Morphology, reproductive biology and molecular genetics

Blackwell Science, Ltd

of ovoviviparous freshwater gastropods

(Cerithioidea, Pachychilidae) from the Philippines,
with description of a new genus Jagora
FRANK KÖHLER & MATTHIAS GLAUBRECHT

Accepted: 10 December 2001
Köhler, F. & Glaubrecht, M. (2002). Morphology, reproductive biology and molecular genet-
ics of ovoviviparous freshwater gastropods (Cerithioidea, Pachychilidae) from the Philippines,
with description of a new genus Jagora. — Zoologica Scripta, 32, 35 – 59.
The new genus Jagora of the family Pachychilidae Troschel, 1857 is described for the type species
Melania asperata Lamarck, 1822 from the Philippines. In addition to J. asperata, a second endemic
species of the new genus, J. dactylus (I. Lea & H.C. Lea, 1850), is recognized on the basis of shell
morphometry and molecular genetic data (cytochrome C oxidase I and 16S rRNA). The taxonomic
history of J. asperata and J. dactylus is revised and the recent distribution documented on basis of
available museum material and the authors’ own field collections. J. asperata occurs on Luzon and
its satellite islands Leyte and Samar, while J. dactylus is restricted to the Visayan islands Bohol,
Cebu, and Guimeras. The morphology of the two species is presented and illustrated in detail,
and compared to taxa of the closely related genus Brotia H. Adams, 1866 to which they
were previously assigned. Among the South-east Asian freshwater Cerithioidea of the family
Pachychilidae, which were previously subsumed under Brotia for their more or less similar shell
morphology and operculum, three distinct lineages can be distinguished, in particular by means
of distinct reproductive anatomy: (1) the species of Brotia sensu stricto from mainland South-
east Asia, Sumatra, Borneo and Java, which all exhibit a subhaemocoelic brood pouch; (2) the
pachychilid species endemic to Sulawesi, currently assigned to the genera Brotia and Tylomelania,
as well as both species of Pseudopotamis endemic to the Torres Strait Islands, which possess a
uterine brood pouch; (3) the females of the Philippine Jagora, which carry egg capsules,
embryos and advanced juvenile stages within the mantle cavity — a unique reproductive feature.
Associated with this mode of ovoviviparity, Jagora is characterized by additional unique pro-
perties of the reproductive system including a deeply incised and long sperm gutter in the medial
lamina, a very short and posteriorly positioned spermatophore bursa formed by the medial
lamina, and a prominent lateral ridge functioning as a seminal receptacle. These characteristics
are exclusive to Jagora and are consequently considered to represent autapomorphies of this
clade which is endemic only to the Philippines. The zoogeographical implications are discussed
in connection with a recently developed palaeogeographical reconstruction.
Frank Köhler & Matthias Glaubrecht, Museum für Naturkunde, Institute of Systematic Zoology,
Humboldt University, D-10115 Berlin, Germany. E-mail: frank.koehler@rz.hu-berlin.de

Introduction judging from the still continuing description of new species,

The Philippine islands are one of the world’s major ‘hot spots’
of species richness and a centre of high endemism, having one
of the highest per-area levels of biodiversity world-wide (Depart-
ment of Environment and Natural Resources 1997; Myers
et al. 2000b; Mittermeier et al. 2000). For example, a total of
519 vertebrate species are endemic to the Philippines, among
them 64% of all mammals (Heaney 1998). However, the doc-
umentation of Philippine biodiversity is far from complete
in particular of invertebrate taxa. Even less ecological, bioge-
ographical or evolutionary information is available for most
representatives of the Philippine fauna and flora. At the same
time, this rich biota is under severe threat due to the accelerating
destruction of natural habitats. This also holds true for fresh-
water ecosystems, which are threatened by pollution, overhar-
vesting, flow regulations and impoundment of rivers, as well as
drainage basin degradation and climatic changes (Dudgeon 2000).
Jagora from the Philippines • F. Köhler & M. Glaubrecht
In addition, the Philippine archipelago, being highly
fragmented and isolated, is one of the most fascinating areas for
biogeographical research. Although much research has been
carried out in recent years, most studies traditionally deal
with vertebrates (e.g. Dickerson 1928; Dickinson 1991;
Ruedi 1995; Parenti & Louie 1998; Heaney 1985, 1986, 1998),
some with insects (e.g. De Jong 1996), whereas other specious
phyla are still underrepresented — for example, gastropods
among the nonmarine molluscs. This stands in contrast to
the fact that land and freshwater gastropods serve as excellent
model organisms for evolutionary and zoogeographical
studies due to a rich and extensive fossil record, limited
dispersal ability and high habitat fidelity in comparison to
birds or mammals (Glaubrecht 1996, 2000).
However, the confusing taxonomy and lack of phyloge-
netic systematics of many (particularly limnetic) gastropods
has previously often hampered further insight into their
evolution, ecology and historical biogeography (for some earlier
approaches, see Davis 1982; Ponder 1988; Glaubrecht 1996).
In recent years, our understanding of molluscan phylogeny
has become much more detailed and substantive due to a
growing body of morphological data, application of cladistic
analyses and the increased application of molecular genetic
techniques, resulting in an increasing number of phylogenies
for gastropods, among them several limnetic groups (e.g.
Reid 1996; Lydeard et al. 1997; Davis et al. 1998; Ridgeway
et al. 1998; Winnepenninckx et al. 1998; Koufopanou et al.
1999; Medina & Walsh 2000; Holznagel & Lydeard 2000).
One of these limnetic groups is Cerithioidea Ferussac,
1819, a basal caenogastropod superfamily with about 17 pre-
dominantly marine, but also some brackish and freshwater,
families. In total, Cerithioidea contains c. 200 genera and
several thousand species. They are pantropically distributed and
of great ecological importance as grazers and detritus feeders
in most tropical to subtropical aquatic ecosystems. Among
them, the so-called ‘melaniids’ or Thiaridae sensu lato repre-
sent one of the richest gastropod faunas in limnetic biotopes
and at the same time one of the least known taxonomically.
These thiarid freshwater cerithioideans were long neglected
by malacologists, resulting in a largely ignored or under-
utilized resource for zoological studies. Their systematics and
taxonomy has hitherto been strongly influenced by the typo-
logical approach of the 19th and early 20th centuries (e.g.
Lamarck 1822; Reeve 1859−60; Brot 1874; Nevill 1884; Von
Martens 1897; Thiele 1929; see also Glaubrecht 1996, 1999
and Köhler & Glaubrecht 2001).
Only recently has our understanding of the phylogeny of
freshwater Cerithioidea been improved by incorporating
new morphological and molecular evidence (e.g. Houbrick
1988; Glaubrecht 1996; Holznagel & Lydeard 2000).
According to these and on-going phylogenetic analyses it has
become evident that the species-rich limnetic Cerithioidea
actually comprises several clades that independently invaded
freshwater habitats and that Thiaridae sensu lato represents a
polyphyletic assemblage among which five different families
should be distinguished (Glaubrecht 1999; Glaubrecht et al.
unpubl. data; Lydeard et al. 2002).
A number of cerithioidean taxa are still poorly known and
a comprehensive phylogenetic analysis of the entire group
requires a thorough re-examination of all its constituent taxa.
In order to contribute to a research program aiming at the
evaluation of the relationships within Cerithioidea, which
would also permit the formulation of hypotheses on the
evolutionary ecology of the freshwater lineages, we have recently
studied the morphology of one of the lesser known groups
representing the South-east Asian pachychilid gastropods
hitherto attributed to the widespread genus Brotia Adams,
1866. Our investigations have revealed that these gastropods
have surprisingly diverse features, particularly their repro-
ductive anatomy. This led us to use reproductive anatomy to
distinguish the species groups (Köhler et al. 2000; Köhler &
Glaubrecht 2001).
One of these groups is represented by the endemic Philip-
pine freshwater gastropod originally named Melania asperata
Lamarck, 1822. We describe in detail the females of this and
a second species that exhibit an unique mode of brooding
which clearly differs from the reproductive biology of other
taxa within Brotia, and discuss the systematic and taxonomic
implications. Since a comparative study on the morphology
of Brotia from the entire distributional range as well as from
those taxa endemic to the Philippines has long been lacking,
the taxonomic status and the biological significance of the
many morphospecies described in the 19th century following
an essentially typological approach has remained unresolved.
In order to clarify the puzzling systematics, the present study
will address the following aspects: (1) whether the anatomy
of the Philippine ‘Melania asperata’ supports a generic
separation from Brotia; (2) the status and relationships of the
described Philippine pachychilid morphospecies; (3) whether
there is only one widely distributed and highly variable spe-
cies or many Philippine species with more limited distribu-
tions within the archipelago; (4) the conclusions to be drawn
from the presented data on the reproductive biology, life
history and zoogeography of these freshwater snails. The
morphologically based conclusions will be supported by
preliminary molecular data.
Materials and Methods
Materials
For this study, in addition to the collections of the Museum
für Naturkunde of the Humboldt University, Berlin (ZMB;
formerly Zoological Museum Berlin), types and supplemen-
tary material were examined from the Australian Museum,
Sydney (AMS), the Natural History Museum, London

