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Rediscovery of Branchipus schaefferi (Branchiopoda: Anostraca) in Belgium - Notes on habitat requirements and conservation management

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Fairy shrimps (Crustacea, Anostraca) are specialized inhabitants of inland water bodies that periodically dry or freeze over. Here we report the first observation since 1997 of a member of this basal crustacean order in Belgium and the first sighting of the species Branchipus schaefferi Fischer, 1834 since 1930. Nineteen populations were found in a restricted area located 55 km sE of Brussels in the Province of Hainaut. Based on a field survey, we discuss the habitat characteristics of these populations. We discuss also the distribution and habitat requirements of the species based on literature and formulate a number of guidelines for the conservation of this species as well as other large branchiopods in densely settled areas with intensive agriculture such as Belgium. Finally, we formulate a number of likely explanations for the lack of recent observations of these organisms in Western Europe and in Belgium.
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Belg. J. Zool., 143 (1) : 3-14
January 2013
Rediscovery of Branchipus schaefferi (Branchiopoda: Anostraca) in
Belgium - notes on habitat requirements and conservation management
Bram Vanschoenwinkel
1, 3,*
, Luc Brendonck
1
, Tom Pinceel
1
, Pascal Dupriez
2
&
Aline Waterkeyn
1, 3
1
Laboratory of Aquatic Ecology, Evolution and Conservation, KU Leuven, Charles Deberiotstraat 32, 3000 Leuven,
Belgium.
2
Natagora Centre-Ouest Hainaut, Rue Marécaux 5, 7333 Tertre, Belgium.
³ BINCO (Biodiversity Inventory for Conservation), Rijmenamsesteenweg 189, 3150 Haacht, Belgium
.
*
Corresponding author: bram.vanschoenwinkel@bio.kuleuven.be
ABSTRACT. Fairy shrimps (Crustacea, Anostraca) are specialized inhabitants of inland water bodies that
                  
          Branchipus schaefferi Fischer, 1834 since
1930. Nineteen populations were found in a restricted area located 55 km SE of Brussels in the Province of

distribution and habitat requirements of the species based on literature and formulate a number of guidelines
for the conservation of this species as well as other large branchiopods in densely settled areas with intensive
agriculture such as Belgium. Finally, we formulate a number of likely explanations for the lack of recent

: fairy shrimp, temporal pools, wheel tracks, conservation management
INTRODUCTION
Fairy shrimps and brine shrimps together make

clam shrimps (Spinicaudata, Laevicaudata
and Cyclestherida) and tadpole shrimps
(Notostraca) they are often referred to as large
branchiopods. Together with a number of extinct
orders they form the class Branchiopoda. Due
to their size, large branchiopods are sensitive
     
occur in aquatic habitats that cannot sustain
     Jeppesen
et al., 2001), either because they are highly
saline, periodically dry or regularly freeze solid.
Brine shrimps (Artemia and Parartemia spp.)
and some members of the fairy shrimp genera
Branchinella, Branchinectella, Branchinecta
and Phallocryptus are typical for salt pans in
different parts of the world. Most fairy shrimp,
on the other hand, generally inhabit temporary
freshwater habitats, ranging from large wetlands,
vernal ponds and marshes to rock pools, wheel
      
Although the class Branchiopoda is an old group
with a near global distribution (
Brendonck et al.,
2008), Belgian records of fairy shrimp and other
large branchiopod populations are extremely
scant. Museum collections in Brussels and Liège
were inventoried by Brendonck (1989a) and
Loneux & Thiéry (1998), respectively, revealing
a historic species richness of seven. These include
three fairy shrimp species: Chirocephalus
diaphanus Prevost, 1803; Eubranchipus
(Siphonophanes) grubii Dybowski, 1860 and
Branchipus schaefferi Fischer, 1834; two clam
shrimp species: Limnadia lenticularis Linnaeus,
1761 and Leptestheria dahalacensis (Rüppel,
1837) and two tadpole shrimp species: Lepidurus
apus Linnaeus, 1758 and Triops cancriformis
Bosc, 1801.
The fairy shrimp C. diaphanus is a widespread

4 Bram Vanschoenwinkel, Luc Brendonck, Tom Pinceel, Pascal Dupriez & Aline Waterkeyn
and North Africa with a range extending
northward into Great Britain and eastward to
the Black Sea. In Belgium, C. diaphanus was
found in Halen in 1903 (Brendonck, 1989A)
and in Sint-Truiden in 1930 (
Loneux & Thiéry,
1998) and was last seen in Hamois in 1998
(
Loneux & WaLravens, 1998). Eubranchipus
(Siphonophanes) grubii is a Central and Eastern
European coldwater species and may have
been observed in Belgium in 1970 in Geel
(
k. WouTers, pers. comm. de Brendonck,
      
Currently, the nearest populations are located
in The Netherlands (North Brabant, Gelderland,
Overijssel and Limburg) (
Brendonck, 1989A;
soesBergen, 2008), Germany (Rhineland-

Maier, 1998;
engeLMann & hahn, 2004) and France (Alsace)
(
defaye et al., 1998). Branchipus schaefferi
is a eurytherm species that is widespread in
Europe and around the Mediterranean basin
with additional records from Northern Africa
and Asia (
BrTek & Thiéry, 1995; aL-sayed &
Z
ainaL, 2005) and was encountered only once
in Belgium (Sint-Truiden, 1930) (
Loneux &
Thiéry, 1998). The closest currently-known
     
 
(
Maier, 1998; engeLMann & hahn, 2004) and
      
(Alsace region) (
defaye et al., 1998). The clam
shrimp L. lenticularis is a Holarctic species
most abundant in northern temperate climates.
In Belgium, it was reported from a marsh in
Genk in 1946 and Zolder-Zonhoven in 1959
(
Brendonck, 1989A). Leptestheria dahalacensis
(Rüppel, 1837) was encountered in Belgium
      
Brussels, presumably having been introduced
inadvertently with mud from temporary ponds
in carp nurseries in Eastern Europe (
Brendonck
et al., 1989B). The tadpole shrimp L. apus is
widespread in Europe and has been reported in
Belgium from Balen or Halen (date unknown),
but nothing else is known about the population
(
Brendonck, 1998). The second tadpole shrimp
species, T. cancriformis, is also widespread in
Europe and historically known from Halen (in
1892 and 1903), Sint Truiden (in 1917, 1929
and 1930), Ferrières (in 1905 and 1906) and
Leuven (date unknown) (
Brendonck, 1998;
Loneux & Thiéry    
currently-persisting population in Belgium is a
T. cancriformis population discovered in 2006 on
a military domain in Brasschaat in the province
of Antwerp (
WiLLeMs & de Leander, 2006).
Despite the relatively intensive monitoring of
many aquatic habitats, fairy shrimps have not
been observed in Belgium since 1998 and clam
shrimps have not been seen since 1989.
In this paper we report the rediscovery of the
order Anostraca in Belgium represented by at
least 19 populations of B. schaefferi, which has
       
