Impacto Presas PDF

Contributing Paper
Molluscan Biodiversity and the

Impact of Large Dams
Mary Seddon
National Museum and Galleries of Wales, UK
Co-chair IUCN Mollusc Specialist Group
Prepared for Thematic Review II.1:

Dams, ecosystem functions and environmental restoration
For further information see http://www.dams.org/
This is one of 126 contributing papers to the World Commission on Dams. It reflects solely the views
of its authors. The views, conclusions, and recommendations are not intended to represent the views of
the Commission. The views of the Commission are laid out in the Commission's final report "Dams and
Development: A New Framework for Decision-Making".
Report to IUCN Species Survival Commission
Molluscan Biodiversity
and the impact of large dams
Prepared by Dr Mary.B. Seddon
National Museum & Galleries of Wales, Cardiff, UK
Co-Chair of IUCN SSC Mollusc Specialist Group
The following have provided data and helpful comments on the
status of the freshwater faunas.
This includes access to unpublished manuscripts which are in press
or in preparation.
Arthur Bogan (North Carolina State Museum), Philippe Bouchet

(Museum dHistoire Naturelles, Paris), David Brown (Natural
History Museum, London), Kevin Cumings (Illinois Natural History
Survey), Olivier Gargimony (Museum dHistoire Naturelles, Paris),
Dai Herbert (Natal Museum, South Africa), Cristian Ituarte (Museo
de la Plata, Argentina), Thomas Kristiansen (Danish Bilharzia
Laboratory, Copenhagen), Richard Neves (US Fish & Wildlife
Service), Winston Ponder (Australian National Museum) Willem
Sirgel (University, South Africa), K. Walker.
Introduction
The phylum Mollusca is currently the second most diverse animal group, with origins which date
before the Cambrian. The phylum includes Snails (Order Gastropoda), clams and mussels
(Order Bivalvia), but also less obvious animals such as slugs and sea-slugs (which have an
internal shell), and octopus and squid (Order Cephalapoda), as well as some smaller orders
which not known to the general public.
1.0. GEOGRAPHIC DISPOSITION OF BIODIVERSITY
A. HOW MANY MOLLUSCS ARE THERE?
There are various estimates of the number of species, with Boss (1973) estimating the number of
species as 80,000, whereas van Bruggen (1995) estimated that the total number of species will
reach 135,000 species. The variation relates to the interpretation and extrapolation of the level of
undescribed species that are currently being published each year.
Table A1 Summary Statistics on Molluscan Biodiversity

Sources: Boss (1973), van Bruggen (1995)
Species
Notes
Marine
31,000 - 100,000
Boss (1973), van Bruggen (1995)
Terrestrial
14,000-35,000
Freshwater
5,000-5,000
Riverine
4000
(Excluding Spring-snails)
Importance of Molluscs to Ecosystem

The biomass of the Mollusca is very important to ecosystems (Summary in van Bruggen, 1995).
In marine environments, molluscs may form the dominant group, especially in their larval stage,
but also on soft substrates (bivalves), and on hard substrates, where some molluscs such as Pearl
oysters (Pinctada) and mussels may be more important in terms of their biomass than corals
(Salvat, 1970). In freshwater environments biomass of molluscs is significant in some
enviroments: for example in temperate countries lake bottoms maybe covered with the small
bivalves from families Sphaeriidae and Pisidiidae. Berrie & Boize (19xx) estimate that 80% of the
biomass of benthic invertebrates at River Thames (at Reading) is composed of freshwater Unioid
mussels.
In tropical areas, the same can be true. Brown (1994) summarises much of the data available on
contribution of molluscs to the biomass of freshwater systems in Africa, and shows that
prosobranchs contribute the major part in Lakes, such as Chad, Malawi, Tanganyika and Victoria.
He cites examples such as Hart (1979; table 6) observations that the species from two
prosobranch genera Bellamya (e.g. Bellamya capillata 0.40 g/m-2) and Melanoides (e.g. Melanoides
tuberculata 1.74 g/m-2) provide a major part of the biomass of Lake Sibaya.
Molluscs in the man-made Lake Kariba made up nearly the entire biomass of the benthic animals
(Prosobranchs 4.1 % and Bivalves 95.8%) (Machena & Kautsky, 1988) and Brown uses their data
to calculate figures of two prosobranch genera Bellamya (e.g. Bellamya capillata 0.28 g/m-2) and
Melanoides (e.g. Melanoides tuberculata 4.08 g/m-2). The latter species, Melanoides tuberculata, is
adapted to maintaining large populations in stable environments with a long mean generation
time and a low intrinsic rate of natural increase (see Pointer et al. 1991; Brown, 1994).
SPECIES AND GENETIC DIVERSITY
Molluscs are characterised by a soft body, usually with an exoskeleton in the form of a shell,
consisting of a body, a single muscular foot, a dorsal mass with a visceral sac, fleshy skin fold (the
mantle, which is contains the glands which form the external shell). The alimentary canal
contains a scraping tongue, the radula, which can be used as a character in identification. The
freshwater fauna can be divided into three main groups:
Freshwater Gastropods (Prosobranchia)
Freshwater Gastropods (Pulmonata)
Freshwater Bivalves (Bivalvia).
The greatest freshwater species diversity and endemism is found within the Prosobranchia and
the Bivalvia groups.
Freshwater prosobranchs
These range in size from large prosobranchs such as the apple-snails of the Ampullaridae (up to
40 mm in size) to the small springs-snails (1-8 mm).
The Afrotropical fauna differs from that of India and SE Asia in its greater richness of species in
the families Ampullaridae, Bithyniidae, Thiaridae and Planorbidae (Brown 1994). Brown notes
the lack of any Hydrobioid radiation in the Afrotropical region, contrasting it with the radiations
reported for the Mekang River (Brandt, 1974; Davis, 1982). Davies (1982) suggests that this is due
to the lack of limestone regions, but Brown (1984) notes that there are suitable perenical river
systems, but the water tends to be low.
The origins of these fauna evolved when the area was part of Pangaea (which broke up 150
million years ago) and then Gondwanland which Davis (1982) suggests accounts for the presence
of some snail groups in Africa, as well as South America and SE Asia.
These groups can be summarised as :
1) South America, Africa, India/Asia.
Family
Pomatiopsidae
Ampullariidae
Thiaridae (Genera Potadoma, Pachychilae and Brota)
2) Africa India SE Asia.
Family
Viviparidae (especially Genus Bellamya)
Thiaridae (Genus Thiara, Melanoidea)
Planorbidae (Subfamily Bulininae)
Brown (1994) attributes the success of families such as the Ampullaridae to their liking for
habitats such as swamps and muddy slow flowing rivers. The genus endemic to Africa, Lanistes,
shows that some adaptations may be related to their predators (fish), the sandy substrate for
feeding and life in deep water (Brown 1994).
The Hydrobiidae are a diverse group of prosobranchs, occuring in lakes, springs, seeps, marshes
and lotic waters. These species are often found in very high densities, and thus can provide a
high component of the benthic epifauna. Some of the genera (e.g. Fontigens) exist in subterranean
waters and springs (Hersler et al 1990), and those species which have highly isolated and
geographically separated populations are currently being studied to determine their phylogentic
relationships.
Freshwater pulmonates
Freshwater pulmonates can be found in all microhabitats in freshwater systems. The group
exhibits the least levels of diversity in most freshwater systems. The freshwater pulmonates are
thought to be less diverse in Africa due to their need to fill the lung with atmospheric air,
reducing their ability to invade deep-water biotypes in lakes (Brown, 1994). These taxa are
excellent colonists, as they are often able to live at high densities in shallow, even seasonal, water
bodies, and as self-fertilising hermaphrodites they are able to disperse and colonise new water
bodies.
In terms of the life-cycle, the species are usually are oviparous, laying eggs capsules of different
types according to family. The number of eggs per capsule varies from one (e.g. the limpet
groups Ferrissia) to over one hundred (e.g. Lymanea natalensis) (Brown, 1994). Water temperature
is generally an important influence on growth rate, although other factors, such as changes in
biotope, or food shortages can also have an impact on pulmonate growth rates (Brown, 1994).
Freshwater bivalves
In order to understand the potential impacts that river channels changes and impoundments have
on bivalves it is useful to know more about the life cycle strategies of these groups. Bivalves,
other then freshwater mussels, typically produce either free-swimming larva (trochophere or
veliger) or have direct development releasing juveniles. The freshwater mussels (Unionoidea)
have reproductive strategy involving a larval stage (called the Glochidia), which is retained in the
female brood pouch or gills and released for their intermediate stage as a parasite of a host fish
before being transformed to bottom dwelling juveniles (or in one case in the USA an aquatic
salamander: the mudpuppy, Necturus sp. ; Bogan, 1998; Watters, 1994). A few US species
deviate from this system, for example Obliquaria reflexa, which can bypass the parasitic phase
(Howells et al., 1996). Most freshwater mussels have separate sexes, although some species are
known to be hermaphrodite.
Spawning, egg deposition and incubation.
Male discharge sperm direct into the water. Females inhale waterborne sperm during normal
feeding and respiration, and egg fertilisation takes place. The fertilised eggs and developing
glochidia are retained in brood pouches in the gills (not all gills have brood pouches, this varies
from species to species).
In the US mussels, long term (Bradytictic) breeders spawn and fertilise eggs in late spring,
summer or early fall producing mature glochidia by late fall or winter, however, the glochidia
may not be released until spring or early summer of the following year. In contrast short term
(Tachyticitic) breeders spawn, fertilise eggs, develop and release glochidia from late spring to
early fall.
Glochidia may be released singly or in groups. Some species discharge conglutinates which
mimic warm, grubs or fish; and thus are consumed by the host fish. The glochidia cannot swim
but must drift with water currents. If they do not find a suitable host fish within a few days they
will die. Similarly if they attach to a none-host species or are incorrectly on a host species, again
within a few days they will die (Howells et al., 1996).
The infection on the host-fish is usually light, and encysted glochidia row little during the
parasitic stage, hence the impact on the fish is minimal. Released juveniles are often similar in size
to the glochidia. Consequently, on release, if they do not become lodged in a suitable substrate
they will not survive.
Feeding
Freshwater mussels are filter feeders requiring a rich and plentiful supply of diatoms, desmids,
filamentous algae and other algal species.
Factors influencing vulnerability
Freshwater mussels are especially vulnerable to habitat disturbance. The mussels of Unionoidea
with their obligate parasitic state, requiring host fish are especially threatened. As at present, only
about 25% of the host fish for the mussels in the USA have been correctly identified, it is more
difficult to predict the impact on damming, as clearly the movement of fish within the river
systems will affect the reproductive success of these mollusc species.
Many species in USA and Europe have extended life cycles, some of which span over 100 years,
where maturity is delayed until 6 15 years (Bauer, 1993; Chesney & Oliver 1998). Some species
also have reduced powers of dispersal, high juvenile mortality and long turnover times.
The prolonged lifecycle of the freshwater mussels does mean that populations may appear
secure, when in fact no active recruitment is taking place, and as such these populations may well
end up being functionally extinct. For those species which have few extant populations, it results
in species (e.g. ) which are Functionally extinct. Chesney & Oliver (1998) have a figure which
illustrates this point well (Figure x.x)
Figure xx Hypothetical size distributions for recruiting and senescent

populations of Margaritifera margaritifera
TABLE XX
Environmental and biotic factors influencing the various stages of the life cycle of Unioid
Mussels (amended from Chesney & Oliver, 1998). Items highlighted in bold are directly
impacted after impoundment; in italics, indirect impacts of impoundments.
FACTOR
Exploitation (pearl fishing, shells for
buttons/seeding)
Fish host population size
Mussel population size
River Bed
LIFE CYCLE STAGES AFFECTED

