Freshwater Large Branchiopods in Portugal · Dos 505 sítios, apenas 87 estão localizados a Norte...

Limnetica 29 (2) x-xx (2011)Limnetica 36 (2) 567-584 (2017) DOI 1023818limn3622ccopy Asociacioacuten Ibeacuterica de Limnologiacutea Madrid Spain ISSN 0213-8409

Freshwater large branchiopods in Portugal an update of theirdistribution

Margarida Machado1 Luiacutes Cancela da Fonseca34 and Margarida Cristo12lowast

1 CCMAR - Centro de Ciecircncias do Mar Universidade do Algarve Campus de Gambelas 8005-139 Faro Portu-gal2 FCT - Faculdade de Ciecircncias e Tecnologia Universidade do Algarve Campus de Gambelas 8005-139 FaroPortugal3 MARE - Marine and Environmental Sciences Centre Laboratoacuterio Mariacutetimo da Guia Av N Sra do Cabo 939750-374 Portugal4 CTA - Centro de Ciecircncias e Tecnologias da Aacutegua Universidade do Algarve Campus de Gambelas 8005-139Faro Portugal

lowast Corresponding author mcristoualgpt2

Received 311016 Accepted 120517

ABSTRACT

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 otherresearchers During this period several studies dealing with freshwater large branchiopods (FLB) were carried out resultingin scientific publications and project reports The distribution of FLB in Portugal was presented in 2 international scientificmeetings but apart from a first paper by Vianna-Fernandes in 1951 and an update done by ourselves in 1999 concerningthe southwest Portugal no other information has been published Therefore this work intends to bring up to date the knowndistribution of this faunal group in freshwater temporary systems This is pertinent because of the recent revision of the taxonTriops cancriformis on the basis of genetic analyses The Portuguese populations were assigned either to the Portugueseendemism T vicentinus or to T baeticus the more widely spread Iberian species both belonging to the T mauritanicuscomplex Furthermore a new species Tanymastigites lusitanica was found and described in Portugal More recently a malefree (or at least a strongly female-biased) metapopulation of T cancriformiswas discovered in a rice field in the central regionof Portugal Here we present an updated status of FLB species of Portuguese temporary lentic systems and their distributionplotted on a UTM (10 times 10 km) grid A total of 505 sites (temporary ponds or assemblages of closely located not individualtemporary pools) have been surveyed (455 by us) In 241 of these (477) at least one species of FLB was found on at leastone occasion Of the 505 sites only 87 are located north of the Tagus River and of these at least one species was found inonly 17 (195) South of the Tagus River (Alentejo and Algarve) 536 of the sites were inhabited by at least one speciesThis list comprises 7 anostracans 2 spinicaudatans and 4 notostracans including 3 species endemic to the Iberian Peninsulaand another 3 to Portugal

Key words Temporary ponds anostracans notostracans spinicaudatans endemic species conservation status

RESUMO

Grandes branquioacutepodes dulccedilaquiacutecolas em Portugal actualizaccedilatildeo da sua distribuiccedilatildeo

O presente trabalho eacute maioritariamente o resultado de 20 anos de trabalho de campo e de laboratoacuterio com campanhas re-alizadas pelos autores e por outros investigadores Durante este periacuteodo foram realizados vaacuterios estudos de que resultaramdiversas publicaccedilotildees e relatoacuterios de projectos O conhecimento da distribuiccedilatildeo dos grandes branquioacutepodes dulccedilaquiacutecolas(GBD) em Portugal baseia-se num primeiro artigo de 1951 de Vianna-Fernandes e de uma actualizaccedilatildeo do conhecimentopublicada por noacutes em 1999 e relativa ao sudoeste de Portugal A sua actualizaccedilatildeo foi posteriormente apresentada em doisencontros cientiacuteficos internacionais mas nunca foi publicada Assim este trabalho pretende actualizar o conhecimento so-bre a distribuiccedilatildeo em Portugal deste grupo fauniacutestico de sistemas lecircnticos temporaacuterios Esta intenccedilatildeo eacute pertinente vistoque na uacuteltima deacutecada o taxon Triops cancriformis foi revisto com base em anaacutelises geneacuteticas As populaccedilotildees portuguesas

17140_Limnetica 36(2) pagravegina 567 28092017

568 Machado Cancela da Fonseca and Cristo

foram atribuiacutedas quer ao endemismo portuguecircs T vicentinus quer a T baeticus a espeacutecie com distribuiccedilatildeo mais alargadana peniacutensula ibeacuterica ambas anteriormente incluiacutedas no complexo T mauritanicus Aleacutem destas uma nova espeacutecie Tany-mastigites lusitanica foi descoberta e descrita em Portugal Mais recentemente foi encontrada num campo de cultivo dearroz na regiatildeo central de Portugal uma metapopulaccedilatildeo de T cancriformis aparentemente composta unicamente por fecircmeasApresenta-se neste trabalho o estado actual do conhecimento sobre a distribuiccedilatildeo dos GBD de sistemas lecircnticos temporaacuteriosportugueses em mapas de quadriacutecula UTM (10 times 10 km) Foram prospectados um total de 505 siacutetios (charcos temporaacuteriosou conjuntos de charcos com localizaccedilatildeo proacutexima natildeo individualizaacutevel) 455 pelos autores Em 241 deles (477) foi obser-vada a presenccedila de pelo menos uma espeacutecie de GBD numa ocasiatildeo Dos 505 siacutetios apenas 87 estatildeo localizados a Norte dorio Tejo e nestes apenas em 17 (195) foi detectada a presenccedila de pelo menos 1 espeacutecie de GBD A sul do rio Tejo (Alentejoe Algarve) em 536 dos siacutetios foi encontrada pelo menos uma espeacutecie de GBD Esta lista inclui 13 espeacutecies (7 Anostraca2 Spinicaudata e 4 Notostraca) entre as quais 3 endemismos ibeacutericos e outros 3 endemismos portugueses

Palavras-chave Charcos temporaacuterios Anostraca Notostraca Spinicaudata espeacutecies endeacutemicas estatuto de conservaccedilatildeo

INTRODUCTION

Large branchiopods are a group of crustaceanspecies (Branchiopoda Anostraca NotostracaSpinicaudata Laevicaudata and Cyclestherida)that inhabit almost exclusively temporary aquaticsystems (Brendonck et al 2008) They play animportant ecological role in lentic systems asdetritivores herbivores carnivores andor scav-engers contributing for example to the control ofmosquitoes and to mitigating eutrophication byfeeding on organic detritus and decaying organicmatter They are also important in the trophicchain as a food source for insects amphibiansand birds They are considered a flagship groupfor freshwater temporary lentic systems (FTLS)and for temporary pond invertebrates (Bren-donck et al 2008) Until a few decades agostudies on these systems were scarce (Hildrew1985 and references therein Boix et al 2012Jeffries et al 2016) most probably due to thegenerally small size of FTLS their ephemeralnature and the relative unpredictability of theduration and timing of their flooding periodsIn Europe the considerable increase of scien-tific interest in these environments over the lasttwo decades (Boix et al 2016b and referencestherein) has been favored by the classificationof Mediterranean Temporary Ponds as a priorityfreshwater habitat under Directive 9243CEEAs a consequence of the increasing scientific in-

terest in temporary ponds namely in their func-tioning and biodiversity many studies on fresh-water large branchiopods (FLB) from severalareas in the world have been published (egBrendonck et al 2008 and references therein)and many new species have been recently discov-ered and described (eg Belk amp Pereira 1982Thieacutery 1988 Brendonck et al 1992 Rogers2001 Belk amp Rogers 2002 Balaraman amp Nayar2004 Olesen amp Timms 2005 Cottarelli et al2007 Cohen 2008 Alonso amp Garcia-de-Lomas2009 Alonso amp Ventura 2013 Machado amp Sala2013 Rogers et al 2013 Timms 2013 Boixet al 2016a Korn amp Hundsdoerfer 2016) InPortugal the first published scientific paper deal-ing with FLB inhabitants of temporary pondsand pools (Vianna-Fernandes 1951) reportedthe occurrence of 4 new species the anostra-cans Tanymastix lacunae Daday 1910 (= T stag-nalis (L 1758)) and Streptocephalus torvi-cornis (Waga 1842) the notostracan Triopscancriformis mauritanicus (Ghigi 1921) andthe spinicaudatan Caenestheriella grubei Daday1915 (= Cyzicus grubei (Simon 1886)) With theexception of one specimen of Apus cancriformisSchaumlffer 1756 (= Triops cancriformis (Lamarck1801)) registered in the catalogue of the inver-tebrate collection of the Zoological Museum ofCoimbra University (Carvalho 1944) no FLBspecies were known to occur in Portugal untilthe publication of this paper

Limnetica 36 (2) 567-584 (2017)


Freshwater Large Branchiopods in Portugal 569

After a second paper by Vianna-Fernandespublished in 1957 the Portuguese FLB faunaremained forgotten during decades likely dueto the minor attention given to their ephemeralhabitats As in other European countries theclassification of Mediterranean Temporary Pondsas a priority habitat has contributed to drawingattention of Portuguese researchers to the FTLSand their communitiesThe work on the Portuguese FLB fauna was

restarted by the present authors 45 years afterthe 1951 publication of Vianna-Fernandes Sincethen several studies were carried out resultingin scientific papers and reports (Reis et al 1997Machado et al 1999a Machado et al 1999bCristo et al 2002 Grosso-Silva amp Soares-Viei-ra 2002 Cancela da Fonseca et al 2008 Pre-sado et al 2010 Gascoacuten et al 2012 Machadoamp Sala 2013 Sousa 2015) Some updates ofits distribution have been presented in 2 interna-tional scientific meetings (Machado et al 2004

Sala et al 2010) Nevertheless updated statusesof FLB in Portugal have not yet been publishedFurthermore in the last decade the taxon Tri-

ops cancriformis have been revised on the basisof genetic analyses In a first step Korn et al(2006) reinstated the subspecies cancriformisand mauritanicus to full species status and raisedto subspecies rank two Northern African lin-eages within T mauritanicus Later four moreIberian endemic lineages within T mauritan-icus were recognized and full species statuseswere attributed to all six T mauritanicus lin-eages (Korn et al 2010) The populations inPortugal were assigned either to the Portugueseendemism Triops vicentinus or to Triops baeti-cus the more widely distributed Iberian speciesThe only specimen of Apus cancriformis (= Tri-ops cancriformis) included in the crustaceancollection of the Zoological Museum of CoimbraUniversity (Carvalho 1944) proved to belong toT baeticus (Machado 2014) More recently a