autecology of the species based on published
literature and the biotic and abiotic characteristics
of the habitats in which it was found. Based on
this we formulate guidelines for more effective
detection, monitoring and conservation of
this species and other large branchiopods in
intensively-developed regions such as Belgium.
MATERIALS AND METHODS
Notes on the ecology and distribution
of the species
Branchipus schaefferi (Fig. 1) is a eurythermic
species that can be found from late spring until fall
in temperate regions (
hössLer et al., 1995; eder
et al., 1997) and in Southern France (
defaye et al.,
1998;
WaTerkeyn et al., 2009) or throughout the
year in warmer regions around the Mediterranean
basin such as Morocco (
BeLk & BrTek, 1995;
BrTek & Thiéry, 1995; Thiéry, 1987; Marrone
& M
ura, 2006). It is most frequently found in
small shallow ponds, puddles or wheel tracks
with turbid water and scarce vegetation (
hössLer
et al., 1995;
peTrov & peTrov, 1997; defaye
et al., 1998;
Boven et al., 2008). The species
can also be found in other habitat types, such
   
peTkovski, 1997; Mura,
2001) and mountain habitats (
eder et al., 1997;
defaye et al., 1998; Mura, 1999; Thiéry, 1987).
5
Branchipus schaefferi rediscovered in Belgium
Fig. 1. – Distribution of the discovered Branchipus schaefferi populations in Hainaut (A)
with thin and thick black lines representing unsealed and sealed roads, respectively. Filled
symbols represent wheel track populations, the empty symbol corresponds to a population in
a farmland pond. (B) Overview of the only two records of this species in Belgium, indicating

Truiden (†). The middle panel shows a typical B. schaefferi wheel track habitat near Binche
(Picture: B. Vanschoenwinkel) (C) and a close up of an adult male showing the antennal

isolate of a wheel track zooplankton community including many fairy shrimps. Females can
be discerned based on the presence of a blue brood pouch (Picture: B. Vanschoenwinkel) (E).
6 Bram Vanschoenwinkel, Luc Brendonck, Tom Pinceel, Pascal Dupriez & Aline Waterkeyn
Exceptionally, it can occur in permanent waters,
as is the case in Germany (hössLer et al., 1995).
It is considered a rather tolerant species, since it
can survive in ponds with short inundations, high
turbidities (Thiéry, 1987), high conductivities
(up to 4500 µS cm
-1
) (WaTerkeyn et al., 2010),
high temperatures (Marrone & Mura, 2006),
eutrophication due to cattle manure (Thiéry,
1987) and high altitudes (up to 2600 m a.s.l.)
(Thiéry, 1987).
Dormant eggs of Branchipus schaefferi
hatch within 1 to 6 days after inundation, while
maturation takes 7 to 30 days, depending on the
temperature and food conditions (
WaTerkeyn
et al., 2009 and references therein). They can
survive for up to 2.5 months (hössLer et al.,
1995; BeLadJaL et al., 2003) and grow up to
20-25 mm (Thiéry, 1991; hössLer et al., 1995;
peTkovski, 1997; defaye et al., 1998; aL-sayed
& ZainaL, 2005). The females have a brightly-
turquoise colored brood sac and can produce up
to 242 dormant eggs per day (maximum reached
21 to 27 days after hatching) (BeLadJaL et al.,
2007). Eggs are typically angular and wrinkled
and more or less spherical ranging from 195 to
290 µm in size (Thiéry et al., 1995). Different
life stages of B. schaefferi can co-occur, probably
due to several hatching peaks triggered by
additional rainfall (hössLer et al., 1995; peTrov
& peTrov, 1997; aL-sayed & ZainaL, 2005).
Branchipus schaefferi often co-exists with one
or several other large branchiopods, such as
the notostracan T. cancriformis (most often
reported), spinicaudatans (Imnadia yeyetta,
I. banatica, L. dahalacensis or L. saetosa),
or other anostracans (Tanymastix stagnalis,
Branchinecta ferox, Streptocephalus torvicornis,
C. diaphanus, C. carnuntanus, C. brevipapis)
(peTrov & cveTkovic, 1997; peTkovski, 1997;
defaye et al., 1998; Maier et al., 1999; Marrone
& Mura, 2006; Boven et al., 2008; WaTerkeyn
et al., 2009).
Study site and sampling protocol
       
branchiopod-like crustacean in the area in 2002
(Dupriez, pers. com.), populations of the species
were discovered in a wheel track North of Binche
(province of Hainaut; Belgium) on 23 July 2012
by Pascal Dupriez during a survey for natterjack
toads (Bufo calamita). The observation was
reported as a potential sighting to the KU Leuven
nationwide large branchiopod survey (http://bio.
kuleuven.be/de/dea/branchiopodhunters/index.
php). Thirty other potential habitats in that area
were surveyed on 25, 26 and 27 July 2012. These
included the two types of temporary ponds present
in the area: wheel tracks as well as a couple
        
Several wheel tracks were dry so the presence of
fairy shrimp in these could not be determined in

populations we used conservative criteria. A set of
tracks that showed obvious signs of connections
     
was considered as a single habitat potentially
housing a single population. Proximate tracks
were only considered as separate habitats if they
were independent depressions separated by a
clear topographic barrier. In total, water quality
variables were measured in 22 habitats (19 of
which contained B. schaefferi). Measurements
were taken between 11.00 and 15.00 under
sunny conditions and included conductivity
(EC; µS cm
-1
), water temperature (T; °C), pH,
oxygen concentration (DO; ppm), total dissolved
solids (TDS; ppm), and total suspended solids
(TSS; ppm) using a HI9828 Multiparameter
Meter (Hanna instruments, Ann Arbor, MI,
USA). Chlorophyll-a concentration (ChlA; mg
L
-1
) and turbidity were determined using a hand
    
Sunnyville, CA, USA). Nutrient concentrations
   
using a Hach DR2400 spectrophotometer (Hach
company, Loveland, CO, USA) by means of the
following methods: total N (persulphate digestion
method), total P (acid persulphate digestion
method using PhosVer® 3), reactive phosphate
(PhosVer® 3 method) and Nitrate (chromotropic
7
Branchipus schaefferi rediscovered in Belgium
acid method). The bottom of many wheel track
habitats was partly or almost entirely covered
with gravel. The gravel coverage (± 10%)
was estimated and included as an additional
predictor variable in the analyses. Habitat size
was assessed by measuring length, width, max
depth and volume calculated using the formula
for the volume of a half ellipsoid. In case pools

the volume was calculated as two separate half
ellipsoids. An aquarium net (mesh 0.5 mm)
was initially used to qualitatively check for the
presence of B. schaefferi. In order to quantify
density of fairy shrimp (number of individuals
per L), quantitative zooplankton samples were
taken by scooping a total of 12 L of water using
         