Adults
Flow regime
Fertilisation, glochidial infection, settlement,

juvenile and adult
Adults, feeding and brooding
Adults, juveniles
Adults, juveniles
Adults, juveniles
All stages
Adults, juveniles
Adults, juveniles
Adults, juveniles
Juveniles
Juveniles
All stages
All stages
Suspended solids
Eutrophication
Nitrogen
Phosphate
Dissolved Oxygen
Conductivity
Calcium
PH
Interstitial particulates
Interstitial water chemistry
Industrial pollutants
Pesticides
Glochidium
Fertilisation, glochidium numbers
Adults, juveniles
Chesney & Oliver (1998) point out that it is important to understand the life cycle of European
freshwater pearl mussel species and to adopt survey, monitoring and management protocols for
all phases. Table 1 has been adapted from Chesney & Oliver (1998) who illustrated the wide
ranging interactions of life cycle stages to environmental parameters for M. margaritifera. In much
of the early survey work in the 1970s in Europe the impact surveys tended focus on the adult
stage which, in pearl mussels, is the most resistant phase to environmental change.
Again, in most handbooks, the habitat preferences are often simplified and for example Chesney
& Oliver (1998) cited the preferences for the Freshwater Pearl Mussel which is often quoted in
gravel patches behind rocks in riffles, mostly in oligotrophic upland rivers. They noted that this
is a gross simplification of where M. margaritifera actually lives, as the species can be found in fast
flowing lowland rivers and in uniform gravel beds in water depths of over a metre and is
sometimes tolerant of calcium rich water ( e.g. Ireland and the Lake District).
In many Unionid bivalves, failure to recruit viable juveniles to populations appears to be the
major obstacle to species recovery (Chesney & Oliver 1998). These authors list several problem
areas for recovery, and all these conditions may exist in the post-impoundment phase:
eutrophication effects are crucial for post-glochidial juveniles, especially on their
interstitial environment requirements.
nutrification, especially raised nitrate and phosphate levels, may have a more
deleterious effect on juveniles.
clogging of the interstitial spaces by organic debris may reduce oxygen as well
preventing access to the sediment.
Correcting these problems will involve:
Catchment management and river restoration
Reviewing and changing agricultural practices to reduce eutrophication.
These will involve raising public (=involved organisations) awareness.
Tackling these problems requires some degree of total river management which is both
contentious and problematic as it conflicts with current practice in agriculture, forestry and waste
management.
Should environmental problems prove impractical to correct in the short term, then many rivers
will lose their remaining senescent populations before recruitment can be restarted. At a future
date such rivers may once again be suitable, but can only be stocked by translocation, a procedure
which is currently not successful. Alternatively if we could captive breed the remaining stock
through to the post juvenile stage we could maintain the adult population and create a larger
time period in which the juvenile environment could be re-established. Currently it is a simple
process to infect young salmonids with glochidia but the subsequent growth through to five years
of age has never been completed with any great success.
It must be borne in mind that in-situ conservation which preserves genetic identity and leads to
recognition of local adaptations is preferable to conservation through translocation as this affects
the gene pool and should only be used in extreme circumstances. Liu, Mitton & Herrmann (1996)
concluded this when evaluating the restocking of the giant floater (Pyganodon grandis) in the
Colorado basin; they concluded that the dgreee of differentiation among drainages was so great
as to require new populations to be founded with individuals from the natuiraql populations
within the same drainage basin.
Key notes
Environment recovery will be complex and may take many years.
Species recovery may rely on re-introduction and captive breeding.
Considerable research and funding needs to be directed at the problems of
translocation and captive breeding.
An overview of Threatened Species
Seddon (1998) gave an overview of the Molluscs included in the 1996 Red List of Threatened
Animals (Baillie & Groombridge, 1996).
The first review of threatened invertebrates was carried out by Collins et al. (1983) and this
highlighted the threatened molluscs of specific regions such as islands e.g. Hawaii, French
Polynesia and Madeira, freshwater species from USA and Lake Baikal as well as the exploited
species such as Giant Clams (Tridacnidae). Global lists have given the status of endangered
molluscs since 1984 through the work of the IUCN Species Survival Commission Mollusc
Specialist Group (IUCN 1987).
Evaluation of freshwater Molluscs in river systems has been a problem using the new criteria as it
can be difficult to establish the range data for linear river systems. Estimating status of riffle
species, for example, as it is extremely difficult to estimate area of occupancy where species occur
in small riffles all the way along the river. As the data on extent of occurrence (i.e. river) are often
difficult to obtain and populations are difficult to assess, a problem which applies to many US
and European freshwater species, some species which were listed in 1994 were not listed in 1996,
except as Data Deficient.
Molluscan extinctions is a significant problem, as the diagram below shows:
Figure 2 Diagram showing the proportion of molluscan extinctions as a total of

all animal extinctions since 1600 AD.
Freshwater Species which are Threatened
Many of the US freshwater species were evaluated on the basis of population decline rather than
range data. Range data provide evidence for assessment of the spring-snails as Critically
Endangered; where there are molluscs with populations in only one known locality (B1). The
spring snails of Austria, Australia and the USA fall into this CE category when there is a threat of
over-abstraction of water from the wells or pollution of the water source (B2a, c). It is likely that
as more endemic spring snails are properly assessed a high proportion of these taxa are likely to
be threatened, given the degree of exploitation of their habitats. Some of these spring snails are
also subject to natural fluctuations (B3) in their populations (Lepitki 1997) consequently a
catastrophic event at a low point in their population cycle increases the possibility of extinction.
Many African freshwater molluscs were included in the 1996 Red List for the first time, as
information on their status is becoming more accessible. These species are threatened by decline
in quality of water, pollution, damming and increased sediment load. Given the generation time
of these species they are accessed at the category Vulnerable B1 and 2c. Freshwater spring snails
frequently fall into the Vulnerable D2 class on the basis of small range.
1.1. RIVER ORIGIN TO MOUTH
Freshwater ecosystems have a range of molluscan habitats within rivers, as well as adjacent
habitats where species rely on the flow of water through the catchment. In general a river is a
one-way system, as many species of mollusc can only move downstream by drifting or being
dislodged by flood events and moved downstream. In some stages of their life cycle those
species which have a larval form can move significant distances upstream with the aid of a third
party (e.g. host fish during a larval stage). Gregarious migrations upstream have been reported
rarely for molluscs, however, Schneider & Frost (1986) did observe that the freshwater neritid
snail Neritina (Clypeolum) latissima was capable of moving upstream by crawling along the bed
of Rio Claro in Costa Rica in groups when dislodged by flood waters (although the water levels
when this was observed were low).
Hynes (1976) showed that in a short clean river there is an increase in invertebrate species
diversity downstream, with increasing numbers of microhabitats, increased eutropification
enhancing productivity and increased sediment imput. More recently specific work on molluscan
diveristy downstream has been attempted. Mouthan (1999) presents the results of a survey of 272
stations in rivers of Franche Cent, France. He found that there were nine different
malacotypes, which could be identified, from the total of 52 species recorded in the survey. He
recognises that there were taxonomic insufficiencies in this work, especially in the spring snail
family Hydrobiidae.
He found that there was a general increase in the number of taxa along the river system, with few
species at the headwaters (Hydrobiidae spp. not separated) to 48 species in the lower reaches of
the river. It must be noted though that there is a decline in the number of species (39) at the
lowest point on the river system. Mouthan (1999) discusses the reasons for this, suggesting that
this may not be a true reflection of the original levels of diversity, but may be related to the
sampling sites. One of the rivers sampled in this study has a series of locks and conurbations
along the course. Six of the species which are not present at this level on the river system are
susceptible to pollution. The lower levels of dissolved oxygen during the summer may also limit
these six species, as well as Pisidium obtusale; thus Mouthan (1999) concludes that the decline in
the lower levels is due to anthrogenic activity including hydropower developments. Mouthon
also noted that the in the areas of highest species richness, a significant number of species also
had their highest abundance in this malacozone.
Mouthan (1999) noted that gastropods (such as the Planorbidae) which favour lentic habitats
with rich vegetation, reach their maximum species diversity and abundance at malacozone 6 on
this hypothetical river, which he again attributes to declining quality of habitats in the lower
reaches of the river.
Valovirta (1998) shows that the density and distribution of mussels along a river can change,
especially on old log floating rivers in Finland. This pattern is also reflected for most other
freshwater species, and as such bed formation and sediments are very important for the large
Unioinid mussels.
Molluscs can be divided into different groups on the basis of their habitat requirements:
Species which are generalists, found in all sizes of water bodies, in all types of water
conditions.
Specialist species requiring permanent water, with specialised microhabitats and/or
specific water conditions
Specialist species requiring temporary water bodies; these species are more
opportunists and tend to be good colonists.
Species that live in the riparian habitats besides large water bodies, requiring a
constant input of water into the habitat. Such species may be marsh dwelling species
(e.g. Vertigo moulinsiana), gallery forest species adjacent to the river.
Species that are spring specialists fed by underground aquifers. These include the
family Hydrobiidae.
In addition the different lifestyle and reproductive strategies that these species adopt influence
the diversity along the river system from origin to mouth.
1.2. Latitudinal gradients
Van Bruggen (1995) demonstrated that latitudinal variability exists in freshwater molluscs in
Africa. Using data from Brown (1980) he shows that the highest numbers of species being found
in the tropics (105 species at 0 - 5S) and lower numbers found in the freshwater systems of north
Africa (5-20 species from desert to mediterranean systems) and southern Africa (38-40 species) (
see Figure 1.2.1).
However, overlying this trend there are some notable deviations at hotspots of endemism and
diversity, where some temperate areas with specialist species swarms have much higher
levels of diversity.
[insert diagram Figure 1.2.1
Picture of Africa, with species diversity across Africa]

1.3. Continent
The highest levels of diversity and endemism within the freshwater molluscan fauna, as presently
recognised, lie in North America (USA: 945 species; Bogan, 1998). Other notable centres of
endemism are found in the rivers of Indo-China, such as the Mekong River and in freshwater
lakes where there have been speciose radiations, such as Lake Baikal (Siberia), Lake Tanganyika
(Africa) and Lake Biwa (Japan) (See below section 1.4). However, these comments reflect the
present state of knowledge, which may well change as our levels of knowledge about
distributions and the taxonomic insufficiency for molluscs in riverine systems in S. America and
SE Asia is improved.
In north America much of the speciation lies in the families Pleuroceridae (Gastropoda) and
Unionidae (Bivalvia). A high proportion of small range endemism lies within the specialist spring
snail groups (Family Hydrobiidae) in Europe, USA (c. 20% of mollusc fauna: see below) and
Australia , and the initial survey work in Sulawesi has shown there are a large number of
undescribed species within this group (Bouchet, 1995; Haas pers. comm). Undoubtly further
work in the less surveyed areas will reveal other areas of high diversity and endemism within
these spring snails.
1.4. Global species hotspots and endemism
(based on Hotspots of freshwater molluscan diversity prepared by Olivier Gargominy, Arthur Bogan,
Philippe Bouchet, Winston Ponder: draft discussion paper from Mollusc Specialist Group)
1.4.1 USA
The current enumeration of about 601 taxa of freshwater gastropods (table 1) and 300 taxa of
unionoid bivalves (Table 2) is based on Turgeon et al. (1988) and their subsequently revised
version (Turgeon et al ,1999?). In the USA freshwater ecosystems are disproportionately
endangered relative to other groups (Master, 1990) with molluscs being the most highly
threatened group within the freshwater organisms.
Table 1
Freshwater molluscan diversity in North America (after Bogan, 1998)
No. of Families
No. of Genera
No. of Species
Gastropoda
14
88
601
Bivalvia
5
57
344
Total
19
145
945
Table 2
North American freshwater bivalve diversity (After Bogan, 1998)
Family
Genera
Species
Margaritiferidae
2
5
Unionidae
48
295
Dreissenidae
2
3
Corbiculidae
1
2
Sphaeriidae
Total
4
57
39
344
Table 3
Table 3
North American gastropod diversity (after Bogan, 1998)
Family
Genera Species
Acroloxidae
1
1
Ampulariidae 2
4
Ancylidae
4
13
Bithyniidae
1
1
Hydrobiidae
37
228
Lymnaeidae
10
58
Neritidae
1
1
Physidae
4
43
Planorbidae
12
47
Pleuroceridae 7
156
Pomatiopsidae 1
6
Thiaridae
2
3
Valvatidae
1
11
Viviparidae
5
29
Total
88
601
1.4.1.1 USA- RIVERS