(BMNH), the Californian Academy of Sciences, San Francisco
(CAS), the Muséum d’Histoire Naturelle, Genève (MHNG),
the Musée National d’Histoire Naturelle, Paris (MNHN), the
Senckenbergmuseum, Frankfurt (SMF), the Überseemuseum,
Bremen (ÜMB), the Zoologisches Museum, Zurich (ZMZ)
and the Zoological Museum Zoological Institute, University
of Hamburg (ZMH). Soft body anatomy was studied using
material preserved in alcohol that was recently collected on
Luzon and Cebu and deposited with ZMB.
Examination of the morphology
Dimensions of shells and protoconchs were measured with a
calliper to 0.1 mm. Shell height (H) is the maximum dimen-
sion parallel to the axis of coiling, breadth (B) the maximum
dimension perpendicular to H, including the aperture.
The length of the aperture (LA) is the greatest length from the
junction of the outer lip with the penultimate whorl to the
anterior lip, the width of the aperture ( WA ) the greatest length
perpendicular to LA. The diameter (DA), height (HA) and
breadth (BA) of the apical whorl were measured as described
by Glaubrecht (1996: 28). Anatomy was studied using a stereo-
microscope with a drawing mirror. Extracted radulae and
protoconchs were cleaned by heating to about 60 °C in 5%
KOH solution and by ultrasound or by soaking in 1% hypo-
chlorite solution and rinsing in distilled water, respectively
(see Reid 1986: 5). They were mounted on specimen stubs
using adhesive splicing tape and coated with gold for examina-
tion with a Jeol JSM 6300 scanning electron microscope. Soft
tissues were prepared using hexamethyldisilazane (Nation 1993).
For histological examination fixed specimens were embedded in
paraffin. Serial sections of 8 µm were prepared using a sliding
microtome and stained with haematoxylin and eosin (HE). The
sex ratio is given as the proportion of males of the entire sample
(number of males/total number of animals examined).
Molecular genetics and sequence analysis
Total DNA was isolated from specimens preserved in ethanol
by application of a CTAB extraction protocol for molluscan
tissues (Winnepenninckx et al. 1993). PCR amplifications
were performed in 25 µL volumes containing approximately
50 ng of DNA, 1 × PCR buffer, 0.5 µM each dNTP, 2.0 mM
MgCl2, 0.5 µM each primer, and 1.25 units of Taq polymerase
(Invitek). After an initial denaturation step of 3 min at 94 °C,
35 cycles of 45 s. at 94 °C, 45 s. at 50 °C, and 60 s. at 72 °C
were performed, followed by a final extension of 5 min at
72 °C. The primers used were LC01490 (F) and a modifica-
tion of HCO2198 (Folmer et al. 1994) with the sequence
5′-AWACTTCTGGGTGKCCAAARAAT-3′ (R) for COI;
modified standard invertebrate primers for 16S (Palumbi
et al. 1991) employed the following sequences: 5′-CCGGT-
YTGAACTCAGATCATGT-3′ (F) and 5′-GTRRGCYT-
AAAATYAGCYAYC-3′ (R) (Wilson unpubl. data). PCR
F. Köhler & M. Glaubrecht • Jagora from the Philippines
products were purified using a Qiagen Purification Kit, and
sequencing was performed on an ABI 377 automated sequencer
using the BigDye Terminator Cycle Sequencing Kit (Applied
Biosystems). For each specimen we directly sequenced a
646-bp fragment of the Cytochrome c oxidase subunit I gene
(COI) and 825 bp of 16S rDNA (16S).
The COI sequences were aligned by eye, the 16S
sequences using Clustal X (Version 1.8) in the multiple align-
ment routine with standard parameters. Maximum parsi-
mony (MP) and neighbour joining (NJ) (Saitou & Nei 1987)
trees were reconstructed with PAUP (Version 4.0b8; Swofford
1998) using standard parameters and the ‘exhaustive search’
option. Pairwise sequence distances, transition and transver-
sion rates were calculated using BIOEDIT for Windows NT
(Hall 1999). The sequences are accessible via GenBank (accession
numbers AY 172438 to AY 172453); alignments can be provided
by the first author.
Systematics
Family placement
Freshwater cerithioideans subsumed under the South-east
Asian genus Brotia, among them the Philippine species
‘M. asperata’, ‘M. dactylus’, ‘M. philippinarum’, ‘M. pulchra’
and ‘M. pagodulus’, were formerly assigned to various families
by different authors (review in Köhler & Glaubrecht 2001).
Traditionally, as with all other freshwater gastropods with an
elongated to turreted shell, these taxa were long considered
as belonging to the large and species-rich family ‘Melaniidae’.
This name, however, is a junior synonym of the valid
family name Thiaridae Troschel, 1857 (for nomenclatorial
details see Glaubrecht 1996: 483–491). Troschel (1856–63)
first suggested differentiating four distinct groups within
the so-called ‘melaniids’, among them the ‘Pachychili’. Later,
Fischer & Crosse (1892: 311) elaborating on Troschel’s
proposal, suggested subdividing the ‘Melaniidae’ into six
taxa, including the Pachychilinae. Thiele (1925: 83; 1928:
381; 1929: 189) then separated this taxon from Thiarinae
on the basis of differences in radular and operculum
morphology. However, he choose the subfamily name
Melanatriinae over Pachychilinae for this group, since he
erroneously considered the generic name Pachychilus Lea
& Lea, 1850 to be preoccupied by Pachychila Eschscholtz,
1831 (Thiele 1921: 142; details in Köhler & Glaubrecht
2001). His view has been followed since then by some
authors (e.g. Wenz 1938; Brandt 1974; Glaubrecht 1996), but
ignored by others (Morrison 1954; Benthem Jutting 1956;
Brandt 1968). Nevertheless, all these authors continued to
place Brotia within Thiaridae. In contrast, Vaught (1989)
suggested a placement within the (otherwise North American)
Pleuroceridae following a suggestion of Morrison (1954) and
Ponder & Warén (1988); for further details see review in
Glaubrecht (1999).
Jagora from the Philippines • F. Köhler & M. Glaubrecht
Based on new data, a comprehensive phylogenetic analysis
of c. 180 morphological characters of Cerithioidea in concert
with a parallel molecular analysis supports the separation of
Pachychilidae (as well as Pleuroceridae) from Thiaridae sensu
stricto and other distinct limnetic clades (Glaubrecht 1996,
1999; Strong & Glaubrecht 1999; Glaubrecht et al. unpubl.
data; Lydeard et al. 2002). Therefore, we here consider the
Oriental genera Brotia, Tylomelania Sarasin & Sarasin, 1897
as well as the Australian Pseudopotamis Martens, 1894, the
African Potadoma Swainson, 1840 together with Melanatria
Bowdich, 1822 from Madagascar and the Neotropical Pach-
ychilus as well as the herein newly described Philippine Jagora
as forming a monophylum; for nomenclatorial reasons the name
Pachychilidae should be preferred over the name Melanatri-
idae and applied to this pantropically distributed clade.
Pachychilidae Troschel, 1857
Jagora gen. n.
Type species. Melania asperata Lamarck, 1822.
Etymology. This genus is named after the German naturalist
and anthropologist Andreas Feodor Jagor (1817−1900), who
between 1859 and 1860 travelled through the Philippine
archipelago ( Jagor 1873). Shells collected on his journey are
deposited with ZMB and facilitated our study on shell
variability and distribution of Jagora.
Diagnosis. Freshwater cerithioidean gastropod with a highly
turreted shell, sculptured by axial ribs and spiral striae. Axial
and spiral elements in some specimens form spiny nodules at
their junctions. Aperture well rounded and slightly angled
below. Operculum multispiral with three to five whorls
increasing in diameter and with subcentral nucleus. Animal
large, comprising up to eight whorls. Head with two moder-
ately long cephalic tentacles, each with a tiny eye at the outer
base. Rather shallow genital groove at the right side of the
head. Flap formed by the mantle roof protruding from the
mantle cavity dorsally to the anterior end of the pallial
gonoduct. Open pallial gonoduct in both sexes. Pallial oviduct
with capsule gland at the base of the oviductal groove and an
extensively elongated free medial lamina on which two sperm
gutters and a short spermatophore bursa develop. Lateral
lamina attached to the body wall, with a papillated and cham-
bered ridge functioning as a sperm storing organ (receptacu-
lum seminis). Females ovoviviparous, retaining egg capsules,
developing embryonic and juvenile stages within the mantle
cavity; egg capsules contain 4 –12 eggs. Long taenioglossate
radula with rachidian teeth possessing a main denticle flanked
by three accessory cusps that taper in size, and a well
developed glabella. Lateral teeth with a main denticle flanked
by two or three smaller cusps. Inner and outer marginals with
two pointed denticles of different size. Stomach with a crys-
talline style, the style sac communicating with the intestine,
a gastric pad and shield, and a sorting area formed by two
transversally folded crescent thickenings.
Taxonomic history. Most classifications of ‘melaniid’ species
were suggested almost exclusively on basis of shell features,
e.g. those by Adams & Adams (1858), Nevill (1884) and
Preston (1915). Troschel (1857) was the first to recognize an
affinity between the Philippine M. asperata and the Neotropical
Pachychilus, noticing shared characteristics of the radula (i.e.
the rachis with one main cusp flanked by accessory cusps that
taper in size, and the glabella) and the multispiral operculum.
Consequently, he treated both as congeneric, referring to
the names P. pulcher and P. dactylus. Later, Fischer & Crosse
(1891−92) described a new genus, Antimelania, also based on
the radula, for the Oriental species M. asperata, M. filocarinata,
M. dactylus, M. cancellata Benson, 1836, and M. variabilis
Benson, 1836. Subsequently, the latter species, which is a
synonym of Brotia costula (Rafinesque, 1833), was designated
the type species of Antimelania by Pilsbry & Bequaert (1927:
300). Thus, Antimelania is rendered a synonym of Brotia, as
was pointed out by Morrison (1954), Brandt (1974) and
Köhler & Glaubrecht (2001). It was Abbott (1948), who first
explicitly grouped Lamarck’s M. asperata with Brotia. Since
then, this suggestion has been followed in the few publications
available (e.g. Burch 1980).
A comprehensive overview of the Philippine taxa described
or previously mentioned in the literature is given under the
respective species below. Most of these names are frequently,
albeit ambiguously used in museum collections. The existing
taxonomic confusion can be illustrated by some examples.
Reeve (1859: pl. 2, fig. 6) listed a ‘Melania reirvardii de Haan’
[sic!] as a synonym of M. inquinata Defrance, 1825, previ-
ously also referred to as ‘Melanoides reenivardii de Haan’ by
Adams & Adams (1858: 297). Later, Brot (1874) treated the
former name as a synonym of Melania perfecta (Mousson 1849)
(while he assigned M. inquinata as a synonym to M. asperata).
However, we were able to trace neither a formal description
nor any potential type material of M. reenivardii or M. reirvardii,
and therefore consider these names as nomina nuda.
Brot (1874) synonymized M. pulchra and M. pagodulus
with M. asperata, as well as M. philippinarum with M. dactylus.
He also described a new species, M. filocarinata Brot, 1874.
He distinguished between three forms, each characterized
by a different shell sculpture: M. asperata with a dark
shell, pronounced axial ribs and prominent spiral lirae,
M. dactylus with a yellowish to greenish colour and closely
spaced axial ribs as well as fine (‘microscopic’) spiral striae,
and M. filocarinata exhibiting more distinct axial ribs and a
‘remarkable thin spiral ridge at the suture’ (Brot 1874: 76).