Table 1 Species of FLB in the mainland of Portugal Total number of sites with FLB number of sites where each species co-occurswith n other species (higher values in bold) percentage of sites with each species in relation to the total number of sites with FLBnumber of sites with x number of species mean species number per site Espeacutecies de GBD em Portugal continental nuacutemero totalde siacutetios com GBD nuacutemero de siacutetios com coocorrecircncias de GBD (valores mais elevados a negrito) percentagem de siacutetios em queocorre cada espeacutecie em relaccedilatildeo ao nuacutemero total de siacutetios com GBD nuacutemero de siacutetios com x nuacutemero de espeacutecies nuacutemero meacutedio deespeacutecies por siacutetio

SpeciesNo of sites where the species co-occur with n other species Sites

Total no

Sites of Total

with FLBn = 0 n = 1 n = 2 n = 3 n = 4 n = 5

C diaphanus 41 31 24 21 3 1 121 502

T stagnalis 18 43 9 13 2 1 86 357

B cortesi 12 38 9 11 3 0 73 303

T baeticus 8 22 16 16 3 1 66 274

C grubei 1 7 10 14 1 1 34 141

T vicentinus 1 7 11 4 0 0 23 95

Tanymastix sp 1 8 1 4 2 0 0 15 62

B schaefferi 7 2 1 2 0 1 13 54

S torvicornis 2 0 3 6 1 1 13 54

T lusitanica 8 1 0 0 0 0 9 37

M maroccana 0 3 1 1 1 0 6 25

T cancriformis 1 0 1 0 0 0 2 08

L apus 0 1 0 0 0 0 1 04

No of species per site X = 1 X = 2 X = 3 X = 4 X = 5 X = 6 Total

No sites with FLB 107 78 30 22 3 1 241

Mean no of speciessite 19

Limnetica 36 (2) 567-584 (2017)



male free (or at least a strongly female-biased)population of T cancriformis was discoveredin a rice field in the central region of Portugal(Machado et al 2016)This study is the result of 20 years of survey

efforts during which the environmental charac-terization of the FTLS was carried out on mostoccasions This characterization comprisingestimation of the plant coverage and measure-ments of pond surface and depth temperaturepH oxygen concentration percentage of dis-solved oxygen conductivity and turbidity of thewater has been mostly presented and discussedby Cancela da Fonseca et al (2008) Neverthe-less the distributions of the FLB in Portugalhave not been updated since the survey of largebranchiopods in the southwest part of the coun-try published in 1999 (Machado et al 1999a)which was the start of the systematic inventoryof these organisms Since then a huge amountof FLB new locations were recorded Conse-quently this study aims to present the updatedFLB inventory and distribution in Portugal andto evaluate the current conservation status of thePortuguese FLB fauna

MATERIAL ANDMETHODS

Data on presenceabsence of Freshwater LargeBranchiopods were mainly obtained from sev-eral surveys carried out from March 1996 toApril 2016 Many of the FTLS were sampledduring the course of several projects both in-stitutional and personal Some were discoveredbased on personal information or through pub-lished work (eg Grosso-Silva amp Soares-Vieira2002 Sousa 2015) Others were detected fromsatellite images from Google Earth sometimescomplemented by the inspection of militarycharts with located ponds or even by chancewhen in the field The FTLS were visited at leastonce during the flooded period and wheneverpossible close to its beginning The geographiccoordinates of Portuguese freshwater temporarylentic systems with FLB species were registeredusing either a Magellan GPS ProMARK XTMor a Garmin-GPS72

Figure 1 Distribution of Branchipus cortesi and Branchipusschaefferi plotted on the map of Portugal on a UTM (10 times 10km) grid baseDistribuiccedilatildeo de Branchipus cortesi e Branchipusschaeffer em Portugal (quadriacutecula UTM-10 times 10 km)

Sampling of the FLB community was per-formed by trawling with a round hand net(305 cm mouth diameter 1 mm2 mesh size) inall possible microhabitats FLB were identifiedby eye to genus level in the field Neverthe-less from each FTLS and at least once a small

Limnetica 36 (2) 567-584 (2017)



sample of each species found was fixed in 4formaldehyde or in 70-998 ethanol and takento the lab for identification to species level Al-though some FTLS were not sampled by us allthe collected FLB were studied by ourselvesThe identification was based on Cottarelli amp

Mura (1983) Thieacutery (1987) Petkovski (1995)Alonso (1996) Brtek amp Mura (2000) Korn et al(2006 2010) and Machado amp Sala (2013) Thenomenclature followed Brtek (2002) Weekers etal (2002) and Machado amp Sala (2013)The collected data were complemented by

the information published by Vianna-Fernandes(1951) Grosso-Silva amp Soares-Vieira 2002Korn et al (2006 2010) Cancela da Fonseca etal (2008) and Machado et al (2016)Amapwith a UTMgrid (10 times 10 km) adapted

from Franco amp Afonso (1982) was georefer-enced into the Portuguese coordinate system(Portuguese_National_Grid Projection Trans-verse_Mercator Datum D_Lisbon) using thesoftware package ArcGIS 92 (ESRI 2006) Thegeographic coordinates of all sites where at leastone FLB species has been found were inserted ina Geographic Information System as an ArcGISdatabase (Booth amp Mitchell 2001) and plottedon the map

RESULTS

A total of 505 sites (temporary ponds or assem-blages of closely located not isolated temporarypools) were surveyed (455 by us) in mainlandPortugal In 241 of those (477) at least onespecies of FLB was found on one occasion Ofthe 505 sites only 87 are located North of TagusRiver and at least one species was found in17 (195) of these South of the Tagus River(Alentejo and Algarve) 536 of the sites wereinhabited by at least one FLB speciesA total of 13 FLB species were identified

7 anostracans 4 notostracans and 2 spinicau-datans A maximum of six coexisting specieswere found in only one FTLS and the mean num-ber of co-occurring species was 19 (Table 1)Data on the current occurrence of these speciesare detailed in Table 1 The inventory of Fresh-

water Large Branchiopods in Portugal mainlandis

Class Branchiopoda Latreille 1816

Order Anostraca Sars 1867

Family Branchipodidae Milne-Edwards 1840

Genus Branchipus Schaeffer 1766

Branchipus cortesi Alonso amp Jaume 1991

Branchipus schaefferi Fischer 1834

Family Chirocephalidae Daday 1910

Genus Chirocephalus Preacutevost 1820 (in Jurine)

Chirocephalus diaphanus Preacutevost 1820 (inJurine)

Family Streptocephalidae Daday 1910

Genus Streptocephalus Baird 1852

Streptocephalus torvicornis (Waga 1842)

Family Tanymastigidae Brtek 1972

Genus Tanymastigites Brtek 1972

Tanymastigites lusitanicaMachado amp Sala 2013

Genus Tanymastix Simon 1886

Tanymastix stagnalis (Linnaeus 1758)

Tanymastix sp1

Order Notostraca Sars 1867

Family Triopidae Keilhack 1909

Genus Lepidurus Leach 1819

Lepidurus apus (Linnaeus 1758)

Genus Triops Schrank 1803

Triops cancriformis (Lamarck 1801)

Triops baeticus Korn 2010

Triops vicentinus Korn Machado Cristo ampCancela da Fonseca 2010

Order Spinicaudata Linder 1945

Family Cyzicidae Stebbing 1910

Genus Cyzicus Audouin 1837

Cyzicus grubei (Simon 1886)

Family Leptestheriidae Stebbing 1902

GenusMaghrebestheria Thieacutery 1988

Maghrebestheria maroccana Thieacutery 1988

Limnetica 36 (2) 567-584 (2017)



Figure 2 Distribution of Chirocephalus diaphanus plottedon the map of Portugal on a UTM (10 times 10 km) grid base Dis-tribuiccedilatildeo de Chirocephalus diaphanus em Portugal (quadriacuteculaUTM-10 times 10 km)

Figures 1 to 6 show the distribution of these13 FLB species plotted on the map of Portugalon a UTM (10 times 10 km) grid baseThe most common species is C diaphanus

followed by T stagnalis and B cortesi The rarestspecies (lt 10 sites) are L apus T cancriformisM maroccana and T lusitanica (Table 1)

Figure 3 Distribution of Streptocephalus torvicornis andTanymastigites lusitanica plotted on the map of Portugal on aUTM (10 times 10 km) grid base Distribuiccedilatildeo de Streptocephalustorvicornis e Tanymastigites lusitanica em Portugal (quadriacuteculaUTM-10 times 10 km)

Streptocephalus torvicornis shows a scattereddistribution in the Alentejo and Algarve whereasTanymastigites lusitanica was found in only 7UTM grid squares located in a small central areaof South Alentejo (Fig 3)

Tanymastix sp1 is a new species for sciencewhich is currently being described (M Machado

Limnetica 36 (2) 567-584 (2017)



Figure 4 Distribution of Tanymastix stagnalis and Tanymas-tix sp1 plotted on the map of Portugal on a UTM (10 times 10 km)grid base Distribuiccedilatildeo de Tanymastix stagnalis e Tanymastixsp1 em Portugal (quadriacutecula UTM-10 times 10 km)

and J Sala unpublished data) It is roughly con-fined to a rectangular area of 1500 km2 locatedin the central part of southern Alentejo and thenorthern Algarve (Fig 4)

Triops baeticus is the species of this genuswith the largest geographical distribution in Por-tugal whereas Triops vicentinus shows a distri-bution limited to the west and central Algarve

Figure 5 Distribution of Lepidurus apus Triops cancri-formis Triops baeticus and Triops vicentinus on the map ofPortugal on a UTM (10 times 10 km) grid base Distribuiccedilatildeo deLepidurus apus Triops cancriformis Triops baeticus e Triopsvicentinus em Portugal (quadriacutecula UTM-10 times 10 km)

and Triops cancriformis is known only from 2close sites in Portugal Lepidurus apus is therarest FLB species in Portugal being knownonly from 1 FTLS a small shallow puddle cor-responding to flooded tracks left by farm tractorsin the middle of a cultivated field (Fig 5)