The extinction crisis for the US freshwater mussels was recognised last century. Higgins (1858)
observed declines in the Unionoid species which he suggested related to deforestation. Rhoads
(1899) remarked on the decimation of freshwater bivalves in the lower Monongahela River above
Pittsburgh which he believed was due to pollution. Ortmann (1909) reported declines on the
Tennessee River systems due to papermill effluents. However documentation of the extinction
crisis did not really gain momentum until the 1970s (Athhearn, 1970; Heard, 1970; Stansberry,
1970; Taylor, 1970). Bogan (1998) summarises the state of knowledge on the extinctions of the
freshwater molluscan fauna in the USA.
Table 4 Threatened Status of North America Unionoid Bivalves
(after Bogan 1998)
No. of Taxa
No. of Candidate taxa
No. of Threatened taxa
No. of Endangered taxa
No. of Extinct taxa
Total
300
61
5
57
35
Percent
20.33
1.66
19.00
11.66
Table 5 Threatened Status of North American freshwater gastropods

(after Bogan ,1998)
No. of Taxa
No. of Candidate taxa
No. of Threatened taxa
No. of Endangered taxa
No. of Extinct taxa
Total
601
173
0
9
42
Percent
28.8
0
1.5
6.99
The greatest diversity in the regional freshwater faunas are reported for Mobile Bay in Alabama
and the Tennessee River Basin and these are also some of the most threatened faunas. For
example with 7 % of total taxa of prosobranch gastropod fauna found in the Mobile Bay River
basin, and the Tennessee River Basin, now extinct (Bogan, 1998). Most of the extinctions in this
group (38 out of 42 taxa) in the Mobile Bay fauna, occurred when the big river shoal fauna was
impounded and covered by deep standing water and subsequent increased siltation.
Ohio-Tennessee River Basin
In the Tennessee River Basin species are under threat following the construction of dams and the
subsequent regulation of flow. A number of gastropods from the family Pleuroceridae are under
threat, as they persist on clean swept shoal areas below dams on the river (Bogan, 1998), and
three other species have been extirpated from the river Haag & Thorp, J.H. (1991). These authors
also reported drop of 50% in number of freshwater mussels from c.100 species that were present
prior to the multiple impoundments, but noted that the presence of the. Asian Freshwater Clam
(Corbicula fluminalis) had also had an impact reducing the density of native mussels in the areas
where they remained. In terms of the general molluscan community, Haag & Thorp observed
lateral channel changes and seasonal changes in the gastropod and bivalve abundances, which
they believed reflected differences in breeding cycle and reproductive behaviour. These lateral
changes across the channel, relate to the bed type and water depth, for example, typical densities
of the Pleurocids ranged from 947 m2 (4.7m water depth) to 1011.8 m2 (7.6m water depth).
The Cumberland River, in the upper part of the Ohio-Tennessee River basin had a series of
impoundments and locks establised between 1916 and 1923 (Blaock & Sieckel, 1996). Over 85
freshwater mussel species were known on this river system, prior to impoundment. In the reach
of the river directly under Lake Barkely there were 45 species, and in the reach under Kentucky
25 species known. Blaock & Sieckel (1996) point out that these esimates are minumums for each
reach of river, as mussel distributions are very patchy, and sampling few sites (7 19) may mean
species were not recorded. Blaock & Sieckel (1996) describe changes in species diversity and
species abundance. For example although mussel abundance shown a loss of 50% between 1911
and 1994, over 65% of the shells were a single species Quadrula quadrula. Table below shows the
number of species lost in the the Kentucky portion of the lower Cumberland river, which wass
inudated by Lake Barkeley. Most of these species were ones that were recorded at low % of the
total fauna ( 1.28 3.12%), but species such as Plethobasus cordatum comprised 40.7% of the fauna
in 1911, declining to 7.2% by 1981 and is now not recorded in the reservoir and has declined
markedly in the riverine sections.
Original Fauna
New Species
Local extinctions
1911
25
-
1981
15
6
10
1994
4
5 (2 new)
11
The Mississippi River system is expected to lose as much as 50% of the current species over the
next decade, directly related to the invasion of the zebra mussel (Master, 1996). Stein & (1996)
points out that the decline of freshwater mussels this will have a detrimental role to the entire
ecosystem, as the freshwater mussels play an imprtant role in sediment mixing and nutrient
recycling, and given their dominance in terms of biomass, the removal could have long-term
recupercussions, as yet unknown.
Summary Statistics
area R1: Ohio-Tennessee rivers (Mississippi), USA (see WCMC report)
richness
of which:
endemic
extinct or threatened
Gastropods
99
Bivalves
120
16 EX
area R2: Mobile Bay basin (Tombigbee-Alabama rivers), USA
Within the Mobile Bay system there are 6 endemic genera ; greatest species diversity in the
Pleuroceridae (76 spp.)
richness
Gastropods
118
endemic
110
Bivalves
74
40
* US Federal List of Threatened and Endangered Wildlife
of which:
* 38 presumed extinct
* 1 Endangered
* 70 Candidate
25 EX
1.4.1.2. USA- SPRING SYSTEMS

The endemic spring-snails are one of the highly endangered groups in the USA and Mexico.
These species are very restricted in their ranges, are typically very abundant within in their small
ranges, and have a generation time of under a year.
Summary Statistics
Springs: area 1: arid/semi-arid Western USA
richness
endemic
>58
Gastropoda
ca. 100
(only Family Hydrobiidae;
espcially the genus Pyrgulopsis) *
of which:
3 EX
Others candidates on
Federal List of Threatened and
Endangered Wildlife
area S2: Florida, USA

richness
Gastropods
(mainly Family Hydrobiidae)
Bivalves
84
endemic
ca. 43
of which:
1.4.2.1. SE ASIA: MEKONG RIVER

The Lower Mekong river was investigated by Davis (1970), and there is a high radiation within
two groups in the Prosobranchs; the faunal assemblage contains over 120 species, mainly the
pomatiopsid family Triculinae (92 endemic species, 11 endemic genera) and the Stenothyridae (19
endemic species). However, there remains more to survey, as only about 500 km of the lower
Mekong main course (with the tributary Mun River) has been well-studied.
Summary Statistics:
area R3: Mekong River (lower Mekong), Thailand-Laos-Cambodia
of which:
richness
endemic
Gastropods
121
111
Unknown
Bivalves
39
5
Unknown
1.4.3.
S. ASIA:
1.4.3.1 NORTHERN WESTERN GHATS, INDIA

The main literature from this region is given in Subbao Rao (1979). There are some distinctive
taxa within the region: 2 endemic genera Turbinicola, Cremnoconchus. The highly localised genus
Cremnoconchus is of phylogenetic significance, as it is the only member of the family
Littorinidinidae known living in a freshwater/terrestrial environment. Members of this group
are typically found in Mangrove or marine shore environments.
The succineid genus Lithotis is known from two species: L. tumida not collected since its
description in 1870, and L. rupicola only known from a single locality.
At present there are no species listed as Endangered on the Red List (Baillie & Groombridge,
1996), but a recent submission for a species of Unionid Pseudomulleria daleyti Smith, 1898 from the
family Etheriidae
has just been evaluated as Endangered B1, B2c. This freshwater species
lives in rivers attached to rocks, in clusters of 3 to 10 individuals, only occasionally exposed
usually submerged.
This species is currently threatened by the construction of a dam, and it
is anticipated that if construction goes ahead the range will decline, with the possibility of local
extinction. It is only known from one other site, and at present the current status of this other
population is unknown.
Summary Statistics:
area R4: Northern Western Ghats, India
Gastropods
Bivalves
richness
ca. 60
11
endemic
ca. 10
3
of which:
Unknown
1 : see note above
1.4.3.1 Japan
area L2: Lake Biwa, Japan
At present no species for this region has been listed, which is a reflection of the non evaluation
rather than the status of the fauna. Again many of these probably qualify as Vulnerable D2, if not
higher.
richness
of which:
endemic
Gastropods
Bivalves
38
16
19
9
area L9: Chilka Lake [brackish water], India
Gastropods
Bivalves
richness
28
43
endemic
ca. 11
25
of which:
Unknown
Unknown
endemic
9
2
of which:
Unknown
Unknown
area L10: Inl_ Lake, Myanmar

