Lagunas Mediterraneas Temporales2

Mediterranean Temporary Pools Volume 2 Species information sheets Grillas P., P. Gauthier, N. Yavercovski & C. Perennou

Production: Station biologique de la Tour du Valat Design: Tapages Publics Drawings: Antoine Catard, except pages 55, 83, 85, 88, 91 and 118 Translated from French by Janet Clayton and John Phillips Cover: photos Jean Roché

© 2004 Station biologique de la Tour du Valat Le Sambuc - 13200 Arles - France Readers are invited to reproduce texts and drawings featured in this publication provided credit is given to the texts and drawings authors and to the Station Biologique de la Tour du Valat. All photos rights reserved. No photographic part of this work may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopying except as may be expressly permitted in writing from the publisher. ISBN : 2-9103-6850-5

Mediterranean Temporary Pools volume 2 Species information sheets

Editors: Grillas P., P. Gauthier, N. Yavercovski & C. Perennou Associate editors: Jakob C., H. Michaud, G. Paradis & L. Rhazi

Editors, associate editors, authors and collaborators Antonetti P. (CNBMC), Calvière T. (TDV), Catard A (CEEP Var), Cheylan M. (EPHE), Delaugerre (CELRL Corse), Fuselier J. (ADENA), Garnéro S. (CEN-LR), Garraud L. (CBNA), Gauthier P. (TDV), Gendre T. (CEN-LR), Grillas P. (TDV), Hébrard J. P. (Université d’Aix-Marseille III-IMEP), Houssard C. (CEN-LR), Hugonnot V. (ad. perso. 1), Jakob C. (EPHE and TDV), Klewscewski M. (CEN-LR), Lombardini K. (EPHE), Michaud H. (CBNMP), Molina J. (CBNM-LR), Papazian M. (SOF), Paradis G. (Université de Corse, ad. perso. 2), Perennou C. (TDV) Pozzo di Borgo M. L. (OEC), Rhazi L. (université Hassan II), Rhazi M.(TDV and IMEP Université d’Aix-Marseille III), Rombaut D. (CEEP Var), Ruchon F. (AGRN-RH), Samraoui B. (Université d’Annaba), Souheil H. (AGRN-RH), Soulié-Märsche I. (Université Montpellier II, CNRS), Thiéry A. (Université de Provence, AixMarseille I), Titolet D. (Lycée international Georges Duby), Yavercovski N. (TDV). ADENA (Fuselier J.) Association de Défense de la Nature des Pays d’Agde Domaine du grand Clavelet, F-34300 Agde Tél.: +33 (0)4 67 01 60 23, fax: +33 (0)4 67 01 60 29 [email protected] AGRN.RH (Ruchon F., Souheil H.) Association de Gestion de la Réserve Naturelle de Roque-Haute, 1, rue de la Tour, F-34420 Portiragnes Tél/fax: +33 (0)4 67 90 81 16 [email protected] CBNA (Garraud L.) Conservatoire Botanique National Alpin de Gap-Charance Domaine de Charance, F-05000 Gap Tél: +33 (0)4 92 53 56 82, fax: +33 (0)4 92 51 94 58 [email protected]; [email protected] CBNMC (Antonetti P.) Conservatoire Botanique National du Massif central Le Bourg, F-43230 Chavaniac-Lafayette Tél: +33 (0)4 71 77 55 65, fax: +33 (0)4 71 77 55 74 [email protected] CBNMP (Michaud H, Molina J.) 1. siège Conservatoire botanique national de Porquerolles Castel Sainte Claire, F-83418 Hyères cedex Tél: +33 (0)4 94 12 82 30, fax: +33 (0)4 94 12 82 31 [email protected] 2. Unité de conservation et collections (Michaud H.) Le Hameau, F-83400 Ile de Porquerolles Tél.: +33 (0)4 94 12 30 32 , fax: +33 (0)4 94 12 30 30 Courriel: [email protected] 3. Antenne Languedoc-Roussillon (Molina J.) Institut de Botanique, Rue Auguste Broussonet, F-34090 Montpellier Tél.: +33 (0)4 99 23 22 11, fax: +33 (0)4 99 23 22 12 [email protected] CEEP: Conservatoire Études des Écosystèmes de Provence Alpes du Sud-Var 1, place de la Convention, F-83340 Le Luc Tél: +33 (0)4 94 50 38 39 / 06 16 97 82 03 2. CEEP Var ((Catard A., Rombaut D.) 1, Place de la Convention, F-83340 Le Luc Tél: +33 (0)4 94 50 38 39, fax: 04 94 73 36 86 [email protected] [email protected] CEN-LR (Gendre T., Garnéro S., Houssard C., Klewscewski M.) Conservatoire des Espaces Naturels du Languedoc-Roussillon 20 rue de la République, Espace République, F-34000 Montpellier Tél.: +33 (0)4 67 22 68 28, fax: +33 (0)4 67 22 68 27 [email protected] CELRL Corse (Delaugerre M.) Conservatoire des Espaces Littoraux et des Rivages Lacustres Délégation Corse 3, rue Luce de Casabianca, F-20200 Bastia Tél: +33 (0)4 95 32 38 14, fax: +33 (0)4 95 32 13 98 [email protected] [email protected] EPHE (Cheylan M., Jakob C., Lombardini K.) Ecole Pratique des Hautes Etudes Laboratoire de Biogéographie et Ecologie des vertébrés, Case 94, Université de Montpellier II, Place E. Bataillon, F-34095 Montpellier cedex 5 Tel: +33 (0)4 67 14 32 90, fax: +33 (0)4 67 63 33 27 [email protected] [email protected] [email protected]

Lycée international Georges Duby (Titolet D.) 200, rue Georges Duby F-13080 Luynes [email protected] OEC (Pozzo di Borgo M. L.) Office de l'Environnement de la Corse Avenue Jean Nicoli, F-20250 Corte Tél.: +33 (0)4 95 45 04 00, fax: +33 (0)4 95 45 04 01 SFO (Papazian M.) Société Française d’Odonatologie 7, rue Lamartine, F-78390 Bois d’Arcy Fax: +33 (0)1 34 60 68 63, site internet: www.libellules.org TDV (Calvière T., Gauthier P., Grillas P, Jakob C., Perennou C., Rhazi M., Yavercovski N.) Station Biologique de la Tour du Valat, Le Sambuc, F-13200 Arles Tél: +33 (0)4 90 97 20 13, fax: +33 (0)4 90 97 20 19 [email protected], site internet: www.tourdu valat.org Université d’Aix-Marseille III – IMEP1 (Hébrard J.P.) Institut Méditerranéen d'Ecologie et de Paléoécologie - CNRS UMR 6116 Université d'Aix-Marseille III, Faculté des Sciences et Techniques de SaintJérôme, Case 461, F-13397 Marseille cedex 20 Tél: +33 (0)4 91 28 85 35, fax: +33 (0)4 91 28 80 51 Université de Provence, Aix-Marseille I (Thiéry A) E.A. Biodiversité et environnement, Université de Provence, 3 place Victor Hugo, F-13331 Marseille cedex 3 Tél: +33 (0)4 91 10 64 25, fax: +33 (0)4 91 10 63 03 [email protected] CNRS, Université de Montpellier II (Soulié-Märsche I.) Laboratoire de Paléobotanique - UMR 5554 du CNRS, Université Montpellier II, C.P. 062, Place E. Bataillon, F-34095 Montpellier cedex 5 Tél: +33 (0)4 67 14 39 78, fax: +33 (0)4 67 14 30 31 [email protected] Université d’Annaba (Samraoui B.) Laboratoire de recherche des zones humides, Université d’Annaba, 4, rue Hassi-Beïda, Annaba, Algeria [email protected] Université Hassan II (Rhazi L.) Faculté des Sciences Aïn Chock, Laboratoire de Biologie et Physiologie Végétale, BP 5366, Maarif Casablanca, Morocco Tél.: (212) 037 86 33 10, fax: (212) 022 23 06 74 [email protected] ad. perso. 1 (Hugonnot V.) Le Bourg, F-43270 Varennes Saint Honorat Tél/Fax: +33 (0)4 71 00 23 07 [email protected] ad. perso. 2 (Paradis G.) 7, cours Général Leclerc, F-20000 Ajaccio Tél: +33 (0)4 95 50 11 65 [email protected] Aknowledgements The Station biologique de la Tour du Valat would like to warmly thank all the editors, authors and everyone who collaborated on this volume as well as Mohand Achérar (CEN-LR), Joël Bourideys (DIREN PACA), Christine Bousquet (AME), Jean Boutin (CEEP), Maddy Cancemy (OEC), Marie-Luce Castelli (OEC), Paul Chemin (DIREN-LR), Claire Chevin (MEDD), Béatrice Coisman (CEEP), Natacha Cotinaut (Mairie du Cannet-des-Maures), Geneviève Coutrot (TDV), Daniel Crépin (DIREN LR), Florence Daubigney (TDV), Marie-Antoinette Diaz (TDV), Christian Desplats (CELRL PACA), Arnaud Dorgère (TDV), Aude Doumenge (AGRN-RH), Emilien Duborper (TDV), Renaud Dupuy de la Grandrive (ADENA), Roger Estève (CELRL PACA), Sabine Fabre (CEN-LR), Mauricette Figarella (DIREN Corse), Guy-François Frisoni (Réserve Naturelle des Bouches de Bonifacio), Jean Giudicelli (Maison régionale de l’eau, Barjols), Denis Gynouvès (ONF Var), JeanClaude Heidet (CEEP), Bruno Julien (European Commission), Emilio Laguna (Generalitat de Valence, Espagne), Nicolas Leclainche (TDV), Olivier Limoges (Pôle relais Mares et Mouillères), Stéphanie Lieberherr (CEEP Var), Gilles Loliot (CELRLLR), Isabelle Lourenço de Faria (Commission Européenne), Margarida Machado (Université d’Algarve, Portugal), Marc Maury (Ecosphère), Leopoldo Medina, Olivier Nalbone (ARPE), Georges Olivari (Maison régionale de l’eau, Barjols), Eric Parent (Agence de l’Eau RMC), Jean-Claude Pic (TDV), Marc Pichaud (TDV), Marlène Savelli (OEC), Pierre Quertier (ONF Var), Bertrand Sajaloli (Pôles relais Mares et Mouillères), Nathalie Saur (Agence de l’Eau RMC), Alain Sandoz (TDV), Laurine Tan Ham (TDV), Florence Verdier (CELRL LR) and Myriam Virevaire (CNBMP) for their contribution to the LIFE “Temporary Pools” project.

Summary Introductory notice . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

5

Macrocrustaceans. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

83

Imnadia yeyetta Hertzog, 1935. . . . . . . . . . . . . . . . . . . .

83

Plant species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

7

Linderiella massaliensis Thiéry & Champeau, 1988 . . . .

85

Apium crassipes (Koch ex Reichenb.) Reichenb. fil. . . . .

7

Tanymastix stagnalis (Linnaeus, 1758) . . . . . . . . . . . . . .

88

Artemisia molinieri Quézel, Barbero & Loisel. . . . . . . . .

9

Triops cancriformis (Bosc, 1801) (1)

Crypsis schoenoides (L.) Lam . . . . . . . . . . . . . . . . . . . . . .

12

Damasonium polyspermum Coss. . . . . . . . . . . . . . . . . . .

15

Elatine brochonii Clavaud . . . . . . . . . . . . . . . . . . . . . . . .

18

& Lepidurus apus (Linné, 1758) (2). . . . . . . . . . . . . . . . .

91

Odonata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

94

Eryngium pusillum L. . . . . . . . . . . . . . . . . . . . . . . . . . . . .

21

Illecebrum verticillatum L. . . . . . . . . . . . . . . . . . . . . . . .

23

Ischnura pumilio (Charpentier, 1825) (1)

Isoetes duriei Bory (1) & Isoetes histrix Bory (2) . . . . . .

26

& Ischnura genei (Rambur, 1842) (2) . . . . . . . . . . . . . .

Isoetes setacea Lam. (1) & Isoetes velata A. Braun (2) .

30

Lestes barbarus (Fabricius, 1798) (1)

Littorella uniflora (L.) Ascherson . . . . . . . . . . . . . . . . . .

34

& Lestes virens (Charpentier, 1825) (2) . . . . . . . . . . . . .

96

Sympetrum fonscolombii (Sélys, 1840) (1)

Lythrum borysthenicum (Schrank) Litv. (1) & Lythrum tribracteatum Salzm ex Sprengel (2). . . . . .

94

37

& Sympetrum meridionale (Sélys, 1841) (2) . . . . . . . . . . . .

99

Lythrum thymifolium L. (1) & Lythrum thesioides M. Bieb. (2) . . . . . . . . . . . . . . . . . .

41

Marsilea strigosa Willd. . . . . . . . . . . . . . . . . . . . . . . . . .

45

Amphibians . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102

Mentha cervina L. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

49

Bufo calamita, Laurenti 1768 . . . . . . . . . . . . . . . . . . . . . 102

Myriophyllum alterniflorum DC. . . . . . . . . . . . . . . . . . . .

52

Discoglossus sardus Tschudi, 1837 . . . . . . . . . . . . . . . . . 105

Nitella opaca (Bruzelius) Agardh . . . . . . . . . . . . . . . . . .

55

Pelobates cultripes (Cuvier, 1829). . . . . . . . . . . . . . . . . . 108 Triturus cristatus, (Laurenti, 1768) . . . . . . . . . . . . . . . . . 111

Ophioglossum azoricum C. Presl (1) & Ophioglossum lusitanicum L. (2) . . . . . . . . . . . . . . . . .

58

Pilularia minuta Durieu ex A. Braun . . . . . . . . . . . . . . . .

62

Triturus marmoratus, Latreille 1800 . . . . . . . . . . . . . . . . 114

Ranunculus lateriflorus DC. (1) & Ranunculus nodiflorus L (2) . . . . . . . . . . . . . . . . . . . .

65

Ranunculus revelieri Boreau . . . . . . . . . . . . . . . . . . . . . .

69

Genus Riccia L. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

72

Riella helicophylla (Bory & Mont.) Mont.. . . . . . . . . . . .

76

Teucrium aristatum Perez Lara . . . . . . . . . . . . . . . . . . . .

80

Glossary. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116 Macrocrustacean inventory form . . . . . . . . . . . . . . . . . . . . . . 120 Bibliography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121

5

Introductory notice This second volume of “Mediterranean Temporary Pools” presents, in the form of information sheets, a summary of the current knowledge on a selection of the remarkable species of Mediterranean temporary pools (29 plant species, 16 animal species). This knowledge is either bibliographical in origin or was acquired during the LIFE-Nature project “Conservation of Mediterranean Temporary Pools No. 99/72049”, presented on page one of volume 1.

The species presented have been selected according to a number of criteria: • Their dependence on temporary aquatic habitats; • Their rarity (this criterion is, however, very variable from country to country); • The Mediterranean nature of their distribution; • Their presence at one or more of the seven sites of the LIFE project; • Their biological characteristics or their ecological requirements, illustrating various adaptive strategies. Plant species Amphibious vascular plants, flagship species of temporary pools, are the best represented with 26 species, i.e. almost a quarter of the 107 rare plants of the oligotrophic* temporary pools censused in 18 countries of the Mediterranean Basin (see volume 1, table 2). A discreet and often poorly known group, the liverworts are described on two information sheets. One is devoted to a very rare species of temporary pools (Riella helicophylla), the other to a genus particularly well adapted to temporary wetland areas, the genus Riccia. Finally, the choice of a charophyte* (algae) dependent on temporary Mediterranean pools, Nitella opaca, illustrates a poorly known but important group in temporary wetland areas. Macrocrustaceans Of the fifty or so species of the Mediterranean Basin, five have been selected from three different Orders. Two species are very rare in France (Linderiella massaliensis is endemic, Imnadia yeyetta is present on two sites). Water requirements differ among the species chosen (ephemeral habitats for Tanymastix stagnalis, semi-permanent habitats for Lepidurus apus and Triops cancriformis). Twelve Mediterranean countries support one or more of these species. Odonata In the Odonata, the 25 species with a Mediterranean distribution have life-history characteristics which are adapted to temporary flooding and can vary according to the latitude. The six species chosen illustrate restricted (Ischnura genei, endemic to the islands of the western Mediterranean) to wide (Sympetrum fonscolombii) Mediterranean distribution as well as several strategies of adaptation to summer drought, based on the mechanism and dates of egg laying, the existence of diapauses (egg or larva), the periods of emergence of the adults, the number of generations per year, wether or not migratory, the greater or lesser capacity to colonise new pools, etc.

6

Amphibians Among the sixty or so species found in the Mediterranean Basin, five which are adapted to temporary pools have been chosen in two different groups (anurans and urodeles), illustrating the various environmental requirements and biological characteristics: duration of larval cycle, size of clutches, etc.).

Information sheet headings Latin name: current nomenclature Classification: except otherwise stated, the class then the family

of the species (current nomenclature) are mentioned (in small upper-case letters) immediately under the title of each information sheet. Main synonyms: synonyms used in “classic” guides to flora and fauna. Common names: when they exist, the common French, Spanish, Portuguese, Italian, English, Moroccan and Algerian names are given. The names used at a national scale are given, with the exception of Morocco, where certain local names of oral tradition are known265, though it is not possible to give any precise region where these are used. For European countries, the French Internet site www.tela-botanica.org was consulted. In addition, the French names of the plant species were researched in the major flora of Bonnier42 and in the Livre Rouge de la Flore Menacée de France (French Red Data Book of threatened flora)274, the Spanish and Portuguese names in Flora Iberica64, the Italian names in the Italian Red Data book of threatened flora86 and in Flora d’Italia292. Subspecies: the subspecies currently recognised are mentioned. Description/identification criteria: a variety of bibliographic sources have been used, as well as the personal observations of the authors. Similar species: the species with which some confusion is possible are mentioned, as well as their distinctive characteristics.

Distribution - Ecology Distribution/range

The distribution of the species in France, then in other Mediterranean countries is given whenever possible, in particular in the countries of the western Mediterranean (Spain, Italy, Maghreb). Habitat

This paragraph contains, for France, and if possible, for other Mediterranean countries: • A general description of the stations; • The mention of the habitats of “European Community Interest” or priority habitats for the species, included in annexe I of the Habitats Directive12, 119, accompanied by their Natura 2000* code; The mention of French habitats defined according to the CORINE Biotopes39 typology; • For the flora: the list of the phytosociological communities* characterised by the species or to which the species belongs, and where possible the main companion species. Ecology

This paragraph provides the main biological characteristics and environmental requirements useful for the conservation management of the species.

Introductory notice Conservation - Management

Biological characteristics

In France, and if possible in other Mediterranean countries, information relating to the biology and life-history traits characteristics of the species is given. • For the flora: the biological type, methods of sexual reproduction (aerial, aquatic), description of seeds, and the biological cycle (germination, growth, flowering, fruiting, dispersal). • For the fauna: feeding, the site and the mechanisms of egg depositing, the biological cycle, (reproduction, egg laying, development), longevity, distinctive behaviour, interannual and spatial variations in the phenology.

Assessment of populations

In France, and in other Mediterranean countries, the number of sites is indicated (with, for the fauna, an estimation of the population numbers, if possible. Conservation status

The status of the populations (stable, expanding or regressing) is provided, as well as the main sites of recent appearance of the species or of disappearance. Legal status of the species/level of protection

On each information sheet, a figure at the end of this paragraph illustrates the annual phenological cycle of a population and an average hydrological state of the pool.

Example: the phenological cycle of Marsilea strigosa France Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Dec.

Emergence of fronds and vegetative growth ? Formation of sporocarps Withering of fronds

generally flooded phase (or of saturation of the soil for certain species) flooded phase some years dry phase This schematic illustration cannot take into account all interannual variability in the Mediterranean climate (temperature, precipitation) or individual variability in the response of organisms, but it can provide the manager with an average state. Some authors express the variability of the response of a population to variations in the hydrological regime by dots extending the solid line of the different phases of a cycle. Environmental conditions In France, and if possible in other Mediterranean countries, the following are given (within the limits of the information available): • For the flora: the hydrological conditions required (depth of water, duration of flooding and drying out, germination of seeds, chemical characteristics, etc.); the nature of the substrate (acidic, calcareous, granulometry, etc.); sensitivity to interspecific competition (in relation to light in particular); the impact of perturbations such as cutting, grazing, trampling by livestock, etc. • For the fauna: the breeding requirements (duration and date of flooding, depth, type of pool, etc.); the territories, the wintering or summering habitat and the necessity for complementary habitats (terrestrial phase of amphibians, for example); the dispersal distance of the adults and the young after the aquatic phase; environmental perturbations and their impact (rain, drought, soil humidity, etc.); interspecific relations (competition, predation, etc.); the main factors in natural mortality, etc.

A table provides for each species: • Inclusion in the IUCN Red Lists176, 399, www.redlist.org. • Inclusion in the annexes of the Bern Convention87. • Inclusion in the annexes of the “Habitats” Directive119. • The status of protection at national or regional level in France and in other Mediterranean countries: - The protection status at national level for the flora was examined for Italy86, 292, Spain (Internet site: www.mma.es/conserv_nat/ acciones/esp_amenazadas/html/catalogo/flora.htm), France, Algeria (decree no. 93-117 of 23/11/1993 fixing the list of protected non-cultivated plant species). - The protection status at national level for the fauna was examined for France, Spain (Internet site: www.mma.es/conserv_nat/ acciones/esp_amenazadas/html), and Portugal (amphibians59). - The protection status at regional level for the flora and fauna was examined for France, and in some regions of Italy for the fauna (Sardinia and Tuscany). • Inclusion in the Red Data Books and Lists in France and in other Mediterranean countries: - For vascular flora, the national Red Data Books and Lists of France274, Italy86, Spain6, Morocco133, Greece290, and Malta344 were consulted as well as the regional Red Data Book for the Balearics331. For the bryophytes, the Red Data Book of the bryophytes of Europe128, and the Red Data Book for France, which is currently being produced, served as references. - For the fauna, the National Red Data Book242 was consulted for France, as well as the Red Data Lists for Spain (amphibians293), Portugal (amphibians59) and Italy (vertebrates14). Conservation problems/threats

The following factors are examined: • Anthropogenic factors: destruction, infilling, drainage, pollution, etc. • Natural factors: dynamics of the vegetation, etc. • Risks relating to populations: risks of extinction at a site. Management and conservation measures

Current measures The main protected sites where the species are found are listed (Natural Parks, Nature Reserves, Natura 2000 sites, etc.) as well as the management measures in place. Recommendations These concern the management measures to put in place (hydrology, grazing, scrub clearing), modifications in the protection status of the species, protection of the habitat, etc.

7

Mediterranean temporary pools Bibliography

Journal officiel: Journal Officiel de la République Française (offi-

The authors of the references are listed in alphabetical order at the end of each information sheet, with a numbered cross-reference to the complete reference in the general bibliography at the end of the volume.

s.l.: sensu lato (in the broad sense) subsp.: subspecies sp.: Indeterminate species (from the Latin species). For example,

cial bulletin for the public, giving details of laws).

Isoetes sp. signifies an indeterminate species of the genus Isoetes spp.: several species of the genus. For example, Isoetes spp. sig-

Other information Acronyms and abbreviations used in the information sheets CBNBP: Conservatoire Botanique National du Bassin Parisien. CEEP: Conservatoire Etude des Ecosystèmes de Provence - Alpes

nifies “all or some species of the genus Isoetes”. em.: amended by (abbreviation of the Latin emendavit, from the Latin verb emendare: to modify, amend, improve). Used in phytosociology, for example: “the alliance Isoetion Br.-Bl. 1931 of the order Isoetalia Br.-Bl. 1931 em. Rivas-Goday, 1970”. µS.cm-1: micro-siemens per cm (unit of measurement for conductivity of water)

du Sud. CEN-LR: Conservatoire des Espaces Naturels du LanguedocRoussillon. MEDD: Ministère de l’Ecologie et du Développement Durable (France). ONF: Office National des Forêts. PACA (region): Région Provence-Alpes-Côte-d’Azur (France). IUCN: International Union for the Conservation of Nature

8

Glossary

The technical and scientific terms not found in a current dictionary (such as the French dictionary Le Petit Robert), are defined in the Glossary. They are marked in the text with an asterisk* the first time they are encountered in each information sheet.

Plant species

Apium crassipes

(Koch ex Reichenb.) Reichenb. fil.

ANGIOSPERMS APIACEAE Main synonyms

Helosciadum crassipes Koch ex Reichenb. A. inundatum (L.) Reichenb. fil.var. crassipes Paol. A. inundatum (L.) Reichenb. fil subsp. crassipes Landi Sium crassipes (Reichenb.) Sprengel French name: Ache Italian name: Sedano di Sardegna Subspecies

None Description/identification criteria50, 292

• Herbaceous plant, hydrophytic* or amphibious, either hemicryptophytic* or annual. • Stem hollow and grooved, at first briefly prostrate and radicant, then upright, erect or floating, capable of growing up to 40 cm long in water. • Leaves alternate, of two types: basal leaves submerged, twoto four- pinnatisect* and completely divided into long, slender strips; emergent leaves simply pinnatisect with toothed segments. • Inflorescence: umbel composed of three to five partial umbels, no bracts, three to six bracteoles. • Flower with calyx invisible, corolla with five small white petals (0.5 mm) inferior ovary with two loculi, style slightly longer than the stylopodium* (apex of the ovary). • Fruit dry, consisting of two monospermic carpels (mericarps), indehiscent, 1.25 to 1.5 mm long, at the tips of short pedicels which are very swollen below. Similar species

Apium inundatum is distinguished by its slightly larger size and bigger fruits, and especially by the non-swollen fruiting pedicels. The two species only coexist in Sicily (no risk of confusion in Corsica or Sardinia).

1 cm

Apium crassipes

Habitat 1. “Mediterranean temporary pool” habitat General description

In Corsica and Sardinia, Apium crassipes is an important constituent in terms of the biomass* of the flooded phase of many temporary pools. In southern Corsica, A. crassipes, together with various floating species (Ranunculus peltatus, R. ophioglossifolius, Illecebrum verticillatum, Baldellia ranunculoides), is abundant in the temporary pools of Padulellu, Tre Padule de Suartone and Padulu219, 221, 284. In Tunisia, A. crassipes is associated with Isoetes velata, Myosotis sicula, Illecebrum verticillatum, Eryngium barrelieri (E. pusillum), Ranunculus ophioglossifolius298. Habitats Directive

“Mediterranean Temporary Pools” (code 3170). CORINE Biotopes

Distribution - Ecology Distribution/range

Apium crassipes is considered to be a Tyrrheno-Sicilo-Numidian endemic species50. France

The species only occurs in Corsica. Other Mediterranean countries

Occurs in Italy (Sardinia, Pontins marshes and Sicily292) and in North Africa (Tunisia and eastern Algeria). Casper & Krausch63 include the Nile Delta in the range.

22.34 Mediterraneo-Atlantic amphibious communities (Isoetalia). 22.341 Short Mediterranean amphibious swards (Isoetion). 22.3412 Aquatic Mediterranean Quillwort swards: communities containing Isoetes velata in water bodies of variable depth. Phytosociology*

In its aquatic form, Apium crassipes belongs to formations of the submerged phase of the Isoetion, within communities of Isoetes velata. Some authors include the hydrophyte* formations in which A. crassipes is found within the communities of the Potamogetonetea pectinati class284. 2. Other habitats General description

Apium crassipes is also found in ditches and in wet meadows. In these biotopes, which dry out rapidly at the end of winter,

9

Mediterranean temporary pools

A. crassipes grows only to a small size and does not produce slender strap-like leaves.

Conservation - Management Assessment of populations

Phytosociology*

In Algeria, around the Mafragh, Tonga and Oubeira lakes, Géhu et al.156, for seasonally flooded meadows resulting from aggradation, created the association with Paspalum distichum and Apium crassipes which they named Helosciadi crassipes-Paspaletum distichi and which they included in the class Agrostietea stoloniferae, the order Plantaginetalia majoris and the alliance Paspalo-Agrostion. Ecology Biological characteristics

Life form Hydrophyte*, either hemicryptophytic* (perennial) or therophytic* (annual). Reproduction Aerial.

Adaptive strategy (sensu Grime163) Stress-tolerant (S).

Corsica April

May

June

July

Aug.

Sept.

Oct.

Nov.

Dec.
Germination




The populations fluctuate from one year to another, but appear to be stable in the medium term at the Padulellu, Tre Padule and Padulu pools. On the other hand, as a result of the cessation of cattle grazing several decades ago, the Arasu, Mura dell’Unda and Muratellu pools now have small populations. The population of the Chevanu temporary pool, which is very bare and where the substrate is very compacted, is heading for extinction. Legal status of the species/level of protection -

Natural factors

Phenological cycle March

In Corsica

Conservation problems/threats

Description of achenes Length: c. 1.25 to 1.5 mm.

Feb.

Conservation status

IUCN 1997 Bern Convention Habitats Directive National and regional protection National Red Data Books and Red Lists

Flowering In Corsica, from mid-March to late April.

Jan.

In Corsica, probably over 20.

Germination

In Corsica, the likely cessation of extensive cattle grazing in the near future threatens to cause a severe fall in numbers at almost all sites, as a result of the spread of tall herbaceous plants (Schoenus nigricans, Scirpus holoschoenus, Oenanthe globulosa, O. lachenalii, etc.) and woody species (Phillyrea angustifolia, Erica scoparia, Myrtus communis, Pistacia lentiscus, etc.). Management and conservation measures

Vegetative growth Flowering Fruiting

Environmental conditions

Hydrology Flooding in winter, even of short duration, appears to be essential for germination or the reactivation of buds and for the growth of young plants. Substrate Loose substrate, quite thin, (less than 10 cm deep), of varying particle size (sand, silt or clay) and acid. Interspecific competition A heliophilous* (light-demanding) plant, which does not thrive in the shade of tall species. Impact of perturbations • A low level of disturbance (by grazing, trampling), which causes a limited degree of substrate compaction, appears to be beneficial. • Heavy, frequent disturbance, for example due to all-terrain vehicles, that causes severe substrate compaction, are very detrimental (Chevanu pool, in Corsica).

10

Current measures

In Corsica, currently no management measures, as a result of the absence of ownership control over land at the sites. Recommendations

In Corsica • The maintenance (and/or re-establishment) of extensive cattle grazing is recommended, as well as the maintenance of routes for walkers and hunters. • At least in the case of the Chevanu pool, a ban on using the site as a car park for several years in succession would probably enable the numbers of this species to increase again.

Bibliography Anonymous, 199912 ; Bissardon & Guibal, 199739 ; Briquet & de Litardière, 193850 ; Casper & Krausch, 198163 ; Géhu et al., 1994156 ; Grime, 1979163 ; Lorenzoni & Paradis, 1997219, 2000221 ; Paradis et al., 2002284 ; Pignatti, 1982292 ; Pottier-Alapetite, 1952298. Authors: Paradis G. & M. L. Pozzo di Borgo Collaborators: Grillas P. & N. Yavercovski

Plant species

Artemisia molinieri

Quézel, Barbero & Loisel

ANGIOSPERMS ASTERACEAE Main synonyms

None French name: Armoise de Molinier Subspecies

None Description/identification criteria

• Sub-woody plant, very aromatic, with the same scent (like camphor) as Artemisia arborescens, 30 to 60 cm tall. • Stem branching above the neck. Leaves bluish-green, finely hispid, especially on the lower sides. Petiole equal in length to the leaf, which is triangular in outline, bipinnatifid* with threadlike lobes 1-1.5 mm wide. • Capitula fairly small, ovoid (2.5 x 2.5 mm), with 10-15 florets arranged in a dense spike. Inner involucral bracts lanceolatespatulate. The achenes (0.8-1 x 0.2-0.3 mm) are greyish with five ill-defined angles, smooth, slightly longitudinally grooved, and cut off somewhat obliquely at the apex*.

Distribution - Ecology

Similar species

Distribution/range

The genus Artemisia is difficult. Historically, A. molinieri has been confused with A. alba Turra31, which belongs to another section. A. molinieri belongs to the Artemisia section which is characterised by its capitula having a glabrous receptacle and being allogamous* with the outer florets female and the disk florets hermaphrodite, all fertile. Within this section, A. molinieri is close to the species with green leaves and with the petiole lacking basal auricles*. Two of these species may be found in France: • Southernwood, (A. abrotanum L.), a cultivated wormwood whose origins are disputed, and which is distinguished from A. molinieri by its lemon scent, larger globose capitula (3-4 x 3-4 mm) with 25-30 florets arranged in a leafy panicle, and its oval inner involucral bracts. • A. chamaemelifolia Vill. of dry grassland in the mountains of southern Europe and beyond, from Spain to Iran. This wormwood is distinguished from A. molinieri by its sessile paired cauline leaves with the lower lobes clasping the stem, and its slightly larger capitula: 2.5-3 mm. A. molinieri, A. chamaemelifolia and some populations of A. abrotanum have 2n = 18. All the wormwoods are found in dry habitats (rock, steppes, rubble etc.), apart from A. molinieri which is the only wetland wormwood.

1 cm

Artemisia molinieri

Endemic* species of the Centre Var (France), occurring in the communes of Besse-sur-Issole and Flassans-sur-Issole. Habitat General description

Large natural depressions of karstic* origin, flooded irregularly depending on the year: flooding varying from more than six months to completely absent. Habitats Directive

“Mediterranean Temporary Pools” (code 3170). CORINE Biotope

22-342: Tall Mediterranean amphibious swards (Preslion cervinae Br.-Bl. 1931), with a type specific to this habitat: the Artemisia molinieri community245, 307. Phytosociology*

Class: Isoeto-Nanojuncetea Order: Isoetalia Alliance: Preslion cervinae Association*: Artemisia molinieri community307 Companion species

Notes: • According to Couteaux & Pons93, this taxon could be of hybridogenic* origin and its two stations could be artificial. • A small buprestid beetle, (Agrilus lacus), whose larva feeds solely on A. molinieri, has recently been described as endemic to the pools of the Centre Var94.

At Gavoty and Redon: • Schoenoplectus lacustris, Scirpus maritimus, Phalaris arundinacea, Alisma lanceolatum, Eleocharis palustris, Oenanthe globulosa, Rorippa sylvestris, Ranunculus ophioglossifolius, Sisymbrella aspera, Veronica anagalloides. • Annual amphibious companion species of high natural-heritage value (for France), subject to competition from the wormwood

11

Mediterranean temporary pools

and favoured by the opening-up of the wormwood mat (by trampling associated with grazing, or cultivation during dry years): Lythrum tribracteatum, Damasonium polyspermum, Heliotropium supinum, Schoenoplectus supinus, Chenopodium urbicum, Crypsis schoenoides, etc. Ecology Biological characteristics

Substrate Colluvial silts of anthropogenic origin (erstwhile erosion of cultivated catchment areas)93. Meso-eutrophic* substrate. Interspecific competition Artemisia molinieri is heliophilous*, and the establishment of woody plants (especially Fraxinus angustifolia) during dry years, or of helophytes* during wet years, is probably harmful to it (no data).

Life form Artemisia molinieri is a competitive perennial species (chamaeophyte*), which forms dense species-poor stands. The few gaps, deriving from past disturbance, permit the development of a diverse annual flora.

Impact of perturbations This species does not appear to be grazed either by sheep (Redon) or by horses (Gavoty), probably because of its high terpene content during the vegetative stage*241.

Biological cycle Vegetative growth begins in February with the appearance of the first green leaves. Its reproductive cycle takes place in summer (from mid-July to the end of August). The seeds begin to be mature from mid-September. The last green leaves wither in October/November. The plant produces runners, also perhaps layers, and it also readily propagates vegetatively. Couteaux & Pons93 considered that vegetative reproduction was the plant’s only means of propagation. The plant is anemphilous382 but the pollen is very poorly dispersed because the stamens remain contained within the corolla of the hermaphrodite florets*93. In addition, the pollen has a very low viability (10 to 30% of pollen viable depending on the population)382. However, the plant produces abundant achenes, which have no pappus* but are very light and are without doubt wind-distributed. Their production is irregular among individuals and stations307, and germination rates are extremely low382. This latter trait must be related to the irregularity of pollen production in a taxon whose origin probably derives from hybridisation*93, or to the demanding ecological conditions with which the wormwood is confronted (grazing, flooding)382. This is no doubt compensated for by the large number of achenes produced by a single plant382. At Lake Redon, many germinations were visible in 2001 after the water had fallen, in areas which had previously been ploughed.

Conservation - Management

Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Dec.

Leafing and vegetative growth Flowering Fruiting

Environmental conditions

Hydrology The habitat of this amphibious species is characterised by: • A flooding period which is often long, from autumn to spring (up to six or nine months), but in some years may be short or absent. • A maximum water depth of 20 to 70 cm, with the optimum apparently around 40 cm.

Assessment of populations and conservation status

There are two well-known sites, Lake Gavoty and Lake Redon, where the plant is extremely abundant (several hundred thousand aerial shoots). The cultivation of part of Lake Redon, in 2001, destroyed a significant part of the station*. The Artemisia molinieri population has since proved to be healthy, many germinations having been seen during the subsequent years. There are several other depressions, which seem capable of having supported the wormwood in the past307, within a radius of a few kilometres from Gavoty and from Redon. Drainage operations and intensive cultivation could have wiped it out here. At three of these depressions, on fallow land, the recent discovery of small populations of A. molinieri (from one to under ten stalks) raises the question of whether these may be relict or alternatively pioneer populations. Moutte & Triat261 found wormwood pollen in the sediment at a fossile lake northwest of Brignoles. They hypothesised that there was formerly a population of A. molinieri at this lake, which is several kilometres away from the currently known populations. However, Couteaux & Pons93 questioned this hypothesis. Legal status of the species/level of protection IUCN 1997 Bern Convention Habitats Directive National and regional protection

National Red Data Books and Red Lists

“endangered” (E)399 France, PACA regional list: Decree of 09/05/1994/Journal Officiel 26/07/1994 France, national Red Data Book: priority species274

Conservation problems/threats Anthropogenic factors

Reproduction Flowering, which is late, is aerial.

12

The deforestation and subdivision of catchment areas leads to the infilling of depressions and changes in water quality. The cultivation of pools in dry years probably constitutes a risk factor.

Plant species

Natural factors

At Lake Gavoty, woody plants (ash, elm) are becoming established as a result of natural succession. This may eventually transform the depression into woodland, not susceptible to flooding and incompatible with the survival of a healthy Artemisia molinieri population. Risks relating to populations

The risk of extinction at any site is very low in both of the populations known for a long time, which are very large (see vol. I, chapter 3, box 27).

• Taxonomic study of A. molinieri, whose status as a species and as an endemic has been questioned. This type of study may only be considered at the scale of the Abrotanum section. • Retention of the band of low woody plants (Christ’s Thorn/ Jerusalem Thorn Paliurus spina-christi, Small-leaved Elm Ulmus minor and Blackthorn Prunus spinosa) around the pools, at the edge of the flooded zones. This plays a vital role as a natural water filter before it discharges into the depression and as a trap for fine materials which could cause the pools to silt up.

Bibliography Management and conservation measures Current measures

• Management plan in progress in the context of the production of the objectives document for Natura 2000* sites, including the two main pools with Artemisia molinieri. • Land acquisitions completed and in progress at Redon, in the framework of LIFE “Temporary Pools”. • Collection and ex-situ preservation of seeds by the Conservatoire Botanique National Méditerranéen de Porquerolles, and precautionary cultivation of the species on Conservatoire land since 1995.

Aboucaya et al,. 20021 ; Berner, 196831 ; Camus, 190360 ; Couteaux & Pons, 198793 ; Curletti & Ponel, 199494 ; Guinochet, 1982167 ; Loisel, 1976215 ; Masotti et al., 2003241 ; Médail & Quézel, 1994245 ; Moutte & Triat, 1968261 ; Olivier et al., 1995274 ; Quézel et al., 1966307 ; Torrel et al., 1999382 ; Tutin et al., 19641980386 ; Walter & Gilett, 1998399. Authors: Michaud H Collaborators: Rombaut D. & N. Yavercovski

Recommendations

• Research into the role of disturbance in the dynamics and conservation of formations containing Artemisia molinieri and of other priority habitats (Lythrion tribracteati and Heleochloion) subject to competition from the wormwood.

13

Mediterranean temporary pools

Crypsis schoenoides

(L.) Lam.

ANGIOSPERMS POACEAE Main synonyms

Heleochloa schoenoides (L.) Host ex Roem. French names: Crypside ovoïde, Crypside faux choin Italian name: Brignolo ovato English name: Swamp Pricklegrass Subspecies

None Description/identification criteria292

• Annual herbaceous plant (therophyte* scapous*). • Stem branched from the base, 5 to 25 cm long, prostrate (rarely erect). • Leaves very typical of grasses (Poaceae) with glabrous, slightly swollen sheaths*, ligule divided into hairs, blade 1 to 3 cm long and 3 to 4 mm wide, at an angle to the stem. • Inflorescence a cylindrical spike, 5 to 8 mm in diameter and 1 to 3 cm long, comprising many spikelets each consisting of one flower with three stamens.

1 cm

Crypsis schoenoides Habitat General description

Distribution/range

Crypsis schoenoides may be found in summer and early autumn, on the dried-out clay-silt edges of mesotrophic* ponds, on the banks of a few hydro-electric reservoirs (Corsica), and in temporary pools (in France, in Morocco in the dayas of the plains and low mountains etc.). It sometimes occurs on the silty edges of watercourses (Corsica, Algeria), in ditches, and also in poached pastures subject to flooding. Its optimal living conditions are clay-silt substrates which undergo an alternation of a lengthy flooded phase and a drying-out phase during summer (sometimes up to the beginning of autumn). The maximum water depth may vary between a few tens of centimetres (pools or ditches) to several metres (hydro-agricultural dams). This species will tolerate a slightly saline substrate.

The taxon is considered to be Paleo-subtropical292.

Habitats Directive

Similar species

Young stages of Crypsis aculeata and Cynodon dactylon, but at the flowering stage confusion is not possible: • The inflorescence of C. dactylon is formed of digitate spikes, long and very slender. • The inflorescence of C. aculeata has the appearance of a tiny capitulum buried among the bracts*, with two stamens per flower.

Distribution - Ecology

“Mediterranean Temporary Pools” (code 3170). France

This species is found at a number of ponds, watercourses and pools on the Mediterranean coast (Roussillon, Languedoc, Provence) and the coast of Corsica. It is also found on the Atlantic coast, from the Gironde to Finistère.

CORINE Biotopes

22.343 Mediterranean halo-nitrophilous swards (Heleochloion). “Post-aestival slightly halophilic and nitrophilous vegetation of temporarily flooded land, with Crypsis schoenoides, C. aculeata, C. alopecuroides and Centaurium spicatum”.

Italy

Several provinces in peninsular Italy (Trentino, Lombardy, Piedmont, Liguria, Emilia-Romagna, Tuscany, Lazio, Abruzzo), Sardinia and Sicily.

Phytosociology* 54, 240, 250, 277, 281, 286, 318, 319

Spain

In Corsica and Spain Class: Isoeto-Nanojuncetea Order: Nanocyperetalia Alliance: Heleochloion Associations* and communities given in the literature: - Crypsio (aculeatae)-Heleochloetum schoenoidis Oberdorfer 1952 (southern mainland Europe, Corsica).

Scattered throughout the country; most frequent in the centre177. Morocco

Tingitan peninsula, northwest Morocco, Middle Atlas. Algeria

Tell region.

14

Crypsis schoenoides (formerly named Heleochloa schoenoides) is characteristic of Heleochloion formations.

Plant species

- Heleochloo schoenoidis-Fimbristyletum biumbellattae Br.-Bl. & Rivas Goday in Rivas Goday 1956 corr.54 (Spain). - Heliotropio supini-Heleochloetum schoenoidis Rivas Goday 1955 (Spain, Corsica). - Lythro flexuosi-Heleochloetum schoenoidis Rivas Martinez 1966 (Spain). - Crypsio schoenoidis-Cyperetum micheliani Martinez Parras et al., 1988 (Spain, Corsica). - Chenopodio chenopodioidis-Crypsidetum schoenoidis Paradis & Lorenzoni 1994 (Corsica). - Echinochloo cruris-galli-Crypsidetum schoenoidis Paradis & Lorenzoni 1994 (Corsica). - Community with Crypsis schoenoides and Corrigiola litoralis (in Paradis & Lorenzoni277) (Corsica). - Community with Crypsis schoenoides and Cotula coronopifolia (in Paradis & Lorenzoni277) (Corsica).

• During summer: growth of plants on a dry substrate (competitive spring vegetation disappears completely or partly, giving way to the development of summer species), followed by flowering on the substrate which has now become very dry. • At the end of summer and beginning of autumn: fruiting (end of August to end of September), then death of the plants and break-up of the spikes. Corsica Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Dec.

Oct.

Nov.

Dec.

Germination Growth Flowering Fruiting Morocco Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Germination Vegetative growth Flowering and fruiting

In mainland France The Helochloion communities have not been studied in detail; a single community is indicated, that with Crypsis aculeata215, in which C. schoenoides and sometimes Cressa cretica also appear.

Dispersal of caryopses

Environmental conditions

In Morocco Crypsis schoenoides is most often found with Corrigiola littoralis, Heliotropium supinum and Hypericum tomentosum. Note whereas the Isoetion communities appear in spring and are oligotrophic*, the Helochloion communities appear in summer-autumn and are meso-eutrophic*. Ecology Biological characteristics

Life form Therophyte* (annual species). Reproduction Aerial. Flowering and fruiting • In Corsica, from mid-August to the end of September. • In Morocco, between June and August. Adaptive strategy (sensu Grime163) Stress-tolerant ruderal (S-R). Disturbance is caused by flooding and by animal grazing during the summer and early autumn. Stress results from the drying-out of the soil during the summer and early autumn, which is variable depending on the topography and on edaphic* characteristics. Caryopses Length: 0.5 mm maximum. Biological cycle • At the end of spring or the beginning of summer: germination of caryopses followed by seedling development during the dry period (date variable, depending on climate and hydrology), on a substrate which is still damp or under a very shallow layer of water (less than 10 cm).

Hydrology Alternation between a long period of flooding in winter-spring and summer drying appears to be essential. Substrate Meso-eutrophic* substrate with fine particles (clay and silt), rich in organic debris and in nutrients, particularly nitrates. It is sometimes oligotrophic*, in the dayas* of Cork-Oak woods in Morocco. It may be slightly brackish at the surface (often more saline deeper down), which explains the presence of Tamarix spp. close to many coastal Crypsis schoenoides stations. There is usually a fairly significant degree of substrate compaction. Interspecific competition As a light-demanding (heliophilous*) species, Crypsis schoenoides does not readily tolerate shading by tall species such as Phragmites australis, Typha latifolia, T. angustifolia, various Scirpus or Paspalum distichum, hence its restriction to bare areas. Impact of perturbations The potential for the establishment and survival of Heleochloion plant communities depends on denudation, generally due to human activity: • Deliberate cutting of helophytes*, to create shooting areas for example. • Grazing and trampling by cattle: trampling by cattle causes a fairly high degree of substrate compaction, which prevents tall helophytes from establishing themselves and does not hamper the germination and growth of the therophytes of the Heleochloion. • etc.

Conservation - Management Assessment of populations In Corsica

This species occurs around ten sites:

15

Mediterranean temporary pools

• Three on the west coast, in the lower Taravo valley (Etang de Tanchiccia, Etang de Canniccia276, 277, 278, 279, 281 (Paradis unpublished observations, 2003). • Two near the Stabiaccio river at Porto-Vecchio280. • Three on the east coast, Etang de Gradugine283, Alzitone reservoir near Ghisonaccia, Teppe Rosse reservoir near Aléria277, 278.

In mainland France

It occurs at more than twenty stations: • Pyrénées-Orientales: Villeneuve-la-Raho reservoir and alluvial plain of the Agly. • Aude and Hérault: Capestang, Poilhès, Ouveillan, Estagnol ponds and elsewhere in the lower valley of the Aude. Sesquiers marsh at Mèze, pools of Notre-Dame-de l’Agenouillade at Agde, pools and ditches at Roque-Haute, Saint-Martin-de-Londres and Saint-Nazaire de Pézan, etc. • Gard: marsh at Aigues-Mortes, ponds of Pujaut and Capelle, and alluvial plain of the Gardon at Dions. • Vaucluse: flood meadows at Monteux. • Bouches-du-Rhône: in the Camargue (ditches at the Tour-duValat, Verdier marsh). • Var: Lake Redon at Flassans, Badelune basins at Cannet-desMaures, pools at Grimaud and Hyères, ditches at Roquebrunesur-Argens. No recent records from Fréjus. In Morocco

More than twenty stations are known from the Tangier, LaracheCasablanca and Middle Atlas regions. Conservation status In France

The population at Tanchiccia (Corsica) appears to be decreasing in comparison with previous observations by Paradis281. On the other hand, all the other populations on the island are either stable or increasing. In mainland France, populations are large and sometimes in very high numbers in the region of Capestang, Ouveillan, Poilhès, etc. The species appears to have decreased on the coast in the PyrénéesOrientales and has disappeared from the Alpes-Maritimes coast332. It is surviving well elsewhere: the number of new sites largely compensates for unconfirmed sites mentioned in earlier records. It seems to have appeared in the Camargue and in the floodplain of the Rhône between Arles and Tarascon as a result of the desalination which followed the increase of ricefields256. In Morocco

The populations appear to be fairly stable despite the increasing pressures on the sites. Legal status of the species/level of protection

IUCN 1997 Bern Convention Habitats Directive National and regional protection

National Red Data Books and Red Lists

16

France, PACA regional list: Decree of 09/05/1994/Journal Officiel 26/07/1994 France, National Red Book: “to be observed”274

Conservation problems/threats Anthropogenic factors

The infilling of pools and ponds (in Morocco and at Tanchiccia in Corsica), drainage for agriculture or permanent flooding for shooting (mainland France), and overdeepening for the extraction of sediments or rock (Morocco) are the main threats. In Morocco, some sites located near large towns are also threatened with destruction in the mid-term by urban development. Natural factors

This plant is favoured by sheep or cattle grazing (France, Morocco) as well as by certain types of management for hunting, such as the creation of pools in reedbeds. Risks relating to populations

In mainland France, the isolation of the Var populations and of the very small inland pools in Languedoc-Roussillon constitutes a potential threat. Management and conservation measures Current measures

• In Corsica: at present there are no management measures in place, due to the absence of control over land ownership at the sites. • In mainland France: the objectives document for the Natura 2000* site pools of the Centre Var takes this species into account, but no conservation measures are known for the other sites. The Conservatoire Botanique Méditerranéen de Porquerolles collects seeds and preserves them (ex-situ). • In Morocco: the species is not protected and does not benefit from any management measures. Recommendations

• Ensure that management by extensive cattle grazing is carried out and that an appropriate hydrological regime is maintained (mainland France, Corsica, Morocco). • In Morocco, organise site monitoring as well as control of urban development in the short term.