zooplankton net. If fairy shrimp densities were

stored in 90% non-denatured ethanol. Total
population sizes were obtained by multiplying
densities with calculated water volumes.
Analyses
The relationship between measured
environmental variables and fairy shrimp
density was analysed using multiple linear
regression. Due to large variation in fairy shrimp
densities including several outliers, analyses
were performed using density ranks rather than
the untransformed data. This transformation
helped to meet the linear regression assumption
of homoscedasticity. In order to reduce the set
of predictor variables, only variables with a
clear trend of association (Spearman correlation

variable were included in a multiple regression
model. Both stepwise forward and backward
selection procedures were used in order to remove


adjusted r² and Akaike’s information criterion
as decisive factors. First order interactions were
also considered. Associations between fairy
shrimp densities and measured environmental
variables were visualised using principal
component analysis (PCA) triplot. This plot
shows the relative positions of different habitats
along the two dominant axes of environmental
variation (PC1 and PC2) while simultaneously
showing the associations between habitats and
environmental variables (shown as vectors) and
associations between environmental variables.
Environmental variables were centered and
standardised prior to analyses. In order to obtain
an objective representation of the environmental
variation and its relationship with the response
variable of interest (fairy shrimp density), the
latter was plotted as a supplementary variable
that does not affect the ordination (Legendre &
Legendre, 1998). All analyses were performed
in Statistica 10 (Statsoft 2011, Tulsa, OK, USA).
RESULTS
B. schaefferi was found in a total of 18 wheel
tracks present in a local network of unsealed
roads in a rural area covering a total area of
approximately 7 km² North of Binche in the
Province of Hainaut. These roads were separated

wheat) by an elevated ridge of approximately 1.5
m wide and 30 cm high, preventing excessive

the species was found in a single temporary pond

area. Most habitats housed large populations
(Average: 4315; Range: 1 - 44000) of adults of
both sexes. The environmental characteristics of
the different B. schaefferi habitats are provided
in Table 1. In general, habitats were relatively
shallow, turbid and lacked aquatic vegetation.
      
Macroinvertebrate species richness was relatively
low in all habitats. Besides B. schaefferi, the
only branchiopod crustacean present was Moina
branchiata. Other inhabitants included at least
one ostracod species, water bugs (Corixidae),
      
several dipterans (Chironomidae, Culicidae,
Eristalis sp.). Principal components analysis was
used to visualize associations between densities
of B. schaefferi and environmental variables. The
8 Bram Vanschoenwinkel, Luc Brendonck, Tom Pinceel, Pascal Dupriez & Aline Waterkeyn

of total variation. The triplot suggests a positive
association between fairy shrimp population
density and reactive phosphate, and negative
associations with pH, gravel coverage, nitrate
and chlorophyll a (Fig. 2). However, gravel
      
was retained in the constructed regression models
associated with lower fairy shrimp densities
(Fig. 3). Associations with other environmental

DISCUSSION
The current study shows that B. schaefferi is
still present in Belgium after not being reported
    
of fairy shrimp in Belgium since C. diaphanus
was last detected in Hamois in 1998 (
Loneux &
WaLravens, 1998). Observation of B. schaefferi
in summer during a warm period and after heavy
rains is consistent with the known phenology of
this heat-tolerant eurythermic species (defaye et
al., 1998). Most of the remaining populations of
TSS
Fig. 2. – PCA triplot showing associations between
environmental variables (vectors), site scores (circles)
and the supplementary variable Population density
(arrow). Environmental variables were centered and
standardised prior to analysis. Population densities
were transformed to ranks.

relationship between the percentage of surface area
of each habitat covered with gravel and the density of
fairy shrimp found in the active populations during
sampling.
B. schaefferi
known from wheel tracks (Boven et al., 2008;
defaye et al., 1998), probably since these are the
most commonly-remaining temporary aquatic
   
landscapes. The species is well adapted to time
stress, displaying traits such as a short life cycle
and high fecundity (hössLer et al., 1995; peTrov
& peTrov, 1997; defaye et al., 1998). Therefore
it is well adapted for living in these short-lived
temporary aquatic systems, which often hold
water for only a couple of weeks.
In general, population densities in most of
the studied habitats were high (Average: 2.18
± 4.1 ind./L; Range: 0.01-18) suggesting that
these populations are well established. Although
nutrient concentrations were moderate to high,
chlorophyll a concentrations were low indicating
that this could be top-down controlled by the
grazing zooplankton community. Freshwater
zooplankton communities, and fairy shrimp in
particular, can be sensitive to pesticides or to
oxygen stress as a result of eutrophication (
Lahr,
1998; rogers, 1998). As such, the fact that
populations were doing well despite the presence
of intensive agriculture in the immediate vicinity
could illustrate that the buffer zones (elevated
ridge covered with grasses, herbs and small
shrubs) that are present between the wheel
9
Branchipus schaefferi rediscovered in Belgium
Variable Average ± st. dev. Range
Surface (m²) 32.4 ± 75.0 0.2 - 314
Depth (cm) 7.8 ± 3.8 2 - 15
Conductivity (µS/cm) 573.4 ± 274.6 133 - 1335
Dissolved oxygen (ppm) 38.6 ± 1.47 6.20 - 1.35
pH 8.10 ± 0.30 7.69 - 8.57
Temperature (°C) 32.1 ± 2.3 27.73 - 35.7
Total Dissolved Solids (ppm) 286.9 ± 137.2 67 - 668
TSS 222.8 ± 205.0 34.0 - 873.7
Chl A (mg/l) 0.011 ± 0.007 0.002 - 0.03
Total N (mg/l) 4.92 ± 3.45 0 - 10.4
Nitrate (mg/l) 4.51 ± 2.49 0 - 9
Total P (mg/l) 2.01 ± 0.76 0.68 - 3.62
Reactive phosphate (mg/l) 1.06 ± 0.78 0.22 - 2.7
TABLE 1
Environmental variables measured in the ponds containing B. schaefferi (n = 19).
      
However, the presence of a large population in
         
runoff suggests that the species may, in fact,
be quite resistant, as was also suggested by
Thiéry (1987). The species, however, remains
vulnerable to habitat destruction. In many areas
        
  
that in the studied area fairy shrimp population
densities were much lower in habitats with ample
gravel coverage. This effect can have different
origins. Gravel application can reduce the depth
and potential length of inundations (hydroperiod)
of the habitat or alter water chemistry and make
them less suitable for fairy shrimp. However, we
found no indications for associations between
gravel coverage and water levels or any of the
measured environmental variables (Spearman R;

purely physical nature. Gravel application can,
for instance, cover the dormant egg bank and
shield fairy shrimp resting eggs from receiving
hatching cues, such as light, impeding successful
recruitment. Filling of roadside ditches
presumably also led to the disappearance of the
last known Belgian population of C. diaphanus in
Hamois (
Loneux, pers. com.; vanschoenWinkeL,
pers. obs.). Consequently, it is advisable to
refrain from adding additional gravel, debris
or other sediments to existing wheel tracks if
these populations are to be preserved. In some

wheel tracks become too deep to allow passage
of vehicles we propose that they should not be
      
of about 10 cm of standing water after rains, as
was the situation observed for the fairy shrimp
populations in this study. Ideally, the top layer
surface sediment (± 4 cm) should be temporarily
removed prior to graveling and replaced on top of
the gravel to ensure that the resting egg bank will
not be covered and fairy shrimp may continue
to hatch during future inundations. Restricting
access of vehicles altogether is probably not
recommended as the disturbance provided by
passing vehicles is necessary to maintain these
wheel tracks. Additionally, previous research has
shown that walkers and motor vehicles can be
important dispersal vectors for large branchiopod
crustaceans (
WaTerkeyn et al., 2010). Regular
exchange of eggs between populations may
ensure healthy metapopulation dynamics
with recolonization rates compensating for
10 Bram Vanschoenwinkel, Luc Brendonck, Tom Pinceel, Pascal Dupriez & Aline Waterkeyn
occasional extinctions. The spatial organization
of the populations in this study located along an
unsealed road network could be illustrative of

using genetic analyses.
Relict populations or products of a recent
introduction?
      
