J OURNAL OF C RUSTACEAN B IOLOGY, 36(3), 396-401, 2016

COLLECTING AND PROCESSING BRANCHIOPODS

Joel W. Martin 1,∗ , D. Christopher Rogers 2 , and Jørgen Olesen 3

1 NaturalHistory Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, CA 90007, USA
2 KansasBiological Survey and the Natural History Museum (Biodiversity Institute), Kansas University,
Higuchi Hall, 2101 Constant Avenue, Lawrence, KS 66047-3759, USA
3 Natural History Museum of Denmark (Zoological Museum), University of Copenhagen Universitetsparken 15,

DK-2100 Copenhagen Ø, Denmark

K EY W ORDS: Anostraca, Cladocera, culture, Cyclestherida, electron microscopy, Laevicaudata, Noto-

straca, Spinicaudata
DOI: 10.1163/1937240X-00002434

I NTRODUCTION the use of similar collecting and preservation methods

This discussion is the first of a series of articles to be pub-
lished in subsequent issues of Journal of Crustacean Biol-
ogy that will cover methods in the collection, preservation,
culture, and other techniques used in the study of particular
groups of crustaceans. Other articles will cover photography
and particular habitats and techniques.
Branchiopod crustaceans (class Branchiopoda) are com-
mon inhabitants of both temporary and permanent bodies of
standing fresh or saline inland waters on all continents, even
Antarctica; there are also a few marine planktonic species
(Brendonck et al., 2007; Rogers, 2009). They are often di-
vided into cladocerans (Cladocera, or water fleas) and non-
cladoceran “large branchiopods” (fairy, tadpole, and clam
shrimps). This artificial division is convenient for those who
work on these groups, but it masks the large size that some
cladocerans can attain and gives the impression that the large
branchiopods (Anostraca, Notostraca, Laevicaudata, Spini-
caudata, and Cyclestherida) are somehow more closely re-
lated to each other than any of them are to cladocerans.
Relationships within Branchiopoda are still not entirely un-
derstood and are the subject of much study (e.g., Martin,
1992; Walossek, 1993; Olesen, 2000, 2003, 2007; Martin
and Davis, 2001; Weekers et al., 2002; Olesen and Gry-
gier, 2004; Stenderup et al., 2006; Richter et al., 2007), but
enough is known to state that, whereas Cladocera appears
to be a natural group, the grouping of “large branchiopods”
is not. There are approximately 1000 described extant bran-
chiopod species (see Martin and Davis, 2001; Brendonck et
al., 2007; Rogers, 2009, 2013).
Most are of a relatively uniform size except cladocerans,
some of which can be particularly small, and the larger
notostracans (tadpole shrimps) (Fig. 1B) and large predatory
anostracans (fairy or brine shrimps) (Fig. 1A), some of
which are relatively large. Such relative uniform size allows
throughout the various groups of branchiopods.
Understanding the life cycle of branchiopods is important
for collection. The typical life cycle of large branchiopods
involves males, females, and eggs that are typically shed
into the environment and a nauplius larval stage. Many clam
shrimps and tadpole shrimps (and rarely fairy shrimps and
cladocerans) are hermaphroditic. Cladocerans typically have
no free living larvae (the predatory Leptodora is an ex-
ception). Some populations and some species (in Spinicau-
data (Fig. 1C), Cyclestherida, and all cladocerans) repro-
duce parthenogenetically, with males being extremely rare
or even absent. Because they inhabit ephemeral bodies of
water, the duration of the entire life cycle of many bran-
chiopods is amazingly short, with development, maturation,
mating, and egg development sometimes occurring in a span
of only a few weeks or less. In many of these species, eggs
or ephippia are produced that can withstand desiccation and
persist in the environment for years until the next rain cy-
cle provides the water that allows them to hatch and begin
again.
E COLOGY
Branchiopods are unusual among crustaceans in that most
species live in freshwater rather than marine habitats. The
majority inhabits seasonally astatic (often called temporary,
ephemeral, or vernal) pools, puddles, and streams that
for most of the year remain dry. Thus, collecting adult
branchiopods is more a matter of timing than of technique.
Yet the substrate will contain eggs that can be easily
collected and reared. Many cladocerans inhabit permanent
water bodies (ponds and lakes) and can be collected in all
seasons except winter (depending on the locality), and a few
species are marine.
Anostracans occur worldwide in ephemeral ponds, playas,
and salt lakes. Species are known from ultrapure fresh waters

∗ Corresponding author; e-mail: jmartin@nhm.org

© The Crustacean Society, 2016. Published by Brill NV, Leiden DOI:10.1163/1937240X-00002434

