Trapped

in the pond
BIODIVERSITY OF CRUSTACEANS
IN TEMPORARY PONDS

Preface

The preservation of Biodiversity is the critical factor determining success when facing the challenges posed by sustainable development.
The safe path to reach that purpose is by facilitating the increase of scientific knowledge through the development of projects in which the inherent value is
maximized by fostering the cooperation among institutions of recognized merit.
Following this approach, the research project CHARCOScomBIO, developed in
partnership by the Centre for Environmental Biology (CBA) and the Centre for Research on Biodiversity and Genetic Resources (CIBIO), started the phase of scientific divulging with the exhibit Trapped in the pond and with this brochure.
EDP sponsored this project through its Biodiversity Fund and assumed a position of aggregating efforts scattered among several working groups and scientific
institutions.
Congratulations are due to the authors of this scientific endeavour.
The results obtained, highlighting the natural value of temporary ponds in terms of biodiversity and nature conservation, and the attractive and didactic way of
presenting the work to the general public, are factors that stimulate the general
interest on the fragile and complex interdependence among living organisms, and
the importance to preserve that equilibrium.
EDP has, since long, valued the importance of natures biodiversity in a pioneering way, and integrates biodiversity in its various business areas. In 2007, EDP implemented its Policy for Biodiversity with the intent of contributing to the worlds
objective of reducing biodiversity loss.
We are convinced that a strong commitment from society, companies and researchers is required to ensure the good functioning of ecosystems, valuing the
services and products provided by nature, and creating opportunities for the sustainable use of natural resources. We are committed in pursuing this strategy in a
clear and transparent way.

Eng. Neves de Carvalho

Director of Sustainability and Environment, EDP Energies SA.

Contents

INTRODUCTION 4
TEMPORARY MEDITERRANEAN PONDS 5
What are ponds? 5
Ponds are priority habitats! 6
BIODIVERSIT Y OF MEDITERRANEAN TEMPORARY PONDS 7
Pond visitors 7
Permanent inhabitants 8
CRUSTACEANS OF TEMPORARY PONDS 10
What are crustaceans? 10
Micro-crustaceans 11
Copepods 11
Calanoida 11
Cyclopoida 13
Harpacticoida 13
Branchiopods 14
Ostracodes 16
Macro-crustaceans Large Branchiopods 17
Anostraca 17
Notostraca 18
Spinicaudata 19
Crustaceans from the age of Dinosaurs! 20
DYNAMICS OF TEMPORARY PONDS 22
Autumn
22
Winter
23
Spring 26
Summer
27
How do pond animals feed? 27
CONSERVATION OF TEMPORARY PONDS 29
What can we do to contribute to pond conservation? 29

Introduction

Temporary Mediterranean ponds are
freshwater systems with shallow depth
that undergo periods of flooding and
drought, and are isolated from permanent water bodies (ex. lakes or rivers).
Ponds are of great ecological and cultural
importance in the arid and semi-arid
areas of the Mediterranean Region. This
type of habitat exists in areas with land
depressions and varies according to local
geology, geomorphology, depth and water origin (underground or runoff water).
Many natural or man made ponds are
profoundly associated with agriculture
and animal farming, and are structuring
elements of the landscape and of the
history of agriculture. Yet, during the XX
century, a sharp decrease in the number
of these aquatic habitats was observed
throughout Europe, as a result of agriculture intensification and urban development. In Portugal, systematic work on
the conservation status of these habitats
and on the temporal variation of its
quality and quantity is unknown. Additionally, projects especially directed to
the conservation of these wetlands are
scarce, and the civil society is generally
unaware of the importance to protect the
habitats. In this context, the project entitled Investigation, Conservation and
Public Awareness of Temporary Ponds
Biodiversity CHARCOScomBIO aimed
to improve the scientific knowledge on
Mediterranean temporary ponds and on
the biodiversity of crustaceans and amphibians inhabiting the ponds, in articulation with a conservation program and
activities to raise the public awareness
of this biodiversity, and environmental
education.

The project CHARCOScomBIO was

one of the winning projects of the EDP
Fund for Biodiversity in 2009, involving
several institutions devoted to scientific
research.

