,640
mi a.s.l.: 10.10.1994: (ad. 2 $-ads.,. Juv.
(INBio 1485418); 25.11.1995: 1 ad. (INBio
1498708); 18.02.1997: 1 аа. (INBio
1498840) (all leg. Каша Martinez); Sendero
el Río, 10°18’29"М, 84°47’37"W, 1,600 т

FIGS. 189, 190. Helicina monteverdensis n. sp. FIG. 189. Holotype, INBio 3542627, height 6.6 mm.
FIG. 190. Paratype 2, ZMB 103884, height 6.8 mm; scale bar 2.5 mm.

338 RICHLING

a.s.l.: 04.04.1995: 1 ad., 1 s.ad. (INBio
1484664); 04.07.1995: 1 ad., 1 s.ad. (INBio
1484659) (both leg. Alejandro Azofeifa);
15.07.1994: 1 s.ad. (INBio 1480137); 1 s.ad.
(INBio 1480151); 16.09.1994: 2 s.ad. (INBio

1480122); 08.12.1994: 2 juvs. (INBio
1480142); 04.07.1995: 6 ads. (INBio
1485230); 29.10.1996: 1 ad. (INBio
1498835) (all leg. Kattia Martinez); Sendero

el Roble, 10*18'16"N, 84°47’27"W, 1,600 т
a.s./.: leg. Каша Martinez, 08.11.1994: 1 ad.
(INBio 1479363); Sendero el Camino,
10°18’03"М, 84%47'15"W, 1,560 т a.s.l.:
23.05.1994: 2 ads. (INBio 1466975);
14.07.1994: 1 s.ad. (INBio 1479374); 1 ad.
(INBio 1480153); 13.12.1994: 1 ad., 1 s.ad.
(INBio 1484678) (all leg. Kattia Martinez)

OTHER SOURCES

COSTA RICA
Guanacaste: 6 mi NNE Tilaran, on road to
Arenal [about 10°33’N, 84°59'W?],

17.07.1971: 1 ad. (UF 69855)

San José: Alata la Palma [Alto Palma?, about
10°03'N, 84°00’W], 07.08.1971: 1 s.ad. (UF
69856)

Description

Shell (Figs. 189, 190, 3361-J): Conical, thin
and fragile, medium to small sized,
semitransparent, shiny. Color: whorls
unicolored, whitish-opaque, yellowish to
bright yellow or even brownish; apical whorls
sometimes with a crimson spot. Surface
textured with fine and regular growth lines
(Fig. 192), causing the glossy appearance.
Embryonic shell with about 1 whorl; 3% (3°/,—
4) subsequent whorls convex; last whorl very
evenly rounded at the periphery; whorls
equally extending in size or last whorl even
more rapidly increasing in diameter, forming
a pointed spire. Suture moderately
impressed. Aperture oblique and curved
backwards; last whorl slightly more
descending towards aperture and inserting
below periphery. Outer lip independently from
the color of the whorls always whitish-
opaque, slightly thickened and very narrowly
expanded and reflexed. Transition to
columella continuous with a slight notch.
Columella slightly curved, transition to the
body whorl without any groove. Basal callus
very weakly developed and slightly
granulated.

FIG. 191. Axial cleft and muscle attachments of
Helicina monteverdensis n. sp., INBio 3542627
(holotype) ; scale bar 2.5 mm.

Internal Shell Structures: (Fig. 191)

Teleoconch Surface Structure (Fig. 192): The
surface structure is similar to that of Helicina
gemma.

Embryonic Shell. Among the specimens
studied from the populations of Monteverde
and Mirador Gerardo, the embryonic shells
show large deviations in structure, showing
all intermediary variations in pit size from
pronounced pits as in Helicina gemma to
small ones as in H. beatrix (Fig. 193A, B).
For comparison with H. fragilis, a
Guatemalan specimen was studied (Fig.
194), but due to the high variations in H.
monteverdensis n. sp., structural differences
cannot be codified. By the way of contrast,
the size differs remarkably: the type lot and

FIG. 192. Teleoconch surface structure of
Helicina monteverdensis п. sp., 2" whorl; scale
bar 100 pm.

CLASSIFICATION OF HELICINIDAE 339

other Guatemalan specimens show a range
of 580 to 760 um, whereas the embryonic
shell of the Costa Rican specimens of H.
monteverdensis n. sp. is clearly larger (830
to 1,000 рт). These differences clearly
exceed the possible deviations caused by
effects of altitude, at least as far as the
results for H. gemma suggest. Furthermore,
the specimens from UF 189883 with a small
embryonic shell size also originate from an
altitude of 1,300 m. The same is true for the
type lot of H. merdigera, the embryonic shell
of which is also remarkably smaller.

The smaller specimens from the population
of Mirador Gerardo possess somewhat
smaller embryonic shells on the average.
The relation of embryonic shell size to the
shell size is discussed below.

Diameter: 935 um (+ 25) (900-1,000) (п = 21)
(IR 634, IR 844, Monteverde); 897 um (+ 29)
(830-990) (п = 15) (IR 1226, Mirador
Gerardo); 708 um (+ 44) (620-760) (n = 4)
(BMNH 1893.2.4.809-812, type lot of
Helicina fragilis, lectotype: 620 um); 610 um

FIG. 193. Embryonic shell of Не/ста FIG. 194. Embryonic shell of Helicina fragilis:
monteverdensis n. sp. A. Mirador Gerardo, IR scale bar 100 um.
1226. B. Same data; scale bar 100 um.

340 RICHLING

FIG. 195. Operculum of Helicina monteverdensis
n. Sp., INBio 3542627; scale bar 1 mm.

(UF 190045: Guatemala: Alta Verapaz De-
partment, 9 km W of Lanquin, 15°35’03"N,
90°03'20"W, 690 m а.$.1., leg. Е.С. Thompson
et al.); 620 um (+ 20) (600-640) (п = 2) (UF
189883: Guatemala, Alta Verapaz Depart-
ment, 1.5 km SE of (San Juan) Chamelco,
15°24’20"N, 90°18'28"W, 1,300 m a.s.l., leg.
Е.С. Thompson); 608 um (+ 18) (580-630) (п
= 5) (UF 237423: Guatemala: Chama, leg.
A.A. Hinkley, ex coll. Beal-Maltbie); 650 um (+
0) (650) (n = 2) (BMNH 20010725.1-2, type
lot of Helicina merdigera).

Operculum (Fig. 195): Very slightly calcified,
calcareous plate covering only part of the
outer surface, thickened towards the col-
umellar side. Color whitish-yellow to pale
horny-amber, nearly transparent throughout.
Columellar side slightly S-shaped, upper
end acute and pointed, lower end rounded
and continuously changing into outer mar-
gin.

Animal (Fig. 338F, G): The color resembles
that of Helicina gemma, but varies more
strongly. In the individuals originating from
Monteverde, the head-foot is usually much
paler, only the tentacles are tinged grey; in
the mantle pigmentation, the whitish-yellow
part is very prominent, but the dark spots
very often form an irregular band at the pe-
riphery. The pattern in specimens from
Mirador Gerardo more closely resembles
that of H. gemma, but it ranges from nearly
black individuals with some very small yel-
lowish-white spots to the reverse, with few
slender black lines and spots. In this popu- FIG. 196. Radula of Helicina monteverdensis n.
lation, a distinct dark band has never been sp. A. Centrals. B. Comb-lateral. C. Marginals;
observed. Usually the dark part of the head- scale bar 50 um.

CLASSIFICATION OF HELICINIDAE 341

foot is larger and more intensive in speci-
mens with a darker mantle pigmentation.
These considerable variations within a
population seem to be typical. The few
specimens from the Cartago Province are
either dark or pale. The mantle pigmentation
is clearly visible through the thin shell.

Radula (Fig. 196): All centrals lack well de-
fined cusps. Comb-lateral with 8 cusps, only
one exception with 11 or 13 cusps on the
other side respectively (Fig. 196C). Cusps
on marginals rapidly increasing in number.
Radula with about 46-73 rows of teeth.

Female Reproductive System (Figs. 197-

199): The reproductive tract of Helicina
monteverdensis n. sp. is similar to that of H.
gemma and equal in size and proportions
except for the bursa copulatrix. The latter
seems to be consistently smaller, although
specimens collected during both (dry and
rainy) seasons were studied in either spe-
cies to exclude possible physiological
changes. Furthermore, the shape of the
bursa copulatrix varies more in Н.
monteverdensis n. sp. Lobes are not al-
ways distinctly developed and they are less
regular.

FIGS. 197-199. Helicina monteverdensis n. sp. FIG. 197. Female reproductive system, IR 844. FIG.
198. Variability of the female reproductive system, population from Mirador Gerardo, IR 1226. FIG.
199. Variability of the female reproductive system, population from Monteverde, IR 844; scale bar
1 mm (Fig. 197), 2 mm (Fig. 198-199).

342 RICHLING

Mirador Gerardo (n=51/45) en 9
A

Monteverde (n=19/10)

FE RE se +
рр

aja
| E Ss
Monteverde INBio (n=27/14)

Petén (type H. fragilis)

| 2 3 4 5 6 [mm] 7
FIG. 200. Shell height of different populations of
Helicina monteverdensis n. sp. in Costa Rica
according to Table 11; on each line: mean value,
standard deviation, absolute range; number of
individuals given as “п = females/males”; upper
line: females, lower line: males; in between and
shaded: average of both for comparison with
populations of unknown sex; sex of individuals
from Santa Elena and Monteverde INBio not
determined anatomically (see text).

Morphometry and Sexual Dimorphism (Table
11, Figs. 200-205)

Around the type locality, the species was
collected in favorable numbers for the mor-
phometric analysis. Three localities are differ-
entiated from N to S: Mirador Gerardo at the
beginning of the northern slope of the Cordil-
lera de Tilarán, the Santa Elena reserve and
finally samples mainly in and around the
Reserva Biológica Bosque Nuboso
Monteverde. The latter are separated in the
lots that were anatomically investigated (coll.
IR) and INBio's. Both populations included
from the latter collection (Figs. 200-204, be-

Mirador Gerardo (n=51/45)

Monteverde (n=19/10)

Santa Elena (n=9/10)

Monteverde INBio (n=29/16)

Petén (type H. fragilis)

6 [mm] 7

FIG. 201. Minor diameter of shell of different
populations of Helicina monteverdensis п. sp.
in Costa Rica according to Table 11; for
explanations see Fig. 200.

Mirador Gerardo (n=51/45)

Q, to

Monteverde (n=19/10)

Santa Elena (n=7/10)

=

Monteverde INBio (n=27/14)

Petén (type H. fragilis)

FIG. 202. Expansion of outer lip of different
populations of Helicina monteverdensis n. sp. in
Costa Rica according to Table 11; for explanations
see Fig. 200.

low thick line), which could not be analyzed for
their sex, were separated, as in Helicina
beatrix, to avoid the artificial high deviations of
measurements with mixed sexes. For com-
parison in the discussion, the lectotype of H.
fragilis is included in the figures.

Morphometry: The shells of the population
“Mirador Gerardo” are remarkably smaller in
all measurements than the other popula-
tions, which are similar to each other. The
constant pattern of differences of the popu-
lations for the different measurements sug-
gests the same shape of the shell. In
comparison with Helicina gemma closely
resembling the species, it is remarkable that
only the populations with the biggest shells
(“Finca Montaña Grande”, “Cerro Cocorí”)
attain the size of the new species from
Monteverde, whereas in the closer popula-
tion “Mirador Gerardo”, the average shell

Mirador Gerardo (n=51/45)

q

Monteverde (n=19/10)

Santa Elena (n=7/10)

Monteverde INBio (n=27/14)

Petén (type H. fragilis)
>

0 1 2 3 4 5

FIG. 203. Height of last whorl of different
populations of Helicina monteverdensis n. sp. in
Costa Rica according to Table 11; for
explanations see Fig. 200.

CLASSIFICATION OF HELICINIDAE 343

Mirador Gerardo (n=50/42)
eae

Monteverde (n=17/7)

Santa Elena (n=9/10) rs

y?-_— — — ————
Monteverde INBio (n=28/15)
——

Petén (type Н. fragilis)

6 [mm] 7

FIG. 204. Height of columellar axis of different
populations of Helicina monteverdensis п. sp. in
Costa Rica according to Table 11; for explanations
see Fig. 200.

size is clearly smaller than in all H. gemma-
populations. Thus, H. monteverdensis п. sp.
displays greater differences in size in just a
few closely located and ecological similar
populations than does the comparatively
widespread H. gemma in the several popu-
lations investigated. This observation sup-
ports the distinctness of the two species.

Sexual Dimorphism: In both populations, the
females possess bigger shells. The data
overlap only very slightly with the measure-
ments of the males, possibly because of the
comparatively high number of specimens
included, rendering individual high devia-
tions more likely. The separation of both
sexes is additionally shown for the original
set of data of height and minor diameter for
the population “Monteverde” and “Mirador
Gerardo” (Figs. 206-207). In volume, the
males only amount to */, or less than the fe-
males. The differences displayed for the
populations are very constant for each mea-
surement. As explained in H. beatrix, the
well-developed sexual dimorphism allows a

Mirador Gerardo (n=51/4

Q,+o

Monteverde (n=11/10)

1 | 1 1
0 0.02 0.04 0.06 0.08 0.1 [ml] 0.14

FIG. 205. Shell volume of different populations of
Helicina monteverdensis n. sp. in Costa Rica ac-
cording to Table 11; for explanations see Fig. 200.

separation of sets of mixed data (illustrated
for Monteverde INBio, Fig. 208).

Habitat

Helicina monteverdensis n. sp. has been
observed crawling and aestivating on leaves
of several small-leafed plants of the under-
growth. Especially at Mirador Gerardo where
Heliconiaceae are abundant on the forest
margins, it was mainly found on older leaves
of plants that were covered with moss and al-
gae. During rainy and cloudy weather, it
crawls on both the upper and lower side of the
leaves. It has not been seen in leaf litter. In
Monteverde and adjacent areas, the species
occurs sympatrically with H. funcki and
Alcadia hojarasca.

Distribution (Fig. 209)

Helicina monteverdensis n. sp. seems to be
adapted to higher altitudes, most records
around 1,500 m. A single exception is the site
north of Tilaran (about 700 m?), which can
also be located south of or today within the
reservoir, because it was collected before the
construction of the Embalse de Arenal reser-
voir. The species occurs in the Cordillera de
Tilaran and the Cordillera Central. It is well
distributed in the cloud forests of the area of
Monteverde — Santa Elena. The record from
the northeastern slopes of the Cordillera
Talamanca is only tentatively attributed to the
new species; further material is required. As
the record from the Cordillera Central indi-
cates, the species is probably much more
widespread, but many possible sites have ei-
ther been poorly investigated or not at all. Ex-
cept for the Cordillera de Guanacaste, H.
monteverdensis п. sp. replaces H. gemma at
higher altitudes.

Discussion

The population in Monteverde consists of
brightly yellow to whitish shelled specimens,
whereas individuals from the Mirador Gerardo
within a very limited collection area exhibit a
wider range of color, additionally including
brownish specimens. The outer lip is always
whitish.

Helicina monteverdensis n. sp. most closely
resembles H. gemma from which it differs by
the constantly different color of the outer lip
and adjacent part of the last whorl. Besides

344 RICHLING

7 т т T T НН:

female >
. male +
min
diam. + a
[mm]
6 =
5.5 F

|

4 4.5 5 5.5 6 6.5 7 heightImm] 8

FIG. 206. Range of measurements in females and males of Helicina
monteverdensis n. sp. exemplary for height and minor diameter in the
population from Monteverde.

female >
male +

5.5

4.5

4 4.5 5 5.5 6 6.5 7 height [mm] 8

FIG. 207. Range of measurements in females and males of Helicina
monteverdensis n. sp. exemplary for height and minor diameter in the
population from Mirador Gerardo.

CLASSIFICATION ОЕ HELICINIDAE 345

4 4.5 5 5.5 6 6.5 7 height [mm] 8

FIG. 208. Plot of measurements for height and minor diameter for individuals
of Helicina monteverdensis n. sp. of unknown sex, exemplary for the
population of Monteverde INBio and the separation proposed.

3500 - 4000 m
3000 - 3500 m |
2500 - 3000 m |
2000 - 2500 m
1500 - 2000 m
1000 - 1500 т

500 - 1000 m

100 - 500 m |
e coll. К

г e coll. INBio |
© others

FIG. 209. Records of Helicina monteverdensis n. sp. in Costa Rica.

346 RICHLING

TABLE 11. Measurements of different populations of Helicina monteverdensis n. sp. given as mean
value with standard deviation, minimum and maximum value (min, max), and number of specimens;
sex of individuals from Santa Elena and Monteverde INBio not determined anatomically (see text)
(min./max. diam. = minor/major diameter, col. axis = columellar axis); linear measurements [mm],
weight [g], volume [ml].

“Mirador Gerardo” (altitude 1450 т) “Monteverde” (altitude 1500-650 т)
lots IR 929, IR 1416 lots IR 296, 634, 825, IR 844, IR 1196
Mean Mean

Sex value Deviation Min Max Number value Deviation Min Max Number

Height f 5.93 0.24 5.40 6.95 51 6.74 0.23 565 7289 19
Height m 5.31 0:15 2485 575 45. 5.58 031 5 60910
Mai. diam. f 5:56 0.23 74,937 648 51 6.32 0.17 S61 652 19
Ма}. dam. m 4.96 0.14 445 541 45 5.46 0.34 482 602 10
Min. diam. f 5.14 0.20 459 601 51 5:87 OMG 518 6129519
Min. diam. m 4.54 0.12 419 488 45 4.99 0.29 434 5.43 10
Outer lip f 3.50 0.11 32514.06. 351 3.91 0.15 3.60 4.18 19
Outer lip т 3.21 0.10%. 27958 43.527 45 3.48 0.20 3.10 3.80 10
Last whorl f 4.62 0.18 418 540 5 522. 0.23 447 558 19
Last whorl m 4.17 OMS EN 2 AS 4.54 0.21 416 495 10
Col. axis f 4.75 0:20) 437 552 50 5.33 0.207 2445 515
Col. axis m 4.22 0. 13213891457 222 4.56 0.19 4.03 4.80 7
Weight f 0.013 0.002 0.009 0.022 51 0.016 0.002 0.011 0.019 11
Weight m 0.011 0.002 0.007 0.017 45 0.013 0.002 0.008 0.018 10
Volume f 0.058 0.07 0.042 0.090 51 0.084 0.008 0.055 0.094 11
Volume m 0.040 0.003 0.032 0.051 45 0.052 0.008 0.037 0.064 10
“Monteverde INBio” (altitude 1520-1690 т)
lots INBio 1466785, 1466975, 1468138,
1468209, 1479239, 1479363, 1480117,
1480143, 1480153, 1483833, 1484024,
1484659, 1484664, 1484678, 1485229,
1485230, 1485231, 1485418, 1498519,
“Santa Elena” (altitude 1200-1550 т) 1498583, 1498680, 1498684, 1498693,
lots INBio 1479270, 1479273, 1485425, 1498708, 1498821, 1498835, 1498840,
1485431, 1487482, 1498545, 3098462 1501428
Mean Mean
Sex value Deviation Min Max Number value Deviation Min Max Number
Height f 6.72 0.19 646 7.21 74 6.82 0.20; 6.41 7.20 127
Height m 5.90 0.21 543 625 10 5.93 0.20 5.61 643 14
Mai. diam. f 6.29 0.17 592 6.63 8 6.42 0.19 597 6.78 29
Mai. Чат. т 5.58 0.20 501 599 10 5.70 0.16 5:43" 6.10 16
Min. diam. f 5.84 013 2555 26.19 9 5.99 0.15 566 70622729
Min. diam. т 5.13 O17 453 556 10 525 0:12 75.007 5.53 16
Outer lip f 3.83 0.16 7358 АТ 7 3.95 0:12 365 450027
Outer lip m 3.56 0.18 318 385 (10 3.60 OM 333 3.91 14
Last whorl f 5.07 0.24 4.70 5.70 7 5.24 0:19; 480 566 2
Last whorl m 4.61 0237 212515 10 4.66 0.14 432 50014
Col. axis f 5.42 0.10 529 5.68 9 5.51 0.19 5.13 5.89 28
Col. axis m 4.79 0.23 413 515 10 Ara. 0:15, 441 5.08 615

CLASSIFICATION ОЕ HELICINIDAE 347

morphometric differences discussed above,
the whorls of H. monteverdensis п. sp. appear
more inflated. The high variation in body color
of H. monteverdensis n. sp. has not been ob-
served in other species.

Small, thin-shelled, yellowish-whitish-trans-
parent Helicinidae without spiral striations
from Central America (mainly Mexico and
Guatemala) were normally referred to Helicina
fragilis or its subspecies H. fragilis elata with
the synonyms Helicina merdigera and the du-
bious Helicina mohriana (e.g., Fischer &
Crosse, 1880-1902; von Martens, 1890-
1901; Baker, 1922a, 1928). Wagner (1908) did
just this when he studied such specimens in
the collection of the ZMB from the more south-
ern parts of Costa Rica. Those specimens
from the Valle de Talamanca and the neigh-
boring Valle de Estrella were reinvestigated
and partly belong to H. escondida п. sp., Н.
chiquitica or partly remain dubious in their
identification (see under H. chiquitica).

To clarify the classification of the small, frag-
ile, whitish Helicinidae from the Monteverde
area, the type material of the taxa mentioned
above and many lots from Mexico, Guate-
mala, Honduras and Belize were studied,
mainly in the collection of the UF.

Helicina fragilis differs from MH.
monteverdensis n. sp. in having fewer whorls
and a much smaller embryonic shell, although
the general size of the shells is about the
same. The aperture and last whorl is relatively
lower, the spire higher. Furthermore, in many
specimens of H. fragilis, a slight angulation at
the periphery (stronger in juvenile stage) is
maintained to at least the beginning of the last
whorl (e.g., paralectotypes), which in H.
monteverdensis n. sp. is always rounded. The
types were collected under decaying leaves,
whereas the new species is arboreal. Baker
(1928) observed H. fragilis elata on rock
ledges, weeds and low brush, but assumed
the aestivation on the ground.

The lectotype of Helicina merdigera is
slightly angulated at the periphery throughout
the last whorl, which is a little shouldered be-
low the suture. In comparison with H. fragilis
and the new species, the shell surface is more
roughly sculptured with irregular growth lines
and oblique grooves, the outer lip is less re-
flexed and the thickening is shifted a little in-
wards. The lower part of the aperture
protrudes further, forming a nearly rectangular
edge at the transition to the columella,
whereas in H. fragilis and H. monteverdensis

n. sp., the transition is continuously or even
with a little notch. Other features are similar to
H. fragilis providing additionally evidence for
the distinctness from the new species. Fischer
& Crosse (1893) related the observation by
Sallé that H. merdigera covers its shells with
own excrement. It is interesting to note that
the fragmented paralectotype shows traces of
exactly such encrustation on the shell pieces.
If this behavior turns out to be typical for H.
merdigera, it is also in contradiction to the
behavior of the Costa Rican specimens which
were never observed to have agglutinated
anything on their shiny shells. Considering the
differences in the type material, it seems more
appropriate to treat H. merdigera as specifi-
cally distinct from H. fragilis until more mate-
rial is carefully studied, but this is beyond the
focus of the current study.

Von Martens (1890), having compared
Shuttleworth’s typical specimen of Helicina
elata to a Guatemalan sample, only recog-
nized the smaller size and a more slender
peristome. Besides size the syntype at hand
differs in a more globose shell, that is, a less
elevated spire with a blunt apex, the lower
whorls increasing less in diameter. In this re-
spect, it also differs from H. monteverdensis n.
sp. The figure in Fischer & Crosse (1893)
does not match the syntype at all. As de-
scribed by Shuttleworth and pointed out by
Von Martens (1890), H. elata exhibits a denti-
form prominence at the base of the columella
(like H. merdigera), which is lacking in the new
species. Whether or not H. e/ata has to be
treated as a subspecies of H. fragilis is a
question beyond the scope of this study.

Type material of Helicina mohriana could not
be located. L. Pfeiffer probably kept it in his
own collection, which became part of the col-
lection of Dohrn (Dance, 1986). The latter is
said to have been destroyed in the Museum
Stettin, Poland, during World War II (Clench &
Jacobson, 1971). Wagner (1908) depicted the
species for the first time (Alcadia (Leialcadia)
fragilis mohriana), but the source of his mate-
rial is unknown, and the one of two specimens
(Wagner, 1908: pl. 14, figs. 14-16) originating
from the type locality, Orizaba, represent a not
yet fully grown shell. Because he does not
give any explanation for this identification and
completely ignored “merdigera” and “elata”, it
does not contribute to the clarification of the
taxon. Thus, it obviously remains a dubious
species, which however is differentiated from
H. monteverdensis n. sp. by a groove in the

348 RICHLING

umbilical area near the columella mentioned in
the original description (“juxta columellam
brevem excavatus”).

Another dubious species (von Martens,
1891) of this complex is Helicina diaphana
from Honduras. It has only been depicted very
inadequately in Reeve (1874), and it com-
pletely escaped the attention of Fischer 8
Crosse (1880-1902) and Wagner (1907-
1911). Rehder (1966) stressed the specific
dissimilarity to Helicina Боисои!! Crosse &
Fischer, 1869, rendering it more likely to a
closer affinity with the “small, thin, fragile,
whitish species”. The study of the probable
syntype shows, on one hand, that H. diaphana
is clearly different from H. monteverdensis n.
sp., but, on the other hand, that the taxon
does not deserve to be treated as a dubious
species, because the description was not
based on a juvenile shell. H. diaphana 1$
broader than high, appearing depressed, but
the periphery is only very roundly angulated, if
at all. The suture is very weakly impressed,
the apex blunt. The uppermost Y4 whorls of the
teleoconch bear widely spaced spiral ridges,
subsequently the shell is sculptured with ir-
regular growth lines and an ornamentation of
small oblique grooves. The aperture is ob-
lique, inserting below the periphery, the outer
lip developed, but only slightly expanded and
reflexed. The transition to the columella lacks
a notch or denticle.

Considering also H. chiquitica and H.
escondida n. sp., it appears that the Mexican
and Guatemalan taxa discussed are not distrib-
uted as far southwards as previously assumed.

The original material of the records of
Monge-Nájera (1997) for H. oweniana and H.
beatrix was checked in the collection of INBio
and can partially be attributed to H.
monteverdensis n. sp. For differences to these
species, compare also the closely related H.
gemma.

Helicina (“Gemma”) escondida
Richling, n. sp.

Type Material

Holotype: INBio 3542623, female (leg. |.
Richling, 12.03.2001, ex IR 1543)

Paratype 1: INBio 3542624, male (same data
as holotype)

Paratype 2: ZMB 103880, female (same data
as holotype)

Paratype 3: ZMB 103881, female (same data
as holotype)

Dimensions:

Holotype: 6.2/6.0/6.4/5.4/3.8/4.9/5.0 mm
Paratype 1: 5.9/5.6/6.1/5.1/3.6/4.6/4.5 mm
Paratype 2: 6.8/6.1/6.6/5.7/3.8/5.1/5.3 mm
Paratype 3: 6.6/5.9/6.4/5.5/3.7/4.9/5.3 mm

Type Locality

SE-Costa Rica, Limón Province, approxi-
mately 9 km W of Matina, a little upstream on
the Rio Barbilla from the crossing of the road
from Siquirres to Limon, along a tributary of
Rio Barbilla, 10°03’29"N, 83°22’24"W, 70 т
a.s.l., valley of small creek in rain forest (prob-
ably secondary forest)

Material Examined

Lee. |. RICHLING

Heredia: S Puerto Viejo de Sarapiqui, Zona
Protectora La Selva, near OTS-Station,
about 10°25'53"М, 84°00'18"W, 60 m a.s.l.,
05.09.1999: (IR 1056)

Limon: About 9 km W of Matina, road Limon
to Siquirres, a little stream up the Rio
Barbilla, along a tributary of Rio Barbilla, in
the valley of a small creek in rain forest,
10°03'29"М, .63°22'24"W,. 70’ masilla
12.0320012(IR 1543)
№ Shiroles, Cerro Mirador, along trail,
09°36’37"М, 82°57’43"М/, 430 т a.s.l.:
16.03.2001: (IR 1601)

INBio COLLECTION

Limón: Sector Hitoy Cerere: Sendero
Bobócara, 09°40’31"М, 83°00’31"W, 200 т
a.s.l., leg. malacological staff of INBio,
10.01.1993: 1 ad. (INBio 1466441); 400 т
NE de la Estación de Hitoy Cerere, Sendero
la “Finca”, 09”40'36"N, 83°01'26"W, 110 т
a.s.l., leg. Alexander Alvarado Mendez,
27.09.2000: 1 ad. (INBio 3091794)
Reserva Biológica Hitoy Cerere: Cruce entre
Sendero Revienta Pechos y Sendero
Espavel, 09°39’12"М, 83*00'58"W, 600 т
a.s.l.: leg. Alexander Alvarado Mendez,
24.04.1999: 5 ads. (INBio 1497850);
Sendero Bobócara: 09°40'02"N,
83°02’42"W, 500 m a.s.l., 12.06.1999: 1 ad.,
1 s.ad. (INBio 3091132); 09°40’53"М,
83*04'09"W, 798 т a.s.l., 17.06.1999: 6
ads., 1 s.ad., 1 juv. (INBio 3542523) (all leg.
Alexander Alvarado Mendez)
Reserva Indigena Tayni, Sendero Bobocara,
09°40’28"М, 83°02'12"W, 200 m a.s.l., leg.
Alexander Alvarado Mendez, 15.07.1999: 1
ad. (INBio 1498244)

CLASSIFICATION OF HELICINIDAE 349

OTHER SOURCES

COSTA RICA

Limon: Los Diamantas Farm [about 10%11'N,
83°37 Wi], 11.08.1971: 1 ad. (UF 69847);
determination uncertain: Los Diamantes
Farm, 12 mi SE Guapiles [about 10%11'N,
83°37'W], leg. R.W. McDiarmid, 13.08.1971:
1 $.а4.: (UR¿217530)
Road cut, along $ bank of Rio Вапапо, ор-
posite La Bomba, 09°54'49.7"М,
83°03'56.4"W, leg. О.С. Robinson & J.M.
Montoya (Stn. 98 CR-15), 21.09.1998: 2
ads. (APHIS PPQ USDA)

Cartago:

Talamanca, Кю Estrella [about 09°43’N,
83°00’W], leg. Pittier (ZMB 103249); Valle
del Río Estrella, Talamanca [about 09°43’N,
83°00'W], leg. H. Pittier, 111.95 (ZMB 48235)
Valleé de Brabri, Talamanca [Bratsi? about
09°33'40"N, 82°53'28"W], leg. H. Pittier,
V111.98 (ZMB 103248)

Turrialba [about 09°54’30"N,
83°41 W], International American Agricul-
tural Institute, 2000 ft., leg. F.G. Thompson
(FGT-76), 03.08.1963: 2 ads. (UF 214143)

Costa Rica, without locality specified: ex Sowerby

& Fulton: 1 ad. (UF 243507: 1 of 3 spec.)

FIGS. 210-212. Helicina escondida n. sp. FIG. 210. Holotype, INBio 3542623, height 6.2 mm. FIG.
211. Paratype 1, INBio 3542624, height 5.9 mm. FIG. 212. Paratype 2, ZMB 103880, height 6.8 mm;
scale bar 2.5 mm.

350 RICHLING

Etymology

The name represents two aspects of the
species: for a long time it has escaped scien-
tific recognition, its occurrence is very “hidden”
in natural environments, and the small size
and variations of the color render it difficult to
find. | had been searching all day without suc-
cess until | came across some tiny helicinids,
which turned out to represent two new spe-
cies: chiquitica and “escondida” (Spanish) =
“hidden”. The Spanish word is preferred here
in homage to its origin and because the Latin
translations are occupied by other helicinid
species.

Description

Shell (Figs. 210-212, 336К-М): conical,
small, fragile and slightly dull. Color:
unicolored yellow except for the outer lip and
a slight, very thin yellowish-white band di-
rectly under suture. Embryonic shell about 1
whorl; 4'/, (37/,-4°/,) subsequent whorls
equally extending in size; last whorl rounded
at periphery; only slightly convex, giving the
spire a very regular and straight appear-
ance. Periostracum thin, under magnifica-
tion with very fine equally spaced spiral
striations at the periphery (up to about 9
lines) and a texture of fine oblique lines that
makes it appear dull. Aperture oblique and
very straight. Outer lip independently from
color of whorls always yellowish-white,
slightly thickened and reflexed nearly rectan-
gularly to the whorls; transition to columella
only with a very little denticle. Basal callus
very weakly developed, umbilical area finely
granulated with a little groove parallel to the
columella.

FIG. 213. Axial cleft and muscle attachments of
Helicina escondida n. sp., INBio 3542623; scale
bar 2.5 mm.

Internal Shell Structures (Fig. 213):

Teleoconch Surface Structure: The transitional
pattern (Fig. 214A, В) stretches for nearly %
of a whorl, the following structure of oblique
diverging grooves is only weakly developed,
becoming still weaker during growth (Fig.
214C, D), but not disappearing. Further-
more, the surface is sculptured with
periostracal spiral ridges that begin in the
second whorl.

Embryonic Shell (Fig. 215): Except for size,
the embryonic shell closely resembles that
of Helicina funcki. In the specimens under
study, it shows only minor deviations, for
example, the lines of pits starting somewhat
later.

Diameter: 728 um (+ 25) (680-780) (п = 19)
(IR 1543).

Operculum (Fig. 216): Thin and only slightly
calcified. Color yellowish and transparent.
Columellar margin slightly S-shaped, upper
end acute, lower end only weakly angulated,
rounded. Inner surface with a little ridge par-
allel to the columellar margin.

Animal (Figs. 338H, 339A-B): The color of the
soft body is variable. The foot is light whitish-
yellow and becoming brownish-grey up-
wards. Dorsal anterior and posterior end and
tentacles being darkest, but some individu-
als are much lighter than other. The mantle
color varies from a unicolored greenish to a
unicolored dark greyish-brown. Some indi-
viduals exhibit a very distinct dark or light
band with irregular margins at about the pe-
riphery, occasionally the apical part of the
mantle is spotted yellowish.

Radula (Fig. 217): The B-central bears 3-4
well-defined cusps; A- and C-central are
smooth or crenulate. The comb-lateral re-
markably differs from other Costa Rican spe-
cies in its consistently low number of only 6
cusps. Furthermore, the cusps increase con-
siderably in size inwards. The number of
denticles on the marginals increases rapidly.
Radula with about 69-78 rows of teeth.

Female Reproductive System (Fig. 218): The
receptaculum seminis is very small and
spherical. The bursa copulatrix bears a few
regular lobes; in one specimen, it is only bi-

CLASSIFICATION OF HELICINIDAE Sol

FIG. 214. Teleoconch surface structure of Helicina escondida n. sp. A. Embryonic shell and transition
of different sections. В. Immediately after embryonic shell: transitional surface structure. С. 2" whorl.
D. 3" whorl; scale bars 500 um (A), 100 um (B-D).

392 RICHLING

FIG. 215. Embryonic shell of Helicina escondida
n. sp.; scale bar 100 um.

lobed. The provaginal sac is elongated and
its stout stalk joins the sac about the middle
of the long side.

Morphometry and Sexual Dimorphism (Table
12, Figs. 219-223)

The sexes of all specimens from the three
areas were determined. Material from INBio
allowed the assignment without preparation
because of the quite transparent shells.

Gas — rc

FIG. 216. Operculum of Helicina escondida п. sp.,
ZMB 103880; scale bar 1 mm.

ip [22

FIG. 217. Radula of Helicina escondida п. sp. A.
Centrals. B. Comb-lateral. C. Marginals; scale
bar 50 um.

CLASSIFICATION OF HELICINIDAE 353

\
u À
LOROS
SE

VA
paa!
TE
x у Y 2
LT RON
а N
N
ma
\ ee")

FIG. 218. Female reproductive system of
Helicina escondida n. sp., ZMB 103880; scale
bar 1 mm.

Rio Barbilla (n=8/3)

A+
C4

Hitoy Cerere (n=7/7)

Shiroles (n=6/3)

Rio Barbilla (n=8/3)

г >

Hitoy Cerere (n=7/7)

Shiroles (n=6/3)

1 1

6 [mm] 7

FIG. 221. Expansion of outer lip of different
populations of Helicina escondida n. sp. in Costa
Rica according to Table 12; for explanations see
Fig. 219.

Morphometry: The populations show only mi-
nor deviations among each other and for the
different measurements. Only the speci-
mens from Hitoy Cerere are a little more el-
evated (height and height of the columellar
axis).

Sexual Dimorphism: The data show a clear
distinction between the measurements for
sexes, with the males being much smaller
(Figs. 224-226). In interpolation from the
minor diameter, males have an average vol-
ume of about 74% that of the females, re-
sembling Helicina tenuis.

Rio Barbilla (n=8/3)

1 Г 1 L I 1

1
0 1 2 3 4 5 6 [mm] 7

FIG. 219. Shell height of different populations of
Helicina escondida n. sp. in Costa Rica accord-
ing to Table 12; on each line: mean value, stan-
dard deviation, absolute range; number of indi-
viduals given as “n = females/males”; upper line:
females, lower line: males; in between and
shaded: average of both for comparison with
populations of unknown sex.

а +0

Hitoy Сегеге (п=7/7)
=

Shiroles (n=6/3)

1 1
6 [mm] 7

FIG. 222. Height of last whorl of different
populations of Helicina escondida n. sp. in Costa
Rica according to Table 12; for explanations see
Fig. 219.

Rio Barbilla (n=8/3) i =
9 Rio Barbilla (n=8/3) 9
> or y o
Hitoy Cerere (n=7/7) Hitoy Cerere (n=7/7)
Shiroles (n=6/3) = Shiroles (n=6/3)
| >
Па 1 1 1 i. 1 1 L 1 1 fe 1 if 1
0 1 2 3 4 5 6 [mm] 7 0 1 2 3 4 5 6 [тт] 7

FIG. 220. Minor diameter of shell of different
populations of Helicina escondida n. sp. in Costa
Rica according to Table 12; for explanations see
Fig: 219.

FIG. 223. Height of columellar axis of different
populations of Helicina escondida n. sp. in Costa
Rica according to Table 12; for explanations see
Fig:.2 19:

354 RICHLING

|

female +
nié male +
diam. + 7
[mm]

4 4.5 5 5.5 6 6.5 7 height[mm] 8

FIG. 224. Range of measurements in females and males of Helicina
escondida n. sp. exemplary for height and minor diameter in the population
from the Rio Barbilla.

female +
ie male +
diam. +
[mm]

4.5

4 4.5 5 5.5 6 6.5 7 height [mm] 8

FIG. 225. Range of measurements in females and males of Helicina
escondida n. sp. exemplary for height and minor diameter in the population
from the Shiroles.

CLASSIFICATION ОЕ HELICINIDAE

TA = А — 1 ЕТ: IT: т И:
female >
2 |
min. mais
diam. r -
[mm]
6 | =
o
5.5 | ER 5]
o
o
+ o
5 L \ -
ге
+
р
4.5 | 4
4 = — N "1 = ze NA J
4 4.5 5 5.5 6 6.5 7 height [mm] 8

FIG. 226. Range of measurements in females and males of Helicina
escondida n. sp. exemplary for height and minor diameter in the population
from the Hitoy Cerere.

86° | ‘185° —

f

3500 - 4000 m
3000 - 3500 m : : L Y
2500 - 3000 m
2000 - 2500 m
1500 - 2000 m
1000 - 1500 m
500 - 1000 m
100 - 500 т

e coll. IR
ia e coll. INBio

09°

© others

FIG. 227. Records of Helicina escondida n. sp. in Costa Rica.

355

356

RICHLING

TABLE 12. Measurements of different populations of Helicina escondida n. sp. given as mean value
with standard deviation, minimum and maximum value (min, max), and number of specimens (min./
max. diam. = minor/major diameter, col. axis = columellar axis); linear measurements [тт].

“Río Barbilla” (altitude 70 т)

“Hitoy Cerere” (altitude 110-798 т)
lots INBio 1466441, 1497850, 1498244,

lot IR 1543 3542523, 3091132, 3091794
Mean Mean
Sex value Deviation Min Max Number value Deviation Min Max Number
Height f 6.40 019 6.15 6.84 8 6.55 016 6.26 6.98 if
Height m 5.81 0.05 573 589 3 5.89 0.15 568 618 7
Mai. diam. f 5.90 0.10 5.76 6.09 8. 5.85 0.19 5.58 6.26 Г
Ма]. Чат. т 5.44 0.10) 15.32 95159 5 5.30 0.14 498 5.58 7
Min. diam. f 5.44 0.10’ 527 574 8 5.38 0.14 5.12 5175 7
Min. dam. m 4.98 0.08 486 5.08 3 4.85 0.13 460 50 7
Outer lip + 9 72 DION (358336 8 3.66 0.11 3.48 3.85 7
Outer lip т 3.56 0.04 3.52 3.62 3 3.43 0.10 3.28 3.60 7
Last whorl f 4.84 0.14 4.56 5.13 8 4.82 0.10 4.62 5.04 7
Last whorl m 4.52 0.08 4.44 4.64 3 4.28 0.10 412 457 7
Col. axis f 5.09 0.13 4.89 5.34 8 5.23 0.189. 487 552 7h
Col. axis m 4.49 006 444 4.58 3 4.67 0.13 4.47 5.03 fl
“Shiroles” (altitude 430 т)
lot IR 1601
Mean
Sex value Deviation Min Max Number

Height f 6.23 0.22 5.91 6.64 6

Height m 5.72 0.09 559 581 &

Maj. diam. f 5.95 008 5.86 618 6

Maj. Чат. т 538 009 524 5.5 €)

Min. diam. f 5.45 0.11 528 5.63 6

Min. diam. т 4.87 0.09 476 £5.01 3

Outer lip f 3.79 008 73167. 3.93 6

Outer lip m 3.56 008 3.45 3.66 3

Last whorl f 4.84 0.11 469 5.01 6

Last whorl m 4.47 0.10 432 4.58 3

Col. axis f 4.99 0.18 467 5эЭ7 6

Col. axis m 4.49 0.11 432 4.60 8
Habitat Distribution (Fig. 227)

Helicina escondida n. sp. is an arboreal spe-
cies that was found on the lower side, more
seldomly, the upper side, of small-leafed under-
growth plants. lt also was observed aestivating
on fronds of ferns. Near the Rio Barbilla, the
species was only found along a small creek
together with H. chiquitica. North of Shiroles
and probably in Hitoy Cerere, it occurs sympa-
trically with H. beatrix confusa and H. funcki.
There it was found on a ridge with forest cover.

The species occurs on the central and south-
ern Caribbean side of Costa Rica at some dis-
tance from the coast mainly in the slightly
elevated hilly countryside. The most northern
occurrence is from the northern foothills of the
Cordillera Central; to the south, Helicina
escondida n. sp. reaches the Valle de
Talamanca. Between the neighboring valleys
Valle de Estrella and Valle de Talamanca, the
species lives up to altitudes of about 800 m.

CLASSIFICATION OF HELICINIDAE 397

Because undisturbed areas in this region are still
relatively uninvestigated due to their inacces-
sible nature, it is very likely that H. escondida п.
sp. can be found at additional localities.

Discussion

Helicina escondida n. sp. was found in at
least three different colors within one popula-
tion (Figs. 210-212, 336K-M): (1) unicolored
yellow (represented in holotype, Fig. 336K),
(2) reddish-brown, except for the umbilical
area, which 15 whitish (represented by
paratype 1, Fig. 336L), or (3) the upper half of
each whorl is reddish brown, this may be very
light, and the lower half yellow or only with a
yellowish band, the transition of both colors
exactly at the suture/ periphery, so that the
yellow is only seen on last whorl (represented
by paratype 2, Fig. 336M). The ощег lip 1$
constantly whitish-yellowish.

Helicina escondida n. sp. is distinguished
from other small helicinids, such as H. beatrix,
H. talamancensis, H. monteverdensis n. sp.,
H. fragilis, and H. gemma, by the special sur-
face structure of fine oblique lines and spiral
striations. In the species mentioned, it is shiny
and smooth. Furthermore, the aperture 1$
straight and not curved backwards. Finally, the
groove in the umbilical area is unique for H.
escondida n. sp. among comparable species.
(For further discussion of small, fragile, whit-
ish-yellowish Central American Helicinidae,
see the section on H. monteverdensis n. sp.)
In H. beatrix, the whorls are much more con-
vex, and it has a higher spire and a distinct
whitish subsutural band. Helicina gemma has
a orange-scarlet outer lip, whereas in H.
escondida п. sp. it is yellowish-white. At its
type locality, H. chiquitica and H. escondida n.
sp. occur sympatrically, but they are easily
separated by their size, color, shape and shell
surface texture.

Helicina (“Gemma”) chiquitica
(Richling, 2001)

Alcadia (Leialcadia) fragilis — Wagner, 1908:
84-85: Costa Rica: Shirores, Talamanca [in
part] [non Morelet, 1851]

Oligyra chiquitica Richling, 2001: 1-2 (text fig-
ure)

Original Description

See “Description”.