Limnetica 36 (2) 567-584 (2017)



Figure 6 Distribution of Cyzicus grubei and Maghrebesthe-ria maroccana plotted on the map of Portugal on a UTM(10 times 10 km) grid base Distribuiccedilatildeo de Cyzicus grubei eMaghrebestheria maroccana em Portugal (quadriacutecula UTM-10times 10 km)

The current known distribution of Cyzicusgrubei in Portugal is limited to the southern partof the country (southern half of Alentejo and Al-garve) Maghrebestheria maroccana has beenfound in only 6 sites in a very restricted Alentejoarea (Fig 6)

Table 1 highlights the difference in the high-est number of sites where n other species co-oc-curred with each species Most anostracan spe-cies were mainly found alone with the exceptionof T stagnalis and B cortesi which normallyshared their biotope with 1 more FLB speciesas did the notostracan T baeticus and the spini-caudatan M maroccana The notostracan T vi-centinus was mostly found together with 2 morespecies and the anostracan S torvicornis aswell as the spinicaudatan C grubei with 3 morespeciesIn Table 2 the FLB species richness found

in Portugal and also South of the Tagus Riveris compared with that reported from otherEuropean Mediterranean and west Asian coun-triesregions from the Palaearctic

DISCUSSION

During the first half of the last century the oc-currence of only 4 freshwater large branchiopodtaxa (2 Anostraca 1 Spinicaudata and 1 Noto-straca) was reported from Portugal (Carvalho1944 Vianna-Fernandes 1951) Their distribu-tion was restricted to four sites one north andvery close to the Tagus River (Azambuja) andthree (Torratildeo Beja and Castro Verde) in thecentral and south Alentejo (cf Machado et al1999a) The eventual existence of a fifth speciesCyzicus tetracerus (Krynicki 1830) althoughreported by Brtek amp Thieacutery (1995) who didnot mention the source of the information hasnot yet been confirmed in Portugal Since westarted our work on FLB in the SW Portugal thenumber of species has increased from 4 to 7 (4Anostraca 2 Spinicaudata and 1 Notostraca)with greater understanding of their known distri-bution in the country and in the Iberian Peninsula(Machado et al 1999a)Nevertheless the percentage of the surveyed

area in the country was still very low (less than5) Nowadays the total surveyed area is approx-imately 70 of Portugal and we report here theexistence of 13 species (7 Anostraca 2 Spinicau-data and 4 Notostraca) 6 more species than 18years ago In general we found the same species

Limnetica 36 (2) 567-584 (2017)



in the same places (cf Machado et al 1999a)but their distribution has greatly expanded due tothe increased surveying (Figs 1 to 6)

FLB occurrence and distribution

Anostracans

B cortesi was mainly found in the southwestedge of the country where it was first discoveredin Portugal (Machado et al 1999a) Previouslyit had been reported in 40 out of 129 Portuguesesites with at least one FLB species (Cancela daFonseca et al 2008) This species is an Iberianendemism (Alonso 1996) It often co-occurswith T stagnalis (Cancela da Fonseca et al2008) It prefers FTLS with low conductivity(Alonso 1996 Cancela da Fonseca et al 2008)

B schaefferi has been reported from 5 moresites than the previously published data (Can-cela da Fonseca et al 2008) In agreementwith the latter study we found this species inpoorly-vegetated ponds It is a Palearctic species(Alonso 1996) widely distributed in the steppeand wood steppe zones of continental Europeand on the western Mediterranean islands of Sar-dinia Sicily and Malta (Brtek amp Thieacutery 1995)In Portugal as in southern Spain (Alonso 1985Boix et al 2007 Fahd et al 2009 Prunier ampSaldantildea 2010 Prunier et al 2011) the speciesis relatively rare with a scattered distributionNevertheless it is frequent in the northern partof Spain (Alonso 1996) being the most commonFLB species in Catalonia (Gascoacuten et al 2012)and in Comunitat Valenciana (Sahuquillo 2012)

C diaphanus is a euryoecious (eg Mura1999 Cancela da Fonseca et al 2008) Palearc-tic species (Alonso 1996) widely distributed inEurope (Brtek amp Thieacutery 1995) In Portugal it isthe most common anostracan as it is also in sev-eral other southern European countries such asFrance (Defaye et al 1998) Italy (Mura 2001)and Spain (Alonso 1996) In relation to the pre-vious published data (Cancela da Fonseca et al2008) the number of known sites increased from65 to 121 and the known distribution is expandedto the northeast of Portugal

T lusitanica is only known from the verysame sites that were reported when it was firstdescribed (Machado amp Sala 2013) It has alwaysbeen found in small and very shallow puddlesor rainpools on unpaved clayish or sandy roadsTypically the water is very turbid with lowmineralization and poorly colonized by aquaticvegetation (Cancela da Fonseca et al 2008Machado amp Sala 2013) This species the onlyrepresentative of the genus in Europe is alsoendemic to Portugal (Machado amp Sala 2013)

T stagnalis was previously reported from1 Portuguese site located in the central part ofAlentejo (Vianna-Fernandes 1951) and is nowa-days the only FLB species in Portugal with theknown occurrence increasing in percentage inrelation to the former published data (318to 357) Previously it was found in 41 sites(Cancela da Fonseca et al 2008) It is a Palearc-tic (Alonso 1996) widespread (Brtek amp Thieacutery1995) species which has been considered as acold stenothermal species (see Alonso 1995Mura 1999 2001) Nevertheless it often occursin water temperatures of 25-30 oC eg in Aus-tria (Eder et al 1997) and in Portugal (Cancelada Fonseca et al 2008) suggesting that it is aeurythermal species The data here presented ex-pands its distribution to the southwest edge ofIberian Peninsula (Alonso 1985 Machado etal 1999a)Since the last published data (Cancela da Fon-

seca et al 2008) only one more site was addedto the distribution of Tanymastix sp1 To date itseems that this species is also endemic to Portu-gal It prefers shallow pools with short hydrope-riods and relatively low water conductivity (Can-cela da Fonseca et al 2008)The distribution of S torvicornis has broad-

ened in 2 sites relative to the previously pub-lished data (Gascoacuten et al 2012) It prefers smalland deep ponds with moderate to long hydrope-riods and waters with relatively low conductivity(Cancela da Fonseca et al 2008) and seemsto avoid large FTLS It is a Eurasian speciesfound in the Mediterranean region Asia Minorand Central Asia (Alonso 1996) being one ofthe most common anostracans in some Mediter-ranean countries (Beladjal et al 2003) such as

Limnetica 36 (2) 567-584 (2017)



Morocco (Thieacutery 1986) and Algeria (Samraouiet al 2006) In Spain Alonso (1996) have re-ported it as very frequent in the hydrographic

sedimentary basins from the South namely inthe low Guadalquivir river region Neverthelessthis species seems to be rare nowadays (Boix et

Table 2 Total number of sites with FLB total number of FLB species maximum number of co-occurring species and mean speciesnumber per site (X) with FLB found in several studied European Mediterranean and west Asian countriesregions from Palearcticndash information not explicit and impossible to infer from references Area information (km2) whenever absent from references wasfound in the web Nuacutemero total de siacutetios com GBD nuacutemero de espeacutecies de GBD nuacutemero maacuteximo de espeacutecies que coocorrem enuacutemero meacutedio de espeacutecies por siacutetio (X) com GBD encontrados em diversos paiacutesesregiotildees paleaacuterticas europeias mediterracircnicas e doproacuteximo oriente mdash informaccedilatildeo natildeo explicita ou impossiacutevel de inferir dos dados das referecircncias A informaccedilatildeo sobre a aacuterea (km2)sempre que natildeo referida foi retirada da ldquowebrdquo

Geographical ZoneSites with FLB

SpeciesReference

Identification Area no Max nosite X

Iberian Peninsula 582 000 mdash 22 6 mdash Alonso (1996) Sala et al(2016)

Portugal (mainland) 89 000 241 13 6 19 Present studyPortugal (South of Tagus River) 39 850 224 12 6 20 Present studySpain (mainland) 493 000 63 15-16 5 15 Alonso (1985) Korn et al

(2006 2010)Spain (mainland) 493 000 mdash 19 6 mdash Alonso (1996) Korn et al

(2006 2010) Sala (perscom)

Girona Province 5600 75 6 3 12 Gascoacuten et al (2012)Comunitat Valenciana 23255 56 9 6 13 Rueda et al (2006) Zierold et

al (2007) Sahuquillo (2012)Forner i Valls amp Brewster(2013) Soria et al (2016)

Extremadura Province 41 643 25 7-8 4 17 Peacuterez-Bote et al (20062008) Korn et al (2010)

Coacuterdoba province 13 769 81 8 5 15 Prunier amp Saldantildea (2010)Prunier et al (2011)

Murcia Region 11 313 17 3 3 12 Verdiell-Cubedo amp Boix(2014)

Maacutelaga Province 7276 20 4 3 14 Rippol Rodriacuteguez et al(2013)

Dontildeana Natural Area 1200 25 8 6 21 Boix et al (2007) Fahd et al(2009)

Menorca 702 20 4 3 17 Boix et al (2010)France 635 121 mdash 14 mdash mdash Defaye et al (1998)Italy 251 537 mdash 14 mdash mdash Cottarelli amp Mura (1983)

Mura (2001) Beladjal et al(2003)

Former Yugoslavia 288 364 mdash 18 mdash mdash Petrov amp Petrov (1997)Banat Province (form Yugosl) 9300 68 10 7 26 Petrov amp Cvetkovic (1997)Austria (eastern) 42 000 approx mdash 15 mdash mdash Eder et al (1997)Belgium 30 528 mdash 7 mdash mdash Vanschoenwinkel et al

(2013)Denmark 42 400 mdash 6-7 mdash mdash Damgaard amp Olesen (1998)Morocco (West) 445 050 101 16 10 mdash Thieacutery (1986 1991)Morocco North and West) mdash 108 14 6 27 Van den Broeck et al (2015)Algeria 2 382 000 mdash 15 mdash mdash Beladjal et al (1995)

Samraoui et al (2006)Numidia (Algeria) 19 4 2 16 Samraoui amp Dumont (2002)Tunisia 163 610 mdash 11 mdash mdash Turki amp Turki (2010) Korn et

al (2006)Iran 1 648 000 15 9 4 16 Atashbar et al (2014)

