NON-CLADOCERAN BRANCHIOPOD CRUSTACEANS FROM SOUTHWEST PORTUGAL. I. OCCURRENCE NOTES

MARGARIDA MACHADO'), MARGARIDA CRISTO') and LUIS CANCELA DA FONSECA?)

' ) Centro de CiEncias do Mar, Universidade do Algarve, UCTRA, Gambelas, P-8000 Faro, Portugal

') Instituto do Mar and Universidade do Algarve, UCTRA. Gambelas, P-8000 Faro, Portugal

ABSTRACT

The presence of non-cladoceran branchiopods was detected in only 16 out of 83 temporary fresh- water ponds of SW Portugal, which were recently sampled by the authors and other researchers. Two anostracans, Branchipus corresi and Chirocephalus diuphanus; two spinicaudatans. Cy:ic.u.\ yrubei and Maghrebestheriu mrrroccuna; and one notostracan, Triops cc~rzcr(fi,r~ni.s inauritunic~r~s. were identified. All taxa are new records locally, and all but C. grubei and 7: c. mauritatlicus are reported for the first time from Portugal.

La prtsence de Branchiopodes non Cladoceres a 6tC dCtectCe dans 16 seulement des 83 mares temporaires d'eau douce du SO du Portugal, qui ont ete recemment Cchantillonnees par les auteurs et d'autres chercheurs. Deux Anostraces, Branchipus corresi et Chirocephalus diuphtrnus, deux Spinicaudata, Cy,icu,s grubei et Mnyhrebestheria maroccana, et un NotostracC, Triops ccztrcriformi.s ~nnuritanicus, ont et6 identifie5. Tous les taxa sont nouveaux pour la faune locale et tous. sauf C. grubei et 7: c. mauritanicus. sont signalCs pour la premikre fois du Portugal.

INTRODUCTION

With the exception of the anostracan genus Artemia, which is characteristic of hypersaline permanent waters (Alonso. 1996), little is known of the Portuguese branchiopod fauna. Carvalho (1944) mentioned the presence of a specimen of Apus cuncriformis Schaffer, 1756 (= Triops cnncriformis (Lamarck, 180 1)) (No- tostraca), collected at Azambuja (fig. l), in the Portuguese invertebrate collection of the Zoological Museum of the University of Coimbra. In 1951, Vianna- Femandes reported the occurrence of 4 freshwater branchiopod taxa: Triops cancriformis mauritanicus (Ghigi, 1921) and the spinicaudatan Caenestheriellu

0 Koninklijke Brill NV. Leiden. 1999 Crustacrana 7 2 ( 6 )

592 M. MACHADO, M . CRISTO & L. C. DA FONSECA

grubei Daday, 19 15 (= Cyzicus grubei (Simon, 1886)) at Beja and Castro Verde, and the anostracans Streptocephulus torvicornis (Waga, 1842) and Taqmastix lacuncre Daday, 1910 (= Tanymasrix stagr~ulis (L., 1758)), respectively, at Cas- tro Verde and Torrgo (fig. l). The existence of Cyzicus tetracerus (Krynicki, 1830) in Portugal was noted in the updated geographic distribution published by Brtek & ThiCry in 1995. However, these authors did not mention the origin of the information and the authors of the present work could not find the source.

The small number of freshwater branchiopod (non-cladoceran) species found in Portugal so far, contrasts with that found in the remaining Iberian Peninsula. In fact, a total of 8 anostracan, 4 spinicaudatan, and 2 notostracan freshwater species are already known to exist in Spain (Alonso, 1996).

This almost complete lack of information on the Portuguese large branchiopod fauna could be due to the scanty attention paid so far by Portuguese researchers to temporary ponds, which are the characteristic habitat of non-cladoceran bran- chiopods. The ephemeral nature of these habitats by itself, and the relative unpredictability of the duration and timing of their flooding periods are certainly responsible for this lack of interest.