richness
Gastropods
Bivalves
25
4
1.4.4. SOUTH AMERICA

Much of the faunal knowledge in this region is still at the checklist stage, and there is still a need
to carry out investigations of the headwaters, and make inter darinage systems comparisons to
establish the taxonomic status of some of the nominal taxa for the region.
1.4.4.1 SOUTH AMERICA: PARANA RIVER
The surveys prior to the construction of the Parana Dam on the Parana river showed that there
were endemic species present in the rapids. These species were threatened with extinction
resulted from changes in the flow regime at Yacrta-Apipe rapids on the Parana River after the
Parana Dam was constructed (Bertonatti, 1999). These species were taken into a captive breeding
programme, and three taxa (Aylacostoma guaraniticum (Hylton-Scot, 1953) Aylacostoma chloroticum
(Hylton-Scot, 1953) Aylacostoma stigmaticum (Hylton-Scot, 1953) are now extinct in the wild
(Darrigan pers. comm; see Quintana, 1999; pers. Comm. For Action Plan).
Downstream from the now disappeared Guayri Falls (flooded when Brazil and Paraguay built
the Itaip hydroelectric plant), the Upper Paran river runs along the Paraguay/Argentina
border confined between vertical walls up to 100 m high, deeply eroding the stony Serra Geral
plateau. Around 56W, during a remote geologic time, the Paran was temporarily delayed by a
basaltic mantle that obstructed its way to the southwest. An amygdaloid rock called spilite,
showing a vacuolar appearance due to steam bubbles, was originated in contact with freshwater.
Finally, the Paran surpassed this bed, divided in multiple branches that then anastomosed to
form a large and complex system with more than 300 islands (Ibicu, Yacyret, Talavera, Apip
Grande, and lesser islands), many rapids such as Apip, and deeper passages such as Mbaracay.
This portion of the Paran, having a flat bed and serious obstacles for navigation, became
radically different from the preceding river, which was deep and carved between high gullies.
Downstream was also completely different, where the water runs towards the confluence with
the Paraguay River, and also in its last sections, which developed a wide floodplain and finally
reach their end in the Plata estuary. Such a limnic singularity brought about endemic forms of life
in this particular section, which were not able to expand their distribution to other parts of the
basin. Instead, they developed a remarkable ecological fidelity to the river bottom in the rapids
area. This is the case of the prosobranch snails belonging to the genus Aylacostoma, a group of
particular scientific interest within the mollusk fauna of Argentina and Paraguay. These mollusks
have a unique reproductive system characterized by parthenogenesis (reproduction without
males) and simultaneous development of up to three embryos inside a marsupium, or
adventitious pouch, located within the parent's neck. These young snails are born when they
have attained a surprisingly big size, and are thus able to withstand the violence of the rapids.
They feed while protected within a thicket of diatoms, green and red algae growing upon the
adult's shell.
Aylacostoma belongs to a group of mollusks of cenozoic origin, which invaded South America
from the northern hemisphere in late Cretaceous or early Tertiary periods, when both continents
became connected for the first time, some 65 million years ago. An abundant fossil record in
strata from the Paleocene ranging to the southern tip of Patagonia evidences its early and
successful adaptive radiation. Climatic and hydrographic changes, however, restricted its
distribution towards the north. Nowadays, the group is limited to fluvial systems in northern
South America. Just one relict area from its former southern distribution persisted outside the
tropics, in the Plata basin until three years ago: the Yacyret-Apip rapids, now flooded by the
main Yacyret reservoir (1,600 km2).
Those rapids represented the unique habitat of a group of endemic species from Argentina and
Paraguay (Aylacostoma guaraniticum, A. chloroticum, A. stigmaticum, A. cingulatum), where they
were discovered and described as late as the 1950s. This discovery was a late and surprising
addition of a tropical family (Thiaridae) to the faunistic inventory of these temperate countries.
Their populations were abundant but had a very limited range, and were made up completely of
females. Due to their particular mode of reproduction, all individuals are essentially identical
within each population or micro-deme (diversity only shows up as a few mutant or senile
specimens) whereas differences appear only between demes. These populations could be
regarded as clones, since the offspring are not produced by the mating of males and females with
different features, but are exact replicas of a single parent. In cases such as the present one, the
low genetic variability leads to a very low adaptive potential to withstand environmental
changes. Moreover, due to their viviparity and low mobility, these snails do not disperse from
their birth place. Even if reproduction without mating is the rule in these species, this may be
periodically compensated by gene recombination by a still unknown mechanism. The cytological
structure of the individuals (haploid, diploid or polyploid), as well as many biological features of
the Argentine-Paraguayan forms, are still completely unknown.
Although their 4 cm long shell makes them conspicuous to a keen observer, and several
thousands were present in a limited area, these snails were totally unknown to the local people.
However, plentiful shell collections were gathered by occasional visitors during the lowest waters
of this century. This evidences the secret life of these animals and their fidelity to the bottom of
the Paran river in the rapids area, characterized by its difficult access and risky navigation.
Since the Yacyret area was the only place where they were found, their total extinction was
expected on flooding of the reservoir, which would lead to the disappearance of the rapids and
all the organisms so strictly adapted to life in this habitat. Indeed, the former rapids became
covered by more than 10 m of still water. Thus, the rocky, well-illuminated bottom bathed by
clear and oxygen-rich waters promoting a dense layer of algae, changed to its present
appearance: a dark and muddy bottom in which algae were unable to survive. Given this
scenario, in August 1993 a team of biologists from the Argentine Museum of Natural Sciences
(MACN) were permitted by the Entidad Binacional Yacyret, in charge of the dam, to survey the
typical locations of the snails in this portion of the river. Many sites were surveyed, including the
few rapids still existing, since most of them (such as the renowned Apip rapids) were lost during
the early flooding of the dam. A dense Aylacostoma population consisting of five different
morphotypes was found along a 150 m stretch of shore in the rapids area. Of these, three could be
identified as known species, but the other two might be new varieties that, unfortunately, were to
be discovered scarcely before their inclusion in the long list of casualties due to human activities
Summary Statistics:
area R5: Parana River, Argentina-Paraguay
richness
endemic
Gastropods
>7
7
Thiaridae
of which:
3 EW
1.4.4.2 SOUTH AMERICA: URUGUAY & LA PLATA RIVER

One species of terrestrial gastropod (Anthinus albolabiatus) which was formerly endemic to Gallery
Forest adjacent to Uruguay River has been proposed as Extinct following construction of the Salto
Grande Dam (Mansur, 1999; pers. comm.).
The La Plata River malacofauna is also suffering declines relating to the spread of the Southeast
Asian Freshwater mussel (Limonoperna fortunei (Dunker, 1857), which is colonising new areas
moving upstream in the estuarine river system; the species has expanded upsteam north to the
confluence of the Paraguay River and the Parana River, a distance of over 1,100 km since 1991
(Darrigan, 1998).
Summary Statistics:
area R11: Lower Uruguay River and Rio de la Plata, Argentina-Uruguay-Brazil
Number
richness
endemic
Gastropods
54
26
Unknown
Bivalves
39
8
Unknown
This lake is one of the notable hotspots in terms of other species endemism and as such figures on
the Molluscan endemicity have been established. Again the spring-snails for the region have yet
to be investigated. No estimates of threat to the fauna have been established.
area L8: Lake Titicaca, Bolivia-Peru
Gastropods
richness
24
endemic
15
of which:
1.4.5.1 AFRICA: SOUTH OF THE SAHARA

The level of molluscan diversity is not as high as the drainage systems in North America or the
Mekong, but the richest river systems are listed here. There are four main basin drainage systems
in africa, south of the sahara, the Niger, the Nile, Zaire and Zambezi (see Figure xx).
The endemic threatened species are nearly all specialised for rapids in river systems. These
species are often only known from small stretches of the river system.
LAKE VOLTA
Area: 8800 Km2
Maximum Depth: 80m
Year of closure: 1964
The construction of the dam, and the subsequent colonisation of the lake by Bulinus truneatus, a
freshwater pulmonate, lead to an increase in the level of urinary schistomiasis reported in the
region (See list of publications in Brown, 1994). Prior to the construction of the dam, this species
was rare, other species from the same genus such as Bulinus globosus, were more common.
However, the latter species was unable to exploit the new habitat, as it cannot tolerate the lakelevel fluctuations. Bulinus truneatus, on the other hand, uses the equatic vegetation such as
Ceratophyllum demersum, and dense populations survive in these stands. In contrast a smaller lake
created by a dam 20 Km downstream on the Volta River, sustains Biomphalaria pfeifferi. Thus there
has been a change in the species assemblages found pre-impoundment and post-impoundment,
and with the additional consequences of increase in human diseases, associated with the
colonising molluscan species.
The Niger, despite its length does not have any endemic species, although the headwaters have
not been properly explored. Brown (1994) speculates that the lack of endemism maybe due to the
impact of Pleistocene climatic change, leading to fluctuations in river level. The majority of
endemism is found in the Volta basin (six endemic species of the prosobranch family Thiaridae
(Potadoma togoiensis, P. bicarinata, Pseudocleopatra voltana, P. togoensis).
Other notable species in the area include 4 species of the genus Sierraia which are known to live
in rivers that are practically devoid of dissolved chemical content in some seasons.
Summary Statistics for region.
area R6:
Western lowland forest and the Volta basin,
Ghana-Cote d'Ivoire-Sierra Leone-Liberia-Guinea
richness
endemic
Gastropods
>=28
19 (plus 9 'near endemic')
of which:
2 CR
Central-Southern Africa
Zambezi River System.
Upper and Middle Zambezi. (Brown, 1994)
Dam
Prosobranchia
Pulmonates
Bivalves
3
4
No. of
Species
22
3
No. of Endemics
2
Lake Kariba: Zambezi River. (Brown, 1994)

Area: 4300Km; Length, 300Km; Maximum Depth, 125m
Year of closure 1958 (filled 1963)
Brown (1994) lists the known gastropod fauna from the Upper and Middle Zambezi River
system. He notes that two species are endemic to the area, Gabbiella balovalensis and Gabiella
zambica; both are from the family Hydrobiidae, and are found in very few localities. He notes that
the headwaters and low tributaries are insufficiently known. Some species became established in
the lake soon after damming, due to the rapid expansion of the plant Salvinia.
Zaire Basin
The lower Zaire Basin is one of the richer areas in central Africa, including the largest expanse of
tropical lowland forest (Guineao-Congo). There are some classic accounts of the fauna of the
region given in Pilsbry & Bequeart (1927) and Mandahl-Barth (1968). The basin changes along its
length, with mountain torrents in the east, leveling to a sluggish stream, descending abruptly to
the central basin, where on the stretch between Kisangani to Kinshasa there is a drop of 100m
over 200 km. The section beyond Kinshasa have series of rapids to Matadi, a distance of 350 km,
and in these rapids there are species representing 5 endemic 'rheophilous' genera, belonging to
the prosobranch families Bithyniidae (Congodoma, Liminitesta) and Assimineidae (Pseudogibbula,
Septariellina, Valvatorbis). These species adhere to bare rock, resisting fast currents and tolerating
a wide range of water levels (Brown, 1994). Brown (1994) notes the uniqueness of the genus
Septariellina that resemble the freshwater limpets Ancylidae.
area R7: Lower Zaire Basin (downstream of Kinshasa),
Congo-Democratic Republic of Congo
richness
endemic
Gastropods
96
24
area R9: Madagascar

richness
Gastropods
30
Endemic genus Melanatria.
endemic
12
of which:
1 EN
2 VU
of which:
1 EN
area L4: Lake Tanganyika, Burundi-Democratic Republic of Congo-Tanzania-Zambia

richness
of which:
endemic
Gastropods
68
45
32 EN
Bivalves
15
8
area L5: Lake Malawi, Malawi-Mozambique-Tanzanie
richness
endemic
Gastropods
28
16
Bivalves
area L6: Lake Victoria, Kenya-Tanzania-Uganda

richness
endemic
Gastropods
28
13
Bivalves
18
of which:
7 EN
1 VU
of which:
5 EN
1 VU
1.4.6. EUROPE (INCLUDING FORMER SOVIET UNION)

area R10: Zrmanja River, former Yugoslavia
richness
Gastropods *
11 *
* only hydrobioid snails
endemic
5*
Number
Unknown
area S4: Balkans region, former Yugoslavia-Austria-Bulgaria-Greece

of which:
richness
endemic
Gastropods
ca. 190
ca. 180
3 EX
9 CR
10 EN
3 VU
area L1: Lake Baikal, Russia
Nearly180 species are known from Lake Baikal (Sitnikova, 1995), of which about 2/3rds are
endemic to the lake system. The radiations include the hydrobid group, which are largely lake
dwelling in Baikal, with the endemic genus Kobeltocochlea. Many of these species are currently
included in the Russian Red List, but as they have not been evaluated in terms of the new IUCN
criteria, although in all liklihood, many of them will qualify under Vulnerable D2.
Slugina et al (1994) noted that the bivalves, whilst not so numerous provide an important part of
the biomass in the lake system. With endemic species of the family Sphaeriidae dominating the
benthic macrofauna on the bottom near shore area.
richness
Gastropods
Prosobranchs
Pulmonates
Bivalves
147
64
83
31
endemic
114
of which:
14
area L7: Lake Ohrid and Ohrid basin, Albania-Former Yugoslavia

richness
of which:
endemic
Gastropods
72
55
1 EX
1.4.7. AUSTRALIAN FRESHWATER MOLLUSCS