This view was followed, albeit not consistently, by sub-
sequent authors presupposing the existence of at least three
closely related species in the Philippines. Nevill (1884)
distinguished a total of five ‘varieties’ or ‘subvarieties’ of
M. asperata and also described a new var. imbricata of
M. filocarinata using subtle conchological characteristics.
However, these subdivisions were based on material which
was almost entirely collected at the same locality (Pagsanhan,
Laguna Province, Luzon). Sarasin & Sarasin (1898: 37)
described a new subspecies, M. asperata celebicola, from
Sulawesi which was considered a synonym of M. perfecta by
Bollinger (1914) and is supported by our finding that the
pachychilids of Sulawesi form a distinct monophyletic group
(Rintelen & Glaubrecht 1999; Köhler et al. 2000; Köhler &
Glaubrecht 2001).
To complicate matters still further, the Philippine J. asperata
was often confused with M. inquinata, a fossil species from
the Paris Basin, again due to similar shells (see criticism
in Sowerby 1838). However, establishing any connection
between a taxon from marine Tertiary deposits of Western
Europe and a recent limnetic element of the Philippine
fauna is likely to be fruitless. Instead of implicitly inferring
a close relationship of those taxa widely separated both in
space and time, the similar shell of both species should
rather be considered a case of homoplasy.
The present analysis of morphology and molecular
genetics suggests recognition of two Philippine species,
J. asperata and J. dactylus. However, we do not agree with
Brot’s (1874) synonymy of M. philippinarum with J. dactylus,
as the former taxon was described from Luzon. Instead,
M. philippinarum is considered here a synonym of
J. asperata.
Jagora asperata (Lamarck 1822) comb. n.
Melania asperata Lamarck, 1822: 164, no fig. (‘Manille’);
Deshayes & Milnes Edwards, 1838: 639 (‘les rivières de
l’Amerique meridionale? Mon cabinet.’); Reeve, 1859: pl. 1,
fig. 2a–e; Chenu, 1859: 288, 955; Brot, 1862: 45; Brot, 1872:
7–8, pl. 1, fig. 5; Brot, 1874: 73, pl. 8, fig. 1a–g.
Melanoides aspera — H. Adams & A. Adams, 1858: 297, pl. 31,
fig. 5c.
Melania (Melanoides) asperata — Nevill, 1884: 264–265
(including subvar. sublaevigata, var. inquinata and three
additional, unnamed varieties).
Melanoides asperata — Preston, 1915: 21.
Brotia asperata — Abbott, 1948: 298, pl. 3, fig. 1; Knipper,
1958: 49; Köhler & Glaubrecht, 2001.
Antemelania asperata — Cabrera, 1984: 1188.
Melania philippinarum Sowerby 1838: pl. 1, figs 1–3 (‘Philip-
pines, Manila’).
Melania pulchra von dem Busch, 1848 — In: Philippi, 1848:
pl. 5, fig. 1 (‘Philippines’); Reeve, 1859: pl. 4, fig. 19.
F. Köhler & M. Glaubrecht • Jagora from the Philippines
Pachychilus pulcher — Troschel, 1857: 116, pl. 9, fig. 4.
Melanoides pulchra — H. Adams & A. Adams, 1858: 297.
Melania pagodulus Reeve, 1859: pl. 12, fig. 72 (locus typicus
unknown).
Melania inquinata Reeve, 1859 (non Defrance 1825): pl. 2,
fig. 6; Brot, 1862: 45.
Melania filocarinata Brot, 1874: 76, pl. 9, fig. 1 (‘Polillo’).
Melania (Melanoides) filocarinata — Nevill, 1884: 266.
Type material examined. Holotype of M. asperata Lamarck,
1822 (MHNG 1093/60, ‘Manille’, Fig. 1A); lectotype of
M. pulchra von dem Busch, 1848, designated by Knipper
(1958) (ÜMB TK 236/1, ‘Philippines’, Fig. 1B); 3 syntypes of
M. pagodulus Reeve, 1859 (BMNH 19990494, locus typicus
unknown, Fig. 1C); 2 syntypes of M. filocarinata Brot, 1874
(ZMZ 522338, ‘Philippines, Polillo’, Fig. 1D).
Other material examined. Philippines: (ZMB 2222, 77093,
88840, 200012, 200017, 200019, 200022–9, 200031–4,
200041, 200077; CAS 6195–6); Luzon: (ZMB 200014–5;
ZMH; BMNH); Manila (ZMB 26714, 200016); Zambales
mountains, stony mountain stream near Iba (ZMB 200113);
North Luzon, Isabella Province, Panasinang (ZMB 200114);
Laguna Province (ZMB 27268; MHNG; AMS 324130; CAS
11306); Laguna Province, Barangay Masaya near Los Baños,
muddy stream (14°10,5′ N, 121°17,4′ E) (ZMB 200111);
Laguna Province, Barangay Masaya near Los Baños, irriga-
tion channel (14°15,31′ N, 121°24,4′ E) (ZMB 200115);
Laguna Province, Mt. Makinlang near Los Baños, 400 m
above sea level (ZMB 200112); Laguna Province, Antipolo
River (BMNH 1913.11.24.33–37; CAS 3972); Laguna
Province, Sta. Cruz (ZMB 78325); Laguna Province, Calauan
(ZMB 26766; MHNG); Tayabas Province, Sampaloc (SMF
292319); Tayabas Province, Mantrao (SMF 292320); Cama-
rines Province, Mambulao (SMF 292346); Albay Province,
Daragu River (ZMB 26768, 200013); Manvelu (ZMB
200011); Sibul (BMNH 1909.10.12.34–44); Brunot Lake,
San Pedro (AMS 324135); Isl. Marinduque (SMF 292346;
292371). Isl. Tablas (SMF 292351). Isl. Polillo (ZMB 200020;
MHNG). Samar (ZMB 26767; MHNG); Loquilocum (ZMB
26765, SMF 292361); Basey (CAS 23860). Leyte (ZMB
200035). Negros (BMNH).
Description
Shell (Fig. 1A–D, G–H): solid and uniformly dark to olive
brown or yellowish brown. The spire is highly turreted with
an average angle of 25° and comprises up to 12 (mean: 8.3)
whorls. The apex is usually eroded or truncated; the sculpture
is variable consisting of both axial and spiral elements.
Generally, on the upper three to five whorls closely spaced
axial ribs predominate while on subsequent whorls these ribs
become indistinct and spiral elements are more pronounced.
In some specimens up to 1 mm long, spiny nodules are
formed where axial and spiral elements meet. In other
Jagora from the Philippines • F. Köhler & M. Glaubrecht

Fig. 1 A–H. Shells. —A. Holotype of M. asperata Lamarck, 1822 (MHNG 1093/60, ‘Manille’). —B. Lectotype of M. pulchra von dem Busch,
1848 (ÜMB TK 236/1, ‘Philippines’). —C. Syntype of M. pagodulus Reeve, 1859 (BMNH 19990494, locus typicus unknown). —D. Syntype of
M. filocarinata Brot, 1874 (ZMZ 522338, ‘Polillo’). —E. Syntype of M. dactylus I. Lea & H.C. Lea, 1850 (BMNH 19990496, ‘Guimeras,
Philippines’). —F. Shells of J. dactylus from Cebu (Matutinao River, ZMB 200108). —G. Shells of J. asperata from Luzon (Laguna Province,
Calauan, ZMB 26766). —H. Shells of J. asperata from Samar (Loquilocum, ZMB 26765). Scale bar = 5 mm.
 F. Köhler & M. Glaubrecht • Jagora from the Philippines

Table 1 Means and standard deviations (in

Parameters (mm)
parentheses) of shell parameters. Species
Species Population examined (n) H B LA WA BW

J. asperata Luzon, ZMB 200114 14 45.0 (5.7) 16.5(1.3) 12.5 (1.2) 8.2 (0.8) 21.6 (1.7)
Luzon, AMS 324130 6 47.1 (3.0) 16.3 (1.4) 12.5 (0.9) 8.3 (0.8) 22.2 (1.5)
Luzon, ZMB 26726 20 47.8 (6.6) 16.8 (2.2) 13.2 (1.3) 8.3 (1.3) 22.6 (2.6)
Samar, ZMB 26767 17 0.5 (5.7) 17.5 (1.9) 14.2 (1.5) 8.9 (1.2) 24.0 (2.7)
J. dactylus Bohol, AMS 324129 5 48.7 (4.2) 16.8 (0.8) 13.5 (0.4) 8.7 (0.3) 23.4 (1.1)
Cebu, ZMB 200108–10 30 46.7 (5.8) 16.3 (6.1) 11.8 (1.1) 7.4 (1.1) 20.9 (2.3)
Guimeras, BMNH 3 75.4 (3.0) 23.7 (2.5) 18.7 (0.5) 12.4 (1.3) 31.7 (1.5)

n, number of specimens examined; H, shell height; B, shell breadth; LA, length of aperature; WA, width of aperature;
BW, body whorl.

individuals, however, either the spiral or the axial elements
may be almost completely displaced. The aperture is oval and
rounded below; the outer lip has a thin to thick margin. Mean
shell dimensions (standard deviations in parentheses) were:
H = 48.5 (± 6.0) mm, B = 17.3 (± 2.3) mm, LA = 13.4 (± 1.6)
mm, WA = 8.6 (± 1.4) mm, BW = 23.0 (± 2.7) mm (n = 57)
(Table 1).
Protoconch: embryos of one clutch were always found to be
of the same developmental stage and, thus, of about similar
size. The development of the juvenile shell is identical with
J. dactylus as described and depicted below (Fig. 7B–H). The
apical whorl, when calcified, is plug-like and has a chara-
cteristically wrinkled structure while subsequent whorls are
sculptured by axial ribs and tiny spiral growth lines. Only one
female from Luzon was found to contain juveniles in its
mantle cavity. The mean dimensions of these juvenile shells
are: HA = 1.7 mm, BA = 1.2 mm, DA = 628 µm (n = 10).
Operculum (Fig. 2D): the operculum is ovate and multispiral
with three to five rapidly increasing whorls and a slightly
eccentric nucleus. It is solid, measures up to 15 mm in height,
and almost fits the aperture.
External morphology and mantle cavity organs (Fig. 2A–B):
animals are relatively large, with the body comprising up to
eight whorls. The skin is dark grey, occasionally with light
patches. The snout is broad and furrowed; the two cephalic
tentacles are long, each with a tiny eye on the outer side of the
base. At the right side of the head a shallow genital groove
runs down from the end of the pallial gonoduct and ends
beyond the right tentacle.
The mantle edge is smooth. A fleshy flap, formed by the
ventral surface of the mantle roof projects from the mantle
cavity dorsal to the end of the pallial gonoduct and the anus.
The mantle cavity is deep, comprising about one and a half
whorls. The osphradium forms a narrow ridge embedded in
a shallow trench and lies adjacent to the anterior part of the
ctenidium. It is delicate and often slightly undulating. The
ctenidium begins shortly behind the mantle edge, extending
posteriorly almost the entire length of the cavity. It is large,
on average twice as long as the osphradium, broad and tapers
posteriorly. Gill blades are sharkfin-shaped to triangular. The
hypobranchial gland is inconspicuous and lies adjacent to the
large rectum.
Nervous system: the nervous system is epiathroid. The
cerebral commissure is long while connectives between cerebral
and pedal ganglia and pleural and pedal ganglia, respectively,
are short. Pleural and cerebral ganglia are fused. The
suboesophageal ganglion is fused to the left pleural ganglion.
The pedal ganglia are deeply embedded in the muscle of
the propodium. They are closely joined to each other and
numerous crystalline statoconia are located basally. Seven
main nerves branch from each of the cerebral ganglia inner-
vating the snout, tentacles and skin; one of them connects to
the buccal ganglion.
Renal system: the kidney lies behind the mantle cavity
adjacent to the rectum and the gonoduct. It is highly com-
partmentalized, glandular and comparatively large, occupying
about one half of the second whorl extending into the pallial
roof.
Radula (Fig. 3A–B, E–F): the taenioglossate radula is long
consisting of 130–220 rows (n = 8) and has a length of up to
25 mm corresponding to approximately half of the average
shell height. Rachis with one main denticle flanked by three
accessory cusps on each side that taper in size. The glabella
is well developed and broad, rounded at its base and has a
concave lateral edge. The anterior margin of the rachis is
convex. The lateral teeth, with a well developed glabella, have
one large triangular main denticle flanked by three smaller
cusps at the outer side and two cusps at the inner side. The
inner and outer marginal teeth are curved (or ‘kneed’) and
possess two pointed cusps with the outer one being a little
broader. The inner marginal teeth are spatula-shaped; the
outer marginals are narrower. Inner and outer marginals
exhibit a simple, inconspicuous flange at their outer rim.
Alimentary system: the salivary glands are large, consisting
of many, heavily branched, delicate tubules, and are located
anterior to the nerve ring. The oesophagus is longitudinally
folded; transverse septae have not been found. The stomach
exhibits features including a heavily ciliated style sac with a
Jagora from the Philippines • F. Köhler & M. Glaubrecht