Directive 92/43/CEE, signed by Portugal, classifies the Temporary Medi- terranean Ponds as a priority habitat. This fact has greatly contributed to drawing the attention of researchers and institutions to these kinds of habitats and to the great need for acquiring knowledge with regard to their communities and major ecological processes, allowing the definition of correct management measures for their conservation.

As noted by Alcazar (1998), temporary ponds are one of the most striking elements of the landscape of the Portuguese SW coastal plateau. A major part of that plateau belongs to a nature park, the Parque Natural do Sudoeste Alentejano e Costa Vicentina (PNSACV) (fig. 1).

Within the framework of several studies (Alcazar, 1998; Reis et al., 1997; Beja. pers. comm.), 76 out of the 238 temporary ponds recorded in the PN- SACV and some adjacent areas (Alcazar, 1998; Reis et al., 1997) were sampled recently. Non-cladoceran branchiopod species or signs of their presence were detected in only 15 of the 76 ponds (table I, fig. l ).

The survey of large branchiopods in the SW part of Portugal was comple- mented by the inspection of 10 of a group of 12 temporary ponds marked in 1959 charts in a narrow north-south directed band, north-east of Odemira (fig. l ) . Three of the inspected ponds had been drained for agricultural purposes and non- cladoceran branchiopods have been found in only one of the seven remaining ones (Lagoa Pequena da Defeza; table I, fig. l).

PORTUGUESE BRANCHIOPODA

a: Azambuja

b: Torr.30

c: Beja

d : Castro Verde

Ponds with:

0 Branchipus cortesi

B. cortesi Maghrebestheria maroccana

A Triops cancriformis mauritanicus

A T. c. mauritanicus Chirocephalus diaphanus

El T. c. mauritanicus, C. diaphanus Cyzicus grubei

@ T c. mauritanicus M. maroccana

Fig. 1. Distribution of branchiopods (non-cladocerans) in Portugal. Approximate location of inhab- ited temporary ponds in SW Portugal (see table I) and approximate situation of other Portuguese locations mentioned in previous publications. Marked isolines show the distribution of medium

annual precipitation in mm. The PNSACV area is shaded.

Altogether, 5 different taxa were found. All but Cyzicus grubei and Triops cuncrijbrmis mauritanicus constitute new additions to the Portuguese fauna.

TABLE I Presence (*), number of individuals, or number of males (M) andor females (F) of the branchiopod taxa found in each pond and sampling occasion:

a, weekly sampling; b, fortnightly sampling. Pond code follows Alcazar (1998)

Pond Pond Sampling ObservedCollected taxa ObserverICollector Number Codername occasion

B. cortesi Alonso M. maroccana C. diaphanus C. grubei 7: c. mauritanicus & Jaume, 1991 ThiCry, 1988 Desrnarest, 1823 (Simon, 1886) (Ghigi, 1921) 5

R. Alcazar & P. Beja R. Alcazar & P. Beja R. Alcazar & P. Beja P. Beja & R. Alcazar P. Beja & R. Alcazar R. Alcazar & P. Beja

Present authors R. Alcazar

R. Alcazar & P. Beja R. Alcazar & P. Beja

l M ; 4 F Present authors 721 Present authors 73 Present authors

Present authors * Present authors * Present authors * Present authors * Present authors

Present authors

TABLE I (Continued)

Pond Pond Sampling Observed/Collected taxa ObserverICollector Number Codername occasion

B. cortesi Alonso M. maroccana C. diaphanus C. grubei 7: c. mnuritanicus & Jaume, 1991 ThiCry, 1988 Desmarest, 1823 (Simon, 1886) (Ghigi, 1921)

9 G12 13liv196 1 M ; l F Present authors 28/i/97 * * R. Alcazar & P. Beja

13livl98 11 M Present authors 10 G16 13/iv/96 exuvies Present authors 11 G20 13/iii/97 exuvies R. Alcazar & P. Beja