Type Material

Holotype: INBio 3404977, female (leg. 1.
Richling, 12.3.2001)

Paratype 1: INBio 3404981, male (same data
as holotype)

Paratype 2: ZMB 103386a, female (same data
as holotype)

Paratype 3: ZMB 103386b, male (same data
as holotype)

Dimensions (height/greatest diameter):

Holotype: 4.9/4.5 mm

Paratype 1: 4.3/4.2 mm

Paratype 2: 4.6/4.4 mm

Paratype 3: 4.3/4.1 mm

Type Locality

SE-Costa Rica, Limón Province, approxi-
mately 9 km W of Matina, a little upstream on
the Río Barbilla from the crossing of the road
from Siquirres to Limón, along a tributary of
Río Barbilla, 10%03'29"N, 83°22'24"W, 70 т
a.s.l., in the valley of a small creek in rain for-
est (probably secondary forest).

Examined Material

Lec. |. RICHLING

Heredia: S Puerto Viejo de Sarapiqui, Zona
Protectora La Selva, near OTS-Station,
about 10%25'53"N, 84°00'18"W, 60 m a.s.l.,
05.09.1999: (IR 1662)

Limon: About 9 km W of Matina, road from

Limon to Siquirres, a little stream up the Rio
Barbilla, along a tributary of Rio Barbilla, in
the valley of a small creek in rain forest,
10°03 29"N, 832224“, 70 т аз,
12.03.2001: (IR 1539)
S Siquirres, road from Limön to Siquirres,
along footpath stream up Rio Pacuarito, in
the valley of a little northern tributary,
10°0538.N, 83°%2811"W, 110 m a.sil.,
18.03.2001: (IR 1611)

INBio COLLECTION

Cartago: Parque Nacional Barbilla, Sector de la
Estación de Barbilla, leg. Alexander Alvarado
Mendez: 09°58’26"N, 83°27 58"W, 500 т
a.s.l., 28.09.2000: 1 ad. (INBio 3100273);
0958'24"N, 83%27'23"W, 300 т asl.
30.09.2000: 1 ad. (INBio 3104239)
Zona Protectora Rio Pacuare, Sector de la
Estación de Barbilla, 09°58’50"N, 83°27 08"W,
500 т а.$.1., leg. Alexander Alvarado Mendez,
05.09.2000: 1 ad. (INBio 3103323)

358 RICHLING

FIG. 228. Helicina chiquitica, holotype, INBio 3404977, height 4.9 mm; scale bar 2 mm.

OTHER SOURCES

COSTA RICA

Limón: Shirores [Shiroles, 09°35’38"М,
82°57'20"W], Talamanca: leg. H. Pittier
(#269): 1 s.ad. (МНММ, part of the lot); leg.
H. Pittier (#208), 03.1895: 1 ad., 2 s.ads.
(ZMB 48336, part of the lot); leg. Pittier: 1
ad. (ZMB 103250)

Etymology

The species is named for its small size.
“Chiquitica” is a diminutive of “chiquita” (Span-
ish) = “small”. In Latin America it correctly
would be “chiquitita”, but in Costa Rica the di-
minutive syllable “-tito/a” sometimes is
changed to “-tico/a”; the Costa Rican people
call themselves also Ticos.

Description

Shell (Figs. 228, 336N): conical-globose, thin,
small, shiny. Color: unicolored, reddish-

FIG. 229. Axial cleft and muscle attachments of Heli-
cina chiquitica, INBio 3404977; scale bar 2 mm.

brown, more or less transparent, towards
aperture color more intensive. Periostracum
very thin, shiny and smooth, except for very
fine growth lines. Embryonic shell of about 1
whorl; subsequent 3*/, to 4 (holotype: 3%)
whorls slightly convex, equally extending in
size; last whorl rounded at periphery. Suture
slightly impressed. Aperture oblique and in
its middle part slightly curved backwards.
Outer lip dark red, thickened, very narrowly
reflexed. Basal callus thin, in umbilical area
surface scaly-granulated.

Internal Shell Structures (Fig. 229):

Teleoconch Surface Structure (Fig. 230): The
transitional pattern is well developed (about
Y of a whorl), the subsequent zone with
oblique diverging grooves is short. The rest
of the teleoconch is smooth, except for fine
growth lines.

Embryonic Shell (Fig. 231): The structure
does not exhibit peculiarities and closely re-
sembles that of Helicina gemma. Compared
with the shell size of the latter, the diameter
of the embryonic shell of H. chiquitica is only
slightly smaller. The embryonic shell of the
clearly larger H. escondida n. sp. is on aver-
age even smaller (728 pm) than that of H.
chiquitica. This shows that embryonic shell
size does not always depend on the shell
size (see general discussion below).
Diameter: 749 pm (+ 28) (660-800) (п = 25)
(IR 1539).

Operculum (Fig. 232): Very slightly calcified,
calcareous plate not reaching the margin.

CLASSIFICATION OF HELICINIDAE 359

FIGS. 230, 231. Shell structure of Helicina chiquitica. FIG. 230 Teleoconch surface structure. A.
Structure of apical part. В. 2" whorl. С. 1% whorl: occasional sharp transition from oblique diverging

grooves to smooth surface; scale bars 500 um (A), 100 um (B-C). FIG. 231. Embryonic shell; scale
bar 100 um.

360 RICHLING

FIG. 232. Operculum of Helicina chiquitica,
holotype, INBio 3404977; scale bar 1 mm.

Color horny-amber, slightly transparent.
Columellar side S-shaped, upper end acute,
lower edge rounded. On inner side, a little
ridge parallel to columellar margin. Outer
surface granulated.

Animal (Figs. 339C, D): The foot-head region
Shows a similar color to other species in
being greyish-black on the dorsal half includ-
ing the tentacles. The mantle pigmentation is
black throughout or mottled with small pale
dots mainly in the apical part. The yellow-
shelled form is also much paler in the body
color.

Radula (Fig. 233): All centrals lack well-de-
fined cusps and the faces are less pro-
nounced. Comb-lateral with 10-12 pointed
cusps, a high number among the Costa
Rican species. Cusps on marginals rapidly
increasing in number. Radula with about 65—
72 rows of teeth.

Female Reproductive System (Fig. 234): The
ascending limb of the V-organ is very promi-
nent and stout, the receptaculum seminis is
rather small and oblong. The bursa
copulatrix is formed by an irregular ovoid
sac, which appears to be internally subdi-
vided. The provaginal sac is simple and con-
nected by a short duct. The pallial oviduct is
relatively short.

Morphometry and Sexual Dimorphism (Table
13, Figs. 235-239)

FIG. 233. Radula of Helicina chiquitica. A.
Helicina chiquitica is the smallest arboreal Centrals. B. Comb-lateral. C. Marginals; scale
species investigated in this study. The few bar 50 um.

CLASSIFICATION OF HELICINIDAE 361

FIG. 234. Female reproductive system of
Helicina chiquitica, IR 1539; scale bar 1 mm.

| Rio Pacuarito (n=2/2)
a

| Rio Barbilla (n=6/10)

Q, ю

+

6 [mm] 7

|
| un
0 1 2 3 4 5

FIG. 235. Shell height of the two populations of
Helicina chiquitica in Costa Rica according to
Table 13; on each line: mean value, standard
deviation, absolute range; number of individuals
given as “п = females/males”; upper line:
females, lower line: males; in between and
shaded: average of both for comparison with
populations of unknown sex.

Rio Pacuarito (n=2/2)

Rio Pacuarito (n=2/2) +
+ E
Río Barbilla (n=6/10)

L 1 1 Г i | 1

6 [mm] 7

FIG. 237. Expansion of outer lip of the two
populations of Helicina chiquitica in Costa Rica
according to Table 13; for explanations see Fig.
2.33.

specimens of H. chiquitica from two different
localities suggest a smaller shell size for the
site “Rio Pacuarito”, but the sample size is
small.

On the other hand, sexual dimorphism is
undoubtedly indicated. The clear distinction
between the sexes is also illustrated at “Rio
Barbilla” (Fig. 240). In the interpolation from
the minor diameter, males have an average
volume of about 65% of that of females.

Habitat

The species has mainly been found on the
lower side of leaves of Araceae, occasionally
also on the leaves of bushy plants of the un-
dergrowth. At Rio Barbilla and Rio Pacuarito,

Rio Pacuarito (n=2/2
Е + 9
м
Rio Barbilla (n=6/10)
4 1 1 1 1 | 1
0 1 2 3 4 5 6 [mm] 7

FIG. 238. Height of last whorl of the two
populations of Helicina chiquitica in Costa Rica
according to Table 13; for explanations see Fig.
235.

| Rio Pacuarito (n=2/2)

+ Q + 9
(of = o
Rio Barbilla (n=6/10 Rio Barbilla (n=6/10
| ( ) Be | )
| 2
JE L 1 it 4 1 1 | т ГВ 1 1 1 1 1
0 1 2 3 4 5 6 [mm] 7 0 1 2 3 4 5 6 [mm] 7

FIG. 236. Minor diameter of shell of the two
populations of Helicina chiquitica in Costa Rica
according to Table 13; for explanations see Fig.
235.

FIG. 239. Height of columellar axis of the two
populations of Helicina chiquitica in Costa Rica
according to Table 13; for explanations see Fig.
235.

362 RICHLING

5 T T
| female >
min.

male +

3:5. -

3 3.5 4 4.5 5 height [mm] 6

FIG. 240. Range of measurements in females and males of Helicina chiquitica
exemplary for height and minor diameter in the population from the Rio Barbilla.

[86°

3500 -
3000 - 3500 m
2500 - 3000 m |
| 2000 - 2500 m |
| 1500 - 2000 т |
| 1000-1800 | | | RP el
500 - 1000 m |
100 - 500 m |

e coll. IR |
0-100m e coll. INBio |

© others |

FIG. 241. Records of Helicina chiquitica in Costa Rica.

CLASSIFICATION OF HELICINIDAE

363

TABLE 13. Measurements of the two populations of Helicina chiquitica given as mean value with standard
deviation, minimum and maximum0 value (min, max), and number of specimens (min./max. diam. = minor/
major diameter, col. axis = columellar axis); linear measurements [mm], weight [g], volume [ml].

“Río Barbilla” (altitude 70 m)

lot IR 1539
Mean

Sex value Deviation Min Max
Height f 4.87 0.23 456 526
Height m 420 0.06 4.04 4.31
Maj. Чат. + 4.48 013 433 4568
Ма). Чат. m 3.97 O10) 3.73 “416
Min. diam. f 4.16 0.16 398 4.43
Min. dam. m 3.63 0:07 343 3:75
Outer lip f 2.82 0.13 2.64 3.04
Outer lip m 2.49 0.07 227 2.61
Last whorl f 3.61 0.14 3.38 3.84
Last whorl т 3.10 0.05 2.94 3.20
Col. axis f 3.88 0.18 3.60 4.17
Col. axis m 3.32 0.07 3.18 3.44

the occurrence seems to Бе confined to the
vegetation along small, partly steep creeks. At
the former locality, Helicina chiquitica lives
sympatrically with H. escondida n. sp.

Distribution (Fig. 241)

Except for the single record northeast of
the Central Cordiilera, the few records are in
the most northern part of the Caribbean foot-
hills of the Cordillera de Talamanca. Helicina
chiquitica has only been found some dis-
tance from the coast in the hilly countryside
from elevations of 70 m to 500 m. The undis-
turbed areas in this region have scarcely
been investigated, because they are difficult
to reach. The northern record renders it also
very likely that H. chiquitica will be found at
additional localities.

Furthermore, the small size and the rapid
decay of shells in a humid tropical climate pro-
vide grounds for assuming that the species
has a wider distribution than actually been
documented. А study by Barrientos (2000) for
Ovachlamys fulgens (Gude, 1900) suggests
that shells decay in less than five months in
the climate of San José with 5-6 dry months
per year as opposed to the Caribbean slope,
where, lacking dry months, it would take place
even faster.

“Río Pacuarito” (altitude 110 т)

lot IR 1611
Mean
Number value Deviation Min Max Number
6 4.49 0.12 4.37 4.60 2
10 3.99 0.21 3.78 4.20 2
6 4.33 0.06 4.26 4.39 2
10 3-72 0.13 3.59 3.84 2
6 4.01 0.09 3.92 4.09 2
10 3.44 0.09; 3.35 3.53 2
6 2118 0:04” 7274. 72.82 2
10 2.42 0.04 238 2.45 2
6 3.54 0.07 347 3.61 2
10 3.01 0.13288 313 2
6 3.64 0:10. 3.54 3.74 2
10 3.15 017 2:98 39 2
Discussion

In some specimens, the apex is reddish-
brown, but the subsequent whorls are yellow.
In this case the outer lip is yellow too.

In its shape and shell sculpture, Helicina
chiquitica is comparable to H. gemma, Н.
monteverdensis n. sp., and H. fragilis, but H.
chiquitica is much smaller and has a different
color. None of the two species mentioned
shows such a dark color in combination with a
dark outer lip. The size of the new species 1$
exceptionally small for the known Central
American helicinids of this shape. Helicina
strebeli L. Pfeiffer, 1861, is another small
helicinid from Mexico, generally treated as a
small subspecies or variety of Helicina flavida
(see von Martens, 1890; Fischer 8 Crosse,
1893; Baker, 1928), which clearly differs by its
spiral striation. Helicina mohriana L. Pfeiffer,
1861, described from Orizaba, Mexico, 1$ dis-
cussed as a dubious species or perhaps juve-
nile stage by Martens (1891) and Fischer 8
Crosse (1893) or as a synonym of Helicina
fragilis merdigera by Baker (1922a) respec-
tively. According to the original description,
however, it is a little broader than it is high,
whereas in H. chiquitica all specimens show
the reverse relation. Furthermore H. mohriana
seems to have more whorls (5.5) than the new

RICHLING

pecies, for which 5 (4 plus about 1 of embry-
onic shell) whorls are exceptional.

The record of Wagner (1908) for Shiroles in
Costa Rica had been based on the material in
the ZMB, since the specimens carried deter-
minations written by Wagner. Reexamination
of the material revealed specimens of H.
chiquitica, but the lot (and a similar lot of Pittier
stored in the MHNN) consists of two different
species, where, unfortunately, all deviating
specimens are immature. They exhibit very
strong spiral cords, but do not seem to repre-
sent H. escondida n. sp. which has recently
been found at Shiroles, whereas H. chiquitica
has not as yet been discovered there.

Pyrgodomus microdinus
(Morelet, 1851)

Helicina microdina Morelet, 1851: 18 (not fig-
ured)

Helicina microdina — L. Pfeiffer, 1852a: 354

Helicina microdina — L. Pfeiffer, 1852b: 256

Helicina chryseis Tristram, 1862: 5: Guate-
mala: mountain forests of Vera Paz (Salvin)
(not figured)

Helicina microdina — Bland, 1866: 8

Helicina chryseis — Bland, 1866: 10

Helicina chryseis — von Martens, 1890: 39, pl.
|, fig. 14

Helicina microdina — von Martens, 1891: 42
(dubious species)

Helicina (Pyrgodomus) chryseis — Fischer 8
Crosse, 1893: 440, pl. LVII, fig. 6

Нейста (Idesa) microdina — Fischer 8
Crosse, 1893: 438-439, pl. LVI, fig. 9

Helicina chryseis — von Martens, 1900: 606

Eutrochatella (Artecallossa [sic]) microdina —
Wagner, 1908: 138-139, pl. 20, figs. 17-20

Eutrochatella microdina chryseis — Pilsbry,
1920b: 197: Guatemala: Chama

Eutrochatella nicrodina [sic] var. chryseis —
Hinkley, 1920: 52: Guatemala: Alta Verapaz:
Chama between Río Tsalbha and Río Negro
Chama: also in river drift

Eutrochatella (Pyrgodomus) microdina
microdina — Baker, 1922a: 61
Eutrochatella (Pyrgodomus) microdina

chryseis — Baker, 1922a: 61 (may be a sex-
form)

Pyrgodomus microdinus chryseis — Baker,
1928: 45-46

Pyrgodomus microdinus microdinus — Baker,
1928: 45-46

Pyrgodomus microdina — Goodrich & van der
Schalie, 1937: 13, 33: Guatemala: Petén:
region of headwater of Río San Pedro de
Mártir, lower Río de la Pasión; Alta Verapaz:
upper part of Río de la Pasión

Pyrgodomus ? spec. — Monge-Nájera, 1997:
113: Costa Rica

Synonymy
Helicina chryseis Tristram, 1862
Original Description

“T. parvula, conica, transversim minute
striata, spiraliter lirata, flava, sursum
saturatior. Anfr. 6 convexi, ultimo angulato;
columella arcuata, superne callosa,
subdilatata. Apertura obliqua, ovalis, margine
simplici, recto.

FIG. 242. Helicina microdina, lectotype, BMNH 1893.2.3.1986, height 3.8 mm; scale bar 1 mm.

CLASSIFICATION OF HELICINIDAE 365

АНИ. 4. — Diam. 4.
H. vulgaris prov. Vera-Paz.

Primo aspectu H. rupestris Pfr. congruit,
quamvis in universum ab ea dissimilis.”

Type Material

BMNH 1893.2.3.1986-90: Morelet coll., pur-
chased from H. Fulton
The Morelet collection was bought by H. Fulton
and later purchased by the BMNH. Fischer 8
Crosse (1893) studied the original material in
Morelet collection and figured one shell which
can be identified by the mark of a “x”. This shell
is here selected as lectotype of Helicina
microdina (BMNH 1893.2.3.1986) (Fig. 242),
because it is uncertain whether Fischer 8
Crosse's comment in the figure caption (pl. LVI,
figs. 9, Эа, 9b, 9c: “Туре de ГН. microdina”) can
be regarded as a type selection.
Dimensions (height/greatest diameter/minor
diameter):
Lectotype BMNH 1893.2.3.1986: 3.8/3.9/3.5 mm

Type Locality

“Vera-Paz” [Guatemala, Alta Verapaz De-
partment].

Examined Material

Lec. |. RICHLING

Puntarenas: N Neily, road from Ciudad Neily
to San Vito, open area with a few trees,
08°40’23"М, 82°56 44"W, 180 m a.s.l., N
Neily, 23.03.1997: (IR 209)

Fila de Cal, road from Ciudad Neily to San
Vito: S Campo Dos, burned area and ground
with secondary growth and limestone rocks,
08°41’00"М, 82°56'29"W 630 m a.s.l.:
23.03.1997: (IR 192); 07.03.1998: (IR.502);
09.02.2000: (IR 1147); Campo Dos, on
Finca, secondary growth and limestone
rocks, 08°41’16"М, 82%56'38"W, 700 m a.s.l.:
07.03.2001: (IR: 1517)

INBio COLLECTION

Puntarenas: Fila Cal, 24 km de San Vito hacia
Ciudad Neilly, 08°41’36"М, 82*56'36"W, 780
т a.s.l., 14.01.1995: leg. Francisco Alvarado:
1 spec. (INBio 1480755); 1 spec. (INBio
1495177); leg. Annia Picado: 2 spec. (INBio
1481148); leg. Mario Chinchilla: 1 spec.
(INBio 1481257); leg. Socorro Avila: 3 spec.
(INBio 1481361); leg. Ronald Villalobos: 6
spec. (INBio 1481512); leg. Marcos Moraga:
4 spec. (INBio 1481563); leg. Oscar Esquivel:
5 spec. (INBio 1485112); leg. Marcos Madri-
gal: 2 spec. (INBio 1495685); leg. Enia
Navarro: 1 spec. (INBio 1495695);
29.08.1995: leg. Marianella Segura: 11 spec.
(INBio 3307036); 24.5 Km S en la carretera
de San Vito hacia Ciudad Neilly, 08°40’55"N,
82°56'23"W, 600 m a.s.l.: leg. Zaidett
Barrientos, 21.11.1995: 1 spec. (INBio
1485119)

Description
Shell (Figs. 243, 336R): Conical, high-el-

evated, triangular in general shape, solid,
small sized, dull. Color: bright yellow and on

FIG. 243. Pyrgodomus microdinus, Fila de Cal, IR 1517, height 3.8 mm; scale bar 1 mm.

RICHLING

366

245. Shell structure of Pyrgodomus microdinus. FIG. 244. Teleoconch surface structure.

,

FIGS. 244

scale

structure of postembryonic shell;

whorl,

FIG. 245. Embryonic shell; scale bar 100 um.

transitional structure. С. 1°

whorl,
bar 500 um (A); 100 um (В-С).

st

A. apical part. B. 1

CLASSIFICATION OF HELICINIDAE 367

upper side darker colored. Whorls sculp-
tured with spiral ridges, very rough and ir-
regular growth lines and oblique striations.
Embryonic shell with about 1 whorl; about
4% subsequent whorls straight; last whorl
angulated on periphery. Whorls equally in-
creasing in size and rapidly descending, al-
ways inserting a little below periphery,
forming a high, pointed, stepped spire. Su-
ture deeply impressed. Aperture oblique and
rather straight. Outer lip of same color as
preceding whorls, neither remarkable thick-
ened nor shortly expanded or reflected. Col-
umella short and arched. Basal callus
weakly developed.

Internal Shell Structures: Sufficient adult ma-
terial was not available, especially because
the thickness of the whorls would have re-
quired cracking the shell in order to examine
the internal structures.

Teleoconch Surface Structure (Fig. 244): In
Pyrgodomus microdinus, a structured transi-
tional zone similar to that of the Costa Rican
species of Helicina is developed for a certain
distance at the very beginning of the
teleoconch. This zone exhibits a very rough
irregular surface of numerous small den-
ticles. The following whorls bear spiral ridges
crossed by irregular growth lines that are
wrinkled throughout.

Embryonic Shell (Fig. 245): Pyrgodomus
microdinus displays a greatly different em-
bryonic shell surface structure. Coarse ob-
lique diverging grooves cover the embryonic
shell resulting in a very rough surface.
Diameter: 530 рт (+ 13) (510-550) (п = 4)
(IR 192, IR 1517).

Operculum: Calcified portion strongly devel-
oped, horny plate very thin and slightly
larger, columellar side of calcareous plate
slightly convex and thickened. Color whitish
and only very slightly transparent. Nucleus
nearly in central position, and growth lines
almost concentric.

Animal (Fig. 339E): The whole snout and the
underside of the foot is whitish-yellowish, the
latter darkens gradually to grey towards the
upper side. The tentacles are of the same
grey, which becomes diffuse at the bases, FIG. 246. Radula of Pyrgodomus microdinus. A.
so that between the tentacles and behind Centrals. B. Comb-lateral. C. Marginals; scale
the eyes the head region is tinged with the bar 50 um.

36

lighter color of the snout. The mantle bears
dark patches, which shine through the shells
in live specimens and result in a greenish-
grey appearance of the animals.

Radula (Fig. 246): Since sufficient Costa
Rican material was not available, a speci-
men from Honduras (Colón Depto., lime-
stone hill, 3 km WSW La Brea, 100 m a.s.l.,
leg. F.G. Thompson et al. (FGT-5389),
10.03.1994 (UF 221175) was studied.
R-central quadrangular; A- and B-central
each bearing 2 sidewards projecting cusps;
C-central with 3 denticles. Comb-lateral
strong developed and T-shaped with minor
crenulations at the cutting edge. Inner
marginals unicuspid, outwards slowly in-
creasing number of slender, rather terminal
cusps. Radula with about 130 rows of teeth.
Baker (1928) studied the radula of the
smaller Mexican Pyrgodomus microdinus
abditus Baker, 1928, and described bicuspid
inner marginals, which necessitated a
change in the original definition of the sub-
family Vianinae with unicuspid inner
marginals. Furthermore, the centrals bear
each one cusp more, and the denticles on
the cutting edge of the T-lateral are more
pronounced. It remains questionable
whether these deviations are typical for the
Mexican subspecies alone, or whether the

FIG. 247. Female reproductive system of
Pyrgodomus microdinus abditus, reproduced
from Baker (1928: pl. IV, figs. 21-23).

368 RICHLING

single specimen investigated in this study is
representative at all or if they are subject to
individual variation. The increased number
of cusps in the smaller Mexican subspecies
could also be caused by the phenomenon
that smaller specimens/species of
Helicinidae independently from their phylo-
genetic position show a tendency to develop
more cusps (see general Discussion).

Female Reproductive System: In Costa Rica,

no adult live specimens could be found, only
some juveniles, therefore their anatomy
could not be investigated.
Baker (1928) (reproduced here in Fig. 247)
gave a description of the female reproduc-
tive system of Pyrgodomus microdinus
abditus: The apical part of the V-organ is
slightly elongated (apical swelling) and the
small spherical receptaculum seminis is situ-
ated near the middle of the dorsal side of the
descending limb (not visible in his figure).
The bursa copulatrix is formed by a long el-
lipsoid sac; the bigger, rounded triangular
provaginal sac exhibits coarse lobes on its
distal margin. Provaginal duct and vagina
are not explicitly mentioned or shown (Fig.
247, arrow), but they are believed to open
into the hypobranchial duct, which orifice lies
at about */, to '/, of the length of the pallial
oviduct. In the light of the newly discovered
absence of provaginal opening in the Costa
Rican species of Helicina, this question re-
mains open for P microdinus abditus pend-
ing further investigation. The close
relationship to Eutrochatella, which is indi-
cated by a similar embryonic shell structure,
suggests the absence of a provaginal open-
ing, because it is undeveloped in this genus
(see under Eutrochatella below).

Morphometry and Sexual Dimorphism

Pyrgodomus microdinus was not found in
sufficient numbers. In addition, most speci-
mens were not fully grown, making a morpho-
metric investigation impossible.

Habitat

Like related species (e.g., species of the
genus Eutrochatella), Pyrgodomus microdinus
lives on limestone rock faces and this obvious
dependence characterizes the species as
calciphile. Goodrich & van der Schalie (1937)

CLASSIFICATION ОЕ HELICINIDAE 369

reported the species from Guatemala as being
restricted to limestone outcrops. Under dry
weather conditions, it was found aestivating
on the underside of larger pieces of rocks or
on shaded vertical sides and crevices of
rocks.

Live specimens of P. microdinus are per-
fectly adapted to the background, because
they possess a camouflage. Small particles of
the surroundings are glued on the rough shell
surface. This behavior was also observed in
Guatemalan specimens by Goodrich 8 van
der Schalie (1937). Only dead specimens
show the bright yellow color.

Distribution

The Costa Rican occurrence seems to be
limited to the area of the Fila de Cal north of
Ciudad Neily (Fig. 248). In view of its ecologi-
cal requirements, this distribution clearly re-
flects geological conditions. п Costa Rica,
calcareous outcrops are only found in a very
few places, such as the Fila de Cal.

The type locality of P. microdinus 1$ Alta
Verapaz, Guatemala. lt has also been re-
corded from Péten in Guatemala and southern
Veracruz, Mexico (as the subspecies P.
microdinus abditus). For Belize, the different
species, P. simpsoni (Ancey, 1886), is men-
tioned by Haas & Solem (1960). The Costa
Rican populations seem to represent the most
southerly occurrence of P microdinus.

Discussion

Fischer 8 Crosse (1893) figured the species
for the first time on the basis of the original
Morelet material. They remarked on the similar-
ity to Pyrgodomus chryseis, with the difference
that the latter species is more elevated. How-
ever, they placed these species in different sec-
tions (/desa and Pyrgodomus). Wagner (1908)
proposed the synonymy of these taxa. Baker
(1928) morphometrically compared P
microdinus, P. chryseis and populations from
Veracruz, Mexico. Аз the result, he attributed
the taxa to subspecific rank and raised a new

[84° [83°

3500 - 4000 m
3000 - 3500 m ee

2500 - 3000 т
2000 - 2500 т
1500 - 2000 m
1000 - 1500 m
500 - 1000 m
100 - 500 т

В

— he = a — en 09°

|

0- 100 m

e coll. IR
e coll. INBio
o others

FIG. 248. Records of Pyrgodomus microdinus in Costa Rica.

370 RICHLING

subspecies for Mexican specimens, P.
microdinus abditus having a smaller size, fewer
whorls, and a more depressed shape. He
noted that subspecific recognition of Р
microdinus chryseis was uncertain and re-
quired study of a larger series. Ancey (1886)
described a quite similar species from Isla de
Utila off Honduras, P. simpsoni, which Baker
(1928) treated as specifically distinct.

The very few adult Costa Rican specimens
are remarkably elevated. Therefore, they
more closely resemble P. chryseis. Against
this historic background and because P.
chryseis and P. microdinus both typically origi-
nate from the same area in Guatemala, it 1$
more appropriate to regard them as synony-
mous until better knowledge of the distribution
becomes available, thereby treating the Costa
Rican specimens as P. microdinus. The geo-
graphical distance to the northern populations
is uncertain due to the lack of extensive inves-
tigations of the Nicaraguan malacofauna.

Alcadia (Microalcadia)
Richling, n. subgen.

Type Species
Helicina hojarasca Richling, 2001
Diagnosis

Shell very small, fragile, with fine spiral stria-
tions and rows of periostracal hairs; outer lip of
adults not differentiated from the whorl. Cal-
careous layer of the operculum very thin.
Embryonic shell with irregular axial threads
and fine oblique grooves crossing each other.
Comb-lateral of radula with numerous cusps.
Female reproductive system with provaginal
opening; provaginal duct very short, thin;
bursa copulatrix very prominent; ascending
limb of V-organ elongated, curved; receptacu-
lum seminis on descending limb displaced to
the ventral side and directed posteriorly.

Etymology

The name refers to the small size of mem-
bers of this subgenus.

Discussion

The characters of the female reproductive
system, in combination with the embryonic

shell surface sculpture clearly assign the
new subgenus to the genus Alcadia and dis-
tinguish it from Helicina and Schasicheila,
which also may have been considered. De-
tails about the characteristics of these gen-
era as well as on other Central American
supraspecific taxa, are given below.

The only subgenus which has not been
included due to its South American occur-
rence and the absence of material for study
is Trichohelicina Weyrauch, 1966 (type
species by original designation: Helicina
(Trichohelicina) klappenbachi Weyrauch,
1966, NE-Argentina, Misiones Province). п
the light of the unexpected higher
supraspecific diversity among the South
American Helicinidae, such as Angulata,
assignment to Helicina by Weyrauch (1966)
certainly requires a critical reinvestigation,
because it is only based on an impression
in the basal callus. The hairy periostracum
in both subgenera is regarded as a typical
ecological adaptation of several ground-
dwelling species and does not primarily in-
dicate a closer relationship. In the absence
of further morphological data for
Trichohelicina, the main differences are the
peculiar parietal canal in the upper edge of
the aperture and the differentiated outer lip
which are absent in Microalcadia п.
subgen.

Alcadia (Microalcadia) hojarasca
(Richling, 2001)

Helicina hojarasca Richling, 2001: 5-6 (text
figure)

Type Material

Holotype: INBio 3404979, (leg. I. Richling,
14.8.1999)

Paratype 1: ZMB 103387 (same data as holo-
type)

Dimensions (height/greatest diameter):

Holotype: 2.4/2.9 mm

Paratype 1: 2.2/2.8 mm

Type Locality

NW-Costa Rica, Guanacaste Province, Cor-
dillera de Tilarán, about 9 km N of Santa
Elena, near Mirador Gerardo, 10°22’19"М,
84°48'25"W, 1,450 m a.s.l., primary cloud for-
est.

CLASSIFICATION ОЕ HELICINIDAE 371

FIG. 249. Alcadia hojarasca, holotype, INBio
3404979, height 2.4 mm; scale bar 2.5 mm.

Examined Material

Lec. |. RICHLING

Guanacaste: About 9 km N Santa Elena,
Sendero at Mirador Gerardo, 10%22'19"N,
84°48'25"W, 1,450 m a.s.l.: 14.08.1999: (IR
933)

Puntarenas: Punarenas: Zona Protectora
Arenal-Monteverde: Reserva Biológica
Bosque Nuboso Monteverde (about
10°18’08"М, 84%47'41"W, 1,500-1,650 m
a.s.l.): 25.02.2001: (IR 1453)

Etymology

The species is named for its habitat
“hojarasca” (Spanish) = “leaf litter”, it is used
as a noun in apposition.

Description

Shell (Figs. 249, 250, 3360): Very small, glo-
bose, fragile. Color yellowish-brown. Embry-
onic shell with about 1 whorl, without clearly
marked transition to adult shell; subsequent
whorls 3 to 3'/,, regularly increasing in size.
Surface with fine spiral cords that are axially
crossed by coarse periostracal folds that
form hairs at regular distances, so that the
whorls bear spiral rows of hairs; 4 rows
present on body whorl, on previous whorls
the upper 2 rows still present. Hairs rather
thick and towards the end extending in
breadth, spatula shaped (Fig. 259B). Basal
callus whitish and near columella surface

FIG. 250. Alcadia hojarasca, holotype, INBio
3404979, height 2.4 mm; scale bar 1 mm.

FIG. 251. Teleoconch surface structure of
Alcadia hojarasca, 2"* whorl; scale bar 100 um.

granular. Aperture oblique, rather straight.
Outer lip not differentiated from whorl, ap-
pearing like not fully grown.

Internal Shell Structures: Could not be inves-
tigated.

Teleoconch Surface Structure (Fig. 251): The
coarse, irregular axial threads on the inner
curvature of the embryonic shell continue for
about a half whorl on the teleoconch before
transforming to the typical, numerous spiral
cords.

Embryonic Shell (Fig. 252): The inner curva-
ture is sculptured with coarse, irregular axial
threads; the marginal part shows numerous
fine, oblique grooves crossing each other. A
pitted structure does not occur.

Diameter: 566 um (n = 1).

Operculum (Fig. 253): Thin, only slightly calci-
fied, calcareous plate only covering the cen-
tral area. Columellar margin irregularly
S-shaped, upper edge acute, but rounded,
at lower edge columellar margin continu-
ously changing into outer margin. Nearly
transparent, whitish-amber colored. Inner
side with a little ridge parallel to the columel-
lar margin.

Animal (Fig. 339F): The color is not very un-
usual, mantle and upper side of the foot and
head region are greyish, towards the under-
side it becomes paler.

Radula (Fig. 254): Only two specimens were
studied and the mounting procedure was

372 RICHLING

difficult due to the small size and preserva-
tion conditions. B-central with 6 small cusps;
C-central rather crenulate; R- and A-central
not seen. Comb-lateral with 11-12 pointed
denticles, accessory plate relatively larger
than in other species, about the size of the
comb-lateral. Cusps on marginals rapidly
increasing in number. Number of rows not
counted.

Female Reproductive System (Fig. 255): The
ascending limb of the V-organ is elongated
and curved; the receptaculum seminis 1$ FIG. 253. Operculum of A/cadia hojarasca,
translocated to the ventral side of the de- paratype, ZMB 103387; scale bar 0.5 mm.
scending limb and directed upwards poste-
riorly. Bursa copulatrix and provaginal sac
exhibit a simple structure, with the former
being much larger and approximately
reaching the top of the V-organ. Their con-
nections to the reception chamber are very
close to each other; at the same point en-
ters a very slender, relatively short
provaginal duct.

Morphometry and Sexual Dimorphism

On one hand, the material available 1$
scanty, while, on the other, the peculiar lack of
the development of a differentiated outer lip of

FIG. 252. Embryonic shell of Alcadia hojarasca; FIG. 254. Radula of Alcadia hojarasca. A. Comb-
scale bar 100 um. lateral. B. Marginals; scale bar 50 um.

CLASSIFICATION OF HELICINIDAE 373

FIG. 255. Female reproductive system of Alcadia
hojarasca, IR 1242; scale bar 0.5 mm.

86° DO | i

the species renders it impossible to recognize
mature specimens without closer anatomical
studies of the individuals. Furthermore,
whether or not mature specimens still in-
crease in shell size remains an unanswered
question.

Habitat

Alcadia hojarasca is a ground dweller, it was
only found under and between leaves in differ-
ent stages of decay. The species appears to
have a preference for Cecropia-leaves. This
observation was also made by Zaidett
Barrientos for A. boeckeleri.

Distribution (Fig. 256)

The species is only known from the type lo-
cality and adjacent areas. It is found on the
higher elevations of the northeastern slope of
the Cordillera de Tilaran.

RE

| 3500 - 4000 т
3000 - 3500 m

2500 - 3000 m
2000 - 2500 m
1500 - 2000 m
1000 - 1500 т

09°

500 - 1000 т
100 - 500 т

0-100 m |

e coll. IR
e coll. INBio
© others

FIG. 256. Records of Alcadia hojarasca in Costa Rica.

374 RICHLING

Discussion

On the Central American mainland there are
no other known species of this size that have
periostracal hairs persisting in adult stage. The
only species with a hairy periostracum on the
mainiand belong to the genus Schasicheila,
which is easily characterized by the protruding
edges of the operculum, a different embryonic
shell structure, and great differences in the fe-
male reproductive system (see below).

The only Central American species compa-
rable in size or probably shape is Helicina
exigua L. Pfeiffer, 1849, described from Hondu-
ras. But in the original description, there is no
hint given as to periostracal hairs. The surface
is described as “subtilissime punctato-striatula”,
but even eroded periostracal hairs would not
leave punctulations because they originate
from projections of periostracal folds. Further-
more, H. exigua has never been illustrated, and
it is treated by von Martens (1891) as a dubious
species.

It is rather unusual for helicinids or snails in
general not to have a distinctly developed or at
least thickened outer lip at the aperture as a
sign of maturity and not to further grow in size.
In fact, this species looks somewhat immature.
On the one hand, field collections revealed
specimens of different sizes but even the big-
gest specimens did not display a differentiated
aperture. But on the other hand, dissections of
specimens of about the same size as the type
have shown that the reproductive system 1$
fully developed, and in the female's receptacula
seminis sperm is present, as it is in the male's
vas deferens. So there is strong evidence to
suggest that the specimens described above
represent the adult stage, although it cannot be
determined in the case of every individual if it
already has reached maturity, because a nor-
mal size variation combined with sexual dimor-
phism still have to be taken into consideration.

For comparison with A/cadia boeckeleri, re-
fer to that species.

Alcadia (Microalcadia) boeckeleri
(Richling, 2001)

Helicina boeckeleri Richling, 2001: 6-7 (text
figure)

Type Material

Holotype: INBio 3404980, male (leg. 1. Richling,
12.3.2000)

Paratype 1: ZMB 103388, female (same data
as holotype)

Dimensions (height/greatest diameter):

Holotype: 2.2/2.6 mm

Paratype 1: 2.3/2.7 mm

Type Locality

NW-Costa Rica, Guanacaste Province,
Parque Nacional Guanacaste, about 10 km S
of Santa Cecilia, Volcán Orosí, near field sta-
tion Pitilla, 10%59'18"N, 85°25'34"W, 700 т
a.s.l., beginning of Sendero Orosilito, primary
forest.

Examined Material

INBio COLLECTION

Guanacaste: Parque Nacional Guanacaste, La
Cruz, 9 km $ de Santa Cecilia, Estacion Pitilla:
10°59’25"N, 85*25'38"W, 700 т a.s.l.: leg.
malacological staff of INBio, 01.03.1995: 9
spec. (INBio 1498481); leg. Evelio Alfaro,
19.04.1995: 1 spec. (INBio 1483310);
10%59'33"N, 85°25’46"W, 700 m a.s.l.: leg.
Dunia Garcia, 10.08.1995: 1 ad. (INBio
1488038); 7 ads., 1 s.ad. (INBio 1488068);
Sector Finca Nacho, 10*58'43"N, 85°25'49"W,
700 m a.s.l.: leg. С. Moraga, 18.08.1994: 1 ad.
(INBio 1480339); Sendero Mena, 10*59"27"N,
85°25’49"W, 700 т a.s.l.: leg. malacological
staff of INBio, 01.06.1993: 6 spec. (INBio
1466290); Sendero Los Memos, 11°02’00"N,
85°25'20"W, 700 т а. $.1.: leg. Calixto Moraga,
02.04.1995: 1 spec. (INBio 1482793)

Etymology

The species is dedicated to Dr. Wolfgang
Bóckeler who first introduced me to Costa

FIG. 257. Alcadia boeckeleri. A. Holotype, INBio
3404980, height 2.2 mm. B. Paratype, ZMB
103388; scale bar 2.5 mm (A), 1 mm (B).

CLASSIFICATION OF HELICINIDAE 375

258

FIGS. 258, 259. Alcadia spp. FIG. 258. Alcadia boeckeleri, holotype, INBio 3404980, height 2.2 mm.
FIG. 259. Periostracal hairs of A. Alcadia boeckeleri, INBio 3404980. B. Alcadia hojarasca, INBio
3404979; scale bar 1 mm (Fig. 258), 0.5 mm (Fig. 259).

Rica and subsequently so often joined me in
my search for the hidden helicinids.

Description

Shell (Figs. 257, 258, 336P): Very small, glo-
bose and fragile. Color yellowish-brown.
Embryonic shell with about 1 whorl, without
clearly marked transition into adult shell; sub-
sequent whorls about 3, regularly increasing
in size. Surface with fine spiral cords that are

axially crossed by very fine threads. On last
whorl, 5 rows of periostracal hairs; on previ-
ous whorls the upper 2 rows are present.
Hairs thin and at the end sharpened (Fig.
259A). Basal callus weakly developed and
near columella granulated. Aperture oblique,
rather straight. Outer lip undifferentiated from
whorl, appearing as not fully grown.

Internal Shell Structures: Could not be inves-

tigated.

FIG. 260. Teleoconch surface structure of Alcadia

boeckeleri, 1* whorl; scale bar 100 um.

FIG. 261. Embryonic shell of Alcadia boeckeleri;
scale bar 100 рт.

376

Ne

FIG. 262. Operculum of A/cadia boeckeleri,
holotype, INBio 3404980; scale bar 0.5 mm.

Teleoconch Surface Structure (Fig. 260): А
zone of transitional structure of about */, of a
whorl, closely resembling oblique diverging
grooves, is continued by numerous spiral
cords, crossed only by growth lines.

[86° i [85°

RICHLING

Embryonic Shell (Fig. 261): The embryonic
shell surface structure is similar to that of
Alcadia hojarasca.

Diameter: 488 pm (п = 1).

Operculum (Fig. 262): Thin, only slightly calci-
fied, calcareous plate only covering the cen-
tral area. Columellar margin irregularly
S-shaped, upper edge acute, but rounded,
at lower edge columellar margin continu-
ously changing into outer margin. Nearly
transparent, whitish-amber colored. Inner
side with a little ridge parallel to the columel-
lar margin.

Animal (Fig. 339G): The color is similar to
Alcadia hojarasca.

Radula: The radula of Alcadia boeckeleri
could not be investigated, because sufficient
material was lacking.

i — — 83

44°

3500 - 4000 m
3000 - 3500 m

- | MA: 107

2500 - 3000 т
2000 - 2500 т
1500 - 2000 т
1000 - 1500 т
500 - 1000 т
100 - 500 м
0- 100m

09°

| e coll. IR
| @ coll. INBio |
| © others

FIG. 263. Records of A/cadia boeckeleri in Costa Rica.

CLASSIFICATION OF HELICINIDAE 377

Female Reproductive System: The inspection
of a single female revealed similar structures
as in Alcadia hojarasca.

Morphometry and Sexual Dimorphism
See Alcadia hojarasca.
Habitat

The species 1$ a typical ground dweller that
lives in the leaf litter. See also under Alcadia
hojarasca.

Distribution (Fig. 263)

The species is only known from the area of
the type locality located on the northeastern
slope of the Cordillera de Guanacaste, which,
together with the volcano Orosí, forms the
northern limit of this mountain chain.

Discussion

For general discussion, see Alcadia
hojarasca.

Alcadia boeckeleri differs from A. hojarasca
in bearing five instead of four rows of
periostracal hairs. Furthermore, the hairs are
much thinner and they have a different shape
towards the end (Fig. 259). The spire 1$ a little

more elevated in A. boeckeleri.

Lucidella lirata
(L. Pfeiffer, 1847)

Helicina lirata L. Pfeiffer, 1847a: 150 (not fig-
ured)

Helicina lirata — L. Pfeiffer, 1847b: 153

Helicina lirata — L. Pfeiffer, 1848: 83

Helicina unidentata — L. Pfeiffer, 1848: 83
[without description]

Helicina unidentata L. Pfeiffer, 1849: 125 (not
figured): Honduras (Dyson, coll. Cuming)
Helicina rusticella Morelet, 1849: 21 (not fig-

ured): Island Carmen
Helicina lirata — L. Pfeiffer, 1850: 1415, pl. 4,
figs. 40-43: Mexico: Yucatan (Hegewisch)
Helicina unidentata — L. Pfeiffer, 1850: 14, pl.
9, figs. 14-17
Helicina lirata — L. Pfeiffer, 1852a: 341
Helicina unidentata var. — L. Pfeiffer, 1852a:
341
Helicina unidentata — L. Pfeiffer, 1852a: 341
Helicina lirata — L. Pfeiffer, 1852b: 246

Helicina unidentata var. — L. Pfeiffer, 1852b: 246

Helicina unidentata — L. Pfeiffer, 1852b: 246

Helicina lirata — L. Pfeiffer, 1856b: 236:
Mexico: Chiapa (Ghiesbreght)

Helicina lirata — von Martens, 1860: 59: Ven-
ezuela: near Maracaybo or environs of
Merida

Helicina lirata — Tristram, 1864: 413: Guate-
mala: mountain-forests of Vera Paz (Salvin)

Helicina lirata — von Martens, 1865: 67

Helicina semistriata Sowerby, 1866: 281, pl.