Fig. 2 A–D. Soft body anatomy of Jagora asperata (Luzon, ZMB 200111). —A. External anatomy of a female; mantle cavity exposed and filled
with egg capsules. —B. Reconstruction of the external female anatomy. —C. Stomach. —D. Operculum. Abbreviations: cf — crescent fold,
dd — opening to the digestive gland duct, dg — digestive gland, eg — egg capsules, ft — foot, gg — genital groove, gp — gastric pad, gs — gastric
shield, i — intestine, kd — kidney, mf — mantle flap (B), marginal fold (C), mr — mantle roof, ov — ovary, sa — sorting area, st — stomach
(B), opening to the style sac (C), tn — tentacle, ty — typhlosoles. Scale bars: A, B = 10 mm; C, D = 5 mm.

cylindrical crystalline style, a single digestive gland duct, a anterior to the opening of the digestive gland duct being
sorting area formed by two transversally folded crescent fused to the glandular pad. Style sac and intestinal groove
thickenings, a long glandular pad with a cuticular gastric shield, communicate as minor and major typhlosoles are unfused
and a crescent ridge and groove with the inner marginal fold (Fig. 2C). The digestive gland occupies the visceral whorls.
 F. Köhler & M. Glaubrecht • Jagora from the Philippines

Fig. 3 A–F. SEM images of the radula. —A. Segment of a radula of J. asperata from Luzon (ZMB 200114). —B. Detail of A. —C. Segment of
a radula of J. dactylus from Cebu (ZMB 200108). —D. Same radula, lateral view. —E. Segment of a radula of J. asperata from Luzon, lateral
view (ZMB 200114). —F. Detail of E. Scale bars = 0.1 mm.
Jagora from the Philippines • F. Köhler & M. Glaubrecht
Table 2 Sex ratios in the two Philippine Jagora species.
Population Sex ratio
J. asperata Luzon, La Laguna (AMS 324130) 0.33
Luzon, Los Baños (ZMB 200112) 0.33
Luzon, Isabella Province (ZMB 200114) 0.50
Philippines (MNHN) 0.40
Subtotal 0.43
J. dactylus Cebu, Matutionao River (ZMB 200108) 0.42
Bohol, San Isidro (AMS 324129) 0.33
Subtotal 0.39
Reproductive system: J. asperata is gonochoristic. The
average sex ratio among the four populations examined
from Luzon is 0.43 (Table 2). The reproductive anatomy of
all examined specimens from several populations on Luzon
( J. asperata) as well as from Bohol and Cebu ( J. dactylus) was
found to be consistent for both species.
Males are aphallic and possess a prostate basely lined by broad
glandular lamellae extending to the mouth of the mantle cavity.
The testis lies dorsal to the visceral whorls and almost com-
pletely surrounds the digestive gland which occupies only the
interior segment of these whorls. The testis consists of thin
tubules embedded into loose connective tissue (Fig. 4C–D).
In females, the pallial oviduct comprises a deep oviductal
groove dorsally bounded by the free medial lamina. Capsule
gland and albumen gland are not distinguishable by their
colouration in the histological sections. Glandular tissue
generally is dark blue stained in histological sections and
comprises the base of the oviductal groove for its entire
length. The whole surface of the medial lamina and the inner
ventral side bounding the oviductal groove (usually referred
Males : females to as the lateral lamina) is covered by a ciliated epithelium
1:2 (Figs 5A–F and 6). The medial lamina gives rise to two sperm
2:4 gutters: one gutter folds almost at the anterior end of the
4:4 medial lamina (Fig. 5A) and deepens toward the posterior
2:3
end of the gonoduct (Fig. 5B–F). A second gutter forms at
9 : 13
5:7 about half of the length of the pallial oviduct (Fig. 5E–F).
2:4 At the inner ventral side of the gonoduct (or ‘lateral
7 : 11 lamina’) a papillated ridge arises at about half of the length of
the pallial gonoduct and becomes larger posteriorly. This
ridge forms chambers in which orientated sperm was found
(Fig. 5D–F). For that reason, this structure is considered to
have a sperm storing function and herein referred to as sem-
inal receptacle. It extends far into the lumen of the second
sperm gutter (Fig. 5F, see reconstruction in Fig. 6).
At about 3/4 of the oviductal length, a simple, papillated
spermatophore bursa is formed by fusion of the edges of the
second sperm gutter (Fig. 5G). The bursa is short and closes
rapidly. Posteriorly, the oviductal groove closes to a blind
pouch that is lined by glandular tissue — probably the
albumen gland (Fig. 5H). Eventually, the renal part of the
oviduct branches off and the remaining pouch closes com-
pletely. At the posterior part of the pallial gonoduct — ante-
rior to the nephroporus — laminated tissue occupies a part of
the mantle cavity (Fig. 5H). This tissue of unknown function
is formed by the mantle roof but also occupies external
regions of the very posterior part of the gonoduct.
Females retain egg capsules and even shelled juveniles
within the mantle cavity. One of three females collected in
Fig. 4 A–D. Histological sections of visceral
whorls showing the digestive gland and
the gonads ( J. dactylus, Cebu, ZMB 200109).
—A. Cross-section of an entire whorl,
showing the ovary. —B. Detail of A, ovary
embedded within loose connective tissue.
—C. Visceral whorl of a male, showing
the testis embedded within loose connective
tissue. —D. Detail of C, orientated sperm
in the testicular tubules. Abbreviations:
ct — connective tissue, dg – digestive gland,
ov — ovary, te — testis. Scale bars: A = 10 mm;
B−D = 1 mm.
 F. Köhler & M. Glaubrecht • Jagora from the Philippines

Fig. 5 A–H. Histological sections of the pallial oviduct ( J. dactylus, Cebu, ZMB 200108) from anterior (A) to posterior (H). A to F depict
histological sections of an isolated oviduct of a female while in G and H the gonoduct is figured as part of sections of a whole specimen, second
female from the same batch. —A. Most anterior section, near the mantle edge, showing the free medial lamina bounding the oviductal groove
and giving rise to the sperm gutter. —B. Section of the anterior part of the pallial oviduct showing the deep sperm gutter and the capsule gland
comprising the base of the oviductal groove. —C. Section at about one third of the oviductal length. —D. Section at about half of the oviductal
length showing the lateral lamina giving rise to a papillated ridge functioning as a sperm storing organ. —E and F. Sections within the second
third of the oviductal length, showing the rise of a second ciliated sperm gutter and the well developed lateral ridge. —G and H. Sections of
the posterior part of the oviduct at the end of the mantle cavity, with G showing the margins of the second sperm gutter fused to form the
short spermatophore bursa, and the lateral ridge reduced. —H. Lateral and medial lamina are fused and the oviductal groove forms a pouch
lined by glandular tissue (albumen gland?). Laminated tissue with unknown function is formed within the mantle cavity. Abbreviations:
cg — capsule gland, ft — foot muscle, lt — laminated tissue, ml — medial lamina, mr — mantle roof, og — oviductal groove, re — rectum,
sb — spermatophore bursa, sg — sperm gutter, sr — seminal receptacle, ssg — second sperm gutter. Scale bars = 1 mm.
Jagora from the Philippines • F. Köhler & M. Glaubrecht
Fig. 6 Schematic reconstruction of the pallial oviduct of J. dactylus
based on histological sections; arrows and capital letters indicate the
approximate position of the histological sections shown in Fig. 5.
Abbreviations: ag — albumen gland, cg — capsule gland, ft — foot
muscle, ml — medial lamina, og — oviductal groove, ov — oviduct, sb
— spermatophore bursa, sg — sperm gutter, sr — seminal receptacle.
November (AMS 324129) was found to contain 270 juveniles,
while females collected in March 2000 contained neither egg
capsules nor juveniles within their mantle cavities.
Distribution. Mapping of all available samples from museum
collections indicate that the species is restricted to the
northern part of the Philippine archipelago, i.e. Luzon and
some of its satellite islands, such as Polillo and Marinduque,
as well as to Panay, Leyte and Samar (see Fig. 8).
Ecology. The animals inhabit predominantly small mountain
streams with stony to sandy bottoms (Abbott 1948). This was
confirmed by our own field collections in 2000, where these
snails were mostly found buried in the sand or attached to
the underside of leaves, presumably feeding on detritus. At
waterfalls they were also found outside the water grazing on
algae. A report of J. asperata from a lacustrine habitat, Lake
Laguna (Luzon), by Woltereck (1941) is questioned here, as
the occurrence could not be confirmed by our own samplings
in the field. The habitat quality of the muddy, brackish water
lake with abundant vegetation is strikingly different from that
of other known riverine habitats of the species. J. asperata
does, however, inhabit some of the affluent streams, and it is
possible that shells could occasionally be washed into the
lake. J. asperata is first intermediate host of the human lung
fluke Paragonimus westermani (see Cabrera 1984).
Jagora dactylus (I. Lea & H.C. Lea, 1850) comb. n
Melania dactylus I. Lea & H.C. Lea, 1850: 191 (‘Guimeras,
Philippines’); Hanley, 1854–58: pl. 6, fig. 48; Reeve, 1859: pl.
2, fig. 7a–b; Chenu, 1859: 288, 955; Brot, 1862: 45, Brot,
1874: 77–78, pl. 9, fig. 2a.
Pachychilus dactylus — Troschel, 1857: 117, pl. 9, fig. 5.
Melanoides dactylus — H. Adams & A. Adams, 1858: 297.
Melania (Melanoides) dactylus — Nevill, 1884: 265–266.
Antemelania dactylus (sic!) — Cabrera, 1984: 1188.
Type material examined. 3 syntypes (BMNH 19990496,
‘Guimeras, Philippines’, Fig. 1E).
Other material examined. Bohol: Bilar, Bilar river (ZMB
65288; CAS 116978); San Isidro (AMS 324130). Cebu:
Matutinao River at the Kawasan waterfalls (9°48,49′ N,
123°22,1′ E) (ZMB 200108–10). Isl. Guimeras (BMNH
19990496; MHNG). Isl. Siquijor (BMNH).
Description
Shell (Fig. 1E, F): solid and uniformly light brown to yellow-
ish. The spire is highly turreted with an average angle of 25°
and comprises up to 12 (mean 9.3) whorls. The apical whorl
is eroded; the sculpture in general consists of closely spaced
axial ribs and spiral elements. The axial ribs predominate on
the upper three to five whorls, while they become more indis-
tinct on subsequent whorls. Tiny nodules are often formed
where axial and spiral elements meet. The aperture is oval
and rounded below; the outer lip has a sharp to moderately
thick margin. Mean shell dimensions (standard deviations in
parentheses) were: H = 47.0 (± 5.4) mm, B = 15.5 (± 1.8) mm,
LA = 12.1 (± 1.2) mm, WA = 7.6 (± 1.1) mm, BW = 21.3
(± 2.3) mm (n = 38) (Table 1).