Exhibit Trapped
in the Pond
The project CHARCOScomBIO included the production of the exhibit
entitled Trapped in the Pond, at the
National Museum of Natural History and
Science. This exhibit intended to show
to the general public, the diversity of
temporary ponds in Portugal, the most
common and abundant crustaceans species in the ponds, and their adaptations
to the environment and life in the ponds
throughout the year.
The visitors are stimulated to know
the answers to the most relevant questions about this subject, such as: What
are temporary ponds? What is the actual
dimension of the crustaceans living in
ponds? How do crustaceans grow? How
do they breathe? What are their reproduction strategies?
In this brochure, we will explore and
illustrate some of these questions, contributing to a better knowledge of one of
the most important and unknown biodiversity of Portugal and of the Mediterranean Region.

Trapped
in the pond

Temporary Mediterranean ponds

What is a pond?
Temporary ponds are without doubt
some of the most remarkable yet
most threatened habitats in the
Mediterranean region.
Mediterranean Temporary Pools, Quzel P.
In Grillas et al. 2004

Temporary ponds are natural ecosystems of still or slow flowing water
where periods of flood follow periods of
drought. The flood period is dependent

of periods of intense rain and has a short

duration. The water depth may vary from
some centimetres to about 2 meters.

ECOSYSTEM
It is the assemblage of organisms living in
a certain area (biotic components) together
with the physical and chemical factors with
which they interact (abiotic components).
In sum, it is the community of organisms
plus the environment in which they live.

Mediterranean temporary pond from Torro, Alentejo.

Mediterranean
temporary pond
at a fossil dune,
Pond B27, Natural
Park from the
Southwest and
Vicentine Coast
(PNSACV).

Ponds are habitats

of priority
conservation!
Temporary ponds are very important
natural ecosystems in the Mediterranean
region. The large diversity of aquatic
plants, molluscs, crustaceans, insects,
amphibians, reptiles and aquatic birds
that exists in temporary ponds rendered
them the designation of priority habitats
for conservation under the Habitats Directive (Habitat 3170). Temporary ponds
are also included in the Ramsar Convention for wetlands since 2002. Portugal
has 30% of the Mediterranean temporary ponds included in the Natura 2000
Network.
Nevertheless, the biodiversity of
temporary ponds is widely unknown or
ill considered by the civil society, and it
is frequently threatened by pressures of
human origin.

HABITATS DIRECTIVE
Its main objective is to ensure the
preservation of biodiversity in
the European Union through the
conservation of natural habitats
considered threatened and of the
animal and floral species there
inhabiting.

NATURA 20 0 0 NETWORK
It is a network of areas established
to ensure the conservation of
habitats and rare wild species,
threatened or vulnerable in the
European Union. It results from the
implementation of two directives:
Birds and Habitats Directive.

Biodiversity of Mediterranean temporary ponds

As ponds are temporary and only
exist during periods of heavy rainfall,
they are frequently unnoticed in the
landscape. However, the biodiversity of
these habitats is unique, adapted to the
alternating periods of flood and drought.
Numerous species of aquatic plants,
crustaceans and other micro- and macroinvertebrates live exclusively in ponds,
spending there their entire life cycle.
These are the so called permanent inhabitants of ponds. Other organisms,
named visitors, such as amphibians and
most insects, use ponds to spend part of
their life cycle or to feed.
In fact, temporary ponds host a
greater biodiversity, and have a larger
number of rare or threatened species,
than other freshwater systems, and

supply species that colonize artificial

or modified environments such as dam
reservoirs.

Pond visitors
Many insects and amphibians are species that visit ponds only during the reproduction period or as feeding grounds.
It is the case of coleopteran insects
(beetles) like the genus Dytiscus. Their
larvae hatch and grow in the pond where
they feed on crustaceans, other insects
and even amphibian larvae. When the
ponds dry, the adults move to permanent water bodies such as rivers, lakes
and reservoirs.
There are insects belonging to the order Hemiptera that characteristically are
predators that reproduce in the ponds.

Full pond
(pond B26, PNSACV)
Dry pond
(pond B26, PNSACV)

This order of insects includes waterstriders (Gerris sp.), insects that live at
the bottom (water scorpions Nepa sp.)
and swimmers (Notonecta sp.).
The amphibians reproduce in the
ponds and both larvae and insects feed
there. The aquatic birds feed in the

ponds, and both birds and mammals

drink there. As an example, the swallows
collect water with their beaks to drink
and to make their nests of clay.