Bibliography Anonymous, 199912 ; Bissardon & Guibal, 199739 ; BraunBlanquet et al., 195249 ; Brullo & Minissale, 199854 ; Grime, 1979163 ; Loisel, 1976215 ; Maire, 1952-1987230 ; Martinez Parras et al., 1988240 ; Molero & Romo, 1988250 ; Molinier & Tallon, 1974256 ; Paradis & Lorenzoni, 1994277, 278 ; Paradis & Orsini, 1992276 ; Paradis & Pozzo di Borgo, 2000280 ; Paradis & Piazza, 1995279 ; Paradis et al., 2002283 ; Paradis, 1992281 ; Peinado Lorca et al., 1988286 ; Pignatti, 1982292 ; Rivas Goday, 1964318, 1970319 ; Rivas-Martínez et al., 2001320 ; Salanon et al., 1994332 ; Valdés et al., 2002391. Authors: Paradis G. & M. L. Pozzo-di-Borgo Collaborators: Grillas P., H. Michaud, L. Rhazi & N. Yavercovski

Plant species

Damasonium polyspermum

Coss.

ANGIOSPERMS ALISMATACEAE Main synonyms

D. stellatum Thuill. var. polyspermum (Coss.) Loret & Barrandon D. stellatum Thuill. var. polyspermum (Coss.) P. Fourn. D. alisma Mill. proles polyspermum (Coss.) Rouy “D. alisma Mill. var. polyspermum Loret & Barrandon” in Guin. & R.Vilm. D. alisma Miller subsp. polyspermum (Coss.) Maire French name: Etoile d’eau à nombreuses graines Spanish names: Saeta de agua, Cola de Golondrina, Almea,

Azumbar

1 cm

Italian name: Mestolaccia siciliana Moroccan name: Mizmar er raaï English names: Starfruit, Thrumwort

Damasonium polyspermum

Subspecies

None Description/identification criteria

• Small annual plant with the leaves all basal and arranged in a rosette. • First submerged leaves linear and not persisting, subsequently replaced by floating leaves with long petioles and narrow lanceolate blades, attenuated or more or less truncated at the base, with faint transverse venation. • Leaves produced during the non-flooded phase similar to the floating leaves but with a shorter petiole and a thicker blade. • Inflorescence most often with one, sometimes several superposed umbelliform whorls. Flowers hermaphrodite with three persistent green sepals and three deciduous coloured petals. Petals 2 to 2.5 times as long as the sepals (5.4 ± 0.4 mm x 4.1 ± 0.6 mm), white, marked with yellow at the base and more or less tinged with pink at the tip. • Fruit consisting of 6-9 carpels arranged in a star shape (follicles), falling when ripe. Five to more than twenty cylindrical seeds with transverse ridges (dimensions: 0.9-1.2 mm x 0.5-0.7 mm), in each mature carpel.

Baldellia ranunculoides (L.) Parl. is distinguished, at all stages of development, by the scent of coriander that is released when the leaves are crushed. During flowering or fruiting, the arrangement of the many obovate carpels (≈ 2 mm) in a globose head removes any doubt. Alisma spp.: The three Alisma species included in the French flora are usually far more robust than Damasonium polyspermum. At flowering or fruiting they are easily distinguished by their many carpels, which are blunt oval shaped, very tightly packed and whorled in a single row. Unlike Damasonium, these are all perennial* species.

Distribution - Ecology Distribution/range

Western Mediterranean species.

Similar species

France

Damasonium alisma Mill. has only two large seeds (1.7-2.5 mm x 0.8-1.2 mm) per carpel when the fruit is ripe. According to some authors, additional distinguishing characters include the plant’s often larger size and more robust build, leaves rounded to cordate* at the base with oblique transverse veins visible against the light, smaller petals (4.2 ± 0.5 mm x 3.7 ± 0.5 mm) lacking pink colour at the tips, and shorter anthers (0.75 ± 0.006 mm v. 0.82 ± 0.12 mm). However, the relevance of these characters needs to be confirmed, especially in relation to the influence of the environment (depth of water, length of submersion, etc.).

Mediterranean France from Biterrois (Hérault) to the Centre Var (Var).

The two Damasonium are not always separated by botanists. According to Rich & Nicholls-Vuille316, they are also separable on the basis of their largely different geographical ranges and by their chromosome numbers (D. alisma 2n = 28, tetraploid; D. polyspermum 2n = 14, diploid*).

Other Mediterranean countries

North Africa: in freshwater dayas*, especially in mountains, in Cyrenaica, in the High Plateaux of northwestern Algeria and in the Middle Atlas in Morocco. Italy: southwest Sicily in the coastal zone292. Edges of ponds and pools in the Iberian Peninsula, in central and southern Spain as well as Portugal, Greece and Syria. Habitat General description

Land temporarily flooded by fresh or slightly brackish water in the meso-Mediterranean zone: pools, edges of ponds and lagoons.

17

Mediterranean temporary pools

Habitats Directive

“Mediterranean Temporary Pools” (code 3170).

Southern France Jan.

CORINE Biotopes

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Dec.

Germination and vegetative growth

22.32 Euro-Siberian dwarf annual amphibious swards (Cyperetalia fusci). 22.34 Mediterraneo-Atlantic amphibious communities (Isoetalia). 22.341 Short Mediterranean amphibious swards (Isoetion Br.-Bl. 1931).

Flowering Formation and ripening of fruits Seed dispersal

Environmental conditions Phytosociology*

• Euro-Siberian dwarf annual amphibious swards (Cyperetalia fusci): - Lythrion tribracteati (e.g. the Lythrum tribracteatum-Teucrium aristatum association*215, 252. - Elatino triandrae-Eleocharition ovatae (e.g. Elatinetum macropodae). • Southern amphibious communities (Isoetalia): - Short Mediterranean amphibious swards (Isoetion). • Damasonium polyspermum sometimes overlaps into formations appearing later in the year and having a more distinctly eutrophic* or even halophilic* character: Heleochloion schoenoidis.

Hydrology The germination of seeds, which is very variable from one year to another, requires immersion in water (a few centimetres) or waterlogged soil in spring, and little vegetation cover (strong insolation). The plant can remain in the vegetative stage under very shallow water, with characteristic floating leaves, but must emerge quickly for flowering and fruiting. The size and density of individuals and the length of their life cycles exhibit spatial variability within the pool, in relation to hydrological gradients (duration of flooding, dates of flooding and drying out) that are themselves dependent on topographical gradients.

These various vegetation types are frequently found in mosaics with perennial communities based on Eleocharis palustris, Bolboschoenus maritimus, Phragmites australis, Mentha cervina, Artemisia molinieri, Isoetes setacea, etc. (Preslion cervinae, Isoetion, Phragmition, etc.).

Substrate Plant tolerant of a wide range of soils and of slightly saline conditions.

Companion species

The species most often associated with Damasonium polyspermum are: Lythrum tribracteatum at all the French localities, with in addition, depending on the site, L. hyssopifolium, Pulicaria vulgaris, P. sicula, Myosurus minimus s.l., Juncus bufonius s.l., J. tenageia and Herniaria glabra. Ecology

Interspecific competition A very light-demanding (heliophilous*) species, and therefore very sensitive to any plant cover at the germination stage. Impact of perturbations Disturbance which helps to control competing vegetation is beneficial, whereas disturbance resulting in the burial of seeds impede germination (trampling and digging by livestock or wild animals, ploughing, overdeepening or dredging of the pool, etc.) (see Vol. 1, box 38).

Biological characteristics

Conservation - Management Life form Annual species (therophyte*). Reproduction Aerial. Seeds (see above § Description) The lifespan of seeds in the sediment is not known, but observations of the dynamics of natural populations indicate that it is long. Biological cycle Seed germination and growth of leaves during the flooded phase (end of winter, beginning of spring). Flowering in spring (from April to June) up to the first dry periods, but may be prolonged or repeated in autumn if the substrate is sufficiently wet. Ripening of fruits and dispersal of seeds during the dry phase (summer) and before the autumn rains which mark the end of the cycle.

18

Assessment of populations France

The species currently occurs at 10 sites: • Hérault: Vendres/Sauvian (Malhol de l’eau), Agde (Notre-Dame de l’Agenouillade), Montarnaud (pool on the Tamareau garrigues). • Gard: La Capelle-et-Masmolène (Etang de la Capelle). • Bouches-du-Rhône: Arles (Tour-du-Valat and Lanau), SaintAntonin-sur-Bayon (Plateau du Cengle). • Var: Gonfaron (Lake Bonne Cougne), Flassans-sur-Issole (Lake Redon), Besse-sur-Issole (Lake Gavoty). • Absent from Corsica. Italy

Only one site is known, on the coast of Sicily86. Spain (Medina, pers. com.) The species occurs in 13 provinces in the centre and north of the country (between 50 and 100 sites), and three provinces in the south of the country (about twenty sites).

Plant species

Portugal

Natural factors

A single site supports this species (Medina, pers. com.).

As the species is an annual, it does not compete well with colonial perennial species (Artemisia molinieri, Mentha cervina, Isoetes spp. Eleocharis palustris, etc.), this being partly compensated for by its abundant seedbank. In many depressions, the appearance of Damasonium is irregular from one year to another37.

Morocco

The species is not considered to be rare133. Note

As the two species of Damasonium have not always been separated by botanists, some data remain uncertain: for example old publications list D. alisma sensu lato from the closed depressions of the Roussillon plain (between Têt and Tech)351, in the coastal pools at Argelès-sur-Mer155 and in the Montmajour marsh (Arles/Fontvieille) (Jacquemin 1848 in Molinier257). In the French Mediterranean region, D. alisma is only known with certainty from the extreme south of Larzac30 and appears therefore to be absent from the meso-Mediterranean zone. Conservation status

In France, eight populations have disappeared, or have not been seen again, in the communes of Sauvian (Vendres plateau), Portiragnes/Vias (Roque-Haute), Agde (Rigaud pools), Redessan, Jonquières, Manduel (pools of Jonquières, Campuget and Redessan), Bellegarde, La Barben (Estagnolet pool).

Risks relating to populations

This risk is low in the absence of any major perturbations, due to the fecundity of the plant and the longevity of the seeds in the sediment. Management and conservation measures Current measures

The Lanau pool (Bouches-du-Rhône) was acquired by the CEEP in 1998. The only management measures known at present are: • Monitoring of the population of the Lanau pool in the Crau, in relation to grazing management. • A management plan in progress for the pools of the Centre Var, in the context of drawing up the Objectives Documents for Natura 2000* sites. • The ex-situ preservation of seeds by the Conservatoire Botanique National Méditerranéen de Porquerolles.

Legal status of the species/level of protection Recommendations IUCN 1997 Bern Convention Habitats Directive National and regional protection

National Red Data Books and Red Lists

- France, national list: Decree of 20/01/1982/Journal Officiel 13/05/1982, amended by Decree of 31/08/1995/ Journal Officiel 17/10/1995 - France, national Red Data Book: “to be observed”274 - Greece, national Red Data Book: “vulnerable”290 - Italy, national Red Data Book: “vulnerable”86

Conservation problems/threats Anthropogenic factors

Several populations have disappeared following the infilling or drainage of the depressions which supported them, in connection with agricultural developments (Vendres plateau, pools and ponds of the Costière Nîmoise) or with urban development (Rigaud pools at Agde).

• Promotion or maintenance of grazing, which is favourable to the growth of the species, as is any other means whereby perennial vegetation communities are opened up. • Promotion of the acquisition and management of Damasonium sites, which are in addition often rich in other rare species.

Bibliography Bernard, 199730 ; Bigot, 195536, 37 ; Braun-Blanquet, 193548 ; Conti et al., 199286 ; Danton & Baffray, 199596 ; Fennane & Ibn Tattou, 1998133 ; Gautier, 1898155 ; Guinochet & Vilmorin, 1978166 ; Maire, 1952-1987230 ; Médail et al., 1998246 ; Michaud & Molina, 1999249 ; Molinier, 1981257 ; Molinier & Tallon, 1947252 ; Molinier & Tallon, 1948253 ; Olivier et al., 1995274 ; Phitos et al., 1995290 ; Pignatti, 1982292 ; Pouzolz de, 1862100, 101 ; Rich & Nicholls-Vuille, 2001316 ; Rivas-Goday, 1970319 ; Simonneau, 1967351 ; Vuille, 1987398. Author: Michaud H. Collaborator: Yavercovski N.

19

Mediterranean temporary pools

Elatine brochonii

Clavaud

ANGIOSPERMS ELATINACEAE Main synonyms

E. hydropiper L. var. pedunculata (Moris) Fiori E. hexandra (Lapierre) DC. subsp. brochonii (Clavaud) P. Fourn. French name: Elatine de Brochon Subspecies

None Description/identification criteria76, 96, 138, 381

• Herbaceous plant, amphibious, short (2-7 cm) with a relatively long and significant root system. • Leaves opposite, spatulate, more or less petiolate, with a smooth slender stalk, prostrate, much-branched and rooting at the nodes. • Flowers hermaphrodite, actinomorphic*, small (not more than 5 mm), sessile, solitary in the leaf axils, with three free sepals persisting in fruit scarcely longer than the fruit, and three free petals, white with pink veins. Androecium of six stamens arranged on two verticils. • Fruit: spherical, slightly flattened capsule* with three valves. Seeds many, small, reticulate, straight or slightly arcuate, with longitudinal ribs, transversely grooved. • Plant brownish-coloured at the end of the cycle.

On the Atlantic plains of Morocco, temporary pools with E. brochonii are found in Cork-Oak woods (Mamora and Benslimane) under sub-humid to semi-arid climatic conditions. Flooding usually takes place in December-January, solely by rainwater, and they dry out during April-May. E. brochonii also occurs in Morocco in mountain pools in siliceous terrain133, 228, 229, 230.

Similar species

Habitats Directive

• Elatine hexandra has pedicellate flowers (sessile in E. brochonii). • Lythrum borysthenicum has crimson petals (not whitish-pink), sepals longer than the capsules (scarcely longer in E. brochonii), and a rough stem which is not slender.

“Mediterranean Temporary Pools” (code 3170).

0.25 cm

Elatine brochonii

CORINE Biotopes

22.34 Southern amphibious communities (Isoetalia). 22.341 Short Mediterranean amphibious swards (Isoetion). Phytosociology*

Distribution - Ecology

Occurs in the southwest (Gironde, Landes, Pyrénées-Atlantique) and in Corsica81, 136, 219, 274, 346, 392.

Elatine brochonii is found together with Juncus pygmaeus, J. bufonius, Lythrum hyssopifolia, Illecebrum verticillatum, Exaculum pusillum and Isoetes velata, which means that it is included in two alliances, the Isoetion velatae and the Cicendion219: Class: Isoeto-Nanojuncetea Order: Isoetalia Alliance: Isoetion velatae Br.-Bl. 1931 Alliance: Cicendion filiformis Rivas Goday (1961, 1964) Br-Bl. 1967.

Other Mediterranean countries

Ecology

Distribution/range

A western Mediterranean species. France

Spain76, Morocco303, Algeria292, 386. Biological characteristics Habitat General description

In France, Elatine brochonii is found on the edges of temporary pools in southwest France on siliceous substrates with irregular flooding between years. In Corsica, this species occurs at a single site (Padulellu pool) in a depression that cuts into a granitic ridge (105 m altitude) covered with low matorral*. Flooding and drying out, which are irregular from one year to another, take place through rainwater and evaporation. The maximum water level recorded in 1996 (a very wet year) was 80 cm219.

20

Life form Therophyte* (annual). Adaptive strategy Stress-tolerant-ruderals (S-R)163. Reproduction Aerial.

Plant species

Seeds Seeds are abundant (33 to 50 seeds per capsule), long-lived* in the sediment, small (length: 0.63 mm, width: 0.27 mm), and weigh less than 1.4 mg. Biological cycle Amphibious plant with a very short life cycle. It begins its biological cycle under water or just after emergence and ends it out of water. Its appearance, irregular from one year to another and on varying dates, has been observed in southwest France346. In Corsica germination takes place at the end of spring (MayJune), only during wet years. Flowering occurs from May to July, fruiting in June and July, and the seeds are dispersed in July and August. In Morocco, Elatine brochonii germinates in February-March and in April forms flower buds which very quickly develop into flowers. Fruiting takes place from April to June with the dispersal of seeds313.

Conservation - Management Assessment of populations

In mainland France, 13 stations were formerly known in the southwest, of which only one appears to have survived274; there is one site in Corsica (Padulellu pool219), and at least eight in Morocco133, 346. Conservation status

The interannual variability of the species causes difficulty in evaluating the status of the populations. However, it no longer appears to be present at most of its mainland French sites, and the Corsican population remains vulnerable. In Morocco the populations appear to be better preserved, but are subject to rapidly increasing human pressures, especially around Rabat. The high survival rate of the seed stocks* in the soil gives grounds for hoping that populations may still exist in the dormant state.

Corsica Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Dec.

Germination

IUCN 1997 Bern Convention Habitats Directive National and regional protection

Flowering Fruiting Seed dispersal Morocco Jan.

Feb.

March

April

May

June

July

Aug.

Legal status of the species/level of protection

Sept.

Oct.

Nov.

Dec.

Germination Flowering Fruiting Seed dispersal

National Red Data Books and Red Lists

Environmental conditions

Hydrology Seed germination requires the soil to be flooded (a few centimetres of water) or saturated during March, and intense insolation (bare soil). The plant may remain in the vegetative stage under very shallow water, but must emerge quickly for flowering and fruiting314. Substrate Acid, oligotrophic* at Moroccan stations, mesotrophic* in Corsica.

- France, national list: Decree of 20/01/1982/Journal Officiel 13/05/1982, amended by the Decree of 31/08/1995/Journal Officiel 17/10/1995 - France, national Red Data Book274: priority species - Spain, national Red List: “vulnerable”13 - Morocco: “very rare” (RR)133, 181, 381

Conservation problems/threats Anthropogenic factors

Digging out and removal of sediments, frequent vehicle movements creating tracks through the pool (Mamora pool in Morocco), or the formation of a sandy alluvial fan partly covering the Elatine station (Padulellu pool in Corsica). Natural factors

Interspecific competition A very light-demanding species (heliophilous*), Elatine brochonii is very sensitive to any plant cover at the germination stage. Impact of perturbations Perturbations which help to control competing vegetation are favourable to Elatine brochonii, while those resulting in burial of the seeds will impede germination (87% germination rate for seeds on the surface compared with only 13% and 6% for seeds buried in sand to depths of 2 mm and 5 mm respectively). This burial may be caused in the field by the passage or digging of wild and domestic animals, sedimentation (Padulellu) and the movements of vehicles.

Competition from terrestrial vegetation (Cistus monspeliensis, Pistacia lentiscus, Myrtus communis, etc.) or from helophytes (Scirpus maritimus, Eleocharis palustris, etc.) only constitutes a temporary problem, inherent to Mediterranean temporary pools, if the hydrological regime is not altered. Risks relating to populations

Despite its large seedbank, which enables populations to persist, risks of extinction may arise in the future, especially at the sites south of the Mediterranean where the species may be considered to be at the southern limit of its range. The high frequency of dry years in these countries (Morocco) could result in the long term in a decline in the populations of Elatine brochonii. The low number of sites increases the risk of extinction.

21

Mediterranean temporary pools

Management and conservation measures

Bibliography

Current measures

Anonymous, 199912 ; Anonymous, 200013 ; Bissardon & Guibal, 199739 ; Cirujano & Velayos, 199376 ; Clavaud, 188381 ; Danton & Baffray, 199596 ; Fennane & Ibn Tattou, 1998133 ; Fiton, 1916136 ; Fournier, 1936138 ; Grime, 1979163 ; Jahandiez & Maire, 19311934181 ; Lesouëf & Richard, 1995210 ; Lorenzoni & Paradis, 1997219 ; Maire, 1926228, 1932229, 1952-1987230 ; Médail et al., 1996243 ; Olivier et al., 1995274 ; Pignatti, 1982292 ; Quézel & Santa, 1962-1963303 ; Rhazi et al,. 2001313, 314 ; Rivas-Goday, 1970319 ; Schotsman & Bosserdet, 1966346 ; Schotsman, 1985347 ; Titolet & Rhazi, 1999381 ; Tutin et al., 1964-1980386 ; Vanden Berghen392, 1966 ; Vivant, 1960396.

None Recommendations

• In France and in Morocco systematic monitoring of the known populations is recommended, as well as an assessment of the environment, the condition of the populations (including seedbanks), and the possible opportunities for their restoration. • At the Padulellu pool (Corsica), it is a matter of urgency that the process of sedimentation be halted (sand originating from the erosion of the track running next to the pool) and, to this end, that the belt of Myrtle maquis be reinstated; the driving and parking of vehicles in the pool should also be prohibited. • In Morocco, the continuance of grazing appears to be beneficial in controlling the amount of competing vegetation.

22

Authors: Rhazi L. Collaborators: Grillas P., G. Paradis & D. Titolet

Plant species

Eryngium pusillum

L.

ANGIOSPERMS APIACEAE Main synonyms

E. barrelieri Boiss. French name: Panicaut nain de Barrelier Italian name: Calcatreppola di Barrelier Subspecies

None Description/identification criteria

• Short, thistle-like plant, in low-growing clumps. • The clump is composed of a more or less large number of units, depending on its age. A unit consists of a main stem, bearing a rosette of leaves at its base and ending in a capitulum (inflorescence). The stem may branch and give rise to new units. • The young rosette leaves are elongated and not prickly. As they age they become stiff, with thin spines along their edges. • There may be very many capitula on a single clump. Each carries several very prickly bracts. The flowers are very small and bluish. They produce tiny achenes (indehiscent dry fruits)220, 274, 292. Similar species

None

Distribution - Ecology

1 cm

Eryngium pusillum

In spring 22.34 Mediterraneo-Atlantic amphibious communities. Phytosociology*

In summer and early autumn: Mentha pulegium and Eryngium pusillum meadows, classifiable within the Trifolio-Cynodontion alliance (order Plantaginetalia majoris156 and class MolinioArrhenatheretea), which does not appear to correspond to any categories in CORINE Biotopes. Ecology

Distribution/range Biological characteristics

Southern Mediterranean species France

Only in southern Corsica Other Mediterranean countries

Life form Perennial species, hemicryptophyte*. It is formed of sympodial* units. Rosettes of leaves develop at the base of the units, which are monocarpic (only flower once). The different units together form a low clump220.

Sardinia, Sicily, southern Italy, Tunisia, Algeria, Syria, Morocco274, 292. Habitat General description

The Musella depression is a poljé* (karstic* depression) which passes through very different ecophases* over the course of a year, whose two extremes are a winter flooded ecophase and a very dry summer ecophase. Flooding, which is due mainly to the rising water table in the underlying limestone, occurs in winter during wet years and may in exceptional cases last from November to April (e.g. in 1996 and 2001). Drying out begins in March (in April in very wet years), by lowering of the water table. In summer, the clay-silt substrate becomes very cracked as it dries out. Habitats Directive

“Mediterranean Temporary Pools” (code 3170).

Germination Aquatic. Reproduction Aerial for flowering220. Survival strategy of the clumps Stress-tolerant (S) according to the terminology of Grime163 Biological cycle Germination in November if the site is flooded, dispersal of young plants during the flooded period (hydrochory), vegetative growth from November to May, flowering end of May and June, fruiting in June and July, dispersal of achenes for a very short distance from the parent plant and dispersal of the seedlings by floatation (hydrochory)220. During and especially at the end of summer the leaves wilt and disappear, while new leaves appear with the first rain of September-October.

CORINE Biotopes

In winter flooding phase 22.5 Oligo-mesotrophic* calcium-rich waters.

23

Mediterranean temporary pools

Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Conservation problems/threats

Dec.







Vegetative growth Vegetative growth Germination and dispersal of seedlings

Flowering Fruiting and dispersal of achenes

Environmental conditions

Hydrology The alternation of a flooded ecophase (from the end of autumn to the beginning of spring) and a dry ecophase is necessary. In the absence of winter flooding the seeds do not germinate in spring. An absence of germination for several years is, however, not a handicap for this species, as the large amount of germination during wet years, plus vegetative* reproduction, enable the population to rebuild itself. Substrate Mesotrophic*.

Anthropogenic factors

At Musella, changes in the substrate have been observed as a result of ploughing and sowing. In addition, the cessation of cattle grazing by the site manager in 2002 and its replacement with grazing by a flock of sheep poses the risk, for the future, of altering the ecological conditions at the site. At Vix, development works destroyed the population and its habitat274. Natural factors

At Musella, the Eryngium pusillum population should not experience any problems in the short term. However, the development of woody plants (Blackthorn, Ulmus minor and Rubus ulmifolius could be detrimental in the medium term (shading). Management and conservation measures Current measures

Interspecific competition A strictly heliophilous* species, negatively affected by the shade from taller vegetation. Impact of perturbations A prickly species once the site dries out, not (or very little) grazed, but highly stimulated by grazing cattle, which restrict competing species and create bare areas which are favourable for germination.

• Satisfactory at the Musella site up to 2001 as a result of the regular extensive cattle grazing. Moreover, the Musella pool is part of a Natura 2000* site. • Seed collection and ex-situ storage by the Conservatoire Botanique National Méditerranéen de Porquerolles, and precautionary cultivation of the species on Conservatoire land since 1993. Recommendations

As shading has a detrimental effect on the species, surveillance and control of the spread of woody plants is advisable (scrub clearing, maintenance of grazing). Accordingly, keeping a flock of sheep at Musella for part of the year is strongly recommended.

Conservation - Management Assessment of populations

Bibliography

A single site in France, in the Musella depression (Bonifacio limestone plateau, southern Corsica). In Sardinia (Italy) this species occurs at temporary pools of the Giara di Gesturi basaltic plateau258.

Anonymous, 199912 ; Bissardon & Guibal, 199739 ; Fennane & Ibn Tattou, 1998133 ; Géhu et al., 1994156 ; Grime, 1979163 ; Jahandiez & Maire, 1931-1934181 ; Lorenzoni & Paradis, 1998220 ; Mossa, 1986258 ; Olivier et al., 1995274 ; Pignatti, 1982292 ; Walter & Gillett, 1998399.

Conservation status Author: Paradis G.

In Corsica, the Musella population is increasing, slowly but surely220, but it has disappeared from the two Corsican stations at Vix and Vico274. Legal status of the species/level of protection IUCN 1997 Bern Convention Habitats Directive National and regional protection

National Red Data Books and Red Lists

24

- France, national list: Decree of 20/01/1982/Journal Officiel 13/05/1982, amended by the Decree of 31/08/1995/Journal Officiel 17/10/1995 - France, national Red Data Book : priority species274 - Morocco: “very rare” (RR)133, 181

Plant species

Illecebrum verticillatum L. ANGIOSPERMS CARYOPHYLLACEAE Main synonyms

Corrigiola verticillata (L.) Kuntze French name: Illécèbre verticillé Italian name: Corriggiola verticillata Portuguese name: Aranhoes Moroccan name: Souifa diel el ma English name: Coral-necklace

1 cm Subspecies

Illecebrum verticillatum

None Description/identification criteria

• Annual herb with thread-like stem, often radicant, quadrangular, glabrous, pinkish. • Leaves opposite (can appear whorled due to the development of axillary clusters), entire, sub-sessile, obovate, 1.3 to 2.6 mm, slightly fleshy, with stipules small (1 mm) and scarious. • Inflorescence of pseudo-whorls*, each formed of two cymes* per node. Each cyme includes four to six flowers. • Flower: calyx with five sepals (1.5 to 2.5 mm), white, recurved, concave on the interior side, mucronate* at the apex. No corolla. Flower with five very small fertile stamens and five minute sterile stamens. Pistil with small ovoid ovary, with very short style and two stigmas. • Fruit: achene with membranous pericarp, retained within the persistent calyx. • Seed sub-elliptical, shining brown, about 1 mm long. The species exists in two very different forms, one terrestrial and one aquatic63: • The terrestrial form develops in spring when the pool has dried out. The stems are short (about 5 to 20 cm) and prostrate, with very short internodes. The flowers are very abundant. • The aquatic form develops in winter and spring. The stems are upright and unranked and may be up to 60 or 70 cm long, the internodes are very long and the submerged leaves are very fine. In March and April the upright stems develop floating branches at the surface of the water which bear a few inflorescences and resemble the terrestrial form, i.e. short internodes and more obovate leaves. The aquatic form produces very few flowers compared with the terrestrial form.

Distribution - Ecology Distribution/range63, 174, 292, 348

This species is “widely distributed in Europe from southwest Spain to Sweden and Poland, in a band extending 200 to 400 km inland from the Atlantic seaboard”348. It is rarer in the Mediterranean region. France

Western France and Corsica. Other Mediterranean countries

Western half of Spain (absent from the Balearic Islands), Portugal, Italy (Lombardy, Piedmont, Tuscany, Marche, Lazio, Sardinia) and near the coasts of North Africa (Tunisia, Algeria and Morocco). Non-Mediterranean region

Azores, Canaries. Habitat 1. “Mediterranean temporary pool” habitat General description

In Corsica, Sardinia, North Africa and more rarely elsewhere, Illecebrum verticillatum is a component, significant in biomass* terms, of the flooded phase of several temporary pools (aquatic form).

Similar species

The terrestrial form may be confused with Paronychia echinulata, but this lives on dry substrates. In the vegetative stage, confusion is also possible with Corrigiola littoralis, but this species has long basal leaves (up to 6 cm) in rosettes.

I. verticillatum is usually associated here with Apium crassipes, Ranunculus peltatus, R. ophioglossifolius, Myriophyllum alterniflorum, Baldellia ranunculoides, Eryngium barrelieri, Isoetes velata, etc.72, 219, 221, 298, 314 I. verticillatum also occurs in its terrestrial form at the edges of pools, in small transitory rain pools within Mediterranean maquis, and in cupular pools* (for example in Corsica at Evisa). Here it forms communities in association with Anagallis parviflora, Cicendia filiformis, Exaculum pusillum, Kickxia cirrhosa, Radiola linoides, Silene laeta, Solenopsis laurentia etc.54

25

Mediterranean temporary pools

Habitats Directive

“Mediterranean Temporary Pools” (code 3170). CORINE Biotopes

22.34 Southern amphibious communities (Isoetalia). 22.341 Short Mediterranean amphibious swards (Isoetion). 22.3412 Mediterranean aquatic communities with Isoetes: communities including Isoetes velata of water bodies of variable depth.

• Growth during winter in the water body. • Flower branches on the water surface in April. • Flowering (limited) in April and May. • Fruiting during the drying-out of the pool (from May, often June). • Seeds in the dormant stage in summer and until the rains of autumn-winter.

Morocco Jan.

Feb.

Phytosociology*




In its aquatic form Illecebrum verticillatum is a member of the submerged phase of the Isoetion formations within the Isoetes velata communities. Some authors place the hydrophyte* formations to which it belongs in the communities of the Potamogetonetea pectinati class221, 284.

Germination

March

April

May

June

July

Aug.

Sept.

Oct.

2. Other habitats

Illecebrum verticillatum also occurs, in its terrestrial form, in other biotopes which flood temporarily and dry out rapidly at the end of the winter: ruts in tracks over a wide area of Europe, small transitory rain pools in clearings in the Atlantic heathlands, cupular pools (Massif Armoricain), or small bare depressions in overgrazed grasslands. Ecology Biological characteristics

Life form Therophyte (annual). Reproduction Aerial.

Dec.
Germination

Vegetative growth Flowering Fruiting

Corsica Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Dec.
Germination


Germination

The communities and associations* which include I. verticillatum and which are situated in places undergoing brief periods of flooding have received more attention from phytosociologists*. Thus, according Brullo & Minissale54, the phytosociological affinities of such communities and associations are as follows: Class: Isoeto-Nanojuncetea Order: Isoetalia - Alliance Cicendio-Solenopsion laurentiae Various associations, in which I. verticillatum is a species which ranks highly in terms of occurrence - Alliance Agrostion pourretii Association Illecebro-Agrostietum pourretii

Nov.

Vegetative growth Flowering Fruiting

Environmental conditions

Hydrology Winter flooding or saturation of the sediment, even for a brief period, appears to be necessary for the germination of Illecebrum seeds. This germination, which takes place rapidly as soon as the first rain falls, enables the species to appear in the field almost every year314. Substrate Loose, very thin substrate (less than 3 cm deep) acid, siliceous, sandy or gritty and oligotrophic*. Interspecific competition This strictly heliophilous* species does not tolerate cover from other species, hence its confinement to substrates that are very thin and poor in absorbable mineral salts. In very turbid pools (Mamora, Morocco) it only grows in abundance after the water has fallen. Impact of perturbations Disturbance resulting in the opening up of the vegetation is clearly favourable (e.g. trampling).

Conservation - Management Assessment of populations

Flowering From mid-March to the end of May (Corsica, Morocco).

In Corsica

The species probably occurs at more than 20 sites. Adaptive strategy (sensu Grime163) Stress-tolerant-ruderal (SR). Description of seeds Length approx. 1 mm. Biological cycle In the deep parts of temporary pools: • Germination during winter (from December).

26

In Morocco

There are over thirty stations distributed between the Rif, northern Atlantic Morocco, and mid-Atlantic Morocco. These sites are located on siliceous soils of the plains and low mountains subject to semi-arid, sub-humid and humid Mediterranean bioclimates*134.

Plant species

In Spain

The species is present at 76 sites scattered through some twenty provinces177. Conservation status

In Corsica, Spain, Portugal and Morocco the species is not threatened, appearing to be stable despite more or less wide interannual fluctuations depending on the site. Legal status of the species/level of protection IUCN 1997 Bern Convention Habitats Directive National and regional protection National Red Data Books and Red Lists

-

nigricans, Oenanthe globulosa etc.) and woody species (Phillyrea angustifolia, Erica scoparia, Myrtus communis, Pistacia lentiscus etc.). This is true of the temporary pools of southern Corsica, where the Illecebrum verticillatum population sizes are very small (Mura dell’Unda, Arasu, Muratellu pools), probably due to the cessation of cattle grazing several years ago which has led to the establishment of a heavy biomass of herbaceous plants. In Morocco Practically all the sites are grazed, allowing the vegetation to be kept open and preventing any possible spread by the surrounding woody plants. Management and conservation measures Current measures

Conservation problems/threats

At present, in Corsica as well as in Morocco, there are no management measures, due to the lack of control over land ownership at the sites.

Anthropogenic factors

Recommendations

In Corsica Up until now, there appear to be potential anthropogenic threats at very few sites.

• In Corsica, the maintenance (and/or re-establishment) of extensive cattle grazing is advisable, as well as the maintenance of routes for walkers and hunters. • In Morocco control of urban development in the short term is recommended.

In Morocco Urban development and the filling-in of pools are the most important threats. Thus some sites in the Mamora forest have disappeared following the construction of the Rabat-Fez motorway. Some other sites close to large towns (Casablanca, Rabat, Benslimane, Bouznika, etc.) are also threatened by urban development in the medium term. Natural factors

In Corsica There is a risk that the likely discontinuation of extensive cattle grazing in the near future will result in a severe decline at several sites due to the spread of tall herbaceous plants (Schoenus

Bibliography Abbayes des, 1946106 ; Anonymous, 199912 ; Bissardon & Guibal, 199739 ; Brullo & Minissale, 199854 ; Casper & Krausch, 198163 ; Chevassut & Quézel, 195672 ; Fennane et al., 1999134 ; Grime, 1979163 ; Herra 1990174 ; Lorenzoni & Paradis, 1997219, 2000221 ; Paradis et al., 2002284 ; Pietsch, 1973291 ; Pignatti, 1982292 ; Pottier-Alapetite, 1952298 ; Rhazi et al., 2001314 ; Schumacker, 1978348 ; Sissingh, 1957352 ; Valdés et al., 2002391. Authors: Paradis G. & M. L. Pozzo di Borgo Collaborators: Grillas P., L. Rhazi & N. Yavercovski

27

Mediterranean temporary pools

Isoetes duriei Bory (1) & Isoetes histrix Bory (2)

0.35 mm

LYCOPODIOPHYTES300 ISOETACEAE

megaspore

Main synonyms

(1) I. duriaei Bory, I. durieui Bory (2) I. histrix Bory subsp. sicula (Tod.) P. Fourn. I. histrix Bory subsp. Delalandei (Lloyd) P. Fourn.

0.25 cm

French names: (1) Isoète de Durieu, (2) Isoète épineux Italian names: (1) Calamaria di Durieu, (2) Calamaria istrice English name: (2) Land Quillwort

1 cm frond base

Subspecies

None

Isoetes duriei

Description/identification criteria96, 300

• Terrestrial perennial plants with a very short stem forming a kind of underground bulb bearing roots, and many fronds (“leaves”) arranged in rosettes at ground level. The fronds of Isoetes duriei are decurved towards the ground, while those of I. histrix are narrow and tough. • Bulb: usually firmly embedded, ringed with blackish spiny scales corresponding to the sclerified* remains of the bases of the preceding years’ fronds. • Fronds: dark green, 4 to 10 cm long, all fertile (sporophylls*), hollowed out at the base on the inner (upper) side to form a pit containing a sporangium. Outer fronds carrying macrosporangia, covered in a complete veil in I. duriei and a partial veil in I. histrix. Innermost fronds carrying microsporangia. • Macrosporangia containing macrospores (females) and microsporangia containing microspores (mâles).

0.35 mm megaspore

1 cm

Similar species

The two species are similar in appearance and in their ecology. One difference between them visible to the naked eye is the presence around the bulb of spiny scales, very obvious in Isoetes histrix, less conspicuous and briefly tridentate in I. duriei. Examination of the megaspores using a microscope or a good binocular lens allows the two species to be easily distinguished: Macrospores medium sized (0.4 to 0.6 mm) and with many tubercles in I. histrix. Macrospores larger (0.6 to 0.8 mm), with no tubercles and with a honeycombed surface in I. duriei.

0.25 cm frond base

Isoetes histrix rivers in the southern Cévennes); in Corsica, it is fairly common in the coastal zone and at low altitudes up to 300 m (but has been found exceptionally at 1,000 m). Other Mediterranean countries

Distribution - Ecology Distribution/range

Portugal, Spain (west of the country and Catalonia), Balearics (Minorca), Tyrrhenian area of Italy (from Liguria to Calabria), Sardinia, Sicily, Greece, Cyprus, Algeria, Morocco and Turkey.

(1) Isoetes duriei Western Mediterranean species (distribution map in Quézel306).

(2) Isoetes histrix Mediterranean-Atlantic species (distribution map in Quézel306).

France

France

On the mainland, it is common at Les Maures and Estérel, fairly rare elsewhere (Biot Massif, between Béziers and Montpellier, southern catchment of the Caroux, southern foothills of the Montagne Noire, Albères, the Roussillon plain, valleys of the Gardon

It is very well represented along the Atlantic (Brittany, PoitouCharentes, Pays de la Loire) and in Corsica, but local in the Mediterranean part of the mainland where it only occurs at Les Maures (Hyères region and the northern edge of the Plaine des Maures).

28

Plant species

Other Mediterranean countries

Phytosociology*

Central and western Iberian Peninsula (Spain, and all the Portuguese provinces), mainland of Italy, Sardinia, Sicily, Macedonia, Greece, Crete, Malta, Croatia, Morocco, Algeria, Tunisia, Middle East (Syria, Lebanon) and Turkey.

Owing to the difference in ecology between Isoetes velata (of periodically flooded habitats) on the one hand, and I. duriei and I. histrix (of non-flooded habitats) on the other, de Foucault98 split the Isoetion of Braun-Blanquet48 and Rivas-Goday306, 319 into two alliances: The Antinorio agrostideae-Isoetion velatae for topographically low-lying areas, periodically flooded. The Ophioglosso lusitanici-Isoetion histricis for topographically higher areas, not flooded. The Quillwort communities of non-flooded habitats are therefore included in the Ophioglosso lusitanici-Isoetion histricis alliance, order Isoetalia velatae and class Isoetea velatae98, 194.

Habitat General description

The most suitable habitats are short Mediterranean swards, wet or waterlogged in winter and the beginning of spring and very dry in summer. The substrate is usually siliceous (silts and sands) and relatively rich in organic matter. At Les Maures and the Estérel, Isoetes duriei also grows equally well in soils with little organic matter where, however, it grows to a smaller size than in accumulation depressions. These kinds of habitat, which are not (or are only exceptionally) flooded, are located on the edges of temporary pools and streams, on various flat areas and in wet clearings in the maquis48, 233, 296. The communities described based on I. histrix are a little less wet than those defined for I. duriei, and form a stage transitional to the annual swards of drier biotopes (with Tuberaria guttata and Anthoxanthum ovatum).

In the Isoetion s.l. (Isoetalia), the habitat of these two species corresponds to the following communities: (1) I. duriei • Isoetetum duriei association*, defined in Languedoc48 and also described from Spain319. • Association with I. duriei and Juncus capitatus described from Corsica233. • Association with I. duriei and Nasturtium asperum described at Les Maures21.

Companion species

(1) I. duriei • In Corsica: Ophioglossum lusitanicum, various Serapias, Aira capillaris, Bellis annua, Cicendia filiformis, Radiola linoides, Linum bienne, Anagallis arvensis subsp. parviflora, Lythrum hyssopifolia, Juncus bufonius, J. tenageja, J. capitatus, Ranunculus revelieri and Lotus conimbricensis. • In mainland France21, 22, 215: the same species, with in addition Isolepis cernua (=Scirpus savii), Nasturtium asperum, Agrostis pourretii, Airopsis globosa, Veronica acinifolia, Juncus pygmaeus, J. fasciculatus and Mentha pulegium. (2) I. histrix • Species common to the various sites: Radiola linoides, Linum bienne, Anagallis arvensis subsp. parviflora, Serapias lingua, Juncus bufonius, Isolepis cernua (= Scirpus savii), Bellis annua. • At the former sites of the littoral Maures, where the species has not recently been seen: Allium chamaemoly, Romulea columnae and Isoetes duriei22. • In Morocco: Poa annua, Filago (= Logfia) gallica, Rumex bucephalophorus and Bromus mollis275. Habitats Directive

• At the edges of pools (1) and (2): “Mediterranean Temporary Pools” (code 3170). • In the absence of disturbance, humid grassland vegetation often invades the short swards where Isoetes histrix occurs, and it is then included in the following habitat: “Mediterranean tall humid grasslands of Molinio-Holoschoenion” (code 6420). CORINE Biotopes

22.34 Amphibious southern communities (Isoetalia). 22.341 Short Mediterranean amphibious swards (Isoetion). 22.3411 Terrestrial Quillwort communities: formations with Isoetes histrix, I. duriei, of ephemeral aquatic habitats.

(2) I. histrix • In France, in the Serapion alliance, Aubert & Loisel 1971: subassociation with I. histrix of the Serapio-Oenanthetum, described by Barbéro22 from Les Maures. • In Spain, in the Isoetion alliance319: - Association Isoetetum histricis s. l. - Association Wahlembergio-Isoetetum histricis. • In Corsica, in the Ophioglosso lusitanici-Isoetion histricis (Br.-Bl. 1931) de Foucault 1988: community with I. duriei and I. histrix. • In North Africa, in the Isoetion s.l.: association with I. histrix and Radiola linoides, described by Chevassut & Quézel72.

Ecology Biological characteristics

Life form Perennial species: bulb-forming geophyte*292. Adaptive strategy163 Stress tolerant-ruderal (S-R). Reproduction Aerial for maturation of spores. Description of spores (macrospores: see above, § “Similar species”) • (1) granular microspores. • (2) microspores covered with fine entangled fibrils. Biological cycle • In France: growth of fronds with the onset of the first autumn rains21 and in winter when the substrate is very wet; ripening of spores in spring, followed by rapid withering of the fronds at the end of spring, when the habitat is drying out rapidly.

29

Mediterranean temporary pools

• In Morocco: the cycle of Isoetes histrix begins at the end of winter after the first rain and ends at the beginning or towards the middle of spring with the formation of spores.

Isoetes duriei France Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Dec.



and vegetative growth

Germination Maturation of spores Withering of fronds

Conservation - Management Assessment of populations

(1) I. duriei • France (mainland and Corsica): large number of sites (see above, § Distribution), but number of stations within the sites impossible to estimate accurately, given the very scattered nature of the species’ distribution. • Spain: not very widespread (provinces of Cadiz, Huelva, Gerona, Badajoz, and one station on Minorca64, 79). • Portugal: locally present in six provinces. - Italy: fairly rare (Sardinia, Sicily, Capraia, Liguria and west coast of the peninsula)137, 292.

Isoetes histrix France Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Dec.





Germination and

vegetative growth Maturation of spores Withering of fronds Morocco Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Dec.





Germination and vegetative growth

(2) I. histrix • France: large number of sites in Corsica, but only three remaining populations in the Mediterranean part of the mainland (Hyères, La Londe-les-Maures, Le Cannet-des-Maures). • Spain: four stations on Minorca, one on Majorca331, and present in 15 provinces in the west of the peninsula and three provinces in the northeast. • Portugal: in all provinces64, 79. • Italy: rare species (Sicily, Sardinia, and the Tuscany, Lazio and Puglia regions)292.

Maturation of spores Withering of fronds

Conservation status

(1) I. duriei The populations appear to be stable. Environmental conditions

Hydrology A very wet substrate is necessary at the beginning of the cycle but these plants do not like to be submerged for long. Substrate (1) not (or very slightly) oligotrophic* in Corsica, oligotrophic on the mainland (Permian sandstones at the Plaine des Maures, rhyolites at the Colle du Rouet). (2) more or less oligotrophic (usually sand-silt). Interspecific competition Low growing species, light-demanding (heliophilous*) and consequently not very tolerant of competition from taller plants. Impact of perturbations The main perturbations are the “ploughing” of the substrate by wild boars, and trampling (by cattle and people). They are beneficial as they limit the amount of cover and litter produced by taller plants. Temperature It appears to govern the distribution of Isoetes histrix at Les Maures, where it is confined to very warm coastal stations. It is usually replaced by I. duriei away from the coast, in the Maures plain and massif in particular22.

30

(2) I. histrix The populations appear to be stable in Corsica and Morocco as they are fairly resistant to anthropogenic and natural threats. They have declined in mainland France, in Roussillon (extinction due to the destruction of the Saint-Estève pool) as well as at Les Maures (not found recently in the Saint-Tropez region, or at the capes of Lardier, Benat, and Taillat, but recently rediscovered at Hyères, at La Londe-les-Maures and Le Cannet-des-Maures). In Malta, the species has not been seen again since its discovery at the end of the 19th century344. Legal status/level of protection IUCN 1997 Bern Convention Habitats Directive National and regional protection

National Red Data Books and Red Lists

France (1) and (2), national list: Decree of 20/01/1982/Journal Officiel 13/05/1982, amended by Decree of 31/08/1995/Journal Officiel 17/10/1995 - Balearics (2): “endangered” (EN)331 - Malta (2): “presumed extinct” (Ex ?)344

Plant species

Conservation problems/threats Anthropogenic factors

France In Corsica, development projects (such as construction works) are a potential threat to some coastal sites. In Provence, in Les Maures, urbanisation and tourist developments (golf courses) have destroyed a large number of Isoetes duriei stations*. Similarly, development along the coastline has destroyed all the coastal I. histrix stations (region of Saint-Tropez, Le Lavandou, Les Bormettes), and constitutes a threat to the other stations. Morocco Along with urban development, the greatest threat to the I. histrix populations is the infilling of pools with debris. Natural factors

• In the absence of disturbance, the closing-up of the habitat due to the spread of taller plants (Schoenus nigricans, Scirpus holoschoenus, Dittrichia viscosa and especially various maquis species) could constitute a threat for several sites in Corsica. This is equally true at Les Maures where the cessation of grazing is favouring the growth of perennial herbaceous species (Paspalum dilatatum, Juncus conglomeratus, Scirpus holoschoenus), and even maquis species. This is not the case at the Moroccan sites, where grazing is commonly practised. • The frequency of dry years does not affect the Isoetes histrix populations, which sustain themselves by their spores as much as by their bulbiform rhizome.

At Les Maures: • The clearing of firebreaks (provided that it is sensitive, i.e. that it does not upset the microtopography), and especially their grazing, are beneficial. • A Natura 2000* area, in the process of being established in the Plaine des Maures, provides the opportunity to take the requirements of management for the conservation of I. duriei into account, as does a Réserve Naturelle project that includes the majority of the stations. • The LIFE “Temporary Pools” project has enabled experimental testing of the management of plant communities with I. duriei to be carried out, as well as the purchase by the CEEP of land where this species grows. Recommendations

France In Corsica, retention of extensive cattle grazing at the sites where it is still present, and its reintroduction at sites where it is no longer practised. In mainland France: • Retention of horse grazing (one Isoetes histrix station) and pedestrian traffic (two I. histrix stations); avoidance of destructive activities (mountain biking,* scrambling, fly-tipping). • Reintroduction of extensive grazing, beneficial for the conservation of I. duriei, at Les Maures. Morocco Avoidance of repeated cultivation of the catchment areas nearest to I. histrix sites, as the colonisation of the habitat by arable weeds may have a negative effect (competition).

Risks relating to populations

(1) I. duriei No extinctions currently known in Corsica or on the mainland. (2) I. histrix No extinctions currently known in Corsica. On the mainland, the inherent risks to populations are low, but possible, due to the limited numbers and the very marked isolation of each of the populations, which are not very vigorous. Management measures Current measures

No management either in Corsica or on the mainland, or in Morocco, but grazing by free-ranging cattle, wild boar, and the movements of hunters and walkers are unintentional management activities which are beneficial to these Quillworts.

Bibliography Anonymous, 199912 ; Barbero, 196521, 196722 ; Bissardon & Guibal, 199739 ; Braun-Blanquet, 193548 ; Castroviejo, 1986200164 ; Chevassut & Quézel, 195672 ; Cirujano et al., 199279 ; Danton & Baffray 199596 ; Foggi & Grigioni, 1999137 ; Foucault de, 198898 ; Grime, 1979163 ; Julve, 1993194 ; Loisel, 1976215 ; Malcuit, 1962233 ; Olivier et al., 1995274 ; Ould Louleid, 1991275 ; Pignatti, 1982292 ; Poirion & Barbero, 1966295 ; Prelli, 2001300 ; Quézel et al., 1979308 ; Quézel, 1998306 ; Rivas Goday, 1970319 ; Saez & Rossello, 2001331 ; Schembri & Sultana, 1989344. Authors: Paradis G. & M. L. Pozzo di Borgo Collaborators: Catard A., H. Michaud, L. Rhazi, D. Titolet &

N. Yavercovski

31

Mediterranean temporary pools

Isoetes setacea Lam. (1) & Isoetes velata A. Braun (2) 0.25 mm 300

LYCOPODIOPHYTES ISOETACEAE

megaspore

Main synonyms

(1) I. delilei Rothm, (2) I. variabilis Le Grand 0.2 cm

French names: (1) Isoète sétacé, Isoète grêle,

(2) Isoète à voile, Isoète voilé Italian names: (1) Calamaria setacea, (2) Calamaria velata Moroccan name: (2) Lehyet-el-rebb English name: (1) and (2) Quillwort (genus name)

1 cm frond base

Isoetes setacea

Subspecies: (2) Isoetes velata A. Braun subsp. velata

I. velata A. Braun subsp. intermedia (Trabut) Maire & Weiller I. velata A. Braun subsp. tegulensis Batt. & Trabut I. velata A. Braun subsp. perralderiana I. velata A. Braun subsp. tenuissima (Boreau) O. Bolos & Vigo Description/identification criteria96, 299

• Perennial amphibious plants: aquatic at least in winter and early spring, then terrestrial. • Habit is an upright tuft when submerged. Stem very short, forming a kind of bulb which bears roots and many slender, fragile fronds. • Fronds (“leaves”) pale green, long, 10 to 40 cm (1), 5 to 15 cm (2), linear and arranged in a dense rosette. These fronds are fertile (sporophylls*): each has a depression at its base on the interior (upper) side, occupied by a large sporangium. The sporangia of I. setacea are bare, those of I. velata are partly or completely covered by a well-developed veil. • Outer fronds bear macrosporangia (bare or covered with a veil according to species), containing macrospores. The inner fronds bear microsporangia (bare or covered depending on the species), containing microspores, which are very abundant. • Main difference between the two species: a very obvious veil over the sporangia of I. velata, while this veil is absent in I. setacea.