Table 3 Number of eggs and juveniles per female.
No. of eggs
Species Population Collecting time (egg capsules) No. of juveniles
J. asperata Luzon (AMS 324129) Nov. 1975 – 275 (–)
J. dactylus Cebu (ZMB 200108) March 2000 ∼50 (9)
Cebu (ZMB 200108) March 2000 ∼90 (12)
Cebu (ZMB 200108) March 2000 – 3 (+)
Cebu (ZMB 200108) March 2000 – 5 (+)
+, shell completely calcified; –, apex uncalcified.
Protoconch (Fig. 7B–H): as described for J. asperata. Juve-
nile shells kept within the mantle cavity were up to 1.7 mm
high and 1.2 mm broad comprising three whorls. Mean
dimensions (standard deviations in parentheses) of the apical
whorl of the protoconch were: DA = 628 (± 34) µm,
HA = 190 (± 66) µm, and BA = 309 (± 24) µm (n = 5).
Soft body anatomy and radula: as described for J. asperata
(Figs 2 and 3C–D).
Reproductive system: anatomy as described for J. asperata.
The average sex ratio among the two populations examined
from Bohol and Cebu is 0.39 (Table 2). Specimens collected
in March on Cebu were in various reproductive states and
contained between 0 and 90 eggs or juveniles in a wide range
of maturity (Table 3). The eggs are enclosed by an egg
capsule, with each containing about 4 –10 eggs; up to 12 capsules
were found within the individuals (Table 3, Fig. 7A). Eggs
contain a considerable amount of whitish granules, presum-
ably yolk. They measure about 1.0 × 1.5 mm in diameter,
thus exceeding the size of protoconchs in early ontogenetic
stages. Although in younger stages the second whorl is
already calcified, it has been observed that nutritive tissue
originating from the egg is attached to the then still uncalci-
fied apical portion of the juvenile shell (Fig. 7B–E). The
amount of the nutritive tissue gradually decreases while the
embryonic shell grows (Fig. 7B–D). Calcification of the
apical whorl is not complete before this material is entirely
consumed. Eventually, crawling juveniles with complete
shells (Fig. 7F–H) are released from the mantle cavity.
Distribution. J. dactylus occurs on the southern Philippine
islands of Bohol, Cebu, Guimeras and Siquijor (Fig. 8).
Following the description of M. dactylus from ‘small streams on
Guimeras, Mindanao, Luzon and Leyte’ by Lea & Lea (1850),
Mindanao was frequently mentioned by subsequent authors
(e.g. Reeve, 1859) as being within the range of J. dactylus.
However, the locus typicus actually is Guimeras, an island south-
east of Panay, as is evident from the original label of the types.
The occurrence on Mindanao could not be confirmed so far.
Ecology. As described for J. asperata, this species inhabits the
upper course of rivers and settles on soft as well as rocky
F. Köhler & M. Glaubrecht • Jagora from the Philippines
substrates (see Bandel & Riedel 1998 for J. dactylus, misidentified
by them as ‘Brotia aff. costula’). J. dactylus is the first interme-
diate host of the human lung fluke Paragonimus westermani
(see Cabrera 1984).
–
Shell morphology and morphometric analysis
–
The type specimens of all described taxa exhibit unusual
shell characteristics. While M. asperata and M. pulchra have
knobbly spiral ribs, in other species nodules or spines are
absent (M. filocarinata, M. pagodulus), or the ribs are smooth
and very closely spaced (M. dactylus). In addition, the types
exhibit different colours from dark brown (M. asperata,
M. pulchra) to yellowish brown (M. pagodulus, M. dactylus)
(Fig. 1C–E). Examination of a series of specimens from a
number of populations, however, revealed that most of these
forms are bridged by intermediates. Based on our observa-
tions, two taxa treated preliminarily as morphospecies can be
distinguished: ‘J. asperata’ with mostly knobbly ribs and
‘J. dactylus’ with more slender shells sculptured by closely and
regularly spaced axial ribs. To the former taxon all material
from Luzon and Samar is assigned, whereas shells from
Bohol and Cebu belong to the latter. In order to establish
statistically significant differences, we analysed several
morphometric shell parameters, counting the ribs on the third
whorl to assess sculptural variation within and between
populations from Luzon, Samar and Cebu. Furthermore, we
analysed the distribution of different shell colorations among
these populations.
We found that specimens from Luzon as a rule have fewer
ribs (less than 10) than those from Cebu (generally more than
10), while specimens from Samar are intermediate (Fig. 9).
Among the examined populations, a considerable degree of
variability was also found concerning the frequency of shell
colours (dark, medium, and light brown). However, instead
of a significant correlation between sculptural variability
and/or shell coloration and spatial distribution with a clear
cut between J. asperata and J. dactylus, we found that these
parameters change clinally from north to south (Fig. 10).
That also holds true for the distributions of several shell
parameters (H, B, LA, WA, BW), which completely overlap
between the examined populations from Luzon, Samar,
Bohol, Cebu, and Guimeras. The mean values and standard
errors for all parameters are given in Table 1. Evaluating the
indices H/B, H/LA, H/BW and BW/LA, shells from Luzon
and Samar (‘J. asperata’) are separated with a slight overlap
from shells of Cebu and Guimeras (‘J. dactylus’). While they
resemble ‘J. dactylus’, this is not found for shells from Bohol,
which cluster together with those from Luzon and Samar
(Fig. 11).
A one-way ANOVA was conducted on the two phenotypes
‘J. asperata’ and ‘J. dactylus’. This analysis reveals that the
parameters H/B, H/LA and H/BW vary significantly
Jagora from the Philippines • F. Köhler & M. Glaubrecht

Fig. 7 A–H. Ontogenetic stages. —A. Egg capsules removed from the mantle cavity ( J. dactylus, Cebu, ZMB 200109). B–H. SEM images of
juveniles at various developmental stages expelled from the mantle cavities of several individuals of J. dactylus from Cebu (ZMB 200108) (B–
E. showing critical point dried specimens). —B. Early ontogentic stage isolated from an egg capsule with a considerable amount of nutritive
tissue originating from the egg; start of shell formation at the second whorl. —C. Embryonic shell isolated from an egg capsule with increased
shell size and nutritive tissue still attached to the apical area of the shell. —D. Later ontogenetic stage expelled from an egg capsule. —E. Detail
of D. —F. Juvenile expelled from the mantle cavity with complete shell calcification, no nutritive tissue visible. —G and H. Details of F. Scale
bars = 1 mm.

Fig. 8 Distribution of J. asperata (circles) and J. dactylus (squares)
based on the compilation of museum material and authors’ own
collections.
Fig. 9 Number of ribs on the third whorl in J. asperata (L1, Central Luzon,
Laguna Province (ZMB 26766); L2, South Luzon, Albay Province (ZMB
26768); S, Samar (ZMB 26767)) and J. dactylus (C, Cebu; ZMB 200109).
between both groups (Table 4). We performed a discrimi-
nance analyses that correctly classified 82.1% of the individ-
uals (Table 5), thus confirming the significance of the mean
differences between both groupings (Table 6).
F. Köhler & M. Glaubrecht • Jagora from the Philippines
Fig. 10 A–D. Shell colours. —A. J. asperata from Central Luzon,
Calauan, Laguna Province (ZMB 26766). —B. J. asperata from
South Luzon, Albay Province, Daragu River (ZMB 26768). —C.
J. asperata from Samar, Loquilocum (ZMB 26767). —D. J. dactylus
from Cebu, Matutinao River (ZMB 200109).
Fig. 11 Shell morphometries (H/LA). Box plot diagram showing
the median, the 25% and 75% percentile and largest nonextremes
(less than 1.5 times box height) of J. asperata (L1, North Luzon,
Isabella Prov. (ZMB 200114); L2, Central Luzon, Laguna Prov.
(AMS 324130); L3, South Luzon, Albay Prov (ZMB 26768); S,
Samar, Loquilocum (ZMB 26767)), and J. dactylus (B, Bohol, San
Isidro (AMS 324130); C, Cebu, Matutinao River (ZMB 200109); G,
Guimeras (syntypes BMNH 19990496)).
Molecular genetics
The sequence alignments were unproblematic and comprise
646 (COI) and 867 sites with indels at 27 positions (16S),
respectively. When possible, we sequenced COI fragments
from two or three specimens of each locality in order to
Jagora from the Philippines • F. Köhler & M. Glaubrecht

Table 4 Results of the variance analysis (one way-ANOVA). Table 5 Results of the discrimination analysis.