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 MARTIN ET AL.: COLLECTING AND PROCESSING BRANCHIOPODS 397

Fig. 1. A, Eubranchipus bundyi Forbes, 1876 (Anostraca) with unidentified cladocerans and copepods, Grass Lake, Siskiyou County, CA, USA; B,
Lepidurus packardi Simon, 1886 (Notostraca), Olcott Lake, Solano County, CA, USA; C, Cyzicus californicus (Packard, 1874) (Spinicaudata), Olcott
Lake, Solano County, CA, USA; D, Dr. Regina Wetzer using a circular dip net to collect branchiopods from a shallow pool in Inner Mongolia, China; E,
“Wildco” bottom “aquatic kick net” in use; F, rectangular net (WaterMark© bottom “aquatic kick net”) with stainless steel frame and aluminum handle; G,
plankton net with detachable collecting bottle; H, heavy-duty, D-frame “aquatic net”; I, WaterMark© stream drift net, with Nitex nylon mesh net and PVC
collecting tube; J, Field Master© refractometer for measuring aqueous concentrations and, indirectly, salinity. Source of images: Forestry Suppliers; TekTak;
Science First; Wildlife Supply.

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 398 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 36, NO. 3, 2016
in the tundra to extremely turbid and silt-laden puddles.
Most species appear to be filter feeders or deposit feeders,
but some are rapacious predators on other invertebrates.
Notostracans are known from temporary bodies of water
on all continents (except Antarctica), plus New Zealand,
and some oceanic islands (e.g. Galápagos). Anostracans and
notostracans inhabit some truly inhospitable environments
and temperatures; eggs of Triops and Artemia have been
shown to withstand boiling water and even liquid nitrogen
and still develop normally (Persoone and Sorgeloos, 1980).
Notostracans are not filter feeders, but instead appear
to feed on detritus or on other organisms, living or dead,
and have been reported to pursue and catch anostracans
and even small fish. Clam shrimps are found across the
same range as the above two groups, and they are almost
always restricted to temporary waters. There are reports of
species of Lynceidae in “prairie streams” (see Martin, 1992),
and Cyclestheria hislopi (Baird, 1859) is known from some
permanent bodies of water, where it is invariably associated
with aquatic vegetation; species of Cyclestheria are basically
circumtropical (Martin and Boyce, 2004; Schwentner et
al., 2013). Although often described as filter feeders, clam
shrimps actually do a large amount of scraping and tearing
of their food, and they will also scavenge.
Cladocerans are typically much smaller (usually 0.2 to
3.0 mm) than other branchiopods. They are typically plank-
tonic rather than benthic (though many are truly benthic),
but the group is morphologically and ecologically diverse.
Species are found in permanent and temporary freshwater,
estuarine, and sometimes marine habitats (especially Podon
and Evadne).
Although habitats differ by species, any of these taxa
might be encountered in just about any ephemeral pool or
similar habitat (such as watering troughs). Their modes of
distribution, with most species having drought-resistant eggs
that can be carried by waterfowl or blown by the wind,
are such that some species are quite widespread (Rogers,
2014a).
C OLLECTION
Collecting adult large branchiopods is most easily done
using a large dip net or sweep net (Fig. 1D, F, H). Nets
made especially for capturing aquatic insects work well.
These can be purchased in circular, rectangular, D-shaped, or
triangular shapes. The triangular nets are by far the strongest
design, whereas the rectangular and D-shaped nets tend to
collect more specimens per sweep because of their larger
size and flat leading edge. Some of these nets are sold under
the name “kick net” as they are often placed downstream
of rocks that are kicked over to expose and release any
organisms that then drift into the net (Fig. 1E). The better
made nets are expensive but are worth the price in terms of
their durability. These nets eventually take a beating along
the leading edge of the frame, which eventually leads to the
deterioration of the net itself. To prolong the life of the net,
it is helpful to sew, using stout thread or floss and a heavy
needle, a layer of thin rubber, such as what is found in an
inner tube, to the leading edge of the net frame protecting the
canvas when the net is dragged along the bottom of the pool.
Aquatic “drift nets” (Fig. 1I) will also work, as will a seine
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if the mesh is small enough (0.5 mm for large branchiopods,
and down to 0.2 mm for cladocerans), although neither is
quite as portable or as easy to use as a simple dip net. For
cladocerans, some workers recommend placing a large mesh
plate in front of the net aperture to reduce the amount of
vegetation while still collecting cladocerans.
The best way to sample with a dip net is to walk
slowly through the pool sweeping the net through the water
ahead of the pressure wave made by your feet. The net
should be moved at a slightly oblique angle from the
direction you are walking, side to side, to minimize pushing
branchiopods ahead of the net in the pressure wave. This
pressure wave will increase in strength as more material
accumulates in your net. Large predatory anostracans live
in large, often deep temporary playas and lakes and are
sensitive to movement in the water, quickly swimming away
and easily avoiding dip nets. These large-size species are