0.25 mm megaspore

0.2 cm 1 cm

frond base

Isoetes velata France

Similar species

• The more terrestrial Quillworts (Isoetes histrix and I. duriei) during the dry phase have their bulbs surrounded by old, persistent sclerified* frond bases; these scales are absent in I. setacea and I. velata whose bulbs, at least in winter and spring, have a whitish appearance299. • The subspecies I. velata subsp tenuissima is endemic to central France, so there is no risk of confusion with subspecies velata.

Extremely rare plant, occurring in: • The Hérault: temporary pools in the Plateau de Roque-Haute and the Plaine de Béziers (Grand-Bois). • The Pyrénées-Orientales: Torremila pool and the Plateau de Rodès (Fenouillèdes). Note: Quézel’s map306 wrongly includes Corsica for I. setacea. Other Mediterranean countries

Distribution - Ecology

Spain (centre and west of the country and in Catalonia), Balearics (Minorca), Portugal (centre and south), and the Maghreb (Morocco, where it has recently been discovered).

Distribution/range

(1) Isoetes setacea Western Mediterranean species (distribution map in Quézel306).

(2) Isoetes velata Mediterranean-Atlantic species (distribution map in Quézel306). France

Rare on the mainland (Plaine des Maures and Estérel in the Var, Plateau de Rodès in the Pyrénées-Orientales) and fairly well represented in Corsica.

32

Plant species

Other Mediterranean countries

• Spain (in more than half of the country, mainly in the west and on the island of Minorca), Portugal (in almost the whole of the country), mainland Italy, Sicily, Sardinia (subspecies tegulensis)86 and Greece. • Maghreb (Morocco, Algeria, Tunisia, Libya). The subspecies intermedia and tegulensis occur in Tunisia, Algeria and Morocco, subspecies perralderiana in Algeria. The subspecies velata occurs in all the countries of the species’ Mediterranean range. Habitat General description

(1) I. setacea At Roque-Haute, I. setacea lives in pools hollowed out in the Quaternary basalt. (2) I. velata Habitats suitable for I. velata are flooded in winter and spring, and very thoroughly dried out in summer. Their substrate is thin, composed of compact rock (granite, schist, rhyolite) or of very fine sediment with a very low content of organic matter (sand, silt and clay). Most of the sites are on siliceous rock, but the temporary pool of Padulu (Bonifacio, Corsica), has a calcareous subsoil. The floodwater is oligotrophic* and its depth varies between sites and between years: approximately 10 to 60 cm. Companion species

(1) I. setacea Braun-Blanquet48 described an Isoetetum setacei for these pools, with the following companion species: Juncus pygmaeus, Myosotis sicula, Lythrum borysthenicum, Cicendia pusilla, Lotus angustissimus, Lythrum thymifolium and Marsilea strigosa.

Phytosociology*

(1) I. setacea • For France, Braun-Blanquet48 described the Isoetetum setacei (see above). • For Spain, Rivas Goday319 considered I. setacea to be a characteristic species of the Cicendion alliance (order Isoetalia). (2) I. velata Various communities and associations* with I. velata have been distinguished around the Mediterranean, for example: • In North Africa, associations with I. velata and Myosotis sicula, and the Eryngium barrelieri and I. velata association72, 298. • In the Colle du Rouet Massif (northwest of the Estérel), the I. velata and Crassula vaillantii association295. • In Corsica, Lorenzoni & Paradis219, 221 and Paradis et al.284 described some further I. velata communities. Note Owing to the wide ecological differences between, on the one hand, I. setacea and I. velata living in periodically flooded habitats, and on the other hand I. duriei and I. histrix living in nonflooded habitats, de Foucault98 split the Isoetion of Braun-Blanquet48, Malcuit (for I. velata)233 and Rivas-Goday319 into two alliances306, 308: • The Antinorio agrostideae-Isoetion velatae for low topographical levels, periodically flooded. • The Ophioglosso lusitanici-Isoetion histricis for higher topographical levels, not flooded. The I. setacea and I. velata communities are thus included in the Antinorio agrostideae-Isoetion velatae alliance, order Isoetalia velatae, class Isoetea velatae. Julve194 accepted this classification.

Ecology

(2) I. velata Species associated with Isoetes velata are varied, according to the depth of water (zonation) the time of year (phenophase) and also the substrate21, 22, 219, 221, 284, 302: • Aquatic plants: Tolypella glomerata (Charophytes), Callitriche truncata, Ranunculus peltatus s.l., R. ophiogossifolius, Myriophyllum alterniflorum, Illecebrum verticillatum, Pilularia minuta, Apium crassipes, Alopecurus bulbosus, Baldellia ranunculoides, Littorella uniflora, Glyceria fluitans, etc. • Other species, more or less associated with flooding: Crassula vaillantii, Lythrum borysthenicum, Juncus pygmaeus, Myosotis sicula, Laurentia michelii, Ranunculus sardous, Scirpus setaceus, Agrostis pourretii, Bellis annua, Elatine brochonii, Corrigiola littoralis, Hypericum tomentosum, etc. Habitats Directive

Biological characteristics

Life form • Perennial species. However, annual populations of Isoetes velata have been identified in the cupular pools of the Colle du Rouet Massif (Var); they have a very low level of reproductive success295. • (1) hydrophyte*292 and geophyte*48. • (2) hydrophyte292, geophyte48 and sometimes therophyte*295 Adaptive strategy (sensu Grime163) Stress tolerant-ruderal (S-R). Reproduction Maturation of spores: (1) aquatic (2) aerial.

“Mediterranean Temporary Pools” (code 3170). CORINE Biotopes

22.34 Southern amphibious communities (Isoetalia). 22.341 Short Mediterranean amphibious swards (Isoetion). 22.3412 Aquatic Mediterranean Quillwort swards: communities including Isoetes setacea or I. velata of water bodies of variable depth.

Description of spores The megaspores of the two species are fairly similar. Those of I. setacea (0.4 to 0.9 mm) have a few inconspicuous tubercles and those of I. velata (0.4 to 0.5 mm) have more or less numerous tubercles on their various sides. The microspores are echinate when mature.

33

Mediterranean temporary pools

Biological cycle Growth of fronds in the water during winter (from the first autumn rain on the mainland for Isoetes velata); maturation of spores in spring (1), or after the water has receded (2); rapid death of fronds at the beginning of summer, when the habitat is no longer flooded and dries out severely. Isoetes setacea Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Dec.





Germination

and vegetative growth Maturation of spores Withering of fronds

(2) I. velata France

• In Corsica, about twenty sites. • On the mainland, a single station in the Roussillon, with the remaining populations confined to the Var: - In the Plaine des Maures, three small recently discovered stations. - In the Plaine de Palayson, the Catchéou pool and a few temporary streams. - In the Estérel, about fifty small cupular pools* on the rhyolite escarpments of the Colle du Rouet (distributed between about fifteen stations), and the Barres de Roussiveau (a few stations). Morocco

Isoetes velata Jan.

Feb.

More than 50 stations for Isoetes velata.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Dec.

Spain and Portugal



Germination

and vegetative growth Maturation of spores Withering of fronds

The species is scattered throughout the Iberian Peninsula (apart from the northern and western provinces), and on the island of Minorca64. Italy

Environmental conditions

Very rare species (Sicily, Sardinia, Tuscany and Lazio regions).

Hydrology These species grow in wet soil and can tolerate water depths of up to 50 cm. The duration of the periods of flooding (six months or more) and drought (sometimes reduced to four or five months) varies depending on climatic conditions. Spores begin to germinate in autumn (November) in soil which is saturated with water or flooded. The hydrological conditions have a significant effect on the growth of Isoetes setacea.

Conservation status

Substrate Usually non-calcareous, oligotrophic and thin. Interspecific competition Light-demanding species (heliophilous) and therefore intolerant of competition. Impact of perturbations Favourable, by opening up of the habitat.

Conservation - Management Assessment of populations

(1) In mainland France a more or less slow decline of the populations of Isoetes setacea and of the Isoetion plant associations is taking place. In particular, the populations of the pools at Saint-Estève (Pyrénées-Orientales), Grammont (Hérault), and Redessan (Gard) have disappeared. (2) In Corsica as well as in Morocco, at sites with loose, thin, even overgrazed substrates, the populations of I. velata are either stable or expanding. On the French mainland the populations in the Roussillon (Saint-Estève) and the Gard (Redessan, Pazac) have decreased, while they appear to be stable at Les Maures and Estérel.

Legal status/level of protection IUCN 1997 Bern Convention Habitats Directive National and regional protection

(1) I. setacea France

I. setacea occurs in about 100 pools in the Roque-Haute Nature Reserve, in one pool in the Béziers plain, one at Torremila, several pools and ditches at Rodès. Spain and Portugal

It occurs in the centre and west, as well as in Catalonia and on the island of Minorca (13 provinces in Spain and six in Portugal)64.

34

National Red Data Books and Red Lists

- France (1) and (2), national list: Decree of 20/01/1982/Journal Officiel 13/05/1982, amended by Decree of 31/08/1995/ Journal Officiel 17/10/1995 - France, national Red Data Book: priority species274 - Italy, national Red Data Book: (2) “vulnerable” (VU)86 - Balearics: (2) “vulnerable” (VU)331 - Morocco: (1) “very rare” (RR), (2) “rare?” (R?)133, 181, 380, 381

Plant species

Conservation problems/threats Anthropogenic factors

(1) I. setacea At Roque-Haute, the discontinuation of sheep grazing, accumulation of organic matter and significant shading are having an effect on the growth of I. setacea (reduction in weight of bulbs and in number and length of fronds) and on its reproduction (reduction in number of macrosporangia, weight of macrospores, number and weight of microsporangia315) (vol. 1, box 46).

• Diagnosis of management problems and implementation of a protocol for monitoring the cupular pools at the Colle du Rouet as part of the LIFE “Temporary Pools” project. • Classification in the “ecological series” by the ONF* of the Catchéou pools (Plaine de Palayson), as a prelude to enabling the natural-heritage issues associated with these pools to be integrated into forestry management. • Establishment of Natura 2000* areas at the Var sites, providing the opportunity to include the protection of Quillwort in the conservation objectives for these sites. Recommendations

(2) I. velata • In France: the spread of agriculture has destroyed some I. velata stations in the Costière Nîmoise, as well as the principal station in the Pyrénées-Orientales (Saint-Estève pool). In Les Maures and Estérel, urbanisation and developments affecting the nature of the soil (plantations, golf, improvement of grazing in the maquis by overseeding and fertilisation) have caused a reduction in the number of pools capable of supporting this species; the Saint-Raphaël and Fréjus stations, among others, have disappeared. Some small artificial hill reservoirs are apparently becoming suitable for its colonisation. • In Morocco, the problems are mainly associated with infilling, extraction of sediment and drainage of pools. Natural factors

In mainland France and Corsica (but not in Morocco, where grazing still takes place) the closing up of the habitat and the accumulation of litter, resulting from the increase of tall plant species (Schoenus nigricans, Scirpus holoschoenus, Dittrichia viscosa and various maquis species), is deleterious to these species, as is infilling by eroded sediments (Plaine des Maures and Colle du Rouet). Risks relating to populations

(1) No short-term risk of extinction. (2) In Corsica, several small stations are threatened with disappearance as a result of the closing up of the habitat (pools of Muratellu, Arasu and Mura dell’Unda). The recently rediscovered Roussillon population is very vulnerable due to its small size. Management and conservation measures Current measures

(1) I. setacea The Roque-Haute site has legal “Réserve Naturelle” (Nature Reserve) status, which is not being implemented at present due to the impossibility of gaining access to the station (no control over usage of the site). The I. setacea sites have been the subject of a survey addressing water levels, the dynamics of woody vegetation and clonal* species315. (2) I. velata In France and Morocco, the opening up of the tallest vegetation through grazing by cattle (Tre Padule Nature Reserve, Capandula pools in Corsica, Plaine de Palayson on the mainland) and by sheep (Padulu site in Corsica) is beneficial. In France, in the Var, further measures are in progress:

(1) I. setacea • Ensuring that the pools are not kept permanently flooded. • Reintroduction of grazing in the pools with helophytes*, and in the pools with woody plants (after clearing). • Removal of litter from the pools to allow the spore stocks to germinate satisfactorily. • Evaluation of the dynamics of the vegetation following the introduction of grazing (and any modifications of management). (2) I. velata • In France: - In Corsica and on the mainland, maintenance of cattle (and sheep) grazing at the sites where it still takes place and its reintroduction at sites where it is no longer practised, while avoiding intensive practices (overseeding, fertilisation), and following clearing operations at sites which have been heavily colonised by woody vegetation (Corsica). - At the Colle du Rouet, implementation of monitoring at the cupular pools in the context of contractual management which is being put in place by the ONF, the commune and the CEEP, and, in the case of pools which are silting up, planning for minor intervention (removal of litter). • In Morocco: Discouraging any modification of the hydrology of the pools (drainage, infilling) as well as extraction of sediment, which destroys the seedbank.

Bibliography Anonymous, 199912 ; Barbero, 196521, 196722 ; Bissardon & Guibal, 199739 ; Braun-Blanquet, 193548 ; Castroviejo, 1986200164 ; Chevassut & Quézel, 195672 ; Conti et al., 199286 ; Danton & Baffray, 199596 ; Fennane & Ibn Tattou, 1998133 ; Foucault de, 198898 ; Gaudillat & Haury, 2002153 ; Grillas & Tan Ham, 1998162 ; Grime, 1979163 ; Jahandiez & Maire, 1931-1934181 ; Julve, 1993194 ; Lorenzoni & Paradis, 1997219, 2000221 ; Malcuit, 1962233 ; Médail et al., 1998246 ; Molina, 1998251 ; Olivier et al., 1995274 ; Paradis et al., 2002284 ; Pignatti, 1982292 ; Poirion & Barbero, 1965295 ; Pottier-Alapetite, 1952298 ; Prelli, 2001300 ; Quézel & Zevaco, 1964302 ; Quézel et al., 1979308 ; Quézel, 1998306 ; Rhazi et al., sous presse315 ; Rivas Goday, 1970319 ; Saez & Rossello, 2001331 ; Titolet & Oualidi, 2000380 ; Titolet & Rhazi, 1999381. Authors: Paradis G. & M. L. Pozzo di Borgo Collaborators: Catard A., H. Michaud, L. Rhazi, M. Rhazi &

N. Yavercovski

35

Mediterranean temporary pools

Littorella uniflora

(L.) Ascherson

ANGIOSPERMS PLANTAGINACEAE Main synonyms

L. lacustris L. French name: Littorelle à une fleur Italian name: Littorella English name: One-Flowered Shoreweed

1 cm

Subspecies

None

Littorella uniflora

Description/identification criteria63, 292

Perennial herbaceous plant, hydrophytic* or amphibious, with two morphological types: a sterile submerged (aquatic) form and a terrestrial flowering form. Submerged form

It has dense rosettes of leaves and stolons. It does not flower. Each rosette consists of: • A vertical rhizome, very short, very flattened (4-5 mm thick), covered by the remains of dead leaves. • Three to 14 leaves, light green, thick, cylindrical, stiff, glabrous, broadly elliptical in cross section, swollen at the base and finely pointed at the tip, with air vessels, and measuring 3 to 15 cm long and 2 to 4 mm thick. These submerged leaves have no stomata. • Many adventitious roots on the rhizome. From the leaf axils arise stolons, 3 to 60 cm long, bearing leaves that are reduced to scales. At their tips, the stolons form new rosettes of leaves and take root in the soil by forming adventitious roots. Hence the stolons enable the plant to propagate itself vegetatively to a considerable degree under the water. Terrestrial form

This results form the transformation of the aquatic form after the water has dropped. It takes the form of dense rosettes of leaves but does not produce stolons. It produces flowers (the species is monoecious). Each rosette includes: • A very short vertical rhizome with adventitious roots. • Leaves many (5 to 25), small (2.5 to 4-10 cm long) and with many stomata321. The upper surface, especially of the internal leaves, has a longitudinal groove, while the lower surface is semi-cylindrical. The male flowers are borne at the tips of fairly long peduncles with the female flowers at their bases. The male flowers are small (sepals 4 mm long and stamens with filaments 10-20 mm), the scarcely visible female flowers are protected by foliar sheaths and surrounded by long woolly hairs. The flowering of each plant is usually rapid and concentrated over one to two weeks321. Flowering follows the emergence of the substrate from the water. Fruit dry, indehiscent, with one seed (achene), 2 mm in length.

36

Similar species

Littorella is subject to confusion with: • Young stages of Baldellia ranunculoides. However, this species fairly quickly develops lanceolate leaves and has the scent of coriander when crushed. • The well-advanced stages of some Quillworts (such as Isoetes velata), but their lack of stolons enables them to be distinguished.

Distribution - Ecology Distribution/range63, 292

Littorella uniflora is a west European and sub-Atlantic species, absent from the areas around the Mediterranean apart from Corsica, Sardinia, central and southern Iberian Peninsula (Medina, pers. com.) and Morocco. It extends from latitude 38°45’ in the south (at Lisbon) to 68°20’ in the north and, in longitude, from the Azores (31°W) to Karelia (35° E). Over this very wide range, its distribution is discontinuous. France

A species of the Atlantic seaboard, but also occurs in Corsica. One locality, not recently confirmed, in the Crau (Etang du Luquier, Bouches-du-Rhône). Other Mediterranean countries

Morocco in the Rif region, Sardinia (where it is extremely rare and reaches the southern limit of its European range), northern Italian provinces (in a non-Mediterranean climate), Spain and Portugal. Habitat General description

1. Commonest habitats in Europe (non-Mediterranean) Littorella uniflora occurs on the edges of oligo-mesotrophic* water bodies: • Lakes and pools in peaty areas, fish ponds, sand and gravel pits, ditches. • Periodically flooded shallow pools on various substrates (silt, sand, gravel and even stones). The optimum living conditions are provided by water bodies that dry out in summer and have a maximum depth of 1.5 m in winter.

Plant species

2. Mediterranean temporary pools In southern Corsica, Littorella uniflora currently occurs in six temporary pools: four in the Tre Padule de Suartone Nature Reserve and two pools, of artificial origin, on the Frasselli plateau. It is associated with the following species: Isoetes velata, Pilularia minuta, Baldellia ranunculoides and Ranunculus ophioglossifolius.

Corsica Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Dec.

July

Aug.

Sept.

Oct.

Nov.

Dec.

Germination and development of stolons Transformation aquatic
terrestrial form Flowering Fruiting

Habitats Directive

In the Mediterranean region: “Mediterranean Temporary Pools” (code 3170).

Morocco Jan.

Feb.

March

April

May

June

Germination and vegetative growth

CORINE Biotopes

22.34 Southern amphibious communities (Isoetalia). 22.341 Short Mediterranean amphibious swards (Isoetion). 22.3412 Aquatic Mediterranean Quillwort swards: communities containing Isoetes velata in water bodies of variable depth (where I. velata has a much earlier phenology than Littorella uniflora). Phytosociology*

Littorella uniflora is a companion species in the Isoetes velata communities (Corsica). Elsewhere it is characteristic of formations of the class Littorelletea in Spain79 and in northwest Europe where it is also a component of submerged plant associations* of the class Potamogetonetea pectinati 47, 82,127.

Flowering Fruiting

Environmental conditions

Hydrology Winter flooding is essential for germination and for the activation of the buds on the short rhizomes to produce stolons. Emergence from the water is essential for flowering117, 127. Substrate Substrate acid, loose, sometimes pisolithic, fairly thin (less than 10 cm deep), of variable particle size (silt, sand, gravel), without (or with little) organic debris and oligo-mesotrophic.

Ecology Biological characteristics

Life form Perennial species: hemicryptophytic* hydrophyte*. Reproduction Aerial. Flowering In Corsica from mid-May to the end of June, and in April-May in Morocco. Adaptive strategy (sensu Grime163) Stress-tolerant (S). Biological cycle At the beginning of spring, under water 10 to 30 cm deep: • The achenes germinate and give rise to new individuals which send out stolons that produce rosettes (submerged form). • The rhizomes break dormancy and produce short stolons giving rise to new rosettes (submerged form). At the end of spring, when the pool has dried out or in very shallow water (less than 10 cm): • Transformation of rosettes from the submerged form to the terrestrial form (beginning of May). • Flowering of plants that have emerged or are under very shallow water (end of May and June). In summer: • Fruiting (June and beginning of July). • Transition of achenes and short rhizomes into the dormant state.

Interspecific competition A heliophilous* plant, intolerant of cover from tall species (Eleocharis palustris, Phragmites australis, various Scirpus). Impact of perturbations Low-intensity disturbance (grazing, trampling) that causes a small amount of substrate compaction is beneficial127. Drainage, by altering the hydrology of the pools, may have an effect on the germination of the achenes and sexual reproductive success. Light Light-demanding (heliophilous plant), which could explain its absence from turbid pools in Morocco.

Conservation - Management Assessment of populations

• France: - In Corsica, only 6 sites (see above). - In non-Mediterranean France: the species has a scattered distribution in the western, central, eastern and extreme northern regions of the country96. • Italy: several scattered sites in the northern regions (Piedmont, Lombardy, Venetia, Emilia-Romagna); a single site in Sardinia292. • Spain and Portugal: about a hundred sites scattered through the northwestern half of the Iberian Peninsula (Medina, pers. com.). • Morocco: the species has been recorded only in the sub-humid to humid mountainous region of the Rif (Issaguene dayet).

37

Mediterranean temporary pools

Conservation status

• In Mediterranean France: in Corsica the populations fluctuate from one year to another, but appear to be stable in the medium term at the Tre Padule pools. There is a lack of data from the Frasselli pools. The four plants recorded by Lorenzoni225 at the Mura dell’Unda pool (north of Porto-Vecchio) have not been found again (looked for in 2001, 2002 and 2003). Gamisans145 reported its occurrence at the Lac de Ninu (1,800 m altitude), but the species could not be found there during a recent study of the lake. In the Crau, where the species was collected in 1954 at the Etang du Luquier257, it has not been seen again and could have disappeared (changes in water quality, invasion of the banks by Paspalum distichum). • In non-Mediterranean France, it is considered to be scattered and vulnerable96. • In Italy, the species is decreasing in the north of the country86. • In Morocco, the populations appear to be stable in the medium term, despite wide interannual fluctuations (linked with variable rainfall). Legal status of the species/level of protection IUCN 1997 Bern Convention Habitats Directive National and regional protection

National Red Data Books and Red Lists

France, national list: Decree of 20/01/1982/Journal Officiel 13/05/1982, amended by Decree of 31/08/1995/Journal Officiel 17/10/1995 - Morocco: “very rare” (RR)133, 230 - Italy, national Red Data Book: “vulnerable”86, 181 - Spain, regional lists (Medina, pers. com.)

Conservation problems/threats Anthropogenic factors

In Corsica, up until now, these kinds of threats appear to be absent.

38

In Morocco and Spain, drainage is the most important threat, as well as overgrazing in Morocco. In northern Italy, increasing eutrophication* of the water is resulting in the decline of the species86, 292. Natural factors

In Corsica, the thorough drying out at the end of spring and in summer and the shallowness and nutrient-poor status of the loose substrate, as well as extensive cattle grazing at the six pools with L. uniflora, are sufficient to prevent the spread of more competitive species (such as Eleocharis palustris). Management and conservation measures Current measures

Corsica: the Tre Padule site is protected by its legal “Réserve Naturelle” status. Morocco: no management measures. Recommendations

Corsica: continuation of extensive cattle grazing. Morocco: work towards limiting the numbers of livestock.

Bibliography Anonymous, 199912 ; Bissardon & Guibal, 199739 ; Bournérias et al., 200147 ; Casper & Krausch, 198163 ; Cirujano et al., 199279 ; Clément & Touffet, 198382 ; Conti et al., 199286 ; Danton & Baffray, 199596 ; Dierssen, 1983117 ; Duvigneaud, 1971127 ; Fennane & Ibn Tattou, 1998133 ; Gamisans, 1988145 ; Grime, 1979163 ; Jahandiez & Maire, 1931-1934181 ; Lorenzoni & Paradis, 2000221 ; Lorenzoni, 1997225 ; Maire, 1952-1987230 ; Molinier, 1981257 ; Pignatti, 1982292 ; Robe & Griffiths, 1998321. Authors: Paradis G. & M. L. Pozzo di Borgo Collaborators: Grillas P., H. Michaud, L. Rhazi & N. Yavercovski

Plant species

Lythrum borysthenicum (Schrank) Litv. (1) & Lythrum tribracteatum Salzm ex Sprengel (2) ANGIOSPERMS LYTHRACEAE Main synonyms

(1) Peplis erecta Moris, P. hispidula Durieu, P. nummulariaefolia Jord., L. biflorum (DC.) J. Gay. (2) L. Salzmannii Jord., L. dibracteatum Guss., L. hyssopifolium L. subsp. salzmanni (Jordan) Bonnier L. hyssopifolia L. var. pseudo-bibracteatum Tallon French names: (1) Péplis dressé; (2) Salicaire à trois bractées Spanish name: (2) Alheli silvestre177 Italian names: (1) Salcerella a foglie ovali; (2) Salcerella con due

brattee English names: (1) Loosestrife (genus name); (2) Threebract

Loosestrife Subspecies

1.5 mm

None Description/identification criteria (1) Lythrum borysthenicum

calyx 1 cm

• Leaves sessile or nearly so, ciliate when young, rounded oval, opposite, the upper leaves in a spiral. • Stem ciliate and fairly rough at the tip, erect, from 5 to 15 cm in height. • Flowers solitary, subsessile in the leaf axils. • Calyx expanded at the base, in a short tube, almost as wide as long, without prominent ribs and longer than the capsule, with five or six short teeth, blunt, equal or almost equal. • Petals crimson, five or six in number, or absent. • Stamens six, enclosed in each flower.

Lythrum borysthenicum

(2) Lythrum tribracteatum • Green plant, glabrous, low (5 to 30 cm) more or less prostrate. • Leaves elliptical, small, entire, linear to oblong (0.3-1.5 cm long by 0.1 - 0.3 cm wide), sessile, alternate. • Flowers small, solitary in the axils of the branch leaves, subsessile, crimson; five or six petals, equalling approximately half the length of the calyx tube. • Calyx narrowed at the base, very elongated into a tube, much longer than wide (0.5 - 0.6 cm in length), with prominent ribs and 8 to 12 very short, triangular, unequal, blunt teeth, and two linear bracts, green, equal in length to the calyx and resembling leaves, or alternatively very short and scarious. • Stamens five to six. Style enclosed. • Fruit: cylindrical capsule* approximately equal to the calyx. Seeds yellow, slightly constricted. 1.25 mm Similar species

(1) Lythrum borysthenicum may be confused with L. (Peplis) portula, but this species has its leaves clearly petiolate and spatulate, a prostrate stem, petals liliaceous, and calyx shorter than the capsule. (2) L. tribracteatum may be confused with L. thymifolia and L. hyssopifolia, but in both these species, the colouration is glaucous, the habit is upright, the external calyx teeth are longer than the internal, the calyx is tubular and the stem unbranched or very little branched.

calyx

1 cm

Lythrum tribacteatum 39

Mediterranean temporary pools

Distribution - Ecology

Habitats Directive

“Mediterranean Temporary Pools” (code 3170). Distribution/range CORINE Biotopes

(1) Lythrum borysthenicum Sub-Mediterranean species. France

Provence-Alpes-Côte-d’Azur (Plaine des Maures, Colle du Rouet, Bois de Palayson, Hyères islands, Plateau d’Evenos), LanguedocRoussillon (Roque-Haute, Béziers Quaternary plain, Plateau des Fenouillèdes , alluvial terraces of the Agly), and Corsica. Extension beyond the Mediterranean zone into west-central France as far as the Loire valley and the Sarthe. Previously recorded in the Dombes and surrounding area, but the data are however, highly dubious266 (R. Dupré, CBNBP, pers. com.). Other Mediterranean countries

Portugal: absent from the north of the country. Spain: western half, to Gerona and Valencia; absent from a large area of Galicia and Asturias64. Italy: Tuscany, Lazio, Sardinia, Sicily292. Morocco: Atlantic plains and mountains of low to medium altitude, in a sub-humid, semi-arid Mediterranean bioclimate*. Also present in Greece, Cyprus, Turkey, Algeria, Tunisia, Libya, Israel. (2) Lythrum tribracteatum Mediterranean species. France

Atlantic region: départements of Loire-Atlantique, Vendée, Charente-Maritime, Aveyron. Mediterranean region: • Languedoc-Roussillon region: lower floodplain of the Aude, plateaux of Vendres, Roque-Haute and Caux-Fontès-Pézenas, Agde, Littoral ponds of Languedoc, Garrigue pools of the Montpellierais, Etang de Capelle and Costière nîmoise, PetiteCamargue. • Provence-Alpes-Côte-d’Azur Region: Lanau pool in the Crau, pools of Cerisières in the Camargue, Plateau des Quatre-Termes, Plateau du Cengle and Centre Var.

22.34 Southern amphibious communities (Isoetalia) 22.341 Short Mediterranean amphibious swards (Isoetion) (France): (1) and (2) 22.343 Halo-nitrophilous Mediterranean amphibious swards (Heleochloion) (Spain): (2) 22.32 Mediterranean annual amphibious swards (Nanocyperetalia) (France and Spain): (2) Phytosociology*

Morocco Both species belong to the Isoetion (velatae). Juncus pygmaeus, J. bufonius, J. capitatus, Lythrum thymifolia, L. hyssopifolia, Isoetes velata, etc. are associated with them. France continentale (1) In the Var (Plaine des Maures, Plaine de Palayson), L. borysthenicum is characteristic, along with Ranunculus revelieri, of an Isoetion association which is endemic* to this region. In LanguedocRoussillon, it occurs in Isoetes setacea formations (Isoetetum setacei Br.-Bl.) (2) L. tribracteatum is often associated with Damasonium polyspermum as a characteristic species of the Lythrion tribracteati alliance (order Nanocyperetalia flavescentis)153, 215. It also occurs in rice fields in the Camargue, and fairly often together with Crypsis on the edges of ponds in Languedoc. It is also characteristic of the Elatinetum macropodae Br.-Bl. (1931), formerly described from Agde. On the central Atlantic coast, it is characteristic of a Junco hybridi-Lythretum tribracteati which occurs in sub-halophilic362 marshes. Corsica L. borysthenicum is associated with Isoetes velata and Juncus pygmaeus in the Tre Padule de Suartone pools221, with Elatine brochonii and Juncus pygmaeus in the Padulellu pool219, with L. hyssopifolia and Cotula coronopifolia in the coastal temporary pool at Tour d’Olmeto Point282.

Other Mediterranean countries

Portugal, Spain, mainland Italy (in all provinces except Marche, Abruzzo and Basilicata), Sardinia, Sicily, Albania, Greece, Morocco, Algeria, Tunisia, Libya, Turkey, Syria, Lebanon and Egypt.

Spain319 L. tribracteatum is characteristic of the Lythrion tribracteati alliance, as well as the Lythro-Heleochloetum schoenoidis association Rivas Martinez 1966, within the Heleochloion alliance.

Habitat Ecology General description

(1) L. borysthenicum: temporary pools on a non-calcareous substrate (sandstone, granite, quartzite, schist, basalt, etc.), flooded in winter and very dry in summer, with oligotrophic water, but also edges of marshes and pools, or slow-flowing shallow rivulets (in Les Maures). (2) L. tribracteatum: temporary pools in plains, low and medium altitude mountains in sub-humid and humid Mediterranean bioclimates. In western France, L. tribracteatum also occurs in brackish grasslands near the coast in depressions with bare wet soil, and places that are trampled by livestock96.

40

Biological characteristics

Life form Therophyte (annual species); scapigerous*. Reproduction Aerial. Biological cycle In Morocco and Corsica, Lythrum borysthenicum and L. tribracteatum begin to germinate in February if the site is flooded; flowering

Plant species

lasts from the end of April to July for L. borysthenicum and from the beginning of May to July for L. tribracteatum. Fruiting begins at the end of May for L. borysthenicum and mid-June for L. tribracteatum, accompanied by seed dispersal a short distance from the parent plant. In mainland France, flowering is slightly later, taking place in JuneJuly for L. borysthenicum and in May-June for L. tribracteatum. In Corsica, L. borysthenicum has the following cycle: germination in February and March, growth in March and April, flowering and fruiting in May and the beginning of June, death of the plant when the habitat has dried out, in June. Lythrum borysthenicum Mainland France Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Impact of perturbations It is favourable when it restricts competing plants.

Conservation - Management Assessment of populations Morocco

Both species are fairly common. They occur in the northern Atlantic plains (dayas* southwest of Tiflet, dayas of Mamora, Benslimane, Loukkos, etc.), the central plains (Haouz Region) and in mountainous regions (Middle Atlas, High Atlas, Rif, Tangier).

Dec.

France Germination and vegetative growth

Germination and vegetative growth

Flowering Fruiting and seed dispersal Morocco Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Dec.

L. borysthenicum is very rare in Languedoc-Roussillon, commoner but local in the Provence-Alpes-Côte-d’Azur Region (Var), fairly common at the Corsican pools. L. tribracteatum is rare throughout the country, including the Mediterranean region (local in Languedoc, very rare in the PACA Region and absent from Corsica).

Germination and vegetative growth

Algeria

Flowering

L. borysthenicum is a rare species, while L. tribracteatum is fairly common303.

Fruiting and seed dispersal

Lythrum tribracteatum Spain Mainland France Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Dec..

Sept.

Oct.

Nov.

Dec.

Germination and vegetative growth Flowering

These species are not considered to be rare. L. borysthenicum is common in the western half of the Iberian Peninsula, and L. tribracteatum has a wide range over the whole peninsula (27 provinces in Spain, six in Portugal)64.

Fruiting and seed dispersal Morocco Jan.

Feb.

March

April

May

June

July

Aug.

Germination and vegetative growth

Conservation status Morocco

Flowering Fruiting and seed dispersal

Environmental conditions

Hydrology These two amphibious pool species are often situated in more hydromorphic (low-lying) positions than L. thymifolium and L. thesioides. The germination of the seeds requires the soil to be saturated with water (flooding is not essential). At the vegetative stage the plant can tolerate shallow water (<4-9 cm) but it cannot survive for long under greater depths of water. Substrate (1) L. borysthenicum grows on usually acid substrates (siliceous soils). (2) L. tribracteatum occurs on siliceous or calcareous substrates (sand, sand-silt, silt or clay-silt soils). In mainland France, the soil is often silt-clay, rich in active limestone* and sometimes salty (Camargue, coastal pools of Languedoc, west-central France153, 362. Interspecific competition Major growth of tall plants may have negative effects on these heliophilous* species as a result of shading. This is the case in mainland France where grazing has been completely discontinued at the majority of the sites.

The populations are fairly stable but there is the risk of a decline in the short or medium term as a result of increasing threats. France

(1) Lythrum borysthenicum • In the Mediterranean region of the French mainland, L. borysthenicum has decreased more than L. tribracteatum. Its status is precarious in Languedoc-Roussillon, where a number of populations have already disappeared (Petite Camargue, Costière Nîmoise, Grammont, Agde, Lamoure, Saint-Estève pool), while others are in jeopardy (Fenouillèdes, plateaux of Vendres and Torremila). In the siliceous parts of Provence the species is surviving well; however, it has disappeared from the Giens peninsula. • In Corsica, at one of the temporary pools (Chevanu pool), used as a car park from May to September, a very severe decline in the population of L. borysthenicum has been recorded since 1991, undoubtedly due to the crushing of plants just beginning to fruit. • L. borysthenicum has recently been rediscovered at the Lac de Grand-Lieu (Loire-Atlantique)67. (2) Lythrum tribracteatum • L. tribracteatum is surviving well in the coastal Languedoc and in the lower floodplain of the Aude. It has disappeared from the edges of the Etang de Berre, but about ten localities have recently been discovered in the Var, which means that its status is less precarious than is indicated in the Red Data Book274. Some localities are, however, threatened by urban development (Agde) or the restructuration of wetlands (Vendres). The very small station

41

Mediterranean temporary pools

in the Hautes-Alpes is vulnerable. In the Aveyron, this Lythrum has recently been observed as a adventive plant in the valley of the Tarn, but the other stations in this département have not recently been seen. • In the Atlantic area, the populations of L. tribracteatum, formerly fairly numerous, appear currently to be reduced to a small number of stations*274, 362. Spain and Portugal

L. borysthenicum may be increasing due to its ability to colonise artificial new habitats (Medina, pers. com.). Legal status/level of protection IUCN 1997 Bern Convention Habitats Directive National and regional protection

National Red Data Books and Red Lists

France: (1) regional list for LanguedocRoussillon: Decree of 29/10/1997/ Journal Officiel 16/01/1998 (2) national list: Decree of 20/01/1982/Journal Officiel 13/05/1982, amended by Decree of 31/081995/Journal Officiel 17/10/1995 - (2) France, national Red Data Book: priority species274

France Lythrum tribracteatum benefits from the legally protected status of some sites (Nature Reserves of Roque-Haute in the Hérault and Saint-Denis-du-Payré in western France), and L. borysthenicum from that of the Tre Padule de Suartone Nature Reserve (southern Corsica). L. borysthenicum also benefits at the Plaine des Maures (Var) from experimental management measures, in the context of the LIFE “Temporary Pools” Project. L. tribracteatum is the subject of annual monitoring at the Lanau pool (Bouches-du-Rhône), and its seeds are preserved ex-situ by the Conservatoire Botanique National Méditerranéen de Porquerolles. Finally, several sites (Roque-Haute, Les Maures plain and massif, Notre-Dame de l’Agenouillade at Agde, etc.) are included in Natura 2000* areas. Recommendations

• Promoting a more accurate census and a survey of the stations. • Avoiding any alterations to the hydrology of the pools (drainage and infilling). • Gaining control over urbanisation and site management. • Keeping the habitat open to a certain extent by grazing or by maintaining the plant cover via clearing. • Implementing legal protection measures for the species and the sites. • Raising the level of awareness among the managers involved.

Bibliography Conservation problems/threats Anthropogenic factors

The discontinuation of traditional stock rearing, conversion to arable land, urbanisation, infilling with debris, the use of pools that are dry or drying out (e.g. as car parks), and drainage, are the causes of the declines and the vulnerability of the populations of both these species. Natural factors

In Morocco, at sites with a semi-arid bioclimate, a high frequency of dry years, which prevent the renewal of the seed stocks, is a potential threat for both these taxa. In France, the main threat for both species comes mainly from the closing-up of the habitat due to the spread of woody plants. Risks relating to populations

The fairly small sizes of the currently existing populations render them vulnerable. Management and conservation measures Current measures

Morocco There are no management measures.

42

Anonymous, 199912 ; Aubert & Loisel, 197118 ; Barbero, 196521 ; Castroviejo, 1986-200164 ; Chagneau, 200367 ; Danton & Baffray, 199596 ; Fennane & Ibn Tattou, 1998133 ; Fennane et al., 1999134 ; Fournier, 1961139 ; Gaudillat & Haury, 2002153 ; Loisel, 1976215 ; Lorenzoni & Paradis, 1997219, 2000221 ; Lorenzoni et al., 1994226 ; Maire, 1952-1987230 ; Médail et al. 1998246 ; Molina, 1998251 ; Nétien, 1993266 ; Olivier et al., 1995274 ; Paradis et al., 1999282, 2002284 ; Pignatti, 1982292 ; Quézel & Santa, 1962-1963303 ; Rhazi et al., 2001314 ; Rivas-Goday, 1970318 ; Terrisse, 1996362 ; Valdés et al., 2002391 ; Walter & Gilett, 1998399. Authors: Michaud H. & L. Rhazi Collaborators: Paradis G. & M. L. Pozzo di Borgo

Plant species

Lythrum thymifolium L. (1) & Lythrum thesioides M. Bieb. (2) ANGIOSPERMS LYTHRACEAE Main synonyms

(1) L. hyssopifolia L. subsp. thymifolium (L.) Bonnier & Layens L. thymifolia L. (2) L. geminiflorum Bertol. L. thesioides M. Bieb. subsp. geminiflorum (Bertol.) Rouy & E.G. Camus L. purpurascens Châtenier 1 mm

French names: (1) Lythrum à feuilles de thym, Salicaire à feuilles

de thym; (2) Lythrum faux-Thésium, Salicaire faux thésion Spanish names: (1) Salicaria-menor (catalan) Portuguese names: (1) Salicaria-menor Italian names: (1) Salcerella con foglie di timo; (2) Salcerella a

1 cm

calyx

fiori appaiati English names: (1) Thymeleaf Loosestrife; (2) Loosestrife (name

Lythrum thymifolium

of genus) Subspecies

None Description/identification criteria96, 139, 292, 386, 394

(1) Lythrum thymifolium • Annual plant, glaucous, more or less scabrid*, from 3-10 cm in height, with stem glabrous, erect, spindly, very leafy and not very much branched. • Leaves linear, sessileµ, alternate, small (0.5-0.9 cm long x 0.10.2 cm wide), narrow, close together, very finely toothed on the edges. • Flowers very small, solitary in the axils of the leaves on the branches, with short pedicels and with two linear bracts at the base of the calyx that are variable in size from one individual to another, sometimes within a single population234, 386. • Calyx elongated, tubular, short (0.20-0.25 cm long), bearing eight very unequal teeth at the tip: four long (0.5-1 mm) and papillose, alternating with four very short and membranous. • Corolla with four pink petals scarcely extending beyond the calyx teeth. • Two to four stamens per flower. • Fruit: capsule*, equal to or less than the length of the calyx tube. (2) Lythrum thesioides • Very similar, though a little taller (up to about forty centimetres). • Flowers very small, grouped in pairs or threes in the axils of all the leaves (solitary in L. thymifolium). • Calyx short, bell-shaped, with 8-12 teeth, and corolla rosepurple, with 4-6 petals with darker central veining (in L. thymifolium the calyx is cylindrical with eight teeth, and the corolla has four uniformly pink petals).

1 cm 1.5 mm calyx

Lythrum thesioides Similar species

• Lythrum tribracteatum is usually more or less prostrate, with very branching stems, leaves often elliptical, flowers with a narrow tubular calyx with 10-12 not very obvious very short equal triangular teeth and a purple corolla with 5-6 petals. This is a rare species, more hygrophytic* than L. thymifolium, and is found in oligotrophic * to eutrophic* habitats. • L. hyssopifolium L. is a common species in all sorts of humid to flood-prone habitats. It is usually robust, not very glaucous, with lanceolate leaves tapering at the base, and flowers usually

43

Mediterranean temporary pools

hexamerous* and pedicellate. The six petals clearly extend beyond the calyx, which has 12 unequal teeth at the tip (six 0.51 mm long, not very papillose, alternating with six short and membranous). The flower has six stamens. Ambiguous forms, intermediate between L. hyssopifolium and L. thymifolium, have (rarely) been found, and their determination is always difficult. • Frankenia pulverulenta is always a plant of saline habitats, on the coast or inland. It has a prostrate habit, a stem that is slightly woody at the base and very branching, petals purplish pink with serrated elongated limbs, and narrow spatulate leaves slightly curled at the edges.

Distribution - Ecology Distribution/range

(1) L. thymifolium Mediterranean species. France274 Départements of Allier, Puy-de-Dôme, Haute-Loire, Ardèche, Drôme, Hautes-Alpes, Pyrénées-Orientales, Aude, Hérault, Gard, Bouches-du-Rhône, Var and Alpes-Maritimes. Other Mediterranean countries 386, 394 Portugal, Spain, Italy, Greece, Algeria and Morocco.

Habitats Directive

“Mediterranean Temporary Pools” (code 3170). CORINE Biotopes

(1) 22.34 Southern amphibious communities (Isoetalia) 22.341 Short Mediterranean amphibious swards (Isoetion) (1) and (2): 22.32 Annual Mediterranean amphibious swards (Nanocyperetalia) Phytosociology* and companion species

(1) Lythrum thymifolium It belongs to the class Isoeto-Nanojuncetea and to the Cicendion filiformis and Isoetion s.l. alliances274. • In Languedoc: Isoetetum duriaei Braun-Blanquet. • In the Provence-Alpes-Côte-d’Azur (PACA) Region: - Spirantho-Anagallidetum tenellae Aubert & Loisel. - Isoeto duriaei-Nasturtietum (=Sisymbrelletum) asperum Barbero. There are a large number of associated species: Juncus pygmaeus, J. capitatus (especially in Languedoc), J. bufonius, L. borysthenicum, L. hyssopifolia, Isoetes velata, I. duriei (ecologically more similar, in France, to L. thymifolium than to I. velata), Laurentia michellii. In the Crau (Lanau pool), the species is associated with species of the Lythrion tribracteati alliance.

Outside the Mediterranean region Three old records from the Caucasus, lower valleys of the Volga and the Don.

(2) Lythrum thesioides Very few facts are known about the habitat of this species, which has been observed as much in cultivated land in flood-prone areas as on the shores of temporary ponds. At Jonquières-Saint-Vincent192 as well as at La Capelle-et-Masmolène249, L. thesioides was abundantly accompanied by L. tribracteatum. At this station there also occurred Potentilla supina, formerly present at Jonquières. At Meyne, L. thesioides was accompanied by several Centaurium (C. pulchellum, C. spicatum and C. tenuiflorum), Blackstonia serotina, Deschampsia media, Brachypodium phoenicoides, Phleum pratense subsp. serotinum, L. hyssopifolium etc.359. The habitat of L. thesioides thus appears to be quite different from that of L. thymifolium and belongs to the Lythrion tribracteati alliance or even the Deschampsion mediae.

Habitat

Ecology

General description

Biological characteristics

(2) L. thesioides Sub-pontic species. France Lower Rhône valley: Pierrelatte (Drôme), Caderousse and Orange (Vaucluse), Tresques, La Capelle-et-Masmolène, Meynes, and Jonquières-Saint-Vincent (Gard). Other Mediterranean countries Two old records from Italy in the floodplain of the Pau.

(1) Habitats suitable for L. thymifolium include pools that are temporarily flooded or simply saturated in winter, and very dry in summer, and also the edges of streams, marshes and ponds. The substrate is usually siliceous (sandstones, schists, basalts, etc.), the floodwater oligotrophic* and of variable depth between sites and years. (2) L. thesioides is also associated with habitats that are floodprone or very wet during the winter, such as the banks of natural freshwater ponds or the alluvial terraces of the Rhône or its tributaries. On the edges of ponds, L. thesioides occurs in the outer band of vegetation, leaving the centre of the marsh to the low, more hygrophytic formations of the Isoetion (formerly at Pazac, in the Gard) or the Heleochloion (Etang de la Capelle, and probably formerly at Jonquières, in the Gard).

44

Life form Therophytes (annual species). Reproduction Aerial. Description of seeds (1) Seeds glabrous, brownish, near ovoid in shape (length: 0.5 mm; width: 0.4 mm), very finely punctuate and light (weight <0.80 mg). (2) Seeds oblong-obovate, pale, irregularly convex on the outer surface, applanate-concave on the inner. Integument covered in very short appressed hairs when dry192.

Plant species

Biological cycle (1) In Morocco, germination at the beginning of February if the site is flooded, vegetative growth from February to the end of March, flowering in April-May, fruiting end of May and June, and seed dispersal a short distance from the parent plant. In France flowering is slightly later, taking place in May and June. (2) Cycle unknown. Based on examination of herbarium samples, flowering takes place mostly in August-October and rarely in June (one record). The phenology* of this species is therefore later than for L. thymifolium, and closer to L. tribracteatum for example. The stamens have anthers which extend to the same level as the stigma, which raises the a priori possibility of autogamy*. This mode of reproduction is all the more probable as the flowers often lack petals. The capsule, which barely extends outside the calyx, opens at the tip to form four very short valves. Seed dispersal is consequently probably limited. The seeds undoubtedly remain viable for a long time, as the plant may not appear every year and may disappear for many years between appearances at a given place. Lythrum thymifolium

Caux-Fontès-Pézenas and Agde, Quaternary siliceous plains of Béziers and Montpellier in the Hérault, Costière Nîmoise in the Gard, Plateau du Coiron in the Ardèche, the Issoire area in the Puy-de-Dôme, Upaix in the Hautes-Alpes, the Lanau pool in the Bouches-du-Rhône, Plateau de l’Evenos , the Plaine and Massif des Maures, Colle du Rouet and Bois de Palayson, Estérel of Fréjus in the Var and the Biot Massif in the Alpes Maritimes246, 274. • In Spain and Portugal: on the Balearic Islands and distributed over a large part of the peninsula, with the exception of the Cordillera Cantabrica64. • In Italy: in the three regions of Puglia, Basilicata and Lazio, and in Sardinia292. • In Morocco: pools in the Atlantic plains (Benslimane) and pools in the Atlas Mountains. (2) L. thesioides In France the plant has only been found 14 times at seven localities since it was discovered by A. Jordan in August 1841 at the Etang de Jonquières. The Etang de la Capelle is the only site at present (1998). Conservation status

Morocco Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Dec.

Aug.

Sept.

Oct.

Nov.

Dec.

Germination and vegetative growth Flowering Fruiting Seed dispersal

(1) L. thymifolium Populations declining due to the transformation of their biotopes. This species is assumed to have disappeared from some former stations (Allier, Haute-Loire, Saint-André-d’Embrun, Aude, former Etang de Jonquières, etc.)..

France Jan.

Feb.

March

April

May

June

July

Germination and vegetative growth Flowering

? Fruiting

?

Seed dispersal

Environmental conditions

Hydrology For seed germination it is only necessary for the soil to be saturated with water. Flooding is not essential. At the vegetative stage the plant can tolerate shallow water (< 5-7 cm). Substrate Sandy to sandy-silty, even silty for L. thesioides. Substrate above all non-calcareous (silica, basalt) for L. thymifolium.

(2) L. thesioides • In France, the following localities have been profoundly modified and the possibility of Lythrum surviving there these days appears very remote: the Etang de Jonquières, now drained and cultivated, where the species has not been seen since 190258; the Etang de Pazac, drained and converted to vineyards and pasture, where the species was only seen once, in 1951360, 361; the areas around Orange (seen on one occasion in 1892), Caderousse70, Tresque (appeared en masse, but only once, in 1853) and Pierrelatte (1915, not seen since)71, 148. These days all these areas are used for agricultural activities that are not very propitious for the survival of annual Lythrum. The plant has not been seen for a long time elsewhere in Europe or in the former USSR. • In Italy, it is considered to be extinct, its two sites not having produced records since the 19th century274.

Legal status/level of protection

Interspecific competition Deep shade from tall plants may be unfavourable for these heliophilous* species, especially at sites which are not grazed (France). Impact of perturbations Favourable in that the habitat is opened up.