genetic analyses, it is plausible that the
discovered populations represent relicts, rather
than a recent establishment of the species. First
of all, the presence of the species is consistent
with the species’ distribution and its historic
presence in Belgium (
Loneux & Thiéry, 1998).
Currently known populations are present in

(Rhineland-palatinate) at about 200 to 300 km
from the Belgian locality (Loneux & Thiéry,
1998). Secondly, the occurrence of a substantial
number of populations, usually consisting
of numerous individuals, suggests that the
populations are likely to have been in the area for
at least several decades. Finally, the fact that the
populations were found in an old agricultural area
with unsealed roads that are probably more than
100 years old, makes continuous and prolonged
persistence of the species in the area a likely
scenario. An upcoming phylogeographic study
across the species’ range (including specimens
from this study) documenting the phylogenetic
relationships among the remaining European
lineages, will likely provide more conclusive
evidence concerning the origin of the Belgian
populations.
A hidden existence
The current study illustrates that populations
of fairy shrimp can remain undetected, although
individuals are relatively large (1 - 4 cm) and
conspicuous and often characterized by bright
coloration, and even in relatively well-studied
and monitored regions, such as Belgium. The
reasons for this are manifold. First of all, fairy
shrimp and other large branchiopods are typically
     

cues (
Brendonck, 1996). If such cues do not
present themselves, it is common that years will
go by without active populations developing
   WaTerkeyn et al., 2009). This is
possible since they produce long-lived resistant,
dormant eggs. For instance, the most common
    C. diaphanus and
E. (S.) grubii, are usually only present during
the colder winter months and the beginning of
spring (defaye et al., 1998), at a time when there
is typically no monitoring. Secondly, even when
eggs hatch and adults develop, they can easily
remain un-noted as fairy shrimps often inhabit
small and inconspicuous systems, such as wheel
tracks and puddles in meadows and cropland
where few people wander. These habitats are
also often considered of low conservation
interest and are therefore not monitored. Thirdly,
water in wheel tracks is often turbid obscuring
potential inhabitants. Finally, active populations
in the water column often only persist for
several weeks as a result of their short life span
and the gradual increase of predation (by e.g.

inundation (spencer et al., 1999).
Towards effective conservation
Large branchiopods are threatened in many

The main reason for this is the loss of temporary
aquatic habitats as a result of intensive agriculture
and urbanisation, and the few remaining habitats
are often degraded (
BeLk, 1998). Although 29
fairy shrimp species are red listed by IUCN, and
several species are included in local red lists
(e.g. in the Alsace), at the moment there is no
legal basis for protection of large branchiopods
in Belgium. Before a species can be red listed

put into localizing and monitoring populations.
Given the hidden existence of the members
of this group, they are typically overlooked in
    
11
Branchipus schaefferi rediscovered in Belgium
therefore recommend that sampling campaigns
should be strategically planned and undertaken

large branchiopods are highest. For instance,
early spring (February, March) and preferably
        
     
cold water species such as C. diaphanus, S.
grubii and L. apus. On the other hand, a summer
drought followed by heavy rainfall presents ideal
conditions for hatching and development of
warm water species, such as B. schaefferi and T.
cancriformis, which can be detected from about
10 days - 3 weeks after inundation.
Due to the frequent disturbance typical of
ephemeral habitats, local populations may
regularly go extinct. Therefore, in order to persist
regionally, dispersal and recolonization from
nearby populations (metapopulation dynamics)
are likely to be important. Promising localities
     
where temporary water bodies are abundant and
have historically been abundant. Although the
local dispersal potential of large branchiopods is
quite high (
vanschoenWinkeL et al., 2008A,B),
successful long distance dispersal (several km)
events are rare (vanschoenWinkeL et al., 2011).
Therefore, recently-formed temporary water
bodies, such as bomb craters and human-made
temporary ponds, may be suitable in terms of
their environmental conditions, but may not be
colonized, even when large branchiopods are
present in the region. Nevertheless, occasionally
isolated relict populations have been detected
(pauLsen, 2000). Finally, over longer time scales,
temporary pond systems typically accumulate
sediments and disappear. Therefore, physical
disturbances that counteract this process can
     
temporary water bodies by wallowing in them
covering their skin with mud (vanschoenWinkeL
et al., 2008B). These turbid, unattractive systems
often hold a large diversity of branchiopod
crustaceans (nhiWaTiWa et al., 2011). In recent
times, many large branchiopod populations
have been found in habitats that are frequently
disturbed by humans, such as wheel tracks (e.g.
Boven et al., 2008). The last remaining Triops
population in Belgium persists in a muddy
track used by tanks and other military vehicles
(WiLLeMs & de Leander, 2006). Similarly,
military domains in Eastern Europe are known
for their diversity of large branchiopods (Maier
et al., 1998). The presence of natural habitat that
was historically set apart, unsealed roads with
puddles and wheel tracks and regular physical
disturbance by vehicles, makes military domains
particularly suitable areas that may be acting as
refuges for temporary pond fauna, such as large
branchiopods.
      
group of crustaceans may be limited, it is
important to realize that temporary ponds not
only house a unique crustacean fauna, but are
also of vital importance for other endangered
species of plants and animals (
WiLLiaMs, 2006).
    