13/iii/98 exuvies Present authors 12 G35 13/iv/96 2 M ; 3 F Present authors

13livl98 1 M ; l F Present authors 13 G36 13/iv/98 exuvies Present authors 14 G37 3Oliii196 30 M; 9 F 54 M; 22 F Present authors

13/iv/96 48 M; 13 F 19 M; 23 F Present authors 1211-1 lliii197 a) 78 4058 1669 Present authors

l llxiil97 * * * Present authors

12lil98 * * * Present authors 12lii198 * * Present authors

15 L. Budens 13/iv/96 4 M ; 1 F Present authors 13livl98 1 M Present authors

16 L. P. Defeza 19/iii/98 3 M ; 5 F 3 M; 1 F Present authors

M. MACHADO. M. CRISTO & L. C. DA FONSECA

TAXA RECORDED

Order ANOSTRACA

Branchipus cortesi Alonso & Jaume, 199 1

Brunchipus cortu.ri Alonso & Jaume, 1991: 222

This species, first described less then a decade ago, is recognized so far as an Iberian endemic that was known until now from only four Spanish temporary ponds located in the Tagus, Guadiana, and Guadalquivir basins (Alonso & Jaume, 1991; Alonso, 1996). Now the occurrence of Branchipus cortesi in seven more ponds is reported, enlarging its geographical distribution to the SW coast of the Iberian Peninsula (fig. 1).

Environmental features of Portuguese ponds are similar to those in Spain (Alonso & Jaume, 1991 ). These are small ponds (200 to 5000 m2) with shallow water (6 1 m deep) of low to medium conductivity (90-320 PS), situated on sandy grounds (clayish in Spanish ponds) in low altitude areas (50-80 m above sea level) with an annual rainfall between 500 and 700 mm. Waters are slightly acid (pH 6-6.8) and show variable transparency (recorded Secchi disc depths of between 20 and 50 cm, approximately). Their bottoms are bare or, if more or less densely covered by aquatic vegetation, there are always bare zones. The length of the flooding period varies to a great extent from pond to pond (from 2.5-3 to 9-10 months), with about 6 months at Spanish locations.

Portuguese specimens closely fit the original description (Alonso & Jaume, 1991) as well as that given by Alonso (1996).

Chirocephalus diaphanus Desmarest, 1823

Chirocephalus diuphar~ris Desmarest, 1823: 4 16, in Alonso, 1996.

Chirocephalus diaphanus is a Paleartic species (Alonso, 1996), largely dis- tributed in Europe (Brtek & ThiCry, 1995), being the most common anostracan in several southern European countries, such as France (Nourisson & ThiCry, 1988), Italy (Cottarelli & Mura, 1983), and Spain (Alonso, 1985, 1996). Unex- pectedly, it had not yet been found in Portugal.

Chirocephalus diaphanus is eurytopic, living in ditches, temporary pools, and inundated fields (Brtek & ThiCry, 1995). It can be found in lowlands as well as in highlands to over 2000 m above sea level (Cottarelli & Mura. 1983; Alonso,

PORTUGUESE BRANCHIOPODA 597

1985, 1996; Nourisson & Thikry, 1988). It normally occurs in poorly miner- alized waters and seems to be indifferent to the degree of water turbidity and cover of aquatic vegetation (Alonso, 1985, 1996). In SW Portugal it was found in only three temporary ponds (fig. l , table I) of different dimensions (from 7300 to 70000 m2 approximately), depth (0.65-1.2 m) and persistence (3-9 months), which are located at about 60 to 140 m above sea level on sandy-clayish soils in areas with annual rainfall below 500 mm. During Chirocephalus diaphanus' life span, the water is characterized by a pH between 7.1 and 8.6 and usually low to medium conductivity values (210-460 PS). However, in the corresponding phase of the flooding period at pond G37, water conductivity is much higher (800-1400 pS). The amount of aquatic vegetation and water transparency are considerably different from pond to pond (water transparency in G3, G12, and G37: 15-30 cm, 10 cm, and 2.5-25 cm, respectively).