Coleopteran
of the genus
Dytiscus (water
beetle)

Marble newt,
Triturus
marmoratus
Swallow
collecting water

Water-strider
insect, Gerris sp.

Permanent
inhabitants

J. Petronilho

Swimming
insect,
Notonecta sp.

Several species of plants, gastropods (snails), hirudines (leeches), tardigrades and crustaceans are among the
inhabitants that spend their complete
life-cycle in the ponds. Plants like Eryngium coriculatum (pond thistle) colonize
the deepest areas of ponds, while Ranunculus peltatus (buttercups) floats and its
flowers emerge from the water surface
in Spring. Potentilla erecta (common

tormentil) and several species of the

genus Juncus and of Isoetes (quillworts)
colonize the shallow areas of the ponds
and their margins.
Medicinal leech

conditions. When the water disappears

from their environment, they resort
to a suspended life stage; they dehydrate completely becoming a life form
that withstands extreme high and low
temperatures and radiation. Thanks
to this ability, tardigrades have been
used in tests during space missions and
were the first animals to survive lack of
oxygen, extreme negative temperatures
and space vacuum.

Ranunculus
peltatus

Isoetes sp.

Tardigrades (tardi=slow, grada=step)

are related to crustaceans, insects and
spiders and their name derives from
their slow walk over the substrate instead of swimming. Tardigrades do
not possess breathing or circulation
systems and may withstand extreme

Despite their large numbers, crustaceans are the less known organisms
in temporary ponds. In fact, when we
think of crustaceans, we recall shrimp,
lobsters, stone crabs and giant crabs
that live in the sea. Yet, there are many
crustaceans that live in freshwater!
There is a great diversity of crustaceans
that live in temporary ponds, and there
exhibit special adaptations and survival
strategies.

Tardigrade

Crustaceans from temporary ponds

THORAX
HEAD

Body regions of
the crustaceans

ABDOMEN

The community of crustaceans that
inhabits temporary ponds is of paramount importance because it maintains
water quality, controlling the growth of
algae, regulating water transparency and
the type of available nutrients.

What is a crustacean?
Crustaceans are arthropods, such as
insects and spiders. The body is formed
by several articulated segments that are

grouped in three regions: head, thorax

and abdomen. The name crustacean derives from the Latin crusta, since these
animals have the body covered by an
external skeleton or exoskeleton, that
forms a crust. This external skeleton is
replaced when the animal grows through
a process called ecdysis or moult. When
animals hatch, crustaceans have a typical larval form called nauplius that undergoes several development stages.

NAUPLIUS 5

NAUPLIUS 6

NAUPLIUS 4

NAUPLIUS 3

NAUPLIUS 2

NAUPLIUS 1

10

Naupliar development
stages

One of the main characteristics of

crustaceans is the presence of several
pairs of appendages that, with the exception of the first antennae, are divided into two branches or ramous, that is,
they are biramous.
Par of biramous
appendages

INTERIOR RAMUS
OR ENDOPOD

EXTERIOR RAMUS
OR EXOPOD

Crustaceans that swim and live in

the water column are named planktonic.
When they live associated to plants and
on the bottom of the pond are named
benthic.
Crustaceans that live in temporary
ponds may be grouped according to
their dimension: The micro-crustaceans
that can only be observed under the microscope or stereomicroscope, and the
macro-crustaceans that are visible to the
naked eye.

Micro-crustaceans
The micro-crustaceans are the most
abundant crustaceans in ponds and
never exceed 5 mm in length. These
crustaceans are divided into three major
groups: copepods, small branchiopods
and ostracodes.

COPEPODS

from the Greek kope = oar

+ podo = foot
Copepods form a very diverse crustacean group, and are considered the most
abundant pluricellular organisms in the
planet, surpassing even insects in the
number of individuals. There are copepods living as parasites of fish and aquatic
mammals, and free living copepods that
are very numerous in temporary ponds.
Free living copepods belong to the orders
Calanoida, Cyclopoida and Harpacticoida
and normally have 1 to 2 mm in length,
cylindrical body and round head. The
adults keep the central eye from the
larval stage, which inspired the name
of the main freshwater genera Cyclops,
by analogy to the monster described in
Homers Odyssey.
Copepods may escape at great speed
from predators to avoid capture. In reality, they are the fastest animals on the
planet, and may cover a distance 500
times their body length, per second,
which means that 1 mm animals reach
1.8 km/h. Comparing with the cheetah,
that is the fastest animal on land (120
km/h), the cheetah would have to reach
2 700 km/h to be as fast as copepods.

