lagunas mediterraneas temporales2

MediterraneanTemporary PoolsVolume 2Species information sheets

Grillas P., P. Gauthier, N. Yavercovski & C. Perennou

Production: Station biologique de la Tour du ValatDesign: Tapages Publics

Drawings: Antoine Catard, except pages 55, 83, 85, 88, 91 and 118Translated from French by Janet Clayton and John Phillips

Cover: photos Jean Roché

© 2004 Station biologique de la Tour du ValatLe Sambuc - 13200 Arles - France

Readers are invited to reproduce texts and drawings featured in this publicationprovided 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. PerennouAssociate editors: Jakob C., H. Michaud, G. Paradis & L. Rhazi

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), PapazianM. (SOF), Paradis G. (Université de Corse, ad. perso. 2), Perennou C. (TDV) Pozzodi Borgo M. L. (OEC), Rhazi L. (université Hassan II), Rhazi M.(TDV and IMEPUniversité 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, Aix-Marseille I), Titolet D. (Lycée international Georges Duby), Yavercovski N. (TDV).

ADENA (Fuselier J.)Association de Défense de la Nature des Pays d’AgdeDomaine du grand Clavelet, F-34300 AgdeTél.: +33 (0)4 67 01 60 23, fax: +33 (0)4 67 01 60 [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 PortiragnesTél/fax: +33 (0)4 67 90 81 [email protected]

CBNA (Garraud L.)Conservatoire Botanique National Alpin de Gap-CharanceDomaine de Charance, F-05000 GapTél: +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 centralLe Bourg, F-43230 Chavaniac-LafayetteTél: +33 (0)4 71 77 55 65, fax: +33 (0)4 71 77 55 [email protected]

CBNMP (Michaud H, Molina J.)1. siège Conservatoire botanique national de PorquerollesCastel Sainte Claire, F-83418 Hyères cedexTél: +33 (0)4 94 12 82 30, fax: +33 (0)4 94 12 82 [email protected]. Unité de conservation et collections (Michaud H.)Le Hameau, F-83400 Ile de PorquerollesTél.: +33 (0)4 94 12 30 32 , fax: +33 (0)4 94 12 30 30Courriel: [email protected]. Antenne Languedoc-Roussillon (Molina J.)Institut de Botanique, Rue Auguste Broussonet, F-34090 MontpellierTél.: +33 (0)4 99 23 22 11, fax: +33 (0)4 99 23 22 [email protected]

CEEP: Conservatoire Études des Écosystèmes de Provence Alpes du Sud-Var1, place de la Convention, F-83340 Le LucTél: +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 LucTél: +33 (0)4 94 50 38 39, fax: 04 94 73 36 [email protected]@wanadoo.fr

CEN-LR (Gendre T., Garnéro S., Houssard C., Klewscewski M.)Conservatoire des Espaces Naturels du Languedoc-Roussillon20 rue de la République, Espace République, F-34000 MontpellierTél.: +33 (0)4 67 22 68 28, fax: +33 (0)4 67 22 68 [email protected]

CELRL Corse (Delaugerre M.)Conservatoire des Espaces Littoraux et des Rivages Lacustres Délégation Corse3, rue Luce de Casabianca, F-20200 BastiaTél: +33 (0)4 95 32 38 14, fax: +33 (0)4 95 32 13 [email protected]@conservatoire-du-littoral.fr

EPHE (Cheylan M., Jakob C., Lombardini K.)Ecole Pratique des Hautes EtudesLaboratoire de Biogéographie et Ecologie des vertébrés, Case 94, Université de Montpellier II, Place E. Bataillon, F-34095 Montpellier cedex 5Tel: +33 (0)4 67 14 32 90, fax: +33 (0)4 67 63 33 27 [email protected]@[email protected]

Lycée international Georges Duby (Titolet D.)200, rue Georges DubyF-13080 [email protected]