Specimens from these three ponds fit the descriptions by Cottarelli & Mura (1983), Nourisson & Thiiry (1988), and, particularly, that by Alonso (1996).

Order SPINICAUDATA

Cyzicus grubei (Simon, 1886)

Estheriu grubei Simon, 1886: 45 1

Cyzicus grubei is an endemic species of the Iberian Peninsula and Balearic Islands (Alonso, 1996). In the Balearics, it is known from only one temporary pond of Minorca Island, while on the Spanish mainland it has been found in the most arid zones of the Douro, Tagus, and Guadalquivir River basins (Alonso, 1996). As mentioned above, it has been recorded in the central part of Southern Portugal (Beja and Castro Verde; fig. 1). The present record extends the known distribution of this species to the very SW end of the Iberian Peninsula.

According to Alonso (1985, 1996), the characteristic habitats of Cyzicus grubei are shallow and low persistence temporary ponds, as well as pools with aquatic vegetation and highly turbid but poorly mineralized waters.

So far, it has been found in only two SW Portuguese temporary ponds which present some very different features in spite of being close to each other geo- graphically (fig. 1). Both are located on sandy-clayish soils 60 to 75 m above sea level, in areas with a mean annual rainfall of a little more than 400 mm. The pond G3 is approximately 7300 m2 in area and rather deep (maximum depth of approximately 1.2 m), with a flooding period of up to 9 months. The edges of this pond are covered by dense vegetation and it can be almost completely

598 M. MACHADO. M. CRISTO & L. C. DA FONSECA

covered by aquatic vegetation during the Ranunculus sp. blooming period, by late winter and early spring. Along the flooding cycle, water pH, conductivity, and transparency do show important fluctuations (pH 6.8-9.3; conductivity: 200- 3800 PS; transparency: 10-55 cm). Although larger (the water spreads to over 13 000 m2), G37 is shallower (0.7 m deep, at the most) and is a less persistent (3-4.5 months) pond. Aquatic vegetation is scarce. Throughout the flooding cycle its water is alkaline (pH 8.2-8.6), usually very turbid (transparency: 0.5- 25 cm) and with a relatively high conductivity (800-1700 pS). In both ponds, the Cyzicus grubei population is present throughout the flooding period.

Although SW Portuguese populations undoubtedly belong to Cyzicus grubei, some of their morphological features do deviate from the original (Simon, 1886) or Daday's (1914) descriptions, or even from that of Alonso (1996). The differ- ences detected are attributable to variability between populations, as well as to the developmental degree or age of individuals. These Portuguese populations will be morphologicaly characterized in a future paper.

Maghrebestheria maroccana Thiery, 1988

Maghrt~besrheria maroccanc~ Thiery, 1988: 44.

Maghrebestheria rnaroccana was discovered (as a new genus and new species of Leptestheriidae) just over a decade ago, in five temporary ponds situated in the northern half of the occidental fringe of Morocco (ThiCry, 1988).

The species, then erroneously identified as Eoleptestheria ticinensis (Balsamo- Crivelli. 1859) had earlier been found in a temporary pond in the North of Spain (Alonso, 1985). In 1988 it was detected in the Laguna de S. Lhzaro, a temporary pond of SW Spain, located approximately 40 km southwest of Sevilla (Alonso, 1996).

Since some doubts have arisen in relation to the identity of the leptestherid recorded in the ponds B38 and Lagoa Pequena da Defeza (table I), and since Alonso ( 1 996) based his description of Maghrebestheria maroccana on the popu- lation of Laguna de S. LBzaro, samples were taken from this last-mentioned pond in March 1998. M. mnroccana was found there, together with the anostracans Branchipus cortesi and Streptocephalus torvicornis, the spinicaudatan Cyzicus grubei, and the notostracan Triops cancriformis mauritanicus.