Conservation - Management Assessment of populations

(1) L. thymifolium • In France, from west to east: Massif des Fenouillèdes and Plaine de la Têt in the Pyrénées-Orientales, plateaux of Roque-Haute,

IUCN 1997 Bern Convention Habitats Directive National and regional protection

National Red Data Books and Red Lists

(2): strictly protected (annexe I) (1) and (2) France, national list: Decree of 20/01/1982/Journal Officiel 13/05/1982, amended by Decree of 31/08/1995/Journal Officiel 17/10/1995 1) and (2) France, national Red Data Book: priority species274 (1) Morocco: “rare” (R)133, 181, 381 (2) Italy, national Red Data Book: “extinct”86

45

Mediterranean temporary pools

Conservation problems/threats Anthropogenic factors

Urban development, as well as drainage of pools and ponds, their filling in and their conversion to agriculture, constitute the greatest threats for both species. Natural factors

In Morocco, successions of dry years prevent the renewal of the L. thymifolium seedbank*, which could pose a problem for some populations in the long term. In France the closing-up of the habitat, resulting from the steady increase of woody plants, constitutes a threat for this species.

• Control over urbanisation. • Maintenance or promotion of a certain degree of opening of the habitat by grazing or by managing the vegetation cover (clearing). Scraping the surface of the soil appears to be beneficial for L. thesioides. • In France, gaining control over the ownership of plots of land at the Etang de la Capelle, in order to implement management and to remove the potential threats of destruction which are still faced by this last known site for L. thesioides. Looking for the species among the seed stocks at this site. Searching at the former localities where this species has been observed in the past, as well as in the few remaining wetlands in the lower Rhône valley, in an attempt to find new populations.

Risks relating to populations

• (1) In France, the L. thymifolium stations are vulnerable and subject to the risk of local extinction, especially as the species is at the northern limit of its range. • (2) The same is true of L. thesioides whose very insecure status in France is perhaps partly mitigated by the longevity of the seedbank in the soil. However, the very severe decline in the species throughout its entire range exposes the single currently confirmed population to particular risk. Management and conservation measures Current measures

No direct management measures either in Morocco or in France. In France, only collection of the seeds of the two species (preserved ex-situ) has been carried out, by the Conservatoire Botanique National Méditerranéen de Porquerolles. Recommendations

• Avoidance of any alterations to the hydrology of the pools and ponds (drainage, infilling or, conversely, permanently flooding).

46

Bibliography Anonymous, 199912 ; Aubert & Loisel, 197118 ; Cabanès, 190358 ; Castroviejo, 1986-200164 ; Charrel, 191370 ; Châtenier, 192271 ; Conti et al., 199286 ; Danton & Baffray, 199596 ; Fennane & Ibn Tattou, 1998133 ; Fournier, 1961139 ; Garraud, 2003148 ; Jahandiez & Maire, 1931-1934181 ; Jordan, 1847192 ; Lesouëf, 1997210 ; Mandin & Hugonnot, 2001234 ; Medail et al., 1998246 ; Michaud & Molina, 1999249 ; Olivier et al., 1995274 ; Pignatti, 1982292 ; Poirion & Barbero, 1966296 ; Rhazi et al., 2001314 ; Rouy & Camus, 1901327 ; Tallon, 1923-1969359, 1953360, 1967361 ; Titolet & Rhazi, 1999381 ; Tutin et al., 1964-1980385 ; Velayos, 1997394 ; Webb, 1968400. Authors: Michaud H. & L. Rhazi Collaborators: Antonetti P., L. Garraud, J. Molina & G. Paradis

Plant species

Marsilea strigosa

Willd.

POLYPODIOPHYTES300 MARSILEACEAE Main synonyms

M. pubescens Ten. M. fabri Dunal French names: Fougère d’eau pubescente, Marsilée pubescente. Spanish names: Trébol de agua, Trébol de cuatro hojas,

Marsilia180 Moroccan names: Ouarda el maa, Rjel-el-Ketta32, 265 Algerian names: Arbas ourrak, Qoub303 English name: Clover Fern (a part of the genus) Subspecies

None

1 cm

Description/identification criteria2

• Heterosporous* fern (with macrospores and microspores), aquatic, with long stolons and thick downy rhizomes with short, close internodes. • Fronds characteristic, formed from 4 leaflets arranged in a cross at the apex of the petiole (rachis), like a Four Leaves Clover leaf. • Two types of fronds: - The first with a floating blade, glabrous, borne on a long petiole, formed during the aquatic phase. - The second with a rigid, upright petiole and downy blade, appearing during the dry phase. • Fruiting: sporocarps* villous, axillary, subsessile, arranged in two rows along the rhizome. Similar species

Water Shamrock (Marsilea quadrifolia L.; syn.: M. quadrifoliata (L.) L.) has always glabrous leaflets and sporocarps with short pedicels, solitary or clustered in twos or threes. In France, M. quadrifolia grows in the temperate region2, whereas M. strigosa is a strictly Mediterranean species, which reduces the risk of confusion. In Spain, M. strigosa can be confused with M. quadrifolia as well as with an endemic species of the Iberian Peninsula, M. batardae (Medina, pers. com.). The latter species has the rhizomes much longer and branched, and widely spaced sporocarps, distinctly pedicellate, each pedicel bearing a conical tooth300. In Morocco, M. minuta (= M. diffusa) has a slender rhizome and distinctly pedicellate sporocarps, grouped in twos or threes

Distribution - Ecology

Marsilea strigosa

France

It is only found today in three localities: • Hérault: pools in the Roque-Haute plateau and a wet depression in the wine-growing plain of Vendres. • Pyrénées-Orientales: alluvial terraces of the Têt, in the uncultivated land of Torremila. Other Mediterranean countries

Italy: south of the peninsula (several localities in the province of Puglia, region of Taranto) and Sardinia86, 292. Spain: between 0 and 400 m, in 14 provinces (Medina, pers. com.). Above all found in the regions of Catalonia, Levante, Almeria, Extramadura, Castilla y Leon and on the islands of Majorca and Minorca64. Algeria303: temporary pools of the Algerois and the Oranais (coastal sahels, coastal plains, Tellian Atlas). Morocco134: dayas* of the Atlas ranges, up to 900 m in altitude (Middle Atlas, High Atlas and Anti-Atlas) and the Atlantic plains (Benslimane, Sidi Bettache).

Distribution/range160, 386

Egypt: Nile Valley and Delta46. Marsilea strigosa is a western Mediterranean species; it also occurs in some localities to the north of the Caspian Sea. It is present in France, Italy, Sardinia, Spain, the Balearics, Algeria, Morocco, Egypt, Romania and Ukraine (Crimea). Its existence in Portugal has been disproved.

Habitat General description

Marsilea strigosa appears to occupy a relatively wide range of natural habitats and substrates, and to be part of several phytosociological units, some of which remain to be specified2.

47

Mediterranean temporary pools

The species grows mainly in temporary pools with nutrient-poor and poorly mineralised water (Medina, pers. com.). It is found within amphibious vegetation which are submerged for a fairly long time (in France at Roque-Haute, or in Spain), dominated by Isoetes setacea or Mentha cervina, or in Crassula vaillantiiLythrum borysthenicum formations2, 251, 319. In Morocco, in the Atlantic plains, it grows in Isoetes velata, Ranunculus peltatus and Myriophyllum alterniflorum formations312. It also occurs in regularly flooded former vineyards, cleared of vines on siliceous gravel* (Vendres plateau and Plaine du Roussillon, France), often in furrows in which the water collects. Habitats Directive

“Mediterranean Temporary Pools” (code 3170). CORINE Biotopes

22.34 Southern amphibious communities (Isoetalia). 22.341 Short Mediterranean amphibious swards (Isoetion). Phytosociology*

France Marsilea strigosa is considered a characteristic species of the Isoetetum setaceae association* Br.-Bl. (1931) 1935215. On abandoned farmland, the formations have yet to be incorporated into phytosociological units2. These are most likely Isoetion formations heavily disturbed by viticulture. Spain The species is characteristic of the Sisymbrello-Preslietum association (Preslion cervinae alliance319).

Sporocarps Suborbicular, tightly packed, sessile* or with very short pedicels (2 to 2.5 mm) covered in articulated hairs300, with a diameter of 3 to 5 mm. These are highly efficient survival organs due to their capacity to survive several decades, or even over a century of drought. In experimental conditions, a sporocarp aged 103 years, taken from a herbarium, was able to produce 15 embryos83. Size of microsporangia: 50-62 microns180. Size of macrosporangia: 450-500 microns180. Biological cycle The sporocarps*, often saturated with water from the autumn, open in late winter and early spring, releasing the microsporangia and macrosporangia. Fertilisation occurs over the following days. The first, filiform, fronds appear within a few days, followed by fronds with two leaflets, then with four leaflets. They appear under a thin layer of water, during the dry phase of the pool, in winter in Morocco (January-February) and in spring in France (in March-April at Roque-Haute). When the soil is waterlogged, vegetative growth is prolific. The rhizome continues to grow, emitting a large number of fronds (dense tufts) as well as stolons which take root, enabling new rhizomes to develop. The formation of sporocarps and spores has been observed in March-April in Italy86 and from April to June in France. When the pool dries out completely, the plant acquires its terrestrial habit. The fronds wither and disappear towards the end of the summer in Morocco (Rhazi L., pers. com.) and up until the autumn in France. The plant then only survives in the form of sporocarps arranged in two tight rows on either side of the driedout rhizome.

Companion species

France • In temporary pools (Roque-Haute and Torremila): Isoetes setacea, Pilularia minuta, Mentha cervina, Lythrum borysthenicum, L. thymifolium, Eleocharis palustris, Myosurus breviscapus, Juncus pygmaeus, Pulicaria vulgaris and Polygonum romanum subsp. gallicum49 (Grillas & Tan Ham, pers. com. CEN-LR, pers. com. ). • On abandoned farmland, an impoverished form of this community can be found (disappearance of Isoetes, Mentha cervina, Pilularia minuta and Eleocharis palustris) augmented by ruderal species (Conyza spp., Dittrichia viscosa, etc.)2, 246. Morocco Isoetes velata, Ranunculus peltatus Lythrum borysthenicum, Myriophyllum alterniflorum, Eleocharis palustris and Baldellia ranunculoides312. Ecology

France Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Dec.

Emergence of fronds and vegetative growth ? Formation of sporocarps Withering of fronds

Environmental conditions

Hydrology The habitat of Marsilea strigosa is characterised by a fairly long inundation period, between five and eight months: from December to May-July in France (Rhazi M., pers. com.); from December to March in North Africa312, and by a maximum water depth of around 40 to 50 cm. In experimental conditions (not published), the opening of the sporocarps and fertilisation are observed under a thin layer of water, or even on saturated ground.

Biological characteristics

Life form Species considered to be perennial* (radicant hydrophyte*, producing stolons, traditionally considered as a geophyte* with a rhizome), but which appears to behave as an annual (therophyte*)395.

48

Substrate Usually non-calcareous (basaltic plateau at Roque-Haute, siliceous gravels at Vendres and at Saint-Estève). Interspecific competition This species is sensitive to competition by perennial grasses and to the shade of woody plants (heliophilous* species).

Plant species

Impact of perturbations The rhizomes of Marsilea do not appear to be sought by Wild Boar, though they can be dug up by chance by these animals. They are resistant to the turning-over of the top layer of soil in vineyards.

Conservation - Management

Legal status of the species/level of protection IUCN 1997 Bern Convention

Strictly protected species (Annexe I)

Habitats Directive National and regional protection

Annexes II and IV - France, national list: Decree of 20/01/1982/ Journal Officiel 13/05/1982, amended by Decree of 31/08/1995/ Journal Officiel 17/10/1995 - France, national Red Data Book: priority species. - Italy, national Red Data Book: “vulnerable”86 - Spain, regional lists (Medina, pers. com.) - Balearics: “vulnerable”331

Assessment of populations France

National Red Data Books and Red Lists

Very rare species with only three sites known. At Roque-Haute, over an ensemble of 200 pools, Marsilea appears to be disseminate in small populations in 10 to 26 pools depending on interannual variations in rainfall162, 246. On Torremila, the species is present in small scattered populations in three humid areas of uncultivated land and one temporary pool.

Conservation problems/threats

Italy

Anthropogenic factors

Considered as very rare (a few localities). Spain

Fifty or so localities in the regions of Catalonia, Levante, Almeria and Extramadura, including five which have now disappeared; very rare species in the Balearics64 (a few localities in southern Majorca and northern Minorca (Médina, pers. com.). Morocco

Many localities. The species, more common than M. minuta, is not considered to be rare or threatened133.

France These are the main threats. The habitat of the Vendres and Torremila stations has been degraded and partially destroyed by drainage and cultivation. It has returned to a wild state, where the species survives, though it risks being put under intensive cultivation again in the short term66. Only one intact pool survives among the vineyards of Torremila. At Roque-Haute, despite the “Réserve Naturelle (Nature Reserve)” status, the absence of control over land ownership and usage no longer permits management or monitoring of the site, and seriously compromises the conservation of the Marsilea populations.

Algeria

Rare species303. Conservation status

Spain The main threats are the drying-out of the pools (by drainage), overgrazing and climate change179 (Medina, pers. com.).

France

Natural factors

The populations have decreased: • The species has disappeared from the Clape d’Agde pool, where it was found in 1831, and from Aigues-Mortes where it was observed at the end of the 19th century2. • The station at Vendres is very seriously threatened; the plants have not been observed since 1998 but the sporocarps are still present in the soil. • The populations of the Saint-Estève pool and the uncultivated lands of Torremila, discovered in 1968, have been destroyed. The species has disappeared since 2000 from two other sites of uncultivated land discovered in 1996; however, two new stations have since been found on this site.

Vegetation dynamics At Roque-Haute, following the abandonment of grazing several decades ago, the spread of woody plants and invasion by Scirpus are threatening the habitat of Isoetes setacea315, which is also that of Marsilea251. At Torremila, the species is subject to competition from perennial grasses (mainly Deschampsia media), encouraged by the natural dynamics of the uncultivated land. Variability of hydrological conditions The interannual fluctuations in flooding cause fluctuations in the abundance of Marsilea without jeopardising the survival of the populations.

Morocco

The species is considered as stable (Rhazi L, pers. com.).

Risks relating to populations

Spain

Its very reduced capacity for dispersal results in a poor aptitude for the colonisation of new sites, and significant isolation of populations, both geographically and genetically395.

The species is in decline (province of Valencia in particular); 10% of stations are expected to disappear over the next 20 years (Medina, pers. com.).

49

Mediterranean temporary pools

Management and conservation measures Current measures

France The Roque-Haute site is protected by its legal “Réserve Naturelle” status. Marsilea was the subject of regular monitoring here from 1992 to 2001. At Vendres, the site is included in a Natura 2000* area, and the Conservatoire des Espaces Naturels du Languedoc-Roussillon (CENLR) has acquired land to favour the restoration of the hydrological regime. At Torremila, the Conservatoire has drawn up the Objectives Document (DOCOB) of the Natura 2000 site (Natura 2000 FR 9102001 “Torremila Humid Uncultivated Land”). The two sites of Vendres and Torremila have been selected for the setting-up of a LIFE project for the conservation of Alpine and Mediterranean flora of EEC interest at the European scale (France, Spain, Italy). Conservation ex-situ: sporocarps have been collected by the Conservatoire Botanique National Méditerranéen de Porquerolles. Cultivation trials successfully carried out in January 2004, particularly within the framework of a project for the restoration of the Vendres site, show the viability of harvested sporocarps. Spain Only a few localities are included in Natura 2000 areas. Some are the subject of particular programmes (Province of Valencia, Balearics) (Medina, pers. com) such as the creation of two microreserves in the lagoons of Sinarcas (Province of Valencia) and the production of sporocarps ex-situ from fragments of rhizomes with a view to the reintroduction of the species179. The species is classified as “threatened” on the regional lists of the regions where it is present (Medina, pers. com.). Italy The species is cultivated in the botanical gardens at Padua86. Recommendations (for France)

Viticulture Ban intensive viticulture on all sites, as it is totally unfavourable to the survival of Marsilea populations through the impact of various agricultural practices (drainage, burying of sporocarps, herbicides). On the other hand, outside of the temporary-pool habitat, extensive viticulture (no herbicides, light cultivation of

the soil surface) is not incompatible with the conservation of Marsilea populations, provided it permits flooding from autumn to spring. Competition from perennials and woody plants (closing-up of the habitat) • In the Roque-Haute Nature Reserve: implement the management plan and more generally restore a concerted management with a natural-heritage objective: reintroduce grazing, preferably by sheep and goats, together with an appropriate monitoring programme. • At Torremila: promote management of perennial grasses by water-weed cutting and grazing. Isolation of populations395 Genetic isolation has implications for conservation strategies: to properly protect this species, and preserve its adaptive potential, it is essential to protect a large number of its populations. Absence of control over land ownership and uses Restore control over land ownership and/or usage, an important prerequisite for the implementation of conservation management of the populations, particularly at Roque-Haute.

Bibliography Aboucaya et al., 20022 ; Aizpuru et al., 20006 ; Amigo, 198710 ; Anonymous, 199912 ; Baudière & Cauwet, 196823 ; Bertrand, 199132 ; Bissardon & Guibal, 199739 ; Boulos, 199546 ; BraunBlanquet et al., 195249 ; Castroviejo, 1986-200164 ; CEN-LR, 200166 ; Colas et al., 199683 ; Conti et al., 199286 ; Bern Convention,197987 ; Danton & Baffray, 199596 ; Directive 92/43/CEE, 1992119 ; Fennane & Ibn Tattou, 1998133 ; Fennane et al., 1999134 ; Foucault de, 198898 ; Gaudillat & Haury, 2002153 ; Greuter et al, 1984-1989160 ; Grillas & Tan Ham, 1998162 ; Ibars & Estrelles, 1997179 ; Ibars et al., 1999180 ; Lewin, 2000213 ; Loisel, 1976215 ; Médail et al.,1998246 ; Molina, 1998251 ; Nègre, 1956264, 1961-1962265 ; Olivier et al., 1995274 ; Pignatti, 1982292 ; Prelli, 2001300 ; Quézel & Santa, 1963303 ; Quézel, 1998306 ; Rhazi, 2001312 ; Rhazi, 2004315 ; Rivas Goday, 1970319 ; Saez & Rossello, 2001331 ; Tutin et al., 1964-1980386 ; Vitalis et al. 2002395 ; Walter & Gilett, 1998399. Author: Yavercovski N. Collaborators: Garnéro S., C Houssard., M. Klesczewski,

H. Michaud & F. Ruchon

50

Plant species

Mentha cervina L. ANGIOSPERMS LAMIACEAE Main synonyms

Preslia cervina (L.) Fresen. French name: Menthe des cerfs Spanish names: Menta de burro, Poleo de ciervo Moroccan name: Fliyou Subspecies

None Description/identification criteria274, 292, 386

• Perennial plant with rhizome very close to the surface42, recognisable by its very musky mentholated scent, its upright habit, its very narrow (1 to 2.5 cm long and 1 to 4 mm wide) opposite leaves (in the axils of which grow fascicle of smaller leaves borne on very short branches), and above all, by the characteristic palmate shape of the bracteoles* surrounding the whorls of flowers, which evoke the antlers of a deer, and to which the species owes its name. • Upright stems, hollow and angled at the base (10 to 60 cm), square-shaped in section and slightly branched. • Glabrous leaves, sessile* but attenuate* at the base, entire or roughly toothed. • Flowers hermaphrodite, forming compact whorls, widely spaced on the upper half of the stem. Calyx tubular and villous at the throat, with four triangular teeth terminating in a fine point. Corolla twice as long as the calyx, with two equal lips, pink or white. Floral bracts identical to the leaves, but larger.

1 cm

Mentha cervina

Similar species

During and after flowering, no confusion is possible: it is the only mint which has deeply lobed bracteoles and whose calyx has four teeth and not five, hence its former inclusion in the genus Preslia. In the vegetative state, the narrow, sessile leaves are characteristic and differentiate it from the Pennyroyal (Mentha pulegium), frequently present in the same habitats but whose leaves are rounded with short petioles. Before flowering, the young shoots do not yet have their scent of mint, and can be confused with young Hedge Hyssop (Gratiola officinalis) plants.

Other Mediterranean countries

Spain and Portugal: western half of the Iberian Peninsula, with some localities in the eastern part AND in the Balearics (Medina, pers. com.). Italy: once found in the wetlands and temporary pools of Abruzzo, but not seen recently292. Algeria: temporary pools of the high plateaux of the Algerois and the Oranais, Geryville region303. Morocco: dayas* and edges of peat bogs of the Rif regions (Rhazi L., pers. com.).

Distribution - Ecology

Habitat

Distribution/range

General description

Species of the thermo and meso-Mediterranean zone. Western Mediterranean species, found in six countries160, 386. France

Present mainly in Languedoc-Roussillon246. In the PACA region (Vaucluse and Bouches-du-Rhône), its presence has not been confirmed recently306, but it was found in 2003 at the Etang des Aulnes (Willm, pers. com).

- In France, this mint occupies the long-flooded parts of temporary pools and lavognes*, and a number of temporary water courses of the Montpellierais garrigues (Molina, pers. com.). It once occurred in the ox-bows of the Rhône (breaches and launes* around Tarascon, Avignon, etc.), where hydraulic modifications caused its disappearance. It forms, with the often perennial species associated with it, areas of tall-herb grassland of several dm2 to several m2. - In North Africa (Morocco, Algeria), the species grows at an altitude where the rainfall is sufficient (above 1,000 m in the

51

Mediterranean temporary pools

Moroccan Rif). In the temporary pools of the plain, it gives way to the Pennyroyal, which is less water-demanding (Rhazi L, pers. com.).

In Morocco, its development is earlier: the shoots appear in March, flowering commences in May and fruiting in early summer.

Habitats Directive

“Mediterranean Temporary Pools” (code 3170).

France Jan.

Feb.

CORINE Biotopes

22.34 Mediterraneo-Atlantic amphibious communities. 22.342 Tall Mediterranean amphibious swards (Preslion cervinae): vegetation of tall annuals of land covered by deep water for long periods, with Mentha cervina.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Dec.

Sept.

Oct.

Nov.

Dec.

Emergence of shoots and vegetative growth Flowering Fruiting Morocco Jan.

Feb.

March

April

May

June

July

Aug.

Germination and vegetative growth

Phytosociology*

Class: Isoeto-Nanojuncetea Order: Isoetalia Alliance: Preslion cervinae Associations*: • In France49, 215: - Preslietum cervinae: Mentha cervina-Eleocharis palustris association, described from western Languedoc. - Preslio-Trigonelletum ornithopodioides: association described from the Costière Nîmoise (Gard), from where it has disappeared. A related formation is perhaps present in the Crau98. • In Spain319: - Preslietum cervinae - Sisymbrello-Preslietum: Mentha cervina is accompanied by Sisymbrella aspera, Veronica anagalloides, Cyperus Badius, Marsilea strigosa, Eryngium galloides, Callitriche platycarpa, etc. - Preslio-Eryngietum corniculati: M. cervina is present here with Eryngium corniculatus, I. velata, I. setacea, etc.

Flowering Fruiting

Environmental conditions

Hydrology In France, the habitat of this species is characterised by a long period of flooding, from the autumn to spring153 (six to eight months depending on the year), a depth of water of between 20 and 70 cm (or 100 cm), with an optimum of around 40 cm (Rhazi M., pers. com.). Germination takes place under a thin layer of water (non-turbid) or waterlogged ground (Morocco). Substrate Various substrates, oligotrophic*274 or meso-eutrophic*98, on basaltic (Roque-Haute), calcareous (garrigues), marl (Minervois), or granitic (Fenouillèdes) bedrock, or on alluvium (borders of ponds).

Companion species

France At Roque-Haute, the mint grows with Marsilea strigosa and Isoetes setacea, or with Eleocharis palustris in the deepest parts of the pools251. In the Montpellierais and Uzégeois garrigues the companion species are Eleocharis palustris, Sisymbrella aspera, Juncus articulatus and Mentha pulegium. Morocco Eleocharis palustris, Callitriche truncata and Alisma plantagoaquatica are present at its stations. Ecology Biological characteristics

Life form Perennial species with vegetative growth (suckers, rhizomes). Although considered by several authors as hemicryptophyte*, in France this species is more of a rhizomatous geophyte*, as it is not visible before the beginning of the spring, and the young shoots appear towards the month of April. Biological cycle In France, its vegetative growth occurs in the spring, and its reproductive cycle in summer. At Roque-Haute, it flowers from late June to mid-August, fruits from late July to September and withers in October.

52

Interspecific competition This heliophilous* species is sensitive to shade. The water depth limits invasion by woody species, but on the other hand creates conditions favourable to colonisation by fiercely competitive helophytes* (reeds, Scirpus, Reedmace, sedges)153.

Conservation - Management Assessment of populations

• In France, two major sites in terms of the number of populations: the Montpellierais garrigues (Hérault), and the Uzégeois and Bagnolais garrigues (Gard). Then two other less important sites: Fenouillèdes (Pyrénées-Orientales) and Roque-Haute (Hérault). Finally, ten or so secondary sites in Languedoc and the Crau, where the plant forms populations which appear to be isolated, and sometimes very small (a few plants in the Etang de Pujaut, Gard). • In Italy, the species is considered to have disappeared292. • In Algeria, a very rare species303. • In Morocco, fewer than five localities133. Other stations could be discovered by searching in the mountainous regions of the Rif. • In Spain and Portugal, the species is not considered rare.

Plant species

Conservation status

Natural factors

France

The shade linked to invasion by woody species, as a result of the abandonment of grazing at some sites (France), is without doubt unfavourable for this species.

The populations of this species seem to be undergoing significant declines over the whole of its range, except in the Hérault and the Gard. • Gard: the species remains abundant in the Uzégeois-Bagnolais (plateaux of Méjannes-le-Clap, Lussan, etc.), and is surviving in the region of Quissac (pools at Vibrac). It is still present in the marshes of Pujaut157, 246 and further north in a lavogne* in the eastern end of Larzac, but it has disappeared from the Costière Nîmoise (Redessan marsh, Etang de Pazac and around Beauvoisin). • Hérault: the Grammont station has disappeared208, 246, but the other populations are stable (temporary pools and streams of the Montpellierais garrigues, the Bittérois plain, and the plateau of Roque-Haute, the Assignan pool in the Minervois). • Pyrénées-Orientales: the species has disappeared from the Saint-Estève pool (alluvial plain of the Têt) and the Catalan coast246 but survives on the Rodès plateau. • Bouches-du-Rhône and Vaucluse: With one exception (Etang des Aulnes), all the stations of the “Crau humide” (Marais du Vigueirat, Marais de l’Audience, Marais de Raphèle) and the “Crau sèche” (Etang du Luquier) appear to have disappeared246 , as have the stations of the sandy banks of the Rhône157, 158, 254, 255, 311.

Risks relating to populations

In North Africa and in France, with the exception perhaps of the Montpellierais and Uzégeois garrigues and the Roque-Haute plateau, a risk of extinction of populations exists in the medium to long term, because of their small number, their distance from one another, and their reduced size. Management and conservation measures Current measures274

Several stations benefit from protected status: • The Site Classé of the Gardiole (commune of Gigean in the Hérault). • The Roque-Haute Nature Reserve. • The Etang des Aulnes, property of the département of Bouches-du-Rhône, included in the network of Espaces Naturels Sensibles*.

Recommendations Other Mediterranean countries

• In Italy, the Abruzzo populations are considered to have disappeared292. • In Spain and Portugal, the species could be declining due to the destruction of its habitat (Medina, pers. com.). • In Morocco, the populations appear to be stable. Legal status of the species/level of protection IUCN 1997 Bern Convention Habitats Directive National and regional protection

National Red Data Books and Red Lists

France, PACA regional list: Decree of 9/05/1994/Journal Officiel 26/07/1994 - France, national Red Data Book: priority species274 - Morocco: “very rare” (RR)181, 133

Conservation problems/threats Anthropogenic factors

In France, habitat destruction is responsible for the disappearance of several important populations: • Reorganisation of land followed by drainage and cultivation (vineyards of the Costière Nîmoise). • Permanent flooding (Grammont and Saint-Estève pools). • Hydraulic modifications (banks of the Rhône). • Silting up, shading (cessation of grazing and lack of upkeep of the lavognes of the garrigues). • Game management of lavognes (sealing of the pool bed with a layer of concrete). • Extraction of materials: the Plateau de Rodès station is now threatened by the creation of a granite quarry246.

• Maintain a favourable hydrological regime • Maintain open vegetation by cutting or grazing. • Protect the habitat at the remaining stations (Rodès plateau in the Pyrénées-Orientales; Etang de Pujaut, Quissac pools etc, in the Gard). • Raise awareness among the rural public of the interest and richness of the many lavognes of garrigue zones so they can be maintained and prevent their beds from being concreted over. • Intensify prospecting in the areas where only a few populations are known (Minervois, the piedmont plain of the Cévennes: Quissac, Alès etc.). • Carry out inventories and monitoring of these populations (dynamics), which are vulnerable due to the fragility of their habitat.

Bibliography Aizpuru et al., 20006 ; Anonymous, 199912 ; Bissardon & Guibal, 199739 ; Bonnier, 199042 ; Braun-Blanquet et al., 195249 ; Conti et al., 199286 ; Bern Convention, 197987 ; Le Dantec et al., 1998208 ; Directive 92/43/CEE, 1992119 ; Fennane & Ibn Tattou, 1998133 ; Foucault de, 198898 ; Gaudillat & Haury, 2002153 ; Girerd,1990157 ; Goujard, 1997158 ; Greuter et al., 1984-1989160 ; Jahandiez & Maire, 1931-1934181 ; Loisel, 1976215 ; Médail et al., 1998246 ; Molina, 1998251 ; Molinier & Tallon, 1950254, 1950-1951255 ; Molinier, 1981257 ; Olivier et al., 1995274 ; Pignatti, 1982292 ; Quézel & Santa, 1962-1963303 ; Quézel et al., 1979308 ; Quézel, 1998306 ; Reynier, 1883311 ; Rivas Goday, 1970319 ; Tutin et al., 1964-1993386 ; Walter & Gillett, 1998399. Authors: Michaud H. & N. Yavercovski Collaborators: Rhazi L., M. Rhazi, F. Ruchon F. & H. Souheil

53

Mediterranean temporary pools

Myriophyllum alterniflorum

DC.

DICOTYLÉDONES HALORAGACEAE Main synonyms

M. verticillatum L. subsp. alterniflorum (D. C.) Bonnier & Layens French name: Myriophylle à fleurs alternes Spanish name: Ovas de rio Italian name: Millefoglio d’acqua gracile English names: Alternate-flowered Water Milfoil, Slender Water

Milfoil Subspecies

None Description/identification criteria

• Aquatic plant, herbaceous, slender (10 to -50 cm), rooted, glabrous, branching. • Leaves green, 6 to 30 mm in length, sometimes reddish, pinnate hair-like. Submerged leaves whorled, aerial leaves situated on the fertile branches and almost all alternate. • Bracts of upper flowers entire or toothed, shorter than the flowers. • Flowers yellowish, forming a spike, generally with 4 petals. Calyx with a short tube with four deciduous lobes. Flowers few, single-sex, alternate, in a very slender, short, at first drooping spike, consisting of three to four female flowers in a whorl at the base and five to eight male flowers in a spike at the tip (with eight stamens). • Ovary inferior, with four carpels, each one containing four loculi with single ovules. Tetragon-shaped fruit, finely tuberculated. Similar species

• Myriophyllum spicatum: has many pink flowers, all whorled, forming a flowering spike, robust and always upright265. • M. verticillatum: has the bracts of the upper flowers all pectinate and longer than the flowers. The spike is very elongated, terminating in leaves; the flowers are pink.

0.5 cm

Myriophyllum alterniflorum

General description

In Morocco and in Corsica, this species is found only in temporary pools. In Morocco, it is present in the pools on the plains (region of Benslimane) and mountains in the Rif region (Outka and Issaguene), on siliceous soils (sandstone, quartzite) and in a semiarid bioclimate*, sub-humid or humid. These pools are generally filled with rainwater in December-January and dry out in AprilMay. Habitats Directive

“Mediterranean Temporary Pools” (code 3170). CORINE Biotopes

Distribution - Ecology Distribution/range

Broad distribution: Europe, North Africa, North America. Mediterranean countries64, 86, 386: France, Spain and Portugal (dispersed over all the peninsula), Italy (Lombardy, Lazio, Sila, Sicily and Sardinia), Tunisia, Algeria and Morocco. France

• On the mainland, the species is absent from the Mediterranean region, and elsewhere occurs in streams and ponds with a siliceous soil. • In Corsica, it is found in seven temporary pools in the south east of the island: four pools in the Tre Padule de Suartone Nature Reserve and three pools, of artificial origin, on the Frasselli plateau. Habitat

54

This species appears in the aquatic phase of oligotrophic* temporary pools. 22.34 Southern amphibious communities (Isoetalia). Phytosociology*

In Morocco, the species is characteristic of Myriophyllum alterniflorum-Ranunculus peltatus communities. The species associated with this community are Callitriche brutia, Glyceria fluitans, Chara vulgaris, Nitella opaca and Illecebrum verticillatum. In the pools of Corsica (pools with Isoetes velata), the associated species are Ranunculus peltatus, Illecebrum verticillatum and Apium crassipes, as well as Potamogeton pectinatus on the Frasselli pools (which indicates slight eutrophication linked to the presence of livestock).

Plant species

Impact of perturbations Favourable when they facilitate the opening-up of the habitat.

Ecology Biological characteristics

Life form Perennial species in permanent waters (hydrophyte*) but with an annual cycle (therophyte*) in temporary Mediterranean pools.

Light The species is heliophilous*, which explains its absence in the turbid pools of the Cork-Oak forest of Mamora in Morocco and in deep pools (with greater depths of water).

Reproduction Aerial pollination. Flowering in March-April.

Conservation - Management

Description of seeds Seeds: four per fruit, small (length: 1.5 mm; width: 1 mm to 1.25 mm). Biological cycle This begins under the water and often ends out of the water. In Morocco and Corsica, germination takes place in FebruaryMarch, when the pool is under water, and flowering in MarchApril. Fruiting, which begins during the aquatic phase, takes place in May-June, but the maturation of fruits and seeds, and their dispersal a few centimetres from the mother plant, occurs in June-July after the drying-out of the pool. In addition to this sexual reproduction, the plant reproduces vegetatively during the aquatic phase, by division and regrowth.

Assessment of populations

• In Morocco, Myriophyllum alterniflorum has been found in 14 pools133, 230 in the Atlantic plains (Benslimane) and the mountainous region of the Rif. • In Algeria, the species is considered to be rare. There are few sites with M. alterniflorum and these are confined to just two regions, the Algiers region (coastal subregion) and the Kabyle region (Greater and Lesser Kabyle and Numidia)303, 338. • In Mediterranean France, there are seven pools, with M. alterniflorum, three of them artificial, and all of them in Corsica. • On the Iberian Peninsula and the Balearics, the species is widely distributed (40 provinces in Spain, five in Portugal). Conservation status

Morocco and Corsica Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Dec.

In Morocco, in Spain and in Corsica, the populations of M. alterniflorum appear to be stable. In Italy, the stations in Tuscany have disappeared86.

Germination

Legal status of the species/level of protection

Flowering Fruiting Seed dispersal

Environmental conditions

Hydrology The germination of seeds requires flooding or at least waterlogging of the soil. The plant is adapted to submerged conditions (divided leaves, etc.) and can withstand high water levels provided there is sufficient light: tolerance to water depth increases with the clarity of the water (for example 50 cm in the pools of Benslimane in Morocco or Padule Maggiore in Corsica). The plant forms its flowers under the water but they emerge above the surface of the water for opening and pollination (anemophily). The water is oligotrophic* to mesotrophic*. Substrate Acidic to neutral. Interspecific competition M. alterniflorum is abundant in the centre of pools, where herbaceous species such as Ranunculus peltatus, Callitriche brutia, etc. grow, but it becomes less abundant or even absent in areas with helophytes* (Scirpus maritimus, Eleocharis palustris), due to competitive exclusion.

IUCN 1997 Bern Convention Habitats Directive National and regional protection National Red Data Books and Red Lists

- Morocco: “very rare” (RR)133, 181, 381 - Italy, national Red Data Book: “vulnerable”86

Conservation problems/threats Anthropogenic factors

Filling in, urbanisation, cultivation and the modification of the hydrology of pools (drainage) constitute the main threats for this taxon. In Morocco, there is a significant risk of extinction of populations at some stations because of urbanisation and the fillingin of pools. Natural factors

In Morocco, the frequency of dry years limits the appearance of this species313 and reduces the chances of renewal of the seed bank *, which could pose problems in the long term. In Corsica, the closing-up of the habitat (invasion by woody species or by helophytes) is a potential threat for the species if grazing is abandoned.

55

Mediterranean temporary pools

Management and conservation measures

Bibliography

Current measures

Anonymous, 199912 ; Castroviejo, 1986-200164 ; Conti et al., 199286 ; Fennane & Ibn Tattou, 1998133 ; Fournier, 1961139 ; Jahandiez & Maire, 1931-1934181 ; Lorenzoni, 1997225 ; Lorenzoni & Paradis, 2000221 ; Maire, 1952-1987230 ; Marchetti, 1997235 ; Nègre, 1961-1962265 ; Pignatti, 1982292 ; Quézel & Santa, 19621963303 ; Rhazi et al., 2001313 ; Rhazi, 2001312 ; Samraoui & de Belair, 1998338 ; Titolet & Rhazi, 1999381 ; Tutin et al., 19641980386.

In Corsica, the populations of the Tre Padule de Suartone pools benefit from the legal status of the site (Nature Reserve). In Italy, the Sicilian stations are included within a Regional Park86. Recommendations

• Maintain the healthy hydrological functioning of the sites, avoiding drainage and silting up. • Control urbanisation. • Maintain an open habitat by controlling the dynamics of the vegetation (woody species and helophytes), and maintaining or introducing grazing. • Implement legal measures for conservation of non-protected sites.

56

Author: Rhazi L. Collaborators: Paradis G. & M. L. Pozzo di Borgo

Plant species

Nitella opaca

(Bruzelius) Agardh

CHAROPHYCÉES CHARACEAE Main synonyms

Chara opaca Bruzelius 1824 N. flexilis var. flexilis f. opacoides R.D. Wood 1962 French name: none English name: Dark stonewort Modified from Groves & Bullock-Webster164

Subspecies

None Description/identification criteria90, 164, 200, 358

• Dark green plants, small in size (10-30 cm), living entirely submerged. Thallus with whorled structure, entirely without cortication. • Whorls composed of 6-8 primary rays (branchlets), divided once to form 2-3 secondary rays (dactyls). Dactyls always unicellular with a blunt tip. Gametangia* situated in the forks of the primary rays. • Dioecious species, very fertile with a marked sexual dimorphism. • Male plants characterised by dense fertile heads with short dactyls; antheridia* solitary and of large size (diameter 650775 µm). • Female plants with longer primary and secondary rays, giving them a more open appearance; fertile nodes carrying up to three, though usually two oogonia* 600-700 µm long and 500-600 µm wide. Oogonium bearing a coronule* composed of 2 x 5 tiny cells. Oospores laterally compressed, dark brown to black, c. 350-500 µm long and 350-400 µm wide with 6-7 spiral ridges. In the fresh state, the oospores have characteristic “winged” extensions of the spiral ridges. - Incrustations of calcite forming more or lesse regularly spaced rings around yhe axis and branchlets, often occur at stations with a low water level. This feature, known as “banded phenomenon of the Characeae”310, may be attributed to the effect of intense light356. Similar species

Two other dioecious species can ressemble Nitella opaca: • N. syncarpa (Thuillier) Chevalier, but this species grows in summer or autumn, not in spring. • N. capillaris (Krokeil) Groves and Bullock-Webster, which also has spring growth but is distinguished by gelatinous mucus around the male and female gametangia90, and by its oospores which are ornamented with papillae131, 357. N. opaca is, with N. tenuissima, the only representative of this genus so far recognised in Languedoc-Roussillon and southeast France. It is easily distinguished from N. tenuissima which has primary branches with two to four forks (multiple dichotomy).

1 cm

male

female

Nitella opaca

Distribution - Ecology Distribution/range France

Nitella opaca is mainly present in central and western France (Aquitaine andBritany)90. The numerous localities listed by Corillion all date from earlier than 1957 and even from the beginning of the 20th century. A single earlier mention of N. opaca concerns southern France near Hyères in the Var (coll. Boulu 1851, determined and then cited by Hy178). Today, there are only five localities known in the south of France: Lanau pool (Bouches-duRhône), the temporary lakes of Bonne Cougne and Redon, (Var)358, some pools in the natural reserve of Roque-Haute (Hérault) and one pool at the Tour du Valat in the Camargue (Grillas, pers. com.). It has disappeared from the Moulin du Rouet site (Hérault) where it was recorded by W. Krause. Other Mediterranean countries

Rare in Spain200 and Portugal90; unknown in Italy and Greece. Algeria: three localities previous to 1950132. Morocco: five localities in 1974165 and from 1985-1990130. Tunisia: the species has been identified in the Holocene sediments of a fresh water locality close to the Mediterranean coast131. Habitat

In the Mediterranean region, the species has been identified only in temporary freshwater habitats, characterised by flooding at the end of the autumn and drying out in May/June. This type of habitat seems to be consistent both in southern France and the localities in North Africa. Habitats Directive

“Mediterranean Temporary Pools” (code 3170).

57

Mediterranean temporary pools

Environmental conditions90, 200

CORINE Biotopes

22.34 Southern amphibious communities (Isoetalia). 22.341 Short Mediterranean amphibious swards (Isoetion). 22.342 Tall Mediterranean amphibious swards (Preslion cervinae). Phytosociology*

N. opaca is part of the aquatic phase of Preslion and Isoetion communities (Isoetalia). This species is also characteristic of the Nitelletum opacae association, often in monospecific*90 populations or associated with Sphaerochara (personal observation).

Hydrology N. opaca is a fresh water species of habitats with a pH* close to neutral (6.5-7.5). N. opaca can be found in both temporary and permanent habitats and also adapts to weak currents. The minimum duration of submersion necessary to complete a full biological cycle is estimated at 5-6 months. Substrate Indifferent to substrate. Interspecific competition Because of its early appearance, N. opaca is not subjected to much competition from other hydrophytes.

Companion species

At the Lanau pool in the Crau (Bouches-du-Rhône), N. opaca is associated with Sphaerochara intricata355. These two species precede the other hydrophytes* and decompose during the emergence of communities of amphibious annuals with Damasonium polyspermum, Lythrum tribracteatum, Mentha pulegium, etc. At Lake Redon (Var), N. opaca is accompanied by Sphaerochara prolifera. The growth of this species also early and precedes that of Ranunculus ophioglossifolius, Baldellia ranunculoides and Butomus umbellatus. At Roque-Haute, it grows side by side with Isoetes setacea, Mentha cervina and Callitriche brutia in some deep pools. In the pools of Morocco, N. opaca is associated with Myriophyllum alterniflorum, Callitriche brutia, Glyceria fluitans, Chara vulgaris and Illecebrum verticillatum312.

Impact of perturbations Because it is abundantly fertile, the species is not very sensitive to “mechanical” disturbances, and even seems favoured by the trampling of livestock (Lanau pool) and wild boar (the longer lasting wet parts of Lake Bonne Cougne), which limit the terrestrial vegetation. Temperature Germination in cold water, fruiting between 12 and 18°C; N. opaca decays when the temperature of the water exceeds 20°C (personal observation). Light Not very light-demanding; however, sexual reproduction is reduced in deep lakes90.

Conservation - Management Ecology Assessment of populations Biological characteristics France

Life form Nitella opaca is an annual species. Biological cycle This is a vernal species, which grows while the water is cold. Thus, it appears very early in shallow habitats, while in western France (Britany) and in northern Europe, it is found up until the summer in permanent lakes and even down to a depth of 40 m in Sweden90, 200. In the Mediterranean region, it is present from February onwards and fructifies abundantly from the end of March to May. The plants then decompose entirely.

Distribution is concentrated in central and western France where it nonetheless appears to be in decline. An estimate of the number of sites in these regions is not currently possible. Five localities are known in the French Mediterranean region. Conservation status France

In overall decline: more of 32 localities listed by91 for western France, mainly Britany, had only been collected at the beginning of the 20th century. Other European countries

Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Dec.


Germination and


vegetative growth Reproduction

Unevenly distributed in northern Europe; considered as fairly frequent, but nonetheless vulnerable, in Spain79, while Krause200 mentions rare sites in central Spain. The species has been recorded recently in the Balkan countries40.

Decay

Legal status of the species/level of protection Figure established on the basis of monthly observations at the temporary lakes of Bonne Cougne and Redon in 2001358.

58

IUCN 1997 Bern Convention Habitats Directive National and regional protection National Red Data Books and Red Lists

-

Plant species

There is currently no legal protection for charophytes in France and the Mediterranean countries, nor are they included in any Red List of threatened species. Conservation problems/threats Anthropogenic factors

In France: • The hydrological regime and the water quality of the Lake Bonne Cougne (Var), which is partly supplied by a karstic* aquifer, are seriously threatened by a golf-course project. • At Roque-Haute, the absence of control over land ownership and usage (ban on access to the site by owners) is preventing the monitoring and the management of populations of Nitella opaca. Management and conservation measures Current measures

At Roque-Haute, the site benefits from the status of Nature Reserve. The pools underwent monitoring from 1997 to 2001. However, since 2002, the manager of the Nature Reserve has been prevented by the owners of the land from carrying out his monitoring and management activities. The Lanau pool has been acquired by the CEEP and undergoes regular monitoring. It is included in a Natura 2000 area, like the temporary lakes of Redon and Bonne Cougne.

Recommendations

It is essential to maintain the temporary nature of the hydrological regime with flooding during the cold season. In particular, in the Mediterranean region, care must be taken to maintain the natural alternation of flooding/drying out, which favours sexual reproduction and, because of this, enables the plant to spread to new pools. Populations of N. opaca are vulnerable due to the fragility of their habitat and it is thus important that the dynamics of these populations are inventoried and monitored.

Bibliography Blazencic & Blazencic, 200340 ; Cirujano et al., 199279 ; Corillion, 195790, 197591 ; El Khiati et al., 2002131 ; El Khiati, 1995130 ; Feldmann, 1946132 ; Groves & Bullock-Webster, 1920164 ; Guerlesquin, 1978165 ; Hy, 1913178 ; Krause, 1997200 ; Raven et al.,1986310 ; Rhazi, 2001312 ; Soulié-Märsche & Thiéry, 1998355 ; Soulié-Märsche & Vautier, 2004356 ; Soulie-Märsche, 1989357 ; Soulié-Märsche, 2003358 ; Wood, 1962403. Author: Soulié-Märsche I.

59

Mediterranean temporary pools

Ophioglossum azoricum

C. Presl (1) & Ophioglossum

lusitanicum

L. (2)

POLYPODIOPHYTES300 OPHIOGLOSSACEAE Main synonyms

(1) O. ambiguum Cosson & Germ O. polyphyllum auct. non A. Braun (2) O. vulgatum L. subsp. lusitanicum (L.) Bonnier & Layens French names: (1) Ophioglosse des Açores, (2) Ophioglosse du

Portugal Italian names: (1) Ophioglosso delle Azzorre, (2) Ophioglosso

lusitanico Portuguese names: (1) Lingua de cobra, (2) Lingua-de-cobra,

Lingua-de-cobra-menor English names: (1) Small adder’s tongue, (2) Least adder’s tongue Description/identification criteria96, 299

• Perennial plants, delicate and unobtrusive from 5 to 10 cm (1), and 2 to 5 cm (2), present in the form of small, fairly localised populations • Two types of fronds (“leaves”) very distinct, as in all the Adder’s-Tongues: a sterile type, with an extended blade, oval or lanceolate, sometimes lying flat but usually upright and curled over to form a furrow; the other fertile, reduced to a fruiting spike with a peduncle and bearing sporangia stacked up on top of each other in two symmetrical rows (at the origin of the name “ophioglosse” from the Greek glossa “tongue” and ophis “snake”). • Fronds of the two types separated at ground level (the sterile fronds generally sheathing the fertile fronds), light green or yellowish-green in colour, and yellowing completely before wilting, after sporulation (production of spores). • Generally one single sterile frond per plant, but can grow in pairs in O. azoricum, in which the fronds are broader. • Spikes (or fertile fronds) do not invariably appear on every plant.

1 cm

Ophioglossum azoricum & Ophioglossum lusitanicum

Distribution - Ecology Distribution/range (1) Ophioglossum azoricum

Western European species present in France and around the Mediterranean in Spain, Portugal, Sardinia and Italy160. It also occurs locally in Slovakia and in the Czech Republic, the Canaries and Madeira (Azores)300.

Similar species

France

Confusion is easy between the three species of Ophioglossaceae, Ophioglossum azoricum, O. lusitanicum and O. vulgatum. O. vulgatum is the largest (15 to 30 cm) and most widespread. Although present in the Plaine des Maures in proximity to near temporary pools, it is generally found in more eutrophic* habitats, undergrowth or damp grasslands. O. azoricum and O. lusitanicum are closer in size and share habitats that are at first sight similar, but are in fact different in terms of hydrology. The easiest distinction criterion is the autumn-winter growth period for O. lusitanicum, spring for O. azoricum. In early spring the two species can be present at the same time but their stages of development (colour, sporulation) are visibly out of step. Finally, note that the number of chromosomes differs between the three species: O. lusitanicum is diploid* (2n=240 chromosomes), O. vulgatum is tetraploid (2n=480) and O. azoricum hexaploid* (2n= 720), suggesting that this latter species is derived from a cross between the other two.

• Atlantic coast. • Mediterranean region: O. azoricum is present in the Plaine des Maures and the Estérel (Var), in the garrigues of the Hérault, the Cévennes in the Gard and Lozère, and in Les Fenouillèdes (PyrénéesOrientales). In Corsica, this species is only found in rares pozzines* on the Tenda Massif and the Plateau du Coscione142.

60

(2) Ophioglossum lusitanicum

Mediterranean and Sub-Atlantic species present all around the Mediterranean Basin (France, Portugal, Spain, Italy, Yugoslavia, Greece, Crete, Croatia, Algeria, Tunisia, Morocco, Turkey, Lebanon) as well as the Canaries, Madeira and England. France

• Atlantic coast (Béarn, Massif Armoricain and its islands). • Mediterranean region: present in Les Maures and the Estérel, the Biot Massif and Les Albères. Formerly reported from the marshes of Coustière in the Crau. Fairly common in Corsica at low altitude141.