OEC (Pozzo di Borgo M. L.)Office de l'Environnement de la CorseAvenue Jean Nicoli, F-20250 CorteTél.: +33 (0)4 95 45 04 00, fax: +33 (0)4 95 45 04 01

SFO (Papazian M.)Société Française d’Odonatologie7, rue Lamartine, F-78390 Bois d’ArcyFax: +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 ArlesTél: +33 (0)4 90 97 20 13, fax: +33 (0)4 90 97 20 [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 6116Université d'Aix-Marseille III, Faculté des Sciences et Techniques de Saint-Jérôme, Case 461, F-13397 Marseille cedex 20Tél: +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 VictorHugo, F-13331 Marseille cedex 3Tél: +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 Mont-pellier II, C.P. 062, Place E. Bataillon, F-34095 Montpellier cedex 5Tél: +33 (0)4 67 14 39 78, fax: +33 (0)4 67 14 30 [email protected]

Université d’Annaba (Samraoui B.)Laboratoire de recherche des zones humides, Université d’Annaba,4, rue Hassi-Beïda, Annaba, [email protected]

Université Hassan II (Rhazi L.)Faculté des Sciences Aïn Chock, Laboratoire de Biologie et PhysiologieVégétale, BP 5366, Maarif Casablanca, MoroccoTél.: (212) 037 86 33 10, fax: (212) 022 23 06 [email protected]

ad. perso. 1 (Hugonnot V.)Le Bourg, F-43270 Varennes Saint HonoratTél/Fax: +33 (0)4 71 00 23 [email protected]

ad. perso. 2 (Paradis G.)7, cours Général Leclerc, F-20000 AjaccioTél: +33 (0)4 95 50 11 [email protected]

Aknowledgements The Station biologique de la Tour du Valat would like to warmly thank all theeditors, authors and everyone who collaborated on this volume as well asMohand 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 (DIRENCorse), Guy-François Frisoni (Réserve Naturelle des Bouches de Bonifacio), JeanGiudicelli (Maison régionale de l’eau, Barjols), Denis Gynouvès (ONF Var), Jean-Claude Heidet (CEEP), Bruno Julien (European Commission), Emilio Laguna(Generalitat de Valence, Espagne), Nicolas Leclainche (TDV), Olivier Limoges (Pôlerelais Mares et Mouillères), Stéphanie Lieberherr (CEEP Var), Gilles Loliot (CELRL-LR), 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), EricParent (Agence de l’Eau RMC), Jean-Claude Pic (TDV), Marc Pichaud (TDV),Marlène Savelli (OEC), Pierre Quertier (ONF Var), Bertrand Sajaloli (Pôles relaisMares 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.

Editors, associate editors, authors and collaborators

5

Summary

Introductory notice . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

Plant species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

Apium crassipes (Koch ex Reichenb.) Reichenb. fil. . . . . 7

Artemisia molinieri Quézel, Barbero & Loisel . . . . . . . . . 9

Crypsis schoenoides (L.) Lam . . . . . . . . . . . . . . . . . . . . . . 12

Damasonium polyspermum Coss. . . . . . . . . . . . . . . . . . . 15

Elatine brochonii Clavaud . . . . . . . . . . . . . . . . . . . . . . . . 18

Eryngium pusillum L. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

Illecebrum verticillatum L. . . . . . . . . . . . . . . . . . . . . . . . 23

Isoetes duriei Bory (1) & Isoetes histrix Bory (2) . . . . . . 26

Isoetes setacea Lam. (1) & Isoetes velata A. Braun (2) . 30

Littorella uniflora (L.) Ascherson . . . . . . . . . . . . . . . . . . 34

Lythrum borysthenicum (Schrank) Litv. (1)

& Lythrum tribracteatum Salzm ex Sprengel (2). . . . . . 37

Lythrum thymifolium L. (1)