Recent accounts of the freshwater molluscs of Australia are described in Mollusca: a southern
synthesis (Beesley, Ross & Wells, 1998), and the data is here is drawn from there, and other
referenced sources.
The main diversity of the Australian freshwater fauna lies in the family Hydrobiidae (spring
snails), although endemic species are known within the Assimidae, Pomatiopsidae and the
Bithyinidae. Within the bivalves, the largest species are the freshwater mussels, such as the
family Hyriidae (which is also present in South America), and this group was used by McMichael
& Hisock (1958) to draw up biogeographical patterns for freshwater molluscs in Australia. Other
important families are the Corbulidae and the Sphaeriidae. The most cosmopolitan group is in
the pulmonates, similar to patterns of endemism seen in the USA.
There has been a decline in freshwater biodiversity in river systems in Australia, where some of
these groups were well adapted to the strong seasonality of the flow regimes (successive seasons
with flash flooding) are now reducing in abundance and diversity since control of river systems
(Walker, 1985; Ponder, 1998). The best documented cases lie in the Murray-Darling River
Systems. There the freshwater gastropods and bivalves in the system have dramatically declined
in recent decades. There are three reasons given for these declines:
a. predation by the freshwater fish, especially the common carp (Cyprinus carpio) which became
widespread in the 1960's. These fish also disturb the sediments on the river bed, leading to
degradation of the habitats (Fletcher, Morison & Hume, 1985)
b. changes in the flow patterns through intensive flow regulation after impoundments and weir
construction
c. possible changes in the biofilms of algae, bacteria and fungi which are a potential food source
for molluscs (Sheldon, 1994)
area S5: Great Artesian basin, Australia
richness
endemic
Gastropods
of which:
area S6: Western Tasmania, Australia

richness
endemic
Gastropods
of which:
4 EX
1.4.8. SE ASIA: NEW CALEDONIA & SULAWESI

Much work is still being done in this region.
Sulawesi & Indonesia
Bouchet has already recognised the presence of 3 endemic freshwater genera: Tylomelania,
Miratesta and Protanoylus. But there are many species still undescribed. In Lake Poso Bogan &
Bouchet (1998) noted that all ten species currently recorded are endemic, but there are significant
new species such as Posotrea anomiodies which is a member of a new genus of the freshwater clam
Corbiculidae. The new species was found cemented to a hard substrate, showing characters
which are unique, for the region and the family, and as such places a higher conservation value
on the fauna of Lake Poso.
Summary Statistics:
area L3: Sulawesi (Lake Poso and the Malili lakes system), Indonesia
richness
Gastropods
Bivalves
ca.50
5
endemic
ca.40
2
of which:
1 EX
1 EX
endemic
65
of which:
40 VU
area S7: New Caledonia
Gastropods
richness
81
3 EN
1 EX
LAKES
area L2: Lake Biwa, Japan
At present no species for this region has been list, which is a reflection of the non evaluation
rather than the status of the fauna. Again many of these probably qualify as Vulnerable D2, if not
higher.
richness
of which:
endemic
Gastropods
38
19
Bivalves
16
9
area L4: Lake Tanganyika, Burundi-Democratic Republic of Congo-Tanzania-Zambia
richness
of which:
endemic
Gastropods
68
45
32 EN
Bivalves
15
8
area L8: Lake Titicaca, Bolivia-Peru
Gastropods
richness
24
endemic
15
of which:
area L9: Chilka Lake [brackish water], India
Gastropods
Bivalves
richness
28
43
endemic
ca. 11
25
of which:
Unknown
Unknown
endemic
9
2
of which:
Unknown
Unknown
area L10: Inl_ Lake, Myanmar

richness
Gastropods
Bivalves
25
4
The extinction crisis for the US freshwater mussels was recognised last century, where Higgins
(1858) observed declines related to deforestation. Rhoads (1899) remarked on the decimation of
freshwater bivalves in the lower Monongahela River above Pittsburgh due to pollution and
Ortmann (1909) reported declines on the Tennessee River systems due to paper mill effluents.
However documentation of the extinction crisis did not really gain momentum until the 1970s
(Athhearn, 1970; Heard, 1970; Stansberry, 1970; Taylor, 1970). Freshwater mussels (Bivalvia) are
especially vulnerable to habitat disturbance. These species have extended life cycles, some of
which span over 100 years, where maturity is delayed until 6 15 years (Bauer, 1993; Chesney &
Oliver 1998). Some species also have reduced powers of dispersal, high juvenile mortality and
long turnover times. In the USA these mussels are most successful where water velocities are low
enough to allow substrate stability, but high enough to prevent excessive siltation. Similar
conditions occur in Europe, although some species are adapted to surviving in silty/muddy
environments. In the USA most rivers show a patchy distribution of mussels, with some areas
containing densely packed beds, which may be composed of many species, with other patches
of river bed being devoid of any species (Neves & Widlak, 1987)
2.0. UPSTREAM IMPACTS

Frequently the data available for pre/post impoundment is assembled by the provision of
checklists. These often do not indicate the absolute impact of impounments upstream of the site.
It is clear that to some extent the impact depends on the type of flow into the reservoir and the
nature of the channels.
Direct reduction of species upstream relating to changes in river channel
Molluscan survey was undertaken on a braided river that enters a reservoir on the River Inn in
Austria; the river is currently dammed by weirs for hydroelectric production (Foeckler,
Diepodder & Diechner, 1991). The data shows that there has been a decline upstream of the
reservoir lossing 10 species (Table xx) and a drastic decline downstream of 38 species. The
upstream decline is due to changes in the flow regime, as since the braided system has been
managed by channelisation there has been an increase in river gradient of 1.3% deepening, the
bed and consequently reducing the active floodplain area, with some marginal areas only being
inundated once every ten years. This has reduced biotype diversity with consequent loss of
species.
Table xx. Decline in Freshwater Molluscan species within the Inn catchment since 1971 (data
abstracted from Foeckler, Diepodder & Diechner, 1991)
No. of species
New species
Species lost
Current survey
36
1
10
River Salzach
River Inn
below reservoir.
Surveys by Seidel
(1971-3)
45
?38
They recognise the loss of species typical of temporary habitats (Segmentina nitida, Viviparus
contectus), the more demanding species of permanent water (Acroloxus lacustris, Radix ampla,
Gyraulus crista, Anodonta anatina) and species requiring non-polluted water (Unio crassus, Pisidium
terinilineatum).
Interesting they attribute some of these absences to the indirect effects of changed flow relating to
presence of the reservoir downstream.
They predict that the temporary habitat species such as the endangered species Segmentina nitida
will become extinct quickly in the region but most significant impacts are on their alpine special
species, which are not present in all rivers, such as Gyraulus acronicus.
2.1. Movement of species upstream

Dam building activity which blocks migratory fish or changes fish communities can also impact
on the freshwater mussel communities which depend on the fish as glochidial hosts.
Dam construction at Lake Pepin on the Mississippi River lead to the demise of mussels upstream
of the dam, as the runs of skipjack herring, their host species, were blocked (Eddy & Underhill,
1974).
2.2. Movement of nutrients upstream
3.0. LATERAL IMPACTS
The major impact on molluscan biodiveristy is on the terrestrial areas adjacent to the current
river system. Maintenance of areas of contiguous undisturbed land within this system is an
important consideration for molluscan conservation. For example In the steepsided valleys of the
KwaZulu Natal province, dam construction, or any development that results in the loss or
fragmentation of relatively pristine or untransformed habitat is likely to impact upon mollusc
populations. The reservoirs will not only eliminate terrestrial molluscs in the areas to be flooded,
they will also transform the habitat in their immediate neighbourhood. This could also result in
the formation of an in-valley barrier, separating up-stream and down-stream populations
(depending on the width of the valley and the height of the dam). This would obviously impact
upon gene flow and population dynamics in low vagility invertebrates such as molluscs.
Rogers (1995) examined riparian wetlands (including riverine forests and thicket) and concluded
that in southern Africa these habitats were under severe threat, especially from alien biotas.
In South America there are possible species extinctions related to loss of gallery forest adjacent to
rivers which are now submerged following construction of the Salto Grande Dam. The land-snail
Anthinus albolabiatus (Jaeckel, 1927) (Mollusca; Gastropoda:Strophcheilidae) was formerly
endemic to Gallery Forest of Uruguay River and has been submitted for inclusion in the IUCN
Red List of Threatened Animals.
3.1. Movement of species laterally
3.2. Movement of nutrients laterally
3.3. Water table changes
A high number of endemic Spring-snails (Family Hydrobiidae) live in small geographically
restricted habitats located in the headwater of the main channels and tributaries within the water
catchments. This group of species is second most endangered groups within freshwater molluscs.
In Europe, Australia and the USA, this species group usually comprises c. 20% of the total
freshwater fauna.
The species are already under pressure from water abstraction at the springs and from habitat
alteration to facilitate abstraction, although a more indirect threat is from reduction of
groundwater levels. Such changes can result from impoundment and the consequent impact on
the flow regime and groundwater recharge within the catchment.
4.0. RESERVOIR IMPACTS