268, fig. 86

Helicina unidentata — Sowerby, 1866: 281, pl.
268, fig. 87

Helicina lirata — Sowerby, 1866: 281, pl. 268,
figs. 88-89

Helicina lirata — Bland, 1866: 8

Helicina unidentata — Bland, 1866: 8

Helicina (Perenna) lamellosa Guppy, 1867:
260, pl. X, fig. 4: Trinidad: Gulf of Paria, is-
let Cotoras

Helicina semistriata — Tate, 1870: 159: Nicara-
gua: in the woods and cocoanut groves
about Boca del Toro, region Chontales [area
around Acoyapa, NE of Lago de Nicaragua]

Helicina lirata — Strebel, 1873: 21, pl. 1, fig. a,
pl. 2 fig. 8: Mexico: Bajadas, Veracruz and
near Antigua

Helicina lirata — von Martens, 1873: 56: Ven-
ezuela

Helicina lirata — Reeve, 1874: pl. 14, fig. 121

Helicina unidentata — Reeve, 1874: pl. 14, fig.
122

Helicina lyrata [sic] - Angas, 1879: 484: Costa
Rica (Gabb)

Helicina unidentata — Ancey, 1886: 254: Hon-
duras: lle d’Utilla (Dyson, Simpson)

Lucidella lirata — Jousseaume, 1889: 232,
235, 256: Venezuela: Caracas, San-Esteban

Helicina lirata — von Martens, 1891: 4142, pl.
|, fig. 18 (living animal): E-Mexico: Vera
Cruz, at the “bajadas”; S-SE-Mexico:
Chiapas, Teapa and San Juan Bautista in
Tabasco; Yucatan; N-Guatemala: mountain
forests of Vera Paz; S-Guatemala:
Retalhuleu; Venezuela

Helicina lirata var. rusticella — von Martens,
1891: 41: Yucatan: Island of Carmen, in the
Gulf of Campeche

Helicina lirata var. unidentata - von Martens,
1891: 41: Honduras; 607: Honduras: Utila
Island

Helicina lirata var. semistriata — von Martens,
1891: 41: N-Panama: Boca del Toro,
Chiriqui

78 RICHLING

7
ЭГО

Helicina lirata — Pilsbry, 1891: 332: Mexico: N-
Yucatan: Labna; Honduras: Utilla Island
(Simpson)

Helicina (Poenia) lirata — Fischer & Crosse,
1893: 397-399: same data as von Martens,
1891 (for H. lirata and H. lirata var.
unidentata) and Mexico: Yucatan, Labna

Helicina (Poenia) lirata var. rusticella — Fischer
& Crosse, 1893: 397-399: same data as von
Martens, 1891

Helicina (Perenna) lamellosa — Guppy, 1893:
228

Helicina (Helicina Perenna) lirata — Guppy,
1895: 74

Helicina (Helicina Perenna) semistriata —
Guppy, 1895: 74

Нейста (Helicina Perenna) lamellosa -
Guppy, 1895: 74

Helicina lirata — von Martens, 1900: 607: S.E.
Mexico: San Juan Bautista, garden of the
Juarez Institute in the same town; Yucatan:
Labna; N-Guatemala: Panzos; SW-Costa
Rica: Alto de Mano Tigre, 690 m [not local-
ized] (Pittier)

Helicina lirata var. rusticella — von Martens,
1900: 607: SW-Costa Rica: El Pozo, in the
shingle (gravices) of the Rio Grande de
Terraba [not localized: Palmar Norte:
08%57'N, 83°27 W, Puntarenas Province]
(Pittier)

Helicina lirata — Pilsbry, 1904: 782: Mexico:
Veracruz: Antigua (Rhoads)

Helicina lirata — Pilsbry, 1910: 503: Panama:
Canal Zone: Tabernillo (Brown)

Lucidella lirata - Wagner, 1911: 341, pl. 68,
figs. 5-7: S-Mexico, Guatemala, Honduras,
Venezuela

Lucidella lirata lamelllosa — Wagner, 1911:
341342, pl. 68, fig. 4: Trinidad (island)

Lucidella (Perenna) lirata — Pilsbry & Brown,
1912: 585

Lucidella lirata var. lamellosa — Vernhout,
1914: 26-27: Suriname: Environs of
Paramaribo

Lucidella lirata — Hinkley, 1920: 41, 49, 52:
Guatemala: Livingston; Jocolo plantation on
north side of Lake Isabal: lake drift; Alta
Verapaz: Chama between Rio Tsalbha and
Rio Negro: also river drift

Lucidella (Poenia) lirata — Baker, 1922а: 54-
55, pl. Ш, fig. 5, pl. М, fig. 21 (radula):
Mexico-Venezuela; Mexico: Tabasco: San
Juan Bautista: Garden of Juarez Institute
(Rovirosa)

Lucidella (Poenia) lirata — Baker, 1922b: 36:
Mexico: S Vera Cruz, near hacienda de

Cuatolapam (Rio San Juan — Arroyo
Hueyapam, canton of Acayacan (Michigan-
Walker-Expedition)

Lucidella (Poenia) lirata — Baker, 1923: 22-23:
Venezuela: San Esteban, Palma Sola, Aroa,
Estación Táchira, La Fria (Michigan-
Williamson-Expedition)

Lucidella lirata — Pilsbry, 1926a: 59, 71:
Panama: Canal Zone: Tabernilla (Brown),
near Darien and Juan Mina (Zetek), Panama
City and Taboga Island (Zetek), Bocas del
Toro (Gabb)

Lucidella lirata — Pilsbry, 1926b: 127: Costa
Rica: Cahuita [09°44’01"М, 82°49'48"W]
(Olsson)

Lucidella (Роета) lirata — Baker, 1928: 33-34,
pl. II, figs. 9-11 (female reproductive sys-
tem): Mexico: Veracruz: Atoyac, 1300-1475
feet

Lucidella lirata — Pilsbry, 1930: 339: Panama:
Canal Zone: roadside in SE of Empire;
Taboga Island (Pinchot-Expedition)

Lucidella (Poenia) га — Bequaert 8 Clench,
1933: 543: Mexico: Yucatan, Chichen Itzá

Lucidella lirata — Goodrich & van der Schalie,
1937: 12, 14-16, 33: Guatemala: Petén: re-
gion of headwater of Río San Pedro de
Mártir, lower Río de la Pasión; Alta Verapaz:
upper part of Río de la Pasión

Lucidella lirata — Richards, 1938: 176: Hondu-
ras

Lucidella lirata — Richards & Hummelinck,
1940: 12-13: Venezuela: Margarita Island:
Hills SE La Asunción; Cerro del Piache; just
above El Valle; La Sierra, El Valle; Toma de
Agua del Valle; Toma de Agua de Encañado,
San Juan; Los Vagras; between Los Vagras
and coast

Lucidella lirata — Bequaert, 1957: 208: Mexico:
Chiapas: Selva Lacandona: Laguna Ocotal,
950 m, Laguna Ocotal to El Censo, 1,000 m;
Veracruz, Tabasco, Yucatan, Quintana Roo,
Guatemala to Panama

Lucidella lirata lamellosa — Bequaert, 1957:
208

Lucidella lirata — Basch, 1959: 8: Guatemala:
Petén: Tikal National Park, 17°10’N,
89°25'W

Lucidella lirata — Hubricht, 1960: 83: USA: S-
Texas: beach drift

Lucidella (Poeniella) lirata -— Haas & Solem,
1960: 130: British Honduras [Belize]: Rio
Frio Cave, Cayo District

Lucidella lirata — Branson 8 McCoy, 1963:
102-103: Mexico: Campeche: Airport,
Cuidad del Carmen

CLASSIFICATION OF HELICINIDAE 379

Lucidella lirata — Thompson, 1967: 228229:
Mexico: Campeche: 8.1 mi SW Champotón,
5.1 mi NNW Dzibalchén, 4.9 mi W
Hopelchén, 3.4 mi $ Cayal (19°45’М,
JOMOW) 72 MiS: Pixtün; 10:2 пи Е
Escárcega, 19.2 mi Е Silvituc; Quintana
Roo: 4 mi E Xpujil, 7.1 mi NNW Xiatil

Lucidella lirata — Regteren Altena, 1974: 71:
Suriname

Lucidella lirata — Tillier, 1980: 35, 36, figs. 20,
21 (operculum): French Guiana: Aouara,
Saut Sabbat (Abattis)

Lucidella (Poenia) lirata - Thompson, 1982:
fig. 13 (radula), 27-28 (embryonic shell)
Lucidella lirata — Monge-Nájera, 1997: 113:

Costa Rica

Synonymy

Helicina unidentata L. Pfeiffer, 1849
Helicina rusticella Morelet, 1849
Helicina semistriata Sowerby, 1866
Helicina lamellosa Guppy, 1867

Original Description

“Т. orbiculato-conoidea, tenuis, acute et
confertim concentrice lirata, diaphana, albida;
spira conoidea, acuta; anfr. 4,5-5 vix
convexiusculi, ultimus carinatus, basi medio
impressus; apertura obliqua, rotundato-
subtriangularis; columella brevissima, sim-
plex, in callum basalem tenuissimum dilatata;
perist. breviter expansum, margine basali
medio obsolete unidentato.

Diam. 4, alt. 2 */, mill.
Habitat in Mexico, Yucatan (Hegewisch).”

Type Material

Not located (assumed to be in the collection
of L. Pfeiffer, because it was not otherwise
stated, collection L. Pfeiffer having been most
likely destroyed in Stettin Museum, Poland
during World War II).

Type Locality

“Mexico, Yucatan” [not clear, whether it re-
fers to the Mexican State of Yucatán or the
Mexican part of the peninsula of Yucatán].
Examined Material
Lec. |. RICHLING

Limon: Parque Nacional Cahuita, trail from
Cahuita to Puerto Vargas, coastal forest with

coco palms: about 09°43’27"М, 82*50'28"W,
4 m a.s.l.: 02.03.1998: (IR 418); 10.03.1999:
(IR 756); 08.08.1999: (IR 902); 10.08.1999:
(IR 913); 04.03.2000: (IR 1314); (IR 1316);
14.03.2001: (IR 1558); (IR 1559); (IR 1640)
Refugio Nacional de Fauna Silvestre
Gandoca-Manzanillo, S Manzanillo, trail
along coast line to S, coastal forest, about
O9°38'06"N; 62°38'26"W, 50° т aisil.,
14.09.1999: (IR 1098); (IR 1124)

W Liverpool, Mexico, at Rio Blanco, high
water deposit, 09°58’32"М, 8308'32"W, 35
m а.5.1.: 22.02.1997: (IR); 12.03.1997: (IR
118)

N Shiroles, along Quebrada Kirio,
093938, 62`57'20'\\: 7120: таз:
15.03.1997: (В 162); 160 ea. s
03.03.1998: (IR 435)

W Bribri, road to Uatsi, about 09°38’11"М,
82*51'48"W, 30 т a.s.l.: at crossing with Rio
Carbón, 30 m а.5.1.: 17.3.1997: (IR 187);
wooded valley within banana plantation, 50
m’a.s.l.: 15.3.2001; (IR 1586)

Southern road from Bribri to Shiroles, small
banana plantation near creek, 09°35'17"N,
62 52:46'"W, 50 m a.s.l.. 15.03.1997: (IR
1.21)

Puntarenas: Refugio Nacional de Fauna
Silvestre Golfito, rain forest, 08°39’26"М,
83”10'50"W, 100 m a.s.l., 14.02.1999: (IR
568)

INBio COLLECTION

Limón: Parque Nacional La Amistad,
Quebrada Cachabri (toma de agua),
09°29'29"N, 82°59'37"W, 360 m a.s.l., leg.
Gerardina Gallardo, 26.11.1996: 3 spec.
(INBio 1488249)
Reserva Indígena Talamanca: 1 km SW de la
Iglesia de Amubri, 09*30'37"N, 82%57'36"W,
70 т a.s.l.: 19.10.1996: 8 spec. (INBio
1488235); 500 т E de la Iglesia de Amubri,
O9°31/06°N, 92756 50 We 70 т ras:
21.10.1996: 6 spec. (INBio 1488268); Sector
Amubri, 09°30’53"М, 82°57’19"W, 70 m a.s.l.:
29.11.1994: 1 spec. (INBio 1483399);
30.11.1994: 1 spec. (INBio 1483446); 2 spec.
(INBio 1483447); 1 spec. (INBio 1483449); 6
spec. (INBio 1483450); 24.10.1996: 30 spec.
(INBio 1487958); 27.11.1996: 1 spec. (INBio
1487352); Amubri, Sendero Soki,
09`30'53№М 625719” 70 т ‘a-sil::
27.11.1996: 1 spec. (INBio 1487364); 11
spec. (INBio 1488219); 11 spec. (INBio
1493414); Suiri, orillas del Rio Telire,
O933SD0N, 82°5550°W, 30 mas
25.11.1996: 17 spec. (INBio 1487345) (all

380 RICHLING

leg. Gerardina Gallardo), Sector Miramar,
Senderos a Rio Moin, 09°37’44"N,
83°00'32"W, 150 m a.s.l.: leg. Zaidett
Barrientos, 08.11.1994: 4 spec. (INBio
1475236)

1 Km $ de Рита Cocles, 09°38'17"N,
82°43'25"W, 40 т a.s.l., leg. Zaidett
Barrientos, 20.08.1996: 1 spec. (INBio
1487843)

Refugio Nacional de Vida Silvestre
Gandoca-Manzanillo: Sector Gandoca,
09°35’30"М, 82°36'13"W, 0 т a.s.l.:
29.07.2000: 4 spec. (INBio 3091175); Sector
Gandoca, Camino а Gandoca, 09°38'04"N,
82°38'37"W, 10 m a.s.l.: 28.04.2000: 5 spec.
(INBio 3097927); Sector Manzanillo,
Sendero a Gandoca, 09°38'20"N,
82*39'03"W, 2 m a.s.l.: 30.03.2000: 1 spec.
(INBio 3098034) (all leg. Alexander Alvarado
Mendez)

Parque Nacional Cahuita: Sector Puerto
Vargas, 09°43'38"N, 82°49’09"W, O m a.s.l.:
leg. Alexander Alvarado Mendez,
31.08.1999: 1 spec. (INBio 3091727); 4
spec. (INBio 3091733); Sendero del limite W
del parque, 09°44’00"N, 82°50’25"W, 10 т
a.s.l.: leg. malacological staff of INBio,
11.06.1997: 1 spec. (INBio 1488171)
Reserva Indígena Tayni: Sendero
Tepezcuintle, 09°40’22"М, 83*01'46"W, 180
т a.s.l: 22.04.1999: 1 spec. (INBio
3096423); Sendero Bobocara: 09°40’28"М,
83°02'17"W, 260 m a.s.l., 01.06.1999: 1
spec. (INBio 1498178); 09°40’28"М,
83°02'12"W, 200 т a.s.l., 01.06.1999: 7
spec. (INBio 1498248) (all leg. Alexander
Alvarado Mendez)

Reserva Biológica Hitoy Cerere: Sendero
Tepezcuintle: 09°40'22"N, 83°01’40"W, 140
m а.5.1., 25.04.1999: 3 spec. (INBio
1497566); 100 т a.s.l., 07.06.1999: 4 spec.
(INBio 3096478); 09°40’18"М, 83°01’43"W,
140 m a.s.l., 28.04.1999: 1 spec. (INBio
1497840); Sendero Boböcara, 09°40’20"М,
83°03’12"W, 620 m a.s.l.: 14.06.1999: 3
spec. (INBio 3095831) (all leg. Alexander
Alvarado Mendez): Sendero Toma de Agua,
09°40’31"N, 83%01'36"W, 100 т a.s.l.:
19.04.1994: 1 spec. (INBio 1473669); 2
spec. (INBio 1473674); 58 spec. (INBio
1474306); 30 spec. (INBio 1474336);
20.04.1994: 1 spec. (INBio 1473613) 1
spec. (INBio 1473838); 08.09.1994: 49
spec. (INBio 1475430); 70 spec. (INBio
1475444) (all leg. Zaidett Barrientos);
28.02.1994: 4 spec. (INBio 1476129);

17.07.1994: 70 spec. (INBio 1478443); 30
spec. (INBio 1478459); 06.12.1994: 2 spec.
(INBio 1475673) (all leg. Gerardo Carballo);
Estación Hitoy Cerere, 09°40'35"N,
83°01'36"W, 100 m a.s.l.: leg. malacological
staff of INBio, 15.11.1993: 7 spec. (INBio
1463364); Sendero Chato, 09°40’41"N,
83°01'26"W, 100 m a.s.l.: leg. Marianella
Segura, 17.07.1994: 10 spec. (INBio
1478236); 70 spec. (INBio 1478239

400m NE de la Estaciön de Hitoy Cerere,
Sendero la “Finca”, 09°40’35"М,
83°01'26”М/; 150 m a.s.l.: 20.07 1999562
spec. (INBio 1495436); 110 т a.s.l.:
07.05.1999: 3 spec. (INBio 3300038);
27.09.2000: 1 spec. (INBio 3091795) (all
leg. Alexander Alvarado Mendez)

Isla Uvita: frente al muelle de Limón,
09°59’45"N, 83°00’50"W, 5 т a.s.l.: leg.
Alexander Alvarado Mendez, 11.10.2000: 21
spec. (INBio 3315375); 3 spec. (INBio
3315385); lado N, 09°59’50"N, 83°00’46"W,
-10 to 5 m a.s..: leg. A. Berrocal,
06.05.2000: 21 spec. (INBio 3396974)

Puntarenas: Playa Blanca, 08°38’18"М,

83°26 16"W, О т a.s.l., leg. Gillermo Mena,
04.09.1995: 1 spec. (INBio 1479925)

Isla Pelicanos, 08°36’10"М, 83*08'48"W, 2 т
a.s.l., leg. Socorro Avila, 01.11.1997: 1 spec.
(INBio 3399211)

Refugio Nacional de Fauna Silvestre Golfito:
Sendero Las Torres, 08°38’37"М,
83°09'54"W, 60 m a.s.l.: leg. Socorro Avila,
03.12.1997: 4 spec. (INBio 1487712); 150 т
al N del Reserva, 08*39'06"N, 83°10'44"W,
40 m a.s.l.: leg. Alexander AlvaradoMendez,
14.02.1999: 3 spec. (INBio 1501379);
Extremo NW del Aeropuerto de Golfito,
08°39'39"N, 83°11'13"W, 100 m a.s.l.: leg.
Socorro Avila, 03.11.1997: 1 spec. (INBio
1487165), 3 spec. (INBio 1487172)
Parque Nacional Piedras Blancas, Playa
San Josecito, 08”39'49"N, 83°15'35"W, 5 т
a.s.l., leg. Еда Fletes, 27.10.1996: 2 spec.
(INBio 1487318)

Quebrada Benjamin, carretera al tanque del
agua 600 m del Barrio Alemania,
08°58’39"М, 83°28'19"W, 100 m a.s.l., leg.
Socorro Avila, 08.05.1997: 1 spec. (INBio
1487661); 1 spec. (INBio 1487691)
Palmar Norte, Barrio Alemania, Sendero a
Jalisco, 1 Km al NE del Tanque de
Acueductos, 08°59’21"М, 83°28'16"W, 400
т a.s.l., leg. Socorro Avila, 08.05.1997: 2
spec. (INBio 1487736); 3 spec. (INBio
1487756)

CLASSIFICATION ОЕ HELICINIDAE 381

FIG. 264. Lucidella lirata, Cahuita, IR 1559; scale bar 1 mm.

OTHER SOURCES
COSTA RICA
Limón: Pandora [about 09°43’М, 82°58'W],

ish: as in original description only faded
specimens). Whorls sculptured with a vary-
ing number of close-set, prominent, sharp

leg. F.G. Thompson (FGT-100), 05.08.1964
(UF 214773)

7 km from Valle La Estrella, at Hitoy Cerere
National Park [about 09°40'35"N,
8301'36"W], 152 m a.s.!., E.L. Raiser et al.
(ERL 079), 09.08.1994 (UF 41405, UF
41406); E.L. Raiser (ERL 080), 09.08.1994
(UF 41421)

1 km NW of Cahuita, 09°45.5'N, 82°50.9'W,
leg. Е.С. Thompson (ЕСТ-5616), 25.02.1996
(UF 268476)

NICARAGUA

Matagalpa: 4.5 km. $ of Matagalpa, 1,200 т
a.s.l., leg. Е.С. Thompson, 16.07.1956 (UF
127683)

Description
Shell (Figs. 264, 336Q): Orbiculate-conoidal,

depressed, thin, small sized, dull. Color:
dark yellowish to brown, diaphanous (whit-

FIG. 265. Axial cleft and muscle attachments of

Lucidella lirata, IR 1559; scale bar 2 mm.

spiral ridges at about equal distance, upper
a little wider spaced, some smaller ridges in
between. Surface textured with irregular,
strong growth lines crossing spiral sculpture.
Embryonic shell with about 1 whorl; 3°/,—4
subsequent whorls slightly convex; last
whorl rounded with a slight keel on periph-

FIG. 266. Teleoconch surface structure of
Lucidella га, 2"? whorl; scale bar 100 um.

382 RICHLING

FIG. 267. Embryonic shell of Lucidella lirata;
scale bar 100 pm.

ery, umbilical region deeply impressed;
whorls rapidly increasing in size and only
slightly descending, forming a low spire and
a slightly pointed apex. Suture deeply im-
pressed. Aperture oblique and curved back-
wards towards its base. Outer lip of the
same color as the preceding whorls, thick-
ened and narrowly expanded, a slight notch
at insertion to body whorl. Reflection very
narrow, lower palatal margin with a broad

FIG. 268. Operculum of Lucidella lirata, IR 1559; FIG. 269. Radula of Lucidella lirata. A. Centrals.
scale bar 0.5 mm. B. Comb-lateral. C. Marginals; scale bar 50 um.

CLASSIFICATION OF HELICINIDAE 383

tooth that is more or less well developed.
Columella short. Basal callus weakly devel-
oped and granulated.

Internal Shell Structures (Fig. 265):

Teleoconch Surface Structure (Fig. 266): A
transitional pattern is absent, the teleoconch
is structured with the typical spiral ridges
throughout, the interspaces are smooth ex-
cept for fine growth line.

Embryonic Shell (Fig. 267): Contrary to the
pitted embryonic shells described for spe-
cies of Helicina, the pits of Lucidella lirata
are not arranged in distinct spiral lines. The
interspacial distance exceeds the diameter
of the pits, which are more sparcely scat-
tered over the surface. The embryonic shell
is even smaller than in the smaller species
Alcadia hojarasca and А. boeckeleri.
Diameter: 426 um (+ 12) (408-465) (п = 13)
(IR 756, IR 1314).

Operculum (Fig. 268): Outer surface very
slightly calcified, a noticeable ridge only at
the columellar side, which 1$ quite straight.

horny-amber and

Color transparent.

FIG. 270. Female reproductive system of

Lucidella lirata, IR 1314; scale bar 0.5 mm.

Nucleus at a significant distance from the
columellar margin. Shape broadly ovoid and
only truncated towards the columella.

Animal (Fig. 339H): Only the tentacles are
grey, the rest of the body 1$ whitish-yellow.
The mantle may be spotted grey, but this 1$
only visible in individuals removed from their
shell.

Radula (Fig. 269): A-central elongated and
smooth; B- and C-central each bearing about
5-6 cusps. Comb-lateral with 9-12 cusps,
cusps on marginals slowly increasing in num-
ber. Total number of rows was not counted.
Baker (1922a) and (Thompson, 1982) found
fewer cusps on the comb-lateral.

Female Reproductive System (Fig. 270): The

receptaculum seminis between the two
limbs of the V-organ is not developed; it is
replaced by an accessory sperm sac on the
top of the V-organ or the very beginning of
the descending limb respectively, but it 1$
located on the outside of the V-organ. The
bursa copulatrix is relatively small and with-
out any lobes. In contrast, the distal side of
the provaginal sac is deeply lobed, an addi-
tional lobule may occasionally be developed
on its stalk. The bursa copulatrix enters the
reception chamber via the stalk of the
provaginal sac rather than directly. The
provaginal duct that continues from the stalk
of the provaginal sac is very short and slen-
der. The pallial oviduct is much less folded
than in the species of Helicina. It receives an
additional sac a short distance from the re-
ception chamber, which serves for sperm
storage.
The anatomy of the species has already
been studied by Baker (1928), but, accord-
ing to his studies of Lucidella aureola
(Férussac, 1822), he assumed the existence
of a receptaculum seminis as described
above for the species of Helicina (he had
only a single specimen and “it was broken
away ...”). The study of serial sections ex-
cludes the presence of such an organ. Fur-
thermore, the additional sac on the oviduct
escaped Baker's attention.

Morphometry and Sexual Dimorphism (Table
14, Fig.-271)

Sufficient numbers of specimens were found
only at Cahuita. Because morphological differ-

BD

FIG. 271. Measurements of Lucidella lirata
according to Table 14; on each line: mean value,
standard deviation, absolute range; number of
individuals given as “п = females/males”; upper
line: females, lower line: males; in between and
shaded: average of both.

ences between populations cannot be consid-
ered here, only height and minor diameter
were measured.

The values of males and females widely
overlap (Fig. 272), so that statements based
on very few shells will only be correct by coin-
cidence (Baker, 1928: 1 female > a few
males). On average, females are larger than
males. In interpolation from the minor diam-
eter, the males have an average volume of
about 92.5% of that of the females, thereby
representing the smallest degree of sexual
dimorphism found among Costa Rican spe-
cies of Helicinidae.

Habitat

Lucidella lirata is a typical ground dweller,
living in the leaf litter under and between de-
caying leaves, trunks and fruits. In Costa Rica,
it was found abundantly very close to the sea
shore in a coconut palm forest on sandy
gravel bottom. A semiaquatic behavior (in and
at edge of pools, often together with aestivat-
ing Pisidium and Planorbis) described by
Baker (1922b) for populations in southern
Vera Cruz, Mexico, was not observed. Be-
sides mesic and rain forests, Thompson
(1967) also mentions a constantly wet
swampy area in Campeche, Mexico as habi-
tats of L. lirata.

Distribution

Lucidella lirata is very widespread in Central
America. lt ranges from southern Mexico to
Panama and along the northern coast of South
America, where it is found in Venezuela,
French Guyana, and Suriname. It inhabits
some coastal islands, such as Isla de Utila,
Honduras, and Isla de Margarita and Trinidad,
Venezuela, but it is absent from the southern

RICHLING

Lesser Antilles. In Mexico, L. lirata seems to be
restricted to the southeastern states Veracruz,
Chiapas, Campeche, Quintana Roo, and
Yucatán, although it may be rare towards the
extreme tip of the Peninsula de Yucatán, be-
cause it was only recorded twice from the latter
state (Labna and Chichen Itza). According to
Correa-Sandoval (2000), it has not been re-
ported north of Jalapa in central Veracruz.

The records for Costa Rica (Fig. 273) are
concentrated in two regions, the southern
Pacific and the Caribbean plains, but within
these areas the species was found at various
different sites. According to the quantity of
material and the author's own field experi-
ence, L. lirata occurs more abundantly on the
Caribbean side. The distribution stretches
along the coast line, where slightly elevated
areas are mainly inhabited. In Costa Rica, the
species is only occasionally found on altitudes
up to 620 m in Hitoy Cerere or up to 690 m in
Alto de Mano Tigre on the Pacific side respec-
tively, whereas Bequaert (1957) reported L.
lirata up to 1,000 m in Mexico.

Discussion

Because Costa Rican populations represent
only a small area within the wide distribution of
Lucidella lirata, and because it is the only spe-
cies of that genus in Costa Rica, rendering a
comparison with other similar taxa within the
area of the study unnecessary, it seemed ap-
propriate to investigate and discuss the spe-
cies to a lesser extent than the representatives
of the genus Helicina and to accept the inter-
pretations of earlier authors on this subject
(e.g., synonymy). For this reason, only Costa
Rican material is listed above. A comprehen-
sive revision of the species should encompass
samples from the entire area of distribution,
something beyond the focus of the present
study.

Lucidella lirata varies throughout its area of
distribution with respect to the number of spi-
ral ridges and their extent towards the umbili-
cal area, for example, Venezuelan specimens
show all intermediates between the typical
form and /amellosa (Baker, 1923; Regteren
Altena, 1974; Tillier, 1980). Furthermore, the
elevation of the spire and the dentition of the
outer lip are subject to variation. In the Costa
Rican samples, the basal tooth is usually well
developed, and the upper outer Пр 1$ irregu-
larly crenulated, the spiral striation is present
only on about half the distance or less from
the periphery to the umbilical area.

CLASSIFICATION OF HELICINIDAE

2.2

a | female o
male +

o $ o | |

|

|

26 height [mm] 3

FIG. 272. Range of measurements in females and males of Lucidella lirata
for height and minor diameter in the population from Cahuita.

86°

Е:
pl

3500 - 4000 m
3000 - 3500 m
2500 - 3000 m
2000 - 2500 m
1500 - 2000 m
1000 - 1500 m
500 - 1000 m
100 - 500 m
0-100т

e coll. IR

| © others

e coll. INBio |

FIG. 273. Records of Lucidella lirata in Costa Rica.

385

386 RICHLING

TABLE 14. Measurements of the Cahuita-population of
Lucidella lirata given as mean value with standard devia-
tion, minimum and maximum value (min, max), and num-
ber of specimens (min. diam. = minor diameter); linear
measurements [mm].

“Cahuita” (altitude 5-10 т)
lot IR 1314

Mean

Sex value Deviation Min Max Number

Height f 2.51 0.08 2.31 267 25
Height m 2.41 0.07 227 ~262 19
Min. diam. f 3.14 0.07 2.94 3.33 25
Min. dam. m 3.06 0.067 2927 3212 719

QUESTIONABLE SPECIES FOR COSTA
RICA

Helicina (Oligyra) flavida
Menke, 1828

Examined Material (Fig. 274)

COSTA RICA

Limön: Field cleared of forest vegetation (now
soccer field), adjacent to Los Corales Ш,
Puerto Limón, 10%00'06.7"N, 83*02'37.9"W,
leg. D.G. Robinson, Summer 1984: 2 ads.
(dead collected) (APHIS PPQ USDA)

Distribution

The species is only known for Costa Rica
from the two dead specimens collected very
near Puerto Limón (Fig. 275). Otherwise,
Helicina flavida 1$ widely distributed in south-
ern Mexico (states: Puebla, N- to S-Veracruz,
Tabasco, Chiapas, Campeche, Quintana

FIG. 274. Helicina flavida, Puerto Limón, APHIS
PPQ USDA, height 5.9 mm, 5.1 тт; scale bar
2.5 mm.

Roo), northern Guatemala (departments:
Petén, Alta Verapaz, Izabal) and Belize (Cayo
district) (von Martens, 1890-1901; Hinkley,
1920; Baker, 1922b; Bequaert, 1957; Haas 8
Solem, 1960; Thompson, 1967; Correa-
Sandoval, 2000).

Discussion

Helicina flavida was originally described
from Jamaica, but aside from some records
from the Antilles in some mid-19" century pub-
lications, almost all subsequent authors
treated the Central American mainland spe-
cies under this name, because a similar spe-
cies had not been found on the Greater
Antilles. Only Wagner (1910a) still retained the
use of flavida for a non-existent Jamaican
species and applied the younger, synonymous
name Helicina brevilabris L. Pfeiffer, 1856 to
the mainland species.

Literature records of Helicina flavida var. for
Costa Rica (von Martens, 1890-1901) clearly
referred to H. beatrix. Therefore, the species
is newly recorded for Costa Rica, although it
remains doubtful whether it still exists in Costa
Rica or whether it was even indigenous to
Costa Rica in the first place. Having recog-
nized the peculiarity of his discovery, David
Robinson re-examined the locality in 1998 but
by then it was “totally built over by urban ex-
pansion”. His examination of some other lime-
stone spots near Puerto Limón where the
limestone may have made it possible for the
species to exist was unsuccessful (personal
communication). On the other hand, Puerto
Limón is the only Caribbean port of Costa
Rica, where the enormous trade of agricultural

CLASSIFICATION ОЕ HELICINIDAE

products actually takes place and introduction
of foreign species is thereby facilitated.
Robinson (1999) lists H. flavida among those
species that were occasionally imported into
the United States. Until better knowledge be-
comes available, the Costa Rican occurrence
of this species remains doubtful.

MISIDENTIFICATIONS FOR COSTA RICA

Helicina amoena
L. Pfeiffer, 1849

Helicina amoena — Monge-Nájera, 1997: 113:
Costa Rica [non L. Pfeiffer, 1849] refers to
Helicina pitalensis

See under Helicina pitalensis.

Helicina oweniana
L. Pfeiffer, 1849

Helicina oweniana — Monge-Nájera, 1997:
113: Costa Rica [non L. Pfeiffer, 1849] refers

86°

387

to Нейста tenuis, Н. talamancensis, Н.
детта and H. monteverdensis п. sp.

Original Description

Helicina oweniana L. Pfeiffer, 1849: 123 (not
figured); L. Pfeiffer, 1850: 40-41, pl. 7, figs.
33,30

Type Material

BMNH 20010751: 3 syntypes, leg. Mr.
Ghiesbright, Hugh Cuming coll. (in original
description “Ghiesbreght”)

The three syntypes (Fig. 276) are very simi-
lar to each other. Compared to most of the
Costa Rican species the shells are solid.
They are of whitish-yellow color with a very
slight touch of green, the lower margin of the
suture is whitish; the apex is red; only the
outer lip is bright orange. The upper whorls
are very straight; the suture very little im-
pressed, whorls regularly descending and
extending in size. Aperture oblique, outer lip

3500 -
3000 -

4000 m
3500 m

2500 -
2000 -
1500 -

3000 m
2500 m
2000 m
1000 - 1500 m

500 - 1000 m
100 - 500 m
0-100 m

e coll. IR
® coll. INBio |
© others

FIG. 275. Records of Helicina flavida in Costa Rica.

RICHLING

straight and perpendicularly and flatly ex- Helicina oweniana coccinostoma

panded, thickened, basal margin with only a Morelet, 1849

little notch. At the columella, a little groove.

Dimensions Helicina oweniana var. coccinostoma — von

8.3/7.6/8.4/7.1/5.1/5.9/6.6 mm Martens, 1900: 605-606: E-Costa Rica: Las

8.5/7.9/8.6/7.2/5.3/6.2/6.8 mm Delicias, near Santa Clara, 400 т

8.2/7.4/8.1/6.8/5.2/6.0/6.7 mm [10%57'37"N, 85°02’W, 40 m a.s.!., Alajuela
Province] (Biolley) [non Morelet, 1849] refers

Type Locality most likely to Helicina gemma

Chiapas, Mexico”. Original Description
For comments, see under Helicina tenuis, H. Helicina coccinostoma Morelet, 1849: 19
talamancensis, H. gemma, and H. monte- (not figured)

verdensis п. sp.

FIGS. 276-278. Helicina spp. FIG. 276. Syntype of Helicina oweniana, BMNH 20010751, height 8.3
mm; scale bar 2.5 mm. FIG. 277. Lectotype of Helicina coccinostoma, ВММН 1893.2.4.1605, height
8.3 тт; scale bar 2.5 mm. FIG. 278. Lectotype of Helicina anozona, ZMB 25604a, height 7.9 тт;
scale bar 2.5 mm

CLASSIFICATION OF HELICINIDAE 389

Type Material

ВММН 1893.2.4.1605-1608, Morelet coll.

“Guatemala, Peten, Palenque”
The type lot contains four specimens from
the Morelet collection, which was bought by
H. Fulton and later purchased by the ВММН.
To the shell of one specimen there was
glued a small label “type”. Because the ori-
gin of this label is uncertain and it is obvi-
ously not from Morelet himself the specimen
is herein chosen as lectotype. The
paralectotypes do not significantly differ
from the lectotype, except for one specimen
being less elevated.

The lectotype (Fig. 277) very closely re-
sembles that of Helicina anozona in shape.
lts color is lighter, yellowish opaque
throughout, except for a whitish band di-
rectly under the suture and the yellowish
outer lip.

Dimensions:
Lectotype BMNH 1893.2.4.1605

8.3/7.9/8.5/6.1/5.3/6.4/6.8 mm
Paralectotypes ВММН 1893.2.4.1606-1608

8.2/7.5/8.1/6.9/5.1/6.1/6.5 mm

7.6/7.8/8.3/6.9/5.1/5.9/5.9 mm

7.216.7/7.3/6.1/4.6/5.5/5.6 mm

Type Locality

“Petenensis sylvas” [Guatemala, Petén De-
partment].

The status of the taxon is not further dis-
cussed here, because it lies beyond the scope
of this study and requires the examination of
more comprehensive Mexican and Guatema-
lan material.

Helicina oweniana anozona
von Martens, 1875

Helicina oweniana var. anozona — Biolley,
1897: 5: Costa Rica: Tuis, 600 m [about
09°51’М, 83°35’W, Cartago Province] and
las Delicias (Santa Clara), 400 т
[10°57’37"N, 85°02’W, 40 т a.s.l., Alajuela
Province] [non von Martens, 1876] refers
most likely to Helicina gemma

Helicina oweniana var. anozona — von Mar-
tens, 1900: 605-606: E-Costa Rica: Las
Delicias, near Santa Clara, 400 т
[10°57’37"М, 85°02’W, 40 m a.s.!., Alajuela
Province] (Biolley), Tuis, 600 m [about
09°51’М, 83°35 W, Cartago Province]

(Pittier, Biolley) [non von Martens, 1876]
refers most likely to Helicina gemma

Original Description

Helicina anozona von Martens, 1875: 649 (not
figured); von Martens, 1876: 261, pl. 9, fig. 7

Type Material

Lectotype ZMB 25604a (leg. Salvin), 1

paralectotype ZMB 25604b (same data); 4
paralectotypes ZMB 40862 (same data)
(present designation); syntypes (now
paralectotypes) SNG 2192 (Zilch, 1979)
Von Martens based the description on speci-
mens collected by Salvin housed in the collec-
tion of the ZMB, where he was curator at the
time. One lot bears von Martens’ sign for
types, the larger specimen best matches the
description and is here selected as lectotype
(Fig. 278). The figure of Helicina oweniana
anozona in von Martens (1890) is based on a
specimen collected later by Champion (ZMB
103307):

The shell differs from H. oweniana in its gen-
eral shape, a more globular appearance: the
whorls are more strongly inflated, the spire is
lower, the aperture relatively larger and the
whorls are slightly shouldered. Except for the
whitish color, the features of the outer lip are
very similar.

Dimensions:
Lectotype: 7.9/8.4/8.9/7.5/5.4/6.3/6.3 mm

Type Locality

“Guatemala, vicinity of Coban” [Guatemala:
Alta Verapaz]. à

The status of the taxon is not further dis-
cussed here, because it lies beyond the scope
of this study and requires the examination of
more comprehensive Mexican and Guatema-
lan material.

Helicina fragilis
Morelet, 1851

Alcadia (Leialcadia) fragilis — Wagner, 1908:
84-85: Costa Rica: Shirores, Talamanca [in
part] [поп Morelet, 1851] refers partially to
Helicina chiquitica

See under Helicina chiquitica and Helicina
monteverdensis n. sp.

390 RICHLING

MORPHOLOGICAL CHARACTERISTICS
OF RELATED SUPRASPECIFIC TAXA OF
AMERICAN HELICINIDAE

For the discussion of the arrangement of
the Costa Rican species and the comparison
of morphological characteristics, the follow-
ing supraspecific taxa were investigated with
emphasis on the less investigated features,
for example, embryonic shell structure and
the anatomy of the female reproductive sys-
tem. When available, the respective type
species were examined, otherwise species
were chosen that are assumed to be closely
related. The taxa encompass all important
genera and subgenera, which include spe-
cies reported for the Central American main-
land and some selected representatives from
South America and the Caribbean Islands.
Unless otherwise stated, the synonymy given
by Baker (1922a) is accepted and not re-
peated. A detailed listing of the objective syn-
onyms can be found in Keen (1960). From
the following taxa, Helicina, Ceochasma,
Alcadia, Lucidella, Eutrochatella, Pyrgodo-
mus and Schasicheila are commonly recog-
nized at the generic level.

Because the following also gives an over-
view of these supraspecific taxa and their
characteristics, Succincta and “Cinctella”
were added only on the basis of annotated lit-
erature data. Furthermore, references for a
few radula descriptions were given for com-
pleteness.

For the verification of the dissections, histo-
logical sections were studied for Helicina
Jamaicensis, Alcadia hollandi, Lucidella aureola
and Eutrochatella pulchella. Similar attempts
for Helicina brasiliensis and Schasicheila alata
proved to be only partially successful due to the
poor condition of the old material.

FIG. 279. Helicina neritella, IR 3454, height 9.8
mm; scale bar 5 mm.

FIG. 280. Teleoconch surface structure of
Helicina neritella, IR 3857. A. Structure of apical
part. B. Pattern of oblique diverging grooves on
the 1° whorl; scale bar 500 um (A), 100 um (В).

CLASSIFICATION OF HELICINIDAE 391

Helicina Lamarck, 1799
Type species
Helicina neritella Lamarck, 1799
Investigated Species
Helicina neritella (Figs. 279, 340A)
Material

Jamaica: Manchester Parish, Mandeville,
Caledonia Road, 18°02’11"М, 77*30'44"W,
600 m a.s.l., 28./29.05.2001 (IR 3454, IR
3459); St. Ann Parish, N Ocho Rios, Fern
Gully, 310 m a.s.l., 04.06.2001 (IR 3857),
leg. W. Böckeler & I. Richling

Morphological Characteristics

Teleoconch Surface Structure (Fig. 280): The
first half whorl is sculptured with the transi-
tional pattern, which is followed by strongly
developed oblique diverging grooves. This
pattern is maintained throughout the
teleoconch.

FIG. 281. Embryonic shell of Helicina neritella, IR
3459; scale bar 100 um.

7
=

FIG. 282. Female reproductive system of
Helicina neritella, apical complex enlarged, IR
3454; scale bars 2.5 mm (left), 1 mm (right).

Embryonic Shell (Fig. 281): Densely struc-

tured with pits arranged in concentric lines,
pits comparatively large. Diameter about
728 um (п = 3), a little smaller than given by
Thompson (1982), other structures equal to
his description.

Radula: Figured in Baker (1922a: pl. Ill, fig. 6,

pl. IV, fig. 17).

Female Reproductive System (Fig. 282): As-

cending limb of the V-organ elongated,
straight, in natural position overlapping with
the posterior part of the pallial oviduct. The
latter in relation to the apical complex re-
markably long, transversally constricted.
Small, oblong receptaculum seminis enter-
ing inner side of the descending limb. Bursa
copulatrix very prominent and elongated
with numerous, densely arranged, rather
small lobules. Provaginal sac appearing
vestigial, long and slender, only slightly de-
marcated from its much elongated and par-
tially coiled stalk. Provaginal opening
absent.

392 RICHLING

FIG. 283. Helicina platychila, UF 259486, height
6.5 mm; scale bar 5 mm.
Investigated Species

Helicina platychila (von Mühlfeldt, 1816) (Figs.
283, 340B)

Material

Dominica: along trail '/, mi. W of Trafalgar FIG. 285. Female reproductive system of
Falls, upper end of banana plantation, leg. J. Helicina platychila, right figure: dorsal view, UF
P. E. Morrison (JPEM-2610), 11.10.1965 259486; scale bar 1 mm.

(UF 259486)

Morphological Characteristics

Embryonic Shell (Fig. 284): Similar pattern as
in Helicina neritella, but pits smaller and
more numerous. Diameter 890 pm.

Female Reproductive System (Fig. 285): Very
similar to that of Helicina neritella. Ascend-
ing limb of V-organ slightly curved; pallial
oviduct relatively shorter and stronger
folded. Bursa copulatrix prominent, with less
numerous lobes, but the latter larger and
elongated. Provaginal sac small, roundly tri-
angular, stalk slightly curved and relatively
long. Provaginal opening absent.

FIG. 284. Embryonic shell of Helicina platychila, FIG. 286. Helicina orbiculata, IR 3361, height 6.5
UF 259486; scale bar 100 um. mm; scale bar 2.5 mm.

CLASSIFICATION ОЕ HELICINIDAE

Oligyra Say, 1818
Type species
Olygyra orbiculata Say, 1818
Investigated Species
Helicina orbiculata (Figs. 286, 340C)
Material

USA: Florida, Gainesville, Museum Road,
near Dickinson Hall, 05.2001, leg. J.
Slapcinsky 4 |. Richling (IR 3359, IR 3361)

Morphological Characteristics

Teleoconch Surface Structure (Fig. 287):
Transitional pattern for about */, whorl, sub-
sequently sculptured with oblique diverging
grooves up to the aperture. Furthermore
widely spaced, slightly impressed spiral
grooves with periostracal ridges.

Embryonic Shell (Fig. 288): Scarcely sculp-
tured with pits arranged in concentric lines.
Interspaces of lines and pits exceed the size
of the pits. Occasionally more densely pitted
(Fig. 288B). Diameter 835 um (n = 4).

Radula (Fig. 289): All centrals with cusps, A-
central about 3-4, B-central about 5-6, C-
central about 2-4; comb-lateral with 7-8
cusps. Cusps of the marginals rather slowly
increasing in number, cusps more distally
than laterally arranged. Except for minor
deviations, this is in agreement with Baker
(1922a).