& Lythrum thesioides M. Bieb. (2) . . . . . . . . . . . . . . . . . . 41

Marsilea strigosa Willd. . . . . . . . . . . . . . . . . . . . . . . . . . 45

Mentha cervina L. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49

Myriophyllum alterniflorum DC. . . . . . . . . . . . . . . . . . . . 52

Nitella opaca (Bruzelius) Agardh . . . . . . . . . . . . . . . . . . 55

Ophioglossum azoricum C. Presl (1)

& Ophioglossum lusitanicum L. (2) . . . . . . . . . . . . . . . . . 58

Pilularia minuta Durieu ex A. Braun . . . . . . . . . . . . . . . . 62

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

Macrocrustaceans. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83

Imnadia yeyetta Hertzog, 1935. . . . . . . . . . . . . . . . . . . . 83

Linderiella massaliensis Thiéry & Champeau, 1988 . . . . 85

Tanymastix stagnalis (Linnaeus, 1758) . . . . . . . . . . . . . . 88

Triops cancriformis (Bosc, 1801) (1)

& Lepidurus apus (Linné, 1758) (2). . . . . . . . . . . . . . . . . 91

Odonata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94

Ischnura pumilio (Charpentier, 1825) (1)

& Ischnura genei (Rambur, 1842) (2) . . . . . . . . . . . . . . 94

Lestes barbarus (Fabricius, 1798) (1)

& Lestes virens (Charpentier, 1825) (2) . . . . . . . . . . . . . 96

Sympetrum fonscolombii (Sélys, 1840) (1)

& Sympetrum meridionale (Sélys, 1841) (2) . . . . . . . . . . . . 99

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

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

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

Pelobates cultripes (Cuvier, 1829). . . . . . . . . . . . . . . . . . 108

Triturus cristatus, (Laurenti, 1768) . . . . . . . . . . . . . . . . . 111

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

Glossary. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116

Macrocrustacean inventory form . . . . . . . . . . . . . . . . . . . . . . 120

Bibliography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121

Introductory notice

This second volume of “Mediterranean Temporary Pools” pre-sents, in the form of information sheets, a summary of the cur-rent knowledge on a selection of the remarkable species ofMediterranean temporary pools (29 plant species, 16 animalspecies). This knowledge is either bibliographical in origin or wasacquired during the LIFE-Nature project “Conservation ofMediterranean Temporary Pools No. 99/72049”, presented onpage one of volume 1.

The species presented have been selected according to a numberof criteria:• Their dependence on temporary aquatic habitats;• Their rarity (this criterion is, however, very variable from coun-try to country);• The Mediterranean nature of their distribution;• Their presence at one or more of the seven sites of the LIFEproject;• Their biological characteristics or their ecological require-ments, illustrating various adaptive strategies.

Plant speciesAmphibious vascular plants, flagship species of temporary pools,are the best represented with 26 species, i.e. almost a quarter ofthe 107 rare plants of the oligotrophic* temporary pools censusedin 18 countries of the Mediterranean Basin (see volume 1, table 2). A discreet and often poorly known group, the liverworts aredescribed on two information sheets. One is devoted to a veryrare species of temporary pools (Riella helicophylla), the other toa 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 temporarywetland areas.

MacrocrustaceansOf the fifty or so species of the Mediterranean Basin, five havebeen selected from three different Orders. Two species are veryrare in France (Linderiella massaliensis is endemic, Imnadiayeyetta is present on two sites). Water requirements differamong the species chosen (ephemeral habitats for Tanymastixstagnalis, semi-permanent habitats for Lepidurus apus and Triopscancriformis). Twelve Mediterranean countries support one ormore of these species.

OdonataIn the Odonata, the 25 species with a Mediterranean distributionhave life-history characteristics which are adapted to temporaryflooding and can vary according to the latitude. The six specieschosen illustrate restricted (Ischnura genei, endemic to theislands of the western Mediterranean) to wide (Sympetrum fon-scolombii) Mediterranean distribution as well as several strate-gies of adaptation to summer drought, based on the mechanismand dates of egg laying, the existence of diapauses (egg or larva),the periods of emergence of the adults, the number of genera-tions per year, wether or not migratory, the greater or lessercapacity to colonise new pools, etc.