Comparison of Portuguese and Spanish specimens allowed confirmation of their identity. Some morphological aspects of the Portuguese specimens, how- ever, either do not fit the ThiCry (1988) and Alonso (1996) descriptions or they differ from the Spanish individuals observed. This is attributable to variability

PORTUGUESE BRANCHIOPODA 599

between populations and mostly to age differences. Morphological character- istics of the Portuguese populations of Maghrebestheria maroccana and their evolution as a function of age will be reported in a future paper.

The two Portuguese temporary ponds inhabited by Maghrebestheria maroc- cana (fig. l ) are small pools (100-2600 m2) with shallow water bodies ( G 0.5 m deep) of low conductivity (50-90 PS), located at low altitude (60-90 m above sea level) on sandy or sandy-clayish grounds in areas with annual rainfall between 500 and 700 mm. The bottom of pond B38 is covered largely by submerged vegetation and its acid water (pH 6.0) is quite transparent (transparency: 50 cm, approximately). The pond Lagoa Pequena da Defeza has no vegetation and its water is slightly alkaline (pH: 7.4) and very turbid (transparency < 10 cm). Their flooding periods last for 2.5-5 months and are considerably shorter than those of other known M. maroccana habitats (ThiCry, 1988; Alonso, 1996).

Order NOTOSTRACA

Triops cancriformis mauritanicus (Ghigi, 192 1 )

Thriops rnu~rritunicus Ghigi, 192 1 : 175, in Vianna-Fernandes, 195 1 .

Triops cancriformis (Lamarck, 1801) is a widespread species, occurring not only in the Paleartic region but also in India (Longhurst, 1955). However, its subspecies mauritanicus has a restricted geographical distribution, having been recorded so far only from NW Africa, the Balearic Islands (Minorca) and the southern half of the Iberian Peninsula (Longhurst, 1955; Alonso, 1985). The present paper reveals the presence of established populations of Triops cancri- formis mauritanicus in the extreme SW limit of the Iberian Peninsula (table I, fig. l), considerably enlarging its known range.

In SW Portugal, Triops cancriformis mauritanicz~s was recorded from tempo- rary ponds at about 60 to 140 m above sea level, situated mainly in areas with annual rainfall below 500 mm, and almost exclusively on sandy-clayish grounds. Their bottoms always present bare zones, even in the case of ponds which are rich in aquatic vegetation. These are ponds of very different sizes (maximum area usually between 100 and 28 000 m2), normally with clayish (transparency generally between 0.5 cm and 30 cm) and shallow (0.5 to 0.7 m deep) waters. In bigger (to over 72 000 m2) and deeper (> 1 m deep) ponds, this notostracan was frequently found only along the edges or in pools separated from the main body of the pond, where water depth and turbidity are similar to those in smaller ponds. Water pH and conductivity vary considerably from pond to pond (pH 6.4-8.8; conductivity: 50-1700 PS). In each one, these environmental parameters may

600 M. MACHADO. M. CRISTO 8~ L. C. DA FONSECA

also undergo important variations along the flooding phase corresponding to the life span of 7: c. mauritanicus (for instance. pH between 7.1 and 8.8 in pond G3, or conductivity between 800 and 1700 pS in pond G37). Pond flooding normally lasts 3 to 5 months but it can last 9-10 months in the larger andlor deeper ponds.

According to Alonso (1985), the characteristic habitat of this taxon involves low persistence, not very deep temporary ponds, rich in aquatic vegetation, and with poorly mineralized, very turbid waters. The environmental features of SW Portuguese ponds inhabited by this notostracan do not contradict Alonso's ob- servations except for the degree of water mineralization and aquatic vegetation cover. As a matter of fact, the most important 7: c. nlauritanicus population occurs in pond G37, which is very poor in aquatic vegetation and has water of relatively high conductivity.