Plant species

Other Mediterranean countries

• Europe: Portugal, Spain, Italy, Yugoslavia, Greece, Crete and Croatia. • North Africa: Algeria (in the Constantinois Tell), Tunisia and Morocco (Middle Atlas, Rif mountains, north and mid-Atlantic plains). • Turkey, Lebanon. Habitat General description

(1) Ophioglossum azoricum The habitats favourable to this species are very wet in winter and spring but rarely flooded for a long period. These are generally depressions, beds or edges of small streams and springtime temporary runnels on well-exposed hillsides on thin soils (Cévennes and Avants-Monts in Languedoc). Its habitat, in the Mediterranean region, is short swards, open or in a mosaic with the shrubby maquis vegetation, on thin, rather acidic, soil. The habitats found in the Atlantic region are coastal swards, dune slacks, sandstone shelves or other types of rock surfaces. On the mainland, the stations are not found above 700 m in altitude but in Corsica they are found up to 1,500 m in peaty basins (pozzines), which remain damp in the summer. The substrate is not very thick, sandy and often acidic. (2) Ophioglossum lusitanicum The habitats are comparable to those of the preceding species (temporarily wet short swards): rock shelves on coastal hillsides, edges of pools and temporary streams, damp depressions on the maquis and small basins in rocky outcrops. However, the species rarely mixes with the preceding species: it seems to seek stations with even thinner substrates, and seems less demanding of a water supply. It is clearly more thermophilic and is even found away from the low-altitude coastal zone. In Morocco, O. lusitanicum is mainly found in the sandy or stony pastureland of the plains and low hills under a Mediterranean mild semi-arid and sub-humid bioclimate. In Algeria, its presence is limited to sandy soils. (1) and (2): Ophioglossum lusitanicum profits from temporary runnels resulting from autumn rains, which are generally more significant than spring rains, whereas the cumulative effect of the supply of water in the depressions seems better adapted to the demands of O. azoricum. On a typical profile through a temporary pool on mainland France (Maures), O. lusitanicum will thus be situated higher than O. azoricum. In Corsica, the two species have very different distributions: O. lusitanicum is found at low altitude, while O. azoricum is only present in some higher-altitude pozzines. Directive Habitat

Phytosociology*

Mediterranean regions • In a European list, Brullo & Minissale54 include Ophioglossum lusitanicum among the characteristic species of the alliance Cicendio-Solenopsion laurentiae (within the order Isoetalia, class Isoeto-Nanojuncetea). In Spain, Rivas Goday319 has described an Ophioglosso-Cicendietum filiformis association. • In crystalline Provence, Loisel215 described O. lusitanicum as characteristic of the association Isoeto-Nasturtietum. • In Languedoc, in the Avants-Monts, O. azoricum is noted as a participant in a hygrophytic variant of a Scillo-Ranunculetum paludosii98. In the Cévennes, O. azoricum forms part of Isoetion communities with Isoetes duriei which have not been described44, 195. • In Roussillon, O. azoricum is associated with various undescribed Isoetion communities212. O. lusitanicum is associated in temporary runnels with I. duriei 272. Other regions • In western France, Ophioglossum azoricum is part of a northAtlantic Ophioglosso azorici-Agrostietum caninae or a continental thermo-Atlantic Ophioglosso azorici-Isoetetum histricis whereas O. lusitanicum is one of the characteristic species of a coastal thermo-Atlantic Ophioglosso lusitanici-Isoetetum histricis98. In the Natura 2000* Habitats Register, this latter community is divided into two: Romuleo columnae-Isoetetum histricis and a Chamaemelo nobilis-Isoetetum histricis153. Ecology Biological characteristics

Life form Perennial species persisting in the summer through a rhizome (geophyte* with rhizome). Adaptive strategy163 Stress-tolerant (of summer drought). Reproduction Production of spores aerial, fusion of gametes in wet conditions. Biological cycle Growth of fronds with flooding (autumn or spring rains according to the species), maturation of spores in winter for Ophioglossum lusitanicum in late spring/early summer for O. azoricum; death of the leaves after sporulation, which coincides with the summer drying-out period for O. azoricum. The phenology* depends on the region. In the Mediterranean region, the cycles can differ from one year to the next depending on the weather conditions. In Morocco, the biological cycle of O. lusitanicum also begins in autumn (November). It terminates in the spring (May) with the withering of the fronds.

“Mediterranean Temporary Pools” (code 3170). CORINE Biotopes

22.34 Southern amphibious communities (Isoetalia). 22.341 Short Mediterranean amphibious swards (Isoetion).

61

Mediterranean temporary pools

O. azoricum

Conservation status

France Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Dec.

The number of stations is noticeably increasing in France, no doubt because of increased knowledge (the plant is either absent, or very poorly described in older works) and improved surveying, even though the habitats in which this species are found are a priori vulnerable. The species has not been found again at RoqueHaute (Hérault).

Germination and vegetative growth Maturation of spores Withering of fronds

O. lusitanicum France Jan.

Feb.

March

April

May

June

July

Aug.

(1) O. azoricum

Sept.

Oct.

Nov.

Dec.

(2) O. lusitanicum Maturation of spores

Germination and vegetative growth

Withering of fronds

Environmental conditions

Hydrology Germination of spores requires a very wet soil, supplied regularly during this phase with oligotrophic* water. These plants withstand occasional flooding. Substrate Acidic, fairly thin

The number of stations is in noticeable decline in France, particularly in the Atlantic zone. In Morocco, the number of stations is fairly stable, although the populations are slightly increasing at the mountainous stations. In Algeria, the species is considered to be rare. Legal status of the species/level of protection IUCN 1997 Bern Convention Habitats Directive National and regional protection

Interspecific competition Heliophilous* species, which does not thrive in the shade of taller species. Impact of perturbations Generally favorable, but overgrazing constitutes a mid-term threat for Moroccan stations.

Conservation - Management

National Red Data Books and Red Lists

- (1) France, national list: Decree of 20/01/1982/Journal Officiel 13/05/1982, amended by Decree of 31/08/1995/ Journal Officiel 17/10/1995 - (2) France, regional lists: • Languedoc Roussillon: Decree of 29/10/1997/Journal Officiel 16/10/1998 • PACA: Decree of 9/05/1994/ Journal Officiel 26/07/1994 - (1) France, national Red Data Book: priority species274

Assessment of populations Conservation problems/threats France

• In mainland France, the two species seem very local; the number of stations is not exactly known. • In Corsica, Ophioglossum lusitanicum is fairly frequent on the coast, O. azoricum is local. Morocco

Ten or so stations* with O. lusitanicum have been censused, and this number could increase with a more systematic exploration of the more inaccessible regions of the Middle Atlas and the Rif. Spain

O. lusitanicum is present in the western half of the country (19 provinces), and in the northeast in the province of Girona, and in the Balearics. O. azoricum is scattered in several provinces in the centre and northeast64. Italy

O. lusitanicum is fairly widespread on the peninsula, but rare on the islands (Sicily, Sardinia)292.

62

Anthropogenic factors

The very specific requirements of the Adder’s-Tongues, in both trophic and hydrological terms, mean that these plants are species which are very sensitive to modifications in the balance of their environment. The localised nature of the stations accentuates their vulnerability. Modification in the hydrology of the sites is a frequent degradation factor. As the habitats occupied by these species are generally not very fertile for agriculture, the most immediate threat remains the development of sites, in particular for all the coastal stations. The Adder’s-Tongues are often established on thin soils which are sensitive to erosion. Excessive numbers of visitors can contribute to degradation at some stations. Natural factors

The closing-up of the habitat by vegetation can be a limiting factor for populations. However, the sites being generally not very productive (skeletal soils), the growth of the competitive vegetation is slow: in particular, populations of Adder’s-Tongues on rocky shelves are in a virtually climax* situation and not vulnerable to this factor. In addition, the competitive vegetation is often sensitive to perturbations (grazing, scrub clearing, etc.).

Plant species

Management and conservation measures Current measures

France • The Conservatoire Botanique National Méditerranéen de Porquerolles is conserving O. lusitanicum spores ex-situ. • In the Plaine des Maures: - Some areas supporting these species belong to the CEEP or the Conservatoire de l’Espace Littoral et des Rivages Lacustres. - Within the framework of the LIFE “Temporary Pools” project, experimental management measures are in place in several sites supporting these species (Péguière pool, Les Aurèdes site) and a site has been acquired in the Bois de Rouquan by the CEEP. - The grazing management and scrub clearance implemented for the Défense Forestière Contre les Incendies can be favourable for these species. - Several stations are included in the Natura 2000* area. Morocco There are no measures for the management of Adder’s-Tongue sites.

• A more exact census of the stations of these inconspicuous plants is also recommended. In most favourable natural habitats, their presence is undoubtedly underestimated. Inclusion of the sites in various inventories (ZNIEFF*, Natura 2000, etc.) is essential. • From time to time, management measures aiming to maintain an open habitat or re-establish a satisfactory hydrological regime can be useful to their conservation, following evaluation.

Bibliography Anonymous, 199912 ; Barbero, 196521, 196722 ; Bissardon & Guibal, 199739 ; Boudrié, 199544 ; Braun-Blanquet, 193548 ; Brullo & Minissale, 199854 ; Castroviejo, 1986-200164 ; Chevassut & Quézel, 195672 ; Conti et al., 199286 ; Danton & Baffray, 199596 ; Fennane et al., 1999134 ; Foucault de, 198898 ; Gamisans & Guyot, 1991142 ; Gamisans & Jeanmonod, 1993141 ; Gaudillat & Haury, 2002153 ; Greuter et al., 1984-1989160 ; Jahandiez & Maire, 1931181 ; Kessler, 2000195 ; Lewin & Escoubeyrou, 1997212 ; Loisel, 1976215 ; Nozeran & Roux, 1958272 ; Olivier et al., 1995274 ; Pignatti, 1982292 ; Prelli, 2001300 ; Prelli & Boudrié, 1992299 ; Quézel, 1998306 ; Rivas Goday, 1970319

Recommendations

• Management of the sites supporting Adder’s-Tongue populations, through statutory measures guaranteeing the maintenance of ecological conditions favourable to their survival, is the best way of conserving the stations. In particular, the implementation of decrees for biotope protection, notably for certain coastal populations, is recommended.

Author: Catard A. Collaborators: Michaud H., G. Paradis, L. Rhazi & N. Yavercovski

63

Mediterranean temporary pools

Pilularia minuta

Durieu ex A. Braun

POLYPODIOPHYTES300 MARSILEACEAE Main synonyms

P. globulifera L. subsp. minuta (A. Braun) Bonnier & Layens French name: Pilulaire délicate Italian name: Pilularia minore English name: Pillwort (name of genus)

0.5 cm

Subspecies

None Description/identification criteria64, 96, 292, 300

• Amphibious plant, perennating by means of its filiform creeping rhizome, just slightly buried under the surface of the ground. • Fronds (“leaves”) green, filiform, cylindrical, upright from 1 to 3 cm in length and arranged singly along the rhizome, the internodes being around 1 cm. • Sporocarps* (“fruits”) globular, from around 0.75 to 1 mm in diameter, densely downy, with short pedicels (the pedicel is recurved at the apex) and dark brown in maturity.

Pilularia minuta

Habitats Directive

“Mediterranean Temporary Pools” (code 3170). CORINE Biotopes

Similar species

Seedlings of rushes, grasses and Isoetes. The distinctive characters of Pilularia minuta are the filiform rhizome very slightly buried in the soil, the globular sporocarps and the filiform leaves.

22.34 Southern amphibious communities (Isoetalia). 22.341 Short Mediterranean amphibious swards (Isoetion). 22.3412 Aquatic Mediterranean Quillwort swards. Companion species

Distribution - Ecology Distribution/range

The species associated with Pilularia minuta are numerous: Isoetes velata, I. setacea, Myosotis sicula, Elatine brochonii, Juncus pygmaeus, J. bufonius, Illecebrum verticillatum, Lythrum thymifolia, L. borysthenicum, L. hyssopifolia, Laurentia michelii, Exaculum pusillum, etc.

Mediterranean species. Phytosociology* France

Hérault, Alpes-Maritimes and Corsica. Other Mediterranean countries

Portugal, Spain (Minorca and probably in Andalusia and Catalonia)64, Italy (Sardinia, Sicily, Rome region), North Africa (Morocco and Algeria), Greece, Cyprus and Turkey.

• Traditional phytosociological system319: communities with Pilularia minuta belonging to the alliance Isoetion Br.-Bl. 1931 of the order Isoetalia Br.-Bl. 1931 em. Rivas-Goday 1970 and the class Isoeto-Nanojuncetea Br.-Bl. & R. Tx. 1943. Braun-Blanquet48 considers P. minuta to be one of the characteristic species of his Isoetetum setaceae or Isoetes setaceaLythrum borysthenicum association (= Peplis hispidula) of RoqueHaute (Hérault).

Habitat General description

Pools with Pilularia minuta are mainly (mainland France, Corsica, Morocco), confined to low-altitude forests or matorrals* dominated, depending on the site, by Cork Oak (Q. suber), Myrtle (M. communis), Broom Heather (E. scoparia), Mastic (P. lentiscus) and Phillyrea angustifolia. Other pools with P. minuta are situated in degraded vegetation dominated by Woody Fleabane (Dittrichia viscosa). The soil there is sandy to sandy-silty and the bedrock sandstone, quartzite, basalt or schist. In Morocco, floooding usually occurs in December-January, solely by rainwater, and drying out in April-May. At Roque-Haute (France) and in Corsica, flooding begins in autumn (November) and ends between April and July, depending on the pools.

64

• More recent phytosociological system98: communities with P. minuta belong to the Antinorio agrostideae-Isoetion velatae alliance (Br.-Bl. 1931) of de Foucault 1988, of the order Isoetalia velatae (Br.-Bl. 1931) of de Foucault 1988 and the class Isoetea velatae (Br.-Bl. & R. Tx. 1943) of de Foucault 1988. Ecology Biological characteristics

Life form Perennial species: hydrophyte*292.

rhizomatous

geophyte*

or

radicant

Plant species

Reproduction Aerial for the maturation of the spores. Biological cycle Germination of spores in very damp or flooded conditions and growth of the fronds in the water in March-April, followed by the formation of sporocarps. In late spring, maturation of the sporocarps and the spores after complete drying out; withering and rapid death of the fronds.

• In Greece, this species occurs in three coastal pools situated on islands in the western and northern Aegean Sea290. • In mainland France, there are a few stations at the pools of the Roque-Haute Nature Reserve and the Béziers plateau (Hérault)45, 300. • In Corsica, around a dozen of stations have been confirmed or recently discovered (pools of the Tour d’Olmeto, Frasseli, Padulellu, Mura dell’Unda, Arasu, and Tre Padule de Suartone Nature Reserve, etc.)172, 219, 222, 223, 224, 302. Conservation status

Morocco Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Dec.

June

July

Aug.

Sept.

Oct.

Nov.

Dec.

Germination and vegetative growth Maturation of spores Withering of fronds France Jan.

Feb.

March

April

May

Germination and vegetative growth Maturation of spores

Populations vary greatly in size from one year to the next, but appear to be surviving at the various stations. • In mainland France it is, however, possible that P. minuta is vulnerable as it is at the northwestern limit of its range. The species has, furthermore, disappeared from the pool at Biot, destroyed in 1975246. • In Algeria, the known stations in the Algiers area and near Oran97 appear to be threatened302, but recent information is lacking.

Withering of fronds

Legal status of the species/level of protection Environmental conditions

Hydrology Germination of the spores requires the saturation or flooding of the substrate. The plant can remain in the vegetative stage under oligotrophic, very shallow water (<5 cm) and form sporocarps, but the complete drying-out of the habitat is essential for the maturation of the sporocarps96. Substrate Non-calcareous substrate at the Moroccan and French stations.

IUCN 1997 Bern Convention Habitats Directive National and regional protection

National Red Data Books and Red Lists

Interspecific competition A heliophilous* species, Pilularia minuta germinates in the clearest parts of the pool. The species is negatively affected by the shade of woody species, herbaceous perennials (Cistus monspeliensis, C. salviifolius, Myrtus communis, Dittrichia viscosa, Ulmus minor, Fraxinus angustifolius, etc.) and helophytes (Scirpus maritimus, Eleocharis palustris, etc.), as well as by mats of charophytes (such as Tolypella glomerata in Corsica) in years with a very rainy winter (as in 2000-2001).

- Strictly protected species (Annexe I) - France, national list: Decree of 20/01/1982/ Journal Officiel 13/05/1982, amended by Decree of 31/08/1995/Journal Officiel 17/10/1995 - France, national Red Data Book: priority species274 - Balearics: “endangered” (EN)331 - Spain, national list: “vulnerable” (VU)13 - Italy, national list: “vulnerable”86 - Morocco: “very rare” species (RR)133, 134, 181, 381 - Greece, national Red Data Book: “vulnerable”(VU) 290

Conservation problems/threats

Impact of perturbations These are favourable (movements of livestock, wild boar, etc.) as they reduce the biomass of larger plants and expose the soil. Adaptive strategy According to Grime163, the species seems to have a stress-tolerant ruderal (S-R) survival strategy

Conservation - Management Assessment of populations

• In Morocco, there are not many pools with Pilularia minuta133, 134, 230: about 12, confined to the Atlantic plains (Benslimane, south of Rommani, southeast of Tiflet).

Anthropogenic factors

Urbanisation, filling in and the modification of the hydrology of pools constitute the greatest threats for this taxon. In Greece, urbanisation is considered to be the main threat290. Natural factors

The closing-up of the habitat, following the growth of tall vegetation, has a negative effect upon this very small species. In France, at Roque-Haute, the abandonment of grazing results in the colonisation of pools by woody species and helophytes. They impede the growth of this species, both directly by their shade, and directly or indirectly through their litter (mechanical and chemical actions). In Morocco, it is mainly the frequency of dry years which limits the appearance each year of this species314.

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Mediterranean temporary pools

Risks relating to populations

Risk of extinction of populations of Pilularia minuta in mainland France, due to their reduced numbers and their isolation. Management and conservation measures

• Promote legal conservation measures for stations which are not yet protected. • Implement the management plan at Roque-Haute, and more generally restore concerted management with natural-heritage objectives on this Nature Reserve.

Current measures

Bibliography Morocco No current management measures. France • Mainland France: at Roque-Haute, on certain sites with Pilularia minuta, the clearing of woody species and the removal of litter to encourage Isoetes setacea were tested within the framework of the LIFE “Temporary Pools” project. However, the conservation of this population, situated on a Nature Reserve, is now uncertain (ban on access, and unilateral management by owners). • Corsica: the creation in December 2000 of the Tre Padule de Suartone Nature Reserve (Bonifacio) should allow management measures to be put in place which are favourable to this species in the four pools of the Reserve. • Collection and conservation ex-situ of the spores by the Conservatoire Botanique National Méditerranéen de Porquerolles. Recommendations

• Control urbanisation. • Limit the plant cover surrounding the stations of Pilularia minuta, to prevent it spreading and to enable the opening-up of the habitat to a certain extent. To attain this objective, the extensive grazing of cattle should be be maintained or encouraged.

66

Anonymous, 199912 ; Anonymous, 200013 ; Bissardon & Guibal, 199739 ; Boudrié et al. 199845 ; Braun-Blanquet, 193548 ; Castroviejo, 1986-200164 ; Conti et al., 199286 ; Danton & Baffray, 199596 ; Daumas et al., 195297 ; Fennane & Ibn Tattou, 1998133 ; Fennane et al., 1999134 ; Foucault de, 198898 ; Grime, 1979163 ; Hébrard, 1990172 ; Jahandiez & Maire, 1931-1934181 ; Lorenzoni & Paradis, 1997219, 2000222 ; Lorenzoni, 1994223, 1996224 ; Maire 1952-1987230 ; Médail et al., 1998246 ; Olivier et al., 1995274 ; Phitos et al., 1995290 ; Pignatti, 1982292 ; Poirion & Vivant, 1969294 ; Prelli, 2001300 ; Quézel & Zevaco, 1964302 ; Quézel, 1998306 ; Rhazi et al., 2001314 ; Rita, 2000317 ; Rivas Goday 1970319 ; Saez & Rossello, 2001331 ; Titolet & Rhazi 1999381 ; Tutin et al. 1964-1993386 ; Walter & Gillett, 1998399. Author: Rhazi L. Collaborators: Michaud H., G. Paradis & M. Rhazi

Plant species

Ranunculus lateriflorus

DC. (1) &

Ranunculus nodiflorus

L (2)

ANGIOSPERMS RANUNCULACEAE Main synonyms

(1) R. nodiflorus L. subsp. lateriflorus (DC.) P. Fourn. (2) R. nodiflorus L. subsp. nodiflorus R. lateriflorus DC. var. charbonelii Rouy & Foucaud French names: (1) Renoncule à fleurs latérales

(2) Renoncule à fleurs nodales Italian name: (1) Ranuncolo a fiori sessili Spanish name: (1) Ranunculo con flores laterales Subspecies

These two taxa are sometimes considered to be two subspecies of Ranunculus nodiflorus.

1 cm

Description/identification criteria

• Annual herbaceous plants (5-20 cm) pale green with slender, fasciculate roots. • Species with erect fistulous* cylindrical stems, sometimes radicant at the lower nodes, dichotomously branching. • Basal leaves petiolate with elliptical or lanceolate-oval blades, sometimes floating and in that case with long petioles; middle and upper leaves with progressively shorter petioles, entire (or slightly sinuate-toothed), lanceolate* to linear-lanceolate. Petiole extended at the base into a whitish membranous sheath. • Flowers very small, from 2.5 to 3 mm in diameter in Ranunculus lateriflorus, smaller (1.5 to 2.5 mm) in R. nodiflorus, solitary, sessile, (lower sometimes with short pedicels), and located in the forks of the stem or between two leaves on the unbranched parts of the main stem. Five pale yellow petals approximately equal to the sepals. • Achenes not deciduous, with surfaces covered in tubercles, very compressed, 1.5 to 2 mm excluding the beak, which approximately equals the main body of the achene in R. lateriflorus and is very short, not more than one-third of it, in R. nodiflorus. • R. lateriflorus is diploid (2n = 16), R. nodiflorus tetraploid (2n = 32). • Further characteristics for distinguishing the two species, given by some authors: in R. lateriflorus the petals are spoon-shaped, in R. nodiflorus they are flat (however, this criterion is difficult to apply objectively to very small petals which are quickly shed, especially as it is variable on a single flower); also, more over of equal size, R. lateriflorus may be more robust than R. nodiflorus.

Ranunculus lateriflorus

Similar species

Other annual Ranunculus species with entire leaves and small to tiny yellow flowers may grow in identical habitats (R. revelieri, R. longipes, R. ophioglossifolius). However, in reasonably welldeveloped individuals there can be no confusion with the two above species, which are the only ones with sessile flowers in the leaf axils and in the forks of the stem branches. The other species have pedunculate flowers.

1 cm

Ranunculus nodiflorus

67

Mediterranean temporary pools

Distribution - Ecology Distribution/range (1) Ranunculus lateriflorus

Paleo-tropical species with a wide but everywhere patchy distribution, around the Mediterranean and east to central Asia and Siberia: France, Italy, Greece, Cyprus, Turkey, Syria, Lebanon, Israel, Algeria, Morocco and Croatia (?). In France: basaltic outcrops in the Roque-Haute area (Hérault), basaltic plateau of Caux-Fontès-Pézenas (Hérault), Plaine du Regard in the south of the basaltic plateau of the Coiron (Ardèche). In Italy, three currently known localities: in Sicily, in the Apennines, and in the Abruzzo86. (2) Ranunculus nodiflorus

Franco-Iberian endemic. In France, the stations are scattered especially through the centre and west of France: Massif des Fenouillèdes (Pyrénées-Orientales), Massif Central in the southern Aveyron and on the basaltic plateaux (Chaux) of the Puy-de-Dôme, the Haute-Loire and the Cantal, the Paris region around Fontainebleau, the Loiret near Orléans, the Indre in the Brenne and the Massif Armoricain. In the Iberian Peninsula (Spain and Portugal) they occur in the centre and the north. Habitat General description

In France: non-calcareous areas that are temporarily flooded in winter and dry out in spring (pools, ditches or nutrient-poor grassland). (1) Ranunculus lateriflorus It grows in fairly deep pools and ditches, even in poor grasslands prone to flooding. At latitudes further south than France, it is found especially in mountains, sometimes on a peaty substrate (pozzines of the High Atlas, pools of the Taurus, Sicily, etc.). Outside France, R. lateriflorus sometimes grows on a non-siliceous substrate218. (2) Ranunculus nodiflorus Outside the Mediterranean area, it shows a preference for flatbottomed basins, very shallow, covered in a clay-gravely silt of maximum depth 3 to 4 cm and scattered around on plateaux where there is much exposed bare rock47, 106. In the rest of its range, this species is found in similar habitats to R. lateriflorus. In Corsica, the plant is rare141 and known only from the mountains in the south of the island, from about 900 to 1,600 m altitude: • In the pozzines* of the Massif de l’Incudine, in the Plateau du Coscione99, 143, 144. • A stream which dries up from June onwards in the Forêt de Marghèse in the Massif de l’Ospedale-Cagna107. Several of the pools where these Ranunculus grow have partly anthropogenic origins: granite quarries in the Pays Bigouden80, basalt quarries at Roque-Haute, stock watering ponds in the Ardèche234.

68

CORINE Biotope

Outside the Mediterranean region (22-11 to 22-13) x 22-32 Oligotrophic to mesotrophic annual communities, topographically low-lying, of plain levels, with Atlantic affinities, of the class Isoeto-Juncetea. Mediterranean region 22.34 Southern amphibious communities (Isoetalia). 22.341 Short Mediterranean amphibious swards (Isoetion). Habitats Directive

Outside the Mediterranean region Community of EC interest “Oligotrophic to mesotrophic annual communities, acidophile, of medium plain to montane levels at altitudes ranging from plains to mountainous, of the IsoetoJuncetea” (code 3130). Mediterranean region “Mediterranean Temporary Pools” (code 3170). Phytosociology*

Class: Isoeto-Nanojuncetea. Order: Isoetalia. Alliances: Cicendion filiformis, Isoetion. Associations*: (1) Ranunculus lateriflorus: • France: Isoetetum setaceae • North Africa: Sedum nevadense-Juncus pygmaeus community304. • Eastern Iberian Peninsula: Ranunculus lateriflorus-Damasonium polyspermum community98. • Sicily: Ranunculo-Antinorietum insularis and MyosuroRanunculetum lateriflori 53, 98, 354. • Taurus (Turkey): R. lateriflorus-Sedum annuum community305. • Eastern Europe: Ranunculo lateriflori-Limoselletum aquaticae54. (2) Ranunculus nodiflorus • Outside the Mediterranean area: Bulliardio-Ranunculetum nodiflori106 (= Crassulo vaillantii-Ranunculetum nodiflori Abbayes 1946) and Ranunculus lateriflorus var. charbonellii (= R. nodiflorus)Sedum villosum community38. • Mediterranean France: undescribed communities, notably with Isoetes setacea, Les Fenouillèdes. • Corsica: Gamisans144, based on three releves from the pozzines in the Plateau du Coscione, described the Ranunculeto-Juncetum bulbosi association, which should be renamed Ranunculo nodifloriJuncetum bulbosi. Companion species

In France, both species are accompanied by suites of species which depend mostly on the area where they are growing: • Languedoc-Roussillon: Isoetes setacea, Lythrum thymifolia, L. borysthenicum, Myosotis sicula, Juncus pygmaeus, Herniaria glabra, Polygonum aviculare subsp. rurivagum, etc. • Ardèche: Crassula vaillantii, Lythrum thymifolia, Polygonum aviculare subsp. rurivagum, Ranunculus hederaceus, etc. • Massif Central: Sedum villosum, Montia minor, Sagina procumbens, Poa annua • Region of Fontainebleau and Massif Armoricain: Crassula vaillantii, Illecebrum verticillatum, Lythrum portula, Polygonum aviculare, Spergularia rubra, Poa annua, Myosotis sicula, etc.

Plant species

• High Atlas (Morocco): Sedum nevadense, Elatine macropoda, Ranunculus batrachioides, etc. • Iberian Peninsula: Sedum nevadense, Pulicaria paludosa, Illecebrum verticillatum, Exaculum pusillum, Lythrum borysthenicum, Juncus pygmaeus, Lythrum thymifolium, Myosotis sicula, etc. • Sicily: Antinoria insularis, Lythrum portula, Mentha pulegium, etc. • Taurus (Turkey): Sedum annuum, Lythrum thymifolia, Eleocharis palustris, etc.

Interspecific competition Very sensitive to competition from perennial species. In Brittany for example, simply root-stripping of the pool vegetation resulted in an increase from 47 R. nodiflorus plants in 1995 to 3000 the following year at a single site80.

Conservation - Management Assessment of populations and conservation status

Ecology (1) Ranunculus nodiflorus Biological characteristics

Life form Annuals (therophytes*). Biological cycle In France, both these Ranunculus germinate during the winter to produce floating leaves. Flowering takes place in spring when the water level is falling (April-June). The flowers quickly produce fruits: flowering and fruiting are almost simultaneous. The plants die and wither from the beginning of summer. R. nodiflorus is preferentially autogamous and its achenes are very buoyant196. This is probably also true of R. lateriflorus which has the same floral characteristics (very small petals, flowers without scent) and similar-sized achenes. The seed bank* of course plays a vital role in both species196. Ranunculus lateriflorus and R. nodiflorus Southern France Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Dec.

In France, this species occurs in the mountains of southern Corsica and a large population is known from Rodès in Les Fenouillèdes. At Montalba-le-Château, a station near the above and known in the 19th century (Castanier in Herbier Oliver, Montpellier University), has not been seen since. There are old reports from the Pyrénées-Atlantique and the Hautes-Alpes. It has been mistakenly reported from Savoie, the Alpes-Maritimes and the Var. Overall, in France, the species is declining due to the disappearance of pools (urban development) and the growth of vegetation following the cessation of grazing274. (2) Ranunculus lateriflorus

There are three remaining localities in France: Roque-Haute, CauxFontès-Pézenas and Coiron. It disappeared from the Costière Nîmoise in the late 1960s or early 1970s following land ownership reorganisation which led to the filling-in of many of the small pools (laquets) which were scattered across this plateau (formerly at the Laquet de l’Oli at Beauvoisin, on Quaternary siliceous gravels). The populations of Aosta and Calabria (Italy) have not been seen recently86.

Germination

Legal status of the species/level of protection

Growth Flowering Fruiting

Environmental conditions

Hydrology Typically amphibious species that require a fairly long period of winter flooding in the spring, extending into March-April for Ranunculus nodiflorus in Brittany106 and to April-May for R. lateriflorus in Mediterranean France. In a relatively wet climate (Massif Armoricain, Paris Basin) the basins occupied by R. nodiflorus are shallow, allowing rapid warming in spring followed by early drying out (des Abbayes, 1947; Bournérias et al., 2001). In a Mediterranean climate, the habitats occupied by R. nodiflorus or R. lateriflorus may be under several tens of centimetres of water and may dry out later. Substrate These are both species of oligotrophic* pools, which attain their optimum on acid mineral substrates: sandy-gravely soil (pH 5.26.5) on the Roque-Haute basalts49, gravely-clayey silts (pH 5.65.8), very thin (optimum 1-2 cm), lying directly over rock in the Massif Armoricain106, granitic sands at Rodès (Fenouillèdes).

IUCN 1997 Bern Convention Habitats Directive National and regional protection

National Red Data Books and Red Lists

- (1) and (2) France, national list: Decree of 20/01/1982/ Journal Officiel 13/05/1982, amended by Decree of 31/08/1995/ Journal Officiel 17/10/1995 - (1) and (2) France, national Red Data Book: priority species274 - (1) Morocco: “rare” (R) 133, 134, 181 - (1) Italy, National Red Data Book: “vulnerable”86

Conservation problems/threats Anthropogenic factors

The depression with Ranunculus lateriflorus at Costière (France, Gard) was filled in during agricultural developments in the 1960s-1970s. The population at the Plateau de Caux-FontèsPézenas survives only in a system of canals used to drain a natural depression that has been taken into cultivation. The R. nodiflorus population at Les Fenouillèdes is threatened by a quarrying proposal.

69

Mediterranean temporary pools

Natural factors

Recommendations

The natural succession of perennial formations suppresses the development of these Ranunculus; however, they appear to be capable of surviving in clearings, even very small ones, within dense vegetation. In addition, the seed bank is of considerable size.

Work towards the acquisition of land at the richest sites (Fenouillèdes, etc.) as well as conservation management by a competent organisation.

Risks relating to populations

Bibliography

Low risk of local extinction due to the size and longevity* of the seed bank. Management and conservation measures Current measures

In France Only the populations of Ranunculus lateriflorus at Roque-Haute are located within a nature reserve which benefits from a management plan. However, the management plan cannot put into practice at this reserve owing to a dispute with the owners. The Conservatoire Botanique National Méditerranéen de Porquerolles has made collections of R. nodiflorus seeds and is preserving them ex situ. In Italy No measures

70

Biche, 188134 ; Billy, 200238 ; Bournérias et al., 200147 ; Brullo & Grillo, 197853 ; Brullo & Minissale, 199854 ; Citoleux et al., 199180 ; Conti et al., 199286 ; Des Abbayes, 1946106 ; Deschâtres, 1991107 ; Fennane & Ibn Tattou, 1998133 ; Fennane et al., 1999134 ; Foucault de, 198898 ; Gamisans & Jeanmonod., 1993141 ; Gamisans, 1970143, 1976144 ; Gaudillat & Haury, 2002153 ; Jahandiez & Maire, 1931-1934181 ; Kirchner et al., 2003196 ; Litardière de, 195599 ; López González 1986218 ; Maire, 1952-1987230 ; Mandin & Hugonnot, 2001234 ; Médail et al., 1998246 ; Olivier et al., 1995274 ; Quézel, 1957304, 1973305 ; Rouy & Foucaud, 1893328 ; Rouy, 1909329 ; Sortino et al., 1977354 ; Tallon, 1967361. Author: Michaud H. Collaborators: Paradis G. & N. Yavercovski.

Plant species

Ranunculus revelieri

Boreau

ANGIOSPERMS RANUNCULACEAE Main synonyms

R. ophioglossifolius Vill. subsp. revelieri (Boreau) P. Fourn. French names: Renoncule de Revelière (Var subspecies:

Renoncule de Rodié) Italian name: Ranuncolo di Reveillière English name: Crowfoot (name of part of genus) Subspecies

Ranunculus revelieri Boreau subsp. rodiei (Litard.) Tutin (Renoncule de Rodié), and R. revelieri Boreau subsp. revelieri (Renoncule de Revelière). These two subspecies are, however, poorly differentiated16, 85 and are not accepted by all botanists141, 109. Description/identification criteria

• Small plant, 10 to 20-30 cm tall, with the stem upright, fistulous* (rarely in rodiei), branched and more or less hollow. • Basal leaves surrounding the stem, with long petioles and oval lanceolate blades. Stem leaves with shorter petioles and more narrowly lanceolate. • Flowers in a pale yellow perianth, with long pedicels. Petals glabrous, shorter than the sepals (very villous on the lower surface in revelieri and glabrous or slightly downy in rodiei). • Fruits: ovoid achenes, blunt, finely grainy, with a very short beak*96, 274, 386.

1 cm

Similar species

Ranunculus fontanus C. Presl and R. ophioglossifolius (Vill.) Boreau have the petals longer than the sepals96. R. ophioglossifolius in addition has noticeably larger flowers than R. revelieri, and its first lowest two leaves are cordate, not lanceolate16, 85. R. ophioglossifolius, which is diploid (2n = 16) would have given rise to R. revelieri which is tetraploid (2n = 32). R. revelieri would thus be an apoendemic*, i.e. a polyploid* taxon no doubt derived, through evolution, from the diploid taxon R. ophioglossifolius85.

Distribution - Ecology Distribution/range

Species endemic* to the Var (Maures and Estérel), Corsica and Sardinia, up to 1,000 m altitude.

Ranunculus revelieri Habitat General description

In Les Maures, Ranunculus r. rodiei is found in the shallow parts of pools, but also in ditches and on the edges of temporary rivulets. In the Plaine de Palayson, this species occurs in clay-silt pools situated in sandy depressions resulting from the erosion of Permian sandstones246 (R. r. rod.). In Corsica, the R. r. revelieri stations are mainly on the edges of temporary pools, as well as depressions and wet meadows16, 85, 108, 109, 222, 285, 292 . Habitats Directive

“Mediterranean Temporary Pools” (code 3170). France

Ranunculus revelieri rodiei in the Var (Maures, Estérel) and R. revelieri revelieri in Corsica.

CORINE Biotopes

22.34 Southern amphibious communities (Isoetalia). 22.341 Short Mediterranean amphibious swards (Isoetion).

Other Mediterranean countries

R. revelieri revelieri in Sardinia (Italy)16, 85, 96, 292.

Phytosociology*

• In the Var, Ranunculus revelieri occurs in an association* of the Isoetion histricis, the Lythrum borysthenicum and R. revelieri subsp. rodiei association18. This association may also be found under the old name “Peplis erecta (= Lythrum borysthenicum) and R. revelieri association”18, 21, 215, 308.

71

Mediterranean temporary pools

• In Corsica, R. revelieri is frequently located among the following perennials: Schoenus nigricans, Plantago lanceolata var. timbali, Oenanthe globulosa, Cynodon dactylon and Carex serrulata. Associated annuals are: Juncus pygmaeus, J. tenageia, Solenopsis laurentia, Cicendia filiformis, Lythrum hyssopifolium, Illecebrum verticillatum (non-submerged form) and Agrostis pourretii. This type of community should not be included in the Isoetion histricis but in the Cicendion.

Interspecific competition For this heliophilous* species, according to Barbero21, competition frequently causes population variations (instability). Experiments carried out within the context of the LIFE “Temporary Pools” project show that in the absence of grazing, competition with perennial herbaceous plants is very detrimental to this species, in particular on soils whose depth is greater than 20 cm.

Ecology

Impact of perturbations There are probably few direct effects on populations (uprooting of individuals), but a positive indirect effect of irregular disturbance of the substrate (stream dynamics, wild boar, etc) which restricts perennial species or improves the hydrological regime.

Biological characteristics

Life form Therophyte* (annual species) with a scape*96, 274, 292.

Conservation - Management

Reproduction Aerial.

Assessment of populations

Seed characteristics96, 386 Length: about 1.5 mm; width: about 1 mm.

In France

Biological cycle Seedlings may appear as soon as the first rain falls in autumn. They subsequently grow very quickly in April and flower in AprilMay. Flowering is advanced if rainfall comes early and delayed in the opposite case. The plant then spends the summer and part of the autumn in the form of seeds. The Lythrum borysthenicum and Ranunculus revelieri association is the most ephemeral among the Isoetion histricis and lasts for a maximum of two or three weeks: the community, submerged for the winter and a high proportion of the spring, sometimes disappears from the end of May, or more commonly in June with the first hot days21, 96.

• Several stations in the Var: Plaine des Maures (abundant, widely scattered), Massif des Maures, Plaine de Palayson (local), Colle du Rouet (very local). • Fifteen to twenty known sites in Corsica in 2001285: Agriate, Massif de Cagna (at 1,030 m altitude), communes of Pianottoli, Figari, Bonifacio and Porto-Vecchio107, 108, 222, 225, 246, 274 and map in Paradis et al.285. In Italy

Several stations in northern Sardinia, in the Limbara mountains, near Macomer, at Molara, and on San Pietro island west of Cagliari (map in Arrigoni16, Conti et al.86). Conservation status

France Jan.

Feb.

March

April

May

June

July

Germination

Aug.

Sept.

Oct.

Nov.

Dec.

Germination

Vegetative growth Flowering Fruiting

Environmental conditions

Hydrology A hygrophytic* species found in shallow water (0-20 cm) but which can survive, in exceptional winter conditions, at depths of up to 40 cm. Germination takes place under water or on wet soil and the plant can tolerate prolonged immersion of the substrate for several months per year, from mid-autumn to the beginning of spring153. Climatic instability causes interannual fluctuations in the abundance of individuals. Ranunculus revelieri can also grow in slow-flowing streams and rivulets18, 153.

In Corsica, the populations appear to be maintaining themselves, but as with many therophytes, few individuals are visible in years when there is very little rainfall in winter and spring (as was the case in 2001-2002). In Provence, populations appear to be stable overall, despite the destruction of some stations.

Legal status of the species/level of protection IUCN 1997399

Bern Convention Habitats Directive National and regional protection

Substrate It usually grows on siliceous soils. National Red Data Books and Red Lists

72

- Ranunculus r. revelieri: “vulnerable” (V) for Corsica (France), and “endangered” (E) for Sardinia (Italy). - Ranunculus r. rodiei: “vulnerable” (V) for France - France, national list: Decree of 20/01/1982/Journal Officiel 13/05/1982, amended by Decree of 31/08/1995/Journal Officiel 17/10/1995 - France, national Red Data Book: priority species274 - Italy, national Red Data Book: “threatened”86

Plant species

Conservation problems/threats Anthropogenic factors

“Cleaning up”, drainage or infilling of the habitat, urban development96, 274. In Provence, several stations have been destroyed directly or indirectly (disruption of hydrology) by tourism developments (golf), transport infrastructure works and urban development. Natural factors

Natural vegetational succession may lead to the habitat closing up and to processes of aggradation by siltation. For example, biotopes suitable for the Lythrum borysthenicum and Ranunculus revelieri association at the Péguière pool (Plaine des Maures) are progressively changing into drier habitats, which are prone to colonisation by other plant associations153, as a result of infilling with sediments from the catchment area. More generally in Les Maures, the disappearance of grazing favours the development of woody plants and perennial herbs (Juncus conglomeratus, Scirpus holoschoenus), to the detriment of R. revelieri. In Corsica, in the absence of disturbance (by cattle grazing, the activities of wild boar and of hares), there is a risk that the R. r. revelieri swards will in future be invaded by maquis species (Myrtus communis, Phillyrea angustifolia, Pistacia lentiscus). Risks relating to populations

At Ranunculus r. rodiei stations, the populations are not visible very year as their development depends closely on the pattern of spring rainfall. Similarly, in Corsica, populations of R. r. revelieri are only visible in years when the winter-spring rainfall is sufficient to enable germination to take place (as was the case in 2000-2001). However, their instability does not imperil the populations. Management and conservation measures Current measures

Corsica The populations at the Tre-Padule de Suartone pools (Bonifacio) benefit from the site’s “Réserve Naturelle” (Nature Reserve) status (Ranunculus r. revelieri).

Mainland France (Maures) • A Nature Reserve project at the centre of the plain should protect the majority of the stations on this site from anthropogenic

threats (R. r. rodiei) 153, 274. • The creation of a Natura 2000* area should enable the necessary requirements for the conservation of this species to be taken into account. Currently, the grazing which has been reintroduced along the firebreaks is favourable for it. • LIFE “Temporary Pools” has enabled experimental habitat management methods to be tested for this species, as well as the acquisition by the CEEP of two sites where it is present (Bois de Rouquan, Vallon de Sauronne). • Seeds have been collected and are preserved ex-situ by the Conservatoire Botanique National Méditerranéen de Porquerolles. Recommendations

In Corsica Managers should ensure that ground disturbance, due for example to extensive cattle grazing and the movements and “rooting” of wild boar (and hares), is continued. In mainland France • Continuation of: - Station monitoring. - Raising the awareness of managers involved, in the context of taking the species into consideration during physical planning. - Control over management by acquisition or by contractual means. • Creation of protected areas wherever the species is present. • Re-establishment of extensive grazing.

Bibliography Anonymous 199912 ; Arrigoni, 198316 ; Aubert & Loisel, 197118 ; Barbero, 196521 ; Bissardon & Guibal, 199739 ; Contandriopoulos, 196285 ; Conti et al., 199286 ; Danton & Baffray, 199596 ; Deschâtres, 1993108 ; Deschâtres et al., 1991109 ; Gamisans & Jeanmonod, 1993141 ; Gaudillat & Haury, 2002153 ; Loisel, 1976215 ; Lorenzoni & Paradis, 2000222 ; Lorenzoni, 1997225 ; Médail et al., 1998246 ; Olivier et al., 1995274 ; Paradis et al., 2002285 ; Pignatti, 1982292 ; Quézel et al., 1979308 ; Tutin et al., 1964-1993386 ; Walter & Gillett, 1998399. Author: Calvière T. Collaborators: Catard A., P. Grillas, G. Paradis & N. Yavercovski

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Mediterranean temporary pools

Genus Riccia L. MARCHANTIOPSIDA RICCIACEAE Main synonyms

None still in use nowadays. French name: Riccia English names: Crystalwort, Riccia

1 mm

Sous-genres

Two sub-genera occur in the Mediterranean area: - Riccia with a single section, Riccia. - Ricciella (A. Braun) Rchb. with two sections: Ricciella and Spongodes Nees. Description/identification criteria

• Small liverworts with gregarious thalli, forming either complete rosettes or more or less connected linear lobes, from a few millimetres to a few centimetres in diameter, with a tough, compact or spongy consistency. • May be annual or perennial* species. • Lobes usually having the dorsal surface longitudinally grooved and swollen at the edges into more or less prominent tumid ridges, varying widely in colour depending on the species and the growing conditions, from pure white to blackish-violet. Edges of the lobes sometimes with hairs, cilia or papillae and with scales which often have a crimson tint. • In species with a compact thallus (subgenus Riccia), dorsal tissue consisting of uniseriate columns of almost contiguous vertical cells; in species with lacunose thalli (subgenus Riciella) dorsal tissue very loose and with a network of air-filled gaps. • Dorsal epidermis pitted with more or less distinct pores. Ventral tissue more or less compact in all species in the genus. • Some species dioecious, others monoecious. • Male gametangia* (antheridia*) embedded in the dorsal tissue of the thallus, more or less projecting on the upper surface in the form of small upright protuberances. • Female gametangia (archegonia) also contained within the thallus and more or less protruding on the upper surface. Once fertilisation has taken place, capsule development within the thallus tissue; at maturity, the capsule may project from the upper or lower surface or not at all. • Spores released when the thallus decomposes at its base, although it may continue to grow at the tip. No elater* among the spores. Spores coloured at maturity, turning from brown to black and very often exhibiting polarity (the distal surface, which is roughly hemispherical, contrasting with the proximal surface which has three triangular facets). Distal surface (less often the proximal surface) usually provided with alveoli demarcated by walls which may have tubercles in their angles (this spore ornamentation is often very important for identifying species, hence it is always worthwhile to look for mature thalli in the field). - Vegetative* reproduction very rare. Introduction to a few species

The genus Riccia includes over thirty species in the Mediterranean area, making it the most widely represented liverwort genus. The species are often fairly difficult to identify in the field, although some of them are easy to recognise with a little experience. They are not all strictly dependent on temporary pool biotopes but may

74

Riccia sorocarpa

colonise any other habitats which are subject to a dry period/wet period alternation. Riccia sorocarpa, R. macrocarpa, R. glauca, R. sommieri, R. subbifurca, R. warnstorfii, R. nigrella, R. michelii, R. beyrichiana, R. crozalsii, R. crystallina and R. canaliculata for example are fairly typical of the surroundings of temporary pools. For species identification see among others the many works of Jovet-Ast193 and of Schuster349.

Distribution - Ecology Distribution/range France

The genus Riccia is particularly well represented in the Mediterranean Midi but many species have a wider distribution. Various species with Mediterranean affinities are found along the Atlantic seaboard (Riccia crozalsii, R. michelii, etc.) or in the Massif Central (R. trichocarpa). Many taxa are rare and only occur at a very limited number of stations. For example, the Massif des Maures, the Biot area and the Nature Reserve de RoqueHaute are sites that are extremely rich in liverworts of the genus Riccia. Other Mediterranean countries

All the countries around the Mediterranean are rich in Riccia, although many of them have a very uneven distribution at this scale. For example, Riccia macrocarpa is much commoner in the west of the Mediterranean Basin than in the east; R. sommieri is only found in west of Mediterranean Basin; R. atromarginata is much less rarer in North Africa than in the north of the Mediterranean Basin, etc. Habitat General description

Suitable habitats include, for example, short and pioneer grasslands with many gaps in the vegetation cover, thoroughly wetted in winter and spring and drying out completely in summer. The physical characteristics of the substrate are important. Riccia need a substrate which is firstly stable and relatively consolidated and secondly has a certain capacity to retain water. Sands, silts and clays (or mixed substrates) are the most suitable substrates in this respect. Biotopes with Riccia are often located close to temporary pools but not actually at the lowest topographical levels. Generally

Plant species

speaking, it may be said that the Riccia of temporary pools and streams are linked with associations* of medium topographic levels. They are concentrated mainly among populations that are kept damp by capillary action or that emerge from the water early in the year, for example Isoetes duriei populations. Higher plant species that are usually associated are mostly small annuals (for example the dwarf Juncus, Radiola linoides, Laurentia michelii, Aira capillaris, Crassula vaillantii, Lythrum thymifolium, Lotus angustissimus, etc.).

Ecology

Riccia are temporarily hygrophytic*, heliophilous*, thermophilic and terricolous species. However, the individual species’ ecological requirements show wide variations with respect to this general pattern. There are species for which the chemical characteristics of the substrate are immaterial while others are distinctly calcicolous or silicicolous. Biological characteristics

Habitats Directive

Communities rich in Riccia may on the whole be included in “Temporary Mediterranean Pools” (3170) or in “Very weakly mineralised oligotrophic waters of sandy plains in the Western Mediterranean with Isoetes “ (3120), in a very general way. It should be added that the phytosociological status of liverworts such as Riccia within vascular-plant communities is generally complex and cannot be simply expressed by means of these codes.

Life form Generally hepatico-therophytic. Annual or perennial species. The species considered to be annual may also revive after the dry season with the resumption of activity in the apical meristem or sometimes the whole thallus. It is possible to observe a resumption of activity of Riccia meristems that have been kept in a herbarium for over ten years, which is a good illustration of their powers of regeneration!

CORINE Biotopes

These communities with Riccia may be placed in: 22.34 “Southern amphibious communities” (Isoetalia), in particular, 22.341 Short Mediterranean amphibious swards (Isoetion). Phytosociology*

The relationships between liverworts and the vascular-plant communities are often complex. Liverworts and bryophytes* are in general very ephemeral and delicate plants whose populations (and thus the communities which they form) are subject to great variability in space and time. The communities are often patchy at the scale of a single station, where their appearance is frequently associated with openings in the herb layer. Some authors21, 319 have considered the Riccia to be characteristic of the overall Isoetion alliance. Barbéro21 and then Hébrard171 for their part emphasise the closeness of the sociological links which join the communities with liverworts to the vascular-plant communities Isoetion, but they also point out that it is very difficult to give an opinion on the true phytosociological affinities of the bryophytes. In addition, Hébrard171 mentions cases of very clear extensions of bryological communities typical of temporarily wet biotopes into Cistus formations, when the soil humidity is sufficiently high. The Campylopetum introflexi Hébrard171 is an association which unites the bryophytes of the rocky seepage surfaces of crystalline Provence, in which Riccia are considered to be companion species. The Riccietum crustatae is another circum-Mediterranean association, well represented in Spain, characteristic of wet, salty soils which dry out severely in summer326. However, most Riccia cannot tolerate the presence of salt in the substrate. It may be seen that the phytosociological characterisation of bryological communities involving Riccia is only in its early stages and that much descriptive work still needs to be carried out.