Female Reproductive System (Fig. 290): V-
organ rather stout, pallial oviduct moderately
constricted. Receptaculum seminis large,
bulbous, entering at inner side of descend-
ing limb of V-organ. Bursa copulatrix rather
small and weakly lobed, of about the same
size as elongated provaginal sac, which is
slightly constricted at the distal side, its stalk
is short. Provaginal opening absent. The fe-
male system was studied and figured by
Baker (1926) (Fig. 9). Except for the
monaulic (instead of diaulic) conditions, the
structures were confirmed. The bursa
copulatrix differs in the length of its lobes in
the specimens of all three locations (Baker:
Miami County, Florida, and San Antonio,

FIG. 287. Teleoconch surface structure of
Helicina orbiculata, IR 3859. A. Structure of
apical part. B. Pattern of oblique diverging
grooves and spiral grooves with periostracal
ridges on the begin of the 2" and 3 whorl; scale
bars 500 um (A), 100 um (B).

4 RICHLING

i=

(411,48

FIG. 288. Embryonic shell of Helicina orbiculata. FIG. 289. Radula of Helicina orbiculata, IR 3359.
A. IR 3359. B. IR 3361; scale bar 100 um. A. Centrals. B. Comb-lateral. C. Marginals; scale
bar 50 um.

CLASSIFICATION OF HELICINIDAE 395

FIG. 290. Female reproductive system of
Helicina orbiculata, IR 3361; scale bar 1 mm.

Texas), the presently studied specimens
have furthermore a larger receptaculum
seminis. Baker (1926) treated the speci-
mens from Texas as the subspecies Helicina
orbiculata tropica Jan, 1846, but a recent
electrophoretical investigation by Strenth &
Littleton (2000) suggests the conspecificity
of both taxa, which has been discussed re-
peatedly.

Succincta Baker, 1922
Type species
Helicina succincta Martens, 1890
Investigated Species

Helicina succincta (investigated by Baker,
1928)

Morphological Characteristics
Radula: Figured in Baker (1928: pl. IV, fig. 26).

Female Reproductive System: Figured in
Baker (1928: pl. Il, figs. 3, 4): Receptacu-
lum seminis remarkably enlarged and
slightly trilobed; bursa copulatrix reduced in
size and only bilobed; provaginal sac well
developed and distally lobed, stalk quite
long. According to the drawing and the re-
sults for re-examined mainland species, the
provaginal opening is here assumed to be
absent.

FIG. 291. Helicina turbinata, Cordova, ZMB
103315, height 11.7 mm; scale bar 5 mm.

Tristramia Crosse, 1863
Type species
Helicina salvini Tristram, 1861
Investigated Species

Helicina turbinata Wiegmann, 1831 (Figs. 291,
340D)

Material

Mexico: Arisolapa [?], leg. Strebel, #1723
(ZMH 2932)

FIG. 292. Embryonic shell of Helicina turbinata,
ZMH 2932; scale bar 100 um.

RICHLING

lorphological Characteristics

Embryonic Shell (Fig. 292): The surface is con-
centrically pitted, but due to the very small
size of the widely spaced pits, the surface
appears nearly smooth. Diameter 765 um.

Radula: Figured in Baker for the closely related
or synonymous species (von Martens, 1890-
1901; Baker, 1922a) Helicina zephyrina Men-
ke, 1830 (1922a: pl. Ш, fig. 9, pl. М, fig. 13).

Female Reproductive System (Fig. 293): V-
organ normally developed; pallial oviduct
transversally and partially also longitudinally
constricted. Receptaculum seminis very
large and bulbous, entering at the inner side
of the descending limb of the V-organ, but,
due to its unusual size, shifted dorsally.
Bursa copulatrix prominent and deeply
lobed; provaginal sac with a rather slender
stalk (not visible in Fig. 293), large and
slightly constricted at its distal side. The
provaginal opening is absent. Except for the
erroneously assumed provaginal opening
the description by Baker (1928) for Helicina
zephyrina is identical, especially with re-

KT ) f P

? |) д já

> "TT SES
Pr di y y Au de

Y à. |
тут SE
LEE ЕЯ . # 3
FIG. 293. Female reproductive system of

Helicina turbinata, right figure: dorsal view, (on
account of the poor preservation kept in nearly
natural position) ZMH 2932; scale bar 1 mm.

FIG. 294. Helicina amoena, ZMB 103345, height
10.6 mm; scale bar 5 mm.

spect to the shape of the bursa copulatrix
and the enlarged receptaculum seminis.

Oxyrhombus Crosse & Fischer, 1893
Type species
Helicina атоепа L. Pfeiffer, 1849
Investigated Species
Helicina amoena (Figs. 294, 340E)
Material
Guatemala: Teleman (ZMB 103345)
Morphological Characteristics

Embryonic Shell: The structure is similar to
those shown for Helicina funcki and H.
pitalensis (Figs. 15, 43): relatively large pits
in concentric lines with their diameter about
equal to their interspacial distance. Within
the material available, large parts of the
embryonic shell were so badly eroded so
that it did not seem to be worth figuring. Di-
ameter 860 um.

Radula: Figured in Baker (1922a: pl. Ill, fig. 8,
pl. IV, fig. 15).

Female Reproductive System (Fig. 295): In
general similar to that of Helicina turbinata,
but receptaculum seminis much smaller,
bursa copulatrix with an elongated central
axis from which numerous, further subdi-
vided lobes branch off. Provaginal sac much
more flattened and more strongly irregularly
lobed at the distal side, its stalk of moderate
length (not clearly visible in Fig. 295).
Provaginal opening absent.

CLASSIFICATION OF HELICINIDAE 397

FIG. 295. Female reproductive system of
Helicina amoena, right figure: dorsal view, distal

parts omitted, ZMB 103345; scale bar 1 mm.

Pseudoligyra Baker, 1954
Synonym (objective)
Tenuis Baker, 1922, non Barrande, 1881
Type species
Helicina tenuis Pfeiffer, 1849
Investigated Species
Helicina tenuis — see above.
Punctisulcata Baker, 1922
Type species
Helicina punctisulcata von Martens, 1890

Investigated Species

Helicina punctisulcata cuericiensis п. subsp. —

see above.

“Cinctella” Baker, 1922,
non Monterosato, 1884

Type species
Helicina cinctella Shuttleworth, 1852

Investigated Species

Helicina cinctella (investigated by Baker, 1928)

Morphological Characteristics

Female Reproductive System: Figured in
Baker (1928: pl. Il, fig. 5): Receptaculum of
normal size; bursa copulatrix prominent and
having several further subdivided lobes;
provaginal sac weakly lobed at distal side,
flattened. The provaginal opening is shown
in the figure, but its existence is very ques-
tionable. Here it is assumed to be absent.

“Gemma” Baker, 1922,
non Deshayes, 1853

Type species

Helicina gemma Preston, 1903

Investigated Species

Helicina gemma — see above.
Tamsiana Baker, 1922

Type species

Helicina tamsiana Pfeiffer, 1851

Investigated Species

Helicina tamsiana

Material

Venezuela: Porto Cabello, leg. Martin (ZMB
103314)

Morphological Characteristics

Embryonic Shell: Only a single specimen was
studied. It remains uncertain whether the
embryonic shell is partially eroded or whether
the surface is rather smooth, except for very
scarce pits and very slight oblique lines.

Radula: Figured in Baker (1923: p. 20, fig. 20).

Female Reproductive System: Described and
figured by Baker (1923: pl. VI, fig. 14): Re-
ceptaculum seminis of normal size; bursa
copulatrix reduced to a very small, simple
sac; provaginal sac very prominent and an-
teriorly elongated, so that the stalk branches
off about the middle of its long side.
Provaginal opening is shown in the figure, its
existence remains questionable.

398 RICHLING

Trinity Road & Guayaguayare Road
(Rushville), 10°07’32"М, 61°03'28"\М, 12 т
a.s.l., leg. К. Auffenberg et al. (КА-1212),
02.06.1994 (UF 226928)

Venezuela: Margarita Island, Porlamar, just
outside town under trees, ex Guido Poppe,
1988 (HNC 54607)

Morphological Characteristics

FIG. 296. Helicina dysoni, UF 226928, height 5.3 Embryonic Shell (Fig. 297): Surface with a few
mm; scale bar 2.5 mm. concentric rows of small, widely spaced pits.
Diameter 590 um.

Analcadia Wagner, 1908 Radula: Figured by Baker (1923: pl. Ш, fig. 12).

Type species Female Reproductive System (Fig. 298):
Limbs of V-organ rather short, small re-
Helicina dysoni Pfeiffer, 1849 ceptaculum seminis entering at inner side
of descending limb. Bursa copulatrix repre-
Investigated Species senting a very small sac, its connection to
the reception chamber remarkably shifted
Helicina dysoni (Figs. 296, 340F) towards the dorsal side compared to all
other species described in this study.
Material Provaginal sac exceptional large and in-

flated, slightly irregularly constricted at its
Trinidad & Tobago: Trinidad Island, Mayaro distal side; stalk short, stout and branching
County, Trinity District, 0.6 km SW junction off at the middle of the sac. Directly at junc-

| FIG. 298. Female reproductive system of
FIG. 297. Embryonic shell of Helicina dysoni, Helicina dysoni, right figure: dorsal view, UF
HNC 54607; scale bar 100 um. 226928; scale bar 1 mm.

CLASSIFICATION OF HELICINIDAE 399

tion with the reception chamber is an ante-
rior sac-shaped appendage. Additional sac
present at ventral side of the pallial oviduct
which seems to be connected to it (remains
to be checked histologically). Provaginal
opening absent.

Sericea Wagner, 1907
Type species
Helicina sericea Drouet, 1859
Investigated Species
Helicina sericea (Figs. 299, 340G)
Material

Suriname: District Suriname, Boden-
savanne [?], mine synagoge, leg. C.O. van
Regteren Altena (loc. 51), 14.03.1963
(RMNH 8890)

Morphological Characteristics

Embryonic Shell (Fig. 300): Similar to that of
Helicina dysoni. Diameter 755 pm.

Female Reproductive System (Fig. 301): V-
organ and receptaculum seminis similar to
Helicina dysoni. Bursa copulatrix of moder-
ate size and subdivided in numerous short
lobules. Provaginal sac exceptional large
and inflated, nearly kidney-shaped, with its
short and stout stalk branching off at about
the middle of the proximal side. It bears an
anterior basal appendage at the junction
with its stalk, which is roundly constricted
twice. Provaginal opening absent.

FIG. 299. Helicina sericea, RMNH 8890, height
5.1mm; scale bar 2.5 mm.

FIG. 300. Embryonic shell of Helicina sericea,
RMNH 8890; scale bar 100 um.

y A
2 A E
/ f {
4
f N u‘ y
| | k
| | RS
\ \ À ©”
TAS \\ \ — er )
Ay y AU \ ae |
x | Ir
Ре. я ) } / | A
\ = J }
à \
/ ad) =
<< : =
Е oat =
Me
| А МА /
> F я / N J
| ( = ( Y
) \ 4
IN _#
\ -
—=< \
\
Sy
\
=“ = A /
ча ri a NE
EN Le E,
( yf
ОД

FIG. 301. Female reproductive system of Не/ста
sericea; upper, right figure: dorsal view, RMNH
8890; scale bar 1 mm.

400 RICHLING

Ceochasma Thompson, 1968
Type species
Ceochasma phrixina Thompson, 1968
Investigated Species
Ceochasma phrixina
Material
Mexico: Colima, 0.3 km. SE Tamala, 152 m
a.s.l., leg. Е.С. Thompson (FGT-777),
02.08.1966 (UF 20139, Paratypes)
Morphological Characteristics
Embryonic Shell: Not examined.
Radula: Figured in Thompson (1968: 49).
Female Reproductive System: Described and
figured by Thompson (1968: 49), for this study
only reinvestigated with respect to provaginal
opening, which was found to be absent.
Angulata Baker, 1922
Type species
Helicina angulata Sowerby, 1842

Investigated Species

Helicina brasiliensis Gray, 1825 (Figs. 302,
340H)

Material

Brazil: Sta. Catharina, Humboldt District,
Joinville, Flussgebiet von Itaporu [area of

FIG. 302. Helicina brasiliensis, ZMH 2931, height
5.6 mm; scale bar 2.5 mm.

FIG. 303. shell of Helicina

Embryonic
brasiliensis, ZMH 2931; scale bar 100 pm.

the Rio Iguacu], leg. W. Ehrhardt, purchased
21.10.1910 (ZMH 2931)

Morphological Characteristics

Embryonic Shell (Fig. 303): Sculptured with
regular, broad, very slightly raised spiral
bands; interspacial distance smaller than the
width of these bands, otherwise smooth. Di-
ameter 695 um (n = 2).

Female Reproductive System (Fig. 304): Recep-
taculum seminis very small, connected to the
inner side of the descending limb of the V-or-
gan; bursa copulatrix large and deeply lobed.
Provaginal sac elongated and of somewhat
irregular outline, its stalk rather stout.
Provaginal duct seems to branch off from this
stalk at its most anterior point. Due to the poor
preservation of the material, the provaginal
duct and opening could not be observed with
certainty, but its presence 1$ very likely.

Alcadia Gray, 1840
Type species

Helicina major Gray, 1824

CLASSIFICATION ОЕ HELICINIDAE 401

irregularly spaced radial threads. Diameter
1800 pm. The embryonic shell had also
been studied and figured by Thompson
(1982: fig. 26), his specimen shows the ob-
lique grooves within the inner curvature, the
diameter is given with 1.0 mm, but a теа-
surement of the figure reveals a more likely
size of about 2.2 mm.

Radula (Fig. 307): The centrals completely
lack cusps and the cutting edges are rein-
forced. The “comb”-lateral agrees in its
rough outline rather with the denticulated
part of the comb-lateral in Helicina, but the
cutting edge is smooth and thickened and
resembles the T-shaped lateral of
Eutrochatella (vianid radula). The accessory
plate seems to be reduced. The tips of the
marginals are rounded and show minor

FIG. 304. Female reproductive system of Helicina crenulations.

brasiliensis, right figure: dorsal view, (distal parts The vianid condition of the radula has al-

omitted, provaginal duct and opening likely, but ready been mentioned by Boss & Jacobson

not verified) ZMH 2931; scale bar 0.5 mm. (1973) and Thompson (1982), but it has
never been figured and described in detail
before.

Investigated Species
Female Reproductive System (Fig. 308): V-
Alcadia major (Figs. 305, 3401) organ with a slight apical swelling, its as-

Material

Jamaica: Manchester Parish, Silver Grove,
Secondary forest & bordering pasture, lime-
stone & red soil, 18°04.95’М, 77°35.35'W,
880-900 m a.s.l., leg. С. Rosenberg & 1.V.
Muratov (JBS 113), 02.10.1999 (ANSP
19559)

Morphological Characteristics
Embryonic Shell (Fig. 306): Surface with more

or less strong oblique grooves, more pro-
nounced towards the margin, and coarse,

FIG. 305. Alcadia major, ANSP 19559, height FIG. 306. Embryonic shell of Alcadia major,
14.2 mm; scale bar 10 mm. ANSP 19559; scale bar 100 um.

402 RICHLING

Ly [Fp

FIG. 308. Female reproductive system of Alcadia
major, ANSP 19559; scale bar 2.5 mm.

cending limb slightly elongated. Receptacu-
lum seminis equal to Helicina on the inner
side of the descending limb, pedicel well
developed. Bursa copulatrix representing a
very large, irregularly shaped sac that
broadly enters the reception chamber. Close
to this connection is a slender provaginal
duct that extends up to its opening for about
'/, of the length of the pallial oviduct. Close
to the reception chamber, the provaginal
duct receives the stalk of the medium-sized,
oblong provaginal sac. The specimen is-not
very well preserved and perhaps the visible
broad connection of the bursa copulatrix
does not reflect the natural condition nor
does the weakly demarcated distal end of
the reception chamber.

FIG. 307. Radula of Alcadia major, IR 3359. A. FIG. 309. Alcadia hollandi, IR 3579, height 7.4
Centrals. B. Comb-lateral. C. Marginals; scale mm; scale bar 5 mm.
bars 50 um (А, В), 100 um (С).

CLASSIFICATION OF HELICINIDAE 403

Palliata Baker, 1922
Type species
Helicina palliata С. В. Adams, 1849
Investigated Species

Alcadia hollandi (C. B. Adams, 1849) (Figs.
309, 340J)

Material

Jamaica: Manchester Parish, Mandeville, 600
т a.s.l., Marshall's Drive, 25./27.05.2001,
leg. W. Bôckeler & |. Richling (IR 3579)

Morphological Characteristics

Embryonic Shell (Fig. 310): Inner curvature
with predominant irregular axial threads, to-
wards the margin interposing with strong,
very distinct oblique grooves. Diameter 840
um (n = 2).

Radula: Figured in Bourne (1911: pl. XL, fig.
56). Similar to Helicina: with denticulated

FIG. 310. Embryonic shell of A/cadia hollandi, IR
3579; scale bar 100 um.

hollandi
explanations modified).

FIG. 311. Female reproductive system of Alcadia
hollandi, IR 3579; scale bar 1 mm.

centrals and marginals, comb-lateral with
cusps and accessory plate.

Female Reproductive System (Fig. 311): As-
cending limb of V-organ shorter than in
Alcadia major, receptaculum seminis small,
located on the inner side of descending limb.

bursa copulatrix
ascending limb \

receptaculum \_\ У N

seminis TR A primary oviduct
descending limb “\. \\4, \ —ргомадта! sac
of V-organ À À

| provaginal duct
AN and vagina

pallial oviduct

intestine

N
EN

an A

AS y
cloaca +

FIG. 312. Female reproductive system of A/cadia
(reproduced from Bourne, 1911,

404 RICHLING

FIG. 313. Асада jamaicensis, IR 3502, height
8.3 mm; scale bar 5 mm.

Bursa copulatrix representing an oblong sac,
longitudinally folded inside. Provaginal sac
rather small and at the end of the very long
stout stalk or itself stalk-like elongated, en-
tering the reception chamber at nearly the
same point as the bursa copulatrix and the
provaginal duct. The latter is long and slen-
der and opens at about '/, of the way from
the beginning of the pallial oviduct. The fe-
male system was figured by Bourne (1911:
pl. XXXV, fig. 25, reproduced here: Fig.
312), but Baker (1926) questioned its cor-
rectness. Bourne's description is verified by

FIG. 314. Embryonic shell of Alcadia jamaicensis,
IR 3574; scale bar 100 pm.

the present investigation, only that sperm
had not been found within the pallial oviduct,
probably due to different physiological con-
ditions.

Investigated Species

Helicina jamaicensis Sowerby, 1841 (Figs.
313, 340K) (belongs to Alcadia, an assign-
ment to a subgenus, e.g., Palliata, is here
not intended)

Material

Jamaica: Manchester Parish, W Bellefield,
valley near road to Banana Ground,
18°04’46" М, 77°26'27"W, 580 m a.s.l.,
24.05.2001 (IR 3502); Manchester Parish,
Mandeville, 600 m а.5$.1., Marshall's Drive,
25./27.05.2001 (IR 3574), leg. W. Bóckeler
8 |. Richling

Morphological Characteristics

Embryonic Shell (Fig. 314): Very similar to
Alcadia hollandi. Diameter 920 um.

IES

FIG. 315. Female reproductive system of A/cadia

jamaicensis, left figure: natural position, right fig-

ure: slightly lateral view, IR 3502; scale bar 1 mm.

CLASSIFICATION OF HELICINIDAE 405

Female Reproductive System (Fig. 315): V-
organ similar to A/cadia hollandi; recep-
taculum seminis larger. Bursa copulatrix,
provaginal sac and provaginal duct very
closely associated. Bursa copulatrix more
prominent than in A/cadia hollandi, its po-
sition is similar. Provaginal sac of nearly
the same size as the bursa copulatrix,
forming an elongated sac with a short,
stout stalk, which rather connects with the
provaginal duct than with the reception
chamber. Provaginal duct slightly inflated
shortly before its opening at about */, of
the way from the beginning of the pallial
oviduct. The latter only with minor constric-
tions.

Idesa H. Adams 8 A. Adams, 1856
Synonym (objective)

Leialcadia Wagner, 1907

Type species

Helicina rotunda Orbigny, 1841 FIG. 317. Embryonic shell of Alcadia rotunda,
ZMB 90412; scale bar 100 um.
Investigated Species

Alcadia rotunda (Figs. 316, 340L) Radula: Figured by Troschel (1856-1863: pl. V,
figs. 10, 11) and Baker (1923: pl. Ш, fig. 13).
Material Centrals and marginals with cusps, comb-lat-

eral denticulated and with accessory plate.

Cuba: Pinar del Rio, Rangel, leg. M.L. Jaume

(ZMB 90412) Female Reproductive System: unknown.
Morphological Characteristics Microalcadia Richling, n. subgen.
Embryonic Shell (Fig. 317): Sculptured with Type species

fine, oblique diverging grooves. Diameter

690 um. Helicina hojarasca Richling, 2001
Investigated Species
Alcadia hojarasca — see above.

Eutrochatella Fischer, 1885

Type species

Helicina pulchella Gray, 1825

Investigated Species

FIG. 316. Alcadia rotunda, ZMB 90412; scale bar |
2.5 mm. Eutrochatella pulchella (Figs. 318, 340M)

406 RICHLING

FIG. 318. Eutrochatella pulchella, IR 3702,
height 7.9 mm; scale bar 5 mm.

Material

Jamaica: Manchester Parish, W Bellefield, val-
ley near road to Banana Ground, 18°04’46"
М, 77°26'27"W, 580 m a.s.l., 24.05.2001 (IR
3504); Trelawny Parish, near Burnt Hill, along
road to Clarks Town, 18°18'24" N,
77°33’46"W, 510 m a.s.l., 02.06.2001 (IR
3808), leg. W. Böckeler & I. Richling

Morphological Characteristics

Embryonic Shell (Fig. 319): Surface rough,
somewhat irregularly wrinkled. These

Be ME RG og
a = nn ERW, 7.
«

es yg Ove à
> AE fs
` Y

oN

FIG. 319. Embryonic shell of Eutrochatella
pulchella, IR 3505; scale bar 100 um.

coarse ridges and grooves show an orienta-
tion similar to the grooves in A/cadia. Diam-
eter 560 um (n = 3).

Radula: Figured in Bourne (1911: pl. XL, fig.

57) and Baker (1922a: pl. VI, figs. 31-32). All
teeth without cusps, except for the outer-
most marginals, lateral reinforced and T-
shaped, accessory plate reduced.

Female Reproductive System (Fig. 320): V-

organ of moderate size; receptaculum
seminis small and located at the inner side
of the descending limb. Bursa copulatrix
large, oblong, externally not further subdi-
vided, broadly connected with the reception
chamber. Provaginal sac smaller than bursa
copulatrix and flattened, lobed at the distal
side, its stalk short and slender. The transi-
tion of the reception chamber to the pallial
oviduct is externally only weakly visible; fur-
thermore, the pallial oviduct of different in-
vestigated specimens was remarkably less
thickened than in the species of Helicina for
example, although the specimens were all
mature. The provaginal opening 1$ absent.
Bourne (1911: pl. XXXV, fig. 26) did not rec-
ognize the monaulic condition; furthermore,
his figure shows a larger bursa copulatrix.

|

>

NS

ns
/

FIG. 320. Female reproductive system of
Eutrochatella pulchella, IR 3808; scale bar 1 mm.

CLASSIFICATION OF HELICINIDAE 407

Embryonic Shell (Fig. 323): Sculptured with
numerous small pits, which are less regu-
larly arranged than in Helicina. Their diam-
eter is smaller than the interspacial distance.
Diameter: 515 pm (n = 3). This is in full
agreement with the description by Thomp-
son (1982: figs. 24, 25).

Radula: Figured and described by Bourne
(1911: pl. XL, fig. 59), Baker (1922a: pl. Ш,
FIG. 321. Lucidella aureola, IR 3852, height fig. 4, pl. V, fig. 22), and Thompson (1982:

5.1тт; scale bar 2.5 mm. figs. 14, 15).
Female Reproductive System (Fig. 324): V-
Pyrgodomus Crosse 8 Fischer, 1893 organ with a left-sided apical swelling, limbs
rather short. Receptaculum seminis absent.
Type species Bursa copulatrix small, simple, broadly con-
nected with the reception chamber.
Helicina chryseis Tristram, 1861 Provaginal sac large, distal side remarkably
lobed, stalk branching off at about the
Investigated Species middle, short and stout. Provaginal duct
short, joining reception chamber nearly to-
Pyrgodomus microdinus — see above. gether with the stalk of the provaginal sac
and the bursa copulatrix, opening at about
Lucidella Swainson, 1840 distal end of the reception chamber. Poste-
rior part (about % to '/.) of pallial oviduct
Type species largely inflated and with numerous internal,

longitudinal folds, at the distal end of this

Helix aureola Férussac, 1822

Investigated Species

Lucidella aureola (Figs. 321, 340N)

Material

Jamaica: Manchester Parish, Mandeville, 600
m a.s.l., Marshall’s Drive, 25./27.05.2001 (IR
3578); St. Ann Parish: N Ocho Rios, Fern
Gully, 310 m a.s.l., 04.06.2001 (IR 3852),
leg. W. Bóckeler & |. Richling

Morphological Characteristics

Internal Shell Structures: (Fig. 322)

FIG. 322. Axial cleft and muscle attachments of FIG. 323. Embryonic shell of Lucidella aureola,
Lucidella aureola, IR 3578; scale bar 2.5 mm. IR 3852; scale bar 100 um.

RICHLING

| \

&

SIA

FIG. 324. Female reproductive system of
Lucidella aureola, IR 3852; scale bar 0.5 mm.

structure enters an additional, long sac. In
both structures, sperm were found. Baker's
(1926: pl. VII. fig. 19) description based on
badly macerated specimens has to be cor-
rected with respect to the absence of the
receptaculum seminis and the additional
structures of the pallial oviduct.

Perenna Guppy, 1867
Type species
Helicina lamellosa Guppy, 1867
Investigated Species

Lucidella lirata — see above.

FIG. 325. Schasicheila alata, UF 251373, height
8.4 mm; scale bar 5 mm.

FIG. 326. Axial cleft and right muscle attachment of
Schasicheila alata, ZMH 2928; scale bar 2.5 mm.

Schasicheila Shuttleworth, 1852
Type species
Helicina alata Pfeiffer, 1848
Investigated Species
Schasicheila alata (Figs. 325, 3400)
Material

Mexico: Agua Caliente, leg. Strebel (#1725)
(ZMH 2928); Veracruz, 2.7 mi S Orizaba,
3,800 feet, leg. M.L. Paulson et al.,
12.08.1965 (UF 251373)

Morphological Characteristics
Internal Shell Structures: (Fig. 326)

Embryonic Shell (Fig. 327): Sculptured with
very regular and prominently raised axial
folds. Diameter 1015 pm.

Radula: Figured in Baker (1928: pl. V, fig. 27).

Female Reproductive System (Fig. 328): Both
limbs of the V-organ remarkably elongated
and curved, a receptaculum seminis devel-
oped as in Helicina, Alcadia and
Eutrochatella is absent, but in all dissected
specimens an accumulation of sperm was
found within the oviduct proximal to the
weekly separated pedicel. A bursa copulatrix
similar to other genera is not present; instead,
a large, strongly recurved sac, which seems
to be fused with the pedicel extents dorsal to
the V-organ, is present. The provaginal sac is
rather small and flattened, slightly lobed at
the distal side; its stalk is very short.
Provaginal duct also very short, opening at

CLASSIFICATION OF HELICINIDAE 409

FIG. 327. Embryonic shell of Schasicheila alata,
ZMH 2928; scale bar 100 pm.

about the beginning of the pallial oviduct. The
investigation confirms the description given
by Baker (1928: pl. IV, figs. 19, 20).

/ se I )
/ Gl p=
© le 3

FIG. 328. Female reproductive system of
Schasicheila alata, V-organ turned to the left, UF
251373; scale bar 1 mm.

DISCUSSION
Knowledge of Costa Rican Helicinidae

Previous to this study and Richling (2001),
six correctly identified species and one cor-
rectly identified subspecies (Helicina funcki, Н.
pitalensis, Н. tenuis, H. beatrix beatrix, Н.
beatrix confusa, H. gemma, Lucidella lirata) of
Helicinidae were known in Costa Rica. Addi-
tionally, a species of “Pyrgodomus” had tenta-
tively been listed. The present investigation
adds seven new species (H. escondida п. sp.,
Н. echandiensis п. sp., H. talamancensis, Н.
monteverdensis п. sp., H. chiquitica, Alcadia
hojarasca, A. boeckeleri) and two new sub-
species (Helicina punctisulcata cuericiensis п.
subsp., H. beatrix riopejensis п. subsp.). п
addition, previously separated subspecies of
Helicina funcki and H. tenuis were shown to
have fallen within the range of intraspecific
variability. The new and verified record of
Helicina flavida remains doubtful in its inter-
pretation. Furthermore, the re-examination of
original material and records or their critical
consideration demonstrated the absence of
the Mexican and Guatemalan species Helicina
amoena, H. oweniana and subspecies and H.
fragilis from the Costa Rican fauna.

On one hand, the remarkable number of
new species reflects the vague knowledge
about the discrimination of the described taxa
subsequent to the well founded major contri-
butions at the end of the 19" century (e.g., von
Martens, 1890-1901; Fischer & Crosse,
1880-1902), that is, the necessity of the ex-
amination of the type material. On the other
hand, the recent discovery of strikingly differ-
ent species (е.д., Helicina echandiensis п. sp.)
in remote areas or the small species Alcadia
hojarasca and A. boeckeleri dwelling in leaf
litter illustrates the deficits in the inventory of
the fauna, as well the difficulties in finding
specimens at all, be it due to the very low
abundance or to locally restricted ranges.

Distribution of Costa Rican Species and
Faunal Composition

The knowledge of the distribution is limited
by the insufficient investigation of the mollus-
can fauna of the adjacent areas Nicaragua
and Panama. Despite considerable collecting
efforts, information remains fragmentary for
several species within Costa Rica itself. Nev-
ertheless, some general aspects emerge. In
the following, “southern Central America” will

refer to the area from the Nicaraguan depres-
sion to about the Canal Zone.

The Costa Rican helicinid fauna is com-
posed of the following elements:

(1) widespread species: Helicina tenuis,
Lucidella lirata, Pyrgodomus microdinus
(2) species limited to southern Central
America: Helicina funcki, H. pitalensis, H.
beatrix, H. talamancensis, H. gemma, H.
monteverdensis n. sp., H. escondida n.
sp., H. chiquitica
species occurring very locally: Helicina
punctisulcata cuericiensis п. subsp., Н.
echandiensis п. sp., Alcadia hojarasca”, A.
boeckeleri”
distributional pattern assumed for the taxon be-
cause affinities doubtful
distribution too poorly known to be discussed fur-
ther.

The list shows that most species are endemic
to southern Central America. This fact greatly
changes the previous idea about the faunal
composition with more widely spread and less
endemic species due to the exclusion of three
misidentified Mexican/Guatemalan species and
the recognition of several new taxa.

RICHLING

Considering the distribution of these groups
within Costa Rica, it becomes obvious that
only the widely spread species occur on the
Pacific as well as on the Caribbean side.
Pyrgodomus microdinus represents an excep-
tion, because its distribution is mainly influ-
enced by its strict limitation to calcareous
outcrops. The other species can be further
subdivided by their restriction to the:

Caribbean side: Helicina funcki, H. beatrix, H.
gemma, H. chiquitica, H. monteverdensis n.
sp., H. escondida n. sp.

Pacific side: Helicina pitalensis, H. tala-
mancensis.

Mountain region: Helicina echandiensis n.
sp., H. punctisulcata cuericiensis n. subsp.

Only the Caribbean species normally cross
the continental divide in the Cordillera de
Tilaran (e.g., Monteverde) and the Cordillera
de Guanacaste, disappearing towards the
Pacific plains. The Pacific species are re-
stricted to the southern area and to the north-
ern foothills of the Cordillera de Talamanca.
The Peninsula de Nicoya and the central and
northern plains and foothills of the Pacific side
are virtually uninhabited by any species of the

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e all other species
o Helicina tenuis

FIG. 329. Costa Rican records of Helicinidae mapped on the annual precipitation
[mm/year], Helicina tenuis individually marked.

CLASSIFICATION OF HELICINIDAE

Helicinidae, except for Helicina tenuis. This is
clearly related to the drier climate.

In conclusion, it can be stated that whereas
in southern Costa Rica the continuously highly
elevated mountain chain of the Cordillera de
Talamanca obviously represents a barrier for
the distribution of species; it is replaced more to
the north by a dry belt along the northern plains
and foothills up to the Valle Central.

The maps (Figs. 329, 330) show the locations
of all records of Helicinidae in Costa Rica
(Helicina tenuis differently marked) mapped on
the distribution of the vegetation or the annual
precipitation respectively (Ministerio de Agri-
cultura y Ganaderia & Instituto Meteorologico
Nacional, 1985). The meteorological data re-
flect the amount of annual rain and also provide
a rough estimation of the real humidity avail-
able for the fauna, but do not reflect the strong
seasonal changes during the dry period in the
northwestern and central parts of the country.
These conditions are much better represented
by the vegetation, here given in a simplified
map (modified after Tosi, 1969), graduated
solely according to the humidity-related type of
vegetation. As may be expected, the vegetation

411

map actually matches the distribution of the
snails in the northwestern part much better. The
single dot in the dry forest area (or < 1,500 mm/
year rain-area) belongs to an old, subsequently
localized record of Helicina funcki (“10 mi W of
Tilarán”). If it is really correctly plotted, the
specimen may also have come from a more
humid river valley. On the Caribbean side which
lacks such contrasting seasonal changes, a
similar correlation cannot be found and the dis-
tribution of Helicinidae is probably not limited by
climatic conditions.

Morphological Characteristics

In the following section the different charac-
teristics will be discussed under general as-
pects, their applicability for species
differentiation, i. e. mainly for the Costa Rican
species, and their value for higher systematics
within the Helicinidae.

Teleoconch Shape

The shape and color of the teleoconch are
assumed to be directly affected by environ-

86° А 85°. 84° 1 ME | 835

oe e E 11°
®

? . . e

ee
> À |
»
®

Ke.

rain forest

dry forest

o Helicina tenuis

wet forest Боб =
moist forest

e all other species

09°

Rx: eae

FIG. 330. Costa Rican records of Helicinidae mapped on the types of vegetation,

Helicina tenuis individually marked.

412 RICHLING

mental selective pressures and therefore
highly adaptable. On one hand, differences
may therefore characterize different species,
but one the other hand, under different envi-
ronmental conditions a high plasticity may
occur within a single species as well. Like-
wise, there is a high probability of convergent
developments reducing the applicability of
these features in higher systematic.

Aspects of the shell color will be discussed
separately, along with the soft body color. Due
to their practical importance at the species
level, also for the determinations, shell charac-
teristics were already discussed in the species
accounts. In the following only certain general
aspects will be considered.

Despite the outline of the shell, the develop-
ment of the aperture is an important feature
for the differentiation of the Costa Rican spe-
cies. The aperture is always more or less ob-
lique, but may be straight or curved
backwards in its middle portion. The outer lip
is flatly expanded or reflexed. The main func-
tion of the outer lip of the aperture is obviously
the tight attachment to leaves or other sur-
faces of an aestivating individual. This is espe-
cially important for all arboreal species, such
as the Costa Rican representatives of
Helicina. Two different trends are realized to
achieve an optimal attachment: (1) the basal
part of the outer lip near the transition to the
columella is protruded, thus forming a den-
ticle, and is combined with a rather straight
aperture (e.g., Helicina tenuis), or (2) the
basal part is straight or even forms a little
notch, but the middle portion of the aperture 1$
strongly curved backwards (e.g., Helicina
beatrix). Especially the slight basal notch
present in all Costa Rican species that
Wagner (1907-1911) summarized under the
species group “Gemma” was used as indica-
tion for the inclusion in the subgenus
Leialcadia. This classification turned out to be
wrong, and the similarities can be explained
by convergent developments in adaptation to
the arboreal life, as is the feature of colorful
shells included in the description of the
Leialcadia. Another striking example for the
misleading shell characteristics is illustrated
by Alcadia jamaicensis (actual arrangement,
based on features of embryonic shell and fe-
male reproductive system) which has formerly
been classified as Helicina jamaicensis (e.g.,
Wagner, 1907-1911).

By the inclusion of Analcadia and Sericea
into Helicina, the presence of periostracal hairs

adds another example of convergence and
shows the unreliability of shell characteristics
as an indicator for relationships. Within the fam-
ily of Helicinidae periostracal hairs are known to
occur within at least in four different genera,
Helicina, Alcadia, Lucidella (L. adamsiana L.
Pfeiffer, 1849) and Schasicheila, the distinct-
ness of which is strongly supported by charac-
teristics of the female reproductive system and
embryonic shell structures. The periostracal
hairs seem to be related to a life within the leaf
litter, they are also developed in other families
of land snails (e.g., Helicidae).

Teleoconch Surface Structure

Among the Costa Rican representatives of
the genus Helicina, two different traits in
teleoconch surface structure can be recog-
nized, a rough surface with oblique diverging
grooves or a very smooth shell. Only one spe-
cies (Helicina escondida n. sp.) shows an in-
termediate characteristic with the former
structure only very weakly developed. Be-
cause in all species the beginning of the
teleoconch (subsequent to the transitional
structure) displays a pattern of oblique diverg-
ing grooves, the smooth surface is likely to be
a derived condition. The rough pattern is fur-
thermore not unique for Helicina, it can be
observed in different genera worldwide (per-
sonal observation), the only otherwise illus-
trated example is given for an Australian
species of Pleuropoma Möllendorff, 1893 by
Stanisic (1997). The relevance of the struc-
tures for revealing relationships remains
doubtful, especially due to the problems in the
classification of the Central American main-
land species (see below). Nevertheless, since
the two traits in the Costa Rican species are
paralleled by other similarities, although also
with intergrades (e.g., shape of the
teleoconch, details of the female reproductive
system, degree of sexual dimorphism), it may
be of importance as a supporting characteris-
tic on a smaller scale.

The distinct transitional structure (youngest
portion of the teleoconch) is not developed in
all taxa. It is present in all Costa Rican species
of Helicina and the Jamaican Helicina neritella,
Pyrgodomus and Alcadia (Microalcadia). In
Lucidella (L. lirata and L. aureola), the final pat-
tern of the teleoconch starts directly at its origin.
The same applies to Angulata in which the
embryonic pattern is even identical to that of
the teleoconch. The shells available for Alcadia

CLASSIFICATION OF HELICINIDAE 413

major were all eroded. The study of the figures
given for Helicina umbonata Shuttleworth,
1854, H. rhips Thompson, 1982 and H. liobasis
Thompson, 1982 by Thompson (1982) show a
similar situation as in Lucidella and Angulata.
With the exception of A/cadia (Microalcadia),
the absence of the transitional structure 1$ al-
ways combined with a spiral sculpture of the
teleoconch, whereas in the other species fol-
lows an oblique or irregular pattern. Because
the transitional structure in Helicina looks like a
preliminary stage of the pattern of the oblique
diverging grooves produced under different
growth conditions (slower or faster), a similar
effect simply would not become apparent in a
spiral pattern. The lack of knowledge of the life
history of Helicinidae reduces possible expla-
nations for the interpretation of the structures.
Especially the example of Helicina umbonata
and related species (verified in its generic po-
sition by embryonic shell structures) renders
this explanation likely. The systematic rel-
evance of this characteristic by itself is there-
fore not suggested here, only its occurrence in
combination with the pattern of the teleoconch.

Shell and Soft Body Color

Like shell characters, the color of shell and
soft body are believed to reflect directly the
result of selective processes by the environ-
mental conditions. For example, a study by
Johnson (1959) on Helicina orbiculata showed
that the percentage of a lighter or darker color
phase within a population is correlated to the
color of the soil and obviously controlled by
predation. The color is inconspicuous for spe-
cies dwelling in the leaf litter, such as Lucidella
lirata, Alcadia hojarasca, and A. boeckeleri,
which are in fact more or less uniformly
brownish or greyish colored. The color of
Pyrgodomus microdinus supports the camou-
flage of the shell on rock surfaces. All Costa
Rican species of Helicina for which the habi-
tat is reported are arboreal. It is known from
various examples of different families of land
snails, for example, Liguus (Orthalicidae),
Amphidromus (Camaenidae), and Cepaea
(Helicidae) that colorful and varied patterned
or the exceptional greenish shells seem to
camouflage the individuals best. In fact all ar-
boreal species investigated have this appear-
ance, but it is realized in two different ways:
(1) heavy, colorful shells, which are more or

less variable in their color, the mantle sur-

face is usually unicolored: Helicina funcki, H.

pitalensis, H. beatrix, H. talamancensis, H.
punctisulcata cuericiensis п. subsp. (2H.
echandiensis п. sp.).

(2) slight, nearly transparent shells, except for
the colored outer lip, the mantle surface is vari-
ously spotted: Helicina tenuis, H. escondida п.
sp., H. gemma, H. monteverdensis п. sp., H.
chiquitica.

Thus the development of the remarkable color

patterns on the soft bodies of certain species

closely depends on the thickness and struc-
ture of the shell. The best example is given by

Helicina tenuis and H. beatrix confusa, which

nearly equal each other in volume (females),

but the shell weight of H. tenuis amounts to
only 3/.. Because the arboreal life seems to
require a color as described above, the obvi-
ous physiological possibility of replacing the
shell color by mantle pigmentation first makes
the evolutionary development of thin shells
possible. This could represent an adaptation
to a limited availability of calcium carbonate
due to the geological conditions in Costa Rica.

Helicina escondida n. sp. and H. funcki are

similar to each other with respect to the pres-

ence of greenish specimens besides strongly
red tinged (shell) individuals in H. funcki or

variously spotted (mantle) individuals in H.

escondida n. sp. H. chiquitica represents an

exception in so far as most individuals are
nearly unicolored black. This may be related
to the small size of the species, because in the
individuals of H. monteverdensis n. sp. from

Mirador Gerardo, contrary to the larger ones

from Monteverde, the dark share of color pre-

vails. Small juveniles of H. funcki are darkly
mottled too.

In the single case of Helicina talamancensis,
the color of the head-foot seems to be charac-
teristic for the species. Assuming the thick-
ness and transparency of the shell to be
species-specific, the presence of a color of the
soft body as described above 1$ also charac-
teristic because it is shown to be closely cor-
related with shell conditions.

Internal Shell Structures: Axial Cleft and Muscle
Attachment

The absorption of the internal whorls of the
spire is a common feature of the families
Neritidae, Ceresidae, Proserpinidae, and
Helicinidae, which had been described for the
latter two families for the first time by Bland
(1854). Solem (1983), while studying 15 differ-
ent species of Helicinidae (worldwide, but not

114 RICHLING

specifically mentioned), established a relation
between the length of the axial cleft and the
number of whorls, that is, the length increases
with the number of whorls. The total range 1$
about % to nearly % of a whorl, and the length
of the axial cleft was shown to be species-spe-
cific (five specimens of each of two species
investigated). His results for Proserpina
Sowerby, 1839, and Ceres Gray, 1856 (in both
nearly % of a whorl), differ from the description
of the respective families given by Thompson
(1980), with Ceresidae being about '/,, of a
whorl and Proserpinidae about % of a whorl.
Thompson incorporated this characteristic in
his considerations of systematic relationships
between the families.

For the Costa Rican material, the present
investigation confirms the constancy of the
length of the axial cleft for different taxa, but
the relation of the number of whorls to the
length of the axial cleft cannot be sustained. It
may be illustrated at two examples: Helicina
punctisulcata cuericiensis n. subsp. has 3°/,
whorls, with an axial cleft of % whorl (Fig. 91),
whereas H. gemma has 4 to 4'/, whorls, with
an axial cleft of about */, whorl (Fig. 163). For
H. beatrix riopejensis n. subsp., shells of both
sexes are figured (Fig. 133) that differ in Y of
a whorl, but the axial cleft amounts the same,
here 3/, of a whorl. The second example, fur-
thermore, shows that a relation of whorl count
to the length of the axial cleft is actually in
contradiction with the species-specificity, be-
cause most species of Helicinidae exhibit a
sexual dimorphism in size that is accompa-
nied by a difference in whorl count of females
and males. This fact escaped the attention of
Solem (1983), but it was noted by Baker
(1928).

On the contrary, the length of the axial cleft
seems to be characteristic for certain system-
atic units as the data given by Thompson
(1980) suggests, although the differences be-
tween his data and Solem's (1983) remain to
be checked. From the present investigation, it
can be seen that Lucidella lirata remarkably
diverges from the species of Helicina, the axial
Cleft is % to % whorl longer than in all species
of Helicina studied. Also, with respect to the
attachment of the right retractor muscle exclu-
sively on the penultimate whorl, the species
differs from Helicina. The examination of
Lucidella aureola, the type species of
Lucidella, revealed similar conditions (Figs.
265, 322). The same is true for Schasicheila,
in which an axial cleft shorter than Y whorl is

St

always combined with the prominent right
muscle broadly attached only on the body
whorl. According to Baker (1925), the axial
cleft of the primitive genus Hendersonia
Wagner, 1905, encompasses nearly one
whorl. In all species of Helicina studied, the
attachment of the right muscle corresponds to
the beginning of the axial cleft and crosses the
inner suture and therefore encompasses both
whorls.