Limnetica 36 (2) 567-584 (2017)



al 2007 Garciacutea de Lomas amp Garciacutea 2008 Fahdet al 2009 Prunier amp Saldantildea 2010 Prunier etal 2011) with the exception of Murcia Regionwhere it is common in temporary freshwater pondsused by cattle (Verdiell-Cubedo amp Boix 2014)Notostracans

L apus was first discovered by Grosso-Silva ampSoares-Vieira (2002) in Correlhatilde Ponte de LimaSeven years later we have found it in the verysame location (confirmed with Grosso-Silvapers com) in very turbid and low oxygenatedwater (L Cancela da Fonseca M Machado andM Cristo unpublished data) It is a Palearcticspecies (Alonso 1996 Rogers 2001) widelydistributed in Europe (Brtek amp Thieacutery 1995) InSpain it is also quite rare occurring only in thenorth (Alonso 1996)

T cancriformis has been recently discov-ered in rice fields of Vale do Sorraia Ribatejo(Machado et al 2016) This finding representsa new species for the Portuguese FLB faunaenlarging its geographical distribution to theSouthwest edge of the Iberian Peninsula and Eu-rope It is a Palearctic species (Alonso 1996)widely distributed in Europe (Brtek amp Thieacutery1995) and Asia including India and Japan (Kornet al 2006)

T baeticus is a species recently describedfollowing a classification based on genetic se-quences (Korn et al 2010) It is one of the newspecies assigned to the Triops mauritanicuslineage (Korn et al 2006) endemic to the Ibe-rian Peninsula In Portugal as well as in Spain(Korn et al 2010 this study) it is the species ofthe mauritanicus complex with the widest dis-tribution

T vicentinus also one of the recently de-scribed Iberian species of the mauritanicus com-plex is a Portuguese endemism Besides thescarcity of known sites it also has a very re-stricted distribution area in the west and centralAlgarve (Korn et al 2010 this study)

Spinicaudatans

C grubei is an endemism of the Iberian Penin-sula and Balearic Islands (Alonso 1996) In

relation to the previous published data (Cancelada Fonseca et al 2008) the number of knownsites increased from 22 to 34 Although it is oneof the most euryoecious Portuguese species itprefers relative deep ponds with long hydroperi-ods (Cancela da Fonseca et al 2008) In Spainit is also a relatively rare species (Alonso 1985Fahd et al 2009 Boix et al 2007 2010) withthe exception of Extremadura where Peacuterez-Boteet al (2006) found it in 36 of the ponds withFLB

M maroccana is known from just one moresite than those previously reported (Cancela daFonseca et al 2008 Gascoacuten et al 2012) Itoccurs in the northwest of Morocco (Van denBroeck et al 2015) from where the species wasdescribed (Thieacutery 1988) It is also present inSpain (Alonso 1996) where with the exceptionof the Dontildeana Natural Area (Boix et al 2007) ithas been rarely found (Alonso 1996 Peacuterez-Boteet al 2006 Prunier amp Saldantildea 2010 Prunier etal 2011 Gascoacuten et al 2012 Sahuquillo 2012)It is a sensitive species which is absent fromlarge FTLS and prefers ponds rich in aquaticvegetation with low turbidity waters and rela-tively low conductivity (Cancela da Fonseca etal 2008)

FLB faunal richness

Of the 13 FLB species known to occur in Por-tugal all but Lepidurus apus are present in thearea south of the Tagus River (Table 2 Fig 5)whereas north of the Tagus River only 4 specieswere recorded the anostracans Chirocephalusdiaphanus and Tanymastix stagnalis and the no-tostracans Lepidurus apus and Triops baeticus(Figs 2 4 and 5) the last one found very closeto the Tagus marginsThe apparent scarcity of the FLB fauna north

of the Tagus River both in species richness and inoccurrence may be due to the fact that this part ofthe country has been much less surveyed (fewersites visited at most on two occasions) than theareas south of the Tagus River (many more sitessampled some of them on a weekly basis) Hencean effort should be made to intensify surveys inthat part of the country

Limnetica 36 (2) 567-584 (2017)



In the Iberian context the Portuguese FLBfauna should be considered rich since Portugaloccupying just 153 of that region accountsfor 591 of the species known to occur in theIberian Peninsula (see Table 2)The number of species recorded in Portugal

is similar to or higher than reported for FranceItaly Belgium Denmark Tunisia and Iran butlower than for Spain the former Yugoslavia andAlgeria (Table 2) Nevertheless considering thespecies richness in relation to area that reportedfor Portugal is only lower than those of Belgiumand Denmark Comparing the number of FLBspecies recorded south of the Tagus River region(STRr) to those of the other regionsprovincesit is only lower in relation to the eastern half ofAustria and to North-Western Morocco (Table2) However a completely different picture is ap-parent when considering the species richness inrelation to the area with that of STRr being onlyhigher than that reported for Extremadura andMurcia (Spain)Maximum number of co-occurring species

and mean species number per site seem to bebetter indicators of species richness for compari-son purposes and have been used in other studies(Boix et al 2007 2010) The maximum numberof co-occurring species both in Portugal (Tables1 and 2) and in STRr (Table 2) is only lowerthan that reported for the Banat Province andalso for Western Morocco 3 decades ago (Thieacutery1991) Presently the value for North-WesternMorocco is the same as for Portugal (Van denBroeck et al 2015 table 1 Table 2) Regard-ing the mean species number per site Portugalpresents the highest value of all the countries un-der consideration and that found for STRr is onlylower than those relative to Banat Province andto North-Western Morocco being similar to thatinferred for Dontildeana Natural AreaDifferences were found in species coexis-

tence related to order whereas notostracans andspinicaudatans were mostly found with otherspecies anostracans were mainly found aloneThese differences should be related to the di-verse lifespans exhibited by the different groupsof FLB In general anostracan species have theshortest life cycle (Cancela da Fonseca et al

2008) and hence are able to survive in quiteephemeral temporary waters as well as in thosewith larger flooding period (Beladjal et al2003) This is the case of all Portuguese freshwa-ter anostracans (Cristo et al 2002 Cancela daFonseca et al 2008) with the exception of thelonely short-lived T lusitanica (Table 1 see alsoMachado amp Sala 2013) and of the long-lived Storvicornis (Beladjal et al 2003) S torvicor-nis together with C grubei are the species withthe longest lifespans recorded in Portugal (Can-cela da Fonseca et al 2008 M Machado perobs) with the latter typically lasting until thehabitat dries out (Cristo et al 2002) Specieswith extended life cycles are unable to survive inshort-lasting temporary waters needing deeperlonger-lasting ponds to complete their life cyclesand hence to persist in the biotope (Beladjalet al 2003) Many of the FLB with short lifes-pans can also be present in long-lived FTLS Tbaeticus T vicentinus and M maroccana haveintermediate lifespans (Cristo et al 2002 Can-cela da Fonseca et al 2008 present authorsper obs) The positive and strong association al-ready reported to exist between T stagnalis andB cortesi (Cancela da Fonseca 2008) explainswhy these short-lived species were usually foundwith one more species instead of being normallyaloneAnother remarkable fact with FLB in Por-

tugal is the total absence of co-occurrence ofcongeneric species which is not the case in othercountries like Spain (Miracle et al 2008) andMorocco (Peacuteregraves 1939 Thieacutery 1991)

FLB conservation status in Portugal

Portuguese FLB fauna has a high conservationvalue Almost half the FLB currently knownto occur in Portugal are endemic species 3 toPortugal -T lusitanica Tanymastix sp1 and Tvicentinus and another 3 to the Iberian Peninsulandash B cortesi T baeticus and C grubei Never-theless 7 species were only found in 15 FTLS orless 4 of them being known from less than 10sites (see results) Seven of the 13 species have arestricted (C grubei) or very restricted distribu-tion (the 3 Portuguese endemisms plus L apus

Limnetica 36 (2) 567-584 (2017)



T cancriformis andM maroccana) (Figs 1-6)Portuguese FLB species are all exclusive

inhabitants of temporary aquatic systems How-ever the ephemeral nature the small size andthe limited visibility of these systems lead to alack of recognition of their values and functionswhich results in them being readily destroyed ortransformed all around Europe (Gauthier et al2004) and other parts of the world before theopportunity to fully study their communities andfunctioning (eg Belk 1998)A large number of studies have called at-

tention to the increasing rate of disappearanceof FTLS in general and Mediterranean Tempo-rary Ponds in particular (eg Lumbreras et al2016 and references therein) and to the causesof their destruction (eg Machado et al 1999aPresado et al 2010 Boix et al 2012 and ref-erences therein Lumbreras et al 2016) In thesouthwest region of Portugal Ferreira amp Beja(2013) reported the disappearance of 56 of296 temporary ponds between 1991 and 2009due mainly to agricultural activities As statedin Belk (1998) ldquoThe overriding requirement forprotecting invertebrates is habitat conservationrdquoAs their survival is strictly dependent on tem-

porary pond preservation the conservation statusof the Portuguese FLB fauna is therefore of greatconcern because of the scarcity of these habitatsand the threats to which they are currently ex-posed (eg Ferreira amp Beja 2013) even in thecase of the most common species In Portugal asin many other countries none of the FLB speciesis red listed Hence it is urgent to intensify thewarnings to the appropriate national or interna-tional entities in order to ensure implementationof conservation laws concerning the PortugueseFLB fauna As stated by Miracle et al (2008)nowdays ldquolarge branchiopods are endangeredinvertebrates of greatest conservation needrdquo

ACKNOWLEDGEMENTS

We are very grateful to Ana Abreu Ana MariaVidal Antoacutenio Luiacutes Baacuterbara Monteiro CarlosNoivo Carlos Pacheco Cristina Cardoso Deacuteb-ora Nascimento Edgar Gomes Eliana Galioto

Fernando Pereira Horaacutecio Costa Inecircs CardosoJoatildeo Puga Joatildeo Reis Jordi Sala Jorge RoqueLuiacutes Palma Luis Sousa M Joseacute CaramujoManuel Joatildeo Pinto Maacuterio Reis Miguel PaisMirjam van de Vliet Nelson Fonseca NuacuteriaSalvador Patriacutecia Presado Pedro Beja PedroPortela Pedro Rocha Pedro Veiga Rita AlcazarSeacutergio Ribeiro Sofia Castel-Branco SusanaCoelho Tiago Marques Tiago Mourinho andUdo Schwarzner for providing information onsite location andor helping in the field workandor sending FLB samples for identificationThis study received national funds through