Shell parameter Measure Mean square F Sig. Predicted group membership

H between groups 44.88 0.777 0.380 Species J. asperata J. dactylus Total

within groups 57.78 – –
J. asperata 46 (80.7%) 11 (19.3%) 57 (100%)
B between groups 0.01 0.001 0.981
J. dactylus 6 (15.8%) 32 (84.2%) 38 (100%)
within groups 15.55 – –
LA between groups 7.96 2.553 0.113
within groups 3.12 – –
Table 6 Parameters describing the quality of the discrimination
WA between groups 4.83 2.445 0.121
within groups 1.98 – –
function.
BW between groups 7.77 0.871 0.353
No. of functions Wilks’ Lambda Chi-square d.f. Sig.
within groups 8.92 – –
H/B between groups 1.76 35.012 0.000 1 0.653 39.274 2 0.000
within groups 0.05 – –
H/LA between groups 1.76 35.012 0.000
within groups 0.05 – – have identical topologies. Furthermore, both NJ trees correspond
B/BW between groups 0.01 0.392 0.533
to the topology of the MP trees. Separate distance analyses
within groups 0.02 – –
H/BW between groups 0.31 29.977 0.000
(NJ, Kimura-2-parameter) were conducted for COI and 16S,
within groups 0.01 – – resulting in similar pairwise distances (Table 7). Accordingly,
BW/LA between groups 0.03 5.539 0.021 sequence distances are smaller between the populations on
within groups 0.00 – – Luzon (1.1% or below), while they are considerably larger
H, shell height; B, shell breadth; LA, length of aperature; WA, width of aperature; BW, between Luzon and Cebu, ranging from 13.5% to 14.7%.
body whorl. For the 16S sequences pairwise transition (s) and transversion
(v) rates were calculated (Fig. 14). Transition and transversion
rates are minor between the populations from Luzon, but are
estimate the variability within single populations. Owing to about tenfold higher (s ∼ 10%, v ∼ 1%) when comparing any
methodological difficulties, it was not possible to obtain COI population from Luzon with those from Cebu.
sequences from one population (Luzon, ZMB 200112).
Alignments were made for 10 COI sequences (six speci- Discussion
mens from three populations of J. asperata, three specimens Evaluation of morphological characters
of the only population of J. dactylus available, and a single The conchological variation of several freshwater snails confused
sequence of Brotia pagodula (Gould, 1847) as an outgroup earlier authors, misleading them into employing a vast number
representative) as well as for six 16S sequences (one specimen of species names. In some cases, this typological approach has
each of four populations of J. asperata, one population of caused a plethora of synonyms. Consequently, many names
J. dactylus, and one specimen of B. pagodula). The number of merely indicate phenotypic disparity rather then species diversity
variable sites is 161 (COI) and 244 (16S), respectively, while (see e.g. Glaubrecht 1993, 1996 for another cerithioidean taxon,
84 (COI) and 40 (16S) sites are parsimonious informative. Melanopsis). On the other hand, shell characters are often the
Two 16S sequences are identical: ZMB 200111 and ZMB most convenient to use for taxonomic purposes (see Reid 1986:
200115. Cladistic analyses were conducted using maximum 8). Consequently, a thorough assessment of the intraspecific
parsimony (MP) and neighbour joining (NJ). All trees were range of shell characteristics is necessary if using these features
rooted with Brotia pagodula as outgroup. The general topol- a priori as diagnostic criteria.
ogy of all MP and NJ trees is identical with the J. asperata Accordingly, the five nominal taxa from the Philippines
populations from Luzon and the J. dactylus population from were distinguished earlier in a typological approach. In contrast,
Cebu forming distinct clusters. The only most parsimonious our study reveals that all named taxa from Luzon and its
phylogram found for the COI sequences is shown in satellite islands Samar and Leyte, M. asperata, M. pagodulus,
Fig. 12A. For the 16S sequences three MP trees were yielded M. pulchra, and M. filocarinata, fall into the same morphos-
due to the unresolved relative position of ZMB 200115 and pace and neither of these phenotypes can be restricted to a
ZMB 200111, which are identical. The strict consensus tree certain geographical area or population. The high concho-
(Fig. 12B) matches the topology of the COI MP tree with the logical variation observed within and among these popula-
exception that an additional sequence (ZMB 200112) was tions suggests that the absence of some sculptural elements,
included in the analysis of 16S. such as ribs, spines and spiral ridges or the different degree
The trees resulting from a distance-based analysis (NJ) of of their development and the varying shell colour, indicate
the COI fragment (Fig. 13A) and the 16S fragment (Fig. 13B) intraspecific phenotypic plasticity.

Fig. 12 Maximum parsimony trees of Jagora.
—A. MP phylogram based on the analysis of
the partial COI sequences of J. asperata
from Luzon (ZMB 200111, 200113, 200115),
J. dactylus (ZMB 200109), and Brotia pagodula
(ZMB 200208) as an outgroup represen-
tative. —B. Strict consensus of three MP
cladograms based on the analysis of the 16S
fragments of J. asperata (ZMB 200111,
200112, 200113, 200115), J. dactylus (ZMB
200109), and Brotia pagodula (ZMB 200208).
For exact localities of the samples see
Material examined.
The shells from populations on Cebu are not only pheno-
typically distinguishable from those of the Luzon populations
(Fig. 9), but are also geographically isolated. Moreover, both
forms, viz. J. asperata and J. dactylus, can be discriminated by
means of minor but significant morphometrical differences
(Tables 4, 5, Fig. 11). In summary, Cebuan shells ( J. dactylus)
in mean are generally more slender and the mean number of
whorls is slightly higher (9.3 ± 1.3) than in shells from Luzon
and Samar (J. asperata) with 8.3 ± 1.2 whorls. The situation,
however, is complicated by the fact that the shells from the
islands of Samar and Bohol exhibit apparently intermediate
characters. Those from Samar cluster together with the Luzon
populations in respect to their morphometry, but lie interme-
diate between Luzon and Cebu in terms of the number of
ribs (Fig. 9). Meanwhile, specimens from Bohol cannot be
discriminated from J. asperata concerning shell parameters,
although they are very similar to J. dactylus from Cebu and
Guimeras. However, due to the low number of specimens
(n = 5) the latter statement is not statistically significant.
In contrast to the shell, differences in the soft body anat-
omy, including radular morphology (Figs 2 and 3), were not
found between individuals from different regions within the
F. Köhler & M. Glaubrecht • Jagora from the Philippines
archipelago. Several anatomical features, however, are pecu-
liarities common to both Jagora taxa. These features include
fusion of the inner crescent fold to the gastric pad of the
stomach, the entire anatomy of the pallial oviduct and the
retention of eggs in the mantle cavity in concert with the lack
of any brooding structure.
A report by Bandel & Riedel (1998), in which a subhaemo-
coelic brood pouch was mentioned as having been found in a
species tentatively identified as ‘Brotia aff. costula’ from the
Matutinao River, Cebu, stands in sharp contrast to all obser-
vations presented here. Apart from the incorrect species
identification, their finding of a brood pouch seems dubious.
According to the shell figured, their report refers without
doubt to J. dactylus. Our own observation supports the presence
of the latter species at the same locality. Therefore, we
assume that the authors either misidentified large specimens
of the superficially similar and syntopic Tarebia granifera
Lamarck, 1822 with specimens of J. dactylus, or that they
failed to recognize the reproductive peculiarities of Jagora
and that the juveniles in this taxon are actually incubated
within the mantle cavity instead of a brood pouch as found
otherwise, for example, in Thiaridae.
Jagora from the Philippines • F. Köhler & M. Glaubrecht

Fig. 13 Neighbour joining trees of Jagora.

—A. NJ tree based on the analysis of the
partial COI sequences of J. asperata from
Luzon (ZMB 200111, 200113, 200115),
J. dactylus (ZMB 200109), and Brotia
pagodula (ZMB 200208) as an outgroup
representative. —B. NJ trees based on the
analysis of the 16S fragments of J. asperata
(ZMB 200111, 200112, 200113, 200115),
J. dactylus (ZMB 200109), and Brotia
pagodula (ZMB 200208). For exact localities
of the samples see Material examined.

In contrast to Jagora, other South-east Asian pachychilids Table 7 Pairwise sequence distances between the populations.
such as Brotia and Tylomelania possess specific brooding
ZMB 200111 ZMB 200112 ZMB 200113 ZMB 200115
structures (for a detailed description of the reproductive
anatomy of Brotia see Köhler & Glaubrecht 2001). Anatom- ZMB 200109 COI 0.129–0.130 — 0.124–0.126 0.129–0.130
ical differences between Brotia and Jagora were also found in 16S 0.135 0.143 0.142 0.135
ZMB 200111 COI — — 0.009–0.014 0.000
the gonoduct anatomy, since some (but not all) Brotia species
16S — 0.011 0.006 0.000
possess a seminal receptacle which is situated dorsal to the ZMB 200112 COI — — — —
bursa. Jagora, however, lacks such a structure. Here, the 16S — — 0.010 0.011
prominent papillated ridge formed by the lateral lamina is ZMB 200113 COI — — — 0.009–0.014
supposed to have a sperm storing function instead (Figs 5D– 16S — — — 0.006

E and 6). This structure herein is termed seminal receptacle
as well, although having neither the same structural origin
nor a similar anatomy as the receptacle in Brotia. In addition,
size and location of the gonads (Fig. 4) differ between both
genera: In Brotia the gonads are much more conspicuous and
do not entirely surround the digestive gland. As holds true for
Brotia from the South-east Asian mainland, Sumatra, Borneo
and Java, clear differences of the reproductive anatomy exist
between Jagora and the pachychilids endemic to Sulawesi.
COI, 663-bp fragment of the Cytochrome c oxidase subunit I gene; 16S, 825 bp of 16S
rDNA (16S).
These taxa were previously subsumed under Brotia, but have
been shown to represent a separate clade characterized in
particular by a pallial oviduct which is modified into a
segmented brood pouch with a slit-like vaginal opening
(Rintelen & Glaubrecht 1999).