Adaptive strategy (sensu During126) • Type “a”, corresponding to “annual shuttle”: lifespan less than one year, spores large and produced in fairly small numbers; major reproductive investment. • More rarely type “s”, corresponding to “short lived shuttle species” (maximum of a few years): short-lived perennials with high investment in reproduction and few fairly large spores. Reproduction Fertilisation requires a film of water on the surface of the thalli or even the substrate; spore maturation takes place during the wet season and may continue into the beginning of the dry season. Spore production is therefore aerial. Spores They are usually fairly large (of the order 100 µm) and heavy, and are dispersed mostly through the movements of animals (birds, mammals, humans, etc.) but also by flowing water in some cases. Dispersal by anemochory*349 appears to be less important and more unpredictable. In most cases the spores are ornamented and this could facilitate their attachment to vectors, although this hypothesis has not been experimentally confirmed. Fragments of thallus or propagules* (very rare) may also be carried by animals. Biological cycle Riccia are capable of carrying out the whole of their biological cycle very rapidly following the appearance of water in the substrate (rain, runoff) or merely on the thallus (condensation, nocturnal dewfall, etc.), enabling them to avoid the driest period. They are therefore able to tolerate the stresses of the driest months to a certain degree through the suppression of any physiological activity combined with an extreme longitudinal folding of the thallus, and by the production of spores that have very tough walls and that retain their germinative capacity for a very long time. The active phase begins with the germination of the spores, which takes place as soon as the first autumn rain starts to fall. The growth of the thalli continues throughout the whole of the wet season. Gamete production may take place very soon after germination (a few weeks for the fastest species), but may also continue until the beginning of the dry season of the following

75

Mediterranean temporary pools

year. The spores are released by the decomposition of the old parts of the thalli as growth continues at the tips of the lobes. The complete desiccation of the substrate and the thalli causes the virtually complete disappearance of Riccia and the release of all the spores produced; the latent phase then begins, with the survival in the substrate of the spores, which may enter into a dormant state.

Conservation status

The conservation status of Riccia populations is intimately linked with the general health of the temporarily wet biotopes. These liverworts have undergone a major decline following the disappearance of many temporary pool biotopes. In Malta, R. melitensis was known from one locality, where it has never been seen again since its discovery344. Legal status of the species/level of protection

Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Dec.

Germination and vegetative growth Fertilisation and maturation of spores Release of spores Latent spores and thalli

IUCN 1997 Bern Convention Habitats Directive National and regional protection National Red Data Books and Red Lists

Environmental conditions

Hydrology A very wet substrate is necessary at the beginning of the cycle but, in most species, flooding must not be too prolonged. Substrate More or less compacted substrates are definitely the most suitable, whether they are sandy, clayey or silty. Terra Rossa*, more or less muddy alluvia, etc., also provide potential sites. Interspecific competition All the species are pioneers and struggle to survive in the presence of competition from other more colonist bryophytes and especially perennial vascular plants, which will eliminate them fairly readily. Impact of perturbations The main perturbations (“ploughing” of the substrate by wild boar, trampling by cattle and humans) may have a positive effect, due to the elimination of taller plants, the destruction of the litter from these plants and the opening of the herb layer through the creation of bare patches (“tonsures”) which are very suitable for Riccia. Temperature All the species are thermophiles. Light All the species are heliophiles, but very slight shading which moderates the intensity of the solar radiation may be a positive factor in the hottest regions of the Mediterranean Basin.

- Europe: Red Data Book for bryophytes128: Riccia crustata: “endangered” (E) Riccia frostii: “rare” (R) R. huebeneriana: “rare” (R) R. ligula: “rare” (R) R. melitensis: “extinct” (K) R. perennis: “rare”(R) R. sommieri: “rare” (R) R. trabutiana: “rare” (R) - France, Red Data Book for bryoflora (proposed)105: Riccia crustata R. ligula

Further species in this genus are locally rare to very rare, but are not included on the protected lists as they are little known. Conservation problems/threats Anthropogenic factors

All the threats with which temporary pool and stream biotopes are faced in the area around the Mediterranean consequently also threaten populations of liverworts of the genus Riccia. Urbanisation, infilling with rubble, drainage and conversion to agriculture, etc. lead to the irreversible loss of “Ricciological” natural heritage. Natural factors

Conservation - Management Assessment of populations

Practically all temporary pools and streams in France are colonised by Riccia. Some rare species are, however, very local. The temporary pools and streams of the countries surrounding the Mediterranean are equally rich in Riccia. The number of sites is impossible to estimate but is probably very high. Spain, Algeria and Morocco are particularly rich in liverworts of this genus.

76

The absence of any factors that will disturb the vegetation, and the closing-up of the natural habitat, will eventually cause the disappearance of pioneer liverworts. The loss of the temporary character of pools, through infilling or through disruption of the hydrological network, is very deleterious to Riccia as this allows the spread of common and much more competitive species of hygrophytic bryophytes (usually pleurocarp* species). However, the presence of dormant spores in the substrate is a positive factor in the re-establishment of populations following a reopening of the habitat.

Plant species

Extinction risks relating to populations

Major interchanges probably take place between neighbouring or widely separated sites, but there are no experimental results to support this assumption. Management and conservation measures

• Promote the perpetuation of extensive agricultural practices which create moderate disturbance, or management activities having the same rationale. Extensive grazing may be favourable at stations where the vascular vegetation has developed too much. Inputs of excess organic matter should, however, be monitored.

Current measures

The sites at Roque-Haute and the Massif and Plaine des Maures are included within Natura 2000* sites (no. Fr 9101430 and no. Fr 9301622 respectively) which should allow particular attention to be paid to the conservation of the temporary pools and their associated species. The Roque-Haute site also has legally protected “Réserve Naturelle (Nature Reserve)” status, which is currently unenforced due to the ban on access imposed on the manager by the owners. Recommendations

Bibliography Anonymous, 199912 ; Barbero, 196521 ; Bissardon & Guibal, 199739 ; Deperiers-Robbe, 2000105 ; Dierssen, 2001118 ; During, 1979126 ; ECCB, 1995128 ; Hébrard, 1970171 ; Jovet-Ast, 1986193 ; Rivas-Goday, 1970319 ; Ros & Guerra, 1987326 ; Schembri & Sultana, 1989344 ; Schuster, 1992349. Author: Hugonnot V. Collaborator: Hébrard J. P.

• Promote a better knowledge of the distribution of the species (surveys and inventories) and raise the awareness of managers. • Maintain the hydrological network in an intact condition (hydrological regime and water quality).

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Mediterranean temporary pools

Riella helicophylla

(Bory & Mont.) Mont.

MARCHANTIOPSIDA RIELLACEAE 0.4 cm

Main synonyms

Duriaea helicophylla Bory & Mont. (basionyme*) French name: Riella à thalle hélicoïde English names: Liverwort, Riella (genus names) Subspecies

None Description/identification criteria

• Aromatic aquatic liverwort (scent fairly like coriander, sometimes strong), measuring up to 3 cm high but often much smaller, annual, soft-tissued, formed from an upright thallus composed of a main branch with a more or less undulate single-layer membranous wing (a single layer of cells) on one side and small lanceolate scales on the other. Wing more or less helicoid, very fragile, and usually reduced to barely recognisable fragments due to various disruptions and to predation by phytophagous invertebrates. • Thallus fixed into the substrate by many rhizoids* growing from the base (plant appearing to be placed gently on the substrate). • Dioecious species whose male plants bear very small antheridia*, arranged in a row on the edge of the membranous wing of the thallus. • Female plants at maturity bearing bottle-shaped capsules* at the tip of the thallus, pedicellate but lacking hairs, indehiscent, protected by a papillose, membranous involucre without a wing (characteristic of the subgenus Riella). • Spores, released when the wall of the capsule decomposes, measuring 70 to 95 µm in diameter, with surface smooth to slightly granulous, bristling with many large “spines” (6 to 10 µm long) which are truncated and widen at the tip. No elaters* among the spores. The sporophyte (capsule), which is normally attached to the gametophyte*, may become quite independent of it (the thallus), when unfavourable conditions (drought, predation, etc.) result in its premature disappearance. The sporophyte then continues to develop in a completely independent way, which is very unusual, if not unique, among bryophytes*. The formation of single-layered propagules with indeterminate growth, arising from the apical meristem, has been described. These propagules, in contrast to spores, do not have a dormant stage and are therefore capable of growing immediately after their release.

20 µm male spore

female

Riella helicophylla

The species of the subgenus Trabutiella (Riella affinis M. Howe & Underw. and R. cossoniana Trab.) are fairly easily distinguished from R. helicophylla by their longitudinally winged involucre. R. notarisii (Mont.) Mont. is monoecious (often a difficult character to assess) and its spores have a rough surface and shorter spines (4 to 5 µm). R. parisii Gottsche has much smaller spores (about 60 µm), with the spines shorter (5 to 6 µm) and not widening at the tip. R. numidica Trab. also has small spores of about 60 to 70 µm. R. bialata Trab. has the wing divided into two. The species of the genus Riella are relatively poorly known from the taxonomic point of view. Great confusion reigns in the nomenclature regarding various taxa*. A complete and worldwide review of the genus appears to be essential to clarify the many points which remain uncertain.

Distribution - Ecology Distribution/range

Similar species

Confusion is possible in the field with all the other species in the genus. Identification can only be carried out by a thorough examination using appropriate optical equipment. Sterile material is practically impossible to identify to species. Mature sporophytes must therefore be looked for in the field amidst the population. In addition, mixtures of species are relatively common.

78

France

Species endemic* to the Mediterranean Basin. Extremely rare species in France, only in the Hérault, known from two localities, close together: • The Notre-Dame de l’Agenouillade site (Agde), discovered recently in 2001353; it is currently still present at this site.

Plant species

• Station by the Etang de Thau (Marseillan), discovered in 1966124, and very probably disappeared following the siting of a huge refuse tip on the exact site.

“Large shallow inlets and bays”. Insofar as these codes did not originally take the associated bryophyte communities into account, the affinities with one or other suggested code should be explained in detail, so as to avoid any ambiguity or confusion.

Other Mediterranean countries

Species occurs in the Iberian Peninsula (Spain and Portugal) and the Balearics, Malta206, quite common in the Maghreb (Algeria, Tunisia), and present in the Near East (Israel).

CORINE Biotopes

The most appropriate CORINE codes appear to be: • For freshwater communities: 22.34 Mediterranean amphibious communities.

Habitat General description

The ecological characteristics necessary for the development of Riella helicophylla populations are: • A temporary water body (prolonged dry period). • Clear, relatively shallow water. • Soil that is bare or with sparse plant cover. The most favourable habitats corresponding to these criteria are therefore of two types: • Temporary pools, lakes, estuaries etc. with saline to supersaline or brackish water (France, Spain, North Africa). • Possibly, temporary freshwater pools (France). The muddy, sandy or clayey substrate is often base*-rich (pH* ranging from about 7 to 8.5). The depth of the water seems to be very important: often very shallow (a few centimetres) and a maximum of 1 metre. In Spain, Riella helicophylla has been collected from sites with a wide range of salinity and chemical composition. The salinity (total dissolved solids) varies from about two to almost 190 g/l, though the species grows best in water with values typically greater than 10g/l77, 78, 189. Continuous evaporation at some sites leads to the formation of saline crusts on the surface. R. helicophylla appears particularly to prefer waters containing sodium chloride although it also colonises those with magnesium sulphate. The waters in which it grows are subject to considerable variations in salinity over a single year and also from one year to another, depending on rainfall, runoff, evaporation, etc. Companion species

The species most frequently associated with Riella helicophylla are other species of the genus Riella (in particular R. cossoniana and R. notarisii), aquatic vascular plants (Ruppia spp., Althenia spp., Potamogeton spp.,) Characeae (Chara galioides, C. canescens, etc., Tolypella spp., Lamprothamnium spp.), rarely bryophytes (Leptodictyum riparium, Drepanocladus aduncus, etc.), Green Algae (Ulothrix spp., Enteromorpha spp., etc.), Cyanophyceae (Microcoleus, Lyngbya, etc.) and sulphur bacteria. When the substrate becomes completely dried out, the vegetation then changes its appearance, with the frequent arrival of a suite of more or less xerophytic vascular and bryophyte species. Note that Riella only really disappears when the substrate is completely dry.

• For halophilic vegetation types: 11.4 Vegetation beds of brackish waters. Phytosociology*

The Riella helicophylla communities are these days included within the class Rielletea helicophyllae, order Rielletalia helicophyllae, alliance Riellion helicophyllae, association* Rielletum helicophyllae78. These vegetation types represent a transitory cover of species that are highly specialised for the colonisation of a very restrictive temporary biotope, at the stage where other communities, dominated by various groups of species (Charetea, Potametea, Ruppietea, Isoeto-Nanojuncetea, etc.), are absent or at an embryonic stage of development. Ecology

It should be borne in mind that Riella helicophylla is a basiphilous*, halotolerant* species, capable of withstanding hypersaline water; a heliophilous pioneer. Biological characteristics

Life form Hepatico-hydrotherophyte. Annual species (capable of living for more than one year in culture if it is permanently submerged). Adaptive strategy (sensu During126) Type “a”, i.e. “annual shuttle” species (= lifespan less than one year, spores large and produced in fairly small quantities; major reproductive investment). Reproduction Maturation of spores aquatic then aerial; release aerial. Spores exhibiting dormancy and capable of retaining their germinative capacity for at least three months. Germination It is at its maximum with low salt concentrations, whereas the thallus grows better at higher salinities. Dispersal of spores Dispersal by water birds (waders, web-footed species), sometimes migratory, by external or internal ornithochory*. This mode of dispersal is assisted by the very large number of spines on the spores. anemochory* appears never to have been described.

Habitats Directive

The simplest approach is to include the Riella community under code 3170, with the overall title “Mediterranean Temporary Pools” for the freshwater communities. On the other hand, the subhalophilic and halophilic* communities could be included under code 1150, with the title “Lagoons”, or even code 1160,

Biological cycle Germination of the spores takes place after the first autumn rain, followed by the slow growth of the gametophyte during winter and the beginning of spring. The spores are released in about May and constitute the resistive stage during the summer

79

Mediterranean temporary pools

drought. When the ecological conditions are favourable, it is possible to see veritable population explosions of this species, which then excludes other species and attains coverage rates of close to 100%. Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Dec.





Legal status of the species/level of protection IUCN 1997 Bern Convention Habitats Directive National and regional protection

Germination and

vegetative growth

National Red Data Books and Red Lists

Fertilisation and maturation of spores Release of spores Spores dormant

Strictly protected species (Annexe I) Annexe II Malta: protected species since 1993206 - Europe, Red Data Book for Bryophytes128 “endangered” (E) - France, national Red Data Book (proposed)105

Environmental conditions Conservation problems/threats

Hydrology • The optimal water depth ranges between a few centimetres and a few tens of centimetres. • The survival of the species is closely dependent on the annual flooding/drying regime. A disruption of the hydrological regime, a delay in the arrival of the rains or too low a level of annual rainfall may considerably restrict or even prevent the development of the species one or several years in succession (temporary disappearances). • Water quality is important, especially salinity. Riella helicophylla appears to be not at all obligatorily halotolerant* but, being a very uncompetitive species, it finds in salt water the conditions which allow it to exist without or almost without any competition. • The species can tolerate a certain degree of eutrophication* of the substrate and of the water. Impact of perturbations Perturbations due to the weather (rainstorms, frosts, etc.), anthropogenic disruption (trampling, ploughing, etc.), or zoogenic disturbances (grazing, disturbance of the substrate, etc.) may be favourable to the extent that they restore pioneer surfaces, provided they take place during the dry period (i.e. after the maturation of the spores). Interspecific competition Natural succession inevitably leads to the disappearance of pioneer Riella helicophylla communities under increasing pressure from vascular plants and algae, whose increasing biomass* and ground cover result in greater competition for light and other essential resources. R. helicophylla does not survive in competition with other plants.

Anthropogenic factors

Urban development around towns such as Agde, Tunis or Algiers constitutes a key threat. The filling in of pools with debris needs to be monitored, particularly at the Notre-Dame de l’Agenouillade site (Hérault, France). The illegal dumping of rubbish and largescale official tips are leading to the irreversible disappearance of the species. Natural factors

Natural vegetational succession leads inexorably to the spontaneous disappearance of pioneer taxa* such as Riella helicophylla, by increased competition within the vegetation mat. In addition, tall plants on the edges of pools (Tamarix sp., Phyllirea sp., etc.) are unfavourable for visiting birds (especially Anatidae) which are capable of carrying spores from one site to another. This reduction in the “accessibility” of some sites is clearly detrimental to the survival of viable metapopulations*. In addition, a high frequency of dry years may explain the species’ decrease or even extinction at some of these stations. A minimum of wetness suffices to trigger the germination of dormant spores but a sufficient amount of water is absolutely necessary to complete the cycle and to produce spores for a new generation. Several dry years may completely deplete a fairly small stock of spores. Risks relating to populations

The risk of extinction is high, given the low numbers at the Notre-Dame de l’Agenouillade station (Hérault, France). The isolation of the French populations compared with those of North Africa and Spain, and their limited area, are a further risk factor. Management and conservation measures Current measures

Conservation - Management Assessment of populations and conservation status

A single site in France; many sites in Spain, distributed among 13 provinces79; several sites in North Africa. Populations appear to be stable or even increasing in Spain and the Maghreb, but are highly threatened in France due to the pressure of urban development in the Agde area.

80

No specific management measures appear to be taking place at sites supporting Riella helicophylla. In France, the Notre-Dame de l’Agenouillade site (Hérault), is included: • In the European LIFE “Temporary Pools” project, which has enabled proceedings for obtaining control over land ownership to be undertaken, hydrological monitoring to be carried out and protocols for monitoring the pools’ vegetation to be drawn up. • In a Natura 2000* area (site Fr 9101416), which should enable the environmental conditions favourable for the conservation of this species to be maintained.

Plant species

Recommendations

• A major survey exercise should be organised in order to locate any possible new population centres, especially in France. • Any management measures which have the aim of maintaining or restoring the pioneer character of the pools and their surroundings, such as controlled grazing or vegetation clearing, are to be encouraged, especially at sites where these activities have been discontinued. • Complete protection of the biotopes (control over land ownership, etc.) should be achieved in order to prevent any destructive urban development projects. • Dumping of rubbish, infilling, and illegal tourism-related activities should be strictly forbidden.

Bibliography Aboucaya et al., 20022 ; Anonymous, 199912 ; Bissardon & Guibal, 199739 ; Cirujano et al., 198877 ; Cirujano et al., 199279 ; Cirujano et al., 199378 ; Deperiers-Robbe, 2000105 ; Dierssen, 2001118 ; Dubois & Hébant, 1968124 ; During, 1979126 ; ECCB, 1995128 ; Jelenc, 1955189, 1957190 ; Lanfranco & Lanfranco, 1999206 ; Proctor, 1961301 ; Skrzypczak, 2001353 ; Trabut, 1891383 ; Trabut, 1942384. Author: Hugonnot V. Collaborator: Hébrard J. P.

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Mediterranean temporary pools

Teucrium aristatum

Perez Lara

ANGIOSPERMS LAMIACEAE Main synonyms

T. cravense Molinier & Tallon French names: Germandrée de Crau, Germandrée aristée English name: Germander (genus name) Subspecies

None Description/identification criteria

• Small annual plant 10 to 20 cm, stem slender taproot. • Stem square in cross-section, upright, usually branching almost from the base; branches erect-spreading. • Leaves pinnatilobed*, with lobes broad, short and blunt, shed from the time of flowering. • Flowers geminate, with short peduncles. Calyx lobes erect, spreading, terminated by a beard. Corolla villous externally, lavender blue with a dark-blue V-shaped mark on the inner base of the middle anterior lobe; posterior lobes streaked with crimson internally. • Fruits: each flower produces four achenes after fertilisation. Similar species

1 cm

Teucrium campanulatum L. is a species of wet clay soils and flood-prone depressions in North Africa, Spain and southern Italy: it is a perennial* plant with the stems often radicant (and not an annual with upright stems), with a shorter and more flared calyx than in T. aristatum, and with whitish, not lavender blue, flowers.

Teucrium aristatum

Distribution - Ecology CORINE Biotopes Distribution/range

22.34 Southern amphibious communities (Isoetalia)

Endemic species of Iberia and Provence.

Phytosociology*

France

Crau d’Arles in the Bouches-du-Rhône. Other Mediterranean countries

Only in Spain, where it was formerly known (more than 100 years ago) in three provinces in the southwest (Huelva, Cadiz and Seville)177 (Medina, pers. com.), and where it was rediscovered in 2002 in the centre of the country (Morales, pers. com.).

In the Crau the vegetation of the Lanau pool is allied to a specific association*215, 252: Order Nanocyperetalia Alliance Lythrion tribracteati Association: Lythrum tribracteatum-Teucrium aristatum215, 252 At this site, the species also forms part of the Deschampsion mediae alliance. Companion species

Habitat General description

In France, this Germander occurs on the edges of a clayey basin created in the Crau pudding stone, whose water is slightly calcareous252. Habitats Directive

Species associated with Teucrium aristatum in France are: Lythrum tribracteatum, Mentha pulegium, Achillea ageratum, Deschampsia media, Herniaria glabra, Polygonum aviculare subsp. depressum, Damasonium polyspermum, Lythrum thymifolium. Ecology Biological characteristics

“Mediterranean Temporary Pools” (code 3170). Life form Therophyte* (annual species).

82

Plant species

Reproduction Aerial: the floral organs as well as the fruits develop out of water. Seeds Achenes oboval-obovate (2 mm x 1.1 mm), glandular, rounded, and with dense very short curved hairs at the tips. Biological cycle Germination of seeds after the water has receded, on very wet soil (in May in France). Flowering occurs at the end of spring (June) on a dry substrate, followed by the ripening of the fruits and the dispersal of seeds during summer.

Spain

The species is very rare here, with a very limited range (in the provinces of Cadiz, Seville and Huelva), but a few new stations were discovered in 2002 in the centre of the country. Legal status of the species/level of protection IUCN 1997

Bern Convention Habitats Directive National and regional protection

France Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Dec.

National Red Data Books and Red Lists

Germination and vegetative growth

World: “rare” (R) - France: “endangered” (E) - Spain: “rare” (R) - France, national list: Decree of 20/01/1982/Journal Officiel 13/05/1982, amended by Decree of 31/08/1995/ Journal Officiel 17/10/1995 - France, national Red Data Book: priority species274

Flowering Fruiting and ripening of fruits ?

?

Dispersal of seeds

Conservation problems/threats Anthropogenic factors Environmental conditions

Hydrology This Germander is above all a species of the edges of pools, which needs a brief period of submersion to develop. It is probable that the saturation of the substrate with water at a given period (end of April or May) triggers germination. Substrate In France, silt-clay soils, poor in calcium and oligotrophic*. Interspecific competition Very sensitive to competition with perennial herbaceous plants.

The acquisition by the CEEP of the site supporting the only French population has enabled the pool to be saved from possible threats of habitat destruction, and management favourable to this species to be introduced. Natural factors

This species is very sensitive to competition with perennial species that produce dense cover (Carex divisa, Agrostis stolonifera, etc.). Towards the end of the 1990s, it had disappeared from a part of the Lanau pool where the cessation of grazing had facilitated a massive growth of perennial grasses, whereas it always maintained itself in the grazed part of the pool. Risks relating to populations

Impact of perturbations Disturbance (grazing, trampling) is favourable to this heliophilous* species since it controls the cover from perennial grasses. At the higher parts of the Lanau pool, the Germander is most often seen on patches of bare soil: roosting places, cattle tracks, holes made by hoofs, and former ditches, where it forms small dense populations.

Conservation-Management Assessment of populations and conservation status France

A single site is known, Lanau pool in the Crau (Bouches-duRhône). Although it undergoes wide interannual fluctuations, this population has been seen regularly since its discovery in 1946, and is not in danger at present. The temporary reduction of grazing at this site seems to have noticeably reduced the size of the visible population, which is in the process of re-establishing itself.

The unique nature of the population (very isolated) entails a high potential risk of extinction at the site. Management and conservation measures Current measures

France • The CEEP became owner and manager of the site in 1998, thanks to financial support from the “Réseau Ferré de France”, in the context of compensatory measures linked with the routing of the LGV Méditerranée (High Speed Rail Link). • The Conservatoire Botanique National Méditerranéen de Porquerolles carries out regular monitoring of the Germander populations, and of the other rare species at the site. • Seeds have been collected and preserved ex situ by the Conservatoire Botanique. • A grazing management experiment, carried out by the Station Biologique de la Tour du Valat, has been running since 2001. The initial results show a substantial spread of Teucrium aristatum in the grazed areas in 2002 and 2003 (see Vol. 1, box 49).

83

Mediterranean temporary pools

Spain The species and the sites which support it are not subject to any specific measures for protection or management (Medina, pers. com.) Recommendations

France • Study the hydrological regime of the site and prevent any disruption of it. • Maintain grazing (formerly sheep, nowadays cattle) which is favourable to the development of annual species like Teucrium aristatum. • Study the population biology of T. aristatum, in particular the key spring recruitment phase (ecology of germination) and the effects of natural and anthropogenic perturbations.

84

Bibliography Devesa, 1987110 ; Loisel, 1976215 ; Maire et al., 1947232, 233 ; Molinier & Tallon, 1947252 ; Molinier & Tallon, 1950-1951255 ; Olivier et al., 1995274 ; Tutin & Wood, 1972385 ; Vivant, 1980397 ; Walter & Gilett, 1998399. Author: Michaud H. Collaborators: Grillas P. & N. Yavercovski

Macrocrustaceans

Imnadia yeyetta

Hertzog, 1935 2 mm

Class: BRANCHIOPODS Order: SPINICAUDATA Family: LIMNADIIDAE

male

I. banatic Marincek & Petrov, 1982 I. cristata Marincek, 1972 I. panonica Marincek & Petrov, 1984 I. voitestii Botnariuc & Orghidan, 194143 egg (150-190 µm)

French name: none English name: Clam Shrimp (general name for Spinicaudata)

Imnadia yeyetta

Subspecies/variability

No valid subspecies. Intraspecific variability more or less marked43. In Yugoslavia, Marincek239 and Marincek & Petrov237 have included several species as being synonymous with I. yeyetta.

Habitat General description

Description/identification criteria

The Spinicaudata have a bivalve carapace that is generally striated, laterally compressed and protecting the body. But unlike numerous Spinicaudata such as Cyzicus, Leptestheria, etc., the bivalve carapace of Imnadia yeyetta is not striated over its entire surface: the upper part of the valves is smooth, around the umbo, while their outer edges are finely striated. Sexual dimorphism is not very marked: the males have 17 pairs of legs whereas the females have only 14; the males have legs P1 and P2 prehensile, armed with “claspers”, enabling them to clasp the female by the edge of the carapace during mating; the females have the pairs P9 and P10 modified into adhesive cylinder-like structures to agglutinate the eggs between their body and their carapace. The carapace, is yellowish to greenish white in colour, slightly transparent, and measures a maximum of 11 to 12 mm. The eyes are sessile. Similar species

The only other species of Spinicaudata possessing a carapace which is partially smooth around the umbo is a species of the Limnadiidae family, Limnadia lenticularis (L., 1761), found in the Ile-de-France and eastern France. This species is Palaearctic: its range extends over central and northern Europe52. L. lenticularis is also present in Italy.

Distribution - Ecology Distribution/range

I. yeyetta is a species mainly present in central Europe (Austria, Slovakia, Hungary, Serbia and Romania). In the Mediterranean region it is found in northern Greece and in the former Yugoslavia52; in France, it is only found in the Bouches-du-Rhône.

I. yeyetta colonises usually turbid temporary pools, in the Camargue Delta and the Crau. Its presence reflects the dispersal into biotopes of resting eggs carried by migratory birds flying between central and southern Europe, such as the Anatidae, which are particularly abundant in the region. Habitats Directive

“Mediterranean Temporary Pools” (code 3170). CORINE Biotopes

Lanau pool (Crau) and temporary pools in the Camargue: 22.32 Annual Mediterranean amphibious swards (Lythrion tribracteati). Ecology Biological characteristics

Feeding Microphagous* species feeding by filtering water and sediments. Detrivorous* feeding regime. Behaviour Usually benthic* species, living on the sediment or partly buried in superficial sediments. Description of eggs Eggs ovoid, 150 to 190 µm in size, characterised by their spiralled surface. Confusion is possible with the gyrogonites* (résistant forms) of charophytes, which are distinguished by the presence of five spiral-shaped apical cells355. Site and mechanisms of egg laying The eggs, grouped together on the abdomen of the female, are discharged into the water during each moult. Their distribution on sediments depends on the position of the female at the time of the moult, the influence of vegetation and areas of open water as well as the wind, in particular the mistral (northerly wind), which creates movements in shallow water through convection cells. The distribution of the eggs can thus be very variable depending on the biotope.

85

Modified from Defaye et al.102

Main synonyms

Mediterranean temporary pools

Longevity Longevity is estimated at less than eight weeks when conditions are favourable (no premature drying out). However, as for all branchiopods, it is very variable depending on the temperature.

Conservation - Management Assessment of populations and conservation status Number of sites

Biological cycle I. yeyetta is most often present in the spring but can also be found in autumn (November-December). A complete cycle was studied in the Camargue by Nourisson & Aguesse270. The females can lay up to 400 eggs at a time (the size of the clutch depends on the age of the female). This occurs shortly after mating. The chitinous* carapace with two lobes appears rapidly, generally at the metanauplius* stage. This first, or “heilophore”, bivalve stage occurs three or four moults after the nauplius* stage. The morphology of the larval stages has been studied by Eder129. Unlike most crustaceans, and in particular the branchiopods, the carapace is not discarded totally during successive moults, which leads to the appearance of growth lines374.

The species is only present in France in the Camargue Delta and the Plaine de la Crau (Lanau pool). In central Europe, I. yeyetta has a fairly wide distribution52. Populations

The densities of the populations are quite low, with an irregularity in the hatching periods and thus of appearance. Legal status of the species/level of protection IUCN Bern Convention Habitats Directive National and regional protection National Red Data Books and Red Lists

-

Anthropogenic factors Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Dec.



Adult stages (clutches)

Hatching and growth of larvae

No data are available relating to the damage thresholds linked to anthropogenic actions, nor are any toxicological data available. Recent observations (Thiéry, original data) are inconclusive. Natural factors

Egg bank

Environmental conditions

Interspecific relationships I. yeyetta can be found with the anostracans Tanymastix stagnalis, Branchipus schaefferi, Chirocephalus diaphanus and/or the notostracan Triops cancriformis. For more data on the ecological conditions of coexistence of several species of branchiopods in a temporary pool in a Mediterranean climate, see Thiéry373. Requirements for reproduction No data. Environmental perturbations Natural perturbations corresponding to the constraints linked to habitats with fluctuating water levels.

No data. Risks relating to populations

In the Lanau pool, the population appears to be fairly stable, given the state of the cyst bank355; no data on the state of the populations in the central Camargue. Management and conservation measures Current measures

At two known sites, situated in Natura 2000* areas, the populations appear to be stable, and their biotopes safe from any human pressure. The Lanau pool was acquired by the CEEP in 1998 and the temporary pools where this species lives in the Camargue benefit from the status of “Réserve Naturelle” (Nature Reserve) (Tour du Valat estate). Recommendations

Main natural mortality factors Not well known. However, as for most branchiopods, the individuals develop in a cohort from the onset of flooding371, which limits coexistence with predators (larvae of Coleoptera and notonectid Heteroptera, for example) which only colonise the habitats several weeks after inundation (see vol. 1, chapter 3e). Note, however, that there could be a possibility of predation, not yet quantified, by Rhabdocoela (Platyhelminthes), which are common in the temporary stillwater habitats of Provence (Thiéry, original data). The impact of birds is poorly known.

Given the current state of knowledge, no conservation measure can be proposed. Only the maintenance of the biotopes in their current state is recommended.

Bibliography AGRN-RH, 20003 ; Anonymous, 199912 ; Botnariuc, 194743 ; Brtek & Thiéry, 199552 ; Defaye et al., 1998102 ; Eder, 2002129 ; Hertzog, 1935175 ; Marincek & Petrov, 1984237 ; Marincek, 1972239 ; Nourisson & Thiéry 1988269 ; Nourisson & Aguesse, 1961270 ; Soulié-Märsche & Thiéry, 1998355 ; Thiéry & Gasc, 1991365 ; Thiéry & Pont, 1987366 ; Thiéry, 1987371, 1991373, 1996374 ; Thiéry et al., 1995375. Authors: Thiéry A. & T. Calvière

86

Macrocrustaceans

Thiéry & Champeau, 1988364

Class: BRANCHIOPODS Order: ANOSTRACA Family: LINDERIELLIDAE

Crustacean drawing modified from Thiéry & Champeau364

Linderiella massaliensis

2 mm

Main synonyms

None French name: None

male Subspecies/variability

No subspecies. Very low morphological variability; low genetic variability (Thiéry, original data).

egg (240-260 µm)

Linderiella massaliensis

Description/identification criteria

Anostracans have an elongated body without a carapace and swim on their backs in open water. They possess two cercopods* (furca) at the end of the abdomen.

Habitat General description

Linderiella possesses 11 pairs of phyllopods* and two cercopods, orange in colour, borne on the last abdominal segment (telson). The males are distinguished from the females by the morphology of the antennae: in the males, they are much more developed, biarticulate, and their pincer shape enables them to clasp the female during mating. The males possess two spiny ventral hemipenises at the base of the abdomen (genital apparatus). The females, with shorter antennae, have, on maturity, a heartshaped brood pouch*, (bicoloured blue/brown) at the base of the abdomen, in which the eggs acquire their outer envelope. The individuals, creamy-greenish in colour and transparent, measure a maximum of between 10 and 13 mm. The females are slightly larger than the males (morphological dimorphism). Similar species

For the non-specialist, all anostracans are very similarly morphologically: they can only be differentiated with an identification key102, 269. Linderiella massaliensis resembles Linderiella sp. of Spain as well as L. africana which inhabits the temporary pools of the Middle Atlas in Morocco.

Distribution - Ecology Distribution/range

Linderiella massaliensis is found in temporary pools which are flooded in autumn and winter, such as Lake Redon. Generally speaking, L. massaliensis occupies pools of karstic origin (poljés*, dolines), of varying depths. The waters are of low turbidity, poorly mineralised (temperature between 5 and 20°C, pH between 7.2 and 8.4, C20 between 250 and 600 µS.cm-1) and rich in microcrustaceans (more than 25 species of cladocerans and copepods at Bonne Cougne). Habitats Directive

“Mediterranean Temporary Pools” (code 3170). CORINE Biotopes

22.34 Southern amphibious communities (Preslion cervinae) 22.32 Annual Mediterranean amphibious swards (Lythrion tribracteati) Ecology Biological characteristics

Feeding Linderiella massaliensis, like all anostracans, is microphagous* and feeds by filtering suspended food particles from the water (microplankton, micro-organisms and organic material), using its legs which are covered in bristles. Anostracans can also disturb sediments in order to bring particles into suspension.

France

Species endemic to southeastern France (Var). Biogeography

Linderiella massaliensis is a vicariant* species of the Moroccan species L. africana Thiéry 1986370 which inhabits several dayas (temporary pools) of the Middle Atlas on the Azrou Plateau. Three other vicariant species are so far known: two in California, L. occidentalis Dodds, 1923 and L. santarosae Thiéry & Fugate, 1994367, and one in Spain Linderiella sp.7. These species together constitute a case of adaptive radiation through fragmentation of the distribution range of a “mother” species at the time of the opening of the Atlantic and the Thetys.

Description of eggs The eggs are exclusively “resting eggs”, spherical and spiny. They have a diameter of 240 to 260 µm. Longevity One generation (from hatching to the disappearance of the adults) can last, depending on the conditions, for a maximum of three to four months. Site and mechanisms of egg laying The eggs are expelled in small jets by the females and fall to the bottom in a random fashion.

87

Mediterranean temporary pools

Biological cycle Although the nauplius* stage only lasts a few hours, growth is slow as the adults are only present from February to March (breeding period) whereas the eggs hatch in autumn, from October to December. The species disappears from the pools at the end of the winter even if they remain flooded, generally until April-May. Little interannual variability in phenology. As for most anostracans, the population functions in cohorts (hatching once the pool has flooded, synchronous growth of the individuals, death).

Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Rare species: the population densities are between 0.5 and 2 ind.L-1 at Bonne Cougne and Petit Laucien. The distribution, which is often regular, can sometimes become aggregated (influence of beds of charophytes, Ranunculus, etc.). Two studies (McNutt & Thiéry and Meglecz & Thiéry) are ongoing, one to map the distribution of the cysts of L. massaliensis within Lake Bonne Cougne, the other to quantify the consequences, through the genetic bottleneck* effect, of the excavation of sediments reducing the pool’s egg bank (analysis of enzymatic polymorphism, levels of mutation and ‘genetic drift’*).

Dec.


Hatching and larval




Populations

Adult stages (egg laying)

Egg bank

Legal status of the species/level of protection IUCN Bern Convention Habitats Directive National and regional protection National Red Data Books and Red Lists

-

Environmental conditions Anthropogenic factors

Interspecific relationships Linderiella massaliensis can live alone (Petit Laucien, Var) or be found with Lepidurus apus, Chirocephalus diaphanus (Lake Redon) and Branchipus schaefferi373. Requirements for reproduction The species requires an autumn and winter flooding regime which corresponds to its biological cycle (fresh water). Environmental perturbations Early drying-out of the pool, in March for example, does not negatively affect the species as the adults are present from February and have been able to begin breeding. The introduction of Pumpkinseed Sunfish (Lepomis gibbosus, a species introduced from the USA) into Lake Bonne Cougne in 2001-2002 caused a collapse in the population of L. massaliensis during that hydrological cycle. The situation returned to normal after the summer drying out (eradication of the fish) and the autumn flooding. Main natural mortality factors Few natural factors affect the biological cycle of this anostracan. For example, the species is not sensitive to freezing temperatures (population present under the ice at Bonne Cougne in January 2001, or a minimum water temperature of 1.8°C at Petit Laucien, in February 2003, Thiéry, original data). However, the species does not appear to tolerate water which is too warm.

The Lake Bonne Cougne (Centre Var) is seriously threatened by both local activities (overdeepening, dyke construction, filling in with spoil brought in from building work or hospital waste containing contaminants, etc.) and by a project for the creation of a golf course on the land above the depression. Although the planning application deposited at the Var préfecture in Toulon was initially refused, it has just been approved. Generally speaking, the modifications of the water quality due to pumping from the karstic water table, extraction of the run-off waters of the topographical catchment area and to water running into the Lake Bonne Cougne from the golf course above will jeopardise in the short term (estimated at less than 10 years), the populations of branchiopods as well as those of planktonic crustaceans, cladocerans, copepods, etc.377 Natural factors

In the Lake Gavoty (Var), the development of the phytocenosis* considered to be natural, is in fact more likely to be influenced by the nutritional inputs provided by the catchment area (homes without a collective treatment system, livestock raising - llamas, goats, horses, etc.). The increase in nitrogen and phosporus levels reduces the hatching rate of L. massaliensis, whose mechanism depends on osmotic processes374. In addition, the covering of the sediments by a very dense vegetation bed reduces the mobility of the larvae and increases natural mortality. Risks relating to populations

Conservation - Management Assessment of populations and conservation status Number of sites

This endemic species has only been found in five temporary pools in the Var (around Saint-Maximin and Brignoles) including Lake Redon, Lake Gavoty, and the Petit Laucien (type locality).

88

Two populations are threatened (Bonne Cougne and Gavoty) out of the five known. If the possibilities for connectivity* are considered for the five populations which are in a fragmented habitat within this metapopulation*, these threats represent a potential loss of 40% of the genetic heritage entering into the gene flows necessary for the maintenance of the metapopulation. In 2002, an identical situation in California for the vicariant species L. santarosae367 resulted in the implementation of conservation measures. In contrast, no actions, or conservation measures are planned in France.

Macrocrustaceans

Management and conservation measures

Bibliography

Current measures

AGRN-RH, 20003 ; Alonso, 19967 ; Anonymous, 199912 ; Brtek & Thiéry, 199552 ; Defaye et al., 1998102 ; Nourisson & Thiéry, 1988269 ; Thiéry & Champeau, 1988364 ; Thiéry & Gasc, 1991365 ; Thiéry & Fugate, 1994367 ; Thiéry, 1986370, 1991373, 1996374 ; Thiéry et al., 1995375, 2002377.

Most of the sites in the Centre Var are included in Natura 2000* areas. Recommendations

Imperative protection of the sites in their current condition where the species is present. Inclusion of the species on a Red List.

Authors: Thiéry A. & T. Calvière

89

Mediterranean temporary pools

Tanymastix stagnalis (Linnaeus, 1758) 2 mm Class: BRANCHIOPODS Order: ANOSTRACA Family: BRANCHIPODIDAE Main synonyms

Cancer stagnalis Linnaeus, 1758 Gammarus stagnalis (Linnaeus), Fabricius, 1775 Astacus stagnalis (Linnaeus) Pennant, 1777 Branchipus lacunae (Guérin-Méneville) Baird, 1852 Chirocephalus lacunae Frauenfeld, 1873 Tanymastix lacunae Daday, 1910

male

egg (190-430 µm) French name: none Spanish, Italian names: species is present in these two countries

Tanymastix stagnalis

but there is no common name. English name: Fairy Shrimp (general name for Anostraca) Subspecies/variability

No subspecies. Variability of adults: Margalef236 and Nourisson271 report aberrant T. stagnalis with hypertrophied antennae. The lenticular cysts can present some differences in diameter depending on their geographical origin375. An aberrant morphology also exists, corresponding to a deformation of the lenticular shape376.

Distribution - Ecology Distribution/range

T. stagnalis is a Palaearctic species, present throughout the Mediterranean region (Europe and North Africa) but it is very unevenly distributed52.

Description/identification criteria

Body without carapace, eyes pedunculate, 11 pairs of phyllopods*, abdomen without appendages, and two cercopods, vermilion in colour, forming the furca, at the end of the abdomen. The males are distinguished from the females by their prehensile antennae which enable mating to take place. A frontal complex (clypeal) unfurls to play a tactile role and for partner recognition. The genital apparatus of the male (2 retractable hemi-penises) is situated at the base of the abdomen. The female’s brood pouch*, also borne on the ventral side at the base of the abdomen, bears two widely spaced spines. The structure and development of the cyst envelopes have been studied by Garreau de Loubresse149, 150, 151. The males are a pale green colour and the females have a bluebrown to orange oviger*. Their maximum size varies from 6 to 20 mm.

France

It has also been found in the Paris region309, in the Camargue270, the Var363 and the Rhône Valley (Thiéry, original data ). Other Mediterranean countries

Spain7, Italy (including Sardinia )92, Macedonia and North Africa (Algeria)154, 339. Habitat General description

Tanymastix stagnalis is present in habitats which dry out rapidly such as shallow pools (sometimes on rock shelves as in the Fontainebleau Forest), vegetated ditches or sansouires (Salicornia scrub) in the Camargue.

Similar species

Within the genus, three other endemic species are morphologically very similar: T. stellae (Corsica, Sardinia), T. affinis (Morocco) and T. motasi (Romania). Only males can be distinguished with certainty (examination of antennae). Generally speaking, T. stagnalis does not coexist with any of these endemic species.

France • Roque-Haute Nature Reserve in the Hérault260. • Cupular pools of the Estérel Massif and the Colle du Rouet (see vol. 1, box 7). These basins of several dm2 of surface area, fill up from October to May, depending on the year. They are covered with an acidic sandy-silty layer (average pH 5.2) from 5 to 10 cm in depth363. • Oligo-brackish temporary pools on calcareous clayey-silty substrate in the Camargue (Tour du Valat estate). Italy Temporary clear water with sparse vegetation92. Habitats Directive

Mediterranean pools “Mediterranean Temporary Pools” (code 3170).

90

Macrocrustaceans

Salicornia scrub Mediterranean and Thermo-Atlantic halophilous scrubs (code 1420) CORINE Biotopes

T. stagnalis is present in Mediterranean temporary pools as well as in other types of seasonally flooded habitat. Temporary pools 22.34 Mediterraneo-Atlantic amphibious communities.

Environmental perturbations Species present in pools with dry sediments in the summer, without a groundwater capillary link. Any modification in the environment which can cause an increase in the mineral content (conductivity <200 µS.cm-1) reduces its hatching potential.

Ecology Biological characteristics

Feeding Tanymastix stagnalis, like all the anostracans, is microphagous* and feeds by filtering the food particles in suspension in the water (microplankton, micro-organisms and organic material), using its legs which are covered with bristles. Description of eggs The eggs, whose lenticular shape is specific to the genus Tanymastix, are coppery brown in colour and measure between 190 and 430 µm in diameter. Behaviour The adults react to variations in light intensity; behaviour involving fleeing towards the pool bed, or even burying in the mud, have been observed when shade is cast over them. Longevity This varies according to the temperature and the season when the species appears in the habitat (from 30 days in summer to over 60 days in winter). Site and mechanisms of egg laying The eggs are laid in open water, where they usually float, and accumulate towards the edges of the pool262. Biological cycle Tanymastix lives for a period of around three months at all seasons of the year, both in winter and in summer . It grows rapidly and, depending on the temperature of the water, can reach sexual maturity after seven to 40 days. The nauplius* stage only lasts a few hours. The biological cycle of T. stagnalis is very variable between sites and from year to year. The cycle begins with the flooding of the pool and its duration depends on the temperature. Feb.

March

April

May

Interspecific relationships Tanymastix stagnalis can be associated with the anostracans Chirocephalus diaphanus, Branchipus schaefferi, with the notostracan Triops cancriformis and with Imnadia yeyetta (Spinicaudata). On the other hand, it is often the only species present in cupular pools, the other species having longer biological cycles. Requirements for reproduction Pioneer species, colonising clear, fresh and poorly mineralised water.

Salicornia scrub (seasonally flooded) 15.61 Mediterranean salt-meadow scrub.

Jan.

Environmental conditions

June

July

Hatching
larval
adults (egg laying)

Aug.

Sept.

Oct.

Nov.

Main natural mortality factors Perturbations in the habitat, especially the introduction of predators (such as the fish Lepomis gibbosus, Gambusia affinis, etc.) are the main threats for T. stagnalis.

Conservation - Management Assessment of populations France

Between five and ten stations in Provence. Tanymastix is present in the Fontainebleau Forest (Paris region), the Roque-Haute Nature Reserve260, at the Colle du Rouet (Var)363, in the Crau, the Camargue, and in a pool in the Rhône valley, at Avignon on the Ile de la Barthelasse (Thiéry, original data). Italy

Eleven localities, including six in Sardinia and four in the centre of the peninsula (see map in Mura263). Spain

Uncommon species7, 236. Conservation status

Little or no data on changes in distribution. Because the species occupies a very fragmented area, its survival can be threatened by land reorganisation measures (resulting in modifications in Plan d’Occupation des Sols (land-use plans) or infilling. However, because it is one of the most rapid anostracans to reach sexual maturity (less than 15 days), it is capable of laying eggs rapidly and thus of regenerating its egg stocks without too much damage.

Dec.

Hatching
larvae
adults (egg laying)

Egg bank

One cycle per year in general, rarely two, in which case poor breeding success is observed during the second cycle (Lanau pool) (Thiéry, original data).

91

Mediterranean temporary pools

control over land ownership and usage) and the pools of the Colle du Rouet are included in a Natura 2000* site.

Legal status of the species/level of protection IUCN Bern Convention Habitats Directive National and regional protection National Red Data Books and Red Lists

-

Recommendations

It is vital that the small shallow biotopes be maintained in a healthy condition.

Bibliography Anthropogenic factors

In general, the infilling of pools and the use of pesticides in the rural environment threaten populations. Management and conservation measures Current measures

Some Italian pools containing Tanymastix are situated in protected areas263. In France, the Roque-Haute site has a legal “Réserve Naturelle” status (currently ineffective because of the absence of

92

Alonso, 19967; Artom, 192717; Brtek & Thiéry, 199552; Cottarelli & Mura, 198392 ; Defaye et al., 1998102 ; García & de Lomas, 2001146 ; Garreau de loubresse, 1965149, 1974150, 1982151 ; Gauthier, 1928154 ; Margalef, 1958236 ; Médail et al., 1998246 ; Moubayed, 1998260 ; Mura, 1991262, 2001263 ; Nourisson, 1960271 ; Nourisson & Aguesse, 1961270 ; Rabet, 1994309 ; Samraoui & Dumont, 2002339 ; Terzian, 1979363 ; Thiéry & Gasc, 1991365 ; Thiéry, 1991373, 1996374 ; Thiéry et al., 1995375, en préparation376. Authors: Thiéry A. & T. Calvière

Macrocrustaceans

Triops cancriformis

(Bosc, 1801) (1) &

Class: BRANCHIOPODS Order: NOTOSTRACA Family: TRIOPSIDAE

Lepidurus apus

(Linné, 1758) (2)

1 cm

Main synonyms

(1) Limulus cancriformis Lamrck, 1801; Apus cancriformis Bosc, 1802; Apus viridis Bosc, 1802; Triops simplex Ghigi, 1921 (2) Monoculus apus L., 1758; Binoculus palustris Müller, 1776 Apus productus Bosc, 1802; Apus apus (Linnaeus) Latreille, 1802 Lepidurus productus (Bosc) Leach, 1819

Triops cancriformis

French names: (1) Triops cancriforme (sometimes inaccurately

egg (340-400 µm) Crustacean drawings modified from Alonso7

called apus – Apus is a bird genus), (2) Lépidure English name: Tadpole Shrimp (general name for Anostraca) Subspecies

Triops c. cancriformis and Lepidurus a. apus are the only two French subspecies of Notostraca, but two other subspecies of T. cancriformis exist (T. cancriformis simplex and T. cancriformis mauretanicus) as well as subspecies of L. apus including L. apus lubbocki in the countries of the Mediterranean Basin (Spain, Morocco, etc.).

1 cm

Lepidurus apus egg (430-520 µm)

Description/identification criteria

These crustaceans, with a dorsoventrally flattened body, have an oval, streamlined dorsal carapace covering the head, the thorax and a part of the abdomen. They can reach a maximum size of 10 to 11 cm, but the two cercopods* (furca), at the end of the abdomen, are almost as long as the body. The compound eyes are sessile* and dorsal. The phyllopods*, the first two pairs of which are locomotive, are more numerous in Triops (48 to 57 pairs) than in Lepidurus (35 to 48 pairs). The posterior abdominal segments are apodal (no legs): the last four to five segments in Lepidurus, the last four to seven segments in female Triops and the last five to nine segments in male Triops. Female Triops are also distinguished from the males by the structure of the 11th pair of thoracic legs which are transformed into a brood pouch (oostegopod), where the eggs are stored. The first pair of legs have flagelliform endites (extensions) extending some way beyond the edge of the carapace in Triops but less so in Lepidurus. The latter is characterised also by the presence, on the final segment (the telson), of a caudal ramus with toothed edges. The dorsal shield is greenish-brown in Lepidurus and brownish in Triops, in which the thoracic legs can be reddish or greenish.

Distribution - Ecology Distribution/range France

Triops is present over the whole country (including Corsica) and is fairly frequent in the east and in the coastal regions, particularly the Mediterranean regions. Lepidurus appears to be absent from eastern France and Corsica, but is more frequent in the north than Triops. Generally speaking, the two species are allopatric (separate ranges). The distribution of the two species seems clearly determined by precipitation, as is the case in southern France372 (P <600 mm for Triops, P >600 mm for Lepidurus), in Israel201, Algeria154 and Morocco369, 371. The genus Triops is found in drier regions than Lepidurus. These distributions may be erroneous, as the two species seem to have been often confused. Furthermore, their presence at certain stations has not been confirmed since 1950. For further details, consult Nourisson & Thiéry269, Brtek & Thiéry52 and Defaye et al.102. Other Mediterranean countries

Similar species

These are the only two species of French Notostraca. Separation of the two species is easy: telson with or without caudal ramus. Identification of the various subspecies, on the other hand, is very difficult. Specialist advice is needed.