FCT - Foundation for Science and Technologyprojects UIDMulti043262013 UIDMAR042922013 and was also partially funded by LIFE+ Nature Project ldquoConservaccedilatildeo de charcos tem-poraacuterios na costa Sudoeste de Portugalrdquo Estudoda Ecologia das Lagoas Temporaacuterias do Su-doeste de Portugalrdquo-Protocolo de colaboraccedilatildeoCCMAR-PNSACV ICN funding ldquoIdentifi-caccedilatildeo dos Elementos de Conservaccedilatildeo (Faunade Crustaacuteceos Filoacutepodes e Anfiacutebios) nos Char-cos temporaacuterios do Parque Natural do Valedo Guadiana e Aacutereas limiacutetrofesrdquo-Protocolo decolaboraccedilatildeo CCMAR-ICN ICN funding ldquoEs-tudo dos Grandes Branquioacutepodes das LagoasTemporaacuterias na aacuterea de influecircncia dos ParquesNaturais do Tejo Internacional e da Serra deAires e Candeeirosrdquo-Collaboration with thePNTI and PNSAC chiefdoms Consoacutercio BIO3 Matos Fonseca amp Associados Estudos e Projec-tos Lda-ldquoPlano de Conservaccedilatildeo para os CharcosTemporaacuterios Mediterracircnicos na regiatildeo de imple-mentaccedilatildeo do EFMArdquo EDIA funding Additionalthanks are due to Karim Erzini who kindly re-viewed the English of the manuscript We alsothank two anonymous reviewers for their helpfulsuggestions that improved the manuscript

REFERENCES

ALONSOM 1985 A survey of the Spanish euphyl-lopoda Miscelacircnea Zooloacutegica 9 179ndash208

ALONSOM 1996 Crustacea Branchiopoda EnRamos MA (ed) Fauna Ibeacuterica 7 Museo Na-cional de Ciencias Naturales CSIC Madrid

Limnetica 36 (2) 567-584 (2017)



ALONSOM amp J GARCIacuteA-DE-LOMAS 2009Systematics and ecology of Linderiella baetica nsp (Crustacea Branchiopoda Anostraca Chiro-cephalidae) a new species from southern SpainZoosystema 31 807ndash827 DOIhttpdxdoiorg105252z2009n4a4

ALONSOM amp M VENTURA 2013 A new fai-ry shrimp Phallocryptus tserensodnomi (Branchio-poda Anostraca) from Mongolia Zootaxa 3670(3) 349ndash361 DOIhttpdxdoiorg1011646zootaxa367035

ATASHBAR B N AGH G Van STAPPEN amp LBELADJAL 2014 Diversity and distribution pat-terns of large branchiopods (Crustacea Bran-chiopoda) in temporary pools (Iran) Journal ofArid Environments 111 27ndash34 DOIhttpsdoiorg101016jjaridenv201407005

BALARAMANU amp CKG NAYAR 2004 A newspecies of the clam shrimp genus Lynceus (Bran-chiopoda Conchostraca Laevicaudata) from Ker-ala India Crustaceana 77(4) 407ndash416 DOI1011631568540041643373

BELADJAL L T NICO PEIREN TM VANDEK-ERCKHOVE amp J MERTENS 2003 Differentlife histories of the co-occurring fairy shrimpsBranchipus schefferi and Streptocephalus torvi-cornis (Anostraca) Journal of Crustacean Biol-ogy 23(2) 300ndash307 DOIhttpdxdoiorg1016510278-0372(2003)023[0300DLHOTC]20CO2

BELK D 1998 Global status and trends in ephemeralpool invertebrate conservation implications forCalifornia fairy shrimp in CW Witham ETBauder D Belk WR Ferren Jr and R Ornduff(Editors) Ecology Conservation and Manage-ment of Vernal Pool Ecosystems-Proceedings froma 1996 Conference California Native Plant So-ciety Sacramento CA 147ndash150

BELK D amp G PEREIRA 1982 Thamnocephalusvenezuelensis New Species (Anostraca Tham-nocephalidae) first report of Thamnocephalus inSouth America Journal of Crustacean Biology2(2) 223ndash226 DOI1011631937240X82X00356

BELK D amp DC ROGERS 2002 A confusing trioof Branchinecta (Crustacea Anostraca) from thewestern North America with a description of a newspecies Hydrobiologia 486 49ndash55 doi101023A1021374012622

BOIX D J SALA S GASCOacuteN A RUHIacute 2007Prospeccioacuten de branquioacutepodos (Crustacea) en laslagunas de la Reserva Bioloacutegica de Dontildeana Scien-tific report Estacioacuten Bioloacutegica de Dontildeana CSIC

BOIX D A RUHIacute J SALA S GASCOacuteN JCOMPTE amp XD QUINTANA 2010 Inverte-brats aquagravetics In Fraga P Estauacuten I amp CardonaE (Eds) Basses temporals mediterragravenies LIFEBASSES gestioacute i conservacioacute a Menorca 15 Con-sell Insular de Menorca Institut Menorquiacute drsquoEs-tudis Maoacute pp 249ndash283

BOIX D J BIGGS R CEacuteREacuteGHINO AP HULLT KALETTKA amp B OERTLI 2012 Pondresearch and management in Europe ldquoSmall isBeautifulrdquoHydrobiologia 689 1ndash9 DOI101007s10750-012-1015-2

BOIX D J SALA D ESCORIZA amp M ALONSO2016a Linderiella jebalae sp nov (CrustaceaBranchiopoda Anostraca) a new species fromthe Rif mountains (northern Morocco) Zootaxa4138(3) 491ndash512 DOI1011646zootaxa413834

BOIX D J KNEITEL BJ ROBSON C DUCHETL ZUacuteNtildeIGA J DAY S GASCOacuteN J SALA XDQUINTANA amp L BLAUSTEIN 2016b Inverte-brates of freshwater temporary ponds in Mediter-ranean climates In Invertebrates in Freshwa-ter Wetlands An International Perspective on theirEcology D Batzer amp D Boix (Eds) SpringerLondon Chapter 5 141ndash189 DOI101007978-3-319-24978-0_5

BOOTH B amp MITCHELL A 2001 Getting Startedwith ArcGISTM Environmental Systems ResearchInstitute Redlands California 260 pp

BRENDONCK L M HAMER amp A THIERY1992 Occurrence of Tetrahedral Eggs in the Strep-tocephalidae Daday (Branchiopoda Anostraca)with Descriptions of a New Subgenus Parastrep-tocephalus and a New Species Streptocephalus(Parastreptocephalus) zuluensis Brendonck andHamer Journal of Crustacean Biology 12(2)282ndash297 DOI1011631937240X92X00139

BRENDONCK L DC ROGERS J OLESEN SWEEKS amp WR HOEH 2008 Global diversityof large branchiopods (Crustacea Branchiopoda)in freshwater Hydrobiologia 595 167ndash176 DOI101007s10750-007-9119-9

BRTEK J 2002 Taxonomical survey of the Anos-traca Notostraca Cyclestherida Spinicaudata andLaevicaudata Acta Rerum Naturualium MuseiNationalis Slovaci Bratislava 48 49ndash60

BRTEK J amp A THIEacuteRY 1995 The geographicdistribution of the European Branchiopods (Anos-traca Notostraca Spinicaudata Laevicaudata)

Limnetica 36 (2) 567-584 (2017)



Hydrobiologia 298 263ndash280 DOI101007BF00033821

BRTEK J amp G MURA 2000 Revised key to fami-lies and genera of the Anostraca with notes on theirgeographical distribution Crustaceana 73 1037ndash1088 DOI101163156854000505083

CANCELA DA FONSECA L M CRISTO MMACHADO J SALA J REIS R ALCAZAR ampP BEJA 2008 Mediterranean Temporary Pondsin Southern Portugal key faunal groups as man-agement tools Pan-American Journal of AquaticSciences 3(3) 304ndash320

CARVALHO RN 1944 Cataacutelogo da colecccedilatildeo deinvertebrados de Portugal existentes no MuseuZooloacutegico da Universidade de Coimbra Crus-tacea Memoacuterias e Estudos Museu Zooloacutegico daUniversidade Coimbra 160 1ndash15

COHEN RG 2008 A new species of Branchinecta(Crustacea Branchiopoda Anostraca) from the Ar-gentinian Tierra del Fuego Zootaxa 1884 60ndash68

COTTARELLI V amp G MURA 1983 Anostraci No-tostraci Concostraci (Crustacea Anostraca No-tostraca Conchostraca) Guide per il riconosci-mento delle specie animali delle acque interne Ita-liane Consiglio Nazionale delle Ricerche AQ1194 18 1ndash73 figs 1ndash25

COTTARELLI V GMURAGampRS OumlZKUumlTUumlK2007 On two new Anostraca species (Bran-chiopoda) from Turkey Chirocephalus anatolicussp nov and Chirocephalus cupreus sp nov Crus-taceana 80(4) 455ndash474 DOI101163156854007780440902

CRISTO M M MACHADO amp J SALA GENO-HER 2002 Identificaccedilatildeo dos elementos de con-servaccedilatildeo (fauna de crustaacuteceos filoacutepodes e an-fiacutebios) nos charcos temporaacuterios do Parque Naturaldo Vale do Guadiana e aacutereas limiacutetrofes Relatoacute-rio final Protocolo de colaboraccedilatildeo ICN-CCMARUniversidade do Algarve

DAMGAARD J amp J OLESEN 1998 Distributionphenology and status for the larger Branchiopoda(Crustacea Anostraca Notostraca Spinicaudataand Laevicaudata) in Denmark Hydrobiologia377 9ndash13 DOI101023A1003256906580

DEFAYE D N RABET amp A THIEacuteRY 1998 At-las et bibliographie des Crustaceacutes Branchiopodes(Anostraca Notostraca Spinicaudata) de FranceMeacutetropolitaine Museacuteum National drsquoHistoire Na-turelle

EDER E W HOumlDL amp R GOTTWALD 1997 Dis-tribution and phenology of large branchiopods in