AmphibiansAmong the sixty or so species found in the Mediterranean Basin,five which are adapted to temporary pools have been chosen intwo different groups (anurans and urodeles), illustrating the variousenvironmental requirements and biological characteristics: dura-tion of larval cycle, size of clutches, etc.).

Information sheet headings

Latin name: current nomenclatureClassification: except otherwise stated, the class then the familyof the species (current nomenclature) are mentioned (in smallupper-case letters) immediately under the title of each informa-tion sheet.Main synonyms: synonyms used in “classic” guides to flora andfauna.Common names: when they exist, the common French, Spanish,Portuguese, Italian, English, Moroccan and Algerian names aregiven. The names used at a national scale are given, with theexception of Morocco, where certain local names of oral tradi-tion are known265, though it is not possible to give any preciseregion where these are used. For European countries, the FrenchInternet site www.tela-botanica.org was consulted. In addition,the French names of the plant species were researched in themajor flora of Bonnier42 and in the Livre Rouge de la FloreMenacée de France (French Red Data Book of threatened flora)274,the Spanish and Portuguese names in Flora Iberica64, the Italiannames in the Italian Red Data book of threatened flora86 and inFlora d’Italia292. Subspecies: the subspecies currently recognised are mentioned.Description/identification criteria: a variety of bibliographicsources have been used, as well as the personal observations ofthe authors.Similar species: the species with which some confusion is pos-sible are mentioned, as well as their distinctive characteristics.

Distribution - Ecology

Distribution/rangeThe distribution of the species in France, then in other Medi-terranean countries is given whenever possible, in particular in thecountries of the western Mediterranean (Spain, Italy, Maghreb).

HabitatThis paragraph contains, for France, and if possible, for otherMediterranean 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 theHabitats Directive12, 119, accompanied by their Natura 2000* code; The mention of French habitats defined according to the CORINEBiotopes39 typology;• For the flora: the list of the phytosociological communities*characterised by the species or to which the species belongs, andwhere possible the main companion species.

EcologyThis paragraph provides the main biological characteristics andenvironmental requirements useful for the conservation man-agement of the species.

6

7

Biological characteristics

In France, and if possible in other Mediterranean countries, infor-mation relating to the biology and life-history traits characteris-tics of the species is given. • For the flora: the biological type, methods of sexual reproduc-tion (aerial, aquatic), description of seeds, and the biologicalcycle (germination, growth, flowering, fruiting, dispersal).• For the fauna: feeding, the site and the mechanisms of eggdepositing, the biological cycle, (reproduction, egg laying, devel-opment), longevity, distinctive behaviour, interannual and spatialvariations in the phenology.

On each information sheet, a figure at the end of this paragraphillustrates the annual phenological cycle of a population and anaverage hydrological state of the pool.

Example: the phenological cycle of Marsilea strigosa

generally flooded phase (or of saturation of the soil forcertain species)

flooded phase some years

dry phase

This schematic illustration cannot take into account all interan-nual variability in the Mediterranean climate (temperature, pre-cipitation) or individual variability in the response of organisms,but it can provide the manager with an average state. Someauthors express the variability of the response of a population tovariations in the hydrological regime by dots extending the solidline of the different phases of a cycle.

Environmental conditions In France, and if possible in other Mediterranean countries, thefollowing are given (within the limits of the information avail-able):• For the flora: the hydrological conditions required (depth ofwater, duration of flooding and drying out, germination of seeds,chemical characteristics, etc.); the nature of the substrate(acidic, calcareous, granulometry, etc.); sensitivity to interspecificcompetition (in relation to light in particular); the impact of per-turbations such as cutting, grazing, trampling by livestock, etc.• For the fauna: the breeding requirements (duration and dateof flooding, depth, type of pool, etc.); the territories, the winter-ing or summering habitat and the necessity for complementaryhabitats (terrestrial phase of amphibians, for example); the dis-persal distance of the adults and the young after the aquaticphase; environmental perturbations and their impact (rain,drought, soil humidity, etc.); interspecific relations (competition,predation, etc.); the main factors in natural mortality, etc.