    
    
support have been directed at protecting certain
endangered amphibians that use temporary
ponds for breeding, such as the natterjack
toad (Bufo calamita    
toad (Bombina bombina). Temporary pond
restoration and construction projects performed

Bombina       
typical temporary pond organisms too. For
instance, different rare macrophytes were shown
to re-emerge from old seed banks during pond
restoration projects (hiLT et al., 2006). Due to
the prolonged viability of their dormant eggs
(Brendonck, 1996), it is not unlikely that large
branchiopods may emerge from old egg banks
present in the sediment. Consequently, a habitat-
oriented conservation strategy protecting the
few remaining high quality temporary ponds
and increasing temporary pond densities in the
        
large number of organism groups, including

landscape dominated by agriculture, the use of
vegetation buffer zones and ridges is likely to

12 Bram Vanschoenwinkel, Luc Brendonck, Tom Pinceel, Pascal Dupriez & Aline Waterkeyn
pesticides (decLerck et al., 2006), even though
some species such as B. schaefferi may be quite
tolerant. Finally, we would also encourage the
re-evaluation of marginal aquatic systems such
as wheel tracks as they may contain unique biota,
such as B. schaefferi.
CONCLUSIONS
This paper reports the rediscovery and the
    B. schaefferi in
Belgium and analyses the link between habitat
characteristics and population densities. For the
studied wheel track populations it was shown
that extensive gravel application was associated
with lower fairy shrimp population densities,
suggesting that this practice should be avoided
if populations are to be preserved. In terms of
conservation management, we conclude that
a habitat-oriented approach preserving natural
processes of desiccation and disturbance is likely
to be most effective for the conservation of fairy
shrimp as well as other typical temporary pond
organisms.
ACKNOWLEDGEMENTS

  
project 3E110799. Bram Vanschoenwinkel and
  
      
to thank Liselore Vanstallen, Falko Buschke
and Bernard Loison for valuable assistance
       