A. Construction impacts
4.1. Site impacts (effects on river environment & biota at site)

Stresses associated with site disturbance, such as dam contruction resulting in physical
disturbance of the river bed may cause species from the bivalve groups superfamily Unionidea to
prematurely empty their brood pouches of glochidia resulting in reproductive decline (Howells et
al., 1996).
4.2. Off-site impacts (materials for dam, access roads, etc.)
Riparian habitats, such as gallery forest and marsh also hold unique species, which are sensitive
to disturbance during the construction phase. The destruction of habitats for temporary roads
leads to loss of species, which maybe permanent or temporary, depending on the degree of
degradation and the amount of habitat fragmentation.
Increased traffic of heavy lorries on the access roads leads to the increased dust which coats the
local vegetation degrading the adjacent habitats. However, data on such losses are rarely
reported, and hence it is difficult to quantify such claims.
B. Post-construction impacts of reservoir
4.3. Running to still water impacts - reduces lotic, increases lentic biota
There are several sections in this report which observe the extinctions or threats to mollusc
species from rapids on the rivers systems, notably in Africa, North and South America.
Such data is also available for decline in local species richness for Europe, for example Fruget
(1991) found that partitioning with a series of dams on the lower Rhone, had an influence on the
resultant macroinvertebrate community structure and distribution, especially due to changes in
flow velocity. The Lower Rhne has a significant number of low level dams for hydroelectric
production and irrigation schemes. Most were constructed between 1957 and 1985, although
channel management was an ongoing process in the industrialisation period in France. Only 19
species of Mollusc were recorded, although this was the most taxonomically diverse group of
macroinvertebrate in the sampling sites along the river. In terms of relative abundance, molluscs
formed 7% of total fauna, significantly less than the Chironomids (61%), as as such the
significance in this case is the loss of diveristy, rather an impact on the biomass, as is the case with
the Chironomids.
However, the creation of the dam can also reduce lentic taxa, through channel management after
construction. For example, river regulation on the lower Murray River in Australia has altered the
normal flooding regime leading to alienation of the billabongs (low Ox-bow lakes) from the main
channel. These lentic macrohabitats (billabongs and backwaters) are characterised by a high
density of scrapers, especially freshwater gastropods. Since regulation was introduced some
Mollusc species have become extinct (Boulton & Lloyd , 1991).
4.4. Creates new sublittoral and profundal zones
The construction of the dam on the Lake Volta in West Africa, and the subsequent colonisation of
the lake by Bulinus truneatus, a freshwater pulmonate, lead to an increase in the level of urinary
schistomiasis reported in the region (See list of publications in Brown, 1994). Prior to the
construction of the dam, this species was rare, other species from the same genus such as Bulinus
globosus, were more common. However, the latter species was unable to exploit the new habitat in
the littoral zone, as it cannot tolerate the lake-level fluctuations. Bulinus truneatus, on the other
hand, uses the aquatic vegetation such as Ceratophyllum demersum, and dense populations survive
in these stands. Thus there has been a change in the species assemblages found preimpoundment and post-impoundment resulting from changes in the littoral zone, and with the
additional consequences of increase in human diseases, associated with the colonising molluscan
species.
4.5. Water level fluctuation impacts
Brown (1994) describes the gastropod diversity of some of the man made lakes in Africa which
are comparable in size to the larger natural lakes. These lakes are the result of dam construction
for economic reasons; usually hydroelectric power generation. The outflow from these lakes
differs from the natural lakes, with most man made lakes suffering seasonal draw-down as flow
out from the lakes is regulated to ensure that the large inflow during the rainy season can be
contained. This gives a very unstable littoral zone, which in itself will place special stresses on
aquatic life at the margins; a factor which will restrict the number of mollusc species which can
survive in the lake.
4.6. Chemistry changes (mercury, humic acids, etc.)
Reservoir construction may alter water chemistry which may in turn impact on successful
spawning of mussels (Isom, 1971). Bivlave species are very sensitive to water chemistry, as the
translocation experiments on endangered mussel species in Europe has shown, as changes in
weater chemistry can lead to stress triggering release of glochoidea during unsuitable flow/water
conditions.
In Africa, Brown (1994) notes that the salinity of the man made lakes remains quite low, despite
the degree of evaporation, partly related to the annual flow regime (and high in flow).
In contrast the contruction of dams on the Murray-Darling rivers in Australia has led to increase
salinities in the reservoir water, but there is no evidence to suggest that these increases have had
any impact on the freshwater mussels in the system (Vickery 1978; Walker 1981). However the
salinity levels of water in the evaporation basins used to store irrigation return water in the
Murray-Darling system, is listed as having an impact on molluscs (Walker, 1998).
Dissolved oxygen levels below 20% saturation can cause stress to freshwater mussels (Ellis 1937,
Ingram 1957), although some species can withstand brief periods of low oxygen levels.
4.7. Physical changes to the fluvial system
4.7.1.Physical disturbance
Stresses associated with site disturbance, through harvesting, artificial disturbance of bed for
fishing purposes or navigation may cause species from the bivalve groups superfamily Unionidea
to prematurely empty their brood pouches resulting in reproductive decline (Howells et al.,
(1996).
4.7.2. Increased siltation
The greatest diversity in the prosobranch gastropod fauna in the USA is found in the Mobile Bay
River basin, and the Tennessee River Basin, with 7 % of total taxa now extinct (Bogan, 1998).
Most of the extinctions in this group (38 out of 42 taxa) are in the Mobile Bay fauna, when the big
river shoal fauna was impounded and covered by deep standing water and subsequent siltation.
Fuller (1974) considered siltation to be a major factor impacting mussel communities in the US.
Howells et al (1996) suggest that some freshwater mussel species may be unable to survive silt
overlays greater than 0.6cm, in contrast, other species may emerge from silt overlays of 18cm,
although some species need to emerge within a few hours of overlay or they will die. Howells et
al. (1996) also noted that the presence of silt in suspended load may reduce the feeding efficiency
of filter-feeding bivalves.
In Texas Neck (1982) noted that reservoir construction was probably the most significant factor
influencing mussel populations. In Texas only one natural lake existed prior to the construction
of reservoir dams. He gives one possible reason for a reduction in the number of mussel species
in the state as inability to colonise areas which were newly formed silt beds, as the species locally
available were not adapted to living within silt beds.
4.7.3. Temperature
Early work in the US, when the demise of large Unionid bivalves was becoming apparent, led
Isom (1971) to conclude that reservoir construction may alter water temperature which may in
turn impact on successful spawning of mussels. More recent work on freshwater mussels
confirms these findings, where temperature affects growth and the duration of the growing
season (Tevesz & Carter, 1980).
4.7.4 Dissolved oxygen
Dissolved oxygen levels below 20% saturation can cause stress to freshwater mussels (Ellis 1937,
Ingram 1957), although some species can withstand brief periods of low oxygen levels.
4.12 SITE SHADING, AND ADJACENT VEGETATION
Whilst freshwater mussels can survive, in the sort term, removal of the shading trees adjacent to a
channel, will lead, in the long term, to a decline as growth rates are potential controlling agents of
unionid distribution (Morris & Corkum, 1999). Species that grow in rivers dominated with
forested riparian vegetation show slow growth through life, whereas species which dominate in
grassy rivers have rapid early growth, and achieve a smaller maximum size at a younger age.
(Morris & Corkum, 1999). This shading thus effects the species which can in the long term
survive in the channel, and some mussel species are known to attempt to relocate to more
favourable habitats downstream.
4.13 GENETIC VARIABILITY IN RESERVOIR POPULATIONS OF PROSOBRNACH MOLLUSCS IN ISRAEL
The freshwater prosobranch genus Melanopsis is a key group in many studies concerning
systematics, parasitology, ecology, palaeontology and evolution (Heller et al. in press). The
Jordan Valley has highly variable populations of Melanopsis and in fossil sediments (in which
Melanopsis is very abundant).
Heller (in press) describes the shell morpometrics sites within this region. Within Melanopsis
costata Heller et al (in press) distinguish three groups, and tentatively suggest that they may be
subspecies. Shells from the upper Jordan River are elongate (M. c. costata); those from Lake
Kinneret are stout (M. c. jordanica); and in those of the Yarmouk each rib consists of a pronounced
upper tubercule, fused to a pronounced, straight lower ridge (M. c. noetlingi).
Heller et al. (in press) describes the former environment where the water level of Lake Kinneret
used to fluctuate annually at an average of 0.801.00 m (maximum 1.50 m). In 1932 the Kinneret
was dammed and its outlet deepened to construct a hydro-electrical plant, and from that year
onwards annual fluctuations in the water level increased to 3 m. In 1964, when the lake became
Israels major reservoir, its level was increased by 1 m; and was left at peak levels for much longer
periods than in its natural condition (Nun, 1977). The small, stout shell of M. c. jordanica
correlates with a stormy habitat. A stout, globose shell could accommodate more foot muscle and
thus enable a stronger adherence to rocks and boulders during severe storms. Melamposis c.
jordanica is found in Lake Kinneret (170 km2), which differs from the Jordan in its frequent
storms. Daily from April to October, a western breeze from the Mediterranean Sea reaches the
Kinneret at about noon. This breeze causes the lake to become wavy, choppy or stormy until late
in the evening. From December to mid-March there is no western breeze, but strong eastern
winds from the Syrian desert (the Sharkiyeh) then frequently reach the lake causing similar
effects (Heller, 1979a). Two snail-predating fish, Barbus longiceps and Blennius fluviatilis, are found
in Israel mainly in Lake Kinneret (Heller, 1979a); they may perhaps exert some selection on shell
colours. M. c. jordanica occurs only along rocky shores of the Kinneret, consisting of gravel,
cobble, stones and boulders; and is not found on muddy shores consisting of muds, sands and
silts. It occurs down to a depth of 5 m (Tchernov, 1975a; Raanan, 1986). Further, in its natural
condition the lakes salinity was almost 400 mg/l (chlorinity). Recent diverting of haline springs
from the lake has decreased its salinity to about 200 mg/l (Nun, 1977). Todays major habitat of
M. c. jordanica is thus very disturbed. Beyond Lake Kinneret, M. c. jordanica occurs in the upper
reaches of the lower Jordan River, from the exit of Lake Kinneret another 2 km downstream, to
Deganiya (beyond which the Jordan is polluted and no Melanopsis are found for quite a
distance). Specimens in the National Mollusc collection record itspresence also 30 km
downstream (Dalhamya, Shifa).
Raanan (1986) suggested that the black and banded morphs of M. costata of Lake Kinneret may
belong to two separate species, with incomplete reproductive isolation. However conchiometric
and electrophoretic studies do not support this suggestion (Marko, 1984; Altman & Ritte, 1996).
In the small, 1 km-long section of the north-eastern shores of Lake Kinneret, from the Upper
Jordan outlet to the Nahal Meshoshim outlet, shells intermediate between M. c. jordanica and M.c.
costata were found. This small stretch has undergone considerable changes since the draining of
the Hula in 1958: large quantities of sediment that once used to be deposited in the Hula today
reach the Kinneret where they have created a rapidly expanding delta, into which the river bed of
the Jordan now extends about 1 km eastwards, towards the Nahal Meshoshim outlet (Nun, 1977).
Heller et al (op cit) point out that it is noteworthy that among 35 samples of Lake Kinneret whose
isozymes were examined by Altman & Ritte (1996), their two samples from Nahal Meshoshim
outlet had the lowest average genotypic similarity. Their evidence (based upon electrophoresis)
supports Heller et al. (op cit) suggestion (based on conchiometrics) that the Meshoshim outlet
population differs from other populations of Lake Kinneret (which are jordanica) in that it is
intermediate between M.c. jordanica and M.c. costata. In the lower reaches of the lower Jordan
another shell occurs, M. c. noetlingi.
In zones of contact the three species largely remain distinct. Hybrids are found in overlapping
zones of no more than a few hundred m, and at low frequencies. This suggests that the species
maintain their distinct genetic integrities, and that the narrow hybrid zones act as substantial, but
not absolute barriers to gene flow. Tchernov (1975b) notes that the smooth and costated
Melanopsis interbreed, at sites where small water bodies reach the shore of Lake Kinneret.
Situations comparable to Melanopsis, where taxa maintain their integrity yet hybridize at the edge
of their geographical or ecological distributions, have been found in a variety of animals,
including mammals, birds, amphibians, reptiles, orthopterans and butterflies (Grant & Grant,
1992; Harrison, 1993; Werner & Watson, 1996; Jiggins et al. 1996). Among gastropods however,
only few cases of hybrid zones have been reported.
Heller et al. (op cit.) do not comment on the possible role that damming has played in the
changing genetic composotion of these populations, howver, given the nature of this freshwater
genus, it is extremely likely that envionmental changes within the Lake resulting from changes
after damming in 1930s may well have played a part.
5.0. DOWNSTREAM IMPACTS
Neves (1999, pers. comm) is his recent review of impact construction of impoundments on
molluscan species richness provides the data in Table 5.0.1. This shows that there has been a
decline of between 40% and 80% loss from the original diversity levels of these rivers, over a
period of 50 years. The figures in post dam richness also indicate the stretches downstream of
the dam where the bed is devoid of mussels.
Table 5.0.1
Impact of Dam contruction and impoundments on Molluscan biodiversity in the USA (Neves,
1999; conference presentation)
Dam/Date
Center Hill,
Caney Fork
(1948)
Demopolis,
Tombigbee R.
(1954)
Wolf Creek,
Cumberland R.
(1952)
Pre-Dam
Richness
39 species
(pre-1940)
Post-Dam
Richness
7 species
(>12 km)
50 species
(1933-35)
29 species
(1992)
39 species
(1947-49)
11 species (1984)
4 species (1993)
(>18 km)
Table 5.0.2 Mollusca Species present within reservoir region, USA