Within the Costa Rican species of both sub-
genera of Helicina the length of the axial cleft
varies from °/, to % of a whorl. In species of a
véry similar shell shape it may differ (e.g., H.
pitalensis — Н. tenuis) or be of about the same
length (e.g., H. monteverdensis n. sp. — H.
chiquitica). The subspecies H. beatrix
riopejensis n. subsp. represents a difficult
case: it closely resembles H. beatrix to which
it is tentatively assigned, but the length of the
axial cleft is like in H. gemma and unlike H.
beatrix.

Embryonic Shell

Previous to this investigation structures of
the embryonic shell of Helicinidae were only
applied twice for systematic considerations.
Clench 8 Jacobson (1971) stressed features
of the embryonic shell to exclude the subge-
nus Striatemoda Baker, 1940, from the Cuban
genus Emoda H. & A. Adams, 1856. In a sub-
sequent contribution on the genus A/cadia by
Boss 8 Jacobson (1973), the embryonic shell
surface is included in the descriptions of vari-
ous species, but its importance has not been
recognized by the authors. Their descriptions
are inadequate for a comparison with results
gained by SEM studies. Finally, Thompson
(1982) successfully used the feature to differ-
entiate the genera Alcadia, Helicina, and
Lucidella by investigation of the respective
type species.

The results of the present study confirm the
applicability of embryonic shell structures for
higher systematics in Helicinidae. Further-
more, the significance of embryonic shell fea-
tures is not only justified by considerations
about their conservative nature but verified by
well-founded parallel changes in the female
reproductive system in all species investi-
gated. The rearrangement of certain
subgeneric units of Alcadia provides a con-
vincing example. Additional genera
(Eutrochatella, Schasicheila and Angulata,
newly raised to generic level) can be charac-

CLASSIFICATION OF HELICINIDAE 415

terized by their special embryonic shell struc-
tures. The close relationship of Pyrgodomus
to Eutrochatella is confirmed. Within the
subgeneric level of the genus Helicina, embry-
onic shell structures show high similarities or
may vary within a single species to the same
degree as between different species (e.g., H.
monteverdensis n. sp.) so that it can only be
occasionally used as a distinguishing charac-
teristic. Among the Costa Rican species, for
example, only H. talamancensis, H. beatrix
beatrix, and H. beatrix confusa exhibit rela-
tively smaller pits with a more prominent
smooth surface.

Implications of the embryonic shell structure
on relationships of the different groups within
the Helicinidae are by far less obvious and the
following examples provide certain evidence
that the similar structures are not always ho-
mologous developments. The pitted structure
of the embryonic shell is known for the genera
Helicina, Lucidella (pits less regularly ar-
ranged) and Australian species of Pleuropoma
(similar to Helicina) (Stanisic, 1997), but, with
respect to complex characters of the female
reproductive system, Helicina represents a
derived condition compared to Lucidella, and
the genera of the Australasian region (see fe-
male reproductive system, Bourne, 1911).
Furthermore, the pitted embryonic shell
seems to be absent in primitive members of
the family, such as Hendersonia occulta ru-
bella (Green, 1832) (personal observation).
Studies on embryonic shell structure of other
gastropods also point to the difficulties to dis-
tinguishing homologous developments (e.g.,
Ponder & Lindberg, 1997).

Contrary to the structural similarity among
closely related species, size provides more
information. Size differences of the embryonic
shell have not previously been studied for
Helicinidae. According to the measurements
of the Costa Rican species, size seems to
depend on two main factors: (1) the size of the
species, and (2) the altitude of the locality of
the individuals.

(1) The diameter of the embryonic shell
would best be compared with the shell vol-
ume, but because this information is not avail-
able for all species, it is compared with shell
height as well as with the minor diameter of
the shells (Figs. 331, 332) to consider the de-
viations due to different diameter-height-rela-
tions, especially for the species of Lucidella,
Alcadia (Microalcadia) and Pyrgodomus. The
embryonic shell size increases with the shell

size, although deviations from this general
trend are remarkable. On one hand, it is ex-
plained by a certain species-specificity of the
embryonic shell size as is for example shown
for Helicina chiquitica or H. monteverdensis n.
sp.; in the latter species embryonic shell size
along with other arguments could be used to
distinguish the species. On the other hand, the
influence of the altitude (see below) inter-
poses with the relation to shell size. With re-
spect to higher systematics within the family,
first data for Lucidella and Eutrochatella sug-
gest that in these genera (and related like
Pyrgodomus), the embryonic shell size 1$ rela-
tively smaller than for example in Helicina.
The specimens investigated for Lucidella au-
reola equal Helicina gemma in size, but the
embryonic shell is about 285-335 um smaller,
those of Eutrochatella pulchella equal Helicina
beatrix confusa in size, the embryonic shell 1$
about 330 um smaller.

(2) Since the size of the embryonic shell also
seems to reflect a certain species-specificity
and not only the influence by the shell size,
the measurements of the embryonic shell
were directly compared to the altitude (and not
as a relation to shell size), but a similar dia-
gram for shell height and diameter to altitude
is given to show the general independence of
shell size and altitude (Figs. 333, 334). The
diagram illustrates a slight increase of the
embryonic shell size at higher altitudes inde-
pendently of the species within the represen-
tatives of Helicina and also for Alcadia
(Microalcadia), although the results for the lat-
ter subgenus are more suggestive in manner
than supported by sufficient data. On species
level, an increase of the size of the embryonic
shell with the altitude 15 clearly shown for
Helicina funcki and H. gemma.

Faced with a virtually complete absence of
data about the natural history of Helicinidae,
the results cannot be discussed in this con-
text. The knowledge is limited to a single de-
scription of deposited eggs for the species
Viana regina (Morelet, 1849), which 1$ cited
from the observation of a Cuban malacologist
by Clench 8 Jacobson (1968). It seems that
these eggs were calcified, because the ob-
server had to break them to examine the em-
bryonic shell. Furthermore, it is known that
eggs are released from the ovary into an egg
sac at the very beginning of the primary ovi-
duct. In histological sections, these eggs can
occasionally be observed in the primary ovi-
duct and in the ascending limb of the V-organ,

416

1400

RICHLING

T T
em-
bryonic
shell
diam.
[um]
Ben e ° Helicina
2 o
o © o
o o o
o ©
800 2
©
°
600 р
o А. hojarasca
o P. microdinus
° A. boeckeleri
FR o L. lirata
nen L Sl Ll - > 1
0 2 4 6 8 min. дат. [mm] 12
of shell
FIG. 331. Relation of embryonic shell diameter to minor diameter of the shell
for Costa Rican species (all species included for which measurements of the
embryonic shell were given).
1400 — 1 T T LD
em-
bryonic
shell
diam. ®
[um] oi o
a Helicina
1000 у |
o ©
9 o
800 o
600 l
© L. lirata .
_ + P. microdinus
> А. hojarasca
400 ° А. boeckeleri
L L LL L -
0 2 4 6 8 height [mm] 12

FIG. 332. Relation of embryonic shell diameter to shell height for Costa Rican
species (all species included for which measurements of the embryonic shell
were given).

CLASSIFICATION OF HELICINIDAE

1400 T ee le T ЕЕ T

em-
bryonic
shell |
diam. 9
[um] o

Helicina а
1000 +

$ o
o 6 ©
o

To o
o

800

oo ©
aif

600 | -
o А. hojarasca

о Р microdinus |
o А. boeckeleri

L. lirata

0 500 1000 1500 2000 altitude [m] 3000

FIG. 333. Relation of embryonic shell diameter to the altitude of the site of the
Costa Rican species (all species included for which measurements of the
embryonic shell were given).

minor diameter of shell +
de height +
[mm]
10 Я
E ° Неюста |
+ o $
o
+ + o
6-*о + - 4
+ o
o o
E o o >
4 Но + =
P. microdinus
? i e o .
, IES ° A. boeckeleri „A: hojarasca
<————_a—A A e = Le
0 500 1000 1500 2000 altitude [т] 3000

FIG. 334. Relation of shell height or minor diameter respectively to the altitude
of the site for the species included in diagram Fig. 333.

417

418 RICHLING

but distal to this portion (and therefore fertil-
ized) eggs were never seen or reported by
other authors.

Operculum

Wagner's classification represents the first
attempt to incorporate the characteristics of
the operculum to a large extent. For this rea-
son, the opercula of other than the Costa
Rican species are not repeatedly described
and discussed. As for shell characteristics, the
features on which the operculum closely de-
pend, subsequent studies, including the
present one, showed the high flexibility for
adaptations to environmental requirements by
cases of convergent developments reflected
in necessary rearrangements of different taxa.
For the genus Alcadia from Cuba, Boss &
Jacobson (1973) compared the calcification of
the operculum in relation to habitat. In arbo-
real species, the calcareous layer is thinner
than in ground-dwellers, the latter possibly
requiring stronger protection against small
predators.

Aspects of the operculum were successfully
applied to recognize and characterize the
primitive members of the family by strong
traces of a retained paucispiral instead of a
concentric condition (Wagner, 1907-1911;
Baker, 1922a).

The opercula of the Costa Rican species of
Helicina are very uniformly developed. Devia-
tions in shape are due to the different shape of
the aperture of the shells, for example, the
operculum of H. funcki is broader than that of
H. beatrix, because the whorls of the former
species increase more rapidly in size. The
opercula of Alcadia hojarasca and A.
boeckeleri do nat differ from those of Helicina
species, except for the more irregularly S-
shaped columellar edge, reflecting the condi-
tion of the strongly rounded periphery of the
shells. This illustrates in exemplary fashion
that the similarity in the operculum does not
necessarily reflect a close relationship be-
tween the taxa, especially in the case of cer-
tain subgeneric units of Helicina and Alcadia.

Radula

Because radular characteristics strongly in-
fluenced the systematic concepts for: the
Helicinidae, these structures were described
and figured in some detail within the species
accounts.

The interpretation of the capituliform com-
plex requires some remarks, because it has
caused some past confusion. Originally,
Troschel (1856-63) described the accessory
plate as a basal appendage, which is always
fused with the tooth, but appearing somehow
subdivided. Baker (1922a) recognized two
plates and paid much more attention to the
form of the accessory plate and the point and
way of the overlapping of both teeth. His fig-
ures give the impression of separate teeth.
The peculiar reflexed wing of the accessory
plate was corrected in its interpretation by
Baker (1926, 1928) as a deposit cementing
the two plates together instead of being a kind
of a cusp. The plates are said to break upon
separation in certain groups of species,
whereas in others they “pull apart easily
enough”. This correction obviously escaped
the attention of subsequent authors. Adopting
Baker (1922a), Thompson (1980, 1982) still
incorporates the characteristic of the wing or
reflection of the accessory plate enveloping
the end of the comb-lateral for his phyloge-
netic considerations. This wing is actually not
visible in his SEM figures, because it was a
misinterpretation of optical microscopic inves-
tigations. Stanisic (1997) states that, contrary
to certain overseas (= neotropical) species,
the comb-lateral and the accessory plate of
the Australian species of Helicinidae are al-
ways fused together. But in this respect, his
SEM figures do not differ from those of
neotropical taxa, for example, the Costa Rican
species. In natural conditions, the two plates
are fused together.

As previously stated, data of this study were
gained by SEM-investigations. Despite the
great advantages of the higher resolution re-
vealing more exactly the dentition of the
marginals and information from the three-di-
mensional arrangement, the method is ac-
companied by some disadvantages. On
account of the very complex structure of the
radula, the numerous marginals always over-
lap each other, especially the outer ones, ren-
dering an exact count impossible. To a lesser
extent, a similar problem exists in analyzing
the number of transverse rows, because, es-
pecially in small or poorly preserved speci-
mens, the preparation procedure and the
necessary positioning may result in losses of
parts of the whole ribbon. In optical micros-
copy especially, the first point simply becomes
obsolete by the transparency of the radula.
Nevertheless, here the SEM-method is pre-

CLASSIFICATION OF HELICINIDAE 419

ferred because drawings from the light micro-
scope reflect rather the interpretation of the
author than the real and hereby comparable
structures as argued above.

In all Costa Rican species of Helicina, the
radula displays nearly the same amount of
intraspecific and interspecific variations. The
only specific differences clearly documented
were found in the dentition of the comb-lateral
of two species, H. escondida n. sp. and H.
chiquitica. In these species, the number of
cusps is constantly strongly reduced or in-
creased, in the former species, the size of the
cusps is remarkably enlarged towards the
centrals. The differences described for the
change in the number of cusps on the
marginals are in most cases likely to be re-
lated to the size of the species. Evidence is
provided by the differences in the subspecies
of Helicina beatrix. Furthermore, all small-
sized species (Helicina chiquitica, Lucidella
lirata and Alcadia (Microalcadia) hojarasca)
convergently possess an increased number of
denticles on the comb-lateral.

Stanisic (1997), while studying Australian
species of the genus Pleuropoma, in which
the radula in general apparently does not dif-
fer remarkably from those shown in this study,
points out specific differences of the comb-lat-
eral for certain species analogous to those
observed for Helicina escondida n. sp. and H.
chiquitica, and in the cusps of the centrals as
well. He relates the divergence in the radula to
the habitats of the species — arboreal, ground-
dweller and limestone-associated.

The results and interpretation of Stanisic
(1997) agree with my own conclusions of see-
ing the features of the radula more in the light
of speciation and adaptations to the substra-
tum the respective species feed on and thus
to be subject to convergent development
rather than providing a conservative structure
useful to indicate phylogenetic relationships.
This concept does not exclude the possibility
that a certain radiation resulted from the colo-
nization of a special habitat (e.g., Eutrochatella
and related taxa of the West Indies on calcar-
eous rocks), which 1$ therefore characterized
by the obviously necessary adaptation of the
radula (e.g., the reinforced T-shaped lateral).
On one hand, the fact of the rearrangement of
certain groups of Baker's system character-
ized by radula characteristics provides evi-
dence in itself. On the other hand, additional
examples can be given for convergent de-
velopments. Boss 8 Jacobson (1973) and

Thompson (1982) had recognized the diverg-
ing radula of Alcadia major (T-shaped laterals
and other features approaching the “vianid”
radula instead of the “helicinid” radula typical
for other subgroups of A/cadia) but hesitated
to comment on this “not matching” fact. Be-
cause Alcadia major lives on “rock bases”
(Baker, 1934a) like other species with a
“vianid” radula (e.g., Eutrochatella pulchella,
Pyrgodomus microdinus), a convergent forag-
ing structure is very likely and is here favored
as an explanation. Outside of the family
Helicinidae, a convergent development of the
“vianid” radula had already been proposed
and accepted by Thompson (1980), namely
for the Proserpinidae.

Morphometry and Sexual Dimorphism

The morphometric differences were dis-
cussed to some extent in the species account
and proved to be useful on the species level,
for example, to judge the status of certain sub-
species (Helicina funcki costaricensis, H.
tenuis pittieri). Furthermore, a relation to the
altitude seems to be specifically different, but
data were sufficient only for a few species. A
decrease in size at higher altitudes (Helicina
funcki and H. gemma) has also been reported
for the Mexican species H. fragilis elata, H.
zephyrina zephyrina and Pyrgodomus
microdinus abditus by Baker (1928). The
study also shows that morphometric analysis
of Helicinidae always has to take into account
the considerable effects of sexual dimorphism.

Sexual dimorphism in Helicinidae has been
known for a long time, but, with very few ex-
ceptions, it has never been subjected to de-
tailed studies. Probably the most frequently
cited example 1$ the different shape of the
shells of both sexes in Viana regina, with the
males being characterized by a deep notch in
the upper outer lip, a fact that was first recog-
nized in this context by L. Pfeiffer (1856a).
Wagner (1910b) considered the subject theo-
retically because he did not have anatomical
material available. The species discussed by
him were not studied here. Baker (1925) pub-
lished the first accurate data for differences in
size for Hendersonia occulta rubella. Although
he selected extremes for the measurements
expecting the females to be larger, the differ-
ence in volume interpolated from his data
amounts to about 2% less for the males and
both sexes intergrade. Baker (1926) men-
tioned only the relationships of females to

420 RICHLING

males of the specimens he had studied ana-
tomically. Baker (1928) analyzed some Mexi-
can species morphometrically, but, with minor
exceptions, the number of specimens for each
station was limited. According to his results,
the sexes intergrade considerably in shape
and size, but the shells of males increase
more rapidly in diameter resulting in fewer
whorls (% to %) and a more depressed shell
shape, when sexual dimorphism is developed.
The genus Schasicheila seems to encompass
species with sexual dimorphism ($. alata, fe-
males > males) and another with intergrading
sizes (S. misantlensis Fischer 8 Crosse,
1893, females = males). In later studies on the
Jamaican malacofauna, Baker (1934a, b) pro-
vided further information, but unfortunately did
not include numerical data or the sample size.
According to him, all species examined of
Eutrochatella E. pulchella, E. tankervillii
(Gray, 1824), E. nobilis (C.B. Adams, 1852)
and Е. costata (Gray, 1824) — have males
larger than the females, whereas the closely
related genus Pyrgodomus (Baker, 1928: P.
microdinus abditus, examination based on
several shells) shows the reverse relation.
Species of Lucidella show all possible rela-
tions (<, >, =), and Helicina and Alcadia are
similar in having larger females. A single ex-
ception for Helicina is given in “H. (Angulata)
rhynchostoma ernesti” von Martens, 1873
(Baker, 1926), but this species requires re-
examination regarding its systematic affinities.

The data on sexual dimorphism of this study
represent the most comprehensive approach
to date to analyze the phenomenon for a se-
lected group of species. Against the back-
ground of limited material available, the newly
developed method of removing the soft body
with only minor damage to the shell provides
a valuable tool, which made the analysis pos-
sible. All species measured exhibit a sexual
dimorphism in size of the shell, with females
averaging larger than males. Differences are
smallest in Lucidella lirata (percentage of
male's volume about 92%), increase in
Helicina funcki and H. pitalensis (about 80-
85%), H. tenuis, H. echandiensis n. sp. and H.
escondida п. sp. (about 75-78%) to the great-
est values in H. beatrix, H. talamancensis, H.
gemma, H. monteverdensis n. sp. and H.
chiquitica (about 61-67(-72)%). This grouping
is paralleled by similarities in shell shape,
teleoconch surface structure and details of the
female reproductive system, suggesting a cer-
tain value of the degree of sexual dimorphism

with respect to systematic affinities. The group
with the highest differences shows further-
more that the dimorphism is independent of
the size of the species, because the extrema
(Helicina talamancensis and H. chiquitica) dif-
fer in volume by a factor of about 7.

The observation by Baker (1928) that in
case of a sexual dimorphism males are more
depressed, that is, relatively larger in diam-
eter, could not really be confirmed by the
present study. For all species, the different
measurements were tested in various rela-
tions to each other and no significant differ-
ences for both sexes were found. For the
relation of height to diameter (here minor di-
ameter), deviations between females and
males for acceptable sample size usually
range approximately 1% or less for all spe-
cies, highest differences amount up to 3-4%
for very few populations. In more populations,
the relative diameter was larger in males than
in females, but examples for a reverse relation
were found likewise in nearly all species.

The morphometric comparison of different
populations clearly demonstrates a certain
variability of size for most species and indi-
cates that investigations of the sexual dimor-
phism will only work with individuals
originating from the same population.

The knowledge of the range of size varia-
tions within a species due to sexual dimor-
phism allows a much better judgement of the
determination of single specimens, for ex-
ample, type specimens, and morphometric
differences applied for the separation of spe-
cies or subspecies.

Female Reproductive System

When revising the contributions on system-
atics of the Helicinidae with respect to the
anatomy, especially genitalia, all authors
agree in the following: “..., but once they [ana-
tomical structures] evolved very little differen-
tiation of these organs and structures oc-
curred with further radiation of subfamilies and
genera” Thompson (1982: 5), or more strictly
“This emphasizes the conclusions of Bourne
(1911: 777) and Baker (1926: 35) that the gen-
eral uniformity of the genitalia of the
Helicinidae makes them useless for diagnos-
tic purposes” Boss & Jacobson (1974: 6).

The first comprehensive study on the
anatomy of Helicinidae based on several spe-
cies worldwide, combining both dissections
and histological studies, was carried out by

CLASSIFICATION OF HELICINIDAE 421

Bourne (1911). А similar study was provided
only for the primitive species Hendersonia
occulta rubella by Baker (1925). All previous
contributions considered the one or the other
detail of different species (Isenkrahe, 1867:
first rough anatomy of Emoda pulcherrima
titanica (Poey, 1851); von Ihering, 1877: ner-
vous system of Pleuropoma beryllina (Gould,
1847); Bouvier, 1886: nervous system of
Angulata brasiliensis and Emoda sagraiana
(Orbigny, 1842); Thiele, 1902: male reproduc-
tive system of Waldemaria japonica (A.
Adams, 1861); Thiele, 1910: female reproduc-
tive system of Helicina kubaryi [nomen nu-
dum?]). Baker (1926, 1928) studied several
American species with emphasis on the geni-
talia. He clearly states that his intention of find-
ing similar clues for systematic affinities, as in
the radula, partially failed, not exclusively be-
cause of the considerable uniformity of the
structures, but also on account of the scanty
and poorly preserved material. Therefore, his
contribution is a valuable collection of descrip-
tions for several anatomical features, but the
nearly complete absence of conclusions and
discussions renders it subject to misinterpre-
tations, as exemplified in the citation above.
Subsequent to Baker, only Thompson (1968)
described the genitalia of his new genus
Ceochasma and provided a detailed anatomi-
cal study on two species representing the re-
lated families Ceresidae and Proserpinidae
(Thompson, 1980).

The present anatomical studies focused on
the investigation and comparison of the fe-
male reproductive system since, on one hand,
the discovery of important deviations from the
present knowledge promised information rel-
evant for phylogenetic purposes contrary to
former assumptions, but on the other hand, it
required the re-examination of previously
studied species/genera and the assessment
of data within the new context.

The study of the Costa Rican species of
Helicina revealed that all species are similar in
the monaulic condition of the female reproduc-
tive system. This result is in contradiction with
all previous anatomical studies (diaulic sys-
tem), especially those of Baker (1926) for vari-
ous Central American species of Helicina.
Baker (1926) assumed the vaginal opening to
be inside the duct of the hypobranchial gland
also discharging into the mantle cavity. This is
reflected in his figures of the female systems
(e.g., fig. 9, H. orbiculata). In fact, the hypo-
branchial duct and gland is closely associated

with the apical complex of the female repro-
ductive system, partially enveloping these
structures dorsolaterally. In dissection, their
separation is not always easy, but the study of
histological serial sections finally confirmed
the absence of any connection of the recep-
tion chamber or associated structures to the
hypobranchial duct. Other Central American
species, such as Helicina amoena, H.
turbinata, and H. orbiculata, the latter also
studied by Baker (1926), also constantly lack
the provaginal opening. Until more knowledge
is available, these results together with the
only vague presentation of the provaginal
opening in Baker's figures lead me believe
that all Central American mainland species
commonly referred to Helicina are monaulic,
that is, that they properly belong to that genus.

Reexamination of several other species
studied by Bourne (1911) and Baker (1926)
and additional type species of higher system-
atic units revealed a much higher structural
diversity than previously documented. For
example, contrary to the results of Bourne
(1911) Eutrochatella pulchella was found to be
monaulic. For Lucidella lirata as well as for L.
aureola, an additional sac on the pallial ovi-
duct for sperm storage was discovered. Fur-
thermore, the receptaculum seminis on the
descending limb of the V-organ described by
Baker (1926, 1928) turned out to be a miscon-
ception. These results add several peculiari-
ties for Lucidella and differentiate it more
strongly from other genera. The characters of
Schasicheila (Baker, 1926, 1928) and Alcadia
(Bourne, 1911) were confirmed.

The changes in the female reproductive sys-
tem are paralleled by consistently different
structures of the embryonic shell and add
valuable features in the characterization and
differentiation of genera and subgenera, for
example, Alcadia from Helicina. Furthermore,
certain changes can be assessed in the direc-
tion of the development. The basal members
of Helicinidae, such as Hendersonia occulta
rubella, possess a diaulic system. The basal
position 1$ reliably founded on the paucispiral
conditions of the operculum and the presence
of the vestigial right auricle. The monaulic con-
dition is clearly the derived condition. Further-
more, the diaulic state seems to prevail in
most of the genera. Assuming the results of
Bourne (1911) for this part to be correct,
Australasian species are diaulic as well. At
least Aphanoconia pachystoma ponsonbyi
(E.A. Smith, 1884) from Papua New Guinea,

422 RICHLING

was re-examined and the structure could be
confirmed.

The топ- or diaulic state is expected to be
related with functional consequences. But al-
though the morphological structures are fairly
well documented, knowledge of functional
aspects is limited to the interpretation of mor-
phological features, because other data are
not available. Bourne's (1911) observations
were not homogeneous for the different taxa
and point in different directions. The presence
of sperm in the posterior part of the pallial ovi-
duct, the bursa copulatrix, and the receptacu-
lum seminis of Alcadia hollandi suggested that
the pallial oviduct serves as the copulatory
canal. By the way of contrast, two Australasian
species of Aphanoconia had sperm within the
provaginal duct and the provaginal sac, favor-
ing the reception of sperm through this open-
ing. Baker (1925) assumed the provaginal
opening to receive the male products in
Hendersonia occulta rubella, as did Thompson
(1980). The monaulic structure only allows the
reception of sperm through the pallial oviduct,
and therefore demonstrates the physiological
possibility. Following this consideration, the
provaginal opening could also be functionally
vestigial, at least with respect to the reception
of sperm, in species with this opening. Under
the morphological conditions of Lucidella for
example (Figs. 270, 324), the reception of
sperm through the vagina would require a
downward movement to reach the posterior
extended portion of the oviduct and its lower
appendage, in which sperm were found. The
function of the various different structures for
sperm storage is even less understood. In
most of the different systems there are three
structures that are found to be used simulta-
neously (receptaculum seminis, bursa
copulatrix, provaginal sac, ?parts of the pallial
oviduct, the appendage of the pallial oviduct or
other analogous developments). These as-
pects remain subject to further studies, possi-
bly ultrastructural analysis.

The variability or specificity of characteristics
of the female reproductive system at the spe-
cies level was studied in detail for the Costa
Rican species. Here the applicability is limited,
as may be expected for characters useful for
higher systematics. Differences mainly occur
in the shape and size of the bursa copulatrix
and the provaginal sac, as well as in the rela-
tion of the apical complex to the pallial oviduct.
The latter feature seems to depend more on
the size of the species, because the apical

complex is not proportionally larger in larger
specimens, but the absolute distance to the
mantle edge is longer, for example, Helicina
funcki, H. pitalensis. The shape of accessory
structures is nearly independent of the devel-
opmental stage. Baker (1926) observed that
lobules of the bursa copulatrix of immature
Helicina convexa L. Pfeiffer, 1849, were al-
most as well developed as those of adults.
Dissections of immature Helicina funcki con-
firm this assumption (Fig. 19, right drawing).
The main changes connected with maturation
take place in the development of the ovary
that finally covers large parts of the visceral
portion and an enormous thickening of the
epithelium of the pallial oviduct (Bourne, 1911;
personal observations). Furthermore the con-
tent of the accessory structures (empty or
filled) does not remarkably influence the
shape. Therefore the deviations illustrated are
more likely due to intraspecific variability
rather than different physiological conditions.
Similar to the radula for some species, a cer-
tain peculiarity can be recognized, for ex-
ample, the lobules at the upper end of the
provaginal sac of Helicina tenuis, the elon-
gated lobes of the bursa copulatrix of H.
funcki, similarly developed in specimens from
Panama (Baker, 1926).

Arrangement of Central American Mainland
Taxa

Except for the single species Pyrgodomus
microdinus and Lucidella lirata, the main part
of the Costa Rican Helicinidae — namely the
species of Helicina — was controversially clas-
sified and shifted to subgroups of Helicina or
Alcadia. Because this confusion is character-
istic for the two genera involving most of the
American mainland species and a consider-
able portion of the Caribbean species, it must
be treated to some extent. Subsequently, as-
pects of the remaining Central American main-
land genera will be discussed.

The Genera Helicina and Alcadia

In the following, the different classifica-
tions of Helicina and Alcadia and related
subgenera proposed in literature will be pre-
sented, critically summarizing the main dis-
tinguishing characters of the groups that
were stressed by the authors. The respec-
tive systematic position of the Costa Rican
taxa is indicated. Additionally, all relevant

CLASSIFICATION OF HELICINIDAE 423

subsequent contributions on the systematic
classifications will be discussed. Finally, the
differentiation and characterization of the
genera and subgroups and necessary rear-
rangements will be proposed according to
the morphological characters and their par-
tially deviating assessment outlined by this
study.

Wagner (1907-1911) — mainly based on fea-
tures of shell and operculum. The respective
assignment of the Costa Rican species is
highlighted in bold face style.

Alcadia
Subgenus: Eucaladia [= Alcadia]: Jamaica,
Cuba, Bahamas, St. Thomas, St. Jan [=

St. John?], Vieque, Puerto Rico,
Hispaniola, French Guyana, Suriname,
Brazil

Formenkreis “Palliata”: Jamaica, Cuba

Formenkreis “Hispida”: Cuba, Bahamas,
St. Thomas, St. Jan [= St. John?],
Vieque, Puerto Rico

Formenkreis “Intusplicata”: Hispaniola

Formenkreis “Sericea”: French Guyana,
Suriname, Brazil

Formenkreis “/ncrustata”: Cuba

Subgenus: Leialcadia [= Idesa]: Cuba, Ja-
maica, Puerto Rico, Hispaniola, Trinidad,
Venezuela, Colombia, Costa Rica, Nica-
ragua, Guatemala, Mexico

Formenkreis “Megastoma”:
Cuba, Puerto Rico

Formenkreis “Nitida”: Cuba, Puerto Rico

Formenkreis “Mamilla”: Hispaniola, Cuba

Formenkreis “Bellula”: Cuba

Formenkreis “Ampliata”: Jamaica

Formenkreis “Tamsiana”: Trinidad, Ven-
ezuela, Colombia

Formenkreis “Gemma”: Costa Rica, Nica-
ragua, Guatemala, Mexico: gemma,
beatrix, (fragilis)

Subgenus: Analcadia: Guadeloupe,
Martinique, St. Lucia, Dominica, Venezu-
ela, Trinidad, Belize, Bonacca (island off
Honduras), Nicaragua, Hispaniola,
Puerto Rico, Vieque, St. Jan [= St.
John?], Tortola

Subgenus Emoda: Cuba

Helicina
Formenkreis “Angulata” and “Variabilis”:
Brazil
Formenkreis “Concentrica”: Venezuela,
Colombia, Peru, Bolivia
Formenkreis “Punctisulcata”: Mexico, Gua-
temala: punctisulcata

Jamaica,

Formenkreis “Cinctella”: Mexico, USA

Formenkreis “Tenuis” [= Pseudoligyra]:
Mexico, Guatemala, Honduras, Nicara-
gua, Costa Rica, Panama, Bolivia:
tenuis

Formenkreis “Turbinata”: Panama, Costa
Rica, Nicaragua, Honduras, Mexico,
Guatemala: funcki, pitalensis

Formenkreis “Succincta”: Mexico, Guate-
mala

Formenkreis “Festiva”: Hispaniola

Formenkreis “Euneritella” [= Нейста]:
Cuba, Jamaica, Grenada, St. Vincent,
Guadeloupe, Dominica, Bonacca,
Martinique, Trinidad, Barbados, Bermuda

The differences between Helicina and
Alcadia given by Wagner (1907-1911) can be
summarized as consisting mainly of shell char-
acteristics (in Helicina: umbilical area always
and constantly with an impressed line or
groove, no periostracal hairs; Alcadia s.s.: typi-
cal basal notch). Furthermore, the operculum of
Helicina only differs in a less prominent sigma-
edge and a nucleus most closely approaching
the columellar edge (but this is also attributed
to Analcadia, Leialcadia and Alcadia s.s. par-
tially). Wagner (1907-1911) adds that some
species of Alcadia and Mexican and Antillean
species of Helicina intergrade or only weakly
exhibit the typical characters respectively.

With respect to the characteristics of the
operculum, Analcadia differs from Leialcadia
only in a deeper groove near the lower part of
the sigma-edge, other differences apply to
shell characteristics. Leialcadia is typically
characterized by the shiny and _ lasting
periostracum and reduced characteristics of
aperture and operculum. Both subgenera lack
the prominent calcareous plate of the opercu-
lum and the deep notch in the basal part of the
aperture described for Alcadia s.s.

The “Formenkreise” were only used to sum-
marize groups of species, they were intro-
duced without any description or type species.

Baker (1922a) — mainly based on characters
of the radula, in combination with some fea-
tures of the shell and operculum.

Oligyra (tropical and subtropical America)
Subgenus: Oligyra: tropical and subtropical
America
Section: Oligyra s.s.: USA, Bermuda Is-
lands, Mexico
Section: Succincta: Mexico to South
America: gemma, beatrix

424 RICHLING

Subgenus: Alcadia: West Indies to South
America
Section: /desa: West Indies
Section: Analcadia: Antilles to Central
America
Helicina (tropical America)
Subgenus: Helicina: West Indies
Subgenus: Tristramia: mainland [of tropical
America]
Section: Oxyrhombus: Central and E-
Mexico to South America: punctisulcata
Section: Tamsiana: northern South America
Section: Angulata: South America to Cen-
tral America
Section: Tenuis [= Pseudoligyra]: Mexico to
Central America; South America: tenuis
Section: Tristramia: Mexico to Colombia:
funcki, pitalensis

According to Baker (1922a), Oligyra differs
from Helicina by a well-developed wing at the
accessory plate and centrals with always well-
developed cusps. The operculum shows
intergrading characters. Compared with
Alcadia, the genus has a light operculum lack-
ing the inferior point fitting in the correspond-
ing notch in the basal outer lip, and A- and
B-central do not exhibit heavy backs. It was
later maintained that Oligyra and Alcadia be-
long to two diverging lines of evolution. It
would, however, be extremely difficult to name
any very definite characteristics for their sepa-
ration.

Succincta differs from Oligyra in the ten-
dency of reduction of the cusps of the A-cen-
tral, whereas A- and B-central of Oligyra are
with well-developed cusps. Additionally, the
shell is more globose, and marginals show a
wing-like expansions below the tips (otherwise
strictly lingulate).

According to Baker (1923) the centrals of the
radula of Analcadia resemble those of Oligyra
s.s., but shell characters approach those of
Alcadia, suggesting Analcadia as a subgenus
between the subgenera Oligyra and Alcadia.
The radula of Oligyra (Alcadia — section:
Idesa) rotunda is said to agree with Alcadia
s.s. in the centrals and with Sericea or
Analcadia in the comb-lateral.

Tristramia differs from Helicina in strictly
lingulate marginals (not sickle-shaped with
lateral wings near tips), a lacking shelf-like
projection bearing the cusps on the A-central
and a differing accessory plate (operculum
and comb-lateral are equal). The differences
between Oxyrhombus, Tamsiana, Angulata,

Tenuis, and Tristramia include shell character-
istics (presence or absence of a Spiral stria-
tion, periphery angular or not) and radula
characters not worth mentioning.

All the radula differences within Helicina and
Oligyra (because the main difference “the
wing” was a misinterpretation) become re-
duced to minor deviations in cusp develop-
ment (presence and shape) on centrals and
the comb-lateral and the arrangement and an
increase in the number of the cusps on the
marginals (more on tip or laterally).

The author erroneously recognized the
“Formenkreise” of Wagner (1907-1911) as
relevant supraspecific taxa and, by doing so,
made them nomenclaturally available (Zilch,
1948); some became objective synonyms of
older supraspecific taxa, other were synony-
mized, but some were accepted as sections.
Because there is good reason to suppose that
Wagner based the “Formenkreise” on similari-
ties of shell characteristics, it may be said that
for the taxa accepted and kept in their posi-
tion, Baker only attempted to consolidate this
system with questionable radula features, in-
stead of having raised it based on these char-
acteristics.

Baker (1926) — system modified to incorpo-
rate anatomical characters.

Helicina
Subgenus: Helicina
Subgenus: Oligyra
Section: Oligyra $.5$.
Section: Succincta: gemma, beatrix
Subgenus: Tristramia |
Section: Tristramia s.s.: funcki, pitalensis
Section: Tenuis [= Pseudoligyra]: tenuis
Subgenus: Oxyrhombus
Section: Oxyrhombus s.s.: punctisulcata
Section: Angulata
Section: Tamsiana
Alcadia
Subgenus: Alcadia
Subgenus: Analcadia
Subgenus: Sericea
Subgenus: /desa (included after Baker, 1923)

Because Baker assumed the anatomy of
Alcadia s.s. was distinct from Oligyra (he stud-
ied Analcadia and Sericea, but did not have
adequate material of Alcadia s.s. and only
very critically interpreted the [correct] figure
given by Bourne, 1911, as generalized), he
raised Alcadia again to generic level and
pointed to further investigations to show the

CLASSIFICATION ОЕ HELICINIDAE 425

exact line of demarcation between Oligyra and
Alcadia. He stated that shell and radula char-
acters of Succincta (Helicina (Succincta)
cacaguelita Pilsbry & Clapp, 1902, examined)
approach those of A/cadia, but the anatomy
appears closest to that of the subgenus
Tristramia (Helicina (Tristramia) funcki exam-
ined).

Baker pointed out that anatomical and shell
characteristics of Helicina and Oligyra “inter-
grade to such an extent, that one meets con-
siderable practical difficulty in any attempt to
differentiate the two groups.” Therefore, he
included Oligyra with two sections as subge-
nus into Helicina.

When describing the anatomy of Helicina
(Oxyrhombus) cinctella, Baker (1928) re-
marked that the female genital system is inter-
mediate between those of the species of
Oligyra and that of H. concentrica L. Pfeiffer,
1849 [the only representative for Oxyrhombus
studied by Baker (1926)], although the struc-
ture of the bursa copulatrix distinctly ap-
proaches that of Tristramia.

Rehder (1966) discussed the systematic af-
finities of Helicina bocourti Crosse & Fischer,
1869, Honduras (formerly regarded as sub-
species of H. dysoni, Venezuela) applying the
characteristics of the radula and concluded
that both species belong to the subgenus
Tristramia, of Helicina, synonymizing
Oxyrhombus and Tristramia, because the
radula of H. bocourti combines characteristics
of both groups. Regarding a rearrangement of
the subgenus Analcadia, of which H. dysoni is
the type species, to Helicina instead of
Alcadia, Rehder (1966) avoided a direct state-
ment, because he later pointed to the
periostracal hairs and the development of the
operculum of H. dysoni as characteristic ofthe
genus Alcadia and mentioned his uncertainty
“that radular characteristics alone can be used
for subgeneric differentiation”.

Boss & Jacobson (1973) revised the Cuban
snecies of the genus Alcadia. Considering
anatomical studies of the previous authors
and combining their own results on the radula
with Baker's (1922a), they pointed out that
“shell morphology, together with certain fea-
tures of the operculum still constitute the most
reliable method of distinguishing members of
the genus”. They then proceed to differentiate
Helicina and Alcadia mainly by the basal notch
or sinus of the shell and an internal lamella
and groove on the columellar edge of the
operculum. Nevertheless, it remains doubtful

how they define the genus outside of Cuba,
that is, the exact line of demarcation between
Helicina and Alcadia, because Alcadia in their
sense (Boss & Jacobson, 1973: 311-312) oc-
curs on most of the West Indian islands as
well as on the mainland from southern Mexico
to northern South America. Actually, this distri-
bution only agrees with Wagner's version and
not with that of Baker (1922a, 1926), who at-
tributed part of Wagner's Leialcadia, namely
the species groups “Gemma” and * Tamsiana”,
to Helicina or Oligyra respectively and, by do-
ing so, excludes Alcadia from Central America
(but not from northern South America). Boss &
Jacobson (1973) even assumed the origin of
the genus to be in Central America.

Their investigation of the radula of four Cu-
ban species (two of them type species of sub-
genera) and Alcadia major (Jamaica) revealed
that it cannot be used as a diagnostic feature
for Alcadia, at least at the present state of
knowledge. Troschel (1856-63), for example,
noted a different shape of the R-central in
Helicina and Alcadia, but Boss & Jacobson
found it to be too variable in their species stud-
ies of Alcadia.

Thompson (1982), clarifying the systematic
affinities of the species group Helicina
umbonata from the West Indies, used embry-
onic shell structures for the first time, because
they are of conservative character and show
little variations within species or closely re-
lated groups. When judging other characteris-
tics for their applicability to systematics, he
regarded shell and operculum as being di-
rectly affected by evolutionary pressures due
to their direct contact with the environment
and thus as being subject to convergence. By
the way of contrast, Thompson (1982) as-
sumes the radula as a useful, conservative
morphological system, citing the studies of
Baker (1922a), although it is obviously also
under direct selective pressure as foraging
organ, and although Thompson (1980) argues
that the vianid radula (radula with inner lateral
T-shaped) has convergently evolved in
Proserpinidae and Vianinae for similar trophic
activities.

His results with respect to the embryonic
shell show that A/cadia clearly differs from
Helicina s.s. Other generic units placed in
Alcadia as subgenera are said to agree with
Alcadia s.s., whereas some mainland subgen-
era associated with Helicina differ from
Helicina s.s. thus requiring re-examination.
Against the backdrop of the controversial ar-

426 RICHLING

rangement of subgeneric units of Helicina and
Alcadia, it remains doubtful just what Thomp-
son (1982) refers to. He does not specify any
taxa or species on which he based his state-
ment about the subgeneric taxa except for
those genera within the actual scope of his
study (Helicina s.s., Alcadia s.s., Lucidella s.s.
and Poenia).

To summarize the present state of knowl-
edge, it can certainly be stated, that the dis-
crimination of the genera Helicina and Alcadia
and their associated subgenera remains a
controversial topic. Especially the species
groups “Gemma” and *Tamsiana”, the former
encompassing part of the Costa Rican taxa,
were shifted either to the one or other genus.
Radula differences turned out to be wrong in-
terpretations or to intergrade or vary. The fea-
tures of shell and operculum most strongly
and constantly influenced the classification
since they actually also represented the foun-
dation for later concepts. The anatomy has
been regarded as too uniform and conserva-
tive even within the Helicinidae, except for
some primitive members (e.g., Hendersonia).
The only definite and first hint for a clear sepa-
ration of the genera s.s. is given by the struc-
ture of the embryonic shell.

New Proposed Arrangement

From the results of the present study, the
following arrangement is proposed for those
taxa investigated. Details of the assessment of
the different characteristics were discussed in
the foregoing chapter. Contrary to previous
attempts, emphasis is placed on differences in
the female reproductive system, which agree
with changes in the embryonic shell structure.

Helicina’

Subgenus: Helicina s.s. (West Indies)

Subgenus: Tristramia (Synonyms:
Oxyrhombus, Pseudoligyra [= Tenuis],
?"Cinctella”) (Central American тат-
land): funcki, pitalensis, tenuis,
echandiensis n. sp., punctisulcata
cuericiensis n. subsp.

Subgenus: Oligyra (Synonym: Succincta)
(Central American mainland)

Subgenus: “Gemma”: gemma, beatrix,
talamancensis, monteverdensis п.
sp., chiquitica, escondida n. sp.

Subgenus: Ceochasma (Mexico)

Subgenus: Analcadia (northern
America)

South

Subgenus: Sericea (northern South America)

?Subgenus: Tamsiana (northern South
America)
Angulata (South America)
Alcadia”

Subgenus: Alcadia (West Indies)

Subgenus: Microalcadia n. subgen. (Cen-
tral American mainland): hojarasca,
boeckeleri

Subgenus: /4еза (West Indies)

“ the South American taxa will be discussed, but they
are only included as examples, because they were
beyond the actual scope of this study

"the taxon Gemma is preoccupied, but considering the
uncertainty of the this subdivision, it seems unjusti-
fied at present to replace the name

“other Antillean subgenera are not considered

Helicina is characterized by the embryonic
shell structured with pits arranged in concen-
tric lines, the absence of a provaginal opening
(1.е., monaulic), and an externally subdivided
bursa copulatrix in the female reproductive
system.

In Alcadia, the embryonic shell exhibits
more or less strong oblique grooves and
coarse, irregularly spaced radial threads.
The provaginal opening of the female sys-
{ет is present (1.е., diaulic) with an elon-
gated provaginal duct. The bursa copulatrix
is an oblong sac that is externally not dis-
tinctly lobed. Other examples included West
Indian subunits (Palliata, Idesa, and the spe-
cies Alcadia jamaicensis, formerly associ-
ated with Helicina) that will not be judged as
to their final status, but they do show a simi-
lar embryonic shell structure, although much
more strongly developed and have, in prin-
ciple, the same arrangement of the female
organs.

In Helicina s.s., the embryonic shell is very
densely sculptured with large pits. The bursa
copulatrix is the predominant accessory organ
of the apical complex in the female reproduc-
tive system. It is complexly subdivided,
whereas the provaginal sac appears simplified
and much reduced. The ascending limb of the
V-organ is elongated.