Both species are present throughout Europe, as well as in the Mediterranean Basin for Lepidurus apus apus. • Spain: Lepidurus apus apus, Triops cancriformis simplex, T. c. mauretanicus 7 (which replaces T. c. cancriformis in southern Spain). • Portugal: T. cancriformis. • Italy: L. apus (mainland Italy, Sicily, Sardinia ), T. cancriformis, and T. numidicus (= T. granarius), a species of African origin present in Calabria.

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Mediterranean temporary pools

• • • • •

Malta: T. cancriformis205. Croatia: L. apus. Former Yugoslavia: L. apus, T. cancriformis238, 289. Morocco: T. c. simplex, T. c. mauritanicus, L. a. lubbocki 369, 371. Israel: L. a. lubbocki, T. c. cancriformis (Thiéry, original data ).

and 50°N (northern and central Europe) and an equal sex ratio below 45°N (southern, western Europe and North Africa), with exceptions (mainland Italy, Sicily, etc.). The parthenogenetic females possess a hermaphroditic gland but the spermatozoa formed degenerate and play no part in reproduction.

Habitat General description

The two species are found in temporary pools. Whereas Triops can be found in sometimes quite turbid pools, Lepidurus is usually found in fresh, clear water, and sometimes in temporary watercourses with a weak current. Triops is also present in artificial habitats flooded in the summer, such as the ricefields in the Camargue297. Habitats Directive

Description of eggs The eggs acquire their external envelope in the brood pouch*. They are smooth. Their diameter is greater in Lepidurus (430-520 µm) than in Triops (340-400 µm). Site and mechanisms of egg laying The eggs (60 to 70 in Lepidurus) are deposited just before the moult, in clumps, on small pebbles where they remain during the dry period. In Lepidurus, the female can lay eggs on the stems of aquatic vegetation or bury her eggs in the mud.

“Mediterranean Temporary Pools” (code 3170). CORINE Biotopes

Mediterranean temporary pools (1) 22.32 Annual Mediterranean amphibious swards (Nanocyperetalia). (2) 22.34 Southern amphibious communities (Isoetalia). Ricefields (1) 82.41 Ricefields.

Ecology Biological characteristics

Biological cycle After hatching, the nauplius* larvae evolve rapidly into metanauplius* larvae, with the dorsal shield already present. Lepidurus, which is present from January to June (most often from February to April), has a life expectancy of 4 to 6 months. Triops, which can be found in all seasons of the year in the flooded phase, from March to January (most often from May to October), has fairly rapid growth and can produce resting eggs from the 15th day, when it is still small in size.

Triops cancriformis Temporary pool Jan.

Feeding They take a wide range of food and they can be considered as predators, herbivores or detritivores. In turbid waters, they are scavengers: they feed on non-living particles (dead organisms, organic detritus, mineral particles) in suspension in the water, and on the bacteria that that build up on these particles. They dig up the mud with the front part of their carapace.

Feb.

March

April

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June

July

Aug.

Sept.

Oct.

Nov.

Dec.





Hatching and larval

stages adult stages (egg laying)

Egg bank

Lepidurus apus Feeding can vary during the growth of the animal, which can pass from a strictly phytoplanktonophagous* feeding regime, when it measures less than 5 mm, to a herbivorous regime (in the broadest senses); when it is larger than 1 cm, it also feeds on microcrustaceans. As an adult, it is capable of devouring weakened anostracans (usually Branchipus schaefferi), chironomid larvae, Oligochaete worms and tadpoles (Pelobates cultripes). Behaviour They usually swim close to the bottom, but can come up to the surface, ventral side uppermost, if there is a lack of oxygen. Reproduction Mating is ventral-ventral in Lepidurus, whereas it appears to be dorso-ventral in Triops. Parthenogenetic reproduction is frequent in Triops, but when both sexes are present, sexual reproduction is the rule. The sex ratio* varies according to latitude with an absence of males beyond 50°N, a low percentage between 45°

94

Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Dec.





Hatching and larval

stages adult stages (egg laying)

Egg bank

Environmental conditions

Requirements for reproduction Their cycle being relatively long, these notostracans require a submersion period of three to five months. Triops, which can be found from March to January, is more thermophilic than Lepidurus, which is present from January to June. Triops cannot withstand temperatures higher than 35°C and thus from June to August is practically absent from habitats flooded in summer, such as the ricefields in the Camargue.

Macrocrustaceans

Interspecific relationships Triops is usually found with Branchipus schaefferi and more rarely with Chirocephalus diaphanus. In the Camargue, it can also live alongside Tanymastix stagnalis, Imnadia yeyetta and Eoleptestheria ticinensis. In ricefields, when its populations explode (densities reaching 300 individuals per m2 have been measured in the Camargue297), it can become an important part of the diet of the Little Egret Egretta garzetta169. Lepidurus usually lives with Chirocephalus diaphanus but can, in the Var, live alongside Linderiella massaliensis. Territories In notostracans, there is a differential micro-distribution between the sexes in the pool. Main natural mortality factors Early drying out of the pool (case of the Lanau pool, for example) can cut short the otherwise significant life span of these species. Predation, notably by Ardeidae, can play an important role in certain populations (Camargue)169, 297.

Conservation - Management Assessment of populations

In France, they are present at many stations: Triops is found notably at the sites of Roque-Haute (Languedoc-Roussillon), Padulu (Corsica), in the Camargue and the Crau (Lanau pool), while Lepidurus has been found in the pools of the Var, Lake Redon, Lake Gavoty and Saint Maximin (PACA). A description of the distribution of the two species in Languedoc-Roussillon is given by Thiéry372. In Italy, these species are fairly widespread, in particular in the ricefields. Conservation status

In France, these two species are common but Triops seems to have been often confused with Lepidurus and their presence at some stations has not been confirmed since 1950. Legal status of the species/level of protection IUCN Bern Convention Habitats Directive National and regional protection National Red Data Books and Red Lists

Malta: rare species (R)344

Conservation problems/threats Anthropogenic factors

As Lepidurus can be found in the overflow pools of rivers (Loire, for example), physical planning of watercourses and their regulation in order to limit flooding can represent an indirect threat to their populations. Triops is sometimes considered to be a nuisance in ricefields where it can be extremely abundant (it uproots young plants, eats their growing tips, and reduces photosynthesis activity by bringing sediments into suspension). Others estimate that these perturbations are minor and that it acts as an extremely efficient biological control agent to combat invading plants. It is, however, eliminated by insecticides or copper sulfate. More generally, the infilling of pools and the use of pesticides destroys populations. Management and conservation measures Current measures

In France the pools of Roque-Haute and the Camargue are situated in Nature Reserves (at Roque-Haute, however, conflicts with the owners do not allow management of the reserve to take place). The lakes of the Centre Var, the Padulu pool in Corsica, and the Lanau pool, acquired by the CEEP in 1998, are included within Natura 2000* areas. Recommendations

Ensure the conservation of the habitat (hydrological functioning and water quality).

Bibliography Alonso, 19967 ; Anonymous,199912 ; Boix et al., 200241 ; Brtek & Thiéry, 199552 ; Defaye et al., 1998102 ; García & de Lomas, 2001146 ; Gauthier, 1928154 ; Hafner, 1977169 ; Knoepffler, 19781979199 ; Kuller & Gasith, 1996201 ; Lafranco, 1990205 ; Longhurst, 1955217 ; Marincek & Petrov, 1992238 ; Moubayed, 1996259 ; Nourisson & Thiéry, 1988269 ; OEC, 2001273 ; Petrov & Petrov, 1997289 ; Pont & Vaquer, 1986297 ; Scanabissi & Mondini, 2002342, 343 ; Schembri & Sultana, 1989344 ; Thiéry & Gasc, 1991365 ; Thiéry, 1985368, 1986369, 1987371, 1988372, 1991373, 1996374 ; Thiéry et al., 1995375 ; Zaffagnini & Trentini, 1980404 Authors: Thiéry A. & T. Calvière

95

Mediterranean temporary pools

Ischnura pumilio

(Charpentier, 1825) (1) &

Ischnura genei

(Rambur, 1842) (2)

Order: ODONATA Suborder: ZYGOPTERA Family: COENAGRIONIDAE Main synonyms

(1) Agrion rubellum Curtis, 1839; Agrion cognata Sélys, 1841 (2) Agrion genei Rambur, 1842 French names: (1) l’Agrion nain, (2) l’Agrion de Gené English name: (1) Scarce blue-tailed damselfly

male

Description/identification criteria95, 401

Like all the Zygoptera, these species have the eyes well separated and the four wings of exactly the same shape. The family characteristics include the absence of any metallic reflections and the entirely hyaline (transparent) wings, which are petiolate (stalked at the base). Both species have a mostly dark or black abdomen with a bronze tint and, in males, the eighth segment at least partly blue (also the ninth in Ischnura pumilio). The pterostigma (dark mark on the wing) is scarcely longer than wide. In I. genei, the pterostigma is slightly larger on the forewings than on the hindwings, while in I. pumilio it is almost twice as large on the forewings as on the hindwings. Only I. genei has a median tubercule on the prothorax* and this is clearly indented. Abdomen: 18 to 27 mm. Wings: 12 to 19 mm. Subspecies

None Similar species

Ischnura elegans (similar to I. genei in particular).

Distribution - Ecology Distribution/range France

Ischnura pumilio occurs only on the mainland (especially the Rhône Valley, Pre-Alps, Champagne-Ardenne, Centre-Ouest, and Var), and I. genei is only found in Corsica122, 159. Other Mediterranean countries

Ischnura genei occurs only on the islands of the western Mediterranean (Sardinia, Sicily, Capri, Malta, Elba, etc.). I. pumilio is more widely distributed, along the whole of the coast of Mediterranean Europe and part of the North African and Middle Eastern coasts95, 401. Habitat

Both species are pioneer colonisers and are therefore attracted to recently created habitats, temporary or otherwise, usually with still water. They may be found in brackish water, and sometimes in still areas of water associated with watercourses (oxbow lakes, etc.). They are usually seen below 800 m altitude, however some sites are known at up to 2,100 m (Ischnura pumilio) 55, 95, 173.

96

0.5 cm

Ischnura pumilio I. pumilio does not like acid biotopes (Samraoui, unpublished data). It is also known to disappear from some sites after a few years, probably due to habitat succession68, 207. Ecology Biological characteristics

(1) Ischnura pumilio: in southern France, the adults, or imagos, can be seen from the beginning of April to the end of September95, sometimes March to October (Papazian, pers. com.), with ephemeral populations323, while in Spain and northeast Algeria the species appears in two waves, one in August, the other in September/ October (Samraoui, unpublished data). The lifespan of the imagos is fairly short (of the order 40 days) and their sexual maturation is also very rapid. Females are capable of breeding six to ten days after emergence69 and the males possibly even sooner207. This maturation is all the more rapid if the individual emerges late in the season, thereby maximising the chances of breeding207. After mating, the female goes alone to lay the eggs on the stems of plants emerging from shallow water, choosing species with stems which are fairly soft (often pithy), such as certain rushes68. The female stations herself just above the surface of the water and plunges her abdomen in so that she can lay her eggs in the submerged part of the stem. She prefers to lay at the level of the leaf sheaths (rushes), where the growth of the plant tissues will completely cover the eggs; this appears to be advantageous for their survival during dry conditions68. The eggs can survive for at least two weeks in an emergent plant if it has its base in the water. The eggs hatch quickly (after 15 days at an ambient temperature of 25°C68, most often between 20 and 30 days (Papazian, pers. com.). Rapid larval development enables the adults of the second generation to emerge before the pool dries out completely. In temporary habitats, this second generation disperses and awaits the return of the water before laying (August to October, depending on the date of re-filling with water), usually in another pool. The eggs then hatch fairly quickly and the young larvae spend the winter in the diapause* stage.

Odonata

In the Mediterranean region, the short hatching time, rapid larval development, precocious emergence spread over a long period of time (advantageous in view of the interannual variability of the hydrological conditions), and the rapid maturation of newly emerged individuals enable two generations of the species to be produced per year. The rapid dispersal of individuals following emergence (migratory instinct), together with the rapid turnover of generations, enable I. pumilio easily to colonise temporary habitats, forming new colonies69. (2) I. genei: few facts are known regarding this species. In France, its phenology is very similar to I. pumilio (Papazian, pers. com.). In Sardinia, adults are present from the beginning of May to the end of September55. Regarding its ecology, one may refer partly to I. elegans, which is very similar. Like I. pumilio and I. elegans, I. genei has two generations per year, with rapid development of the pre-imaginal* stages5, 95. Simplified phenological cycle of Ischnura pumilio

(oxidation ponds), the marshes and rice fields of the Camargue, on the Durance (small temporary pools at Puy-Sainte-Réparade), in the Crau (Vergière canal) and on the Touloubre at Saint-Chamas (Papazian, pers. com.). (2) I. genei occurs at a number of sites in Sardinia55 and at 16 out of 50 stations visited as part of the dragonfly survey in Corsica159, 323, notably in the temporary pools at Tre Padule de Suartone. Legal status of the species/level of protection IUCN Bern Convention Habitats Directive National and regional protection National Red Data Books and Red Lists

Italy, regional protection (Tuscany): regional law n° 56 of 06/04/2000 -

Conservation problems/threats

Temporary pool in southern France Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Anthropogenic factors

Dec.


Winter larval diapause,

First generation (pool A)


Interference with habitats that dry out in summer (drainage, filling in), and lack of maintenance of pools.

followed by larval growth Emergence of adults, egg laying and migration

Natural factors

Second generation Larval development, emergence and dispersal to other pools

Egg laying (pool B) Hatching of eggs




Larval diapause…

Environmental conditions

Hydrology Both species lay their eggs in submerged stems. The eggs die if the plant is out of the water for more than about fifteen days (no summer diapause at the egg stage), as do the larvae if the pool dries up too early.

Overgrowing of vegetation at water bodies, siltation and overgrowing of wetlands; feeding competition with other dragonflies (such as Ischnura elegans), predation by spiders and natural mortality during heavy rain (very frail species). Risks relating to populations

Early and prolonged drought may result in the complete destruction of all the larvae at a site. Only recolonisation will then enable these Ischnura to return. Management and conservation measures Current measures

Vegetation The sites used are rather sunny and the emergent aquatic vegetation need not necessarily be very well developed173, 204. The plants used by Ischnura pumilio for egg laying have a grasslike form330 such as rushes (Juncus bulbosus, J. tenuis, J. subnodulosus, J. inflexus, J. articulatus) or spike-rushes (Eleocharis palustris) 68,173. I. pumilio prefers to colonise water bodies with clayey beds and fairly shallow average depth (less than one metre). Competition Ischnura pumilio has a low level of tolerance of competition from other dragonfly species. It is a requirement therefore that there are few of these at the sites or at least that some areas are free from competitors (notably I. elegans), or that the vegetation is well-developed enough to allow I. pumilio to find refuge173. Accordingly, the adults often only occupy a limited territory within a suitable biotope330.

Conservation - Management Assessment of populations and conservation status

(1) Ischnura pumilio is less common than I. elegans. In the French Mediterranean region, it is known at the pools in the Plaine des Maures244, the Centre Var377, the Colle du Rouet, Porquerolles

In Corsica, Tre Padule benefits from its legally protected “Réserve Naturelle (Nature Reserve)” status. The sites at the Plaine des Maures, the Centre Var and the Colle du Rouet are included in Natura 2000* areas. Recommendations

The aquatic vegetation should be maintained in an undeveloped condition if the site is not much used by dragonflies, and even more so if there is strong competition. Vegetation management should favour small rushes or club-rushes, with sufficiently low water levels. In addition management should aim for the pool to be flooded until spring (from November to April-May) to enable the larvae to develop.

Bibliography Aguesse, 19685 ; Anonymous, 199711 ; Bucciarelli et al., 198355 ; Cham, 199268, 199369 ; D’Aguilar & Dommanget, 199895 ; Dommanget, 1994122 ; Grand & Papazian, 2000159 ; Heidemann & Seidenbusch, 2002173 ; Landmann, 1985204 ; Langenbach, 1993207 ; Médail et al., 1993244 ; Roche, 1990323 ; Rudolph, 1979330 ; Thiéry et al., 2002377 ; Wendler & Nüss, 1994401. Author: Gendre T. Collaborators: Jakob C., M. Papazian, B. Samraoui & N. Yavercovski

97

Mediterranean temporary pools

Lestes barbarus

(Fabricius, 1798) (1) &

Lestes virens

(Charpentier, 1825) (2)

Order: ODONATA Suborder: ZYGOPTERA Family: LESTIDAE Main synonyms

(1) Agrion nympha Hansemann, 1823 Agrion barbara Fabricius, 1798 Lestes barbara Selys, 1840 (2) Lestes vestalis Rambur, 1842 Agrion virens Charpentier, 1825

Lestes barbarus males

French names: (1) Leste sauvage, (2) Leste verdoyant English names: (1) Shy Emerald Damselfly, Southern Emerald

Damselfly, (2) Small Emerald Damselfly Subspecies

L virens virens (in southern France) L. virens vestalis (in Italy) Description/identification criteria

(1) Lestes barbarus has a clearly bicoloured pterostigma (dark mark in the wing) when mature: brown in its basal half and yellow-white in its apical half. The overall body colour is clear metallic green at first, becoming more or less coppery, even dull brown in old individuals. The lower hind part of the head, the coxa, the metathorax, and the thoracic and abdominal lines are pale yellow. The male has short cerci*. The female has a rounded vulvar scale, untoothed, and ochre-coloured legs in most cases. Measurements of male: abdomen 26 to 35 mm; hind wings 19 to 25 mm. Measurements of female: abdomen 26 to 33 mm; hind wings 21 to 25 mm. (2) Lestes virens has a uniformly pale brown pterostigma, edged with white veins (apart from in juveniles). The general body colour is brilliant bronze-green, the sides of the thorax and the tip of the abdomen are pale blue in mature males. The lower hind part of the head is pale yellow, as in L. barbarus. The male has very short cerci. The female has a vulvar scale with a short, only slightly pointed tooth. L. virens is smaller than L. barbarus. The subspecies from southern France, L virens virens, shows a thin pale yellow line along the anterior part of the metathorax*. Measurements of male: abdomen 24 to 34 mm, hindwings 16 to 24 mm. Measurements of female: abdomen 24 to 30 mm, hindwings 17 to 23 mm95, 173, 322, 402.

1 cm

Lestes virens

Distribution - Ecology Distribution/range France

(1) West and southeast France (but mainly a southern species: coast and Corsica). (2) Southern France (coast and Corsica). Other Mediterranean countries

(1) Southern and temperate Europe, North Africa, Middle East. (2) Western Mediterranean Basin except mainland Italy. Habitat

(1) Lestes barbarus is found in still, sunlit water bodies, clear, shallow, often but not exclusively oligotrophic*, up to 250 m altitude. As a pioneer species, it appears more particularly to prefer habitats which dry out in summer (temporary pools) and brackish coastal marshes, where there are often large populations. (2) L. virens occurs in still bodies of fresh water, including temporary pools, colonised by vegetation, up to 1,000 m altitude. Ecology

Similar species

Lestes barbarus is very similar to L. virens (especially at the juvenile stage) and to its subspecies L. v. vestalis.

Biological characteristics

(1) In southern France, the adults of Lestes barbarus (known as imagos) are fairly common from the end of May (immature at that time, they become mature in June) to the end of September (at which time they colonise new pools). In spring, after emergence, the immature imago leaves the water, to return when it is sexually mature, usually in June-July. The period of maturation between flying away and returning to the pool is estimated to be 15 days on average for Zygoptera89. In other regions maturation

98

Odonata

may take place following a period of aestivation, as is the case in particular of the populations of southern Italy and northern Algeria338, 340, 390. The duration of the maturation period influences the egg-laying period: in southern France laying may take place from the end of June to September (Papazian, pers. com.), while in southeast Algeria (in Numidia) the females become mature in September and no breeding behaviour is seen before October (Samraoui, unpublished data). Mating (forming a heart shape) and the tandem flight which follows, may last for between 30 and 160 minutes (for both species) and usually takes place during late morning387. After mating, the partners continue their tandem flight and the females begin to lay eggs using their ovipositor (egg-laying organ). In both species, eggs are laid into the aerial parts of plant stems4: the female’s ovipositor pierces the plant to insert the eggs, which are often laid in groups in the same stem, with gaps of a few millimetres between them. The distances covered between stems range from a few centimetres to a few metres387. Often, the male breaks away from the tandem formation after a certain time, and the female then continues to lay by herself. The total duration of egg laying may last for up to six hours, depending on the weather conditions and diurnal rhythm. After a period of diapause* inside the plant, which varies according to the date of laying, the eggs hatch in the spring of the following year, and give birth to elongated larvae which develop fairly quickly, taking about two months and moulting about ten times. These larvae are carnivorous, like all dragonfly larvae, and eat twice their own weight in food every day. Thanks to the diapause eggs (a primitive protective characteristic of the species) and to the short larval development period (six to eight weeks) very synchronised emergence may take place in spring, with practically all the imagos emerging over a short time. The sex ratio* (calculated at a Camargue site) is 50% females for larvae and 54.1% females for exuviae4. The adult dies later in the autumn. The maximum adult lifespan of a damselfly such as L. barbarus or L. virens in southern France is one and a half months, while in populations further to the south it extends to more than two and a half months4. (2) The mature adults of Lestes virens may be seen from June to September in southern France, and from May (occasionally end of April) to November in northeast Algeria (Samraoui, unpublished data). The female, who may or may not be accompanied by the male, lays her eggs in pairs, in spring and summer, into plant stems. After a winter egg diapause (often preceded by a summer diapause), larval development takes place in the following spring as with L. barbarus, and lasts for eight to ten weeks402. In the North African part of the Mediterranean Basin, a proportion of Lestes also lays in summer, before the site becomes flooded; the diapause eggs then withstand the drought, then overwinter and do not hatch until the following spring340. In Numidia, this is true of L. virens at lower altitudes (laying in June-July), while at higher altitudes breeding activity in this species does not begin until August (Samraoui, unpublished data). Both species (L. barbarus, L. virens) only have one generation per year.

Phenological cycle of Lestes barbarus Southern France Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Dec.





Eggs in diapause

Larval development Emergence of adults Maturation of adults Egg laying Death of adults

Environmental conditions

Winter temperatures Lestes barbarus and L. virens are adapted to the Mediterranean climate due to their rapid life cycle and their temperature requirements95. However, in southern France, L. barbarus requires a period of cold, of at least fifteen weeks below 10°C, to trigger the hatching of all of the eggs4. Hydrology As far as is known for these two species, the eggs are laid most often in a dry environment; the pool need not be flooded at the time of laying for the eggs to survive4, 340, since they are capable of entering a long period of diapause inside the plant stems. On the other hand, the emergence of adults, which is delayed, must take place before the pool dries out completely in spring or summer, and this is usually the case thanks to the rapidity of larval development. At the time of the emergence of the prolarva*, it sometimes happens that the place where the eggs were laid is dry; the prolarva, which on hatching allows itself to drop off the stem, is then capable of crawling to the water over a distance of a metre or more (Papazian, pers. com.). Vegetation According to Utzeri et al.387, the plants preferred by Lestes virens vestalis for egg laying at a temporary pool in Italy (Rome), are Carex sp. and Juncus effusus (green or dry stems). L. barbarus shows a preference for the same species, as well as Juncus articulatus, Polygonum hydropiper, Alisma plantago and Mentha sp. Samraoui (unpublished data) reports that in Numidia, L. barbarus prefers to lay in the stems of Juncus effusus.

Conservation - Management Assessment of populations Mainland France

Both species are known from the Alpes Maritimes at the Pic des Courmettes (Papazian, pers. com.), as well as in the Var: Plaine des Maures244, Colle du Rouet325, Joyeuse valley at Callas, lakes of the Centre Var377. Lestes barbarus has been observed at several other localities in the Var (Palayson wood, Giens Peninsula)363 (Papazian, pers com.), as well as in the Bouches-du-Rhône in the Crau and in the Camargue363, and in the Hérault at Roque-Haute260. L. virens occurs in the Vaucluse along the Durance (Papazian, pers. com.)

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Mediterranean temporary pools

Corsica

Lestes barbarus, a migratory and pioneer species, appears to be commoner than L. virens, with six stations out of the 50 visited during the survey of dragonflies in Corsica, compared with only two stations for L. virens. Both species are known in particular from Tre Padule and Padule Maggiore159, 323.

• The lakes of the Centre Var, the Plaine des Maures, the Palayson wood and the Colle du Rouet are included within Natura 2000* areas, as well as the Lanau pool, which was acquired in 1998 by the CEEP. Recommendations

Legal status of the species/level of protection

• Attempt to ensure, at temporary pools and fresh or brackish still waters, that there are flooded conditions until spring (AprilMay), suitable for the development of the larvae. • Maintain and promote aquatic vegetation and bands of vegetation around the edge of the pool, suitable for egg laying (rushes, sedges, water plantains, mints, bistorts).

Lestes barbarus and L. virens have no legal protection at the present time227.

Bibliography

Conservation status

No information.

IUCN Bern Convention Habitats Directive National and regional protection National Red Data Books and Red Lists

-

Conservation problems/threats Anthropogenic factors

Aguesse, 19614 ; Anonymous, 199711 ; Carchini & Nicolai, 198462 ; Corbet, 199989 ; D’Aguilar & Dommanget, 199895 ; Donath, 1981123 ; Grand & Papazian, 2000159 ; Heidemann & Seidenbusch, 2002173 ; MNHN, 1993227 ; Médail et al. 1993244 ; Moubayed, 1998260 ; Robert, 1958322 ; Roche, 1990323 ; Rombaut, 1994325 ; Samraoui & Corbet, 2000340 ; Samraoui et al., 1998341 ; Shiemenz, 1954350 ; Terzian, 1979363 ; Thiéry et al., 2002377 ; Utzeri et al., 1976388, 1984390, 1987387, 1988389 ; Wendler & Nüss, 1997402.

Drainage and infilling of the sites used. Natural factors

Siltation and overgrowing of wetlands; feeding competition with other dragonflies (e.g. Sympetrum fonscolombii), predation by water beetles, parasites, interannual variability of hydrological conditions (early drying out may result in the death of all larvae). Management and conservation measures Current measures

In France: • The Tre Padule de Suartone and Roque-Haute sites are legally protected due to their “Réserve Naturelle (Nature Reserve)” status (at Roque-Haute, the management of the site is compromised as there is no control over land ownership or the activities for which the site is used).

100

Authors: Jakob C. & J. Fuselier Collaborators: Papazian M., B. Samraoui & N. Yavercovski

Odonata

Sympetrum fonscolombii (Sélys, 1840) (1) & Sympetrum meridionale (Sélys, 1841) (2) Order: ODONATA Suborder: ANISOPTERA Family: LIBELLULIDAE

males

Main synonyms

(1) Libellula erythroneura Schneider, 1845 Libellula insignis Brittinger, 1850 (2) Libellula hybrida Rambur, 1842 Libellula nudicollis Hagen, 1850 Libellula meridionalis Sélys, 1841

1 cm

French names: (1) Sympetrum de Fonscolombe,

Sympetrum fonscolombii

(2) Sympetrum méridional English names: (1) Red-veined Darter, (2) Southern Darter Description/identification criteria

Sympetrum have six to eight veins in the forewings. The bases of the hindwings have a yellow mark which is fairly well-developed in S. fonscolombii (but often absent in S. meridionale). (1) In S. fonscolombii, the legs are black with a yellow stripe on the outer edge. Measurements of male: abdomen 22 to 29 mm, hindwings 26 to 30 mm. Measurements of female: abdomen 22 to 28 mm, hindwings 26 to 31 mm. (2) S. meridionale has yellowish or brown thorax with practically no black, and yellowish legs with scarcely any black markings. The pterostigma is pale yellow to reddish, edged with black veins. The abdomen is red in the male and brown-yellow in the female. Measurements of male and female: abdomen 22 to 28 mm, hindwings 20 to 30 mm95, 322, 401. Similar species

No confusion possible with Sympetrum fonscolombii, whereas S. meridionale may be confused with S. vulgatum ibericum and S. sinaiticum.

1 cm

Sympetrum meridionale

Habitat

(1) Sympetrum fonscolombii prefers still, occasionally brackish water (pools, ponds, gravel pits, marshes, boggy edges of lakes), from sea level up to 2,000 m (sometimes much higher for wandering individuals). It prefers wetlands that are very little vegetated (or vegetated with helophytes* and hydrophytes*), well exposed to the sun, and of very variable water depth.

Distribution - Ecology Distribution/range

(2) S. meridionale also prefers still water, more or less colonised by vegetation, up to 1,800 m altitude95, 173.

France

Ecology

(1) S. fonscolombii is found especially in the Mediterranean area, including Corsica, and in the Rhône Valley; it is much rarer in the rest of the country. (2) S. meridionale has the same range but appears to be commoner in the rest of the country122, 401. Other Mediterranean countries

The whole of the Mediterranean region95.

Biological characteristics

The eggs of Sympetrum often enter a diapause* phase after being laid, and do not hatch until they have spent the summer or winter in egg form89. However, in S. striolatum in Corsica it is also possible to observe larvae hatching out before the winter and these spend the winter in the water of the pool (Papazian, unpublished data.) In both cases, this type of annual cycle, with the eggs laid in autumn, corresponds to the flooding cycle of Mediterranean temporary pools340. S. meridionale and S. fonscolombii lay directly into the water (a highly evolved characteristic) but have partly retained the diapause at the egg stage.

101

Mediterranean temporary pools

(1) Adult S. fonscolombii show well-marked exploratory behaviour and undertake movements over great distances (migratory species)95, 122, which explains their sudden appearance in more or less large numbers322. The imagos are usually seen from March (an early species) to December, mainly from March to June and from August to December in temporary habitats. The mass movements of various Sympetrum that are observed in the countries around the Mediterranean are explained as being flights between pools which dry out in summer and aestivation sites338. In this species, there are at least two and even three generations per year in the Mediterranean region: the first emerges in March and, following a period of rapid maturation, lays eggs which hatch after very short incubation period; the larvae grow quickly before the water body dries up completely, and the emergence of the second generation takes place340 (Papazian, pers. com.). This second generation lays at the end of summer or in autumn, as soon as the water returns. In southern France, either the eggs (or a proportion of the eggs) of the second generation hatch quickly and the larvae spend the winter in diapause, or they do not hatch until the following spring, depending on biological and ecological factors (Papazian, pers. com.). In northeast Algeria (Numidia)340, embryonic or larval diapause does not take place; the second generation eggs hatch and the larvae undergo an autumn and winter growing phase which is prolonged due to the low temperatures; in addition, if there is water available in summer the second egg laying takes place earlier, and the species manages to lay for a third time late in the season; this is also the case in southern Europe in ricefields340. Laying usually takes place in tandem (the male still clasping the female). In full flight, the female, who like all Libellulidae has no ovipositor, taps the water surface with her abdomen: the eggs thus released into the water remain caught up on a support (an aquatic plant in most cases), before falling to the bottom. In France the larval stage of S. fonscolombii (first generation) is among the shortest observed among Odonata in France (along with Hemianax ephippiger). In the Mediterranean region, possibly after a diapause due to the drying out of the pool or to wintry temperatures in the water, the eggs hatch in a short time (rapid incubation). Emergence is usually nocturnal, extending from March in the Mediterranean region95, to November. (2) In southern regions, adult S. meridionale may be seen from the beginning of June (a late species) to mid-October95. At pools in northeast Algeria, Samraoui et al.341 have noted the presence of adult S. meridionale around the pools in May-June (immature imagos) and in September-October (mature). In temporary habitats, the adults have just enough time to emerge before the summer drought. They mature over the summer. At this time in northeast Algeria340, they disperse to refuges (woodland) at high altitudes where they slowly develop sexual maturity (imaginal diapause); adults may travel up to 35 km to aestivate at altitudes greater than 1,000 m. After the summer phase of maturation, the adults return, at the end of summer or in autumn, to the egg laying sites, which by now are usually flooded, to breed. In southern France (Papazian, pers. com.), the eggs may either hatch rapidly, the larvae then spending the winter in diapause, or delay hatching until the following spring. On the other hand in northeast Algeria larval

102

development continues until the emergence of adults in the spring of the following year; there is no embryonic diapause if water is available, but if it happens that the species lays in habitats that are still dry, the eggs can then remain resistant to desiccation until the first rains340. Contrary to S. fonscolombii therefore, S. meridionale only has one generation per year in temporary habitats (southern France, North Africa)341 (Papazian, pers. com.). The eggs (up to 550) are laid in places with shallow water, among reeds and Carex, even in puddles. S. meridionale is a migratory species like S. fonscolombii. The lifespan of the adults which survive to aestivation is among the longest known among Anisoptera. Simplified phenological cycle of Sympetrum fonscolombii Southern France Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

First generation

Dec.







Diapause of eggs

or larvae, followed by larval development Emergence of adults, sexual maturation, followed by egg laying and migration

Second generation Larval development, emergence of adults, migration Aestivation and maturation of adults Eggs laying and migration

Environmental conditions

Hydrology • Larval development: Aguesse4 showed that laying does not always take place at times and places that are favourable for the development of the larvae. Among Libellulidae such as Sympetrum, females are frequently seen laying eggs on the dry outer edge of a pool, which will not be submerged at a suitable time, or on the water surface at a pool which dries out a few weeks later. While larval development is jeopardized by the failure of the laying site to refill with water or by early drying out of the pool, these laying strategies remain nonetheless effective overall thanks to egg diapause (facultative), which takes place if the site is dry and which enables the eggs to survive the dry period. It should be noted that this group of Odonata has the widest geographical range4. • Adult diapause: For North Africa, Samraoui et al.341 described a prolonged, diapaused adult phase in S. meridionale (3-4 months during the summer dry period), when there is no reproductive activity. Von Hagen (in Samraoui et al.341) described the same phenomenon in S. striolatum in southern Spain. Papazian (pers. com.) has observed it in Corsica and in southern France.

`

Odonata

Conservation - Management

Management and conservation measures

Assessment of populations and conservation status

Current measures

Mainland France

Both species are fairly common near the Mediterranean, especially in the PACA region (more local in the north): • Var: Plaine des Maures244 and lakes of the Centre Var377; Sympetrum fonscolombii has in addition been recorded from the Colle du Rouet325, the Plaine de Palayson363, the oxidation ponds at Porquerolles and the Quinson reservoir on the Verdon, and S. meridionale on the Giens peninsula (Papazian, pers. com.). • Bouches-du-Rhône: S. fonscolombii is recorded from the Crau363 and the Camargue (Papazian, pers. com.) • Hérault: S. fonscolombii is known from the Roque-Haute plateau3.

In France, some sites are wholly or partly legally protected by their Réserve Naturelle status (Roque-Haute, Camargue, Les Tre Padule de Suartone); others are included in Natura 2000* areas (Colle du Rouet, Plaine des Maures, lakes of the Centre Var, Lanau pool), and among these some have been the subject of operations of land ownership control in the context of the LIFE “Temporary Pools” project (some parts of the Plaine des Maures and the Lanau pool). Recommendations

While Sympetrum fonscolombii is a common species in the French Mediterranean region, S. meridionale is much more confined to the coastal fringe, and the survival of these populations therefore depends above all on the conservation of coastal pools.

Corsica

Sympetrum fonscolombii occurs at six out of the 50 stations visited during the survey of the dragonflies of the island, and S. meridionale occurs at ten of these 50 stations323. Notably, both are present at Padule Maggiore and Tre Padule. Legal status of the species/level of protection IUCN Bern Convention Habitats Directive National and regional protection National Red Data Books and Red Lists

-

Bibliography AGRN-RH, 20003 ; Aguesse, 19614 ; Aguesse, 19685 ; Anonymous, 199711 ; Corbet, 199989 ; D’Aguilar & Dommanget, 199895 ; Dommanget, 1994122 ; Heidemann & Seidenbusch, 2002173 ; Médail et al., 1993244 ; Robert, 1958322 ; Roche, 1990323 ; Rombaut, 1994325 ; Samraoui & Corbet, 2000340 ; Samraoui et al., 1998341 ; Terzian, 1979363 ; Thiéry et al., 2002377 ; Wendler & Nüss, 1994401. Authors: Jakob C., T. Gendre & J. Fuselier Collaborators: Papazian M., B. Samraoui & N. Yavercovski

Conservation problems/threats Anthropogenic factors

(1) The presence of people at water bodies is a constraint on Sympetrum fonscolombii, which is very timid322. Natural factors

(1) Shading of water bodies and the growth of very dense aquatic vegetation is probably harmful to Sympetrum fonscolombii. (2) Male S. meridionale are often parasitized by mites (Acarus libellulae De Geer) 322.

103

Mediterranean temporary pools

Bufo calamita

Laurenti 1768 1 cm

AMPHIBIA BUFONIDAE Main synonyms

None French names: Crapaud calamite, Crapaud des joncs Spanish name: Sapo corredor English name: Natterjack Toad Subspecies

Bufo calamita

None Description/identification criteria

The length varies between 40 and 80 mm (sometimes up to 90 mm in the female). The build is sturdy with prominent paratoid glands* arranged in parallel, a horizontal pupil and pale greenishyellow iris, and a greenish to brownish dorsal surface generally crossed by a pale yellow mid-dorsal line125, 333. The breeding male develops brown nuptial pads on the toes. The species has a characteristic mode of locomotion: unlike most amphibians the Natterjack does not jump, but walks rapidly or runs. It is easy to spot thanks to its far-carrying song, which may be heard from up to a kilometre away. Similar species

The Green Toad (Bufo viridis) has no yellow dorsal line; it has handsome green markings contrasting with its ash-grey back. The Common Toad (B. bufo) has a golden-red iris.

Distribution - Ecology Distribution/range France

The Natterjack is found in all parts of France except for the high mountains (maximum 1,500 m in the Pyrenees) and Corsica. This species extends all the way into built-up areas in some towns. It is abundant in the south but its distribution becomes more discontinuous towards the north. Other Mediterranean countries

The Natterjack occurs in Spain (except for part of the northwest and the Pyrenees) and in Portugal168, but is absent from Italy.

Habitat

The Natterjack is typically an amphibian of open habitats, with short vegetation and few trees. It prefers loose substrates, either sandy (sandy back dunes) or stony (Plaine de la Crau, Plaine des Maures), degraded garrigue and maquis and also cool heathlands (Plateau de l’Aubrac). It adapts to man-made sites: parks and wasteland on the edges of built-up areas. Its aquatic habitat is fairly varied: live stock-watering ponds, flooded areas among sand dunes, puddles, Mediterranean wadis, quarry lakes, etc.

104

These habitats are typified by often shallow water (a few tens of centimetres at most), an absence of predators, and sparse or even completely non-existent aquatic vegetation. Ecology Biological characteristics

This is a pioneer species, which is able rapidly to colonise recently created habitats such as temporary pools. It prefers loose, sandy soils. In search of suitable habitats, Natterjacks may easily cover distances of more than 2.5 km268. The males may live for seven years while females can sometimes live for 17 years125. Like almost all European amphibians, the Natterjack has terrestrial and aquatic phases. Breeding usually starts in March in the Southern France with the young toads leaving the water at the start of summer (June), extending into July further north. In the south, a second breeding season is sometimes observed at the end of summer, depending on the storms of August and September. Young frogs from this breeding season emerge at the end of September or beginning of October. Males form choirs and sing in shallow water to attract the females. After mating, the adults quickly leave the pool, and the eggs, which are laid in strings, (25004000 at a time) hatch after a few days. Aquatic larval development is relatively brief (four weeks to three months). Their rapid development does not always protect the young Natterjacks from falling victim to early drying out184, 185. The juveniles are very small at metamorphosis compared with other species, but grow quickly. Sexual maturity is generally attained when they are three years old. The tadpoles, which are completely black, may feed on algae or other plants as well as detritus. They are sometimes cannibalistic268. Both adults and juveniles are predators of terrestrial insects.

Amphibians

Annual phenological cycle

Legal status of the species/level of protection

Southern France Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Dec.

Normal cycle Arrival Spawning Larval development then metamorphosis
hibernation

Departure

IUCN Bern Convention Habitats Directive National and regional protection


Aestivation by adults

then terrestrial phase and

National Red Data Books and Red Lists

Exceptional cycle Spawning Larval development hibernation

Departure, terrestrial phase and

Environmental conditions

Hydrology Flooding must take place at the latest at the beginning of May, the limiting date for breeding in the Mediterranean region. However, a brief period of flooding (two to five months) does not disrupt the process of larval development and the year’s breeding success. Substrate for spawning This is one of the rare species capable of breeding in the complete absence of any aquatic vegetation, in concrete-lined ponds, rocky wadis or bare sandy areas. Interspecific competition This species avoids temporary pools used by other species, especially the Green Frog (Rana perezi) or the Common Toad; the digestive tract of some species contain an alga which inhibits the larval growth of Natterjack Toad tadpoles191. Impact of perturbations This species is highly adapted to unstable habitats and as a result is very tolerant of perturbations in the aquatic environment. Its short larval development period, tolerance of high temperatures, ability to cover great distances to breed and production of a large number of eggs make it a species well-adapted to the risks of drying out and for the colonisation of new sites. On the other hand, it needs open habitats for feeding and moving around, and declines when the habitat becomes wooded. Shade The Natterjack prefers pools with strong insolation and rather short terrestrial vegetation, the larvae being capable of withstanding high temperatures, up to 40°C for brief periods.

Conservation - Management Assessment of populations and conservation status

The Natterjack Toad is fairly abundant in the Mediterranean region, rarer and more sparsely distributed in northern Europe125. It occurs at about 29% of Réserves Naturelles de France land and 10% of Réserves Naturelles Volontaires. However, if only those Réserves Naturelles located within its French range are included, it is present at about 48% of them135.

Annexe II Annexe IV - France, national list: Decree of 22/07/1993/Journal Officiel 09/09/1993 - Spain, national list293 - Portugal, national list59 - Europe: “to watch”26 - France, national Red Book: “to watch”242 - Spain, national Red Book: “low concern” (LC)293 - Portugal, national Red Book: “not threatened” (NT)59

Conservation problems/threats

The Natterjack Toad is a relatively little-threatened species in the Mediterranean region, aside from the impact of increasing amounts of woodland. Anthropogenic factors

The transformation of a shallow temporary pool into a deep or even permanent pool poses problems for this species. Such a pool can support greater numbers of invertebrate predators than a shallow pool, and also enables other amphibian species to breed, thereby impeding or preventing spawning among Natterjacks. However, it is able to breed in some places alongside the Western Spadefoot (Pelobates cultripes), Parsley Frog (Pelodytes punctatus), Midwife Toad (Alytes obstetricans), Marbled Newt (Triturus marmoratus), Great Crested Newt (T. cristatus), Palmate Newt (T. helveticus), Painted Frog (Discoglossus pictus) or Stripeless Tree Frog (Hyla meridionalis). This is the case for example at the Valliguières pool and the Opoul pool, where temporal segregation from the other species allows it to breed successfully. The introduction of fish causes the decline and disappearance of this species. Finally, the degradation, destruction or modification of the terrestrial habitat has harmful effects, by destroying the aestivation and hibernation sites which are essential for the survival of amphibians in the Mediterranean (stumps of trees or bushes, rocks, vegetation near the water bodies). Natural factors

If the aquatic and terrestrial habitats become overgrown through natural succession, this poses problems for pioneer species of open habitats like the Natterjack. The increase of terrestrial plant cover, in particular, has been proved to be a factor in its decline: on a very small scale, plant cover provides shelter, but on a larger scale it may form an obstacle to moving around, and a bare habitat is greatly preferred (see the Plaine de la Crau, where the species reaches extremely high densities!). Risks relating to populations

The disappearance of shallow temporary water bodies and the destruction of breeding sites increase the risk of local extinction as a result of the increasing isolation of habitats.

105

Mediterranean temporary pools

Management and conservation measures Current measures

Recent scrub clearing around shallow pools such as those at Roque-Haute (terrestrial vegetation) is improving the quality of the habitat and migratory routes. The overdeepening of pools, frequently undertaken in the case of pools which have become very shallow through aggradation*, has a negative impact on the population56, as does the creation of permanent pools next to shallow temporary pools35: both these measures favour other, competing species of amphibians.

methods and are often more prudent than deepening the pool. Population reinforcement may be achieved by creating a series of temporary pools with a range of varying depths around a site27, 35.

Bibliography

Recommendations

Beebee, 197724, 198325, 198526, 199627, 199728 ; Beebee et al., 199329 ; Biggs et al. 200135 ; Buckley, 200156 ; Cabral et al., 199959 ; Duguet & Melki, 2003125 ; Fiers, 1998135 ; Günther, 1996168 ; Jakob et al., 1998185, 2003184 ; Joly, 1994191 ; Maurin, 1994242 ; Nöllert & Nöllert, 1996268 ; Plueguezuelos, 2002293 ; Salvador & Garcia-Paris, 2001333

In most cases, the reinforcement of populations or the substitution of recreated habitats for destroyed habitats are effective

Author: Jakob C.

106

Amphibians

Discoglossus sardus

Tschudi, 1837

AMPHIBIA DISCOGLOSSIDAE Main synonyms

None French name: Discoglosse sarde Italian name: Rana Tirrenica English name: Tyrrhenian painted Frog

1 cm

Subspecies

None

Discoglossus sardus

Description/identification criteria

This Painted Frog is a small amphibian of slender appearance with a strong resemblance to a frog of the genus Rana. The males, which are slightly larger than the females, may reach 5 cm in length from snout to cloaca. The colour of the upperside is very variable, most often brownish but sometimes reddish, greyish or even greenish, more or less heavily marked with irregular dark areas all over the body. A pale band runs between the eyes across the top of the head. This pattern gives the animal the appearance of a dead leaf, which makes it very cryptic on the ground and in the water. The hind legs have pale and dark bands. The underside is a rather uniform pearly cream colour. The pupil is shaped like a water droplet and the eardrum is not visible. Eggs are laid in tens at a time, separately or in the form of a sheet at the bottom of the water. They are greyish or bicoloured, with a diameter of 1 to 1.5 mm, and are contained within a 3 to 4 mm gelatinous capsule. The tadpoles are small and dark-coloured, fairly similar to Bufo tadpoles. Like all the Discoglossidae, they have a ventral spiracle*.

Habitat

Similar species

Biological characteristics

The Tyrrhenian Painted Frog forms a species pair with the Corsican Painted Frog, with which it was confused until 1984. Compared with this species the hind leg is shorter: if extended along the body, the heel usually reaches as far as between the eye and the tip of the snout, but no further. The snout is more pointed and the part between the eye and the nostril is slightly sloping.

Distribution - Ecology Distribution/range

This Painted Frog is restricted to the French and Italian Tyrrhenian islands: Sardinia and some of its islets, Corsica and the Lavezzi Islands, the Tuscan Archipelago (Giglio and Montecristo Islands), the island of Monte Argentario, and the eastern Hyères Islands (Port-Cros and Levant). This isolation is undoubtedly very longstanding, such that each of these populations has evolved distinctive and specific characteristics. In Corsica, it ranges over a good part of the island, from sea level up to at least 1300 m. At PortCros it is found in all the streams on the island, notably the Vallon de la Solitude and the Vallon Notre-Dame. It seems to be rare on the Île du Levant where surveys are still required to ascertain its precise status.

This species lives in a wide range of habitats, from low maquis to mature oak or pine woodland. It is undemanding in its choice of breeding sites: springs, coastal marshes, canals, brooks and streams, depressions cut off from the main channels of rivers, reservoirs and fountains. In particular it uses impermanent bodies of water which in coastal regions are fed by unpredictable events (isolated puddles, temporary runnels, temporary pools, etc.). It is not uncommon for the eggs to fail to hatch, or to be dried out by evaporation or carried away by floods. This species can breed in relatively brackish water and, in contrast with the Corsican Painted Frog, can adapt to disturbed or modified habitats (polluted streams on the edges of villages, artificial reservoirs). Outside the breeding season, the species may be observed under piles of leaves or under trunks of fallen trees, several hundred metres from the nearest aquatic habitat. Ecology

This species is active for a good part of the year, by day as well as by night, but comes out mainly during wet weather in autumn (October-November) and at the end of winter (February-April). Breeding may be observed from October to April, usually following heavy rain (two to three nights after the rain starts). The males’ song is quiet, not very far-carrying and restricted to the breeding seasons. Males arrive at the breeding sites first and remain there longer than the females. Mating is brief and the eggs are laid in bundles of 20 to 50. The small tadpoles hatch out after about three days. These tadpoles grow rapidly and metamorphose after four to six weeks. Some, originating from eggs laid in autumn, grow much more slowly (20-25 weeks) during the winter. In Corsica as well as at Port-Cros, the eggs are laid over a very long period of time. Several generations of tadpoles coexist in a single place and it is common to see eggs, tadpoles of all sizes and juvenile frogs all present simultaneously. At metamorphosis individuals are very small (about 8 mm) and resemble adults in every respect. They reach 16 mm a year later, 30 mm after two years and 45 mm at the end of the third year. The lifespan may certainly extend to nine or ten years. The diet consists of small invertebrates caught on land: woodlice, spiders and various insects.

107

Mediterranean temporary pools

the species to be common during the 1950s-60s, but threatened by the deforestation of the island, the destruction of watering places and the impact of the Viperine Snake on the larvae and the adults. No surveys have been carried out since then to assess the current status of this population. Most of the occupied sites are situated on steep islands that are not very suitable for agriculture and are well preserved overall.

Phenological cycle Southern France Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Dec.

Usual cycle Extended egg laying Larval growth and metamorphosis

Legal status of the species/level of protection Departure and

terrestrial phase

Occasional cycle
Larval growth and metamorphosis


Egg laying (sometimes) then

IUCN Bern Convention Habitats Directive National and regional protection

Environmental conditions

Hydrology This species is perfectly adapted to the unpredictability of the Mediterranean hydrological regime (very extended breeding period, rapid development of larvae) reducing its vulnerability to climatic hazards. Aquatic habitat This Painted Frog uses almost all types of aquatic habitat for breeding, with the exception of flowing water and very large water bodies. This species ranges widely over the countryside, without specialising in any one type of habitat. Pools are therefore not essential for the survival of its populations, in contrast with other amphibians. A slight degree of salinity is tolerated, as well as a high degree of turbidity (muddy puddles). Organic pollution does not seem to have any affect on breeding. In general, this species uses small temporary water bodies, preferably exposed to sunlight, although a fairly large amount of plant cover may be tolerated. Terrestrial habitat Almost all kinds of habitat may be used: dense woodland, maquis, or partly built-up areas.

Conservation - Management

National Red Data Books and Red Lists

Annexe II Annexes II and IV - France, national list: Decree of 22/07/1993/Journal Officiel 09/09/1993 - Italy, regional lists (Sardinia, Tuscany): regional law of 29/07/1998 - France, national Red Data Book: “rare”242 - Italy, national Red Data Book14

Conservation problems/threats Anthropogenic factors

Owing to its lack of specialisation, the threats to this species are not very serious. Dams interrupt the distribution of populations and reduce the number of breeding sites; water pollution undoubtedly affects the eggs and larvae. One threat which cannot be ignored is that of forest fires, which as well as destroying adults have the effect of profoundly altering the hydrological regime of the watercourses (more torrencial regime) and the nature of the breeding sites (input of ash and sediment). Coastal populations are no doubt affected by anti-mosquito treatments (Corsica) and by the spread of the Italian Pool Frog (Rana bergeri), introduced probably recently into Corsica. The introduction of fish into closed water bodies (small reservoirs, ponds) is recognised as constituting a threat (for example the case of the Parc de Port-Cros). It is acknowledged that the introduction of alien competitive or predatory species in particular should be controlled (American crayfish, fish, frogs), as these will undoubtedly have catastrophic effects on this species.