caught more easily with a two-person minnow seine. Many
fairy shrimps, especially in clear water, have strong predator
avoidance responses, and upon detecting movement will
quickly dive for cover in vegetation. These species often
must be hunted individually or collected by pulling up
quantities of vegetation or debris, placing it in an enamel
pan, and sorting out the individual shrimps. If access to the
pool is difficult or if the body of water is very small, smaller
nets used for the aquarium trade or for fish bait also will
work well.
Tadpole and clam shrimps typically take refuge at the
bottom, and tadpole shrimps will burrow shallowly into the
substrate. As a result they are often undersampled by typical
dipnetting. By standing in one spot in a pool and moving
the dipnet through a figure-eight pattern several times, many
specimens can be drawn up from the bottom and into the dip
net.
An aquatic light trap can be effective, especially if the
habitat is very deep. Branchiopods are positively phototropic
and will come readily to an aquatic light trap. The downside
of this method is that one or two predatory insects can reduce
or mangle branchiopods. A bright light shone on a pool at
night also will attract large numbers of branchiopods.
For planktonic cladocerans, collecting devices can include
plankton nets (Fig. 1G) and tubes designed for taking
midwater samples from shore or boat (e.g., Schindler-Patalas
plankton traps, Clarke-Bumpus tow nets, Kemmerer and Van
Doren water bottles, and Pennak style tubes and collectors;
see Pennak, 1962; Wissmar and Rabe, 1970; George and
Owen, 1978; Swanson, 1978; Amoros, 1980; DeVries and
Stein, 1991; Paggi et al., 2001). Many cladocerans live
associated with vegetation in shallow water. A simple and
effective method to collect some of these forms is to drag
a plankton net slowly through the vegetation several times.
The first hauls will disturb the fauna, causing them to lose
their association with the vegetation and/or the bottom;
the fauna can then be collected during the next sweep of
the net. Care should be taken to reduce the amount of
detritus in the sample (try not to stir up the bottom too
much during sampling). Because cladocerans tend to be
very small, sorting is best undertaken under a dissecting
microscope after fixation/preservation has taken place.
 MARTIN ET AL.: COLLECTING AND PROCESSING BRANCHIOPODS
Associated data should always be recorded as well; these
include the date, collectors, site coordinates and a general
description of the site (including a digital photograph if
possible), water temperature, and the refractive index (using
an optical refractometer (Fig. 1J), an indirect measure of
salinity).
R EARING AND C ULTURE
Often the habitat where branchiopods are known to exist is
dry, and/or it is not possible or convenient to visit the site
while it contains water. It is nevertheless possible to collect
some of the dry mud from the habitat bottom for culturing in
the lab. Collecting mud can be done almost any time, with
the eggs hatched out at the researcher’s convenience. Dry
branchiopod eggs can remain viable for many years, and the
upper limit of their viability is unknown, with reports of dry
eggs remaining viable for more than a century. Once water is
added to the eggs, the animals will begin to hatch. Modifying
the light/dark cycles and altering other physical parameters
(water quality, heat, type of dirt, turbidity) can improve
the likelihood of hatching (for anostracans see Brendonck
et al., 1990; Rogers and Fugate, 2001; Rogers, 2014b; for
notostracans see Scott and Grigarick, 1978; Fry-O’Brien and
Mulla, 1996; and the Triops rearing web site of Dr. Steven C.
Weeks, available online at http://www.tadpoleshrimp.info/
rearsug.html).
Substrate is very important for most species. The eggs as-
sociated substrate contains the specific geochemical compo-
nents that the animals need to survive (Rogers, 2014b), as
well as the propagules of the various algae, bacteria, and mi-
crometazoans that the developing branchiopods feed upon.
The substrate (especially for cold-water species) should al-
ways be dry when collected. Damp or moist substrate col-
lected into a plastic bag or bottle often harbors fungi. The
container can act as a greenhouse, and the fungi kill and con-
sume the eggs. Typically, only the top centimeter or so of
substrate needs to be collected; most of the egg bank is usu-
ally in the top sediment layers. Some tadpole shrimps will
attach their eggs in a clump to a rock or some other hard
substrate, if available, and the anostracan Streptocephalus
oviposit their eggs in the substrate (Kraus et al., 2004).
Temperature is very important in culturing eggs. Cold-
water species must be cultured in a refrigerator or culture
chamber with a chiller, and often need daily fluctuations
of temperatures to develop properly. The lower temperature
also maintains the higher dissolved oxygen levels that the
species require (Rogers and Fugate, 2001). Warm-water
species often require a water temperature above 27°C and
additional weak aeration from a bubbler or air stone. The
container should be large enough to allow the animals to
move away from the action of the rising bubbles. Fairy
shrimps especially are soft bodied and are easily damaged
in the bubble current.
Culture containers for most species should hold 5 liters or
more and should be shallow, only a few centimeters deep,