With respect to the above-mentioned char-
acteristics the subgeneric units Tristramia,
Oxyrhombus, Pseudoligyra, “Cinctella”,
Succincta and Oligyra were confirmed in their
association with Helicina.

The species group “Gemma” sensu Wagner
(1908) sharing shell characteristics with sub-

CLASSIFICATION OF HELICINIDAE 427

groups of A/cadia is now clearly distinguished
from that genus and belongs to Helicina s.l.

Tamsiana, the second group in questionable
position, could not finally be assessed in its re-
lationship since adequate material was not
available. A single embryonic shell studied
(ZMB 103314) is partially eroded, and the sur-
face can only be described with uncertainty as
very scarcely pitted and crossed by very slight
oblique lines. The figure of the female system
given by Baker (1922a) shows a much re-
duced bursa copulatrix and a very large
provaginal sac. Assuming the provaginal ori-
fice to be incorrect, these characteristics
would approach those of Analcadia and sub-
ordinate Tamsiana to Helicina, but a reexami-
nation of the female system is still required.

The northern South American subunits
Analcadia and Sericea, both studied by the re-
spective type species, have to be rearranged
from Alcadia to Helicina. They have a less
densely pitted embryonic shell in common
with a tinge of oblique lines and a strongly
enlarged provaginal sac with a basal append-
age. Furthermore, Analcadia and Sericea
share the feature of a hairy periostracum,
which is absent in other subgroups of Helicina.
These similarities probably indicate a close
relationship of both taxa, but, on account of
the differently developed bursa copulatrix and
the conspicuous sac on the middle portion of
the pallial oviduct of Helicina (Analcadia)
dysoni, the taxa are tentatively recognized as
separate subgenera of Helicina.

Study of the Brazilian species “Helicina”
brasiliensis, closely related to the type species
of Angulata (Helicina angulata), shows a very
different embryonic shell structure (broad, regu-
lar, concentric lines instead of pits or oblique
grooves and radial threads), thus providing suf-
ficient reason to raise Angulata to generic level.
Older available names do not seem to exist.
The anatomy of the female system resembles
Helicina with respect to the bursa copulatrix,
provaginal sac, receptaculum seminis, and the
V-organ, but unfortunately the most important
feature, the mon- or diaulic condition could not
properly be determined due to the poor preser-
vation of the material available. Preliminary
studies on other South American species sug-
gest a higher diversity of the female system
than in the Central American mainland species
of Helicina and render the presence of a
provaginal opening more likely than its ab-
sence. Beside the above-mentioned taxa
Tamsiana, Analcadia and Sericea subgrouped

to Helicina, only four other supraspecific taxa
have been based on South American species
— Angulata, Variabilis Baker, 1922; Concentrica
Baker, 1922; and Trichohelicina. Variabilis and
Trichohelicina have not yet been investigated,
the latter is discussed above under Alcadia
(Microalcadia) n. subgen. Radula and female
reproductive system of Helicina concentrica
were studied by Baker (1923, 1926). Although
the existence of a provaginal opening remains
to be re-examined, the anatomical structures
with a weakly lobed bursa copulatrix and a
strongly enlarged provaginal sac show more
similarities to the other species/ subgenera of
northern South America (Analcadia, Sericea,
Tamsiana) than to Angulata.

Because of the similarities of the female re-
productive system of Ceochasma to Helicina
and the absence of other distinguishing fea-
tures, except for the outstanding and charac-
teristic development of the deep slit-like sinus
at the suture of the body whorl, the genus is
hereby tentatively regarded as a subgenus of
Helicina.

A subdivision of the Central American main-
land species of Helicina remains difficult. The
following available, non-synonymous
supraspecific taxa (unless otherwise stated
the synonymy given in Baker (1922a) for Cen-
tral American mainland supraspecific taxa is
accepted here) have to be considered:
Oligyra, Succincta, Tristramia, Oxyrhombus,
Punctisulcata and Pseudoligyra. The preoccu-
pied “Formenkreis” names of Wagner Gemma
and Cinctella were accepted by Baker (1922a)
and type species were designated. Because
Baker treated both as synonyms, differentiat-
ing features were never formulated.

As shown above, Baker (1926) could not
find any anatomical characteristics distin-
guishing Helicina and Oligyra. Radula charac-
teristics also do not contribute much to the
differentiation of the taxa given above, for ex-
ample, Pseudoligyra is said to differ from
Tristramia only in the dentition of the C-central
(4 cusps or rounded hook).

With respect to the Costa Rican species
studied, two groups can be distinguished en-
compassing the following species:

15 group: Helicina funcki, H. pitalensis

2™ group: Helicina beatrix, H. talamancensis,

H. gemma, H. monteverdensis n. sp., H.
chiquitica

Remaining: Helicina tenuis, H. escondida п. sp.

H. echandiensis n. sp., H. punctisulcata
cuericiensis n. subsp.

428 RICHLING

The groups can be characterized as follows:
1° group: (a) embryonic shell: diameter of
pits equal to interspaces, (b) surface
structure of teleoconch with oblique di-
verging grooves, (c) cusps of marginal
teeth slowly increasing in number, (d)
provaginal sac irregularly lobed at distal
side, (e) bursa copulatrix with numerous,
often further subdivided lobes, central
axis or lobes elongated, (f) males in vol-
ume a little more than 80% of females (in
Helicina pitalensis not known).

2" group: (a) embryonic shell: in some spe-
cies diameter of pits smaller than
interspaces and less densely pitted, (b)
surface structure of teleoconch smooth,
except for fine growth lines, (c) cusps of
marginal teeth rapidly increasing in num-
ber and cusps more laterally arranged,
(d) provaginal sac smooth at distal side,
(e) bursa copulatrix with tendency to less
numerous lobes, (f) males in volume
about 62-70% of females.

Helicina tenuis, H. echandiensis п. sp. and Н.
punctisulcata cuericiensis n. subsp. fit in the
first group except for (e) the rather simple bursa
copulatrix and in (f) being intermediate between
the groups with a male's volume about 75% of
the female's (in H. echandiensis n. sp. only few
individuals investigated, in H. punctisulcata
cuericiensis n. subsp. unknown).

Helicina escondida n. sp. rather approaches
the second group except for b) a surface struc-
ture similar to that of the 1* group although very
slightly developed and in (f) being intermediate
between the groups and equal to H. tenuis with
a male's volume about 75% of the female's.

With respect to the radula, all the species
have the common trait that at least the A-cen-
tral is without well-defined cusps. Only occa-
sionally it is crenulate or, п a single specimen,
even denticulate, but not consistently for any
species. Other deviations appear rather spe-
cies-specific, for example, comb-lateral т
Helicina escondida n. sp.

A comparison of the species investigated for
the subgeneric units proposed by earlier au-
thors (the type species of Oligyra,
Oxyrhombus, Pseudoligyra and “Gemma”,
possibly related species for Tristramia and
Punctisulcata, for Succincta and “Cinctella”
only literature data were available) does not
resolve clearly differentiated groups. At this
level, the detailed embryonic shell structure
does not seem to be applicable, because it
already intergrades among the groups of the

Costa Rican species otherwise separated by
different characteristics. For the female repro-
ductive system, two main trends can be recog-
nized in the relative development of the bursa
copulatrix and the provaginal sac and its stalk,
but intergrades can also be found among the
Costa Rican species. On one hand, the bursa
is relatively large and more complex in its
structure, and the provaginal sac 1$ long-
stalked and irregularly lobed at its distal side
or end. On the other hand, the bursa
copulatrix is simply subdivided and more or
less reduced in the number of lobes, and the
provaginal sac is smooth and more distinct in
its outline. The first trend 1$ significantly found
in Tristramia and Oxyrhombus, “Cinctella” and
to a lesser extent also in Punctisulcata and
Pseudoligyra, sharing also the teleoconch
surface structure of oblique diverging grooves.
This group most closely resembles Helicina
s.s. from the West Indies. The other trend is
developed in “Gemma” in combination with a
very smooth shell. Oligyra and Succincta rep-
resent an intermediate stage, with a lobed
provaginal sac and a remarkably reduced
bursa copulatrix (although not to the same
degree within different populations of Helicina
orbiculata). Furthermore, they share the fea-
ture of an enlarged receptaculum seminis and
a shell sculptured with spiral grooves. Addi-
tionally, at least H. orbiculata exhibits the pat-
tern of oblique diverging grooves on the
teleoconch. According to Baker (1928),
Helicina (Succincta) flavida combines similar
characteristics.

Data of Baker (1928) allow the interpolation
of the degree of sexual dimorphism for Helicina
zephyrina (to Tristramia) of about 82% (portion:
male's of female's volume) resembling the 1*
group of the Costa Rican species.

Summarizing, the features of the embryonic
shell and female reproductive system are
helpful on the generic level, rather than for dif-
ferentiating within this group of species, al-
though the latter characteristics show
tendencies that are probably worth following
up for other species. Except for the trends in
the denticulation of the marginals, which
seems to be more influenced by the specimen
size, differences in the radula appear more
subjective rather than objective, or they are
limited to single species. Although in the data
predominantly limited to Costa Rican species,
a correlation of shell similarities and the de-
gree of sexual dimorphism is obvious. This
fact provides evidence that these characteris-

CLASSIFICATION OF HELICINIDAE 429

tics, especially the features of the shell, are
useful in recognizing relationships at the
subgeneric level when the assignment to the
genus is also verified by other features.

Considering the practical taxonomical ne-
cessity of assigning the Central American
mainland taxa to certain subgroups and since
other subgenera of Helicina can be recog-
nized properly, although final definitive differ-
entiating characteristics could not be found,
the proposed arrangement 1$ tentative and
follows the similarities outlined above.
Tristramia, Oxyrhombus, Pseudoligyra and
2”Cinctella” (the latter not studied) are as-
sumed to be synonymous including the 1*
group of the Costa Rican species. The name
Tristramia has priority. Furthermore, Oligyra
and Succincta are regarded as synonymous
with Oligyra being the older name, but Oligyra
and Tristramia diverge from the group of
“Gemma”, which encompasses the 2" group
of the Costa Rican species. This name will
tentatively be used for the separate subgenus,
although it will have to be replaced in case this
subdivision must be modified by additional
data. Presently, the proposal of a new name
seems inappropriate.

The well-defined differences between
Helicina and Alcadia and other subgenera
mentioned above, together with the uniformity
among the species from the Central American
mainland with respect to otherwise distin-
guishing features, suggest a much closer re-
lationship within the mainland species than
was previously assumed. The genus Alcadia
has been shown to be absent from the Central
American mainland and northern South
America, except for the newly discovered
small species Alcadia (Microalcadia)
hojarasca and A. (M.) boeckeleri, which is dis-
tinguished from the Alcadia s.s. and examples
investigated from other West Indian species
by the peculiarities outlined in the description
of the new subgenus. The presence of only a
limited number of small-sized species of a
genus on the mainland, as is here shown for
Alcadia, is paralleled in the genus Lucidella
and the new world Vianinae (according to the
definition of Thompson, 1980; the subfamilial
arrangement will not be discussed here) with
their main radiation in the West Indies. In this
case, only the species L. lirata and L. midyetti
Richards, 1938, or Pyrgodomus microdinus
and Р simpsoni respectively occur оп the
mainland. By way of contrast, Helicina repre-
sents the predominant genus on the mainland

and is spread over the West Indian Islands,
although the exact distribution still remains
subject to further studies. This is due to the
previous confusion with A/cadia according to
the characteristics of the post-embryonal shell
and the operculum, upon which the only clas-
sification including species of the West Indian
fauna had been based. But the species
Helicina platychila from Dominica (Lesser
Antilles), included as an example, and the
type species Helicina neritella from Jamaica
(Greater Antilles) clearly confirm the wider dis-
tribution.

Other Central American Mainland Genera

Due to insufficient material, the genus
Pyrgodomus could not be examined for fea-
tures of the female reproductive system, but
the similarities in shell shape and surface
structure, embryonic shell and radula confirm
the close relationship to the Antillean genus
Eutrochatella. Especially the size of the em-
bryonic shell appears to be characteristically
reduced in these genera compared with
Helicina and Alcadia. The examination of the
type species of Eutrochatella revealed
monaulic conditions in the female reproductive
system rendering a closer affinity to Helicina
likely. Up to now there has been no evidence
for assuming that the monaulic condition
evolved more than once. The different embry-
onic shell structure of Helicina as well as the
radula characteristics of Eutrochatella were
discussed as being subject to convergent de-
velopments. Whether or not the differentiation
of Pyrgodomus at the generic level is justified,
depends on further investigations of the West
Indian species and the final re-examination of
the anatomy of Pyrgodomus. The traditional
treatment is therefore tentatively maintained,
although, according to the present data, the
divergence from Eutrochatella probably does
not exceed those differences of the mainland
subgenera of Lucidella and Alcadia to their
West Indian subunits.

The same applies to Lucidella as to
Pyrgodomus, namely that the main portion of
the species inhabits the West Indian Islands
and only a few species, such as Lucidella
lirata, occur on the mainland. Therefore, the
discussion will be limited to the typical subge-
nus and Perenna based on the presently in-
vestigated species from Costa Rica.
According to the system of Keen (1960),
which is adopted here, two additional subgen-

430 RICHLING

era, Poenia and Poeniella, are established on
a species from Jamaica or the Lesser Antilles
respectively. The investigation of the female
reproductive system of Lucidella lirata as well
as of L. aureola required the corrections of
important details given by Baker (1926, 1928).
Therefore, it does not only confirm the affini-
ties of Perenna to Lucidella but also allows the
clear differentiation of the genus by peculiar
characteristics of the female anatomy.
Lucidella lacks the receptaculum seminis on
the inner side of the descending limb of the V-
organ, which bears apical swellings, but pos-
sesses an additional sac-like structure for
sperm storage at the posterior portion of the
pallial oviduct. In the absence of the recep-
taculum seminis and in the shortness of the
provaginal duct, Lucidella resembles
Schasicheila. The differences in the embry-
onic shell structure outlined by Thompson
(1982) were confirmed and, as in other gen-
era, are parallel in the anatomical features.
Furthermore, the investigation of the internal
shell structures showed the peculiar attach-
ment of the right portion of the retractor
muscle on the penultimate whorl and a com-
parable long axial cleft. As stated above,
Perenna, although diverging in shell shape,
generally agrees with Lucidella, s.s., with re-
spect to embryonic shell and female system,
but the bursa copulatrix is more closely asso-
ciated with the stalk of the provaginal sac than
directly with the reception chamber. Addition-
ally, the posterior portion of the pallial oviduct
is less inflated and internally folded, thus pro-
viding further reasons for retaining the
subgeneric separation.

Finally, the genus Schasicheila does not
occur in Costa Rica and seems to be limited to
Mexico and Guatemala, but it is included here
to take all Central American mainland genera
of the Helicinidae into account. Schasicheila is
characterized by several peculiarities of the
postembryonic shell and operculum (Summa-
rized by Wagner, 1907-1911). Its radula does
not diverge remarkably from the typical den-
ticulated type of Helicina, for example. Con-
cerning its anatomy, Baker (1926, 1928)
recognized the genus as one of the most ab-
errant groups of Helicinidae. Reexamination of
the type species confirmed all of Baker’s ob-
servations, especially with respect to the
diaulic condition of the female reproductive
system. The embryonic shell structure and the
internal shell structure added further distin-
guishing features.

ACKNOWLEDGEMENTS

| am deeply indebted to Dr. Wolfgang
Bockeler for giving me the opportunity to dedi-
cate my thesis to the helicinids and for all his
continuing support during this time.

It is a great pleasure for me to thank Dr.
Vollrath Wiese for the numerous discussions
and the never-ending assistance on all as-
pects of my research and for placing the vari-
ous facilities of his museum at my disposal.

Dr. Zaidett Barrientos, Malacologia of the
INBio, allowed me to study the helicinids in the
collection of INBio and helped me to obtain the
permits for my field work in Costa Rica. She
and Maribel Muniga sacrificed their time to
trace down some obscure, old collecting sites
for me. Julio Magaña Cubillo provided help in
many practical things. Socorro Avila Araya
and Alexander Alvarado-Mendez accompa-
nied me for some days in the field.

Dr. Fred G. Thompson kindly allowed me to
study the comprehensive collection of the UF
and greatly facilitated my stay at Gainesville;
John Slapcinsky gave practical help, espe-
cially with respect to the search for Helicina
orbiculata and in handling the subsequent
loans to me, Liath Appleton shared her house
for that time with me; all of them greatly con-
tributed to the nice time | had in Gainesville.

Dr. David Robinson always impressed me
by his immediate and frank help. Furthermore,
he allowed me to include his Costa Rican
record of Helicina flavida.

Dr. Matthias Glaubrecht, Berlin, offered me
his time for an occasional discussion once in
a while and provided access to the collection
and library in his charge.

| thank the following curators and private
people for making material of their collection
available for my studies, although not all mate-
rial has finally been included in this study: Dr.
Zaidett Barrientos (Santo Domingo), Dr. Robert
Cowie (Honululu), Steffen Franke (Düsseldorf),
Dr. Matthias Glaubrecht (Berlin), Zamira
Guevara М. (Nicaragua), Dr. Margret Gosteli
(Bern), Shingo Habu (Japan), Dr. Jean-Paul
Haenni (Neuchatel), Dr. Bernhard Hausdorf
(Hamburg), Dr. Robert Hershler (Washington,
D.C.), Dr. Ronald Janssen (Frankfurt/Main),
Regina К. Kawamoto (Honolulu), Dr. Günther
Kohler (Frankfurt), Wim Maassen (Leiden),
Dr. Igor Muratov (Philadelphia), Dr. Fred
Naggs (London), Prof. Dr. Adolf Riedel
(Warszawa), Dr. David G. Robinson (Phila-
delphia), Dr. Menno Schilthuizen (Malaysia),

CLASSIFICATION OF HELICINIDAE 431

FIG. 335. A-C. Helicina funcki. A. Río Barbilla. В. Manzanillo. С. Santa Elena. D-E. H. pitalensis. D.
Bajo Bonito. E. Peninsula de Osa. Е-1. H. tenuis. F-H. Cabo Blanco. |. La Selva. J-K. H. echandiensis
п. sp., campamento Echandi. L-M. H. punctisulcata cuericiensis п. ssp., Estación Cuerici; scale bars
4 mm (A-E), 3 mm (F-M).

43: RICHLING

FIG. 336. A. Helicina beatrix beatrix, Guayacán. B-C. H. b. confusa. В. Uatsi. С. Shiroles. D. Н. b.
riopejensis п. ssp., Rio Peje. Е. H. talamancensis, Bajo Bonito. F-H. H. gemma. Е. Cacao. С. Las Pavas.
H. Siquirres. |-J. H. monteverdensis п. sp., Monteverde. К-М. H. escondida n. sp., Río Barbilla. N. H.
chiquitica, Río Barbilla. O. Alcadia hojarasca, Mirador Gerardo. Р. A. boeckeleri, Pitilla. Q. Lucidella lirata,
Cahuita. В. Pyrgodomus microdinus, Fila de Cal; scale bars 3 mm (А-М), 2 mm (N-P), 1.2 mm (Q-R).

CLASSIFICATION OF HELICINIDAE 433

FIG. 337. Living animals of. A. Helicina funcki, Cahuita. В. H. funcki, juvenile, Uatsi. С. H. pitalensis,
Bajo Bonito. D. H. tenuis, Cabo Blanco. E. H. tenuis, La Selva. F. H. beatrix confusa, Uatsi. С. H.
beatrix confusa, Shiroles (photograph: V. Wiese). H. H. beatrix riopejensis п. ssp., Río Peje.

434 RICHLING

FIG. 338. Living animals. А. Helicina beatrix beatrix, Guayacän. В. H. talamancensis, Bajo Bonito. С.
H. gemma, Cacao. D. H. gemma, Las Pavas. E. H. gemma, Siquirres. F. H. monteverdensis n. sp.,
Monteverde. G. H. monteverdensis n. sp., Mirador Gerardo. H. H. escondida n. sp., Shiroles.

CLASSIFICATION OF HELICINIDAE 435

FIG. 339. Living animals. А. Helicina escondida п. sp., Shiroles. В. H. escondida п. sp., Rio Barbilla.
С. Н. chiquitica, Rio Barbilla. D. H. chiquitica, Rio Pacuarito. Е. Pyrgodomus microdinus, Fila de Cal
(photograph: V. Wiese). Е. Alcadia hojarasca, Mirador Gerardo. С. A. boeckeleri, Pitilla. H. Lucidella
lirata, Cahuita.

RICHLING

bi

FIG. 340. A. Helicina neritella, Jamaica. B. H. platychila, Dominica. C. H. orbiculata, Florida. D. H.
turbinata, Mexico. E. H. amoena, Guatemala. Е. H. dysoni, Trinidad & Tobago. С. Н. sericea, Suriname.
H. Angulata brasiliensis, Brazil. |. Alcadia major, Jamaica. J. A. hollandi, Jamaica. К. A. jamaicensis,
Jamaica. L. A. rotunda, Cuba. M. Eutrochatella pulchella, Jamaica. N. Lucidella aureola, Jamaica. O.

Schasicheila alata, Mexico; scale bar 5 mm.

CLASSIFICATION OF HELICINIDAE 437

Enrico Schwabe (München), Dr. Luis Ricardo
Simone (Sao Paulo), John Slapcinsky
(Gainesville), Dr. John Stanisic (South
Brisbane), Dr. Fred G. Thompson (Gainesville),
Dr. Vollrath Wiese (Cismar), Prof. Dr. Andrej
Wiktor (Wroclaw), and Richard Williams
(London).

Stefan Pfeiffer, Center for Microscopy of the
Biozentrum, University of Kiel, shared his
great knowledge about the SEM techniques
enabling me to make satisfying SEM photo-
graphs of helicinid radulae. Antje Thomas,
Zoological Institute, introduced me into histo-
logical methods.

| am greatly indebted to David Hingston,
Hamburg, for the thorough English correction.
Jürgen Guerrero Kommritz, Hamburg, trans-
lated the Spanish abstract.

This study was made possible through a
doctoral scholarship of the Studienstiftung des
Deutschen Volkes e. V.

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Revised ms. accepted 14 September 2003

RESEARCH NOTES

MALACOLOGIA, 2004, 45(2): 443-450

REPRODUCTIVE PERIOD AND GROWTH RATE OF THE FRESHWATER SNAIL
HELEOBIA PARCHAPPII (d'ORBIGNY, 1835) (GASTROPODA: RISSOOIDEA)
INA SHALLOW BRACKISH HABITAT (BUENOS AIRES PROVINCE, ARGENTINA)

Claudio С. De Francesco'& Federico |. Isla?

ABSTRACT

Heleobia parchappii (d’Orbigny, 1835) is a rissooidean snail very widespread in freshwa-
ter environments of Argentina, occurring at salinities < 1%o. п 1998, we found a population
of this species inhabiting a brackish canal (10-34%) located close to the headwaters of the
Mar Chiquita coastal lagoon (37%31'07"S, 57%18'30"W, Buenos Aires, Argentina). The aim
of the present work 1$ to analyse the reproductive period and growth rate of this species
in this saline condition. Monthly collections were made from August 1998 to August 1999.
Two hundred snails per sample were measured and size-frequency distributions (SFD)
constructed. Data on reproductive periods, growth rates and shell sizes were estimated
from comparisons of SFD diagrams along the year. Our results show that H. parchappii is
able to develop permanent populations in brackish waters. The species exhibits a seasonal
cycle of reproductive activity from spring to fall, with two main spawning peaks. Maximum
growth rate takes place in fall, whereas it is lower during the rest of the year, and snails
attain smaller sizes than in freshwater environments. These results may explain the great
abundance of autochthonous concentrations of H. parchappii in past estuarine habitats from
the same region.

Key words: Heleobia parchappil, reproductive period, growth rate, brackish water, Buenos

Aires Province, Argentina.

INTRODUCTION

Heleobia parchappii (d'Orbigny, 1835) is a
rissooidean snail that is very widespread in
freshwater environments of Argentina, being
particularly abundant in rivers, creeks, shal-
low lakes, ponds and streams of the Pampa
Region (Gaillard 8 Castellanos, 1976;
Castellanos & Landoni, 1995). In these fresh-
water habitats, the species has an annual
cycle of reproductive activity with develop-
ment direct to a benthic juvenile. Research
on reproduction indicates that it displays a
high natality rate towards the late spring and
a minor peak in winter (Cazzaniga, 1981a).
During this period, egg capsules are laid in
the substratum and over shells of the same
species (Cazzaniga, 1982). Snails occur on
different substrata, such as submerged veg-
etation, pebbles, muds and serpulid reefs
(Cazzaniga, 1981a, b; Darrigran, 1995; De
Francesco & Isla, 2003). They are omnivo-

rous, preferentially feeding on the epipelic
diatoms, ostracodes, rotifers, ciliates and
chironomid larvae associated to the periphy-
ton (Cazzaniga, 1981b).

The genus Heleobia Stimpson, 1865, be-
longs to the family Cochliopidae Tryon, 1866,
according to Wilke et al. (2001). However,
this genus was included for many years in
the worldwide family Hydrobiidae Troschel,
1857, and there are some authors who main-
tain that they should remain in this family
(see Liu et al., 2001). Nevertheless, snails
resemble hydrobiids in general features of
head/foot and genitalia, as well as in distribu-
tional ecology and biology (De Francesco,
2002). For that reason and to avoid future
taxonomic confusion, we use here the infor-
mal term hydrobioid (that groups all
rissooidean snails) for general considerations
(Kabat & Hershler, 1993), being clear that the
studied snails may belong to either of these
two rissooidean families.

"CONICET - Facultad de Ciencias Exactas y Naturales, Universidad Nacional de La Pampa, Uruguay 151, 6300 Santa

Rosa, Argentina; cgdefrancesco@exactas.unlpam.edu.ar

¿CONICET - Centro de Geología de Costas y del Cuaternario, Universidad Nacional de Mar del Plata, CC 722, 7600 Mar

del Plata, Argentina; fisla@mdp.edu.ar

444 ОЕ FRANCESCO & ISLA

Heleobia parchappii is also found in abun-
dance in holocene freshwater sequences of
the Pampa Region, accompanied by other
freshwater snails, such as Chilina parchappii
(d'Orbigny, 1835), Biomphalaria peregrina
(d'Orbigny, 1835), B. tenagophila (d'Orbigny,
1835), Lymnaea viatrix (d'Orbigny, 1835), and
Pomacea canaliculata (Lamarck, 1822)
(Ameghino, 1889; Frenguelli, 1945a, b; De
Francesco & Prieto, 1999; Zárate et al., 1998).
Although H. parchappii does not inhabit estua-
rine environments (De Francesco & Isla,
2003), it is commonly found in estuarine se-
quences outcropping along the southwestern
Atlantic coast. Here, H. parchappii gives rise
of extensive monospecific deposits (Isla et al.,
1986; Farinati & Zavala, 1995; De Francesco
& Zarate, 1999; De Francesco, 2002;
Espinosa et al., 2003; De Francesco & Isla,
2003). Variations in stable isotope composition
of these fossil shells suggested that H.
parchappii may have been adapted in the past
to wide salinity fluctuations as a consequence
of the drying of small ponds (Bonadonna et
al., 1995). Ecological studies, however, indi-
cate that H. parchappii cannot settle perma-
nent populations in mesohaline environments

ra |
Rio de La Plata 3
|
T
+ MAR DEL
PLATA
S
: &
MIRAMAR ©
RY
On
y
NECOCHEA 0 20 100km
59° 00’ W 575 00' W

FIG. 1. Мар of the sampling site.

38° 30'5

(Cazzaniga, 1982). In such conditions, size
decreases and density fluctuates greatly
(Cazzaniga, 1981a). These differences be-
tween the habitat of living and fossil snails
have led to controversial interpretations and,
elsewhere, to different paleoenvironmental re-
constructions. In this regard, the good preser-
vation, wide range of sizes and low
fragmentation observed in H. parchappii fossil
shells suggest autochthonous concentrations
deposited under low energy estuarine condi-
tions (Farinati & Zavala, 1995; De Francesco
8 Zárate, 1999; Espinosa et al., 2003). How-
ever, their presence in littoral marginal depos-
its where today they are absent leads some
authors to interpret them as representing
allochthonous material (Aguirre & Farinati,
2000; Aguirre 8 Urrutia, 2002) transported by
currents from creek and river headwaters.
Fortunately, the finding in 1998 of an artificial
canal with a high salinity content inhabited by
H. parchappii provided an opportunity to
analyse the reproductive biology and ecology
of this species in this saline condition. In pre-
vious work (De Francesco & Isla, 2003), we
observed that H. parchappii was present dur-
ing the whole sampling period in this brackish

Mar ee
Chiquita | к.
Сапа!

"Las
° Gallinas
: creek *

Mar
Chiquita

Coastal
Lagoon

НЕЕ CYCLE ОЕ HELEOBIA PARCHAPPII IN BRACKISH WATER 445

canal at mean values between 17% and 23%
(even up to 34%), and pointed out that this
species should be classed equally as a fresh-
water-brackish species. The purpose of this
paper is to analyse in more detail the basic as-
pects of its life cycle in order to demonstrate
the capacity of this species to tolerate and
settle permanent populations in brackish wa-
ters.

MATERIAL AND METHODS

The brackish canal (87°31 07"S,
57°18'30"W) is located close to the mouth of
the Las Gallinas creek (Fig. 1), a shallow
stream that flows into the headwaters of the
Mar Chiquita coastal lagoon, Buenos Aires
Province. It is an artificial canal, approximately
1 min depth and 3 т in width, characterised
by a typical brackish fauna. Reef-like aggre-
gates of the serpulid polychaete Ficopomatus
enigmaticus (Ten Hove & Weerdenburg,
1978), crabs and shorebirds are among the
most abundant organisms. Although the canal
is located far from the marine influence, it is
carved into holocene marine deposits (Fasano
et al., 1982). Consequently, salinity remains
relatively high, varying in relation to the bal-
ance between precipitation and evaporation in
the area.

Snails were collected from reefs of Е
enigmaticus using three replicate cores of
6 cm diameter x 11 cm height. Samples were
sieved gently through 0.35 mm mesh (45
ASTM), and placed in plastic cups. All plastic
cups were filled with water from the canal in
order to keep snails alive for a correct taxo-
nomic determination in the laboratory. Monthly
collections were made from August 1998 to
June 1999. Water temperature, salinity, pH,
and dissolved oxygen at the time of sampling
were also measured (De Francesco & Isla,
2003). In the laboratory, 200 snails per sample
were measured under a Wild M5A stereo-
scopic microscope and size-frequency distri-
butions (SFD) constructed. Total shell length
(the distance from the apex to the anterior
margin of the aperture) was used as an esti-
mate of size. Estimation of reproductive peri-
ods, growth rates and shell sizes were made
from comparisons of SFD diagrams during the
year. Climatic conditions did not allow us to
sample in September 1998 and February
1999. In addition, data on water temperature
and salinity could not be recorded in March

1999 because the canal became dry. All speci-
mens are housed in the Micropaleontology
Laboratory of the Coastal Geology Research
Centre (University of Mar del Plata).
Size-frequency distributions were analysed
to recognise dominating size groups using the
procedure described by MacDonald & Pitcher
(1979). This analysis, termed MIX, uses initial
estimates of the distribution mixture by maxi-
mum likelihood to find the best fit to the data
assuming a normal distribution for each com-
ponent. The program estimates the proportion
of the mixture and the means and standard
deviations of the component distributions, as
well as the goodness of fit of the model to the
data. Results from the MIX were employed to
separate cohorts and follow cohort-specific
growth. Growth pattern was determined by
graphical analysis of the progression of de-
tected components in successive SFD.

RESULTS

Monthly size-frequency distributions of
snails fitted a normal distribution in August,
November, December 1998, and January
1999, whereas they were bimodal during the
rest of the year (Fig. 2). Three cohorts were
detected (Table 1, Fig. 3A). The breeding pe-
riod of H. parchappii took place from spring to
fall (Figs. 2, 3A). Two main reproductive peaks
were recorded, one in early spring and the
other during the fall. Egg capsules, however,
were present during the whole period, at-
tached to shells of living specimens and over
the reef surface. The second peak was more
important, according to the higher abundance
of youngest snails found (Fig. 2). Both peaks
started at temperatures of 20°C and salinities
between 20% and 22%o (Fig. ЗВ). In general
terms, temperature and salinity varied during
the whole sampling period between 9.45°C
and 26.25°C and 10.6% and 34%, respec-
tively (Fig. 3B). The pH was very basic, vary-
ing between 7.66 and 9.15 (De Francesco &
Isla, 2003).

From August 1998 to January 1999 (late
winter-summer) the growth rate of cohorts 1
and 2 was 0.40 mm month”. There was a ре-
riod of minor growth between November and
December (0.08 mm month") that was only
recorded in cohort 2 because of the disap-
pearance of cohort 1. This growth trend corre-
lated with the variation in water temperature
(Fig. 3B). Significant growth was not recorded

446 DE FRANCESCO & ISLA

between January and March 1999 (0.05 mm
month"), which coincided with the local
drought. After that, the growth rate of cohort
2 rose to the highest values of the cycle (0.52
mm month") from March to Мау 1999 (fall).
During the same period, the mean size of ju-
veniles from cohort 3 decreased due to mor-
tality. Finally, during the late fall and early
winter, the growth rate of cohort 3 was low
(0.085 mm month"), and the size of cohort 2
gradually decreased (mortality) (Fig. 3A).
Size of adult snails varied between 2.11 +
0.64 mm and 4.02 + 0.82 mm (Table 1),
which corresponded to specimens of 4 and 5
whorls (Fig. 4). The newly hatched snails
found in October 1998 and March 1999 at-
tained sizes between 0.65 + 0.28 mm and
0.83 + 0.62 mm (Table 1). Density of snails
varied between 0.2 and 4.1 individuals per
cm’ during the year.

AUG 1998

cli.

OCT 1998

one. aa A Е

NOV 1998

Number of individuals

Jl.

dll.

DISCUSSION

Our results show that H. parchappii is able to
develop permanent populations in brackish
waters with mean salinities between 17% and
23% during the year. In this circumstance, the
species exhibits a seasonal cycle of reproduc-
tive activity from spring to fall, with two main
spawning peaks. This reproductive pattern is
very similar to that of the congeneric species
H. conexa (Gaillard, 1974) found in Mar
Chiquita coastal lagoon, which is mostly influ-
enced by water temperature (De Francesco,
2002). Cazzaniga (1981a) found a similar re-
productive pattern with two spawning peaks in
H. parchappii from freshwater environments of
the southern Buenos Aires Province. This
similarity observed in life cycles under differ-
ent saline conditions invites some speculation
about the relative importance of this factor in

DEC 1998 APR 1999

“ooo loa

JAN 1999 MAY 1999

|
La

MAR 1999 JUN 1999

Shell length (mm)

FIG. 2. Size-frequency distributions of Heleobia parchappii in the Mar Chiquita canal.

НЕЕ CYCLE OF HELEOBIA PARCHAPPII IN BRACKISH WATER 447

TABLE 1. Mean shell length, standard deviation and proportion of each cohort of Heleobia parchappii by
month in the Mar Chiquita canal based upon MIX analysis of percent size-frequency histograms (Prop =
proportion of a component, SL = mean shell length, SD = standard deviation, X* = goodness-of-fit test
between observed and expected, df = degrees of freedom, p = significance level for goodness-of-fit test).

Cohort 1 Cohort 2 Cohort 3
Month Prop SL SD Prop SL SD Prop SL SD X df п p

Aug 1998 1100 ° 22171 0.64 - - 6.13 4 200 0.19
Oct 0.23 295 1.39 0.77 0.65 0.28 é 4.43 5 200 0.49
Nov - 1.00 1.05 0.38 - 8.32 6. 200: 0:22
Dec - 1.00 1.21 0.29 - 9.66 4 200 0.05
Jan 1999 - 1.00.) 2.28 "0:70 - 12.80 5 200 0.03
Маг - 025’ 247 057 015 083 062 1.54 2 200 0.46
Арг = 0.16 327 055.084 066 028 2:39 3 200 0.50
Мау - 0.30 4.02 0.82 0.70 0.40 0.30 3.83 6 200 ‘070
Jun - 0.39 3.98 0.85 0.61 0.57 0.30 5.73 6 200 0.45

6 А

5 e Cohort 1 о Cohort2 A4 Cohort3

44 No data No data

available

ix TL

| $8 ea

Shell length (mm)
WwW

0
40 B
NE No data e Temperature (°C)
available Я о Salinity (%o)
30 |
%
© 25 E o о
5 20 8 о ®
1 tt
10 o O d e
No data
5 available
0 =}

Months

FIG. 3A. Mean shell length and standard deviation from Table 1 of each component of all samples
during the sampling period (August 1998-June 1999) based upon MIX analysis of percent size-
frequency histograms. B: Annual distribution of temperature and salinity in the Mar Chiquita canal.

448 ОЕ FRANCESCO & ISLA

conditioning the reproductive activity of snails.
Cazzaniga (1982) pointed out that Н.
parchappii experiences a marked fluctuation
in density, an unstable size structure and high
abundance of young snails in mesohaline en-
vironments. In the Mar Chiquita brackish ca-
nal, we found that even though the density
fluctuates greatly, the size structure remains
relatively stable and cohorts can be followed
monthly during the year. Thus, it appears that
salinity does not have a marked influence on
the reproductive activity of H. parchappii but
does affect population structure.

Cazzaniga (1982) found that sizes between
2.5 mm and 4.1 mm corresponded to pre-re-
productive subadults (4 whorls), while adults
ranged between 4.1 mm and 5.5 mm (5-6
whorls). The adult snails from the brackish ca-
nal show shell sizes that resemble those of
pre-reproductive subadults, whereas they co-
incide with adults in whorl number. These re-
sults demonstrate that snails from the brackish
canal attain smaller sizes than those found in
freshwater environments from the southern
Buenos Aires Province. This effect could be

Number of whorls

0 2

related to the influence of the higher salinity

content present in the canal. Salinity has been

shown to be a key environmental variable con-

trolling the growth and distribution of
hydrobioid gastropods in northern Europe.

Here, snails inhabiting brackish water reached

sexual maturity at a smaller size than those.
living in fresh water (Forbes, 1991; Jacobsen

& Forbes, 1997).

Heleobia parchappii appears to follow a spe-
cial strategy in the unstable environment of
Mar Chiquita canal. It is adapted to drastic en-
vironmental fluctuations, such as drying
events. It can be seen from the analysis of
cohorts that the size structure was not signifi-
cantly altered by the drying event recorded in
February—March 1999. Snails only showed a
temporary growth cessation but continued
their natural growth after the canal flooded
again. It has been seen that snails can remain
in an inactive condition in stressed environ-
ments for up to a month (De Francesco, un-
published data). Probably, snails remained
quiescent among the reef tubes (reefs retain
humidity) during the drying period but rapidly

a 6

Shell length (mm)

FIG. 4. Scatter diagram for the number of whorls and shell length of Heleobia

parchappii in the Mar Chiquita canal.

LIFE CYCLE ОЕ HELEOBIA PARCHAPPII IN BRACKISH WATER 449

spread as soon as climatic conditions became
favourable.

The tolerance of H. parchappii to waters with
high salinity content may explain the great
abundance of autochthonous fossil concentra-
tions of this species found in coastal outcrops
on the southeastern Buenos Aires Province
(Farinati & Zavala, 1995; De Francesco &
Zárate, 1999; De Francesco, 2002; Espinosa
et al., 2003). The reason why H. parchappi ac-
tually occurs only in freshwater environments
far away from the marine influence cannot at
present be explained. One likely factor in the
modern restriction of H. parchappii to freshwa-
ter habitats is the interspecific competition with
H. conexa. This species actually inhabits
coastal environments similar to those occu-
pied by H. parchappii during the Holocene.
However, Н. conexa was not present in this
area in the past (De Francesco 8 Zárate,
1999; De Francesco, 2002), and that probably
facilitated the access of the opportunistic H.
parchappii to estuarine environments. There is
evidence that interspecific competition signifi-
cantly affects the distribution pattern of estua-
rine hydrobioids mostly in coastal lagoons
(Fenchel & Kofoed, 1976; Cherril & James,
1987; Gorbushin, 1996; Grudemo & Bohlin,
2000; Barnes, 1999, among others).

Although it is not possible to consider the
generality of the ideas presented in this study
with the scattered observations presented
here, these data indicate that H. parchappii
has, in at least some situations, the capability
to tolerate and sustain stable populations in
environments with high salinity (up to 34%) in
spite of the fact that it generally occurs in
freshwater environments. These results dem-
onstrate a greater tolerance than 1$ usually
assumed for this species, with important con-
sequences for the reconstruction of past envi-
ronments.

ACKNOWLEDGEMENTS

This research is part of the first author's doc-
toral thesis in the University of Mar del Plata.
Financial support for this work was provided
by the project Evolución de ambientes
sedimentarios durante los ultimos 20,000
anos from Universidad Nacional de Mar del
Plata (15/EO65) and by FONCYT (N*01617).
Fieldwork was greatly aided by Luis Cortizo.
Andrea Gavio, and Sandra Obenat are grate-
fully acknowledged for assistance with the

MIX analysis. We are indebted to Mrs.
Romano for permission to sample on her
lands. We also acknowledge two anonymous
reviewers for improving this manuscript with
helpful comments. CGD was funded by the
National Council of Scientific and Technical
Research of Argentina (CONICET) through a
fellowship.

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CAZZANIGA, N. J., 1981a, Estudios bio-
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relacionados con la invasión de canales por
malezas acuáticas. Doctoral Thesis, Univer-
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CAZZANIGA, М. J., 19816, Caracterización
química y faunistica de canales de drenaje del
valle inferior del río Colorado (partido de
Villarino y Patagones, provincia de Buenos
Aires). Ecosur, 8 (15): 25-46.

CAZZANIGA, М. J., 1982, Notas sobre hidróbidos
argentinos. 5. Conquiliometría de Littoridina
parchappii (D'Orbigny, 1835) (Gastropoda
Rissoidea) referida a su ciclo de vida en
poblaciones australes. /heringia, Serie
Zoologia, 61: 97-118.

450 ОЕ FRANCESCO & ISLA

CHERRIL, А. J. & К. JAMES, 1987, Evidence for
competition between mudsnails (Hydrobiidae):
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DARRIGRAN, G., 1995, Distribución de tres
especies del género Heleobia Stimpson, 1865
(Gastropoda, Hydrobiidae) en el litoral
argentino del Río de La Plata y arroyos
afluentes. /heringia, Série Zoologia, 78: 3-8.

DE FRANCESCO, С. С., 2002. Significado
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concentraciones holocenas de Heleobia (Gas-
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Universidad Nacional de Mar del Plata, Argen-
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DE FRANCESCO, С. С. & Е. 1. ISLA, 2003, Dis-
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a mixed estuary and a coastal lagoon, Argen-
tina. Estuaries, 26 (3): 790-797.

DE FRANCESCO, С. С. & A. К. PRIETO, 1999,
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37 (4): 8R-9R. ;

DE FRANCESCO, С. С. & М. А. ZARATE, 1999,
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holocenos del rio Quequén Grande (Provincia
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paleoambiental. Ameghiniana, 36 (3): 297-310.

ESPINOSA, M. A., С. G DE FRANCESCO & Е.
|. ISLA, 2003, Paleoenvironmental reconstruc-
tion of Holocene coastal deposits from the
southeastern Buenos Aires Province, Argen-
tina. Journal of Paleolimnology, 29 (1): 49-60.

FARINATI, Е. А. & С. ZAVALA, 1995, Análisis
tafonómico de moluscos y análisis de facies en
la serie holocena del río Quequén Salado,
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del VI Congreso Argentino de Paleontología y
Bioestratigrafía, 117-122.

РАЗАМО, J., М. HERNANDEZ, Е. 1. ISLA 4 E.
SCHNACK, 1982, Aspectos evolutivos y
ambientales de la laguna Mar Chiquita
(provincia de Buenos Aires, Argentina).
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FENCHEL, T. 8 L. H. KOFOED, 1976, Evidence
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FORBES, V. E., 1991, Response of Hydrobia
ventrosa (Montagu) to environmental stress:
effects of salinity fluctuations and cadmiun ex-
posure on growth. Functional Ecology, 5: 642-
648.

FRENGUELLI, J., 1945a, El Piso Platense.
Revista del Museo de La Plata, Sección
Geología, 2: 287-311.

FRENGUELLI, J., 1945b, Las diatomeas del
Platense. Revista del Museo de La Plata,
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GAILLARD, M. C. 8 CASTELLANOS, Z. A. de,
1976, Mollusca Gastropoda Hydrobiidae. Pp.
1-40, in: R.A. RINGUELET, ed., Fauna de agua
dulce de la República Argentina. FECIC,
Buenos Aires.

GORBUSHIN, А. M., 1996, The enigma of mud
snail shell growth: assymmetrical competition
or character displacement? Oikos, 77 (1): 85-
92.

GRUDEMO, J. & T. BOHLIN, 2000, Effects of
sediment type and intra- and interspecific com-
petition on growth rate of the marine snails
Hydrobia ulvae and Hydrobia ventrosa. Jour-
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ogy, 253: 115-127.