Source: Neves, 1999.
Reservoir
Preimpoundment
Postimpoundment
Date
Richness
Richness
Norris,
40 species
12 species
Clinch R.
(1935-37)
(1990s)
(1937)
Center Hill,
39 species
2 species
Caney Fork
(pre-1940)
(1993)
(1948)
Cumberland,
59 species
16 species
Cumberland R.
(1947-49)
(1961)
(1952)
Wheeler,
>60 species
38 species
Tennessee R.
(pre-1935)
(1960-69)
(1936)
18 species
(1991)
Demopolis,
50 species
8 species
Tombigbee R.
(1933-35)
(1954)
Demopolis/Warrior, Warrior
48 species
13 species
R.
(pre-1950)
(1972-75)
(1954/57)
Vaughan & Taylor (1999) looked at molluscan distributions along a 240 km stretch of the Little
River in Oklahoma finding that there was a mussel extinction gradient downstream from large
impoundments, where with increasing distance from the dam there was a relative increase in
mussel species richness and species abundance. The general distribution of mussels was nested
reflecting the distance from the dam on the river and its tributaries, with only those stretches
furthest from the dam containing the relatively rare species. After the next dam downstream, the
same trends were apparent, but apparently weaker, with reduced mussel abundance overall.
5.1. Seasonality of flow
5.2. Change in flow regime

Australian freshwater mussels Alathyria jacksoni and Velesunio ambiguus have different habitat
preferences in the Murray-Darling river system, the former being a large river species, which is
less tolerant of environmental changes, and is particularly vulnerable to dehydration (Walker,
1998). Velesunio ambiguus occurs in temporary pools, lakes and minor streams and can survive out
of water for up to a year (Walker, 1998). The flow regime in Murray-Darling basin is influenced
by dams and weirs with the consequence that the habitats occupied by these species are less
distinct. However, below the Darling confluence the dams have been replaced by weirs, which is
a habitat which is more favourable for Velesunio ambiguus and Walker (1981) found from
archaeological studies that Alathyria jacksoni has declined in abundance, whereas Velesunio
ambiguus has increased in abundance.
5.2.1. Increase in flow
High water velocities can displace the setting of juvenile mussels in freshwater systems before
they have an opportunity to settle on the bed (Layzer & Madison, 1993; Chesney & Oliver, 1998).
It may also have indirect effects, causing:
more bedload movement damaging molluscs ( juvenile mussels are especially
vulnerable; Young & Williams, 1983).
increased bed erosion moving material (and contained mussels downstream)
decreasing habitat complexity through closing oxbow type channels ( see above
sections)
Williams et al (1993) suggest that sediment stability might be one of the most critical habitat
requirments for freshwater mussels.
5.2.1. Decreases in flow
Long periods of low flow below dams, can result in stranding of mussels (Fisher and Lavoy, 1972;
Chesney & Oliver, 1998). Mussels are unable to repond quickly to changes in water flow, as
although they are able to relocate if conditions become poor, they move slowly. If the musslke
survive the stranding, the stresses will have lead to a loss of the glochidia, and may lead to a
longer term reproductive decline ( McMahon, 1991; Chesney & Oliver 1998).
5.3. Physical changes (temperature, oxygen, & other water quality measures)
5.3.1 Water temperatures downstream of dam
Water releases downstream during summer months in US river systems are colder than the water
entering the reservoir. In some river systems this has been shown to reduce, or even eliminate
mussel populations (Ahlstedt, 1983; Miller et. Al. 1984 Lydeard & Mayden, 1995). This is due to
the suppression of the mussel metabolic rates at a rtime when growth should be high, which in
turn leads to a suppression of reproduction (Vaughn & Taylor, 1999).
There are other indirect results of water temperature changes relating to the changing fish
distributions. Vaughn and Taylor (1999) described the replacement of warm-water fish with
more tolerant species and introuduced coldwater species such as Rainbow (Oncorhyncus mykiss)
and Brown trout (Salmo trutta) as porviding a potential permanent colonisation barrier to
mussels, given the fact mussels species require specific host species, and to a large extent most of
host fish species are unknown.
5.3.3. Salinisation
In Australia, there has been dam construction and weirs in the Murray-Darling basin, changing
the flow regimes, there is less distinction between the occupied habitats of two mussel species,
but below the Darling confluence the dams have been replaced by
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mussel species Hyridella glenelgensis ( Bivalve: Unionidae) which lives in the Glenelg River system
of south-west Victoria, is cited as being at risk through water diversions and salinisation (Walker,
1998).
Not all salinisation is the direct effect of dam construction, but additional factors such as land
clearence is given as a possible reason for the decline of the species Westralunio carteri from the
Avon river system in western Australia (Kendrick 1976).
5.4. Turbidity and sediment
In the Tennessee River Basin a number of Pleurocerid gastropod species are under threat, as they
persist on clean swept shoal areas below dams on the river (Bogan, 1998). These areas are at risk
from increased sedimentation and changes in flow regime.
5.5. Water velocity
5.6. Downstream lakes
5.7. Changes to food chain
The gastropod fauna of the River Murray has declined from 18 species (recorded from various
sources: Museum collections, Publications and Archaeological middens) to a single abundant
native species , a limpet from the genus Ferrissia and an introduced species Physa acuta (Sheldon
& Walker, 1993). The survival of this limpet is attributed to shape of the radula, and the small
size of the animal, which Botting (1995) believed would allow the species to exploit the minute
particles associated with the biofilms, and that most of the larger species would not have access
to this food source. Sheldon (1994) believes that there have been changes in the biofilms of algae,
bacteria and fungi in the Murray River which are a potential food source for molluscs, and that
this factor, along with predation and flow regulation accounts for the decline of species in the
system.
Most of these species are still present elsewhere in the Murray-Darling basin, but one species for
the family Viviparidae Notopala hanleyi, is believed to be extinct in its natural habitat, remaining
as an abundant species in three irrigation pipelines in South Australia, where it is regarded as a
pest species, which requires control to allow water flow for irrigation.
8.2. Agriculture, forestry, industry and municipal effects
The reservoirs created by dams are often used for recreational activities such as fishing and
boating. New species can be introduced by these activities into the catchment area. In the case of
Mollusca, fish stocking by species outside their native catchment areas can introduce new
freshwater mussels, where juveniles, which had their glochidial stage on the fish are released in
the reservoir, and conditions there are suitable for the survival of the species. Howells et al (1996)
cite two cases in Texas, where ponds and reservoirs have new distribution records of the floater
freshwater mussel groups.
8.3. Medical implications of changing molluscan species in rivers
Hydropower and irrigation projects carry the risk of increasing the local prevalence of snailtransmitted diseases in both humans and livestock. These diseases are caused by two main types
of parasitic helminthes: flukes (Trematoda) and roundworms (Nematoda). Both these groups of
organisms have complex life cycles that involve periods in larval hosts, such as snails and an
adult phase in birds or mammals. These hosts are usually very specific, and the increase in
disease results from decline of the native species, and increase in abundance of the host species,
sometimes replacing the natives as there were alien to that specific catchment area.
Burch & Upatham (1989) list those snail species which are known to be intermediate hosts, and
indicates their distributions and parasites in the Meokng River basin. Although there are over 50
species of Pomatiopsidae recognised in the Mekong basin, only two are though to be
intermediate hosts for human parasites (Neotricula species).
Schistomiasis:
In the Mekong river Schneider et al. (1984) believed that these was an increase in parasite
transmission resulting from proposed impoundments which change river habitat and ecology.
This is because the species which can act as hosts become more prolific as they are adapted to the
new environments whereas other similar species which are not vectors, (usually in the same
genus or closely related species) decline in abundance, and maybe lost to the river system
(Woodruff & Upatham, 1992). In Thailand in 1992 the Pak Mun hydropower and irrigation
project caused controversy as to whether the blood flukes which pass on Schistomiasis would
become established from the nearest known source in Laos (Usher, 1991).
Paragonimiasis:
The knowledge in 1992, suggested that molluscs were not the source of this disease in the
Mekong basin, as the creation of new habitats has not yet led to the expansion of the known snail
vectors into Thailand. However, there remains a chance that the snails may become established
and impact on the local species of the genus Brotia.
Opisthorchiasis:
There are 12 species of the freshwater prosobranch Bithynia recognised in Thailand, but only the
three species in the subgenus Digniostoma transmit Opisthorchis viverrini. These species can live at
very high densities and are usually found in rice fields, canals, ponds and lakes. River
impoundment has led to an increase in the abundance of these three species, and disease
outbreaks were noted at the time of construction of the Ubolratana (or Nam Pong) Dam in NE
Thailand (Woodruff & Upatham, 1992).
Echinostomatiasis:
Schneider et al. (1984) predicted that proposed impoundments would lead to an increase in the
incidence of infections and increasing disease severity.
Table 8.3.1
Human diseases, where Molluscs act as intermediate hosts, which may be increasing as the result
of impoundation in Thailand
(Data abstracted from Woodruff & Upatham, 1992)
Disease
Schistomiasis
Blood-Fluke:
Schistoma japonicum
Paragonimiasis
Paragonimia heterotremus
( N.B. P. westamani has not
Intermediate hosts
Molluscs:
Oncomelania spp.
Neotricula spp.
Source of Information
Woodruff & Upatham (1992)
Molluscs:
Brotia asperata
been recorded from humans in

the Mekong basin yet)
Melanoides tuberculata
Tarebia granifera
Opisthorchiasis:
Molluscs:
Opisthorchis viverrini
Bithynia funiculata
Bithynia siamensis
Bithynia goniomphalos
Echinostomatiasis
Numerous
Echinostome
species
Molluscs:
Various hosts including Pila
ampullacea,
P. polita, Radix
rubiginosa,
Indoplanorbis
exustus, Filopaludina spp.
9.0. PRIORITY OF IMPACTS