AustriaHydrobiologia 359 13ndash22 DOI101007978-94-017-3177-5_2

ESRI 2006 ArcGIS Rcopy 9 What is ArcGIS 92 Envi-ronmental Systems Research Institute RedlandsCalifornia

FAHD K M ARECHEDERRA M FLORENCIOD LEOacuteN amp L SERRANO 2009 Copepods andbranchiopods of temporary ponds in the DontildeanaNatural Area (SW Spain) a four-decade record(1964ndash2007) Hydrobiologia 634 219ndash230 DOI101007s10750-009-9889-3

FERREIRA M amp P BEJA 2013 Mediterranean am-phibians and the loss of temporary ponds are therealternative breeding habitats Biological Conser-vation 165 179ndash186 DOIhttpsdoiorg101016jbiocon201305029

FORNER i VALLS E amp E BREWSTER 2013First observation of Triops (Crustacea Branchio-poda Notostraca) in the Natural Park of the SerradrsquoIrta (Peniacutescola el Baix Maestrat) NEMUS 3101ndash110

FRANCO JA amp MLR AFONSO 1982 Distri-buiccedilatildeo de Pteridoacutefitos e Gimnospeacutermicas em Por-tugal Serviccedilo Nacional de Parques Reservas ePatrimoacutenio Paisagiacutestico Lisboa

GARCIacuteA-DE-LOMAS J amp CM GARCIacuteA 2008Observaciones de Branchiopoda en lagunas tem-porales de la provincia Revista de la sociedadGaditana de Historia Natural 5 145ndash151

GAacuteSCON S M MACHADO J SALA L CAN-CELA DA FONSECA M CRISTO amp D BOIX2012 Spatial characteristics and species nicheattributes modulate the response by aquatic pas-sive dispersers to habitat degradation Marine andFreshwater Research 63 232ndash245 doiorg101071MF11160

GAUTHIER P P GRILLAS amp M CHEYLAN2004 Menaces sur les mares temporaires meacutediter-raneacuteennes In Grillas P P Gauthier N Yavercov-ski amp C Perennou (Eds) Les mares temporairesmeacutediterraneacuteennes Volume 1-Enjeux de conserva-tion fonctionnement et gestion Station biologiquede la Tour du Valat Le Sambuc Arles 61ndash68

GROSSO-SILVA JM amp P SOARES-VIEIRA 2002Primeiro registo de Lepidurus apus (Linnaeus1758) para Portugal (Crustacea BranchiopodaNotostraca Triopidae) Boletiacuten de la Sociedad En-tomoloacutegica Aragonesa (SEA) 30 176

HILDREW AG 1985 A quantitative study of thelife history of a fairy shrimp (BranchiopodaAnostraca) in relation to the temporary nature of

Limnetica 36 (2) 567-584 (2017)



its habitat a Kenyan rainpool The Journal of Ani-mal Ecology 54 99ndash110 DOI1023074623

JEFFRIESMJ LB EPELE JM STUDINSKIamp CF VAD 2016 Invertebrates in temporarywetland ponds of the temperate biomes In Inver-tebrates in Freshwater Wetlands An InternationalPerspective on their Ecology D Batzer amp D Boix(Eds) Springer London Chapter 4 105ndash139 DOI101007978-3-319-24978-0_4

KORNM F MARRONE JL PEacuteREZ-BOTE MMACHADO M CRISTO L CANCELA DAFONSECA amp AK HUNDSDOERFER 2006Sister species within the Triops cancriformis lin-eage (Crustacea Notostraca) Zoological Scripta35(4) 301ndash322 doi101111j1463-6409200600230x

KORNM A J GREEN M MACHADO J GAR-CIacuteA-DE-LOMAS M CRISTO L CANCELADA FONSECA DA FRISCH JL PEacuteREZ-BOTE amp AK HUNDSDOERFER 2010 Phy-logeny molecular ecology and taxonomy of south-ern Iberian lineages of Triops mauritanicus (Crus-tacea Notostraca) Organisms Diversity and Evo-lution 10(5) 409ndash440 doi101007s13127-010-0026-y

KORNM amp AK HUNDSDOERFER 2016 Mole-cular phylogeny morphology and taxonomy ofMoroccan Triops granarius (Lucas 1864) (Crus-tacea Notostraca) with the description of twonew species Zootaxa 4178(3) 328ndash346 doi1011646zootaxa417832

LUMBRERAS A JT MARQUES AF BELO MCRISTO M FERNANDES D GALIOTO MMACHADO A MIRA P SAacute-SOUSA R SIL-VA LG SOUSA amp C PINTO-CRUZ 2016 As-sessing the conservation status of Mediterraneantemporary ponds using biodiversity a new tool forpractitionersHydrobiologia 782(1) 187ndash199 DOI101007s10750-016-2697-7

MACHADOM 2014 Amendment to the register ofthe specimen nAcirco 212 in Carvalho R N 1944Cataacutelogo da colecccedilatildeo de invertebrados de Portugalexistentes noMuseu Zooloacutegico da Universidade deCoimbra Crustacea Memoacuterias e Estudos MuseuZooloacutegico Universidade de Coimbra 160 1ndash15

MACHADOM M CRISTO amp L CANCELA DAFONSECA 1999a Non-Cladoceran BranchiopodCrustaceans from Southwest Portugal I Ocur-rence notes Crustaceana 72(6) 591ndash602 DOI101163156854099503636

MACHADOM M CRISTO J REIS amp L CAN-CELA DA FONSECA 1999b Biological data onTriops cancriformis mauritanicus (Ghigi 1921)and Cyzicus grubei (Simon 1886) ndash CrustaceaBranchiopoda ndash in SW Portugal temporary pondsLimnetica 16 1ndash7

MACHADOM M CRISTO amp L CANCELA DAFONSECA 2004 Known distribution of freshwa-ter large branchiopods in Portugal Reacutesumeacutes dela Confeacuterence Les mares temporaries mediter-raneacuteennes de la connaissance agrave la gestion et aacutela restauration (Rochebrune-sur-Argens Franccedile25ndash28 Maio 2004) p 49

MACHADOM amp J SALA 2013 Tanymastig-ites lusitanica sp nov (Crustacea BranchiopodaAnostraca) from Portugal first representative ofthe genus in Europe Zootaxa 3681(5) 501ndash523DOIhttpdxdoiorg1011646zootaxa368151

MACHADOM LG SOUSA L CANCELA DAFONSECA ED GALIOTO amp MJ CARA-MUJO 2016 First record of the tadpole shrimpTriops cancriformis (Lamarck 1801) (CrustaceaBranchiopoda Notostraca) in Portugal Book ofAbstracts XVIII AIL Congress Tolosa Cataluntildea4-8 July 2016 pp 102

MIRACLE MR M SAHUQUILLO amp E VI-CENTE 2008 Large branchiopods from freshwa-ter temporary ponds of Eastern Spain Verhandlun-gen Internationale Vereinigung fuumlr theoretischeund angewandte Limnologie 30 501ndash505

MURA G 1999 Current status of the Anostraca ofItaly Hydrobiologia 405 57ndash65 DOI101023A1003701004970

MURA G 2001UpdatingAnostraca (Crustacea Bran-chiopoda) distribution in Italy Journal of Limnol-ogy 60(1) 45ndash49 DOIhttpsdoiorg104081jlimnol200145

OLESEN J amp B TIMMS 2005 Caenestheriellamariae sp nov (Crustacea Branchiopoda Spini-caudata Cyzicidae) a new clam shrimp fromWest-ern Australia Zootaxa 824 1ndash8

PEacuteREgraveS J-M 1939 Contribution agrave la connaissancedes Notostraceacutes du Maroc Bulletin de la Socieacuteteacutedes Sciences Naturelles du Maroc 19 23ndash32

PEacuteREZ-BOTE JL A MUNtildeOZ JM GARCIacuteAEP RODRIacuteGUEZ AJ ROMERO P CORBA-CHO amp J FERNAacuteNDEZ 2006 Distribucioacutenestatus y conservacioacuten de los grandes branquioacutepo-dos (Crustacea Branchiopoda) en Extremadura(SO de la Peniacutensula Ibeacuterica) Boletiacuten AsociacioacutenEspantildeola de Entomologia 30(1-2) 41ndash57

Limnetica 36 (2) 567-584 (2017)



PEacuteREZ-BOTE JL A MUNtildeOZ JM GARCIA SRODRIGUEZ amp AJ ROMERO 2008 Distribu-cioacuten y abundancia de los huevos de resistencia deTriops cancriformis mauritanicus Ghigi 1921 yBranchipus cortesi Alonso amp Jaume 1991 (Crus-tacea Branchiopoda) en una laguna temporal delsuroeste de la peniacutensula Ibeacuterica Limnetica 27(1)57ndash64

PETROV B amp DM CVETKOVI 1997 Communitystructure of branchiopods (Anostraca Notostracaand Conchostraca) in Banat province in Yugos-laviaHydrobiologia 359 23ndash28 DOI101007978-94-017-3177-5_3

PETROV B amp PETROV I 1997 The status of Anos-traca Notostraca and Conchostraca (CrustaceaBranchiopoda) in Yugoslavia Hydrobiologia 35929ndash35 DOI101007978-94-017-3177-5_4

PETKOVSKI S 1995 On the presence of the genusTanymastix Simon 1886 (CrustaceaAnostraca) inMacedoniaHydrobiologia 298(1) 307ndash313 DOI101007BF00033825

PRESADO P MOURINHO T CANCELA DAFONSECA L amp MACHADO M 2010 Novosparadigmas do recreio e lazer e Conservaccedilatildeo daNatureza Um caso de estudo na Costa SudoestePortuguesa Actas do 6oCongresso a APDEA e IVCongresso de Gestatildeo e Conservaccedilatildeo da Natureza(Comunicaccedilatildeo ID73 editado em CD) 13p PontaDelgada APDEA-Universidade dos Accedilores SMiguel Accedilores

PRUNIER F amp S SALDANtildeA 2010 Grandes Bran-quioacutepodos (Crustacea Branchiopoda AnostracaSpinicaudata Notostraca) en la Provincia de Coacuter-doba (Espantildea) (Ano hidroloacutegico 20092010) Bole-tiacuten de la SociedadEntomoloacutegicaAragonesa (SEA)47 349ndash355