ISLA, F. 1., J. Е. FASANO, Е. FERRERO, М. A.
ESPINOSA & E. J. SCHNACK, 1986, Late
Quaternary marine-estuarine sequences of the
southeastern coast of Buenos Aires Province,
Argentina. Quaternary of South America and
Antarctic Peninsula, 4: 137-157.

JACOBSEN, R. 8 V. E. FORBES, 1997, Clonal
variation in life-history traits and feeding rates
in the gastropod, Potamopyrgus antipodarum:
performance across a salinity gradient. Func-
tional Ecology, 11: 260-267.

KABAT, А. R. & К. HERSHLER, 1993, The
prosobranch snail family Hydrobiidae (Gas-
tropoda: Rissooidea): review of classification
and supraspecific taxa. Smithsonian Contribu-
tions to Zoology, 547: 1-94.

LIU, H. P., К. HERSHLER 8 Е. G THOMPSON,
2001, Phylogenetic relationships of the
Cochliopinae (Rissooidea: Hydrobiidae): an
enigmatic group of aquatic gastropods. Mo-
lecular Phylogenetics and Evolution, 21 (1):
17-25.

MACDONALD, P. D. & T. J. PITCHER, 1979,
Age-groups from size-frequency data: a
versatile and efficient method of analysing
distribution mixtures. Journal of Fisheries
Research Board of Canada, 36: 987-1001.

WILKE, T., G. M. DAVIS, A. FALNIOWSKI, F.
GIUSTI, M. BODON & M. SZAROWSKA,
2001, Molecular systematics of Hydrobiidae
(Mollusca: Gastropoda: Rissooidea): testing
monophyly and phylogenetic relationships.
Proceedings of the Academy of Natural
Sciences of Philadelphia, 151: 1-21.

ZARATE, М. А., М. ESPINOSA & L. FERRERO,
1998, Palaeoenvironmental implications of а
Holocene diatomite, Pampa Interserrana,
Argentina. Quaternary of South America and
Antarctic Peninsula, 12: 135-152.

Revised ms. accepted 13 July 2003

MALACOLOGIA, 2004, 45(2): 451-452

FIRST REPORT OF A TERRESTRIAL SLUG (ARION FASCIATUS)
LIVING IN AN AQUATIC HABITAT

Roger J. Haro, Rick Gillis & Scott T. Cooper

Department of Biology, University of Wisconsin-La Crosse, La Crosse
Wisconsin 54601 USA; haro.roge@uwlax.edu

The banded slug, Апоп fasciatus (Nilsson,
1823), was introduced into North America
from Europe (Chichester & Getz, 1969),
where it is commonly found in damp areas
and wet meadows adjacent to streams
(Pfleger & Chatfield, 1988). Although best
known as farm and garden pests, slugs oc-
cupy a variety of habitats, often playing a ben-
eficial role similar to that of many terrestrial
snails in the decomposition of forest litter (Ma-
son, 1974; Petersen 4 Luxton, 1982). In fact,
slugs are commonly described as “snails with-
out shells”, an appropriate moniker given that
they often replace ecologically similar species
of snails in habitats with insufficient supplies of
calcium for shell formation (Barnes, 1987).

Although there are a number of freshwater
pulmonate snails, most of these must come to
the surface to obtain air for respiration (Barnes,
1987). Kinzie (1992), however, observed the
predaceous snail Euglandina rosea (Férussac)
immersed in Hawalian streams and docu-
mented them feeding on aquatic snails in the
laboratory. Furthermore, several species of
deep-water lymnaeid snails appear to have
abandoned air breathing entirely, filling the
mantle cavity with water, and a few have even
evolved a secondary gill (pseudobranch) that
consists of series of folds of the mantle near the
pneumostome (Pennak, 1953; Barnes, 1987).
While the ability of an air-breathing pulmonate
to survive underwater was confirmed by Lowell
8, Carriker (1946) in an experiment in which 20
generations of the pond snail Lymnaea
stagnalis appressa were raised entirely under
water, no species of terrestrial slug has ever
been recorded as living in freshwater. However,
Rollo 8 Shibata (1991) did find some individu-
als of the slug Deroceras leave Müller sub-
merged on the underside of floating logs in a
flooded wetland.

We have observed A. fasciatus at three dif-
ferent sites in the headwaters of Spring Cou-
lee Creek and at one site in the headwaters of
nearby Poplar Creek, all of which are near the
village of Coon Valley (43°40’М, 90°54’W),

451

Vernon County, Wisconsin. The majority of our
observations of A. fasciatus living in the
aquatic habitat are from a single, 2 m? riffle
site in the upper reaches of Spring Coulee
Creek (USCC). These coldwater streams,
which are perennial and remain ice-free during
the winter, are located in the Driftless Area of
southwest Wisconsin, a region that was not
covered with ice during the latest period of
Pleistocene glaciation.

Although at first we thought that these aquatic
slugs might belong to an undescribed species,
dissection of the reproductive tract by J.B.
Burch of the Mollusk Division of the University
of Michigan Museum of Zoology showed them
to be A. fasciatus. Voucher specimens have
been deposited in the University of Michigan
Museum of Zoology (UMMZ 300110). This tax-
onomy was supported by genetic analysis of
PCR amplified DNA from the mitochondrial cy-
tochrome c oxidase subunit | gene (GenBank
accession number AY321295), which showed
that all 584 bases matched perfectly with those
previously described for two specimens of A.
fasciatus (GenBank accession numbers:
AY094598 from Lithuania and AF239735 from
Georgia, USA).

Slugs at the USCC site were checked at least
quarterly since being discovered in 1996 by the
first author (RJH). Arion fasciatus (normally 15-
30 individuals) are observed at USCC through-
out the year, although casual observations
suggest that slug densities decline during the
winter. Annual water temperature fluctuation at
this site is very small (~1C°), with temperatures
between 9-10°C (Deuschle, 2001). Current
velocities in this riffle are moderate and stable
throughout the year (mean + SD = 24.4 + 7.7
cm/s) (Deuschle, 2001). Dissolved oxygen con-
centrations in midsummer average 10.56 +
0.09 mg ОЛ. We believe the USCC site is well
aerated throughout the year; brook trout
(Salvelinus fontinalis) and slimy sculpin (Cottus
cognatus), two fish that require high dissolved
oxygen tensions, are commonly observed.
Slugs were observed at depths between 0.5-

452 HARO ЕТ AL.

10 cm. We have observed both adults and ju-
veniles of all sizes (total body length range:
0.7—6 cm), suggesting that these slugs may
complete their life cycles within this stream
system. Although we have not yet searched
for eggs in the stream, experiments done by
Rollo 8 Shibata (1991) revealed that the eggs
of some slug species have a high hatching
success when reared underwater.

Sites where slugs have been observed are,
with one exception, always beneath shaded
highway bridges, and no specimens have been
seen in locations exposed to direct sunlight.
Individuals occur on submerged rocks as well
as sand substrata in gently flowing water. From
the grazing trails we observed, the slugs ap-
pear to be feeding on periphyton growing on
rocks and fallen leaves. We have not observed
slugs on the stream bank or in the adjacent
woods, although several individuals were seen
on rocks partially submerged at the water-air
interface. Specimens brought into the labora-
tory and placed in an experimental flume
(Vogel 8 La Barbera, 1978) have survived over
two weeks entirely submerged (until removed
for histological examination) and made no at-
tempts during this time to climb onto the sides
of the flume or rocks that projected above the
water line. Slugs maintained for several days in
a plastic jug, however, left the water within
12 h of capture and climbed onto the sides of
the container, suggesting that the ability to re-
main submerged is dependent on a continuous
supply of flowing, well-oxygenated water.

These findings run counter to the long-stand-
ing presumption that pulmonate slugs are en-
tirely terrestrial. In spite of an extensive
literature search, we cannot find a single refer-
ence that suggest otherwise, nor are any of the
authorities in the field with whom we have spo-
ken aware of a terrestrial slug living in water. As
such, this population of stream-dwelling slugs
poses a number of interesting questions that
remain to be studied in more detail.

LITERATURE CITED

BARNES, К. D., 1987, Invertebrate zoology, 5"
ed. Saunders College Publishing, New York,
New York. 893 pp.

CHICHESTER, L.F. & L. Е, (GETZ; 1969, the
zoogeography and ecology of arionid and
limacid slugs introduced into northeastern
North America. Malacologia 7: 313-346.

DEUSCHLE, О. R., 2001, Effects of deposited
sediment on a keystone grazer, Glossosoma
intermedium (Trichoptera: Glossosomatidae)
and its associated macroinvertebrate commu-
nity in southwestern Wisconsin streams.
Master’s Thesis, University of Wisconsin — La
Crosse, 63 pp.

KINZIE, R.A., Ш, 1992, Predation by the intro-
duced carnivorous snail Euglandina rosea
(Ferussac) on endemic aquatic lymnaeid
snails in Hawaii. Biological Conservation 60:
149-155.

LOWELL, E. N. & M. R. CARRIKER, 1946, Ob-
servations on the biology of the snail Lym-
naea stagnalis appressa during twenty
generations in laboratory culture. American
Midland Naturalist 36: 467-493.

MASON, С. F., 1974, Mollusca. Pp. 551-591,
in: С. H. DICKENSON & С. J. Е. PUGH, eds., Biol-
ogy of plant litter decomposition. Academic
Press, Inc., New York.

PENNAK, R. W., 1953, Fresh-water inverte-
brates of the United States. The Ronald Press
Company, New York, 769 pp.

PETERSEN, H. & L. LUXTON, 1982, A com-
parative analysis of soil fauna populations
and their role in decomposition processes.
Oikos 39: 288-354.

PFLEGER, V. 8 J. CHATFIELD, 1988, A guide to
snails of Britain and Europe. The Наптуп Pub-
lishing Group Limited, London, UK, 216 pp.

ROLLO, C. D. & D. M. SHIBATA, 1991, Resil-
ience, robustness, and plasticity in a terres-
trial slug, with particular reference to food
quality. Canadian Journal of Zoology 69: 978-
987.

VOGEL, 5. & M. LA BARBERA, 1978, Simple
flow tanks for research and teaching.
BioScience 28: 638-643.

Revised ms. accepted 16 July 2003

MALACOLOGIA, 2004, 45(2): 453—458

POPULATION STRUCTURE IN DREPANOTREMA KERMATOIDES AND
D. СМЕХ (GASTROPODA, PLANORBIDAE) IN NATURAL CONDITIONS

Alejandra Rumi', Diego E. Gutiérrez Gregoric?,
М. Andrea Roche? 8 Mónica P. Tassara'

División Zoología Invertebrados, Museo de La Plata, Facultad de Ciencias Naturales y
Museo, Universidad Nacional de La Plata, Paseo del Bosque s/n”, 1900
La Plata, Buenos Aires, Argentina; alerumi@museo.fcnym.unlp.edu.ar

ABSTRACT

Drepanotrema kermatoides and D. cimex are present in the Multiple Use Natural Reserve
“Isla Martín García”, Argentina. Although these species are endemic to the Neotropical
Region, their biology and ecology have been little studied. The purpose of this study was
to analyze the main demographic trends of Drepanotrema in different seasons under natural
conditions. Relative abundance and population structure in nature were compared. Sea-
sonal samples were taken from six environments between 1995 and 1997. Each species
was found in three different environments. Drepanotrema kermatoides was more abundant
in The Stream and D. cimex in Tank Quarry. From the population structure observation, we
found that the dominant class corresponded to individuals at the onset of reproduction.

Key words: demography, Planorbidae, Drepanotrema, Río de la Plata Basin.

INTRODUCTION

The planorbid genus Drepanotrema 1$ en-
demic to the Neotropical Region and includes
nine species, with six found in Argentina: D.
anatinum (d'Orbigny, 1835), D. cimex (Mor-
icand, 1839), D. depressissimum (Moricand,
1839), D. heloicum (d'Orbigny, 1835), D. ker-
matoides (d'Orbigny, 1835), and D. lucidum
(Pfeiffer, 1839). While they are abundant, their
biology and ecology have been scarcely stud-
ied (Bonetto et al., 1990; Rumi, 1991; Ham-
man et al., 1993; Rumi et al., 1997), and their
morphology and classification have not re-
ceived as much attention as have those of
other planorbids, especially species of
Biomphalaria, which are important for human
health.

The purpose of this study was to analyze the
main demographic trends shown by
Drepanotrema in natural conditions in the dif-
ferent habitats and seasons by analyzing
changes in relative abundances and popula-
tion structure.

'Researchers of CONICET.
“Fellowship of CIC and UNLP.

The study was conducted in the Multiple Use
Natural Reserve “Isla Martín García” (IMG),
Argentina, where intensive biodiversity re-
search has already been conducted (Lahitte 8
Hurrell, 1988, 1994; Rumi et al., 1996;
Armendáriz et al., 2000; Armendáriz 8 Cesar,
2001). Freshwater gastropods found in the
IMG include species of Ampullariidae,
Hydrobiidae, Physidae, Chilinidae, Ancylidae,
and Planorbidae. The Planorbidae are repre-
sented by the genera Biomphalaria Preston,
1910; Antillorbis Harry 8 Hubendick, 1963;
and Drepanotrema Fischer 8 Crosse, 1880
(Rumi et al., 1996).

MATERIALS AND METHODS

The IMG 1$ located in the Upper Río de la
Plata, south of the mouth of the Uruguay River
(34°11’25”5; 58°15’38”W) (Fig. 1). It is the only
island in the Río de la Plata system that consti-
tutes an outcrop of the Brazilian massif of Pre-
cambrian crystalline basement rocks. The

“PhD candidate in Natural Sciences, Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata, Buenos

Aires, Argentina.

454 RUMI ET AL.

0 280 mts

a
F Окт

FIG. 1. Isla Martin Garcia map with sampling
stations: 1. Rubbish Quarry; 2. Tank Quarry; 3.
Big Quarry; 4. Buoy Quarry; 5. The Stream and
6. Intangible Zone.

other deltaic islands are sedimentary. Eight
seasonal samplings were carried out between
the summer of 1995 and the autumn of 1997 in
six environments (Fig. 1). One of them, the
Stream is small and drains into coastal pools. It
had a depth of no more than 1 m, and a domi-
nant floating plant carpet, especially of Pistia

stratiotes and Salvinia biloba. Another four sta-
tions sampled, the quarries used for mineral
exploitation of the crystalline basement rock
many years ago are now artificial ponds. Ex-
cept for the Rubbish Quarry (named by local
inhabitants because it is close to a garbage
dump), they were covered by carpets of free-
floating macrophytes, especially the Lem-
naceae species, Salvinia biloba, S. minima,
Azolla filiculoides, Limnobium laevigatum, and
Pistia stratiotes. The quarries showed strong
eutrophication and desiccation in summer. Fi-
nally, the Intangible Zone (restricted area) 1$ a
flooded sector, which is frequently devoid of
water. Iris pseudacorus was dominant and sur-
rounded by a xerophylous forest.

Snails were collected in the littoral zone of
the stations using sieves of 15 cm diameter and
0.14 mm mesh size. Snail relative abundance
was calculated as captures per unit effort
(CPUE), that is, specimens/30 min/person.
Maximum shell diameter of all specimens was
measured with calipers to 0.02 mm precision.
For graphic representation of seasonal popula-
tion structure for each species and environ-
ment, numbers of individuals in each 1 mm size
class were used. To estimate class differences
of each season among the years, the Student's
“t” test (two means of matching samples; two
tales) was employed. If the obtained differences
were not significant, the mean value for each
class was used. In cases when the abundance
was lower than the number of classes consid-
ered, the sample was not taken into account.

RESULTS

Two species were recorded: Drepanotrema
kermatoides and D. cimex (Fig. 2).

FIG. 2. Dorsal and lateral view of shells of A. Drepanotrema kermatoides, diameter = 5.40 mm, and B.

D. cimex, diameter = 5.25 mm.

POPULATION STRUCTURE IN DREPANOTREMA SPP. 455

TABLE 1. Drepanotrema kermatoides. Mean estimations of snail abundance by season and habitat.
(-): without samples; М: number of considered seasons. *зате value for М register.

Intangible Zone

Mean Range N Mean
Summer 0 0,3 18.33
Autumn 24 0-48 2 4.5
Winter - - -
Spring 21:9 5-50 2 53

Big Quarry Stream
Range N Mean Range N

0-55 3 314 314*
0-9 2 36 - 1

a 1 x =
4-102 22 1773 59-296 2

Drepanotrema kermatoides (Table 1, Fig. 3):
This species inhabited three very different
environments. Big Quarry (the largest on the
island), despite water-level fluctuations, had
sufficient water to maintain snail populations.
However, during most of the year, very low
densities were recorded. They were some-
what higher in the spring. Considering popula-
tion structure, adults between 4 and 6 mm
predominated. The Stream had, in summer,
the most abundant population numbers, in
spite of high fluctuations. In cases in which the
population structure/season was compared
among different years, the results did not

Intangıble Zone

Nx Nx
= 30 60
= 20 40
Е 10 Li 20
0 0?
5 30 60
= 20 40
core 10 20
0 0
op 30 60
a 20 7 40
A 10 - 20 4
0 | i 0
5 30 60
= 20 40
5 10 - 20 -
A 0 | i | | | | 0
012345678 0 1

шт

show significant differences: summer1995/
1997: t = 0, df = 8, P > 0.05 and spring: t =
2.24, df = 8, P > 0.05. Juveniles occurred al-
most year round. Finally, the Intangible Zone
maintained a very low population level. As in
Big Quarry, there was a greater representation
of adults.

Drepanotrema cimex (Table 2, Fig. 4):
Populations of this species were found in
three quarries: Tank, Rubbish and Buoy. As
with D. kermatoides, abundances varied
widely. The Tank Quarry showed the most
abundant and seasonally stable population.
The highest abundance of individuals was

Big quarry Ne Stream
120
80
40
120
80
40

80
o №
0
= y a
0

2. E EN TS

012345678
mm mm

FIG. 3. Drepanotrema kermatoides. Population structures by season.

456 RUMI ЕТ AL.

TABLE 2. Drepanotrema cimex. Mean estimations of snail abundance by season and habitat. (-):
without samples; N: number of considered seasons.

Tank Quarry

Rubbish Quarry

Buoy Quarry

Mean Range N Mean Range N Mean Range М
Summer 497 52-942 2 25.39 0-69 3 - - -
Ащитп 392 0-784 2 184.5 1-386 2 - - -
Winter 276 - 1 74 - 1 E = Е
Spring 86 50-122 2 26:5 4-49 2 52 - 1

observed in summer, decreasing slowly to-
wards the spring. Population structure/sea-
son did not differ significantly among different
years: summer 1996/1997 t = 1.86, df = 8,
Р > 0.05 and spring: t = 0.81, df = 8, Р > 0.05.
Although adult sizes dominated, all juvenile
sizes were represented. At the Rubbish
Quarry, densities were lower than at the Tank
Quarry, with D. cimex more abundant in au-
tumn. In 1995, the population was well repre-

Rubbish quarrv

Nx Nx

180 500
er 375
E ee 250
5 60 125
| 0 0
_ 180 500
2 120 375
E = 250
> 60 125
> du =
„ 180 500
Ss 150 375
= 60 250
A 2 125
180 500
ВУЛ 315
| = |

= 60 1
2 0 PRES A

AZ S24 0: 7 8
mm

Tank quarry

sented all year. However, in the summer
1996, the quarry dried out and the abun-
dance become very low. The population did
not recover. Adults and juveniles, although in
a lower proportion, were represented. The
Buoy Quarry (called that because of a large
red marine buoy half sunken in the center),
only sampled in the spring 1996, showed a
medium abundance with all sizes repre-
sented.

Buoy quarry

50 7 10
25 ee
() | = 0 1 AA ЕЯ

072343078 0123456018

mm mm

FIG. 4. Drepanotrema cimex. Population structures by season.

POPULATION STRUCTURE IN DREPANOTREMA SPP. 457

DISCUSSION

Drepanotrema kermatoides was most abun-
dant at spring and summer in The Stream.
This abundance was associated with The
Stream's shallow depth and proximity to the
river, which backed up into the stream during
floods (because of the strong SE winds,
mainly in winter), thus favoring the migratory
movements of snails, and causing high fluc-
tuations in abundance. By contrast, in the In-
tangible Zone abundance was very low. This
could be due to the fact that this environment
during dry seasons suffered total desiccation
in summers and autumn 1997 when only
empty shells were detected. Recruitment be-
tween spring and summer was drastically re-
duced because of the desiccation effects and
negating recovery of population size. The Big
Quarry (the largest on the island) with its sur-
face totally covered by carpets of free floating
macrophytes, had the lowest snail abun-
dance. This was apparently due to the sum-
mer 1995, when the quarry had considerable
plant decay, mainly produced by macrophytes,
resulting in near anoxia (0.2 mg/l), with high
concentrations of soluble reactive phosphorus
(SRP) (1,086 g/l), ammonium (338 g/l) and
organic matter (13 mg/l), and with low сопсеп-
tration of nitrates (7 g/l).

Drepanotrema cimex was most abundant
throughout the sampling periods, especially
between summer and autumn. The Tank
Quarry showed the most abundant and sea-
sonally stable population. Only once, in au-
tumn 1997, was there a strong desiccation
event.

From the observation of population structure
relative to size (Figs. 3, 4), we found that the
dominant classes (3 to 5 mm) corresponded
to the onset of reproduction. In general,
Drepanotrema species showed the greatest
recruitment in autumn and spring. These re-
sults agree with reproductive strategies found
in other freshwater mollusks in these areas of
Argentina (Hamann et al., 1993; Rumi, 1993).

The population structure (Fig. 4) in Buoy
Quarry, although having only one sample,
seemed to have two cohorts. One presumably
originated in autumn (according to the size)
and another, of adult individuals in their last
growth stages, perhaps carried over from the
previous year. Thus, it can be inferred that lon-
gevity of this species would exceed one year
in natural conditions. Other observations of
the Planorbidae (e.g., Biomphalaria peregrina

(d'Orbigny, 1835)), have also indicated
longevities of two years, although under labo-
ratory conditions (Rumi, 1993).

ACKNOWLEDGEMENTS

This study was funded by the Facultad de
Ciencias Naturales y Museo, Universidad
Nacional de La Plata, and the Agency of Sci-
entific Promotion (BID 1201/OC-AR PICT 01-
03453), and supported by the Office of Natural
Resources and Protected Areas, Ministry of
Agriculture, Cattle Raising and Feeding,
Buenos Aires Province. We specially thank to
George M. Davis for checking the English and
comments on an early draft of the manuscript.
We also thank the rangers of the IMG Reserve
for theirs valuable collaboration and assess-
ment.

LITERATURE CITED

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distribution and ecology of littoral Oligochaeta
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ARMENDARIZ, |. C., I. I. CESAR & М. С.

DAMBORENEA, 2000, Oligoquetos en
ambientes lénticos de la Reserva Natural е
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ВОМЕТТО, A. A., A. RUMI & М. P. TASSARA,
1990, Notas sobre el conocimiento limnológico
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HAMMAN, М. I., A. RUMI 8 М. OSTROWSKI de
NUNEZ, 1993, Aspectos biológicos sobre los
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Drepanotrema spp. (Mollusca, Planorbidae) en
un biotopo lenítico del nordeste argentino.
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arbórea y arborescente de la Isla Martín García.
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RUMI, A., 1991, La familia Planorbidae
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(8): 51pp. home |

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DARRIGRAN, 1996, Moluscos de agua dulce
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RUMI, A., М. P. TASSARA & А. A. BONETTO,
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1-19.

Revised ms. accepted 13 October 2003

MALACOLOGIA, 2004, 45(1-2): 459-479

Таха in bold are new; pages in italic

indicate figures of taxa.

abditus, Pyrgodomus microdinus 368-370,
368, 419-420
Abrina 169
Achatina musaecola 185-186, 185
acuminata, Ampullaria 81
acuta, Ampullaria 56, 81
Pomacea 56
adamsiana, Lucidella 412
adusta, Ampullaria 80
africana, Corbicula 3
Afropominae 42
Afropomus 42
Agrimonia eupatoria 127
alata, Helicina 408
Schasicheila 390, 408, 408-409, 420, 436
Alcadia 196-200, 203, 370, 375, 390, 400,

404, 406, 408, 412, 414, 418-427, 429

(Analcadia) platychila 196, 198
beatrix 297
boeckeleri 373-374, 374-376, 376-377,

383, 409-410, 413, 418, 426, 432, 435

hojarasca 214, 343, 371-373, 373, 375,

376-377, 383, 405, 409-410, 413, 418,

426, 432, 435

hollandi 390, 402-403, 403-405, 422, 436

jamaicensis 412, 426, 436
(Leialcadia) beatrix 283
(Leialcadia) beatrix confusa 297
(Leialcadia) beatrix nicaraguae 284
(Leialcadia) fragilis 357, 389
(Leialcadia) fragilis mohriana 347
(Leialcadia) gemma 318

major 401, 401-402, 403, 412-413, 419,

425, 436
(Microalcadia) 370, 412-413, 415, 427
(Microalcadia) boeckeleri 195, 197, 215,
274, 429
(Microalcadia) hojarasca 195, 197, 215,
370, 419, 429
palliata 213
rotunda 405, 405, 436
aldersoni, Pila (Pomacea) 56
Pomacea 56
Amauropsina 58
amazonica, Ampullaria 56
Pomacea 43, 56
americanista, Ampullaria 78
Pomella 78
amoena, Helicina 195, 197, 246, 254, 387,
396, 396-397, 409, 421, 436

INDEX

459

Amphidesma nuculoides 169
Amphidromus 413
ampla, Lymnaea 141
Radix 141-147, 143, 145
Ampliata 423
Ampluria rotundata 81
Ampullaria 42-43, 46, 49-51, 66, 73, 81,
121
acuminata 81
acuta 56, 81
adusta 80
amazonica 56
americanista 78
angulata 56
aperta 80
archimedes 57
armeniacum 57
aulanieri 57, 75
auriformis 57
aurisformis var. ocanensis 70
aurostoma 57
australis 57
autumnalis 57
avellana 57, 81
baeri 52
balteata 52
batabana 58
belizensis 58
bilineata 78
borealis 81
brasiliensis 58
bridgesii 58
brownii 46
bruguieri 80
buccinoidea 81
bulimoides 81
bulla 58
buxea 58
caliginosa 58
canaliculata 42, 58, 81
canalifera 81
cassidiformis 59
castanea 52
castelloi 59
castelnaudii 59
catamarcensis 59
cerasum 59
chemnitzii 59
chiquitensis 50
chlorostoma 52
cincta 60
cingulata 52
citreum 60

460

columbensis 60
columbiensis 60
columellaris 51, 60
conica 52, 81

conoidea 60

consolatrix 60
contamanoensis 60
cornucopia 60
costaricana 60

cousini 61

crassa 46-47, 81

crassa var. monticola 47
crassatina 81
crocostoma 52
crosseana 61

cubensis 61

cumingii 61

cuprina 42, 53
cyclostoma 47

dacostae 61

decussata 61

decussata var. commissionis 60
delattrei 61

depressa 62, 81
disseminata 62

dolioides 62

dolium 62

dorbignyana 62
dorbignyi 62

dubia 53

dysoni 62

effusa var. conica 52
electrina 63

elegans 63

elongata 81

erogata 63

erronea 63
erythrostoma 63
eumicra 63

excavata 81

exigua 80

eximia 63

expansa 53

fasciata 51, 53, 63-64, 75, 79
fasciata var. dilatata 62
fasciata Var. elongata 63
faujasii 81

(Felipponea) elongata 49
(Felipponea) neritiniformis 49
ferruginea 64

figulina 64

figulina var. semperi 75
flagellata 64

flagellata var. guatemalensis 65
flatilis 64

INDEX

fragilis 81

fumata 64
galloprovincialis 81
georgii 64

geveana 53

geveana var. suprafasciata 55
gevesensis 53
ghiesbreghti 64
gibbosa 79

gigantea 65

gigas 65

gigas var. unicolor 76
gossei 65

granulosa 47
guadelupensis 44, 54
guaduasensis 65
gualtieri 65

guinaica 71

guyanensis 65
haemastoma 65
hanleyana 66

hanleyi 66

haustrum 66

hepataria 79
hondurasensis 66
hopetonensis 66
hybrida 81

immersa 66

imperforata 79

impervia 47

innexa 66

insularum 66
intermedia 54

interrupta 66

intropicta 67

knorrii 51

labiosa 67

laevigata 81

lamarckii 67

lattrei 67

lattrei chamana 59
lemniscata 67
leucostoma 67

levior 67

linnaei 67

livescens 67
luteostoma 51, 54
lymnaeaeformis 67
malleata 68

malleata var. arata 57
malleata var. chiapasensis 59
malleata var. exculpta 63
malleata var. oajacensis 70
malleata var. prasina 73
malleata var. strebeli 75

INDEX

marginatra 68
martinezi 68
media 81
megastoma 78
melanocheila 68
mermodi 69
meta 69
metcalfei 69
mexicana 69
miamiensis 69
miltocheilus 69
miltochilus 69
modesta 70
monachus 70
monstrosa 70
naticoides 48
neritoides 78
nigrilabris 70
nobilis 70
notabilis 70
novaegranadae 70
nubila 48
nucleus 79
oblonga 70
obtusa 79
ocanensis 70
occlusa 71
ochracea 71
olivacea 71
olivieri 48
orinoccensis 55
ormophora 48
oviformis 71
pachystoma 55, 79
pallens 80
palmeri 71
paludinoides 80
paludosa 71
papyracea 71
patula 71, 81
patula catemacensis 59
pealiana 71
репезта 72
peristomata 72
pernambucensis 72
perovata 81
pertusa 72
petiti 48
phaeostoma 72
physis 72
physoides 72
picta 72

pinei 72
planorboides 79
planorbula 55

461

poeyana 72
pomatia 72
pomum 72
(Pomus) crassa var. oblonga 48
(Pomus) gigas var. minor 69
(Pomus) martensiana 68
ponderosa 81
porphyrostoma 73
proboscidea 81
producta 73
prunella 81
prunulum 55
pulchella 48
pulchella gallardoi 47
puncticulata 73
puntaplaya 73
purpurascens 73
pygmaea 81
pyrum 73
quercina 73
quinindensis 55
quitensis 74
reflexa 74

reflexa var. melanostoma 69
retusa 74

reyrei 74
rhodostoma 55
robusta 74

roissii 48

rosea 81

rotula 51
rotundata 81
rufilineata 80
rugosa 74
scalariformis 81
scalaris 74
scholvieni 74
schrammi 78
semitecta 74
semperi 75
sigaretina 81
simplex 75

sloanii 48

solida 49

sordida 75
sowerbyi 49
spirata 75, 81
spixii 49

sprucel 75
storeria 49
superba 75
swainsoni 75
swainsonii 75
tamsiana 56
tasmaniae 81

462

tenuissima 76
teres 56
testudinea 76
tristis 56, 80
tristrami 76
trochulus 80
venetus 76
vermiformis 76
vexillum 76
violacea 77
welwitschiana 77
woodwardi 77
yatesii 77
yucatanensis 77
yucatanensis var. yzabalensis 77
zonata 49, 78

Ampullariidae 40, 43, 46-47, 62, 64, 78,

80-81, 179, 453
Ampullariinae 42
Ampullarius 42, 51
Ampullospiridae 58

Analcadia 196, 198, 398, 412, 423-427

anatinum, Drepanotrema 453
Ancylidae 37, 453
Angulata 196, 198, 400, 412-414,
423-424, 426-427
brasiliensis 421, 436
angulata, Ampullaria 56
Helicina 400, 427
Pomacea 56
Pomus 57
annularis, Pomacea 81
anozona, Helicina 388, 389
Helicina oweniana 296, 332, 389
Antillorbis 453
aperta, Ampullaria 80
Aphanoconia 422
pachystoma ponsonbyi 421
apparine, Galium 127
appressa, Lymnaea stagnalis 451
Araceae 301
arata, Pomacea 57
archimedes, Ampullaria 57
Pomacea 57
Arion fasciatus 451
armeniacum, Ampullaria 57
Pomacea 57
Asolene 41-42, 46-50, 60, 78
brownii 46
crassa 46-47
cyclostoma 47
elongata 49
exumbilicata 47
fasciolata 47
gallardoi 47

INDEX

granulosa 47

iheringi 49

impervia 47

monticola 47

naticoides 48

neritiniformis 49

nubila 48

oblonga 48

olivier! 48

ormophora 48

petiti 48

platae 48

pulchella 48

roissii 48

roissyi 48

sloanii 48

solida 49

sowerbyi 49

spixii 49

storeria 49

(Surinamia) fairchildi 78

zonata 49
aspersa, Helix 125
aspersum, Cornu 125-128, 130-131
Asteraceae 126
atlanticus, Oxychilus 121-123

Oxychilus (Drouetia) 121, 122
aulanieri, Ampullaria 57, 75

Pomacea 57
aureola, Helix 407

Lucidella 390, 407, 407-408, 412,

414-415, 421, 430, 436

auricularia, Radix 141-147, 143-144

Lymnaea 142

Lymnaea (Radix) 141
auriformis, Ampullaria 57

Pomacea 57

aurisformis var. ocanensis, Ampullaria 70

aurostoma, Ampullaria 57
Pomacea 57

australis, Ampullaria 57
Corbicula 2, 7, 29-32, 35
Pomacea 57

autumnalis, Ampullaria 57
Pomacea 57

avellana, Ampullaria 57, 81
Pomacea 57

Azolla filiculoides 454

baeri, Ampullaria 52
Pomacea 52
balteata, Ampullaria 52
Pomacea 52
balthica, Radix 141-147, 143, 145
batabana, Ampullaria 58

Ротасеа 58
beatrix, Alcadia 297
Alcadia (Leialcadia) 283
Helicina 275, 283-284, 283, 288-291,
294, 296-297, 307-308, 311-312, 314,
316, 318, 325-326, 334, 338, 342-343,
348, 357, 386, 410, 412-414, 418—420,
423—424, 426—427
Helicina beatrix 285-288, 287-292,
294-296, 300-301, 303, 305, 307, 311,
314, 316, 409, 415, 432, 434
Helicina (Gemma) beatrix 283
Helicina (Oligyra) 283
Oligyra 423
Oligyra (Succincta) beatrix 283
var. sensu, Helicina 284, 291, 294
belizensis, Ampullaria 58
Pomacea 58
Bellula 423
beryllina, Pleuropoma 421
Beta 128
maritima 126-130
bibliana, Pomacea 81
bilineata, Ampullaria 78
biloba, Salvinia 454
Biomphalaria 163, 453
glabrata 101—104, 107, 149-152, 157,
160, 161, 162-164
peregrina 444, 457
pfeifferi 150, 163
tenagophila 444
boeckeleri, Alcadia 373-374, 374-376,
376-377, 383, 409—410, 413, 418, 426,
432, 435
Alcadia (Microalcadia) 195, 197, 215,
374, 429
Helicina 196, 198, 374
borealis, Ampullaria 81
boucourti, Helicina 348, 425
Brachytheciaceae 126
Brachythecium 128
rutabulum 126-130
brasiliensis, Ampullaria 58
Angulata 421, 436
Helicina 390, 400, 400-401, 427
Pomacea 58
brevilabris, Helicina 386
bridgesii, Ampullaria 58
Pomacea 42, 58
Bromus hordeaceus 127-128
Brotia 36
Broussaisia 175
brownii, Ampullaria 46
Asolene 46
bruguieri, Ampullaria 80

INDEX

buccinoidea, Ampullaria 81

bugensis, Dreissena 118

bulimoides, Ampullaria 81

Bulimus sinamarina 78
tristis 80
urceus 51

Bulinus 163

bulla, Ampullaria 58
Pomacea 58

buxea, Ampullaria 58
Pomacea 58

cacaguelita, Helicina (Succincta) 425

caliginosa, Ampullaria 58
Pomacea 58

Camaenidae 413

camena, Pomacea 58

463

campbellorum, Semelina 169, 171-172,

172-173

canaliculata, Ampullaria 42, 58, 81

Pomacea 42-43, 58, 444
canalifera, Ampullaria 81
Carduus 128

tenuifloris 126-127, 129

caroliniana, Polymesoda 6-7, 30-31

carota, Daucus 127

cassidiformis, Ampullaria 59
Pomacea 59

castanea, Ampullaria 52
Pomacea 52

castelloi, Ampullaria 59
Pomacea 59

castelnaudii, Ampullaria 59
Pomacea 59

catamarcensis, Ampullaria 59

Pomacea 59

catemacensis, Ampullaria patula 59

Pomacea 59
Catinella rotundata 175
celebensis, Corbicula 6

Ceochasma 196, 198, 390, 400, 421,

426-427
phrixina 400
Cepaea 133, 413
hortensis 133, 137

nemoralis 121, 125, 133, 137

vindobonensis 133-137, 134, 136, 139

cerasum, Ampullaria 59
Pomacea 59

Ceratodes 49-51
fasciatus 50-51

Ceres 414

Ceresidae 413-414, 421

chamana, Ampullaria lattrei 59

Pomacea 59

464

chaquensis, Ротасеа 59
Pomacea canaliculata 59
chemnitzii, Ampullaria 59
Pomacea 59
chiapasensis, Pomacea 59
chiapensis, Helicina 254-257, 258, 271
chiappensis, Helicina 255
Chilina parchappii 444
Chilinidae 453
chiquitensis, Ampullaria 50
chiquitica, Helicina 195, 197, 302, 347-348,
350, 356-358, 358-362, 360-364, 389,
409-410, 413-415, 419-420, 426-427,
432, 435
Helicina (Gemma) 215, 357
Marisa 50
Oligyra 357
chlorostoma, Ampullaria 52
Pomacea 52
chryseis, Eutrochatella microdina 364
Eutrochatella (Pyrgodomus) microdina 364
Helicina 364, 407
Helicina (Pyrgodomus) 364
Pyrgodomus 369-370
Pyrgodomus microdinus 364, 370
Cibotium 175
cimex, Drepanotrema 453-457, 454
cincta, Ampullaria 60
Pomacea 60
Cinctella 196, 198, 390, 397, 423, 426-429
cinctella, Helicina 397
Helicina (Oxyrhombus) 425
cingulata, Ampullaria 52
Pomacea 52
citreum, Ampullaria 60
Pomacea 60
clappi, Varicella 185
coccinostoma, Helicina 388, 388
Helicina oweniana 296, 332, 388
columbensis, Ampullaria 60
Pomacea 60
columbiensis, Ampullaria 60
Pomacea 60
columellaris, Ampullaria 51, 60
Pomacea 60
commissionis, Pomacea 60
Concentrica 423, 427
concentrica, Helicina 425, 427
Conchylium 51
conexa, Heleobia 446, 449
confusa, Alcadia (Leialcadia) beatrix 297
Helicina beatrix 205, 214, 284, 289-294,
297, 297-301, 300-303, 307, 356, 409,
413, 415, 432-433
Helicina (Gemma) beatrix 214, 297

Oligyra (Succincta) beatrix 297
conica, Ampullaria 52, 81
Pomacea 52
conoidea, Ampullaria 60
Pomacea 60
consolatrix, Ampullaria 60
Pomacea 60
contamanoensis, Ampullaria 60
Pomacea 60
contrarius, Planorbis 50
convexa, Helicina 422
Corbicula 1—4, 5, 6-7, 11-13, 17-19,
20-23, 24, 25, 29-36, 38
africana 3
australis 2, 7, 29-32, 35
celebensis 6
ducalis 8
fluminalis 2-3, 6-7, 29-32
fluminea 1-4, 6-7, 9-10, 11, 13, 16-17,
21, 29-36, 38
gustaviana 4
japonica 2-3, 7, 30-31, 33, 35
javanica 1, 4, 6-8, 9-10, 11-12, 16, 21,
29-33, 35-36, 38
lacustre 13, 14, 16
lacustris 6, 11, 13, 16
[еапа 2-3, 7, 30-32, 35
lindoensis 17
linduensis 1, 4, 11-12, 17, 18-21, 21,
33-36, 38
loehensis 1, 4, 7, 11-12, 17, 18, 24-26,
25-27, 29-36, 38
madagascariensis 3, 7, 29-32
mahalonensis 21-22
masapensis 26
matannensis 1, 4, 6-7, 11-12, 17-19, 17,
21-23, 22-23, 25, 27-36, 38
moltkeana 13
moltkiana 1, 4, 5, 7, 11-13, 14-18,
16-17, 21, 26-27, 29-36, 38
possoensis 1, 4, 6-7, 11-12, 17, 24, 26,
26-36, 28, 38
pullata 4
rivalis 4
sandai 2-3, 7, 17, 30-35, 37
subplanata 4, 6, 21-22, 26-27, 35-36
sumatrana 4, 6, 11, 13, 14, 16
tobae 4, 35
towutensis 21-22
tumida 4
verbecki 13
Corbiculidae 1-2, 4, 33
cornuarietis, Helix 49-50
Marisa 50
Cornu aspersum 125-128, 130-131

INDEX 465

cornucopia, Ampullaria 60 Deroceras leave 451
Pomacea 60 reticulatum 125
costaricana, Ampullaria 60 diaphana, Helicina 335, 336, 348
Pomacea 60 diffusa, Pomacea 62
Succinea 244 Pomacea bridgesii 62
costaricensis,Helicina funcki 195, 197, dilatata, Pomacea 62
215-217,217, 226, 234, 236, 244-245, dioica, Уса 126-129, 135
419 disseminata, Ampullaria 62
Helicina (Retorquata) funcki 215 Pomacea 62
Helicina (Tristramia) funcki 215 dolioides, Ampullaria 62
costata, Eutrochatella 420 Pomacea 62
cousini, Ampullaria 61 dolium, Ampullaria 62
Pomacea 61 Pomacea 62
crassa, Ampullaria 46-47, 81 dorbignyana, Ampullaria 62
Asolene 46-47 Pomacea 62
var. monticola, Ampullaria 47 dorbignyi, Ampullaria 62
var. oblonga, Ampullaria (Pomus) 48 Pomacea 62
crassatina, Ampullaria 81 Dreissena bugensis 118
crocostoma, Ampullaria 52 bugensis profunda 118
Pomacea 52 polymorpha 109-110, 111-112, 117-119
crosseana, Ampullaria 61 Drepanotrema 453, 457
Pomacea 61 anatinum 453
cubensis, Ampullaria 61 cimex 453-457, 454
Pomacea 61 depressissimum 453
cuericiensis, Helicina punctisulcata 195, heloicum 453
197, 273-274, 278-282, 281-283, 397, kermatoides 453-455, 454, 457
409-410, 413-414, 426-428, 431 lucidum 453
Helicina (Tristramia) punctisulcata dubia, Ampullaria 53
214,277 Pomacea 53
cumingi, Pomacea urabaensis 76 ducalis, Corbicula 8
cumingii, Ampullaria 61 dysoni, Ampullaria 62
Pomacea 61 Helicina 398-399, 398, 425, 436
Pomacea sanjosensis 74 Helicina (Analcadia) 427
cuprina, Ampullaria 42, 53 Pomacea 62
Pomacea 42, 53
Cyanea 175 echandiensis, Helicina 195, 197,
Cyanocyclas 6 272-276, 273, 275-277, 281, 409-410,
cyclostoma, Ampullaria 47 413, 420, 426-428, 431
Asolene 47 Helicina (Tristramia) 214, 271
Cyrena orientalis var. javanica 8 echioides, Picris 127-130
cythereoidea, Semelina 170 edulis, Mytilus 119
Effusa 42, 51-54, 61, 73
dacostae, Ampullaria 61 effusa var. conica, Ampullaria 52
Pomacea 61 Nerita 53
Dactylis glomerata 127-128, 130 Pomacea 53
Daucus carota 127 elata, Helicina 335, 336, 347
decussata, Ampullaria 61 Helicina fragilis 347, 419
var. commissionis, Ampullaria 60 electrina, Ampullaria 63
Pomacea 61 Pomacea 63
delattrei, Ampullaria 61 elegans, Ampullaria 63
Pomacea 61 Pomacea 63
deppeana, Helicina 245 elongata, Ampullaria 81
depressa, Ampullaria 62, 81 Ampullaria (Felipponea) 49
Pomacea 62 Asolene 49

depressissimum, Drepanotrema 453 Pomacea 63

466

Elytrigia 128
repens 126-130
Emoda 414, 423
pulcherrima titanica 421
sagraiana 421
enigmaticus, Ficopomatus 445
equestris, Pila 79
ernesti, Helicina (Angulata) rhynchostoma
420
erogata, Ampullaria 63
Pomacea 63
erronea, Ampullaria 63
Pomacea 63
erythrostoma, Ampullaria 63
Pomacea 63
escondida, Helicina 195, 197, 274-275,
277, 301-302, 347-348, 349-353,

350-357, 355, 358, 363-364, 409-410,

412-413, 419-420, 426-428, 432,
434-435
Helicina (Gemma) 215, 348
Eucaladia 423
Euglandina rosea 451
eumicra, Ampullaria 63
Pomacea 63
Euneritella 423
eupatoria, Agrimonia 127