Fig. 14 Diagrams showing pairwise rates of
transitions (s) and transversions (v) of
sequences of J. asperata (Luzon), J. dactylus
(Cebu), and Brotia pagodula (Thailand) vs.
the sequence distances based on the
Kimura-2-parameters model. —A. 16S
sequences. —B. COI sequences.
Next to clear anatomical differences, Jagora and Brotia
share a peculiar modification of their ontogenesis: in the
early stages soft tissue protrudes from the apical whorl of the
forming juvenile shell (Fig. 7B–D). This tissue is believed to
have a nutritive function for the encapsulated embryo. The
so-called ‘soft apex’ has been reported for some, but not all,
Brotia species and has been correlated with the development
of a wrinkled apical shell (Morrison 1954; Solem 1966; details
in Köhler & Glaubrecht 2001). Accordingly, the wrinkles are
caused by the shrinking visceral mass of the embryo during
the retarded process of shell calcification. A similar storage
structure, a so-called hepatic lobe, was described for numerous
‘prosobranchs’ (Carrick 1938) functioning as a substructure
for the formation of the digestive gland (Fioroni & Schmekel
1976: 129ff ).
Systematic conclusions
The peculiar reproductive anatomy of both Jagora species is
believed to be synapomorphic and sets them apart from other
Pachychilidae, especially from Brotia and the endemic species
of Sulawesi. The distinctiveness of the Philippine taxa is
supported by characteristics of the stomach (fusion of the
inner crescent ridge to the gastric pad), the radula (convex
upper margin of the rachidian) and the reproductive biology
(egg capsules containing more than one egg). Molecular
genetic data from the COI and 16S sequences of South-east
Asian pachychilids (see Köhler et al. 2000; and unpubl. data)
support the monophyly of this Philippine clade with a rather
basal position. Thus, a generic status of the Philippine clade
independent from Brotia reflects its monophyly as well as its
evolutionary and morphological distinctiveness. A substan-
tial hypothesis on the phylogeny of the South-east Asian
Pachychilidae awaits cladistic analysis of further molecular
and morphological data.
We deliberately synonymize M. filocarinata, M. pagodulus,
M. philippinarum and M. pulchra with J. asperata, thus agreeing
F. Köhler & M. Glaubrecht • Jagora from the Philippines
with Knipper (1958) and (in part) with Brot (1874), as these
named forms cannot be unequivocally correlated with distinct
populations. These aforementioned morphospecies rather
delineate extremes of a single, albeit conchologically variable
species, instead of representing natural entities or biospecies.
Although the differences are not very pronounced, shells
of J. dactylus from Cebuan populations can be distinguished
from those of J. asperata. The morphological differences
between both taxa are, however, obviously restricted almost
exclusively to shell characteristics, as the soft body and
radular anatomy are identical. This also holds true for many
aspects of their ecology, except for the seasonal variation of
their reproductive activity (see below).
However, cladistic analyses of molecular data consistently
separate J. asperata from J. dactylus, and the NJ trees reveal
considerable genetic distances between both groups. This
provides additional evidence for the separation of two sibling
species in the Philippine archipelago. We anticipate that the
inclusion of more sequence data from additional localities
and islands will facilitate this analysis on the separation
between the taxa. Unfortunately, suitable material is not yet
available. Thus, in the absence of additional data from other
Visayan islands, it cannot be completely resolved whether
we deal with two closely related parapatric (in parts of their
distribution perhaps even sympatric) species, or only with a
single variable species.
Nevertheless, simply subsuming J. dactylus under J. asperata
would cover the existing discontinuity found in our study.
Therefore, we here refrain from synonymizing both taxa and
treat them as different biological entities instead until evidence
has surfaced in favour of an alternative hypothesis.
Reproductive biology and life history strategy
The placement of all species which were previously sub-
sumed under Brotia within Thiaridae (e.g. Morrison 1954;
Solem 1966; Davis 1971; but see Glaubrecht 1996 for critics
Jagora from the Philippines • F. Köhler & M. Glaubrecht
on the erroneous former concept of Thiaridae s.l.) has fuelled
speculation that parthenogenesis is the predominant mode of
reproduction in these cerithioideans (Benthem Jutting 1956;
Brandt 1968). However, as documented here, reproduction
in Jagora is bisexual and the sex ratio is constantly more or
less balanced (Table 2). Thus, we are able to correct our earlier
assumption, which was based on a smaller sample, that this
ratio may be skewed towards females (Köhler & Glaubrecht
2001). As, similar to several species of Brotia, a balanced sex
ratio was found in Jagora, our data do not support the
assumption that parthenogenesis is the predominant mode of
reproduction among (ovo)viviparous pachychilids of South-
east Asia.
The females of Jagora produce exceptionally large eggs
containing a considerable amount of yolk, and they are the
only representatives of South-east Asian pachychilids to
produce egg capsules containing more than a single egg.
Comparable egg sizes were reported from some other
freshwater gastropods, e.g. Brotia (Köhler & Glaubrecht
2001), Ampullaria and Viviparus (Fioroni & Schmekel 1976),
while most caenogastropods produce significantly smaller
eggs (see e.g. Fretter & Graham 1994; Fioroni & Schmekel
1976).
The average number of retained juveniles in Jagora is
assumed to correspond to the number of retained eggs (see
Table 3). In those cases where only a few juveniles were found
in a female, they always had fully developed and completed
shells and were ready to be released from the mantle cavity.
Thus, they are likely to be the last remaining members of
already released clutches. Similar numbers of eggs and
juveniles were also observed in many viviparous species
of Brotia (Abbott 1948; Davis 1971; Dudgeon 1989;
Kruatrachue et al. 1990; Köhler & Glaubrecht 2001) and
some viviparous Thiaridae, e.g. Melanoides tuberculata (Müller,
1774) (Abbott 1948; Starmühlner 1969, 1976; Glaubrecht
1996: 119) and Thiara scabra Müller, 1774 (Riech 1937: 45).
The considerable intraspecific variability of egg numbers
observed in all these taxa has been attributed to variable
habitat conditions, individual nutrition status and the
size or age of the females (e.g. Dudgeon 1989 for Brotia
hainanensis).
Despite the scarcity of data on the ecology and life history
of the two species of Jagora, some general conclusions can be
made:
1 We assume that the reproductive cycle of J. asperata is
synchronized with the seasonal changing monsoon climate.
The reproductive activity of this species is clearly restricted
to the rainy season in Luzon between June and December.
In Cebu, however, with a smaller monsoonal influence, rainfall
tends to be continuous throughout the year. Our observations
suggest that the reproductive activity of J. dactylus is not
restricted to a particular season.
2 In contrast to viviparous Thiaridae, where brooding
females possess secretory epithelia in the brood pouch in
order to nourish their embryos (Muley 1977; Glaubrecht
1996), in Jagora the growth of the retained embryos seems to
be sustained only by the large amount of yolk delivered with
the egg, since any nutritive structures of female tissue have
not been found. Evidence for the existence of nurse eggs or
feeding on retarded embryos (cannibalism), also reported to
be adaptations connected to ovoviviparity (Fretter 1984;
Fretter & Graham 1994), was not found. Otherwise, either
the number of eggs could be expected to be much higher than
observed, or the number of released juveniles should be
significantly lower.
3 The increased amount of yolk delivered by the egg is
correlated with a remarkable modification of shell formation
during ontogenesis. In general, shell formation in gastropods
starts at the ectodermal area of the embryonic visceral mass
(shell field) with the production of a conchiolin layer that
increases in diameter and remains uncalcified. Later, the shell
gland secretes primarily isolated calcium carbonate crystals
on this layer (Fretter & Graham 1994: 577; for ontogenesis
in some limnetic gastropods see Riedel 1993). We hypothesize
that in Jagora the agglutination of calcium carbonate crystals
in the periostracum is retarded and that the apical shell is
not formed before the large yolk sac has been consumed to
a considerable degree. The typical wrinkled apical shell
structure is believed to result in two overlapping processes:
shrinking of the visceral mass of the embryo and the
simultaneous agglutination of shell material. This phenomenon
has been extensively described by Riedel (1993) for the thiarid
Melanoides tuberculata and was connected to the typically wrinkled
apical shell known from several viviparous cerithioidean taxa,
such as Brotia, Melanoides, Tarebia H. & A. Adams, 1854,
Thiara, Lavigeria Bourguignat, 1888 and Potadomoides Leloup,
1953 (see Riedel 1993; Glaubrecht 1996; Köhler &
Glaubrecht 2001). However, thiarids generally lack a protruding
yolk sac as observed in Jagora and some species of Brotia.
Hence, it is the mode of embryonic nutrition via a large yolk
sac and not the wrinkled apical shell itself that is unique to
both pachychilid taxa.
4 The development of brooding structures in freshwater
gastropods, such as the brood pouches of Thiaridae and
Pachychilidae, has been viewed as a preadaptation for the
exploration of freshwater habitats by originally marine gastropod
groups (see Discussion in Glaubrecht 1996). However, among
the South-east Asian Pachychilidae we found three groups
with clearly distinct and relatively complex brooding structures
(Rintelen & Glaubrecht 1999; Köhler et al. 2000; Köhler &
Glaubrecht 2001) while other non-Oriental pachychilids,
such as Melanatria and Pachychilus, are oviparous. Preliminary
molecular genetic data suggests that the Neotropical
Pachychilus is in a more basal position than the brooding