Calanoida
Calanoid copepods are planktonic
animals with long antennae and the
ability to produce resting or resistant
eggs. The eggs are transported in a sac
until the female releases them. Several
species of various dimensions inhabit
temporary ponds.
In temporary ponds from the coastal
region of continental Portugal, the most

11

common species are Dussartius baeticus,

Diaptomus kenitraensis and Hemidiaptomus roubaui. In some ponds we can also
find Copidodiaptomus numidicus that was
very successful in colonizing dam reservoirs and is very abundant in the reservoirs of central and southern Portugal.

Hemidiaptomus
roubaui

Hemidiaptomus
(Gigantodiaptomus) roubaui
(Richard 1888)
Maximum Length: 6 mm
It is the largest and most colourful
copepod in Portugal. It is usually reddish to blue because of the presence of
carotenoid pigments (such as the ones
giving colour to carrots and tomatoes).
After the first rains of autumn, the resting eggs in the sediment hatch originating the single annual generation
inhabiting the ponds from November to
February.

12

Dussartius baeticus (Dussart 1967)

Maximum Length: 1.23 mm
It is a small filter feeding copepod of
bluish colour that inhabits ponds in the
south-western Iberian Peninsula, from
where it is endemic.

Dussartius
baeticus

Diaptomus kenitraensis
(Kiefer 1926)
Maximum Length: 2.6 mm
It is a copepod that usually accompanies Hemidiaptomus roubaui in the
coastal ponds of southern Portugal. It
is generally orange in colour and it is
omnivorous.

Diaptomus
kenitraensis

Cyclopoida

Acanthocyclops robustus
(G.O. Sars 1863)
Maximum Length: 1.7 mm
It is the cyclopoid with the largest
distribution in Portuguese ponds and
reservoirs.

Cyclopoid copepods are both planktonic and benthic. They have shorter
antennae than calanoids and females
carry two sacs of eggs that hatch a few
days after laying. When the ponds dry,
the larval stages and adults of some
species migrate to the sediment where
they fold upon themselves and enter a
stage or dormancy or diapause. In each
pond there are usually three to six species of cyclopoid copepods varying from
0.8 mm of Tropocyclops prasinus to 3 mm
of the species Megacyclops viridis and
Macrocyclops fuscus.
Macrocyclops fuscus
(Jurine 1820)
Maximum Length: 3.1 mm
It is a large copepod and adults are
carnivorous, frequently preying upon
other crustaceans and insect larvae.
Females carry two egg sacs with numerous eggs that hatch a few days after
laying.

Harpacticoida

U. Hopp

Macrocyclops fuscus

Cyclopoid
copepod,
Acanthocyclops
robustus

Harpacticoid copepods have generally short antennae and their body is

more cylindrical (or worm like) than
that of cyclopoids. They usually live
associated with the sediment and submerged plants. When preparing to enter
the resting stage or diapause, the larval stages or adults may exude mucus
to which particles aggregate, and thus
form a cyst that protects them when the
pond dries. In ponds, the most common
harpacticoids belong to the species Attheyella (Neomrazekiella) trispinosa,
Bryocamptus minutus and Canthocamptus
microstaphylinus.

13

Attheyella
trispinosa

Attheyella (Neomrazekiella)
trispinosa (Brady 1880)
Maximum Length: 0.95 mm
Lives associated with submerged
plants and may swim in the water column.

BRANCHIOPODS

from the Greek branchia + podos =

gills on feet
The class Branchiopoda is considered the most primitive of all crustacean
classes. It is assumed that this group
evolved in the sea, and was forced to
move to temporary ponds and hypersaline (salty) lakes to avoid predation
when fish appeared. These crustaceans
are very sensitive to the degradation
of water quality and are only found in
well preserved ponds. All crustaceans
breathe oxygen dissolved in the water
through the body surface, yet branchiopods breathe also through their
limbs that have a flat appearance and