with a large water surface area for gas exchange.
Many species can be cultured outdoors where local
climate conditions are similar to the substrate collection site.
Plastic wading pools, placed where they will collect natural
rainwater and will receive little allochthonous material (such
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399
as leaf litter, which may change the chemistry of the culture),
can be very effective.
To start the culture, place the substrate in the container and
spread it out as much as possible. Ideally one liter of sub-
strate for every 5 liters of water will ensure that the substrate
maintains the chemical nature of the original habitat. Next
add 2 liters of rainwater or distilled or deionized water. For
cold-water species, the water should be near freezing with-
out ice formation. For warm-water species the water should
be between 30 and 35°C because many of these species can-
not survive below 30°C. After adding the water, gently mix
the culture and begin aeration. After 24 hours, add another
2 liters of water, stir gently, and wait. Cold-water taxa should
be started around 2°C, and after 48 hours the temperature
should be raised to 5°C. Some habitats will have cold-water
species swimming during the winter and warm-water species
swimming during the summer, so individual cultures run at
different temperatures are needed to obtain hatches of both
groups.
Most cultures will need light. It is best to have the
light source at one corner of the culture container to
create a thermocline, providing a slight range of light and
temperatures that the developing branchiopods can occupy.
Cold-water species will need light that will not increase
the temperature of the system. Some tropical species need
darkness or black substrates to survive after hatching.
Most cultures of fairy shrimps, clam shrimps, and clado-
cerans, as well as juvenile tadpole shrimps, are easily raised
on a diet of unicellular algae, finely ground (powdered) com-
mercial tropical fish plant-flake food, or yeast. If sufficient
substrate is present, however, the native microorganisms in
the branchiopod’s diet also will be present. It is easy to
over feed using yeast or commercial fish food. Only enough
should be added to the culture to start bacteria or the yeast
as a culture within the culture. Champagne maker’s yeast is
preferable to baker’s yeast because it has higher protein con-
tent. Typically, a pinch dissolved in 10 ml of water is needed
per 5 liters of water once every two weeks.
The large predatory fairy shrimps need a constant supply
of smaller fairy shrimps on which to feed. These predators
need to capture and kill their own prey items and will not
eat dead or moribund prey (Rogers et al., 2006). Tadpole
shrimps need a variety of food, including yeast, metazoans
(bloodworms, mosquito larvae, copepods, fairy shrimps),
and some plant matter; some survive well on carrots.
P RESERVATION
Because of their small size, branchiopods can be dropped
directly into 70% ethanol. Because of their high water
content, the alcohol should be replaced after 24 hours
to avoid dilution of the preservative and decomposition.
Alcohol is a preservative rather than a fixative, so if the
specimens are needed for histological or morphological
purposes, such as scanning electron microscopy, it is best
to expose them briefly to 5% formalin or other fixative
prior to rinsing them in freshwater and placing them in 70%
ethanol for storage. If specimens are needed for molecular
work (and if freezing is not an option), they can be placed
directly in 95% ethanol. It is often the case that when
one finds one branchiopod, the entire adult population is
 400 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 36, NO. 3, 2016
also found so there is usually no shortage of specimens.
This makes it relatively easy to preserve specimens in
multiple ways for multiple uses, some fixed and some
frozen or alcohol-preserved for molecular work. Collecting
cladocerans with a plankton net yields small samples of
water with a high concentration of specimens. In these cases,
and if material for genetic analyses is not required, the
simplest preservation method is to fix the batch by adding
a small volume of high percentage formaldehyde to the
sample. A final concentration of 2-4% of the sample is
desirable. Branchiopods dropped into ethanol or formalin
often contort, and may not make acceptable subjects for
photography. They are nevertheless easily anesthetized and
relax in carbonated water or lemon-lime soda. Because
anostracans are soft bodied and have a high water content,
they typically shrink after preservation anywhere up to one
third their original size (Rogers, 2002).
P HOTOGRAPHY AND P REPARATION FOR
E LECTRON M ICROSCOPY
Photographic and SEM methods used for branchiopods
do not differ appreciably from those used for other small
crustaceans (Felgenhauer, 1987). Typical fixatives for SEM
use are osmium tetraoxide (1%), formaldehyde (3.7%),
glutaraldehyde, and Bouin’s solution (e.g., see Felgenhauer,
1987; Møller et al., 2003; Olesen et al., 2003). Olesen et
al. (2003) reported that Bouin’s solution results in the least
shrinkage of tissue. It is rare that the collector has these
solutions on hand in the field, and various alternatives,
including rum, whiskey and other alcoholic products, have
been used occasionally with mixed results but sometimes
successfully.
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