Marcel Moncousin, Marius Loison, José Godin
and André Pourtois, who made the initial
observations in 2002 hinting at the potential
presence of anostracans in the region.
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Received: September 28th, 2012
Accepted: February 2nd, 2013
Branch editor: Schön Isa
... Many pioneer species of aquatic insects and other invertebrates specialize in early successional stage habitats, found in seasonal or newly created water bodies. These pioneer species mostly prefer open sunny habitats without aquatic or riparian vegetation (Ruhí et al., 2009;Vanschoenwinkel et al., 2013;Sroka et al., 2016). As the habitats age, these pioneer species are gradually replaced during the succession by other taxa including generalists and species requiring more densely vegetated water bodies (Corbet, 2004;Kadoya et al., 2004). ...
... Early successional habitats are often disproportionately hit by human activities as they are highly susceptible to eutrophication and direct habitat loss, e.g., by the channelization of rivers. Many pioneer species are thus considered rare or endangered (Harabiš et al., 2013;Vanschoenwinkel et al., 2013), and the availability of suitable manmade habitats may contribute to their conservation (Harabiš and Dolný, 2015;Kolář et al., 2017;Vojar et al., 2016). A prime example of such habitats in Central Europe and elsewhere are ponds in sand and gravel pits in alluvial sediments, which are created or spontaneously form after sand and gravel mining (Rademacher, 2011;Ř ehounková and Prach, 2008;Vebrová et al., 2018). ...
Article
Man-made freshwater habitats are an important part of the European landscape, especially in areas with mostly absent or degraded natural habitats. To assess the role of different man-made standing waters in anthropogenic landscapes, we surveyed adult odonate communities in a cluster of 20 water bodies including fishponds and sandpit ponds in early and ongoing successional stages. We found 35 odonate species (i.e., 47% of the fauna of the Czech Republic), but their presence differed significantly among the three habitat types. The highest species diversity, driven mainly by the presence of generalists, was found in fishponds. Sandpit ponds in an early suc-cessional stage hosted the least diverse communities dominated by pioneer and vagrant species. Specialist species occurred in both types of sandpit ponds, especially those in an ongoing successional stage, more than in fish-ponds. Although the dragonfly biotic index did not differ among the three types of localities, all four species from the national Red list recorded during the study occurred only in sandpit ponds. The main environmental drivers of local odonate communities included the coverage of shoreline by emergent vegetation, water depth and bottom substrate; the latter two characteristics largely corresponded to the distinction between sandpit ponds and fishponds. We conclude that both sandpit ponds and fishponds play an important role in maintaining freshwater biodiversity that requires a mosaic of habitats in different successional stages.
... Persistence and reproduction in ephemeral ponds are independent of population density and depend on the timing of the hydroperiod and the organisms' life cycle [39]. A winter temperature increase may reduce both the developmental time of crustaceans and hydroperiod, due to evaporation [35,81]. In this scenario, if the hydroperiod shortens much more than the life cycle, the result will be a demographic decline with a depletion of the egg bank proportional to the hatching rate of the resting eggs. ...
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Organisms respond to climate change in many different ways and their local extinction risk may vary widely among taxa. Crustaceans from freshwater temporary ponds produce resting eggs to cope with environmental uncertainty and, as a consequence, egg banks have a fundamental role for population persistence. The egg bank dynamics of six clonal lineages of Heterocypris incongruens (Ostracoda) from Northern Italy were simulated. Clonal lineages W1 and W2 are the most common “winter ecotypes”, clonal lineages S1 and S2 are allochthonous “summer ecotypes” and clonal lineages I1 and I2 are relatively rare and generalist in terms of seasonality. Fecundity and proportion of resting eggs vary by clonal lineage, temperature and photoperiod. The clonal extinction risk was estimated in present climate conditions and under climate change. For comparison, and to assess the potential colonization of northern ponds, clonal lineages from Lampedusa Island (Southern Italy), L, were considered. Cohen’s general model was used for simulating egg bank dynamics and the extinction rate of each clonal lineage was estimated with uncertainty analysis. A 30 year simulation in present and climate change conditions was carried out. Extinction rates were lower in climate change conditions than in present conditions. Hydroperiod, hatching rate and egg deterioration rate were the critical factors that affected extinction rates. Extinction rates varied among clonal lineages. This suggests that H. incongruens might be able to have multiple responses to climate change due to its genetic diversity. In climate change conditions, W clonal lineages underwent a niche expansion, while a mismatch between photoperiod and hydroperiod might generate a detrimental effect on the phenology of summer S clonal lineages that might cause their extinction. Southern clonal lineages L, showing an intermediate extinction rate, might colonize northern temporary ponds.
... We found that the animals tolerate wide variations in temperature (ranges 1 to 39 °C), and survival is possible only during short-term exposure to 40 °C which was registered as maximum temperature tolerance by B. schaefferi. The highest recorded natural water temperature is ~35 °C in relatively shallow habitats (Vanschoenwinkel et al., 2013). Nauplius, 29: e2021029 In general, a reduction in ambient temperature is immediately followed by a decrease in muscular performance in Daphnia O.F. Müller, 1785. ...
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The behavioural responses of Branchipus schaefferi males and females to short-term thermal stresses in six different rearing conditions are studied. The animal’s performance was tested in pure tap water and water collected from males and females culture medium. The animal's behaviour was recorded using a high-definition digital video camera mounted approximately 30 cm above the experimental containers. The swimming track and the thoracopods beating rates were recorded manually for each animal at different temperatures. The results indicated that in most cases, a significant increase in active swimming and limbs beating rate with increasing temperature. However, the animals tested in the culture medium were significantly less active compared to those in tap water under similar thermal stress. Animals seem to produce substances in culture media that influence their locomotor behaviour under thermal stress.
... Both factors were negatively correlated to the model output. Actually, persistence and reproduction depend on the timing of the organims' life cycle and the pond hydroperiod (Pyke, 2005;Vanschoenwinkel et al., 2013;Brendonck et al., 2015). If the hydroperiod is shorter than the life cycle, the result will be a demographic decline with a depletion of the egg bank proportional to the resting eggs hatching rate. ...
Article
Resting life stages (e.g. dormant seeds and resting eggs) have important implications for ecological and evolutionary processes. In this study, we simulated the impact of different environmental scenarios on the dynamics of resting eggs that make up an “egg bank” of a common fresh water ostracod, Heterocypris incongruens (Crustacea). Our goal was to investigate how the persistence and the wind-mediated spatial distribution of the species in vernal temporary ponds on Lampedusa Island (Southern Italy) were affected. A general model for selection on seed germination in unpredictable environments was used to simulate within pond egg bank dynamics. Metapopulation dynamics were simulated using Levin's and Hanski's models assuming three generalized spatial patterns of pond distribution (random, aggregated along the main wind direction, evenly spaced along the main wind direction) and two dispersion processes (random walk and wind shear). We applied global sensitivity and uncertainty analysis (GSUA) to the models. We estimated the egg bank growth rate based on 30-year simulations under present climatic conditions, and assuming a 2°C rise in winter temperature under global climate change. Hatching rate and deterioration rate were the most important input factors for the dynamics of the egg bank. In warmer winter conditions, the probability that a pond water balance is positive, a reliable hydroperiod estimation, was the most important factor in the egg bank simulation dynamics. Regular distribution of ponds along the wind gradient and wind shear, had the highest dispersal and colonizing potential, considering the percentage of empty ponds reached (60 %). Levins’ model predicted that the equilibrium varied between 0 and 8 % of colonized ponds while Hanski's model predicted values between 0 and 20 %. In Hanski's model the rescue effect increased the probability of occupied ponds. Potential colonizing resting eggs and extinction rate were positively and negatively correlated to the percentage of colonized ponds, respectively. Our simulations can be generalized to aquatic invertebrate taxa that inhabit temporary ponds, have egg banks, colonize habitats using few propagules and disperse passively by wind.
... Our results have further implications regarding the importance of anostracans as IG predators for the entire community. Anostracans can reach high densities in their habitats (Daborn, 1977;Horváth, Vad, Vörös, et al., 2013b;Vanschoenwinkel, Brendonck, Pinceel, Dupriez, & Waterkeyn, 2013; and they act both as competitors and predators of smaller zooplankton (Jocque, Vanschoenwinkel, & Brendonck, 2010;Lukić et al., 2018;Waterkeyn et al., 2011), which suggests a high impact on the zooplankton community. Even though B. orientalis hatches very early after inundation/ice break (Lukić, Vad, & Horváth, 2016;Petkovski, 1991), it is probably predominantly herbivorous in the early stages of its life (similar to other anostracans; Daborn, 1975;Fryer, 1983), which gives some time for zooplankton communities to establish at the beginning of the wet phase of their habitats. ...