14

are called phyllopods. Branchiopods

are divided into two groups: the large
branchiopods that may reach 80 mm
in length, and the small branchiopods
that never exceed 5 mm in length. The
small branchiopods are usually called
cladocerans, resemble a flattened disc
and have one single compound eye. The
most well known cladoceran belongs to
the genus Daphnia. Daphnia are usually
called water fleas because they swim
propelled by a pair of antennae on their
head and seem to jump in the water, as
terrestrial fleas do on land.
Unlike the large branchiopods that
are found exclusively in continental waters (temporary ponds and hyper-saline
lakes), the cladocerans are also marine.
As a consequence of their transparent
exoskeleton, it is possible to observe on
the microscope all the body parts that
constitute the animal. It is possible to
see the beating heart at the dorsal region and the embryonic development
in the incubating chamber. When the
juveniles hatch they are a miniature of
the adult.
These organisms are very sensitive to
aquatic environmental changes and are
used as bio-indicators, that is, they are
used to evaluate the toxicity of water in
areas affected by effluents of industrial,
agricultural or domestic origin.
In Portugal, there are many cladoceran species in ponds and, up until now,
seven species of Daphnia, three species
of Ceriodaphnia and two species of Simocephalus were identified.

It is an Iberian endemism that occurs

in weakly mineralized waters. The body is
oval and laterally compressed. The head
has a large helmet (cephalic helmet) that
is typically rounded.

Chydorus
sphaericus

Daphnia obtusa (Kurz 1875)

Maximum Length: 2.5 mm
This species of Daphnia shares several
ponds of the south-western coast with
Daphnia hispanica.

Chydorus sphaericus
(Mller 1776)
Maximum Length: 0.23 mm
It is the smallest known crustacean
in Portuguese ponds. The animals usually have the shape of a sphere flattened
laterally. Unlike Daphnia that are strictly
planktonic, Chydorus may swim in the
water column but usually live associated
with submerged plants or on the bottom
of ponds (they are benthic).
Daphnia (Ctenodaphnia) hispanica
(Glagolev & Alonso 1990)
Maximum Length: 3.5 mm

A. Petrusek

Daphnia
hispanica

Daphnia obtusa

15

Ceriodaphnia reticulata
(Jurine 1820)
Maximum Length: 1.3 mm
The body is shaped like a globe and
has a reddish colour when living in
transparent water. It has a very large
compound eye.

Scapholeberis rammneri
(Dumont y Pensaert 1983)
Maximum Length: 2.5 mm
It is a rare crustacean in Portugal.
The ventral part of the carapace forms
a flat surface with setae (like feathers),
which allows it to swim inverted at the
water surface; it swims with the belly
up. The ventral area is heavily pigmented
from brown to red, which offers protection against ultra-violet radiation when
the animals swim beneath the water
surface.

Scapholeberis
rammneri

A. Guilln, Proyecto Agua

Ceriodaphnia reticulata

Simocephalus exspinosus
(DeGeer 1778)
Maximum Length: 2.8 mm
It feeds mainly on detritus and may
live in the ponds during the whole flood
period, associated with the submerged
vegetation.

OSTRACODES

from the Greek strakon = shell

They have a laterally flattened body
protected by a carapace similar to that
of a clam. Ostracodes are the most abundant crustaceans in the fossil record.
They appear in sedimentary rocks since
the Cambric (between 540 and 500 million years ago) until our days. Ostracodes have a widespread use in palaeontolOstracode

Simocephalus exspinosus

16

ogy because of their large geographical

distribution, small dimension and good
preservation of carapaces.
Some ostracodes have a light organ
that produces chemical luminescence.
Most of them use light as a defence
against predators and some may use it
for mating.
Ostracodes are poorly known in Portugal where a species inventory is lacking. In general, a pond is inhabited by
three to seven species.

Macro-custaceans
Large Branchiopods

ANOSTRACA

These transparent animals are usually called fairy shrimp because of

their delicate appearance. Characteristically they do not possess a carapace

J.F. Cart

The macro-crustaceans of temporary

ponds in Portugal are the large branchiopods.

The large branchiopods differ from

the small branchiopods (cladocerans)
because of the presence of a pair of compound eyes and a larger number of limbs.
The time required for these organisms to complete their life-cycle differs
according to species. Species like Tanymastix stagnalis can complete their cycle
in ponds of short flooding period while
species of the genera Triops and Cyzicus
require a long flood period (more than
4 months) to complete their life-cycle.
In Portugal, the large branchiopods
are represented by three sub-groups or
orders:

Compound eyes of Triops

17

Chirocephalus
diaphanus

Chirocephalus diaphanus
(Desmarest 1823)
Maximum Length: 13 mm
It is a Mediterranean species widely
distributed in Europe. The animals are
translucent, with red tips in the abdomen and limbs. Its life cycle is very fast,
and can only survive in the absence of
predators. The eggs tolerate complete
dryness and hatch when in contact with
water.