PRUNIER F R SOSA amp S SALDANtildeA 2011Grandes Branquioacutepodos (Crustacea BranchiopodaAnostraca Spinicaudata Notostraca) en la Provin-cia de Coacuterdoba (Espantildea) (Ano hidroloacutegico 20102011)Boletiacuten de la Sociedad Entomoloacutegica Arago-nesa (SEA) 49 223ndash226

REIS J M MACHADO M CRISTO amp L CAN-CELA DA FONSECA 1997 Ocorrecircncia de Tri-ops cancriformis mauritanicus Ghigi e de Cyzicusgrubei (Simon) ndashCrustacea Branchiopoda ndash emlagoas temporaacuterias do Sudoeste de Portugal Re-latoacuterio final Protocolo de colaboraccedilatildeo CCMAR-PNSACV

RIPPOL RODRIGUEZ J M DE LAS HERASCARMONA JM MORENO BENITEZ F PRU-

NIER amp F Soacutelano 2013 Grandes branquioacute-podos (Crustacea Branchiopoda Anostraca No-tostraca) en la provincia de Maacutelaga Espantildea (antildeohidroloacutegico 20122013) Arxius de MiscelmiddotlagraveniaZoologravegica 11 163ndash177

ROGERS DC 2001 Revision of the neartic Lepidu-rus (Notostraca) Journal of Crustacean Biology21(4) 991ndash1006 DOIhttpdxdoiorg1016510278-0372(2001)021[0991ROTNLN]20CO2

ROGERS DC W THAIMUANGPHOL N SAEN-GPHAN amp L SANOAMUANG 2013 Currentknowledge of the South East Asian large bran-chiopod Crustacea (Anostraca Notostraca Laevi-caudata Spinicaudata Cyclestherida) Journal ofLimnology 72 (s2) 69ndash80 DOIhttpsdoiorg104081jlimnol2013s2e5

RUEDA J JA AGUILAR-ALROLA amp F MEZ-QUITA I JUANES 2006 Contribucioacuten al conoci-miento de los crustaacuteceos (Arthropoda Crustacea)de las Malladas de la Devesa del Parque Natu-ral de la Albufera (Valencia) Boletin AsociacionEspantildeola de Entomologia 30(1-2) 9ndash29

SAHUQUILLOM 2012 Distribution and diver-sity of crustacean communities in Mediterraneanponds (Eastern Spain) relationship with climaticand limnological factors PhD Thesis Universityof Valencia Spain

SALA J F AMATDBOIXLCANCELADAFON-SECA M CRISTO M FLORENCIO J GAR-CIacuteA-DE-LOMAS S GASCOacuteN M MACHADOMR MIRACLE JL PEacuteREZ-BOTE J RUEDAA RUHIacute M SAHUQUILLO amp L SERRANO2010 Updating the distribution and conservationstatus of large branchiopods in the Iberian Penin-sula and Balearic Islands Programme and Abstractsof the 4th Conference of the European Pond Con-servation Network (EPCN) ldquoEyes of the land-scape-Value ponds in the 21st centuryrdquo Berlin(Erkner) Germany 1st-4th June 2010 p 115

SALA J M ALONSO F AMAT L CANCELADA FONSECA M CRISTO M FLOREN-CIO J GARCIacuteA DE LOMAS S GASCOacuteNM MACHADO MR MIRACLE A MIROacuteJL PEacuteREZ-BOTE JL PRETUS F PRUNIERJ RIPOLL J RUEDA M SAHUQUILLO LSERRANO M VENTURA D VERDIELL-CU-BEDO amp D BOIX 2016 Species richness andcomposition patterns of large branchiopods as-semblages in the Iberian Peninsula and BalearicIslands Book of Abstracts XVIII AIL CongressTolosa Cataluntildea 4-8 July 2016 pp 123

Limnetica 36 (2) 567-584 (2017)



SAMRAOUI B amp HJ DUMONT 2002 The largebranchiopods (Anostraca Notostraca and Spini-caudata) of Numidia (Algeria) Hydrobiologia486 119ndash123 DOI101023A1021338432186

SAMRAOUI B K CHAKRI amp F SAMRAOUI2006 Large branchiopods (Branchiopoda Anos-traca Notostraca and Spinicaudata) from the saltlakes of Algeria Journal of Limnology 65(2) 83ndash88 DOIhttpsdoiorg104081jlimnol200683

SORIA JM J RUEDA A ESCRIVAacute M SE-GURA E GARCIacuteA S ROMO F MESQUITAJOANES amp X ARMENGOL 2016 El resurgirde San Benito (Este Peninsular) tras la inundacioacutende 2015 Book of Abstracts XVIII AIL CongressTolosa Cataluntildea 4-8 July 2016 pp 127

SOUSA LG 2015 Conservaccedilatildeo de charcos tempo-raacuterios mediterracircnicos sob maneio agriacutecola umaavaliaccedilatildeo usando anfiacutebios MSc Thesis in BiologyConservation Universidade de Eacutevora

THIEacuteRY A 1986 Les Crustaceacutes BranchiopodesAnostraces Notostraces et Conchostraces du Ma-roc occidental I Inventaire et repartition Bulle-tin de la Socieacuteteacute drsquoHistoire naturelle de Toulouse122 145ndash155

THIEacuteRY A 1987 Les crustaces BranchiopodesAnostraca Notostraca amp Conchostraca des mi-lieux limniques temporaires (Dayas) au Maroc Ta-xonomie Biogeographic Ecologic Thegravese Docto-rat egraves SciencesUniversiteAix-Marseille III 405pp

THIEacuteRY A 1988 Maghrebestheria maroccana ngen n sp nouveau repreacutesentant des Leptestheri-idae au Maroc (Conchostraca) Crustaceana 5443ndash56 figs 1-9 DOI101163156854088X00627

THIEacuteRY A 1991 Multispecies coexistence of bran-chiopods (Anostraca Notostraca amp Spinicaudata)in temporary ponds of Chaouia plain (westernMorocco) sympatry or syntopy between usuallyallopatric species Hydrobiologia 212 117ndash136DOI101007978-94-011-3366-1_15

TIMMS BV 2013 A Revision of the Australianspecies of Lynceus Muumlller 1776 (Crustacea Bran-chiopoda Laevicaudata Lynceidae) Zootaxa 3702

(6) 501ndash533 DOIhttpdxdoiorg1011646zootaxa370261

TURKI S amp B TURKI 2010 Copepoda andBranchiopoda from Tunisian temporary watersInternational Journal of Biodiversity and Conser-vation 2(4) 86ndash97

VANDENBROECKM A WATERKEYN L RAI-ZI amp L BRENDONCK 2015 Distribution co-existence and decline of Moroccan large branchio-pods Journal of crustacean biology 35(3) 355-365 DOI1011631937240X-00002316

VANSCHOENWINKEL B L BRENDONCK TPINCEEL P DUPRIEZ amp A WATERKEYN2013 Rediscovery of Branchipus schaefferi (Bran-chiopoda Anostraca) in Belgium ndash notes on habi-tat requirements and conservation managementBelgian Journal of Zoology 143(1) 3ndash14

VERDIELL-CUBEDO D amp D BOIX 2014 Prime-ros datos sobre la distribucioacuten de grandes bran-quioacutepodos (Crustacea Branchiopoda) en la Re-gioacuten de Murcia (SE Espantildea) Anales de Biologiacutea36 65ndash69

VIANNA-FERNANDESAM 1951 Contribuiccedilotildeespara o estudo dos Filoacutepodes portugueses I Pri-meiras espeacutecies encontradas em Portugal Arqui-vos Museu Bocage 22 75ndash85

VIANNA-FERNANDESAM 1957 Sur la validiteacutedu nom Streptocephalus torvicornis (Waga) BairdRevista Portuguesa de Zoologia e Biologia Geral1 29ndash32

WEEKERS PHH G MURUGAN JR VANFLE-TEREN D BELK amp HJ DUMONT 2002 Phy-logenetic analysis of anostracans (BranchiopodaAnostraca) inferred from nuclear 18S ribosomalDNA (18S rDNA) sequencesMolecular Phyloge-netics and Evolution 25 535ndash544 DOIhttpsdoiorg101016S1055-7903(02)00289-0

ZIEROLD T B HANFLING amp A GOacuteMEZ 2007Recent evolution of alternative reproductive mo-des in the lsquoliving fossilrsquo Triops cancriformis BMCEvolutionary Biology 7 161 DOI1011861471-2148-7-161

Limnetica 36 (2) 567-584 (2017)


Con la colaboracioacuten de


foram atribuiacutedas quer ao endemismo portuguecircs T vicentinus quer a T baeticus a espeacutecie com distribuiccedilatildeo mais alargadana peniacutensula ibeacuterica ambas anteriormente incluiacutedas no complexo T mauritanicus Aleacutem destas uma nova espeacutecie Tany-mastigites lusitanica foi descoberta e descrita em Portugal Mais recentemente foi encontrada num campo de cultivo dearroz na regiatildeo central de Portugal uma metapopulaccedilatildeo de T cancriformis aparentemente composta unicamente por fecircmeasApresenta-se neste trabalho o estado actual do conhecimento sobre a distribuiccedilatildeo dos GBD de sistemas lecircnticos temporaacuteriosportugueses em mapas de quadriacutecula UTM (10 times 10 km) Foram prospectados um total de 505 siacutetios (charcos temporaacuteriosou conjuntos de charcos com localizaccedilatildeo proacutexima natildeo individualizaacutevel) 455 pelos autores Em 241 deles (477) foi obser-vada a presenccedila de pelo menos uma espeacutecie de GBD numa ocasiatildeo Dos 505 siacutetios apenas 87 estatildeo localizados a Norte dorio Tejo e nestes apenas em 17 (195) foi detectada a presenccedila de pelo menos 1 espeacutecie de GBD A sul do rio Tejo (Alentejoe Algarve) em 536 dos siacutetios foi encontrada pelo menos uma espeacutecie de GBD Esta lista inclui 13 espeacutecies (7 Anostraca2 Spinicaudata e 4 Notostraca) entre as quais 3 endemismos ibeacutericos e outros 3 endemismos portugueses

Palavras-chave Charcos temporaacuterios Anostraca Notostraca Spinicaudata espeacutecies endeacutemicas estatuto de conservaccedilatildeo