taxa Jagora and Brotia (Köhler et al. 2000; and unpubl. data).
The hypothesis that brooding among limnic Cerithioidea
has evolved as a so-called ‘key innovation’ prior to the
colonization of freshwater habitats requires the assumption
that (ovo)viviparity subsequently has been lost in (then
secondarily) oviparous freshwater taxa. In contrast, the more
parsimonious explanation is that brooding has arisen a
posteriori to the habitat shift, and perhaps several times
independently, among freshwater cerithioidean taxa.
Consequently, the different brooding structures within
the South-east Asian Pachychilidae are not considered
homologous, and we assume a multiple origin of brooding
within this clade. Brooding seems to be strongly favoured by
selection in pachychilids, although to date we have no good
explanation for this. Given that the ability of the parent to
supply its offspring with nutrients is limited and correlated
with its own size and feeding status, the number of retained
eggs should be correlated with its size or comparable efforts
for embryonal nourishment (Calow 1978). Since the pro-
duction costs per egg are increased for the large amount
of nutritients delivered with it, each egg is most valuable
for the parent and loss due to predation, for example, is less
tolerable. Therefore, brooding and retention of developing
embryos may clearly increase the survival chance of each
offspring and, thus, the fitness of the parent, resulting in a
‘k-selective’ strategy. This scenario is only one of many realized
among freshwater gastropods in order to cope with the
challenges of their environment.
An alternative, rather ‘r-selective’ life history strategy is
pursued by some freshwater Thiaridae (for details see
Glaubrecht 1996). Although thiarid species also incubate
their juveniles in special brood-pouches, the free-swimming
larval stages have not been reduced in some taxa. Instead,
thousands of veligers are released from the brood pouch of
the parent. The planktotrophic larvae, however, usually develop
in a marine environment and are not exposed to the ‘harsh’
conditions of freshwater habitats. Due to their amphidro-
mous life cycle, these thiarids such as Stenomelania Fischer,
1885, are restricted to estuarine habitats and/or the lower
course of rivers where they rely on passive dispersal of their
marine veligers for the colonization of new habitats. In addi-
tion, the marine phase in their life cycle enables such species
to colonize even remote oceanic islands (Glaubrecht 1996,
2000; Myers et al. 2000a).
Zoogeographical implications
Members of the Pachychilidae exhibit a pantropical distribu-
tion indicative of a Gondwanan origin, and therefore pro-
bably represent an old Mesozoic lineage (Glaubrecht 2000).
The South-east Asian representatives of this family are
believed to form a monophyletic group. Although details of
their relationships are still insufficiently understood, both
F. Köhler & M. Glaubrecht • Jagora from the Philippines
the ancient history of the family and the monophyly of the
South-east Asian clade are supported by molecular data
(Köhler et al. 2000; and unpubl. data). As the dispersal
abilities of pachychilids seem to be limited, their recent
distribution is assumed to particularly reflect relatively
ancient vicariance events like the migration, separation or
amalgamation of terranes rather than younger events related
to the Pleistocene sea-level changes, while active dispersal is
believed to play a minor role if at all (Glaubrecht 2000;
Köhler & Glaubrecht 2001).
Accordingly, the geological history of the region since the
Cretaceous and during the Tertiary provides an important key
to the understanding of the biogeography of South-east Asian
Pachychilidae in general and the systematic position of Jagora
in particular. However, the palaeogeography of the Philippines
is complex and unfortunately not yet completely understood,
although considerable advances have been made recently in
particular in reconstructing regional tectonic events. Reviews
are presented in Hall & Blundell (1996; see also literature
therein), Hall (1996, 1998), Packham (1996) and Pubellier
et al. (1991) on which the following scenario is largely based.
It is generally accepted that the Philippine archipelago is
an assembly of several independent terranes or microplates
that have drifted to their present position since the Mesozoic
from various source areas and that most of the islands formed
part of an arc at the southern edge of the Philippine Sea plate
before the early Miocene (Hall 1996, 1998 and maps of
palaeogeographical reconstructions therein). Since the Miocene
the Philippine continental fragments have moved in a very
narrow zone, mainly as part of the Philippine Sea plate, resulting
from their general movement into and between other plates
and leading to the composite nature of the archipelago as we
know it today. In terms of their geology, the Philippines are
considered an ephemeral feature that ended up as an arc
terrane attached to the Eurasian continental margin.
The archipelago straddles the convergent boundary between
the Eurasian and Philippine Sea plates. Further to the east,
the geologically young Philippine Trench is situated and it is
generally agreed that much of the archipelago was developed
on the Philippine Sea plate rather the Eurasian margin.
While the eastern part substantially comprises rock built up
by island arc processes from the Cretaceous (Late Jurassic)
onwards, continental terranes lie mostly west of this line and
include Mindoro, west Panay and west Mindanao. The
terranes in western Mindanao include the Zamboanga
and Daguma Range and were juxtaposed before the early
Pliocene by strike-up movement. The continental blocs that
are parts of the Asian margin were incorporated by the long-
term clockwise rotation of the Philippine Sea plate terranes.
It is believed that this plate has rotated clockwise during the
past 50 Myr and that this process has brought the arc facing
south to south-west northward where it collides obliquely
Jagora from the Philippines • F. Köhler & M. Glaubrecht
with the Asian margin. During the Miocene this rotation may
have been initiated by the collision of the northern edge of
the Australian craton with the Philippine Sea plate (for the
general location of the Philippines see schematic reconstruc-
tions in Packham 1996: 139, fig. 13). In addition, in the
middle to late Miocene the Palawan Block on the southern
flank of the South China Sea collided with the Philippine arc.
Middle to upper Miocene volcanic rocks in western Mindanao
that find their southern extension through the Sangihe arc to
volcanic rock of similar age in eastern SW Sulawesi have been
interpreted (although this is not undisputed), as an east-facing
arc on the western side of the Molucca Sea.
Summarizing the above reconstruction, Mindoro, western
Panay, North Palawan and western Mindanao (Zamboanga)
are substantially continental terranes originating from the
Asian margin of the Eurasian plate, while the northern and
eastern parts of the Philippines with Luzon have emerged
from and rotated with the Philippine Sea plate to their
present position since the early Eocene (see Hall 1996;
Packham 1996). Unfortunately, as is also true for other areas,
it is unresolved at which times the islands emerged above sea
level and if they were permanently subaerial for continuous
periods during the Cenozoic.
Biogeographical evidence for the origin of the faunal and
floral elements of the Philippine biota has to date mainly
dealt with those patterns for which much later, essentially
post-Pliocene, events in connection with fluctuating sea-
levels during glacial periods can be attributed (Dickinson 1991;
Jong 1996; Heaney 1985, 1986, 1999). Thus, various parts of
the archipelago were isolated from the Asian mainland and/
or adjacent islands, such as Borneo, Sulawesi and Taiwan, due
to earlier tectonic events as well as later changing sea-levels.
Accordingly, its faunal and many floral elements could have
reached it at different times and in various ways: (1) drifting
in on individual continental terranes (2) active dispersal,
using existing land-bridges during low sea-level stands or
(3) passive and random over-water dispersal.
Without doubt, many Recent elements of the Sundaland
flora and fauna dispersed within the Philippines during the
Pleistocene, about 135 000 years ago, when sea level fluctua-
tions provided the most recent opportunity for a simplified
overland faunal exchange throughout South-east Asia
(Dickinson 1991). During this time, the Philippines were
mainly colonized from Borneo via Palawan or the Sulu
Archipelago and, to a lesser degree, from Sulawesi via the
Sulu sea (Heaney 1985, 1986; Dickinson 1991; Jong 1996). In
contrast to the mobile taxa, the freshwater snails currently being
studied in various islands in Wallacea, in particular Sulawesi
and the Philippines, may have an early origin (see Köhler &
Glaubrecht 2001). We suggest that the Recent distributional
patterns of South-east Asian pachychilids were more likely
caused by the above stated palaeogeographical events during
the earlier Cenozoic than by Pleistocene events in connection
with glacial periods, as found for many vertebrate taxa.
Molecular data provide a suitable tool not only for inferring
phylogenetic relationships but also to estimate a lineage’s age,
even when the latter is controversial (review in Nei et al. 2000).
The assumption of a molecular clock requires constant muta-
tion rates and a calibration with fossil or palaeogeographical
data. Our NJ trees (Fig. 13A–B) suggest almost constant
mutation rates within Jagora, although a calibration cannot
be conducted here. However, for the 16S gene in Littorina
rates of 1.5% transitions and 0.13% transversions per Myr
were found (Reid et al. 1996: 887, Fig. 5). Under the assump-
tion of similar substitution rates in Jagora, the populations
from Luzon (J. asperata) and Cebu (J. dactylus) would have
been isolated from each other 6.6–7.7 Myr ago. This might
only provide a very rough age estimate, since mutational rates
could be very different in Jagora, for example, due to varying
effective population sizes, and different metabolism and
mutation rates. Nevertheless, it is consistent with our sugges-
tion that tentatively hypothesize Jagora as representing an
ancient element of the Philippine fauna. Its ancestor might
have reached the archipelago on one of those continental
terranes that have since the Mesozoic drifted away from
the northern margin of Gondwana to subsequently build the
marginal rim of South-east Asia long before Pleistocene
events shaped the distributional pattern found, for example,
predominantly in the vertebrate fauna. Accordingly, we
anticipate that dispersal during periods of low sea-level stands
would have played only a minor role and is less important
for the distribution patterns found today in the Philippine
pachychilids.
Irrespective of the arrival time of a potential ancestor of
Jagora in the Philippines, it is reasonable to assume that it
originated with faunal elements occurring in areas that later
formed Borneo or (West-) Sulawesi (for the composite nature
of Sulawesi see Hall 1996, 1998). This route implies that
Palawan, Mindanao or the Sulu Archipelago have, at least for
some time span, been within the distributional area of the
ancestor of Jagora and its closest relatives among the pachy-
chilids of the genus Brotia from the Asian mainland and
Sulawesi, respectively. The only known pachychilid from this
area is ‘Melania soolooensis’ Reeve, 1859 from the Sulu Islands.
Since only a few dry shells of the species are available, any
potential affinity to Jagora remains speculative. Alternatively,
an ancestral species originating from the South-east Asian
mainland, moving via emergent islands that drifted to their
present positions as part of Luzon, Panay or Mindanao is
possible, as suggested by De Jong (1996) for butterflies.
Any substantive discussion of the historical biogeography
of Jagora, has, however, to await the identification of the
sister taxon in the course of a phylogenetic analysis of the
South-east Asian pachychilids in general. A more comprehensive

analysis of morphological and molecular data is urgently
required in order to establish a more reliable basis for the
calibration and testing of a molecular clock, and provide a
more detailed biogeographical scenario.
Acknowledgements
We are most grateful to the curators P. Bouchet (MNHN),
Y. Finet (MHNG), B. Hausdorf (ZMH), R. Janssen (SMF),
E. Kuster-Wendenburg (ÜMB), T. Meier (ZMZ), W. Ponder
and I. Loch (AMS), and K. Way (BMNH), who provided the
types and other material of their collections. We especially
thank Y. Finet and D. Reid (BMNH) for indispensable advice
in tracing literature and material, and A. Sumalde from the
Museum of Natural History, Los Baños, for offering support
in the field in the Philippines. S. Schütt was a pleasant field
companion and also helped with laboratory work. W. Mey
(ZMB) kindly provided some material from Luzon, which
facilitated this study. We are indebted to A. Meyer (University
Konstanz) for logistical help, to A. Wilson for providing
primers, and to T. vs. Rintelen for his help with molecular
genetic work and analysis and for his permission to use some
additional sequence data. B. Geist helped with histology and
illustrations. C. Ratton (MNHG), P. Lozouet (MNHN), V.
Heinrich (ZMB), and the BMNH kindly provided photo-
graphs. Furthermore, we thank two anonymous referees
for their helpful comments on the manuscript. The work of
the first author was funded by a postgraduate scholarship
of the Konrad-Adenauer-Stiftung (Germany). Visits to the
malacological collections of the BMNH and the MNHN
were funded by the BIORESOURCE- and PARSYST-
programme, respectively, of the EU. The project was sup-
ported through a grant to the second author by the Deutsche
Forschungsgemeinschaft (GL 297/4).
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