Tanymastix stagnalis
(Linnaeus 1758)
Maximum Length: 9 mm
Its the fastest growing branchiopod
that inhabits short duration ponds with
only a few centimetres depth. They live
in temporary ponds all over Europe, yet
nowadays they are threatened by the

18

J.F. Cart

protecting their long and segmented

body. They have a large number of swimming legs. They swim inverted, with the
dorsal region directed to the bottom
and the ventral region facing the water
surface.
Several species are represented in
Portugal: Chirocephalus diaphanus,
Branchipus cortesi, B. schaefferi, Streptocephalus torvicornis, Tanymastigites
sp. and Tanymastix stagnalis.

disappearance of shallow ponds. Females

transport their very colourful eggs in a
spherical sac.
NOTOTRACA

The notostracans are considered

living fossils because they maintained
the same body shape since the Triassic,
around 220 million years ago. They are
present in all continents with the exception of Antarctica and live at the bottom
of temporary ponds and brackish lakes.
In Portugal there are three species of notostracans: Triops baeticus, T. vicentinus
and Lepidurus apus.
The most known and numerous genus is Triops. The first larval stages are
herbivorous and planktonic, when they
grow they start to capture small organisms and the adults may hunt other
macro-crustaceans, insects and even
amphibian larvae. They have a very fast
development and may reach the adult
stage within a week after hatching. Their
main predators are birds, but nowadays
the exotic Louisiana crayfish is becoming
very abundant and is a serious threat to
these crustaceans.

Tanymastix
stagnalis

Triops baeticus

Triops baeticus (Korn 2010)

Maximum Length: 80 mm
Triops baeticus is the largest crustacean in Portuguese ponds that frequently shares with other large branchiopods
like Chirocephalus diaphanus and Cyzicus
grubei.
SPINICAUDATA

These animals are characterized by

the presence of a carapace that has two
valves or shells and covers the whole
body in a fashion similar to the shell of
clams. They spend most of the time at the
bottom of ponds and may partially bury
in the sediment. The organisms react to
danger contracting the muscle of the
valves to close them tight and remaining
motionless at the bottom until the danger disappears. Unlike most crustaceans,
the Spinicaudata dont substitute their
carapace completely during the several
moults, which leads to growth lines in
their shells. They feed mostly on detritus and algae that they filter at the

bottom and around submerged plants.

In Portugal, two species are present:
Cyzicus grubei and Magrebestheria maroccana.
Cyzicus grubei (Simon 1886)
Maximum Length: 12 mm
It is the branchiopod that requires
longer to complete its life-cycle (up to
120 days). Their populations are rare
and localized. Cyzicus populations, like
those of the genus Triops inhabit the
earth since the Jurassic.

19

Cyzicus grubei

Triops
cancriformis

Crustaceans from the

time of Dinosaurs!
Before the dinosaurs appeared,
crustaceans with the shape of Triops and
Cyzicus already existed, and some fossils
with forms similar to these crustaceans
date back to 416-359 million years ago
(Devonian period).
Triops cancriformis is one the most
ancient species of animals that inhabit
the earth with the same body form since
the Carboniferous period (359-299 mil-

20

 J. F. Cart
Hans Hillewaert

Atlantic
horseshoe
crab, Limulus
polyphemus
(Linnaeus 1758)

lion years ago). The horseshoe crab is

the only known species that is older than
Triops, and this species also suffered
few alterations in the last 450 million

of years. Curiously, and although it is an

arachnid and much larger that Triops, the
horseshoe crab is extraordinarily similar
to this crustacean.

21

Dynamics of temporary ponds

In temporary ponds, species succeed each other during the hydrological period. Some species such as the
copepod Hemidiaptomus roubaui or the
large branchiopod Tanymastix stagnalis establish themselves after the first
rains. Other species appear only after a
few weeks of flooding such as Daphnia
and the small copepod Dussartius baeticus. Crustaceans may arrive at the pond
from other places, brought by birds or
other visiting organisms. However, usually the crustaceans in a pond originate
from resistance forms that are stored
in the sediment at the bottom of that
pond.