9.1.3 Loss of keystone species
Stein & Flack (1996) points out that the current decline of freshwater mussels in the Mississippi
Basin will have a detrimental role to the entire ecosystem. They point out that the freshwater
mussels play an important role in sediment mixing and nutrient recycling, and given their
dominance in terms of biomass, their removal could have long-term recupercussions, as yet
unknown.
9.1.4 Loss of stocks of economic species
Many of the declining mussel species were traditionally used in the button industry, and
increasingly for the use as seeding material for pearl production. Thus ultimately there will be a
economic loss to the region.
In terms of the economic losses, Bowen et al. (1994) estimated that the annual value of unionid
shell exports to Japan exceed $40 million, and the Zebra mussel, which will continue to exist in
the river systems, is not a suitable substitute.
9.2. Highest priority impacts
These can be ordered in terms as those that are not reversible; that is the direct loss of habitat, as
mitigation and habitat creation rarely recreates the existing habitat, in terms of its original species
composition.
9.2.1 Loss of habitat under areas of impoundment
a. Extinctions in Mollusca directly associated with dam construction occur with the loss of
rapid habitats within river systems, as frequently these contain endemic species.
b. Local extirpation of species in the adjacent riparian forests, which can with localised species
lead to fragmentation of populations ultimately threatening the species.
Care should be taken during Environmental Impact Assessments to make sufficient provision for
invertebrate surveys of these habitats.
9.2.2. Preventation of migratory routes for fish hosts
After dam construction and impoundment, unless sufficient provision is made for migratory
species, another severe impact is the preventation of fish movments leading to decline in the
mussels that utilise the fish as a host during the life cycle.
Care should be taken during design phase to design suitable fish passes which will not stress the
fish, which may impact on the mussel species.
9.2.3 Changing bed characteristics
Siltation after the impoundment leads to loss both of species within the reservoir area, and on the
steam bed downstream of the dam.
10.0. CUMULATIVE EFFECTS
10.1. Are there other dams on the river or drainage basin?
Some of the river systems here have more than one dam, which has in itself contributed to local
extinctions.
If several large dams are required on a single river system, this could have severe impacts on
molluscan diversity and abundance, and considerable lengths of undisturbed beds are required
to overcome the effect of a single impoundment. To prevent extinctions where large dams are
essential, preference should be given to designs that favour long lengths of undisturbed stream
bed.
10.2 What are additive or synergistic effects of the planned dam in combination?
10.2.1 Extinctions and Decline in Biodiversity
The impact of dam contruction on biodiversity is a combined function with other factors
including:
Pollution:
in the USA and Europe water pollution, especially in large river systems has been a serious factor
causing decline in mollusc biodiversity since the beginnings of large-scale industry. These
industries used river systems to abstract water for power, dispose of liquid waste and discharge
used water. In more recent decades industrialisation has had a similar impact on rivers in SE
Asia, India and Australia.
Exploitation:
Direct harvesting of freshwater bivalves providing shell for the button industry has been a major
factor contributing to the decline of mussels in the USA (Bogan, 1998) and India (Subbao Rao,
1978). In addition shell in both of these regions is now used for seeding pearl oysters, and the
fishing of mussels for their own pearls has been an on-going industry for over 2000 years, as
Roman exploitation is known.
Alien Species:
In the USA and South America a new factor, alien species is causing extinctions of an already
depleted fauna. The combination of the Zebra mussel (Dreissena polymorpha) and the Asian
Freshwater clam (Corbibula fulminalis) is causing decline of many native freshwater species (Stein
& Flack, 1996).
In recent years, the spread of the Zebra mussel in particular has had the most profound impact in
terms of threatening US freshwater mussels. The Zebra mussel has a different life cycle, partly
due to the lack of a need for a host fish, and partly as it can anchor. The native mussels are large
enough to be used as a anchoring point for Zebra mussels and thus they become smothered by
high densities of the non-native species.
Ricciardi, Neves & Rasmussen (1998) estimated the decline period for various lakes and rivers in
North America. They show that after colonisation of Dreissena polymorpha to Lake Erie, Lake
Oneida and Lake Wawasee decline of the native mussels happened within 4 years, whereas in
Lake St Clair and Detroit River decline took eight years. They noted that most of the Great
Lake/St Lawrence and Hudson River species were not endemic, thus the losses do not represent
global extinctions. However, as already noted the Mississippi river system, could lose 60
endemic species. Of these species c. 65% only exist in large channels and mainstem channels (38
species) which are those parts of the river system which are most vulnerable to invasions of
Dreissena polymorpha. However, although the smaller channels may not be as vulnerable to the
invasion of D. polymorpha, Ricciardi et al (1998) point out that the fragmentation of these
populations in these channels may increase the likelihood of extinction from other anthropogenic
effects such as regulation of flow after impoundments.
Ricciardi et al (1998) also point out that as survivorship of the glochidia is already low and that
the larval dispersal is restricted by dams (Williams et al. 1992), the recovery potential through
either immigration or recolonisation for the large Unioids is reduced.
Based on the known losses of freshwater mollusc species, Ricciardi et al. (1998) provide estimates
of projected extinction curves, one extrapolating that without the Zebra Mussel the species loss
would have equated to 1.2% per decade, but with the Zebra Mussel the rate reaches 12% per
decade. However, when the number of functionally extinct species are taken into account the
predicted species loss, without the Zebra Mussel increases to 4.2% per decade. This led Ricciardi
et al. (1998) to conclude that the estimates of 12% per decade, are in fact conservative, as other
factors such river regulation are not diminishing, thus unless major river restoration programmes
are undertaken, these species are in grave danger of extinction.
[Insert Figure 1 from Ricciardi ?]
10.2.2. Increase in Medical problems
Recommendations
1) Use stabilisation methods for the riverbed which do not impact on the invertebrate
communities.
2) Protect water supplies including ground water.
3) Protect adjacent riparian forests, preventing development of monocultures, especially those
of alien species.
4) Preserve fish fauna, allowing free movement beyond impoundments.
5) Chesney & Oliver (1998) point out that in-situ conservation which preserves genetic identity
and leads to recognition of local adaptations is preferable to conservation through
translocation as this affects the gene pool and should only be used in extreme circumstances.
Put more funding and effort into at the problems of translocation and captive breeding.
MBS: Material to incorporate, then dump

Thailand has 170 species, with 96 brackish water species recognised, however few of these species
transmit human diseases at present.
Order
Families
Genera
Freshwater species
Brackish water species
Number
6
23
75
170 +
96
Pest species
< 20
Burch & Upatham (1989) list those species which are known to be intermediate hosts, and
indicates their distributions and parasites.
Although there are over 50 species of Pomatiopsidae recognised in the Mekong basin, only two
are though to be intermediate hosts for human parasites (Neotricula species).
MBS: Material to incorporate, then dump

Woodruff D.S. & Upatham E. Suchart (1992) Snail transmitted diseases of medical and vetinary
importance in Thailand & the Mekong Valley Journal of Medical and Applied Malacology, 4: 1
12
Burch, J.B. & Upatham E. Suchart (1989) Medically important mollusks of Thailand Journal of
Medical and Applied Malacology, 1: 1 9.
Hydropower and irrigation projects carry the risk of increasing the local prevalence of snailtransmitted diseases in both humans and livestock. These diseases are caused by two main types
of parasitic helminthes: flukes (Trematoda) and roundworms (Nematoda). Both these groups of
organisms have complex life cycles that involve periods in larval hosts, such as snails and an
adult phase in birds or mammals. These hosts are usually very specific, and the increase in
disease results from decline of the native species, and increase in abundance of the host species,
sometimes replacing the natives as there were alien to that specific catchment area.
Schistomiasis: Schneider et al. (1984) believed that these was an increase in parasite transmission
resulting from proposed impoundments which cahgne river habitat and ecology. This is because
the species which can act as hosts become more prolific as they are adapted to the new
environments whereas other similar species which are not vectors, (usually in the same genus or
closely related species) decline in abundance, and maybe lost to the river system. In Thailand in
1992 the Pak Mun hydropower and irrigation project caused controversy as to whether the blood
flukes which pass on Schistomiasis would become established from the nearest known source in
Laos (Usher, 1991).
Paragonimiasis: The knowledge in 1992, suggested that molluscs were not the source of this
disease in the Mekong basin, as the creation of new habitats has not yet led to the expansion of
the known snail vectors into Thailand. However, there remains a chance that the snails may
become established and impact on the local species of the genus Brotia.
Opisthorchiasis: There are 12 species of the freshwater prosobranch Bithynia recognised in
Thailand, but only the three species in the subgenus Digniostoma transmit Opisthorchis viverrini.
These species can live at very high densities and are usually found in rice fields, canals, ponds
and lakes. River impoundment has led to an increase in the abundance of these three species, and
disease outbreaks were noted at the time of construction of the Ubolratana (or Nam Pong) Dam in
NE Thailand Woodruff & Upatham, 1992).
Echinostomatiasis: Schneider et al. (1984) predicted that proposed impoundments would lead to
an increase in the incidence of infections and increasing disease severity.
Table XX
Human diseases, where Molluscs act as intermediate hosts, which may be increasing as the result
of impoundation in Thailand (Data abstracted from Woodruff & Upatham, 1992)
Disease
Schistomiasis
Blood-Fluke:
Schistoma japonicum
Paragonimiasis
Paragonimia heterotremus
( N.B. P. westamani has not
been recorded from humans in
the Mekong basin yet)
Intermediate hosts
Molluscs:
Oncomelania spp.
Neotricula spp.
Source of Information
Molluscs:
Brotia asperata
Melanoides tuberculata
Tarebia granifera
Opisthorchiasis:
Molluscs:
Opisthorchis viverrini
Bithynia funiculata
Bithynia siamensis
Bithynia goniomphalos
Echinostomatiasis
Numerous Echinostome
species
Molluscs:
Various hosts including Pila
ampullacea, P. polita, Radix
rubiginosa, Indoplanorbis
exustus, Filopaludina spp.

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Contributing Paper

Molluscan Biodiversity and the

Prepared for Thematic Review II.1:

For further information see http://www.dams.org/

Report to IUCN Species Survival Commission

Arthur Bogan (North Carolina State Museum), Philippe Bouchet

Table A1 Summary Statistics on Molluscan Biodiversity

Boss (1973), van Bruggen (1995)

Boss (1973), van Bruggen (1995)

Boss (1973), van Bruggen (1995)

Importance of Molluscs to Ecosystem

Figure xx Hypothetical size distributions for recruiting and senescent

LIFE CYCLE STAGES AFFECTED

Fertilisation, glochidial infection, settlement,

Catchment management and river restoration

Reviewing and changing agricultural practices to reduce eutrophication.

These will involve raising public (=involved organisations) awareness.

Figure 2 Diagram showing the proportion of molluscan extinctions as a total of

Picture of Africa, with species diversity across Africa]

1.4.1.1 USA- RIVERS

Table 5 Threatened Status of North American freshwater gastropods

1.4.1.2. USA- SPRING SYSTEMS

area S2: Florida, USA

1.4.2.1. SE ASIA: MEKONG RIVER

1.4.3.1 NORTHERN WESTERN GHATS, INDIA

area L9: Chilka Lake [brackish water], India

area L10: Inl_ Lake, Myanmar

1.4.4. SOUTH AMERICA

1.4.4.2 SOUTH AMERICA: URUGUAY & LA PLATA RIVER

1.4.5.1 AFRICA: SOUTH OF THE SAHARA

Lake Kariba: Zambezi River. (Brown, 1994)

area R9: Madagascar

area L4: Lake Tanganyika, Burundi-Democratic Republic of Congo-Tanzania-Zambia

area L6: Lake Victoria, Kenya-Tanzania-Uganda

1.4.6. EUROPE (INCLUDING FORMER SOVIET UNION)

area S4: Balkans region, former Yugoslavia-Austria-Bulgaria-Greece

area L7: Lake Ohrid and Ohrid basin, Albania-Former Yugoslavia

1.4.7. AUSTRALIAN FRESHWATER MOLLUSCS

area S6: Western Tasmania, Australia

1.4.8. SE ASIA: NEW CALEDONIA & SULAWESI

area S7: New Caledonia

area L9: Chilka Lake [brackish water], India

area L10: Inl_ Lake, Myanmar

2.0. UPSTREAM IMPACTS

2.1. Movement of species upstream

4.0. RESERVOIR IMPACTS

4.1. Site impacts (effects on river environment & biota at site)

Table 5.0.2 Mollusca Species present within reservoir region, USA

5.1. Seasonality of flow

5.2. Change in flow regime

Woodruff & Upatham (1992)

been recorded from humans in

Woodruff & Upatham (1992)

Woodruff & Upatham (1992)

9.0. PRIORITY OF IMPACTS

10.2.2. Increase in Medical problems

MBS: Material to incorporate, then dump

MBS: Material to incorporate, then dump

Woodruff & Upatham (1992)

Woodruff & Upatham (1992)

Woodruff & Upatham (1992)

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