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1. Omnivory is widespread in food webs, with an important stabilising effect. The strength of omnivorous trophic interactions may change considerably with changes in the local environment. 2. Shallow temporary waters are often characterized by high levels of inorganic turbidity that may directly limit the food uptake of filter-feeding organisms, but there is little evidence on how it might affect omnivorous species. Anostracans are key species of temporary waters and recent evidence suggests that these organisms are omnivorous consumers of both phyto-and zooplankton. 3. Using Branchinecta orientalis as a model species, our aim was to test how turbidity affects the feeding of an omnivorous anostracan. To do this, we used short-term feeding experiments and stable isotope analyses, with animals collected from soda pans in eastern Austria. In the feeding experiments, algae and zooplankton were offered as food either separately or in combination. The prey type treatments were crossed with turbidity levels in a factorial design. 4. There was a pronounced decrease in the ingested algal biomass with increasing turbidity. Conversely, ingestion rates on zooplankton were less affected by turbidity. Stable isotope analyses from field material supported our experimental results by showing a positive relationship of the trophic position of anostracans and the trophic niche of the communities with turbidity. 5. Our results show that turbidity modulates the intraguild trophic relationship between anostracans and their prey by shifting the diet of anostracans from more herbivorous in transparent to more carnivorous in turbid waters. Thus, inorganic turbidity might also have a community shaping role in plankton communities of temporary waters through altering trophic relationships.
... Current range of this species is limited to patches of suitable habitats clustered in regions of appropriate land morphology -especially in military training grounds, where habitats of this species depend on the specific form of land use (e.g. Gołdyn et al. 2012), and so that such areas may act as refuges for large branchiopods (Vanschoenwinkel et al. 2013). During the last 30 years, due to changes in European geopolitical situation as well as remodeling of military strategies, many training grounds have been abandoned and their ecosystems have changed dynamically -mainly due to the secondary plant succession (Jentsch et al. 2009, Cizek et al. 2013, Zentelis & Lindenmayer 2015. ...
Article
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Although large branchiopods are undoubtedly an important part of the temporary water bodies, they remain the least studied group of macroinvertebrates inhabiting freshwaters. One of these species is the fairy shrimp Branchipus schaefferi. Its known occurrences across the central Europe are limited almost exclusively to temporary pools on military training grounds. Thus, the species is particularly threatened by changes in military activities, especially by the decline in the number of temporary pools caused by abandoning of training grounds followed by the secondary plant succession. In the present study, we performed genetic analyses of known populations of B. schaefferi in military training grounds of Poland using nuclear 18S ribosomal DNA sequences. No genetic diversity was observed among nine populations from four military grounds. Further, very close phylogeographical relationships among B. schaefferi from distant geographic areas (Poland, Italy and Algeria) were found (genetic distance of 0.001-0.002 substitutions per site).
... during summer (Beladjal, Mertens & Dierckens, 2007;Sarma, Beladjal, Nandini, Cer on-Mart ınez & Tavera-Briseño, 2005). Temporary freshwater habitats range from large wetlands, vernal pools and marshes to rock pools, wheel tracks, small puddles and flooded rice fields (Vanschoenwinkel, Brendonck, Pinceel, Dupriez & Waterkeyn, 2013). To survive to the next generations, fairy shrimps produce resting eggs which are able to withstand extreme environmental conditions (Mertens et al., 2008), such as drought or frost (Reniers, Vanschoenwinkel, Rabet & Brendonck, 2013). ...
Article
The variable quality and high price of Artemia (Leach 1819) cyst products, used worldwide as live food, motivate aquaculturists to find an appropriate alternative, especially for fresh/brackish water organisms. In this study, Branchinecta orientalis (G. O. Sars 1901), a common fairy shrimp in north-western Iran, was reared for 15 days using effluent from trout ponds enriched with effluent filtrate as sole feed, or co-fed with microalgae (Scenedesmus sp.). The effluent filtrate was replaced by algae at different ratios (25% and 50%), and feeding experiments were designed at densities of 100, 200 and 400 individuals/L in 3-L containers and at 100 individuals/ L in 20-L containers. The results indicated that, at a certain density, the final length and survival were not significantly affected by different feeding regimes (p > .05). In 3-L containers, the highest length and survival were observed at the lowest density. B. orientalis contained the highest amounts of proteins, carbohydrates and lipids, though, when co-fed algae, although the differences with the treatment fed solely effluent filtrate were often limited. Inclusion of algae in the diet also resulted in higher levels of a number of PUFAs. Our study proves that B. orientalis can be mass-cultured successfully using trout effluent as culture medium without additional microalgae. Fish pond effluent is massively available as a cheap food source. Recycling aquaculture effluent for this type of live food production contributes to lowering the use of natural resources and to less discharge of nutrient loads into natural water bodies. KEYWORDS: Anostraca, Branchinecta orientalis (G. O. Sars 1901), fairy shrimp, fish effluent, mass culture
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1. Intense anthropogenic disturbance threatens temporary pond ecosystems and their associated fauna across the Palearctic. Since fairy shrimps (Crustacea, Branchiopoda) are endemic to temporary ponds, populations are declining due to habitat loss and it is important to define adequate units for conservation. 2. Phylogeographic reconstructions, based on genetic variation, provide valuable information for defining evolutionary and conservation units, especially for organisms with high levels of cryptic diversity like many fairy shrimps. We studied a total of 152 individuals of the fairy shrimp Branchipus schaefferi from 79 populations across the Palearctic and used mitochondrial (CO1) and nuclear (ITS1) DNA data to reconstruct the phylogeography of the species. 3. Our results show that B. schaefferi comprises four highly diverged (10.3-16.5%) evolutionary clades. The present-day haplotypes within each of the clades likely diverged from lineages that were maintained in separate refugia during the Pleistocene ice ages. While two clades represent distinct geographic regions, the two remaining clades have more wide and overlapping ranges. In addition, a limited number of shared haplotypes among populations from geographically distant regions within three of the clades suggest recent long distance dispersal events. 4. Overall, the studied B. schaefferi dataset comprises high levels of genetic differentiation, without a clear morphological signal. Phylogenetic searches and pairwise genetic distances suggest that the studied lineages belong to a complex of four morphologically cryptic species. Since these four evolutionary old clades persist (± 2 my), despite overlapping geographic ranges and since they span a variety of ecological conditions, they should be considered as separate Evolutionary Significant Units for conservation.
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Freshwater large branchiopods in Portugal: an update of their distribution This study is based largely on 20 years of field and laboratory work, with surveys conducted by the authors and some other researchers. During this period several studies dealing with freshwater large branchiopods (FLB) were carried out, resulting in scientific publications and project reports. The distribution of FLB in Portugal was presented in 2 international scientific meetings, but apart from a first paper by Vianna-Fernandes in 1951 and an update done by ourselves in 1999 concerning the southwest Portugal, no other information has been published. Therefore, this work intends to bring up to date the known distribution of this faunal group in freshwater temporary systems. This is pertinent because of the recent revision of the taxon Triops cancriformis on the basis of genetic analyses. The Portuguese populations were assigned either to the Portuguese endemism T. vicentinus, or to T. baeticus, the more widely spread Iberian species, both belonging to the T. mauritanicus complex. Furthermore, a new species, Tanymastigites lusitanica was found and described in Portugal. More recently, a male free (or, at least, a strongly female-biased) metapopulation of T. cancriformis was discovered in a rice field in the central region of Portugal. Here we present an updated status of FLB species of Portuguese temporary lentic systems and their distribution, plotted on a UTM (10 × 10 km) grid. A total of 505 sites (temporary ponds or assemblages of closely located, not individual temporary pools) have been surveyed (455 by us). In 241 of these (47.7%) at least one species of FLB was found on at least one occasion. Of the 505 sites, only 87 are located north of the Tagus River and of these, at least one species was found in only 17 (19.5%). South of the Tagus River (Alentejo and Algarve), 53.6% of the sites were inhabited by at least one species. This list comprises 7 anostracans, 2 spinicaudatans and 4 notostracans, including 3 species endemic to the Iberian Peninsula and another 3 to Portugal.
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Słowa kluczowe: duże skrzelonogi, Branchiopoda, astatyczne zbiorniki wodne, koleiny. Opisano nowe stanowisko zadychry pospolitej Branchipus schaefferi, rzadkiego gatunku skorupiaka należącego do gromady skrzelonogów Branchiopoda. Zadychrę znaleziono 12 czerwca 2011 roku w dwóch kałużach, położonych ok. 70 m od koryta rzeki Raby, w miejscowości Gdów (49°53,703′N, 20°11,698′E). W wypełnionych wodą koleinach na drodze gruntowej nie występowały rośliny wodne, nie było też drapieżnych bezkręgowców (poza nielicznymi formami imaginalnymi chrząszczy). Obserwowano natomiast liczne larwy komarów i małżoraczki. Zadychra zajmowała głównie głębsze części kałuż. Na opisywanym stanowisku, poza zadychrą, 8 sierpnia 2012 roku zanotowano również obecność wszystkich stadiów rozwojowych (tj. jaj, larw, osobników młodocianych oraz 4 osobników dorosłych) kumaka górskiego Bombina variegata – gatunku, który również preferuje tego rodzaju siedliska.
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Macroinvertebrate populations from constructed and natural vernal pools on the same land forms and in close proximity, were compared quantitatively to determine colonization and temporal trends. Population trends in the natural pools were used to establish the functional success criteria for constructed pools. Success was evaluated by the quantitative and qualitative similarities of the constructed pool invertebrate populations to those of the natural pools. Most constructed pool invertebrate populations were found to mirror existing populations of vernal pool "obligate" species in natural vernal pools within two years. A combination of factors (i.e., over abundance of Glyceria sp., ponding depth, amount of organic matter in pool) may have contributed to the deviation of invertebrate populations in a few constructed vernal pools from those in natural pools. Quantitative and qualitative monitoring will continue for seven more years.