Autumn
During the first rains in Autumn, the
ponds begin to fill and the crustaceans
that were trapped in the sediment try
to rapidly dominate the habitat.
The resistance eggs hatch and from
them exit:
Miniatures of adults in the case of
cladocerans that try to establish
large populations.
or
Larval forms named nauplius in the
case of copepods and large branchiopods.

22

The females of the cladoceran Daphnia hatch from resistance eggs and procreate without fertilization from males
they form clones of females through a
type of reproduction called without fertilization or parthenogenesis. Males only
appear when the environmental conditions are not favourable.
Hatching of
Daphnia from a
resistance egg

Copepod larval forms awake from

their dormant stage (diapause) and become active in the ponds.
When the females of copepods that
entered diapause in the previous season

resume their active life, they start immediately to produce hundreds of eggs. This
ability gives them advantage over species
that spent the dry period as larval stages
and still have to develop to the adult stage
before starting to reproduce.

Larval stages of
Triops sp.

24 HOURS

15 HOURS

48 HOURS

7 HOURS

3 HOURS

23

Winter
When the pond fills at the beginning of
winter, the micro-crustaceans and the macrocrustaceans reach their maximum diversity.

Scapholeberis rammneri
(Dumont & Pensaert 1983)

Dussartius baeticus
(Dussart 1967)

Macrocyclops fuscus
(Jurine 1820)

Triops baeticus
(Korn 2010)

Simocephalus
exspinosus
(DeGeer 1778)

Macro-crustaceans
Micro-crustaceans

24

Chirocephalus diaphanus
(Prevost 1803)

Daphnia hispanica
(Glagolev & Alonso 1990)

Hemidiaptomus roubaui
(Richard 1988)

Ceriodaphnia reticulata
(Jurine 1820)

Ostracoda
Cyzicus grubei
(Simon 1886)

25

Spring

Reproduction of
Daphnia sp., that
includes a parthenogenic
and sexual cycle

Certain crustaceans like Daphnia

have different reproductive strategies
according to the environmental conditions. When it is necessary to build large
populations to colonize the pond, the
females that hatch from resistance eggs
procreate without being fertilized by
males they form clones of females using a type of asexual reproduction without fertilization or parthenogenesis.
When the pond begins to dry and
the environmental conditions degrade,
females continue to produce eggs by

parthenogenesis that now originate

males. The males are formed only when
there isnt sufficient food, the environmental conditions are adverse or there
is overpopulation. Under these conditions, females start to produce eggs
that when fertilized by males originate
eggs that are encased in a thick wall and
become resistance eggs or ephippia. In
the next pond cycle, the resistance eggs
will hatch into females and all restarts.
Ephippia (plural of ephippium) ensure
the survival of populations during the
dry periods and may also be dispersed
by the wind or birds visiting the pond.

PARTHENOGENETIC REPRODUCTION

SEXUAL REPRODUCTION

Adverse
environmental
conditions

26

Summer
As water disappears from the pond,
crustaceans prepare themselves to face
the period of drought, before becoming
food to insects, amphibians and birds.
During the dry period, crustaceans
remain as resting stages, in a state of
latent life or diapause. Most of them
remain at the bottom of the pond as resistance eggs, yet some species remain
dormant as resistant stages in a state
of suspended life.
The resistance eggs are very resistant
to drought. Some eggs of Triops may be
viable after 15 years in the sediment
which may have contributed to the survival of the genus on earth while dinosaurs disappeared millions of years ago.

How do pond animals

feed?
The origin of the energy in the food
or trophic web and the way by which energy is transferred between the links in
the web is one of the most interesting
questions in ecology. In aquatic trophic
webs, the energy may either originate
from the sun and the producers are
plants (or algae), or from organic matter of terrestrial origin that arrives at the
aquatic ecosystem.

TROPHIC WEB
Is the way to express the food
relationships among organisms of a
community/ecosystem, starting at
the producers (plants or bacteria) and
connecting with herbivores, predators
and decomposers. There is a transfer of
energy through the trophic web under the
shape of nutrients (the available energy
diminishes along the trophic web), always
from producers to decomposers.