INTRODUCTION

Large branchiopods are a group of crustaceanspecies (Branchiopoda Anostraca NotostracaSpinicaudata Laevicaudata and Cyclestherida)that inhabit almost exclusively temporary aquaticsystems (Brendonck et al 2008) They play animportant ecological role in lentic systems asdetritivores herbivores carnivores andor scav-engers contributing for example to the control ofmosquitoes and to mitigating eutrophication byfeeding on organic detritus and decaying organicmatter They are also important in the trophicchain as a food source for insects amphibiansand birds They are considered a flagship groupfor freshwater temporary lentic systems (FTLS)and for temporary pond invertebrates (Bren-donck et al 2008) Until a few decades agostudies on these systems were scarce (Hildrew1985 and references therein Boix et al 2012Jeffries et al 2016) most probably due to thegenerally small size of FTLS their ephemeralnature and the relative unpredictability of theduration and timing of their flooding periodsIn Europe the considerable increase of scien-tific interest in these environments over the lasttwo decades (Boix et al 2016b and referencestherein) has been favored by the classificationof Mediterranean Temporary Ponds as a priorityfreshwater habitat under Directive 9243CEEAs a consequence of the increasing scientific in-

terest in temporary ponds namely in their func-tioning and biodiversity many studies on fresh-water large branchiopods (FLB) from severalareas in the world have been published (egBrendonck et al 2008 and references therein)and many new species have been recently discov-ered and described (eg Belk amp Pereira 1982Thieacutery 1988 Brendonck et al 1992 Rogers2001 Belk amp Rogers 2002 Balaraman amp Nayar2004 Olesen amp Timms 2005 Cottarelli et al2007 Cohen 2008 Alonso amp Garcia-de-Lomas2009 Alonso amp Ventura 2013 Machado amp Sala2013 Rogers et al 2013 Timms 2013 Boixet al 2016a Korn amp Hundsdoerfer 2016) InPortugal the first published scientific paper deal-ing with FLB inhabitants of temporary pondsand pools (Vianna-Fernandes 1951) reportedthe occurrence of 4 new species the anostra-cans Tanymastix lacunae Daday 1910 (= T stag-nalis (L 1758)) and Streptocephalus torvi-cornis (Waga 1842) the notostracan Triopscancriformis mauritanicus (Ghigi 1921) andthe spinicaudatan Caenestheriella grubei Daday1915 (= Cyzicus grubei (Simon 1886)) With theexception of one specimen of Apus cancriformisSchaumlffer 1756 (= Triops cancriformis (Lamarck1801)) registered in the catalogue of the inver-tebrate collection of the Zoological Museum ofCoimbra University (Carvalho 1944) no FLBspecies were known to occur in Portugal untilthe publication of this paper

Limnetica 36 (2) 567-584 (2017)









Total no

Sites of Total

with FLBn = 0 n = 1 n = 2 n = 3 n = 4 n = 5

C diaphanus 41 31 24 21 3 1 121 502

T stagnalis 18 43 9 13 2 1 86 357

B cortesi 12 38 9 11 3 0 73 303

T baeticus 8 22 16 16 3 1 66 274

C grubei 1 7 10 14 1 1 34 141

T vicentinus 1 7 11 4 0 0 23 95

Tanymastix sp 1 8 1 4 2 0 0 15 62

B schaefferi 7 2 1 2 0 1 13 54

S torvicornis 2 0 3 6 1 1 13 54

T lusitanica 8 1 0 0 0 0 9 37

M maroccana 0 3 1 1 1 0 6 25


L apus 0 1 0 0 0 0 1 04


No sites with FLB 107 78 30 22 3 1 241


Limnetica 36 (2) 567-584 (2017)





MATERIAL ANDMETHODS




Limnetica 36 (2) 567-584 (2017)







RESULTS





















Tanymastix sp1
















Limnetica 36 (2) 567-584 (2017)









Limnetica 36 (2) 567-584 (2017)








Limnetica 36 (2) 567-584 (2017)







DISCUSSION



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Anostracans








Limnetica 36 (2) 567-584 (2017)







SpeciesReference



















Limnetica 36 (2) 567-584 (2017)








Spinicaudatans




FLB faunal richness



Limnetica 36 (2) 567-584 (2017)











Limnetica 36 (2) 567-584 (2017)







ACKNOWLEDGEMENTS




REFERENCES



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Limnetica 36 (2) 567-584 (2017)










Total no

Sites of Total

with FLBn = 0 n = 1 n = 2 n = 3 n = 4 n = 5

C diaphanus 41 31 24 21 3 1 121 502

T stagnalis 18 43 9 13 2 1 86 357

B cortesi 12 38 9 11 3 0 73 303

T baeticus 8 22 16 16 3 1 66 274

C grubei 1 7 10 14 1 1 34 141

T vicentinus 1 7 11 4 0 0 23 95

Tanymastix sp 1 8 1 4 2 0 0 15 62

B schaefferi 7 2 1 2 0 1 13 54

S torvicornis 2 0 3 6 1 1 13 54

T lusitanica 8 1 0 0 0 0 9 37

M maroccana 0 3 1 1 1 0 6 25


L apus 0 1 0 0 0 0 1 04


No sites with FLB 107 78 30 22 3 1 241


Limnetica 36 (2) 567-584 (2017)





MATERIAL ANDMETHODS




Limnetica 36 (2) 567-584 (2017)







RESULTS





















Tanymastix sp1
















Limnetica 36 (2) 567-584 (2017)









Limnetica 36 (2) 567-584 (2017)








Limnetica 36 (2) 567-584 (2017)







DISCUSSION



Limnetica 36 (2) 567-584 (2017)





Anostracans








Limnetica 36 (2) 567-584 (2017)







SpeciesReference



















Limnetica 36 (2) 567-584 (2017)








Spinicaudatans




FLB faunal richness



Limnetica 36 (2) 567-584 (2017)











Limnetica 36 (2) 567-584 (2017)







ACKNOWLEDGEMENTS




REFERENCES



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Limnetica 36 (2) 567-584 (2017)






MATERIAL ANDMETHODS




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RESULTS





















Tanymastix sp1
















Limnetica 36 (2) 567-584 (2017)









Limnetica 36 (2) 567-584 (2017)








Limnetica 36 (2) 567-584 (2017)







DISCUSSION



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Anostracans








Limnetica 36 (2) 567-584 (2017)







SpeciesReference



















Limnetica 36 (2) 567-584 (2017)








Spinicaudatans




FLB faunal richness



Limnetica 36 (2) 567-584 (2017)











Limnetica 36 (2) 567-584 (2017)







ACKNOWLEDGEMENTS




REFERENCES



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Limnetica 36 (2) 567-584 (2017)



















Limnetica 36 (2) 567-584 (2017)





















Limnetica 36 (2) 567-584 (2017)








RESULTS





















Tanymastix sp1
















Limnetica 36 (2) 567-584 (2017)









Limnetica 36 (2) 567-584 (2017)








Limnetica 36 (2) 567-584 (2017)







DISCUSSION



Limnetica 36 (2) 567-584 (2017)





Anostracans








Limnetica 36 (2) 567-584 (2017)







SpeciesReference



















Limnetica 36 (2) 567-584 (2017)








Spinicaudatans




FLB faunal richness



Limnetica 36 (2) 567-584 (2017)











Limnetica 36 (2) 567-584 (2017)







ACKNOWLEDGEMENTS




REFERENCES



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Limnetica 36 (2) 567-584 (2017)










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Limnetica 36 (2) 567-584 (2017)







DISCUSSION



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Anostracans








Limnetica 36 (2) 567-584 (2017)







SpeciesReference



















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Spinicaudatans




FLB faunal richness



Limnetica 36 (2) 567-584 (2017)











Limnetica 36 (2) 567-584 (2017)







ACKNOWLEDGEMENTS




REFERENCES



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Limnetica 36 (2) 567-584 (2017)









Limnetica 36 (2) 567-584 (2017)







DISCUSSION



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Anostracans








Limnetica 36 (2) 567-584 (2017)







SpeciesReference



















Limnetica 36 (2) 567-584 (2017)








Spinicaudatans




FLB faunal richness



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Limnetica 36 (2) 567-584 (2017)







ACKNOWLEDGEMENTS




REFERENCES



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Limnetica 36 (2) 567-584 (2017)








DISCUSSION



Limnetica 36 (2) 567-584 (2017)





Anostracans








Limnetica 36 (2) 567-584 (2017)







SpeciesReference



















Limnetica 36 (2) 567-584 (2017)








Spinicaudatans




FLB faunal richness



Limnetica 36 (2) 567-584 (2017)











Limnetica 36 (2) 567-584 (2017)







ACKNOWLEDGEMENTS




REFERENCES



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Limnetica 36 (2) 567-584 (2017)






Anostracans








Limnetica 36 (2) 567-584 (2017)







SpeciesReference



















Limnetica 36 (2) 567-584 (2017)








Spinicaudatans




FLB faunal richness



Limnetica 36 (2) 567-584 (2017)











Limnetica 36 (2) 567-584 (2017)







ACKNOWLEDGEMENTS




REFERENCES



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Limnetica 36 (2) 567-584 (2017)








SpeciesReference



















Limnetica 36 (2) 567-584 (2017)








Spinicaudatans




FLB faunal richness



Limnetica 36 (2) 567-584 (2017)











Limnetica 36 (2) 567-584 (2017)







ACKNOWLEDGEMENTS




REFERENCES



Limnetica 36 (2) 567-584 (2017)





















Limnetica 36 (2) 567-584 (2017)

























Limnetica 36 (2) 567-584 (2017)





















Limnetica 36 (2) 567-584 (2017)



















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Limnetica 36 (2) 567-584 (2017)









Spinicaudatans




FLB faunal richness



Limnetica 36 (2) 567-584 (2017)











Limnetica 36 (2) 567-584 (2017)







ACKNOWLEDGEMENTS




REFERENCES



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ACKNOWLEDGEMENTS




REFERENCES



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ACKNOWLEDGEMENTS




REFERENCES



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Freshwater Large Branchiopods in Portugal · Dos 505 sítios, apenas 87 estão localizados a Norte...

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Transcript of Freshwater Large Branchiopods in Portugal · Dos 505 sítios, apenas 87 estão localizados a Norte...