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Large branchiopods are threatened worldwide by the loss and degradation of their temporary aquatic habitats owing to drainage and intensive agriculture. Sound ecological knowledge of their diversity and distribution is a prerequisite to formulate effective conservation measures. In the present study, large branchiopods were collected from 82 temporary freshwater pools belonging to five habitat types in Kiskunsag (Hungary). Dormant propagule bank analysis complemented the field survey. Eleven species were found, with large branchiopods occurring in more than half of the study systems. The high regional species richness and occurrence frequency of large branchiopods make Kiskunsag a true 'hot spot' of large branchiopod diversity. The local environment was more important than spatial factors (isolation) in explaining the presence of the most common species. Dispersal was most likely not limiting for the large branchiopods in the study area and colonisation success of different species was differentially affected by local conditions, possibly invertebrate predation risk and hydroperiod. Meadow pools and wheel tracks contributed most to regional species richness through the presence of rare and exclusive species. To conserve branchiopod diversity, we stress the importance of high habitat diversity in the landscape and the need to conserve neglected habitats such as wheel tracks.
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The somatic growth, longevity, and reproduction of two Moroccan anostracan populations from different climatic areas were studied under standardized laboratory conditions. Both populations were subjected to allozyme analyses covering four loci, and molecular analyses of the variable regions, the Internal Transcribed Spacers (ITS1 & ITS2) intervening the nuclear ribosomal genes (18S, 5.8S, 28S rDNA). The ecological characteristics of the life cycle of each population are presented, together with their genetic differences and phylogenetic relationships.
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Branchipus schaefferi and Streptocephalus torvicornis commonly co-occur in ephemeral ponds throughout the Mediterranean Region. We compared survivorship, growth, and reproduction. Our results show statistically significant differences in all three parameters under laboratory conditions at 25°C, reflecting different life history strategies between these species. Branchipus schaefferi grows quickly to 18 mm body length, producing roughly 1700 cysts during an average lifespan of 24 days, while S. torvicornis lives an average of 120 days (length 24 mm), laying 2400 cysts. This suggests that S. torvicornis is better adapted to deep longer-lived pools, whereas B. schaefferi may survive in small, more ephemeral pools as formed from spring melt water and autumn rains. While the lifespan is equal for both sexes in B. schaefferi, the males of S. torvicornis live 2.8 times longer than females (325 days versus 114) compelled into a long postreproductive period by lack of females in their environment.
Article
The proportion of predatory animal species is often believed not to vary systematically across communities. However, we predict that larger temporary freshwater pools, and pools that are more permanent, will contain a higher proportion of predatory animal species. In 24 temporary rockpools in Northern Israel (supporting communities dominated by ostracods, copepods, cladocerans, flatworms, dipterans and amphibians), the mean proportion of macroscopic predatory species (averaged over a series of samples) increased with increasing pool area. For the highest possible proportion of predatory species (including microscopic species with uncertain diets), the relationship with pool area was not statistically significant. We did not find significant relationships between permanence and the proportion of either macroscopic or all possible predatory species. Larger pools and pools that were more permanent had more species. Species richness and the proportion of macroscopic predators were positively correlated. These patterns imply that species-poor ecosystems are likely to be functionally different from species-rich systems.
Article
1. The notion that the spatial configuration of habitat patches has to be taken into account to understand the structure and dynamics of ecological communities is the starting point of metacommunity ecology. One way to assess metacommunity structure is to investigate the relative importance of environmental heterogeneity and spatial structure in explaining community patterns over different spatial and temporal scales. 2. We studied metacommunity structure of large branchiopod assemblages characteristic of subtropical temporary pans in SE Zimbabwe using two community data sets: a community snapshot and a long-term data set covering 4 years. We assessed the relative importance of environmental heterogeneity and dispersal (inferred from patch occupancy patterns) as drivers of community structure. Furthermore, we contrasted metacommunity patterns in pans that occasionally connect to the river (floodplain pans) and pans that lack such connections altogether (endorheic pans) using redundancy models. 3. Echoes of species sorting and dispersal limitation emerge from our data set, suggesting that both local and regional processes contribute to explaining branchiopod assemblages in this system. Relative importance of local and regional factors depended on the type of data set considered. Overall, habitat characteristics that vary in time, such as conductivity, hydroperiod and vegetation cover, best explained the instantaneous species composition observed during a snapshot sampling while long-term species composition appeared to be linked to more constant intrinsic habitat properties such as river connectivity and spatial location.
Article
1.Several human activities, such as actions for nature conservation, research and recreational activities, are closely associated with inland aquatic habitats that are usually considered as isolated island habitats. In this study, the possibility of unintentional dispersal of aquatic invertebrates among water bodies via footwear and motor vehicles was investigated.2.Mud samples collected from boots and from the tyres and wheel cases of cars used for field work by biologists (Camargue, Southern France) were hatched under laboratory conditions and also checked for the presence of unhatched propagules. A large number of organisms hatched and invertebrate propagules from a wide range of taxa were encountered (including Artemia, freshwater large branchiopods, Cladocera, Ostracoda, Rotifera, Turbellaria, Nematoda, etc.). The results also demonstrated that different research groups tend to transport the aquatic invertebrates typical for their respective study systems.3.Human dispersal of aquatic invertebrates has been studied mainly on large continental scales, such as in the case of transoceanic transport via ballast water in ships. This study provides evidence that dispersal via footwear and motor vehicles may result in frequent dispersal of aquatic invertebrates on a local scale, and we presume also occasionally over longer distances. Given the rapid spread of invasive zooplankton species (e.g. Artemia franciscana encountered in this study), we promote caution and recommend cleaning before transport of any equipment which comes in contact with water or aquatic sediment. Copyright © 2010 John Wiley & Sons, Ltd.
Article
1. To monitor the diversity and distribution patterns of large branchiopods and the effects of local and regional processes, 30 temporary wetlands in a nature reserve in the Camargue (southern France) were sampled and characterised during three consecutive inundations (2005–08). Additional species were added to the list for each wetland by hatching animals from the resting egg bank, after determining the optimal hatching conditions. 2. A total of five species were found, representing 28% of the species known in France and 56% of the known Camargue species. Tanymastix stagnalis, Branchipus schaefferi, Chirocephalus diaphanus (Anostraca), Triops cancriformis (Notostraca) and Imnadia yeyetta (Spinicaudata) were distributed over a total of 19 wetlands. 3. More than one species was present in 79% of the wetlands containing large branchiopods. Individual wetlands harboured on average 2.8 species, with a maximum of five coexisting species. Large branchiopod assemblages were temporally variable, differing among the three inundations with different climatological conditions. 4. The most important habitat factor influencing the distribution of large branchiopods was salinity, adversely affecting the density and survival of hatchlings. The persistence of large branchiopods in these temporary waters may be threatened by increasing salinisation driven by intensive water management and climate change.
Article
1. Many invertebrates inhabiting insular aquatic habitats rely on external agents or vectors to disperse. Besides water connections and wind, waterfowl and amphibians are known to mediate passive dispersal of freshwater invertebrates. However, the possibility of dispersal by terrestrial mammals has been largely overlooked. 2. We investigated the potential of both external and internal zoochorous dispersal of aquatic invertebrates by the wild boar (Sus scrofa) in Mediterranean wetlands in the Camargue (France). As wild boar frequently visit wetlands for feeding and wallowing purposes, we hypothesized that they may be important passive dispersal vectors of aquatic invertebrates at a local scale. Dried mud was collected from selected ‘rubbing trees’ used by boars to dispose of parasites. Additionally, faecal pellets were collected from different locations in the wetland area. 3. Seventeen freshwater invertebrate taxa including rotifers, cladocerans, copepods and ostracods hatched from sediment obtained from ‘rubbing trees’, while invertebrates hatching from dried faeces (10 taxa) were mainly rotifers. Dispersing invertebrates were collected up to 318 m from a nearest potential dispersal source. Both abundance and richness of invertebrates significantly decreased with dispersal distance. 4. Our results demonstrate that large mammals such as wild boar can act as dispersal vectors of aquatic invertebrates at a local scale in the wetland area of the Camargue and suggest that external transport may be quantitatively more important than internal transport. As wallowing (mud bathing) is common in many terrestrial mammals, this mode of dispersal may be quite widespread.