Egg of
Tanymastix
stagnalis

Egg of
Chirocephalus
diaphanus

In the classic pond trophic web, the

sun is the source of energy. The energy
is used by algae belonging to the phytoplankton to produce organic matter
(biomass) that serves as food to herbivores (consumers). Detritus in suspension, dissolved organic matter and waste
material from the feeding process are
incorporated by bacteria that will serve
as food to protozoans and thus form a
microbial loop in the classic web. The
protozoans also feed on algae and serve
as food to herbivores.

27

H ER B I VO R ES

C ARNIVO RES

Protozoans

Dissolved organic matter

P ROTO Z OAN S

P H Y TO P L AN K TO N
Bacteria

MICROBIAL LOOP

D I S S O LV ED O RG AN I C MAT T ER

Aquatic trophic web that

includes the microbial
loop

28

Conservation of temporary ponds

Temporary ponds have been valued by human communities for millennia because of their functions as
water reservoirs for cattle drinking
water during the rainy season, and as
a source of fresh grass in the dry season. As natural ecosystems, temporary
ponds were used as resting and feeding
grounds by local and migratory aquatic
birds. Additionally, they serve as water purifying systems for groundwater
reserves. However, nowadays there
is a sharp decrease in the number of
Mediterranean temporary ponds. The
main problems of this habitat are its
short duration and small size, which
renders them unnoticeable in the landscape. Other problem derives from the
present agricultural and cattle raising
practices that no longer require the
services offered by ponds because of
the intensification of the production
means. Therefore, because of their

limited visibility and the lack of recognition of its value, the habitat is easily
destroyed or transformed.
The most common threats to ponds
result from deficient management as a
result of decrease in traditional agricultural practices, forestation and land
use incompatible with the presence of
these habitats such as removal of surrounding vegetation, extraction and
drainage of water, deepening of ponds
and its transformation into permanent
water reservoirs. Management of temporary ponds is directed towards restoring their ecological function, correcting
the negative effects of human activities
that disturb them.
Conservation measures are mainly
related to grazing management, control of agricultural activities in the surrounding areas and control of invasive
and exotic species (ex. ice-plants, wattles, Louisiana crayfish).

29

Recommendations
for biodiversity
conservation
in temporary ponds:
1. Ponds should not be disturbed when
full.
2. When the ponds dry, walking and
driving through them should be
avoided to prevent the destruction
of eggs and resistance stages stored
in the sediment.
3. Fish should not be introduced in the
ponds since they feed on crustaceans
and destroy their populations.
4. The pond area should not be used in
agriculture.
5. Fertilizers should be avoided in the
ponds surrounding areas. Fertilizers
stimulate the growth of algae that are

30

responsible for the increase in water

turbidity and oxygen depletion, thus
causing the death of aquatic animals.
6. The natural vegetation surrounding
the ponds should be kept.
7. Avoid levelling the ground of ponds.
8. The ponds should not be deepened.
9. Temporary ponds should not be
transformed into permanent water
reservoirs.
Remember that this habitat is very
vulnerable and has in its sediments
a unique fauna.

REFERENC ES

P RO M O T O R S

Alonso M. (1996) Crustacea. Branchiopoda. In Ramos

M.A. (Ed) Fauna Iberica, vol. 7, Museo Nacional de
Ciencias Naturales, CSIC, Madrid. 486 p.
Quzel P. (2004) In Gauthier P., Yavercovski N.,
Perennou C. (Eds) Mediterranean Temporary Pools,
Volume 1 Issues relating to conservation, functioning
and management. Station biologique de la Tour du
Valat, Arles, France. 122 p.

JULY 2013
AUTHOR S

Maria Jos Caramujo

Carina Cunha
Carla C.C.R de Carvalho
Cristina Lus

F U N D I NG

EXEC UTI VE PRODUC TI O N

Marta Fonseca
PHO T OGRAPHY

Carla C.C.R. de Carvalho

with the exception of photos where
other authors are indicated

PA R T N E R S

I LLUS TRATI ONS

Maria Jos Caramujo

Daphnia adapted from Miguel Alonso (1996)
GRAPHI C DESI GN

TVM designers
I SBN

978-989-98300-2-8
ACKNOWLEDG EMENT S

Adam Petrusek
Antonio Guilln, Proyecto Agua
Hans Hillewaert
Joo Petronilho
Jean Franois Cart
Lus Cancela da Fonseca
Margarida Machado
Patrcia Garcia Pereira
Ulrich Hopp

SUPPORT