Assessment of populations and conservation status Natural factors

Most of the populations are large, except on some islets which have practically no fresh water such as the Lavezzi islets. Corsica undoubtedly supports the largest numbers. In Sardinia this species seems to be less common and it is only known there from a small number of sites. There are no quantitative data available for the Tuscan archipelago, but according to Knoepffler (1962), the islands of Giglio and Montecristo have very large populations due to the presence of abundant fresh water. On Port-Cros, Knoepffler estimated the population size at around 5000 adults at the end of the 1950s. Recent observations indicate that numbers may be lower at present. The introduction some years ago of fish into the artificial reservoir where a high proportion of the island’s breeding adults were concentrated had reduced the breeding capacity of the population. The eradication of the fish in 2001 made the site once again available for the Painted Frogs, which bred there in numbers in 2002-2003. (Joyeux, pers. com.). On the Levant Islet, Knoepffler198 considered

108

Natural threats are few in number apart from the problem of fire, which may in some cases constitute a serious threat (small wooded islands such as Port-Cros). Reafforestation is known to constitute a threat to some populations by its effect on the breeding sites (shading of open water, and reduction of open water due to the accumulation of litter). Risks relating to populations

It is necessary to maintain sufficient numbers to counter the risk of chance extinctions, which is particularly high in an island situation. Management and conservation measures Current measures

The Tyrrhenian Painted Frog occurs at many sites in France and Italy which benefit from official protection either specifically for

Amphibians

wildlife (Parc National de Port-Cros, Nature Reserves of Etang de Biguglia, Scandola and Bouches de Bonifacio in Corsica, Parc Naturel Régional de Corse, Tuscan Islands National Park) or as a result of laws relating to land use (in France, sites belonging to the Conservatoire de l’Espace Littoral et des Rivages Lacustres and Forêts domaniales de Corse). The inclusion in the Natura 2000* Network of most of these areas, where the conservation of the species is usually taken into account in the management plan, is recommended. The species is the subject of research in the Parc National de Port-Cros and in Corsica. An inventory of the populations of the Ile du Levant is planned. Recommendations

• It would be useful to clarify the status of some populations which are still poorly known: the islets of the Tuscan archipelago (Italy) and the Ile du Levant (France). • On Port-Cros, it would be worthwhile to contain the growth of forest cover around the main breeding sites by selective streamside felling. In addition, it would be worth reassessing the dry

stone dams along the watercourses, which were constructed in the early years of the Park to hold back sediment: the piles of leaves accumulated over the years have had the effect of blocking the beds of the island’s streams, tending to cause the water to run underground, with consequent losses of the amount of open water available for use by the aquatic fauna. The number of breeding sites has decreased significantly as a result.

Bibliography Anonymous, 200414 ; Capula & Corti, 199361 ; Cheylan & Massemin, 199973 ; Cheylan, 198375 ; Delaugerre & Cheylan 1992103 ; Delaugerre 1999104 ; Gasc et al., 1997152 ; Knoepffler, 1962197, 1973198 ; Maurin, 1994242 ; Pesme, 2001287 ; Salvidio et al., 1997335, 1998337, 1999336 ; Vandenbrouck, 1996393

Authors: Cheylan M. & M. Delaugerre

109

Mediterranean temporary pools

Pelobates cultripes

(Cuvier, 1829)

AMPHIBIA PELOBATIDAE Main synonyms

None French names: Pélobate cultripède, Crapaud à couteaux Spanish name: Sapo de espuelas Portuguese name: Sapo-de-unha-negra English names: Western Spadefoot, Iberian Spadefoot Toad

1 cm

Pelobates cultripes

Subspecies

None Description/identification criteria

The Western Spadefoot is a fairly large amphibian (10-11 cm) with a squat appearance. The head is short and surmounted by large protuberant eyes. The eardrum is not visible. The skin is smooth, and variable in colour from greenish-yellow to brown, strongly marked with more or less contiguous brown blotches, giving each individual its own unique pattern. In the field it can be immediately recognised by its upright stance. There is little sexual dimorphism, apart from the presence in the male of a yellow lens-shaped protuberance on the forelegs. On the heel there is a down-turned black spade used for digging in the soil. The tadpoles, which are very large when fully-grown (up to 12 cm) are distinguished by a bulky oval body and a lead-grey to greenish colouration, with fine golden markings. The caudal crest* is lanceolate shaped, pointed at the tip and practically unmarked. When they leave the water the young measure on average 25 mm and weigh 1.5 to 2 g. The spawn takes the form of a long string, 12 to 20 mm in diameter and 1.1 metres long on average. Spawn from different females is often mixed up, tangled together at the edge of a water body, usually caught up in vegetation 0 to 20 cm deep. It may be distinguished from the spawn of “true” toads by the irregular distribution of the eggs within the string and by the larger number of eggs. Similar species

Impossible to confuse with other species, except in the case of the tadpoles.

Distribution - Ecology Distribution/range

This Spadefoot is found in the Mediterranean regions of southwest Europe: Spain, south and southwest France, as well as in Portugal. In France the populations form two distinct nuclei: a Mediterranean nucleus which extends eastwards to the Fréjus area in the Var département and to the Ardèche in the Rhône Valley, and an Atlantic nucleus extending from the mouth of the Gironde to the Guérande peninsula in the département of Loire-Atlantique. Habitat

The Spadefoot especially prefers open habitats: dry plateaux, sandy areas, degraded garrigue and maquis, grazed meadows and

110

Mediterranean arable land, up to 800 metres altitude in southern France and 1,700 m in the Iberian Peninsula. In Spain it is common in the dehesa* country typical of southwest Iberia and around the mouths of rivers (Marismas del Guadalquivir). The breeding sites are fairly varied: permanent or temporary pools, dune slacks, marshland ditches, concreted lavognes*, gravel pits, hill reservoirs. Ecology Biological characteristics

Like most Mediterranean amphibians, the Western Spadefoot is active especially in winter and spring, from November to May in southern Spain, from March to June and then from the end of August to November in France. Strictly nocturnal, they are especially active during rain, whenever the temperature reaches at least 8°C. In the daytime the animal remains buried in the soil at a depth of 6 to 20 cm, up to 40 cm or deeper in winter. The breeding season varies depending on the local climatic conditions. It extends from October to February in the south of the Iberian Peninsula and from the end of February to the beginning of May in central Spain. In southern France the breeding cycle is markedly bimodal with one more or less consistent breeding period in autumn (September to November) and another in spring, usually between the third week in February and the end of April, sometimes to the beginning of May. The onset of this event is triggered by continual rain over several days (20 mm), with an air temperature of 9 to 16°C. The breeding adults arrive at the breeding sites at dusk and remain under a few centimetres of water immediately next to the water’s edge. The largest males arrive first, sometimes before the females, and remain for one to eight nights. The females usually stay for one to three nights at the breeding place. During amplexus*, which lasts an average of 72 hours, the male and the female both give a call which resembles the clucking of a chicken. In Spain, the average number of eggs laid is 2318, while at a site at the Ile d’Oléron it is 960 eggs. The strings of eggs are wound around the vegetation and are laid in shallow water, usually less than 20 cm below the surface. The tadpoles feed mostly on plant material, both algae and higher plants, but also detritus. When food becomes scarce they may eat the tadpoles of other species or of their own species. Larval development usually takes three

Amphibians

and a half months for eggs laid in spring, up to eight months for autumn eggs. Metamorphosis takes place mostly in June and July. Some tadpoles do not metamorphose in summer and are still in the water in autumn. In this case, they probably do not undergo metamorphosis until they are one year old. The diet of the adults consists entirely of invertebrates: spiders, harvestmen, beetles, bugs, hymenopterans, butterflies and moths, grasshoppers, diptera, ants and insect larvae. Some individuals may live for 10-15 years; sexual maturity appears to develop at two years old. They have many aquatic predators: dragonfly larvae, water beetles, water snakes, herons. Phenological cycle Southern France Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Dec.

Usual cycle Arrival

Conservation - Management Assessment of populations and conservation status

The Spadefoot is currently declining over the whole of its range. This decline is due to the destruction of its breeding sites or terrestrial habitats, the widespread use of pesticides, and the increasing fragmentation of its populations. In France, the total number of known sites is not greater than 150. This species is particularly threatened at the edges of its range, especially in the Var département where only two out of the four historically known sites still remain, but also in the Bouches-duRhône, the Vaucluse and all the Atlantic coastal départements. Most of the breeding sites hold fewer than 100 individuals, a few tens being the norm. Six Conservatoire de l’Espace Littoral et des Rivages lacustres sites and four Nature Reserves support the species. In Spain, the situation is a cause for concern in most provinces. The decline seems to have been particularly steep over the course of recent decades. In many cases, the survival of the species depends on the preservation of a single breeding site.

Spawning Larval development and metamorphosis

Legal status of the species/level of protection

Departure, aestivation and terrestrial phase

Occasional cycle Arrival Spawning





Larval development and metamorphosis

IUCN Bern Convention Habitats Directive National and regional protection

Environmental conditions

Hydrology The length of the larval cycle constitutes a major constraint for this species, particularly in view of the unpredictability of the Mediterranean climate. Many breeding attempts are therefore doomed to failure, whether due to the breeding site drying out earlier than usual or to delayed flooding. Ideally, the site should start to fill with water at the end of October and remain flooded until the end of July. Aquatic habitat It is usually characterised by fairly deep water, most often deeper than 70 cm. The spawn is laid preferably on submerged vegetation, very close to the water’s edge, in water containing not more than 5 g of salt per litre (the tadpoles will tolerate slightly higher salinities, up to 10 g.l-1). The breeding sites are typically poor in aquatic vegetation: the highest densities of tadpoles are found in sites with hydrophyte biovolumes of less than 10 kg/m3. Terrestrial habitat The presence of open areas appears to be a major factor. Otherwise the habitat may be very varied: coastal dunes, dry grasslands, open heathlands, garrigue, oak or pine woodland, dry plateaux, arable land, on sandy, stony or even rock substrates. In sandy areas, the presence of rabbits appears to be favourable for the Western Spadefoot: it uses their burrows for shelter in the daytime. Grazing, especially by sheep, is also beneficial.

National Red Data Books and Red Lists

Annexe II Annexe IV - France, national list: Decree of 22/07/1993/Journal Officiel 09/09/1993 - Spain, national list293 - France, national Red Data Book: “vulnerable”242 - Spain, national Red Book: “almost endangered” (NT)293 - Portugal, national Red Book: “not threatened” (NT)59

Conservation problems/threats Anthropogenic factors

Impacts of an anthropogenic nature include the destruction of both terrestrial habitats (conversion to arable, urban development) and aquatic habitats (infilling of pools), the introduction of competitive or predatory alien species (fish, American crayfish, Marsh Frogs), the use of toxic chemicals (pesticides) and the destruction of the animals (road traffic). Coastal populations are especially affected by tourism developments and by anti-mosquito treatments. Further inland, it is the abandonment of farmland and the spread of woodland which are the two principal causes of decline. Natural factors

The main threats are due to weather conditions, either when the breeding site dries out before the tadpoles have metamorphosed, or when a sudden dry period lowers the water level to below the strings of spawn, which are usually laid at water level at the water’s edge. Some predators (notably Polecats) can have a major impact on the adults during the breeding season.

111

Mediterranean temporary pools

Risks relating to populations

Essentially, it is necessary to maintain high enough numbers to overcome stochastic risks, as well as a dense enough network of connections between populations to ensure interchanges of individuals and hence gene flow. Distances of less than a kilometre between sites would appear to be desirable. Management and conservation measures

setting up management agreements, etc. As far as management is concerned, the most important actions are the following: to control the introduction of undesirable species (Louisiana Crayfish, fish, terrapins etc.), to ensure that the hydroperiod* is compatible with the species’ breeding cycle, to control plant cover over an area of radius 100 to 200 m by grazing and/or scrub clearing, and to place shelters by the edges of the pools to promote the survival of the newly metamorphosed young.

Current measures

In France, if some of this species’habitats are situated in protected areas, the species is not the subject of specific conservation measures, apart from the monitoring protocols set up at some sites in Charente-Maritime, in the Camargue (Tour du Valat estate), at the Valliguières site in the Gard département, in the RoqueHaute Nature Reserve in the Hérault, in the Parc du Lubéron in the Vaucluse and at the Suze-la-Rousse site in the Drôme. Recommendations

It would be helpful, in the first place, to maintain the spatial cohesion of the breeding sites, if necessary by recreating suitable sites in recently abandoned areas. This could be attempted experimentally at a few sites: the Giens peninsula and the Plaine de Palayson in the Var, the Vallée du Calavon in the Vaucluse, the Plateau d’Aumelas in the Hérault. In addition, it would be useful to carry out some activities at certain breeding sites: providing information for owners and communes,

112

Bibliography Alvarez et al., 19908 ; Busack & Zug, 197657 ; Cabral et al., 199959 ; Cei & Crespo, 197165 ; Cheylan & Poitevin, 199974 ; Conservatoire de l’Espace Littoral et des Rivages Lacustres, 199884 ; Diaz-Paniagua & Arrizabalaga, 1987111 ; Diaz-Paniagua, 1983112, 1985113, 1986114, 1988115, 1990116 ; Dohogne, 1999120 ; Doménech, 1994121 ; Duguet & Melki, 2003125 ; Fiers, 1998135 ; Garcia-Paris, 1990147 ; Gasc et al., 1997152 ; Lescure, 1984209 ; Lizana et al., 1994214 ; Maurin, 1994242 ; Nöllert & Nöllert, 1995267 ; Petit & Lomont, 1958288 ; Pleguezuelos & Lizana, 2002293 ; Rodriguez, 1988324 ; Salvador & Garcia-Paris, 2001333 ; Salvidio & Quero, 1987334 ; Thirion, 1996378, 2001379. Author: Cheylan M.

Amphibians

Triturus cristatus

(Laurenti, 1768)

AMPHIBIA SALAMANDRIDAE

male

Main synonyms

None French name: Triton crêté Italian name: Tritone crestato English names: Great crested Newt, Warty Newt Subspecies

1 cm

Triturus cristatus

None Description/identification criteria

Large newt (total length of adults: 19 cm), dark brown or almost black in colour, finely spotted with white on the flanks. The underside is yellow/orange, marked with black. The skin is very characteristically warty. During the breeding season, the male develops a high, spiky dorsal and caudal crest. This has a silvery streak along its side. Females and juveniles have an orange dorsal line. The throat is greyish, spotted with white. Similar species

• Blasius’ Newt (originally described as a separate species, Triturus blasii) is a hybrid between the Great Crested and Marbled Newts. It is distinguished from the Great Crested by its ventral surface, which is greenish black above and yellowish with black markings and fine white spots below. • The range of the Italian Crested Newt (T. carnifex) is almost completely separated from that of the Great Crested Newt. Its feet are longer, its body is more elongated and its upperside is paler, showing many black marks; the male’s crest is lower and less deeply indented.

Distribution - Ecology Distribution/range France

This species is found throughout the northern half of France, north of a line from Charentes to Savoie. In the southern half of France it is currently known only from a few sites: one at Arles, Bouches-du-Rhône, in the Gard51, two in the Drôme and one in the southern Ardèche. Other countries

The range extends from Great Britain in the west to the Ural Mountains in Russia and, north-south, from southern Scandinavia to Austria and northern Romania. This species is absent from Spain and Portugal. In the southern Mediterranean countries, it is replaced by two vicariant species formerly treated as subspecies of Triturus cristatus: • T. carnifex (Laurenti, 1768) in Italy and the Balkans; • T. karelinii (Strauch, 1870) by the northwest coasts of the Aegean Sea (Turkey, Greece).

Habitat General description

The Great Crested Newt is a species of lowlands which lives mainly in fish-free permanent pools and/or temporary pools which dry out in summer. The three sites in the Mediterranean region take the form of: • a marlpit in a calcareous area covered with pine woods, garrigue with Holm Oak, heathland with juniper and vineyards (Valliguières in the Gard, see vol. 1, box 26), • a small quarry pool fringed with Yellow Flag Iris and reeds (Peyreguil, in the Gard), • a marshy area with willows and reeds, surrounded by built-up areas and degraded (Trinquetaille, at Arles). Ecology Biological characteristics

In the Mediterranean region, the only information concerning the species’ life cycle comes from the Etang de Valliguières. At this site the aquatic phase is variable in length between years, depending on the flooding period. In “normal” conditions, the newts arrive at the pool in November/December and leave again around May. The length of time spent in the water is about 8090 days33. The adults feed mostly on aquatic and terrestrial insects19. The larvae feed on aquatic prey (especially microcrustaceans). Breeding begins in April-May. The female lays 200 to 500 eggs per year248, placed one by one underneath the leaves of submerged plants which she then folds up with her hind feet (mainly species with broad leaves such as Ranunculus gr. peltatus, Glyceria fluitans, etc.). Embryonic development takes from two to three weeks depending on the water temperature. It is followed by a two- or three-month larval stage which ends with metamorphosis, usually at the end of the summer (mid-July to August, exceptionally September if the pool has not completely dried up). The young newts then leave the water and seek out a damp refuge in the surrounding area, where they hide and feed. After two years, in optimal conditions, they attain sexual maturity (males usually one year earlier than females140, and return to the water to breed. Immature T. cristatus tend to appear in the pool later than the adults. Adult survival is fairly high247: between 60 and 75% depending on the year. The maximum lifespan in the wild is from 14 to 18 years.

113

Mediterranean temporary pools

The Great Crested Newt is known for its fairly strong fidelity to breeding sites, local variations in breeding success being compensated for by its long life expectancy. The newts do not appear to colonise new breeding sites more than 400 m away from their natal pool20. Juveniles may disperse for up to a kilometre15, 202.

been subject to conservation measures for three years but its isolation, low numbers (about 200 individuals) and irregular breeding mean that its chances of survival in the medium to long term are questionable. The population at Peyreguil (Gard) is still poorly known. The presence of fish here constitutes a serious threat.

During the terrestrial phase, the individuals hide underneath stones, under pieces of wood or in crevices in the ground, a few hundred metres, or further, from the breeding sites. Fidelity to terrestrial sites appears to be linked to the different “scent complexes” which distinguish them, orientation being carried out by means of olfaction170.

Legal status of the species/level of protection IUCN Bern Convention Habitats Directive National and regional protection

National Red Data Books and Red Lists

Annexe II Annexes II and IV - France, national list: Decree of 22/07/1993/Journal Officiel 09/09/1993 - France, national Red Data Book: “vulnerable”242

Phenological cycle Southern France Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Conservation problems/threats

Dec.



overwintering (aquatic phase) ?

Arrival and (juveniles, adults) Eggs laying Larval growth Larval metamorphosis Departure, terrestrial phase and aestivation

Anthropogenic factors

The destruction of breeding sites (filling-in of pools), changes to the farmed environment resulting from new management practices, the isolation of different populations through habitat fragmentation203, and the introduction of competitive alien species (Marsh Frogs, fish, American crayfish) are the main causes of decline. Natural factors

Environmental conditions

Hydrology Successful breeding depends on the presence of water up to the end of July, which is essential for the metamorphosis of the juveniles.

The repeated drying-out of breeding sites may eventually threaten the survival of the population161; the effect of wild boar during the emergence of the young newts also represents a threat.

Management and conservation measures Current measures

Aquatic habitat The site must be relatively large and deep (>1 m) open to the sun and lacking fish. The presence of macrophytes* is essential for egg laying and for the development of an abundant invertebrate fauna; however, vegetation should not cover more than 20% of the water surface (maximum tolerated by breeding individuals)88. Terrestrial habitat The presence of wooded areas immediately adjacent to the site appears to be necessary for the survival of adults (and newly emerged young) during the terrestrial phase188. The presence of natural refuges (logpiles, piles of stones, stone walls) is also a favourable factor. Ditches and hedges provide biological corridors which facilitate dispersal.

Conservation - Management Assessment of populations and conservation status

Due to their isolation, the populations of Mediterranean southern France are highly endangered in the short term. The site at Trinquetaille (Bouches-du-Rhône) is situated within very degraded semi-urban surroundings. The population at Valliguières (Gard) has

114

The population of the Etang de Valliguières has been monitored since 2000216 as part of the LIFE “Temporary Pools” project. Among the planned work (see vol. I, Box 50) are included: • Digging out the pool in order to extend the period of flooding and thereby to encourage more regular breeding. • The creation of an artificial pool close to the existing site, in order firstly to encourage more regular breeding and secondly to create a second breeding site with the aim of improving the probability of survival of the population. • The control of invasive species (fish, crayfish, Marsh Frog). Recommendations

• Produce a complete inventory of the sites used by the species in the Mediterranean region. • Establish a network of potential breeding places in close proximity to one another (a few tens or hundreds of metres). • Control introduced species. • Maintain “biological corridors” around the pools. • Educate the public so that they do not fill in or bank up pools. • Control the impact of wild boar. For this, various solutions may be considered: installing electric fences around all or part of the site during the period when the young are emerging, digging out a substitute pool, planting bramble bushes around the pool to increase the number of refuges for the young newts.

Amphibians

• Urgently introduce conservation measures at the Trinquetaille/ Arles site in the Bouches du Rhône (translocation of individuals to a more suitable site could be considered).

199488 ; Francillon-Vieillot et al., 1990140 ; Griffiths & William, 2000161 ; Hayward et al., 2000170 ; Jehle, 2000188 ; Kupfer & Kneitz, 2000202 ; Laan & Verboom, 1990203 ; Lombardini et al., 2002 216 ; Maurin, 1994242 ; Miaud, 1990247, 1991248.

Bibliography

Authors: Lombardini K. & M. Cheylan

Arntzen & Teunis, 199315 ; Avery, 196819 ; Baker & Halliday, 199920 ; Besnard, 200233 ; Brogard et al., 199651 ; Cooke et al.,

115

Mediterranean temporary pools

Triturus marmoratus Latreille 1800 AMPHIBIA SALAMANDRIDAE

male

Main synonyms

None French name: Triton marbré Spanish name: Triton jaspeado Portuguese names: Tritão-marmorado, Tritão-verde English name: Marbled Newt

1 cm

Triturus marmoratus

Subspecies

T. m. pygmaeus in the Iberian Peninsula, recently considered to be a species in its own right. Ecology Description/identification criteria333, 405

Large newt (total adult length: 16 cm) black to brown in colour with green marbling on the back. Females and juveniles have an orange-red line down the spine. In the breeding season, the male develops a smooth crest with vertical black and yellowish-white bars. The underside is black, dotted with small white spots (orange-yellow with large black markings in the Great Crested Newt). Similar species

Pygmy Marbled Newt (Triturus pygmaeus), Blasius’ Newt (T. blasii), Great Crested Newt (T. cristatus). The Great Crested Newt differs in its orange underside marked with black. Blasius’ Newt is a hybrid between T. marmoratus and T. cristatus. It may show morphological characters of both species, but it is completely absent from the Mediterranean region. The Pygmy Marbled Newt, an endemic species of southern Spain, has a black underside like the Marbled Newt, but is smaller.

Distribution - Ecology Distribution/range

Biological characteristics

Like almost all the European amphibians the Marbled Newt has a bimodal cycle, with a terrestrial phase and an aquatic phase. At all stages of development, they are carnivorous. The larvae feed mostly on aquatic invertebrates405, the adults and juveniles are predators of tadpoles and of aquatic and terrestrial insects. The lifespan is long, up to 14 years in the wild345. In Mediterranean pools, the aquatic phase occurs mainly in winter owing to the unfavourable condition of the terrestrial habitat in summer. In southern France the period for which they are in the water extends from mid-November to mid-June, with considerable variation between sites and years. Breeding takes place in February-March in the south of France, October to May in the Iberian Peninsula. This phenology is slightly variable depending on interannual variations in the flooding period. Larval development takes a relatively long time (two to three months140). A single female may take several weeks to lay all her eggs. The eggs are laid one by one, in a fold in a leaf of an aquatic plant, with a preference for species with broad, flexible leaves such as Mentha pulegium (Pennyroyal), Ranunculus spp. or Callitriche sp. 9.

France

This species occurs in the western half of the country, as far north as the Seine. The eastern limit of its range lies in the Sommières area, in the Gard département. Other Mediterranean countries

The terrestrial habitat in evidently very important for the aestivation of amphibians in the Mediterranean as the animals run the risk of desiccation. Juveniles and adults hide under stones or among the roots of bushes. They do not move far, travelling partly underground during the summer drought.

Iberian Peninsula, north of the river Tagus, from the coast up to 2,100 m in the central mountains. Phenological cycle Habitat Southern France

The Marbled Newt, during its aquatic phase, is found in small bodies of still water, lacking fish: Mediterranean temporary pools (code 3170), shaded woodland pools, sometimes fed by springs333. The terrestrial habitat is varied: arable land or pasture, open grassy areas (Causses) and woodland.

Jan.

Feb.

March

April

May

June

July

Aug.

Sept.

Oct.

Dec.



Hibernation

Arrival and feeding (juveniles, adults) Egg laying Larval development

Larval metamorphosis Departure, then terrestrial phase and aestivation (juveniles, adults)

116

Nov.

Amphibians

Environmental conditions183, 184, 405

Conservation problems/threats

Hydrology Breeding sites must be holding water by February at the latest to allow the eggs to be laid. A flooding period of five months is then necessary for completion of larval development and successful breeding. Breeding failure in a given year does not constitute a major problem for the population, unless it recurs too frequently.

Anthropogenic factors

Aquatic habitat The water should be of a reasonable depth (about 0.8 m) and provide sufficient growth of aquatic vegetation to support the development of sufficient invertebrates for the feeding requirements of the larvae and adults. Terrestrial habitat It may be very varied in character (open heathland, garrigue, oak or pine woodland, arable or even built-up areas). Hedges and copses around the edge of breeding sites provide important terrestrial habitats and migration routes135.

Impacts of an anthropogenic nature include the destruction of habitats, both terrestrial (grubbing out of hedges and woods, destruction of old stone walls) and aquatic (filling-in of pools), the introduction of competitive or predatory alien species (fish, American crayfish, Marsh Frog (Rana ridibunda), and destruction of the animals (road traffic). Natural factors

Too short a period of flooding (i.e. through silting up) may prevent the completion of larval development, which will jeopardise the presence of the newt at a site. Risks relating to populations

The destruction of breeding sites and terrestrial habitats brings the risk of extinction by reducing the interconnections between populations.

Management and conservation measures

Conservation - Management Assessment of populations and conservation status

In Spain there are large populations which are relatively stable, apart from in the Ebro Valley and in north Catalonia where it appears to be more threatened. In France, its status gives no cause for concern although some local disappearances have been recorded, due mainly to the filling-in of pools or to land consolidation (destruction of hedges). It occurs at about 13% of Réserves Naturelles and 3% of Réserves Naturelles Volontaires. However, if only those Réserves Naturelles located within the newt’s French range are included, it occurs in 42%182.

Legal status of the species/level of protection

IUCN Bern Convention Habitats Directive National and regional protection

National Red Data Books and Red Lists

Annexe III Annexe IV - France, national list: Decree of 22/07/1993 Journal Officiel 09/09/1993 - Spain, national list293 - Portugal, national list59 - France, national Red Data Book: “vulnerable”242 - Portugal, national Red Data Book: “not threatened” (NT)59 - Spain, national Red Data Book: “low concern” (LC)293

Current measures

No specific measures are currently being applied at the RoqueHaute Nature Reserve (Hérault, France). The effect of scrub clearing around the pools (terrestrial vegetation) on the survival of adults and their migration routes remains to be assessed. Recommendations

• Maintain the cohesion of populations at the local and regional scales. • Control the introduction of fish and American crayfish into breeding sites. • Maintain well-developed aquatic vegetation. • Do not burn the area around pools or the sites used as refuges in the terrestrial phase. Dry years, when breeding does not take place, do not present real problems for this long-lived species, as long as the aestivation refuges enable good numbers of adults to survive. The maintenance of vegetation corridors around the pools is strongly recommended.

Bibliography Alvarez et al., 19899 ; Cabral et al., 199959 ; Fiers, 1998135 ; Francillon-Vieillot et al., 1990140 ; Gasc et al., 1997152 ; Hilton-Taylor, 2000176 ; Jakob, 2001182 ; Jakob et al., 2003183, 184, 1999186; Jehle & Arntzen, 2000187 ; Maurin, 1994242 ; Pleguezuelos et al., 2002293 ; Salvador & Garcia, 2001333 ; Schoorl & Zuiderwijk,1981345 ; Zuiderwijk, 1997405.

Author: Jakob C. Collaborator: Cheylan M.

117

Glossary Actinomorphic: (= regular) describes a flower, or a group of

identical organs within a flower, having radial symmetry. Active limestone: very fine particles of limestone in the soil,

which are easily dissolved by the action of carbon dioxide. The limestone ions thus released are capable of being absorbed by plants. Aggradation: Situation resulting from gradual infilling by accumulated material (soil, silt, sand, gravel, etc.). Allogamous: having a mode of sexual reproduction involving cross-fertilization among plants. Amplexus: among amphibians, mating without internal fertilisation. The eggs and sperm are emitted simultaneously and fertilisation is external. Depending on the species, amplexus may be lumbar (the male clasps the female around the lower part of the abdomen) or axillary (the male clasps the female under the armpits). Anemochory: mechanism of dispersal of seeds, spores, eggs, etc. of certain animal and plant species by the wind. Anemochory: mechanism of dispersal of seeds, spores, eggs, etc. of certain animal and plant species by the wind. Antheridium: organ in which the male gametes are produced, in algae, mosses, liverworts and ferns. Apoendemic: describes a new plant taxon which has been produced in a given region by polyploidisation (by becoming polyploid*) from diploid taxa* which are more or less widely distributed in the neighbouring regions. Association (phytosociological): see volume 1, chapter 2, box 4. Attenuate: width gradually decreasing (tapering). Auriculate: having auricles (in botany, small lobes situated at the base of a leaf, petiole, ligule, etc.). Bases: all the exchangeable positive ions in the soil which raise the pH*. Basionym: original name under which a species was first described. Basiphile: inhabiting an environment (soil, water) rich in bases*. Beak: in botany, a stiff pointed projection on the end of a dry fruit. Benthic: inhabiting the bed of a wetland or an ocean. Bioclimate: climatic conditions which characterise the various types of continental environment, either at the scale of natural regions of limited area or at the scale of stations occupying small areas. Biomass: total mass of living material, animal and vegetable, in a defined biotope at a given time. Bipinnatifid: used of a pinnatifid* leaf whose divisions are themselves pinnatifid. Bract: more or less modified leaf situated in the axil of a flower or an inflorescence. Bracteole: small bract situated in the axil of the ray of a branched inflorescence or in the axil of each of its constituent flowers. Brood pouch: see oviger Bryophytes: group comprising both mosses and liverworts. Capsule: in mosses and liverworts, designates a hollow dry-walled organ of the sporophyte, in which the spores develop. Among higher plants, designates a dry dehiscent fruit (i.e. opening spontaneously when ripe) containing more than one seed. Caudal crest: membrane on both sides of the tail in tadpoles (in the Anura) and larvae (in Urodela) used for propulsion. In some species, this crest extends up the back to the back of the head (Tree Frog tadpoles for example).

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Cercopods: (= cerci) in arthropods and crustaceans, appendages

located on the last abdominal segment (usually two in number, one on each side). Chamaeophyte: (= chameophyte) low woody plant whose buds are at least 25 cm above the soil (see Figure) Charophytes: specialised algal group consisting of one family, the Characeae, characterised by the whorled structure of the thallus and by the highly complex structure of the reproductive organs (antheridia and oogonia). Chitinous: formed of chitin, a molecule involved in the structure of the arthropod cuticle. Climax: used of a plant community which has developed to a sustainable state of equilibrium with the climatic and edaphic conditions of the environment, in the absence of human intervention. Clonal: used of an organism (animal or plant) derived from a single cell or a single individual, by asexual reproduction (known as vegetative propagation* in plants) Cohort: set of individuals which have experienced the same event at the same time (individuals born at the same time or breeding at the same time in pools, for example). Connectivity: facilitation of the movement of individuals of a species between local sub-populations to form a single functional demographic unit. Cordate: describes a heart-shaped flat organ. Coronule: short apical cells which develop a crown shape at the tip of the oogonium* in the Characeae. Cupular pool: small temporary pool located in bowl created in rock by natural erosion (see volume 1, chapter 2a). Cyme: branched inflorescence in which each flower is located at the end of a branch. Daya: a temporary pool in Morocco. Dehesa: Spanish name used to describe a landscape specific to certain regions in the centre and south of Spain, consisting of pasture or cereal cultivation scattered with evergreen oaks or Cork Oaks, which are usually cut. Detrivore (or Détritivorous): feeding on dead organic material. Diapause: period during which metabolic activity and the development of an insect is suspended at a particular stage (egg, larva, nymph or adult), as a result of the action of internal or external factors. Diploid: describes an organism whose cell nuclei have a double set of chromosomes (2n). Echinulate: covered in small spines. Ecophase: during its life cycle, a species passes through different stages (egg, larva, juvenile, etc.). An ecophase corresponds to one of these stages having a different ecology from the other parts of the cycle. Edaphic: refers to ecological factors linked with the soil as well as to their relationships with plants. Elater: elongated cell attached to a spore, and capable of flexible movements. Endemic: used of a species exclusively confined to a given biogeographical area, often of limited extent. Espace naturel sensible (Sensitive Natural Areas): areas bought by each département in France since 1985 “to preserve the quality of sites, landscapes and natural habitats” in their territories, using a tax levied on building. Eutrophic: used of water rich in nutrients (nitrates, phosphates) and with little oxygen at depth. Opposite of “oligotrophic*”. Eutrophication: process of enrichment of a water body with mineral nutrients (phosphates, nitrates) often accompanied by a

Glossary

proliferation of algae and/or aquatic higher plants, leading to oxygen depletion of the deeper water as well as a build-up of organic matter. Fistulous: hollow and cylinder-shaped. Gametangium: organ of the gametophyte* in which the gametes are formed. Gametophyte: in the life cycle of algae, mosses, liverworts and ferns, designates the haploid* phase of the organism, which produces gametes; in higher plants refers to the haploid organ which produces the gametes. Genetic bottleneck: sudden decrease in the size of a population associated with a decrease in the total genetic variability. Genetic bottleneck: sudden decrease in the size of a population associated with a decrease in the total genetic variability. Genetic drift: within small populations, chance fluctuations in gene frequency with each generation, enabling rare genes (which would be eliminated by selective pressure within large populations) to be continuously expressed. Geophyte: plant species which withstand the unfavourable season thanks to the presence of bulbs, rhizomes or any other type of underground reserve organ (see Figure). Gravel bed/Gravels: loose sedimentary formation consisting of rock fragments. Gyrogonites: calcified female fructifications produced by charophytes, corresponding to fossil or living forms after dispersal. They are invariably made up of five cells in the form of lefthanded spirals, joined at the tips. Halotolerant: tolerant of a certain amount of salt in the environment (water, soil, etc.). Haploid: organism or organ whose cell nuclei each contain a set of n chromosomes, comprising a single copy of each chromosome. Heliophilous: used of a plant which grows in conditions of strong sunlight. Helophyte: marsh plant whose budding parts, which enable it to survive during the bad season, are laid down in the sediment, while in the good season they develop an aerial structure which extends above the water surface (Reed, for example). Hemicryptophyte: perennial herbaceous plant whose budding parts, which enable it to survive during the bad season, remain on the surface of the soil, at the very base of the stems (or of the tuft for caespitose Graminae) (see Figure). Heterosporous: producing spores of two types (in algae, mosses, ferns and liverworts): microspores and macrospores. Hexamerous (6-merous): describes a plant organ the number of whose constituent equivalent parts (petals, sepals, stamens, etc.) is six or a multiple of six. Hexaploid: said of an organism whose cell nuclei contain three pairs of each homologous chromosome. Hybridogenic: taxon derived from a cross between two or several different taxa usually different species or sub-species. Hydroperiod: period during the year when the pool contains water. Hydrophyte: plant that lives in an aquatic environment (Water Milfoils, Water Lilies). Hydrophytic: refers to a hydrophyte* plant. Hygrophilous: organisms dependent on biotopes characterised by high soil water content. Karstic: relating to karst (the whole of the surface and subterranean structure of a limestone massif, which results from the dissolution of rock and is characterised by a wide-ranging system of underground water flow).

Laune (or lône): a local term meaning a former branch of a watercourse which can reconnect with the present watercourse when water levels are medium or high. Lavogne: local term meaning depressions dug out for watering livestock, in the Causses of the Cevennes and the Uzégeois and Montpellierais garrigues (France). Life form: five life forms (see Figure) were defined by Raunkiaer in his ecological classification of plants based on their strategy for surviving through the unfavourable season. Each life form is defined by the position of the buds in the architecture of the plant, as well as by their degree of protection during this season. Longevitous: living for a long time Macrophytes: generic term including all plants that are visible to the naked eye. Matorral: word of Spanish origin, in areas with a Mediterranean climate usually applied to a bushy vegetation community, adapted to drought and often dominated by evergreen species, often with small leaves. Meso-eutrophic: see Mesotrophic*. Mesotrophic: intermediate between eutrophic* and oligotrophic*. Metanauplius: larval stage following the nauplius* stage in crustaceans, characterised by the beginnings of body segmentation. Several moults, over a few weeks or a few months, are then necessary to attain the adult stage. Metapopulation: set of populations that are interconnected via migration events (gene flow) and are subject to extinction and recolonisation. This concept may be extended to include any set of populations developing in a more or less independent way which are, however, interconnected through rare instances of migration. Metathorax: posterior part of the thorax in insects, to which are attached the last pair of legs (and sometimes the second pair of wings). Microphagous: feeding on small prey. Mucronate: terminating abruptly in a sharp point. Natura 2000: ongoing European policy, with the aim (through the implementation of the Habitats Directive119 and of the Council Directive 79/409/EEC of 2 April 1979 on the conservation of wild birds) of reconciling human activities and natural habitats within a network of sites having great ecological interest, some designated for their diversity of birds (Special Protection Areas) others for the rarity of their fauna, their flora and their natural habitats (Special Conservation Areas). Nauplius: first larval stage of crustaceans, followed by the metanauplius* stage. Oligotrophic: describes nutrient-poor water (low in nitrate, phosphate, sulphate): its opposite is “eutrophic”. Oogonium: reproductive cell of lower plants (algae, mushrooms) which produces the female gametes. Ornithochory: mechanism of seed* or spore dispersal by birds. It may be external (carried on the body) and/or internal (absorption and passage through the digestive tract). Pappus: plume of hairs crowning an achene. Parotoid glands: glands located at the back of the head among some species of amphibians (true toads, salamanders), producing a poisonous secretion as a deterrent against predators. Perennial: in botany, describes a plant species which lives for more than one year (as opposed to annual). Phenology: description of the various phases in the life cycle of a species. Phytocenosis: collection of plant species having a homogenous appearance and colonising the same habitat (syn.: plant community).

119

Mediterranean temporary pools

Phytoplanktonophagous: feeding on phytoplankton. Phytosociology: see volume 1, chapter 2, box n° 4. Pinnatilobed: used of a divided leaf when the divisions cut into

the lamina for less than half its width. Pinnatipartite: used of a pinnate leaf when the divisions cut into the lamina for more than half its width. Pinnatisect: used of a pinnate leaf when the divisions cut completely into the lamina as far as the midrib. Pleurocarp: describes a bryophyte* whose female gametangia* (and hence sporophytes) appear in a lateral position and not at the ends of the gametophyte* stems. Polje: karstic* plain or subsidence valley, resulting from the coalescence of several karstic erosion systems (“ouvalas”) and covered in decalcified clays. A polje is subject to more or less prolonged seasonal flooding which sometimes occurs suddenly, depending on the underground karstic network. Polyploid: condition of an organism whose cells contain more than two sets of homologous chromosomes, i.e. more than 2n chromosomes. Pozzines: high altitude swards dominated by Carex, growing on a peaty substrate and mostly dotted with pools (Corsica, Pyrénées, Sierra Nevada, etc.). Pre-imaginal stage: final larval stage before the adult stage, in insects.

Prolarva: temporary larval stage (of very short duration) imme-

diately after hatching. Propagule: any part of an organism, produced by asexual or sex-

ual reproduction, capable of producing a new individual. Prothorax: in Odonata, the anterior part of the thorax, very

short, bearing the head and the first pair of legs. Pudding stone: consolidated sedimentary rock of the conglom-

erates group, of continental or marine origin, composed of pebbles bound together with a limestone or sandstone cement. Recruitment: the adding of new individuals to a population. Recruitment takes place through reproduction, immigration and restocking. Rhizoid: hair-like rooting structure, particularly among mosses. Scabrid: rough (covered in protrusions). Scape: leafless peduncle of a flower (or inflorescence). Sclerified: impregnated with lignin (component of the wall of certain cells, in particular in bushes, trees and shrubs, making it rigid and impermeable). Seed stock (of the soil): all the viable seeds and spores in the soil. Seedbank: all the viable seeds in the soil. Sessile: Botanical: an organ (leaf, flower) having no petiole or peduncle. Zoological: microorganism attached to a support (stem, rock, etc.). Sex-ratio: ratio of the numbers of male and female individuals in a defined population.

Figure. The main biological types, according to Raunkiaer**

Pistacia lentiscus

Artemisia molinieri

Eryngium pusillum

Ophioglossum lusitanicum

Isoetes histrix

Ranunculus revelieri

In perennial plants, position of the buds surviving the bad season

seeds

Chamaeophytes

Hemicryptophyte

Annual plants **: Raunkiaer, C, 1934. The life form of plants. Oxford, Clarendon Press. 632 pages

120

Cryptophytes Geophytes with bulb

Cryptophytes Géophytes with rhizome

Therophytes (surviving by seeds)

Perenial plants

N. Beck

Phanerophytes

Glossary

Sheath: a leaf base expanded to form a more or less cylindrical

Therophyte: synonym for an annual plant, a herbaceous plant

envelope, split longitudinally or not and clasping the stem. Sorus: group of sporangia*, arranged in a distinctive pattern on the lower surface of the fronds of some ferns. Spiracle: in tadpoles, opening which enables water to be ejected during respiration. In tadpoles of Painted Frogs, Midwife Toads and Yellow-bellied Toads, it is located in a median position on the underbelly. In other European amphibians it is situated on the sides of the abdomen. Sporocarp: in some ferns (Marsilea, Pilularia, etc.), a particular type of sorus*, closed and with tough walls, enclosing the spores. Sporophyll: in ferns, specialised leaf or frond, bearing one or more sporangia. Station (of plants): homogenous area of land where a plant species (or a particular plant community) is found. Stylopodium: small fleshy nectariferous disc which surmounts the fruit in Apiaceae (= Umbellifers), bearing two separate styles. Sympodic: describes the parts of a plant resulting from sympodial growth, i.e. lengthening of the stem in a succession of segments produced by lateral budding of the stems which have flowered in the previous year. Terra Rossa: type of soil found in Mediterranean regions, derived from the breakdown of limestones and characterised by an accumulation of ferric oxides.

with a very short reproductive cycle, lasting a few months or in certain cases a few weeks, which survives the bad season in the form of seeds. (see Figure). Therophytic: describes a therophyte*. Trophic: everything relating to nutrition among plants and animals. Vegetative reproduction: mode of reproduction of a plant species using vegetative organs (stolons, rhizomes, tubers, etc.). Vegetative stage: part of the life cycle of plants relating to germination, growth and propagation (vegetative reproduction), excluding flowering and fruiting. Verticil (whorl): all the homologous organs arising from a stem at the same point and radiating out from the stem. Vicariant: used of animal or plant species that are taxonomically closely related and which inhabit environments with similar ecological characteristics in different geographical regions. ZNIEFF: (Zone naturelle d’intérêt écologique floristique et faunistique) one of a list (begun in 1982 by the MEDD) of French natural areas, both terrestrial and marine, whose interest lies either in the stability and the richness of the ecosystem or in the presence of rare and threatened animals and plants.

121

Macrocrustacean inventory form Please send to: Danielle DEFAYE, Muséum National d’Histoire Naturel, Laboratoire de Zoologie - Arthropodes (Crustacés), 61, rue Buffon, F-75005 Paris

1. Essential data A) Source

or Biblio. ref. or code:

Observer or code:

or Coll. ref. or code:

Surname, first name: Adress:

Author(s): Date: Title:

Where deposited: Inventory number:

Name of form compiler if different from observer:

Periodical: Volume, number, page:

Name of form compiler if different from observer:

or

B) Taxon Number of reference list:

Genus

Species

Subspecies or variety Taxon

determined by: verified by: v.c.

C) Place

or

Coordinates – in grades (Paris meridian)

and

Commune + locality:

and

Altitude:

Longitude:  E, W

Name of pool or water body:

metres

At: INSEE code: IGN map number:

Latitude:

or coordinates – in degrees (Greenwich meridian)

D) Date

Longitude:  E, W ‘ °

‘’

Latitude °

‘’

‘

hr 1

hr 2

day

month

year

2. Additional data Type of contact:  seen  photo  captured/released  found dead  preserved  exuviae  ..............................

Abundance and stage: 1 2-10 +10    male    female    oviger    sexe = ?    juvenile    larvae    cyst

3. Additional information

Biotopes:  clay  sand  rock surface  clear water  turbid water  grassy ground  bare soil  algae  vegetation beds  ..............................

 salt marsh  salt water  fresh water

 Isoetes  Ranunculus  ....................

Method of collection:  by hand  shrimp net  drag net  plankton net  bird stomach  amphibian stomach  fish stomach  ..............................

(in note form to aid validation)

Depth of biotope: Duration of flooding: Colour on the living creature: Systematic notes, associated fauna, biometry, etc.:

Use reverse side of the form if necessary

122

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26. Beebee, T. J. C., 1985. Discriminant analyses of amphibian habitat determinants in south-east England. Amphib. Reptilia 6: 35-43. 27. Beebee, T. J. C., 1996. Ecology and Conservation of amphibians. Chapman and Hall, London. 214 pages. 28. Beebee, T. J. C., 1997. Bufo calamita in Atlas of amphibians and reptiles in Europe, J.P. Gasc & al. (Eds.), SEH & MNHN (IEGB/SPN), Paris. pp. 120-121. 29. Beebee, T. J. C., L. V. Fleming, & D. Race, 1993. Characteristics of natterjack toad (Bufo calamita) breeding sites on a scottish saltmarsh. Herpetol. J. 3: 68-69 30. Bernard, C., 1997. Flore des Causses (Premier supplément). Bull. Soc. Bot. Centre-Ouest. 28: 61-76. 31. Berner, L., 1968. Une herborisation à l’”étang” de Capestang (Languedoc). Monde Plant. 361: 5-6. 32. Bertrand, P. Y., 1991. Les noms des plantes au Maroc. Actes éditions, Institut Agronomique Vétérinaire Hassan II (IAVHII), Rabat, Maroc, 168 pages. 33. Besnard, A., 2002. Dynamique de la population de Triton crêté (Triturus cristatus) de l’étang de Valliguières (Gard). Une étude prospective. Ecole Pratique des Hautes Etudes, Montpellier, 30 pages. 34. Biche, B., 1881. Flore des environs de Roque-Haute, communes de Vias et de Portiragnes. Bull. Soc. Ét. Sci. Nat. de Béziers 5: 119-144. 35. Biggs, J., G. Fox, P. Nicolet, M. Whitfield & P. Williams, 2001. Dangers and opportunities in managing temporary ponds. Freshwater Forum 17: 71-80. 36. Bigot, L., 1955. Une Alismacée rare de Camargue : Damasonium stellatum (Lmk.) Rich. polyspermum Coss. Actes de la Réserve de la Camargue. Terre Vie 4: 238-243. 37. Bigot, L., 1999. Sur la réponse de Damasonium polyspermum Cosson (Alismataceae) aux variations édaphoclimatiques, d’après un suivi de 43 ans (1954-1996) dans une mare temporaire de la Tour du Valat (Camargue, Bouches-du-Rhône, France). Bull. Soc. Linn. Provence 50: 3-88. 38. Billy, F., 2002. Végétations pionnières en Basse-Auvergne. Bull. Soc. Bot. Centre-Ouest 22: 1-131. 39. Bissardon, M. & L. Guibal, 1997. CORINE Biotopes version originale : types d’habitats français. ENGREF, Muséum National d’Histoire Naturelle, Nancy, pp.32-37. 40. Blazencic, J. & Z. Blazencic, 2003. An overview of the existing data on living charophytes (Charales) of the Balkan Peninsula. Acta Micropal. Sin. 20/2: 103-110. 41. Boix D., J. Sala & R. Moreno-Amich, 2002. Population dynamics of Triops cancriformis (Crustacea: Branchiopoda: Notostraca) of the Espolla temporary pond in the northeastern Iberian peninsula. Hydrobiologia 486: 175-183. 42. Bonnier, G., 1990 (1934). Flore complète illustrée en couleur de France, Suisse et Belgique, 4 vol. Belin. 43. Botnariuc, N., 1947. Contribution à la connaissance des Phyllopodes Conchostracés de Roumanie. Nat. Biol., Bucarest 5(1-3): 68-158 + 10 pls. h. t. 44. Boudrié, M., 1995. Découverte d’Ophioglossum azoricum C. Presl. dans les Cévennes. Monde Plant. 454: 14-15. 45. Boudrié, M., H. Michaud, J. Molina & J. Salabert, 1998.- Les Ptéridophytes du département de l’Hérault. Monde Plant. 462: 11-19. 46. Boulos, L., 1995. Flora of Egypt, Checklist. Al Hadara Publishing, Cairo, Egypt, p 2. 47. Bournérias, M., G. Arnal & C. Bock, 2001. Guide des groupements végétaux de la région parisienne. Belin, pp. 178-181 et 195-200. 48. Braun-Blanquet, J., 1935. Un joyau floristique et phytosociologique “l’Isoetion” méditerranéen. Bull. Soc. Sci. Nat. Nimes, SIGMA 42(47): 1-43. 49. Braun-Blanquet, J., N. Roussine & R. Nègre, 1952. Les groupements végétaux de la France méditerranéenne. CNRS, Paris, pp. 81-82. 50. Briquet, J. & R. de Litardière, 1938. Prodrome de la Flore corse, vol 1/3. Lechevalier, Paris, pp. 87-89. 51. Brogard, J., M. Cheylan & P. Geniez, 1996. Découverte du Triturus cristatus Laurenti, 1768) (Amphibia, Caudata) dans la région méditerranéenne française. Bull. Soc. Herp. Fr. 80: 5-8. 52. Brtek, J. & A. Thiéry, 1995. The geographic distribution of the European Branchiopods (Anostraca, Notostraca, Spinicaudata, Laevicaudata). Hydrobiologia 298: 263-280. 53. Brullo, S. & M. Grillo, 1978. Ricerche fitosociologiche sui pascoli dei Monti Nebrodi (Sicilia settentrionale). Not. Fitosoc. 13: 23-61. 54. Brullo, S. & P. Minissale, 1998. Considerazioni sintassonomiche sulla classe Isoeto-Nanojuncetea. Itinera Geobotanica 11: 263-290. 55. Bucciarelli, I., A.P. Galetti & M. Pavesi, 1983. Attuali conoscenze sul popolamento odonatologico della Sardegna- Lav. Soc. Ital. Biogeogr. 8: 465-544.

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