Eutrochatella 195-199, 203, 368, 390, 401,

405, 408, 414-415, 419-420, 429
(Artecallossa) microdina 364
costata 420
microdina chryseis 364
nicrodina var. chryseis 364
nobilis 420
pulchella 390, 405, 406, 415, 419-421,
436
(Pyrgodomus) microdina chryseis 364
(Pyrgodomus) microdina microdina 364
tankervillii 420
excavata, Ampullaria 81
exculpta, Pomacea 63
exigua, Ampullaria 80
Helicina 374
eximia, Ampullaria 63
Pomacea 63

var. dilatata, Ampullaria 62
var. elongata, Ampullaria 63
fasciatus, Arion 451
Ceratodes 50
Marisa 50
fasciolata, Asolene 47
Helix 47
faujasii, Ampullaria 81
Felipponea 41-42, 49-50
ferruginea, Ampullaria 64
Pomacea 64
Festiva 423
Festuca rubra 127-128, 130
Ficopomatus enigmaticus 445
figulina, Ampullaria 64
Pomacea 64
filiculoides, Azolla 454
flagellata, Ampullaria 64
Pomacea 64
var. guatemalensis, Ampullaria 65
flatilis, Ampullaria 64
Pomacea 64
flava, Pomacea 64
Pomacea paludosa 64
flavida, Helicina 283, 296, 363, 386-387,
386-387, 409, 430
Helicina (Oligyra) 215, 386
Helicina (Succincta) 428
var. beatrix, Helicina 283
fluminalis, Corbicula 2-3, 6-7, 29-32
fluminea, Corbicula 1-4, 6-7, 9-10, 11, 13,
16-17, 21, 29-36, 38
fontinalis, Lymnaea 147
Lymnaea (Peregriana) 147
fragilis, Alcadia (Leialcadia) 357, 389
Ampullaria 81
Helicina 195, 197, 335, 338-339, 339,
342, 347, 357, 363, 389, 409, 423
Freycinetia 175
fucki, Helicina 215
fulgens, Ovachlamys 176, 244, 363
fumata, Ampullaria 64
Pomacea 64
funcki, Helicina 205, 210-211, 211,
214-216, 216, 224-228, 225-226,

Exotica 169

expansa, Ampullaria 53
Pomacea 53

exumbilicata, Asolene 47
Helicina 47

228-229, 234-236, 238, 239-247, 244,
249-250, 252, 254, 262, 264, 268, 270,
274, 281, 286, 301-302, 306, 315-316,
322, 324-325, 328, 343, 350, 356, 396,
409-411, 413, 415, 418-420, 422-424,
426-427, 431, 433

Helicina (Retorquata) 215

Helicina (Tristramia) 214-216, 425

Helicina (Tristramia) funcki 215

funckii, Helicina 215

fairchildi, Asolene (Surinamia) 78
Pomella 78

falconensis, Pomacea 63

fasciata, Ampullaria 51, 53, 63-64, 75, 79

funki, Нейста 215-216

Galium apparine 127
molugo 127
gallardoi, Ampullaria pulchella 47
Asolene 47
galloprovincialis, Ampullaria 81
garciae, Pomacea 64
Pomacea paludosa 64
Gemma 196, 198, 397, 412, 423, 425-429
gemma, Alcadia (Leialcadia) 318
Helicina 205, 209, 210, 214, 275, 277,
296, 302, 307-308, 318-319, 319,
321—326, 322, 324-332, 331, 334,
338-343, 348, 357-358, 363, 367,
387-389, 397, 409-410, 413-415,
419-420, 423-424, 426-427, 432, 434
Helicina (Gemma) 215, 318
Oligyra 423
Oligyra (Succincta) 318
georgii, Ampullaria 64
Pomacea 64
Geranium molle 127
robertianum 127
geveana, Ampullaria 53
Pomacea 53
var. suprafasciata, Ampullaria 55
gevesensis, Ampullaria 53
Pomacea 53
ghiesbreghti, Ampullaria 64
Pomacea 64
gibbosa, Ampullaria 79
gigantea, Ampullaria 65
Pomacea 65
giganteus, Pomacea 65
Pomus 65
gigas, Ampullaria 65
Pomacea 42—43, 65
var. minor, (Pomus) Ampullaria 69
var. unicolor, Ampullaria 76
glabrata, Biomphalaria 101-104, 107,
149-152, 157, 160, 161, 162-164
glauca, Helix 51, 54
Pomacea 54—55
glomerata, Dactylis 127-128, 130
gossei, Ampullaria 65
Pomacea 65
granulosa, Ampullaria 47
Asolene 47
guadelupensis, Ampullaria 44, 54
Pomacea 54
guaduasensis, Ampullaria 65
Pomacea 65
gualtieri, Ampullaria 65
Pomacea 65

467

guatemalensis, Pomacea 65

guinaica, Ampullaria 71

gustaviana, Corbicula 4

guyanensis, Ampullaria 65
Pomacea 65

haemastoma, Ampullaria 65
Pomacea 65
hanleyana, Ampullaria 66
Pomacea 66
hanleyi, Ampullaria 66
Pomacea 66
haustrum, Ampullaria 66
Pomacea 43, 66
Heleobia 444
conexa 446, 449
рагсварри 443-449
Helicidae 133, 412—413
Helicina 195-200, 203, 211, 213-214, 217,
301, 335, 370, 383-384, 388, 390-391,
401-403, 406-408, 412-430
alata 408
amoena 195, 197, 246, 254, 387, 396,
396-397, 409, 421, 436
(Analcadia) dysoni 427
angulata 400, 427
(Angulata) rhynchostoma ernesti 420
anozona 388, 389
beatrix 275, 283-284, 283, 288-291,
294, 296-297, 307-308, 311-312, 314,
316, 318, 325-326, 334, 338, 342-343,
348, 357, 386, 410, 412-414, 418-420,
423-424, 426-427
beatrix beatrix 285-288, 287-292,
294-295-296, 300-301, 303, 305, 307,
311, 314, 316, 409, 415, 432, 434
beatrix confusa 205, 214, 284, 289-294,
297, 297-301, 300-303, 307, 356, 409,
413, 415, 432-433
beatrix nicaraguae 296, 296, 307
beatrix riopejensis 195, 197, 211, 214,
277, 290-291, 293, 302-307, 305-308,
334, 409, 414, 432-433
beatrix var. sensu 284, 291, 294
boeckeleri 196, 198, 374
boucourti 348, 425
brasiliensis 390, 400, 400-401, 427
brevilabris 386
chiapensis 254-257, 258, 271
chiappensis 255
chiquitica 195, 197, 302, 347-348, 350,
356-358, 358-362, 360-364, 389,
409-410, 413-415, 419-420, 426-427,
432, 435
chryseis 364, 407

468

cinctella 397

coccinostoma 388, 388

concentrica 425, 427

convexa 422

deppeana 245

deppeana parvidens 215-217, 217, 245

diaphana 335, 336, 348

dysoni 398-399, 398, 425, 436

echandiensis 195, 197, 272-276, 273,
275-277, 281, 409-410, 413, 420,
426-428, 431

elata 335, 336, 347

escondida 195, 197, 274-275, 277,
301-302, 347-348, 349-353, 350,
354-357, 355, 358, 363-364, 409-410,
412-413, 419-420, 426-428, 432,
434-435

exigua 374

exumbilicata 47

flavida 283, 296, 363, 386-387, 386-387,
409, 430

flavida var. beatrix 283

fragilis 195, 197, 335, 338-339, 339, 342,
347, 357, 363, 389, 409, 423

fragilis elata 347, 419

fragilis merdigera 363

fucki 215

funcki 205, 210-211, 211, 214-216, 276,
224-228, 225-226, 228-229, 234-236,
238, 239-247, 244, 249-250, 252, 254,
262, 264, 268, 270,274, 281, 286,
301-302, 306, 315-316, 322, 324-325,
328, 343, 350, 356, 396, 409-411, 413,
415, 418—420, 422-424, 426-427, 431,
433

funcki costaricensis 195, 197, 215-217,
217, 226, 234, 236, 244-245, 419

funckii 215

funki 215-216

gemma 205, 209, 210, 214, 275, 277,
296, 302, 307-308, 318-319, 319,
321-326, 322, 324-332, 331, 334,
338-343, 348, 357-358, 363, 367,
387-389, 397, 409-410, 413-415,
419-420, 423-424, 426-427, 432, 434

(Gemma) beatrix beatrix 283

(Gemma) beatrix confusa 214, 297

(Gemma) beatrix riopejensis 214, 303

(Gemma) chiquitica 215, 357

(Gemma) gemma 215, 318

(Gemma) monteverdensis 215, 334

(Gemma) escondida 215, 348

(Gemma) talamancensis 215, 308

(Helicina) tenuis 256

heloisae 284

INDEX

hojarasca 196, 198, 370, 405

(Idesa) microdina 364

jamaicensis 196, 198, 390, 404, 404, 412

kubaryi 421

lamellosa 379, 408

lindeni 254, 258, 270-271, 271

lindeni var. minor 255

liobasis 413

lirata 377

lirata var. rusticella 377-378

lirata var. semistriata 377

lirata var. unidentata 377-378

lyrata 377

major 400

merdigera 282, 334, 335, 340, 347

microdina 364-365, 364

mohriana 334, 347, 363

monteverdensis 195, 197, 282, 334,
337-341, 338-339, 341-348, 345, 357,
363, 387-389, 409-410, 413-415, 420,
426-427, 432, 434

neritella 390-391, 391-392, 412, 429, 436

nicaraguae 284, 284

(Oligyra) beatrix 283

(Oligyra) flavida 215, 386

(Oligyra) тает! 255

orbiculata 203, 213, 392-395, 393, 413,
421, 428, 430, 436

orbiculata tropica 395

oweniana 195, 197, 256, 271, 296, 308,
316, 318, 328, 334, 348, 387, 388, 389,
409

oweniana sensu 316

oweniana anozona 296, 332, 389

oweniana coccinostoma 296, 332, 388

oweniana var. anozona 318, 389

oweniana var. coccinostoma 318, 388

(Oxyrhombus) cinctella 425

palliata 403

(Perenna) lamellosa 377-378

(Perenna) lirata 378

(Perenna) semistriata 378

pitalensis 203, 244-246, 246, 248-249,
249-252, 253, 254, 315-316, 387, 396,
409-410, 413-414, 420, 422-424,
426-428, 431, 433

platychila 392, 392, 429, 436

(Poenia) lirata 378

(Poenia) lirata var. rusticella 378

pulchella 405

punctisulcata 277, 277, 282, 397,
423-424

punctisulcata cuericiensis 195, 197,
273-274, 278-282, 281-283, 397,
409-410, 413-414, 426-428, 431

punctisulcata punctisulcata 283

punctisulcata zunilensis 271, 278, 278,
282-283

(Pyrgodomus) chryseis 364

raresulcata 282

(Retorquata) funcki 215

(Retorquata) funcki costaricensis 215

rhips 413

rotunda 405

rupestris 365

rusticella 377, 379

salvini 395

semistriata 377, 379

sericea 399, 399, 436

strebeli 363

succincta 395

(Succincta) cacaguelita 425

(Succincta) flavida 428

talamancensis 195, 197, 252, 308,
310-312, 311-317, 318, 357, 387-388,
409-410, 413, 415, 420, 426-427, 432,
434

tamsiana 397

tennuis 256

tenuis 195, 197, 214, 254-256, 256, 258,
262, 262-264, 265-271, 271, 315-316,
353, 387-388, 397, 409-410, 410-411,
411-414, 420, 422-424, 426-428, 431

tenuis tenuis 256

tenuis pittieri 195, 197, 245, 255-256,
258, 259, 266-267, 271, 419

tenuis var. chiapensis 255

tenuis var. lindeni 255

(Tenuis) tenuis 255

terryae 308, 309, 316

(Trichohelicina) klappenbachi 370

(Tristramia) echandiensis 214, 271

(Tristramia) funcki 214-216, 425

(Tristramia) funcki costaricensis 215

(Tristramia) funcki funcki 215

(Tristramia) funcki parvidens 215

(Tristramia) lindeni 255

(Tristramia) pitalensis 214, 245

(Tristramia) punctisulcata cuericiensis
214, 277

(Tristramia) tenuis 214, 254-255

tuncki 215

turbinata 395-396, 395-396, 421, 436

umbonata 413, 425

unidentata 377, 379

vernalis 254-257, 257, 263, 266, 270

vernalis verapazensis 255, 271

zephyrina 396, 428

zephyrina zephyrina 419

469

Helicinidae 195-197, 199-200, 203,
205-206, 207, 208, 210-214, 213, 244,
270, 282, 316, 332, 347, 357, 384, 390,
409, 410-411, 411-415, 418-422, 426,
430

Heliconiaceae 218, 243, 277, 301, 306, 343

Helix aspersa 125
aureola 407
cornuarietis 49-51
fasciolata 47
glauca 51, 54
lineata 67
neritina 54
oculuscommunis 54
platae 46, 48

heloicum, Drepanotrema 453

heloisae, Helicina 284

Hendersonia 414, 426
occulta rubella 415, 419, 421-422

hepataria, Ampullaria 79

Hispida 423

hojarasca, Alcadia 214, 343, 371-373, 373,

375, 376-377, 383, 405, 409-410, 413,
418, 426, 432, 435

Alcadia (Microalcadia) 195, 197, 215,
370, 419, 429

Helicina 196, 198, 370, 405

hollandi, Alcadia 390, 402-403, 403-405,
422, 436

hollingsworthi, Pila (Pomacea) 66

hondurasensis, Ampullaria 66
Pomacea 66

hopetonensis, Ampullaria 66
Pomacea 66

hordeaceus, Bromus 127-128

hortensis, Cepaea 133, 137

hybrida, Ampullaria 81

Hydrobiidae 443, 453

Idesa 369, 405, 423-424, 426

ignota, Pila 79

iheringi, Asolene 49

immersa, Ampullaria 66
Pomacea 43, 66

imperforata, Ampullaria 79

impervia, Ampullaria 47
Asolene 47

Incrustata 423

innexa, Ampullaria 66
Pomacea 66

insignis, Luntia 185-186, 185

insularis, Pomacea 42

insularum, Ampullaria 66
Pomacea 42, 66

470

intermedia, Ampullaria 54
Marisa 51
Pomacea 54

interrupta, Ampullaria 66
Pomacea 66

intropicta, Ampullaria 67
Pomacea 67

Intusplicata 423

Iris pseudacorus 454

jamaicensis, Alcadia 412, 426, 436
Helicina 196, 198, 390, 404, 404, 412

jamesi, Semele 173

japonica, Corbicula 2-3, 7, 30-31, 33, 35
Waldemaria 421

javanica, Corbicula 1, 4, 6-8, 9-10, 11-12,
16, 21, 29-33, 35-36, 38

kermatoides, Drepanotrema 453-455, 454,
457
klappenbachi, Helicina (Trichohelicina) 370
knorrii, Ampullaria 51
Marisa 51
kubaryi, Helicina 421

labiata, Radix 141-147, 143-144
labiosa, Ampullaria 67
Pomacea 67
lacustre, Corbicula 13, 14, 16
lacustris, Corbicula 6, 11, 13, 16
laevigata, Ampullaria 81
laevigatum, Limnobium 454
lagotis, Lymnaea 141
Radix 143, 147
lamarckii, Ampullaria 67
Pomacea 67
lamellosa, Helicina 379, 408
Helicina (Perenna) 377-378
Lucidella lirata 378, 384
lanceolata, Plantago 127
Lanistes 42
latifolia, Silene 127
lattrei, Ampullaria 67
Pomacea 67
leana, Corbicula 2-3, 7, 30-32, 35
leave, Deroceras 451
Leialcadia 405, 412, 423, 425
Lemnaceae 454
lemniscata, Ampullaria 67
Pomacea 67
Leptinaria 185
leucostoma, Ampullaria 67
Pomacea 67
levior, Ampullaria 67
Pomacea 67

Liguus 413
Limacidae 121
Limnobium laevigatum 454
Limnopomus 47, 51, 59-60, 66, 68
limosa, Neocorbicula 6-7, 30-31, 34
lindeni, Helicina 254, 258, 270-271, 271
Helicina (Oligyra) 255
Helicina (Tristramia) 255
var. minor, Helicina 255
lindoensis, Corbicula 17
linduensis, Corbicula 1, 4, 11-12, 17,
18-21, 21, 33-36, 38
linearis, Pomacea 81
lineata, Helix 67
Pomacea 42-43, 67
linnaei, Ampullaria 67
Pomacea 67
liobasis, Helicina 413
lirata, Helicina 377-378
Helicina (Perenna) 378
Helicina (Poenia) 378
Lucidella 195, 197, 214, 270, 377-379,
381-383, 383-386, 385, 408—410,
412-414, 419-420, 422, 430, 432, 435
Lucidella (Регеппа) 215, 378
Lucidella (Роета) 378-379
var. lamellosa, Lucidella 378
var. rusticella, Helicina 377-378
var. rusticella, Helicina (Poenia) 378
var. semistriata, Helicina 377
var. unidentata, Helicina 377-378
lirulata, Semele nuculoidea 170
livescens, Ampullaria 67
Pomacea 67
loehensis, Corbicula 1, 4, 7, 11-12, 17, 18,
24-26, 25-27, 29-36, 38
Lucidella 196-200, 203, 213, 383, 390,
407, 412-415, 421-422, 426, 429-430
adamsiana 412
aureola 390, 407, 407-408, 412,
414-415, 421, 430, 436
lirata 195, 197, 214, 270, 377-379,
381-383, 383-386, 385, 408-410,
412-414, 419-420, 422, 429-430, 432,
435
lirata lamellosa 378, 384
lirata var. lamellosa 378
midyetti 429
(Perenna) lirata 215, 378
(Poenia) lirata 378-379
lucidum, Drepanotrema 453
Luntia 185
insignis 185-186, 185
lutea, Pomacea 67
luteostoma, Ampullaria 51, 54

=

=

Ротасеа 54
Lymnaea 141
ampla 141
auricularia 142
fontinalis 147
lagotis 141
ovata 141-142
peregra 141-142, 147
peregra f. ampla 141-142
peregra f. ovata 141-142
peregra f. typica 141-142
(Peregriana) fontinalis 147
(Radix) auricularia 141
(Radix) peregra 141
stagnalis 123
stagnalis appressa 451
typica 141
viatrix 444
lymnaeaeformis, Ampullaria 67
Pomacea 67
Lymnaeidae 37, 141
lyrata, Helicina 377

Macoma 169
Mactra veneriformis 119
maculata, Pomacea 41, 43, 51, 68
madagascariensis, Corbicula 3, 7, 29-32
mahalonensis, Corbicula 21-22
major, Alcadia 401, 401-402, 403,
412-413, 419, 425, 436
Helicina 400
malleata, Ampullaria 68
Pomacea 68
var. arata, Ampullaria 57
var. chiapasensis, Ampullaria 59
var. exculpta, Ampullaria 63
var. oajacensis, Ampullaria 70
var. prasina, Ampullaria 73
var. strebeli, Ampullaria 75
Mamilla 423
manco, Pomacea 68
manetou, Pila 68
Pomacea 68
mansoni, Schistosoma 101-102, 104, 150,
152
marginatra, Ampullaria 68
Pomacea 68
Marisa 41-42, 49-51, 77
chiquitica 50
cornuarietis 50
fasciatus 50
intermedia 51
knorrii 51
planogyra 51
rotula 51

INDEX

471

maritima, Beta 126-130
martensiana, Ampullaria (Pomus) 68
Pomacea 68
martinezi, Ampullaria 68
Pomacea 68
masapensis, Corbicula 26
matannensis, Corbicula 1, 4, 6-7, 11-12,
17-19, 17, 21-23, 22-23, 25, 27-36, 38
media, Ampullaria 81
Medicago minima 127
Megastoma 423
megastoma, Ampullaria 78
Pomella 78
melanocheila, Ampullaria 68
Pomacea 68
melanostoma, Pomacea 69
Melantho ponderosa 47
merdigera, Helicina 282, 334, 335, 340, 347
Helicina fragilis 363
meridaensis, Pomacea 69
Pomacea (Limnopomus) 69
mermodi, Ampullaria 69
Pomacea 69
meta, Ampullaria 69
Pomacea 69
metcalfei, Ampullaria 69
Pomacea 69
Metrosideros polymorpha 175
mexicana, Ampullaria 69
Pomacea 69
miamiensis, Ampullaria 69
Pomacea 69
Microalcadia 196, 198, 370, 405, 426
(Microalcadia) Alcadia 370, 412-413, 415,
427
microdina, Eutrochatella (Artecallossa) 364
Eutrochatella (Pyrgodomus) microdina
364
Helicina 364-365, 364
Helicina (Idesa) 364
Pyrgodomus 364
microdinus, Pyrgodomus 195, 197, 215,
364, 365-367, 367-370, 369, 407, 410,
413, 419, 422, 429, 432, 435
Pyrgodomus microdinus 364
midyetti, Lucidella 429
miltocheilus, Ampullaria 69
Pomacea 69
miltochilus, Ampullaria 69
Pomacea 69
minima, Medicago 127
Salvinia 454
minor, Pomacea 69
minuscula, Pomacea 54
Miratesta 37

472

misantlensis, Schasicheila 420
modesta, Ampullaria 70
Pomacea 70

mohriana, Alcadia (Leialcadia) fragilis 347

Helicina 334, 347, 363
molle, Geranium 127
moltkeana, Corbicula 13
moltkiana, Corbicula 1, 4, 5, 7, 11-13,
14-18, 16-17, 21, 26-27, 29-36, 38
molugo, Galium 127
monachus, Ampullaria 70
Pomacea 70
Monstera 243, 301
monstrosa, Ampullaria 70
Pomacea 70
Montacuta subquadrata 169-170, 171

monteverdensis, Helicina 195, 197, 282,
334, 337-341, 338-339, 341-348, 345,
357, 363, 387-389, 409-410, 413-415,

420, 426-427, 432, 434

Helicina (Gemma) 215, 334

monticola, Asolene 47

Musa 243

Musaceae 243

musaecola, Achatina 185-186, 185
Streptostele 185-186
Streptostele (Tomostele) 185-186,
185-186

Mysella 170
subquadrata 170

Mytilus edulis 119

nais, Pomacea 70
Naticidae 52, 58, 75
naticoides, Ampullaria 48
Asolene 48
nemoralis, Cepaea 121, 125, 133, 137
Neocorbicula 6
limosa 6-7, 30-31, 34
nuculoidea, Semele 169
Nerita effusa 53
urceus 51, 76

neritella, Helicina 390-391, 391-392, 412,

429, 436
Neritidae 413
neritina, Helix 54

Pomacea 54

neritiniformis, Ampullaria (Felipponea) 49

Asolene 49
neritoides, Ampullaria 78
Pomella 78

nicaraguae, Alcadia (Leialcadia) beatrix 284

Helicina 284, 284
Helicina beatrix 296, 296, 307

nicrodina var. chryseis, Eutrochatella 364

nigrilabris, Ampullaria 70
Pomacea 70

Nitida 423

nobilis, Ampullaria 70
Eutrochatella 420
Pomacea 70

notabilis, Ampullaria 70
Pomacea 70

novaegranadae, Ampullaria 70
Pomacea 70

nubila, Ampullaria 48
Asolene 48

nucleus, Ampullaria 79

nuculoides, Amphidesma 169
Semelina 169-170, 171, 172-173

oajacensis, Pomacea 70

Obione 130

oblonga, Ampullaria 70
Asolene 48
Pomacea 70

obtusa, Ampullaria 79

ocanensis, Ampullaria 70
Pomacea 70

occlusa, Ampullaria 71
Pomacea 71

ochracea, Ampullaria 71
Pomacea 71

octona, Subulina 244

oculuscommunis, Helix 54
Pomacea 54

Oleacinidae 185

oligista, Pomacea 55

Oligyra 196, 198, 393, 423-429
beatrix 423
chiquitica 357
gemma 423
(Succincta) beatrix beatrix 283
(Succincta) beatrix confusa 297
(Succincta) gemma 318
talamancensis 308

olivacea, Ampullaria 71
Pomacea 71

olivieri, Ampullaria 48
Asolene 48

Olygyra orbiculata 393

orbiculata, Helicina 203, 213, 392-395,

393, 413, 421, 428, 430, 436

Olygyra 393

orientalis var. javanica, Cyrena 8

orinoccensis, Ampullaria 55
Pomacea 55

ormophora, Ampullaria 48
Asolene 48

Orthalicidae 413

INDEX 473

Ovachlamys fulgens 176, 244, 363 pernambucensis, Ampullaria 72
ovata, Lymnaea 141-142 Pomacea 72
Radix 141-143 perovata, Ampullaria 81
oviformis, Ampullaria 71 pertusa, Ampullaria 72
Pomacea 71 Pomacea 72
oweniana, Helicina 195, 197, 256, 271, petiti, Ampullaria 48
296, 308, 316, 318, 328, 334, 348, 387, Asolene 48
388, 389, 409 pfeifferi, Biomphalaria 150, 163
sensu 316 phaeostoma, Ampullaria 72
var. anozona, Helicina 318, 389 Pomacea 72
var. coccinostoma, Helicina 318, 388 philippiana, Pomacea 55
Oxychilus atlanticus 121-123 Physidae 453
(Drouetia) atlanticus 121, 122 physis, Ampullaria 72
Oxyrhombus 196, 198, 396, 424-429 Pomacea 72
physoides, Ampullaria 72
Pachychilidae 37 Pomacea 72
pachystoma, Ampullaria 55, 79 Picris 128
Pomacea 55 echioides 127-130
pallens, Ampullaria 80 picta, Ampullaria 72
Palliata 404, 423, 426 Pomacea 72
palliata, Alcadia 213 Pila 41-43, 46, 49, 51-53, 70, 78-81, 179
Helicina 403 equestris 79
palmeri, Ampullaria 71 ignota 79
Pomacea 71 manetou 68
Paludina pulchra 73 periscelis 72
paludinoides, Ampullaria 80 (Pomacea) aldersoni 56
paludosa, Ampullaria 71 (Pomacea) hollingsworthi 66
Pomacea 42, 71, 179-181 sepulta 80
papyracea, Ampullaria 71 Pilidae 41, 46
Pomacea 71 pinei, Ampullaria 72
parchappii, Chilina 444 Pomacea 72
Heleobia 443-449 Pipturus 175
parvidens, Helicina deppeana 215-217, Pisidium 384
217,245 Pistia stratiotes 454
Helicina (Tristramia) funcki 215 pitalensis, Helicina 203, 244-246, 246,
pattersoni, Pomacea 55 248-249, 249-252, 253, 254, 315-316,
patula, Ampullaria 71, 81 387, 396, 409-410, 413-414, 420,
Pomacea 71 422-424, 426-428, 431, 433
pealiana, Ampullaria 71 Helicina (Tristramia) 214, 245
Pomacea 71 pittieri, Helicina tenuis 195, 197, 245,
penesma, Ampullaria 72 255-256, 258, 259, 266-267, 271, 419
Pomacea 72 planogyra, Marisa 51
peregra f. ampla, Lymnaea 141-142 Planorbidae 453, 457
f. ovata, Lymnaea 141-142 Planorbis 384
f. typica, Lymnaea 141-142 contrarius 50
Lymnaea 141-142, 147 planorboides, Ampullaria 79
Lymnaea (Radix) 141 planorbula, Ampullaria 95
Radix 141-143, 145, 147 Pomacea 55
Peregriana 141 Plantago lanceolata 127
peregrina, Biomphalaria 444, 457 platae, Asolene 48
Perenna 408, 429-430 Helix 46, 48
periscelis, Pila 72 platychila, Alcadia (Analcadia) 196, 198
Pomacea 72 Helicina 392, 392, 429, 436
peristomata, Ampullaria 72 Pleuropoma 412, 415, 419

Pomacea 72 beryllina 421

474 INDEX

Poaceae 126, 128, 301 columbiensis 60
Poenia 426, 430 columellaris 60
Poeniella 430 commissionis 60
poeyana, Ampullaria 72 conica 52
Pomacea 72 conoidea 60
Polymesoda caroliniana 6-7, 30-31 consolatrix 60

polymorpha, Dreissena 109-110, 111-112, contamanoensis 60

117-119
Metrosideros 175

Pomacea 41-43, 47, 49-51, 56, 60, 63,

72278. 11, 19)81
acuta 56
aldersoni 56
amazonica 43, 56
angulata 56
annularis 81
arata 57
archimedes 57
armeniacum 57
aulanieri 57
auriformis 57
aurostoma 57
australis 57
autumnalis 57
avellana 57
baeri 52
balteata 52
batabana 58
belizensis 58
bibliana 81
brasiliensis 58
bridgesii 42, 58
bridgesii diffusa 62
bulla 58
buxea 58
caliginosa 58
camena 58
canaliculata 42-43, 58, 444
canaliculata chaquensis 59
cassidiformis 59
castanea 52
castelloi 59
castelnaudii 59
catamarcensis 59
catemacensis 59
cerasum 59
chamana 59
chaquensis 59
chemnitzii 59
chiapasensis 59
chlorostoma 52
cincta 60
cingulata 52
citreum 60
columbensis 60

cornucopia 60
costaricana 60
cousini 61
crocostoma 52
crosseana 61
cubensis 61
cumingii 61
cuprina 42, 53
dacostae 61
decussata 61
delattrei 61
depressa 62
diffusa 62
dilatata 62
disseminata 62
dolioides 62
dolium 62
dorbignyana 62
dorbignyi 62
dubia 53
dysoni 62
effusa 53
electrina 63
elegans 63
elongata 63
erogata 63
erronea 63
erythrostoma 63
eumicra 63
exculpta 63
eximia 63
expansa 53
falconensis 63
fasciata 63-64
ferruginea 64
figulina 64
flagellata 43, 64
flatilis 64

flava 64

fumata 64
garciae 64
georgil 64
geveana 53
gevesensis 53
ghiesbreghti 64
gigantea 65
giganteus 65
gigas 42-43, 65

INDEX

glauca 54-55
gossei 65
guadelupensis 54
guaduasensis 65
gualtieri 65
guatemalensis 65
guyanensis 65
haemastoma 65
hanleyana 66
hanleyi 66
haustrum 43, 66
hollingsworthi 66
hondurasensis 66
hopetonensis 66
immersa 43, 66
innexa 66
insularis 42
insularum 42, 66
intermedia 54
interrupta 66
intropicta 67
labiosa 67
lamarckii 67
lattrei 67
lemniscata 67
leucostoma 67
levior 67
meridaensis 69
linearis 81

lineata 42-43, 67
linnaei 67
livescens 67
lutea 67
luteostoma 54
lymnaeaeformis 67
maculata 41, 43, 51, 68
malleata 68
manco 68
manetou 68
marginatra 68
martensiana 68
martinezi 68
melanocheila 68
melanostoma 69
meridaensis 69
mermodi 69
meta 69
metcalfei 69
mexicana 69
miamiensis 69
miltocheilus 69
miltochilus 69
minor 69
minuscula 54
modesta 70

monachus 70
monstrosa 70
nais 70

neritina 54
nigrilabris 70
nobilis 70
notabilis 70
novaegranadae 70
oajacensis 70
oblonga 70
ocanensis 70
occlusa 71
ochracea 71
oculuscommunis 54
oligista 55
olivacea 71
orinoccensis 55
oviformis 71
pachystoma 55
paludosa 42, 71, 179-181
paludosa flava 64
paludosa garciae 64
pattersoni 55
penesma 72
periscelis 72
peristomata 72
pernambucensis 72
pertusa 72
phaeostoma 72
philippiana 55
physis 72
physoides 72
picta 72

pinei 72
planorbula 55
poeyana 72
pomatia 72
pomum 72
porphyrostoma 73
prasina 73
producta 73
prourceus 73
prunulum 55
pulchra 73
puncticulata 73
puntaplaya 73
purpurascens 73
ругит 73
quercina 73
quinindensis 55
quitensis 74
reflexa 74

retusa 74

геуге! 74
rhodostoma 55

475

476

robusta 74
rugosa 74
sanjosensis 74

sanjosensis cumingii 74

scalaris 74
scholvieni 74
semitecta 74
semperi 75
simplex 75
sordida 75
spirata 75
sprucei 75
strebeli 75
superba 75

suprafasciata 55

swainsoni 75
swainsonii 75
tamsiana 56
tenuissima 76
teres 56
testudinea 76
tristis 56
tristrami 76
unicolor 76
urabaensis 76
urceus 41, 76
variegata 81
venetus 76
vermiformis 76
vexillum 76
vickeryi 76
villata 56
violacea 77

welwitschiana 77

woodward 77
yatesii 77
yucatanensis 77
yzabalensis 77
zeteki 77
zischkai 77
zonata 78

pomatia, Ampullaria 72

Pomacea 72

Pomella 41-42, 50, 78

americanista 78
fairchildi 78
megastoma 78
neritoides 78
schrammi 78
sinamarina 78

pomum, Ampullaria 72

Pomacea 72

Pomus 51

angulata 57
giganteus 65

INDEX

ponderosa, Ampullaria 81
Melantho 47
ponsonbyi, Aphanoconia pachystoma 421
porphyrostoma, Ampullaria 73
Pomacea 73
possoensis, Corbicula 1, 4, 6-7, 11-12, 17,
24, 26, 26-36, 28, 38
Possostrea 37
phrixina, Ceochasma 400
prasina, Pomacea 73
proboscidea, Ampullaria 81
producta, Ampullaria 73
Pomacea 73
profunda, Dreissena bugensis 118
Proserpina 414
Proserpinidae 413-414, 419, 421, 425
Protancylus 37
prourceus, Pomacea 73
prunella, Ampullaria 81
prunulum, Ampullaria 55
Pomacea 55
pseudacorus, Iris 454
Pseudoceratodes 42
Pseudoligyra 196, 198, 397, 423-424,
426-429
pulchella, Ampullaria 48
Asolene 48
Eutrochatella 390, 405, 406, 415,
419-421, 436
Helicina 405
pulchra, Paludina 73
Pomacea 73
pullata, Corbicula 4
puncticulata, Ampullaria 73
Pomacea 73 -
Punctisulcata 196, 198, 397, 423, 427-428
punctisulcata, Helicina 277, 277, 282, 397,
423-424
Helicina punctisulcata 283
puntaplaya, Ampullaria 73
Pomacea 73
purpurascens, Ampullaria 73
Pomacea 73
putris, Succinea 175
pygmaea, Ampullaria 81
Pyrgodomus 196-200, 210, 364, 369, 390,
407, 409, 412, 415, 420, 429
chryseis 369-370
microdina 364
microdinus 195, 197, 215, 364, 365-367,
367-370, 369, 407, 410, 413, 419, 422,
429, 432, 435
microdinus abditus 368-370, 368,
419-420
microdinus chryseis 364, 370

INDEX 477

microdinus microdinus 364 Catinella 175
simpsoni 369-370, 429 rubella, Hendersonia occulta 415, 419,
pyrum, Ampullaria 73 421-422
Pomacea 73 rubra, Festuca 127-128, 130
rufilineata, Ampullaria 80
quercina, Ampullaria 73 rugosa, Ampullaria 74
Pomacea 73 Pomacea 74
quinindensis, Ampullaria 55 rupestris, Helicina 365
Pomacea 55 rusticella, Helicina 377, 379
quitensis, Ampullaria 74 rutabulum, Brachythecium 126-130

Pomacea 74
sagraiana, Emoda 421

Radix 141, 143, 147 Salicornia 130

ampla 141-147, 143, 145 Salsolaceae 126

auricularia 141-147, 143-144 salvini, Helicina 395

balthica 141-147, 143, 145 Salvinia biloba 454

labiata 141-147, 143-144 minima 454

lagotis 143, 147 sandai, Corbicula 2-3, 7, 17, 30-35, 37

ovata 141-143 sanjosensis, Pomacea 74

peregra 141-143, 145, 147 sativa, Vica 127
raresulcata, Helicina 282 Saulea 42
reflexa, Ampullaria 74 scalariformis, Ampullaria 81

Pomacea 74 scalaris, Ampullaria 74

var. melanostoma, Ampullaria 69 Pomacea 74
regina, Viana 415, 419 Schasicheila 196-199, 370, 374, 390, 408,
repens, Elytrigia 126-130 412, 414, 420-421, 430
reticulatum, Deroceras 125 alata 390, 408, 408-409, 420, 436
retusa, Ampullaria 74 misantlensis 420

Pomacea 74 Schistosoma mansoni 101-102, 104, 150,
геугег, Ampullaria 74 152

Ротасеа 74 scholvieni, Ampullaria 74
rhips, Helicina 413 Pomacea 74
rhodostoma, Ampullaria 55 schrammi, Ampullaria 78

Pomacea 55 Pomella 78
riopejensis, Helicina beatrix 195, 197, Semelangulus 169

211, 214, 277, 290-291, 293, 302-307, Semele 173
305-308, 334, 409, 414, 432-433 jamesi 173

Helicina (Gemma) beatrix 214, 303 nuculoidea 169
Rissooidea 443 nuculoidea lirulata 170
rivalis, Corbicula 4 sirulata 170
robertianum, Geranium 127 virginiana 169
robusta, Ampullaria 74 Semelidae 169

Pomacea 74 Semelina 169
Rochefortia subquadrata 170 campbellorum 169, 171-172, 172-173
roissil, Ampullaria 48 cythereoidea 170

Asolene 48 nuculoides 169-170, 171, 172-173
roissyi, Asolene 48 subquadrata 169-170, 171, 172-173
rosea, Ampullaria 81 semistriata, Helicina 377, 379

Euglandina 451 Helicina (Perenna) 378
rotula, Ampullaria 51 semitecta, Ampullaria 74

Marisa 51 Pomacea 74
rotunda, Alcadia 405, 405, 436 semperi, Ampullaria 75

Helicina 405 Pomacea 75
rotundata, Ampluria 81 var. figulina, Ampullaria 75

Ampullaria 81 sepulta, Ampullaria 80

478

Зепсеа 196, 198, 399, 412, 423—424,
426—427
sericea, Helicina 399, 399, 436
sigaretina, Ampullaria 81
Silene latifolia 127
simplex, Ampullaria 75
Pomacea 75
simpsoni, Pyrgodomus 369-370, 429
sinamarina, Bulimus 78
Pomella 78
sirulata, Semele 170
sloanii, Ampullaria 48
Asolene 48
solida, Ampullaria 49
Asolene 49
sordida, Ampullaria 75
Pomacea 75
sowerbyi, Ampullaria 49
Asolene 49
spirata, Ampullaria 75, 81
Pomacea 75
spixii, Ampullaria 49
Asolene 49
sprucei, Ampullaria 75
Pomacea 75
stagnalis, Lymnaea 123
Stagnicola 143
Stenogyridae 185
storeria, Ampullaria 49
Asolene 49
stratiotes, Pistia 454
Straussia 175
strebeli, Helicina 363
Pomacea 75
Streptaxidae 185
Streptostele musaecola 185-186
(Tomostele) musaecola 185-186,
185-186
Striatemoda 414
Suaeda 130
subplanata, Corbicula 4, 6, 21-22, 26-27,
35-36
subquadrata, Montacuta 169-170, 171
Mysella 170
Rochefortia 170
Semelina 169-170, 171, 172-173
Subulina octona 244
Subulinidae 185
Succincta 196, 198, 390, 395, 423-428
succincta, Helicina 395
Succinea costaricana 244
putris 175
thaanumi 175-176
Succineidae 175
sumatrana, Corbicula 4, 6, 11, 13, 14, 16

INDEX

superba, Ampullaria 75
Pomacea 75

suprafasciata, Pomacea 55

Surinamia 78

swainsoni, Ampullaria 75
Pomacea 75

swainsonii, Ampullaria 75
Pomacea 75

talamancensis, Helicina 195, 197, 252, 308,
310-312, 311-317, 318, 357, 387-388,
409-410, 413, 415, 420, 426-427, 432,
434

Helicina (Gemma) 215, 308
Oligyra 308
Tamsiana 397, 423-427
tamsiana, Ampullaria 56
Helicina 397
Pomacea 56

tankervillii, Eutrochatella 420

tasmaniae, Ampullaria 81

Tellinidae 169

Tellinoidea 169, 173

tenagophila, Biomphalaria 444

tennuis, Helicina 256

tenuifloris, Carduus 126-127, 129

Tenuis 397, 423-424

tenuis, Helicina 195, 197, 214, 254-256,
256, 258, 262, 262-264, 265-271, 271,
315-316, 353, 387-388, 397, 409-410,
410-411, 411-414, 420, 422-424,
426-428, 431

Helicina (Helicina) 256
Helicina (Tenuis) 255
Helicina (Tristramia) 214, 254-255 .
Helicina tenuis 256
var. chiapensis, Helicina 255
var. lindeni, Helicina 255
tenuissima, Ampullaria 76
Pomacea 76
teres, Ampullaria 56
Pomacea 56
terryae, Helicina 308, 309, 316
testudinea, Ampullaria 76
Pomacea 76

thaanumi, Succinea 175-176

titanica, Emoda pulcherrima 421

tobae, Corbicula 4, 35

Tomostele 186

towutensis, Corbicula 21-22

Trichohelicina 370, 427

tristis, Ampullaria 56, 80

Bulimus 80
Pomacea 56
tristrami, Ampullaria 76

INDEX 479

Pomacea 76 Pomacea 76
Tristramia 196, 198, 395, 424-429 Viana regina 415, 419
trochulus, Ampullaria 80 Vianinae 425, 429
tropica, Helicina orbiculata 395 viatrix, Lymnaea 444
tumida, Corbicula 4 Vica sativa 127
tuncki, Helicina 215 vickeryi, Pomacea 76
Turbinata 423 villata, Pomacea 56
turbinata, Helicina 395-396, 395-396, 421, vindobonensis, Cepaea 133-137, 134, 136,
436 139
Tylomelania 34-35, 37 violacea, Ampullaria 77
typica, Lymnaea 141 Pomacea 77
virginiana, Semele 169
umbonata, Helicina 413, 425 Viviparidae 41
unicolor, Pomacea 76 Viviparus 143
unidentata, Helicina 377, 379
urabaensis, Pomacea 76 Waldemaria japonica 421
urceus, Bulimus 51 welwitschiana, Ampullaria 77
Nerita 51, 76 Pomacea 77
Pomacea 41, 76 woodwardi, Ampullaria 77
Urtica 128 Pomacea 77
dioica 126-129, 135
Urticaceae 126 yatesii, Ampullaria 77
Utricularia 180 Pomacea 77
yucatanensis, Ampullaria 77
Variabilis 423, 427 Pomacea 77
Varicella 185 var. yzabalensis, Ampullaria 77
clappi 185 yzabalensis, Pomacea 77
variegata, Pomacea 81
veneriformis, Mactra 119 zephyrina, Helicina 396, 428
venetus, Ampullaria 76 Helicina zephyrina 419
Pomacea 76 zeteki, Pomacea 77
verapazensis, Helicina vernalis 255, 271 zischkai, Pomacea 77
verbecki, Corbicula 13 zonata, Ampullaria 49, 78
vermiformis, Ampullaria 76 Asolene 49
Pomacea 76 Pomacea 78
vernalis, Helicina 254-257, 257, 263, 266, Zonitidae 121
270 zunilensis, Helicina punctisulcata 271, 278,

vexillum, Ampullaria 76 278, 282-283

MALACOLOGIA

International Journal of Malacology

Vol. 45(2) 2004

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VOL. 45, NO. 2 MALACOLOGIA 2004

CONTENTS

CLAUDIO G. DE FRANCESCO & FEDERICO 1. ISLA
Reproductive Period and Growth Rate of the Freshwater Snail Heleobia
Parchappii (d’Orbigny, 1835) (Gastropoda: Rissooidea) in a Shallow

Brackish Habitat (Buenos Aires Province, Argentina) ................. 443
ROGER J. HARO, RICK GILLIS, & SCOTT T. COOPER

First Report of a Terrestrial Slug (Arion Fasciatus) Living in an Aquatic

O A и O A 451

IRA RICHLING
Classification of the Helicinidae: Review of Morphological Characteristics
Based on a Revision of the Costa Rican Species and Application to the
Arrangement of the Central American Mainland Taxa (Mollusca: Gastropoda:
Nello psa acia aire meets pia ai e ly Sa 195
ALEJANDRA RUMI, DIEGO E. GUTIÉRREZ GREGORIC, M. ANDREA ROCHE,
8 MONICA P. TASSARA
Population Structure in Drepanotrema Kermatoides and D. Cimex
(Gastropoda, Planorbidae) in Natural Conditions ..................... 453

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ААА SS

ИИА

VOL. 45, NO. 2 MALACOLOGIA

CONTENTS

IRA RICHLING 23
Classification of the Helicinidae: Review of Morphological Characteristics
Based on a Revision of the Costa Rican Species and Application to the
Arrangement of the Central American Mainland Taxa (Mollusca: Gastropoda:
Nenlopsina) essa ads ata de ato a а
RESEARCH NOTES
CLAUDIO G. DE FRANCESCO & FEDERICO I. ISLA
Reproductive Period and Growth Rate of the Freshwater Snail Heleobia
Parchappii (d’Orbigny, 1835) (Gastropoda: Rissooidea) in a Shallow
Brackish Habitat (Buenos Aires Province, Argentina) .................
ROGER J. HARO, RICK GILLIS, & SCOTT T. COOPER
First Report of a Terrestrial Slug (Arion Fasciatus) Living in an Aquatic
| о...
ALEJANDRA RUMI, DIEGO Е. GUTIÉRREZ GREGORIC, M. ANDREA ROCHE,
& MONICA P. TASSARA
Population Structure in Drepanotrema Kermatoides and D. Cimex
(Gastropoda, Planorbidae) in Natural Conditions ....................

354 АЙ À

09/18/05 19812 ©

2004

195

443

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