Specimens from SW Portugal closely fit the taxonomic observations of Long- hurst (1955) and Alonso (1985); nevertheless they show some differences in rela- tion to Alonso's ( 1 996) description, which are attributable to variability between populations. Morphological characterization of SW Portuguese populations of Triops carzcrijbnrzis mauritunicus will be presented in a future publication.

COMMENTS

Through the present work the number of freshwater non-cladoceran branchio- pod species recorded in Portugal has increased from 4 to 7 (4 Anostraca, 2 Spini- caudata, and 1 Notostraca), adding also to the knowledge of the distribution of five of these in the Iberian Peninsula.

Although these branchiopod species were detected in less then 20% of the ponds prospected, the existence of a larger number of non-cladoceran branchio- pod species or, at least, a larger number of inhabited ponds is expected, since only about 30% of the ponds of SW Portugal were inspected. Furthermore, this kind of habitat is not limited to this part of the country. It is desirable, then, for purely scientific purposes. to complete the survey of Portuguese temporary ponds and their fauna, particularly non-cladoceran branchiopods.

Such research is very urgent as it is known that, nowadays, this kind of habitat is extremely vulnerable, being subject to innumerable threats. These in- clude draining, reclamation for agricultural or urban development, filling up with garbage or waste, pollution by fertilizers, pesticides or garbage, unsustainable agriculture or cattle raising, and even bad management decisions for conserva- tion purposes. The latter reflects the existing deficient knowledge of temporary pond ecology (Biggs et al., 1994; Collinson et al., 1995; Lanfranco, 1996). As Alonso (1985: 179) has pointed out, some non-cladoceran branchiopods "are

PORTUGUESE BRANCHIOPODA 60 1

in danger of extinction because of the rapid alteration and destruction of their habitats". This is especially the case of species with a restricted range.

Although a considerable area of SW Portugal is inside PNSACV, some of the above mentioned threats are present there, too. Branchipus cortesi ponds are in risk of being reclaimed for touristic construction purposes, as well as being cultivated or drained for cattle raising or agricultural uses (e.g., pond B38) or even for forestry (Pinto, 1997). Furthermore, they may be deepened for irriga- tion purposes, leading to the loss of their ecological features. Since 1996, three of the known ponds in this area have already been deepened (Alcazar, pers. comm.). As mentioned above, B38 and Lagoa Pequena da Defeza are the only Portuguese ponds where Maghrebestheria maroccaiza has been recorded. Lagoa Pequena da Defeza is risking to be drained for improving vehicle circulation on the dirt road which crosses it, or for planting Eucalyptus globulus La Billardikre, 1800 for the paper industry. The ponds of the extreme Southwest, inhabited by Chirocephalus diaphanus, Cyzicus grubei, andlor by Trioys cancrijbrmis mnuri- tanicus, are threatened by garbage or other waste deposition, artificial deepening for cattle raising or agricultural purposes, sand and gravel extraction, or even planting of Eucalyptus spp. (Pinto, 1997).

Thus, it is urgent to intensify studies on these kinds of ecosystems and their communities, allowing the definition and implementation of conservation mea- sures. Survival of the branchiopod species discussed in the present paper is entirely dependent on this, since they are exclusively inhabitants of temporary ponds.

ACKNOWLEDGEMENTS

We are very grateful to PNSACV for financial support and for providing infor- mation on environmental parameters of some ponds; to Prof. Pedro Beja and Dr. Rita Alcazar for the inestimable help in specimen collecting and for providing useful information related to some branchiopod habitats; to Dr. Luis Palma for helping to locate some ponds; to Dr. Joiio Reis and Dr. Ana Abreu for valuable assistance in field work; and to Prof. Pedro Beja, Prof. Karim Erzini. Dr. Joiio Reis, and Dr. Luis Palma for their critical reading of the manuscript, valuable suggestions, and corrections of the English.

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First received 9 June 1998. Final version accepted 1 September 1998.