Conservation - Management

Assessment of populationsIn France, and in other Mediterranean countries, the number ofsites is indicated (with, for the fauna, an estimation of the pop-ulation numbers, if possible.

Conservation status The status of the populations (stable, expanding or regressing) isprovided, as well as the main sites of recent appearance of thespecies or of disappearance.

Legal status of the species/level of protectionA 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 Franceand in other Mediterranean countries: - The protection status at national level for the flora was exam-ined for Italy86, 292, Spain (Internet site: www.mma.es/conserv_nat/acciones/esp_amenazadas/html/catalogo/flora.htm), France, Alge-ria (decree no. 93-117 of 23/11/1993 fixing the list of protectednon-cultivated plant species). - The protection status at national level for the fauna was exami-ned 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 faunawas examined for France, and in some regions of Italy for thefauna (Sardinia and Tuscany).• Inclusion in the Red Data Books and Lists in France and inother Mediterranean countries:- For vascular flora, the national Red Data Books and Lists ofFrance274, Italy86, Spain6, Morocco133, Greece290, and Malta344 wereconsulted as well as the regional Red Data Book for the Balea-rics331. For the bryophytes, the Red Data Book of the bryophytesof Europe128, and the Red Data Book for France, which is cur-rently being produced, served as references. - For the fauna, the National Red Data Book242 was consulted forFrance, as well as the Red Data Lists for Spain (amphibians293),Portugal (amphibians59) and Italy (vertebrates14).

Conservation problems/threatsThe following factors are examined: • Anthropogenic factors: destruction, infilling, drainage, pollu-tion, 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 wellas the management measures in place.

RecommendationsThese concern the management measures to put in place(hydrology, grazing, scrub clearing), modifications in the protec-tion status of the species, protection of the habitat, etc.

Jan. Feb. March April May June July Aug. Sept. Oct. Nov. Dec.

Emergence of fronds and vegetative growth

Withering of fronds

Formation of sporocarps?

France

Introductory notice

8

Bibliography

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

Other information

Acronyms and abbreviations used in the information sheets

CBNBP: Conservatoire Botanique National du Bassin Parisien.CEEP: Conservatoire Etude des Ecosystèmes de Provence - Alpesdu Sud.CEN-LR: Conservatoire des Espaces Naturels du Languedoc-Roussillon.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

Journal officiel: Journal Officiel de la République Française (offi-cial bulletin for the public, giving details of laws).s.l.: sensu lato (in the broad sense)subsp.: subspeciessp.: Indeterminate species (from the Latin species). For example,Isoetes sp. signifies an indeterminate species of the genus Isoetesspp.: several species of the genus. For example, Isoetes spp. sig-nifies “all or some species of the genus Isoetes”.em.: amended by (abbreviation of the Latin emendavit, from theLatin verb emendare: to modify, amend, improve). Used in phy-tosociology, for example: “the alliance Isoetion Br.-Bl. 1931 ofthe order Isoetalia Br.-Bl. 1931 em. Rivas-Goday, 1970”. µS.cm-1: micro-siemens per cm (unit of measurement for con-ductivity of water)

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

Mediterranean temporary pools

9

Plant species

ANGIOSPERMS

APIACEAE

Main synonymsHelosciadum crassipes Koch ex Reichenb.A. inundatum (L.) Reichenb. fil.var. crassipes Paol.A. inundatum (L.) Reichenb. fil subsp. crassipes LandiSium crassipes (Reichenb.) Sprengel

French name: AcheItalian name: Sedano di Sardegna

Subspecies None

Description/identification criteria50, 292

• Herbaceous plant, hydrophytic* or amphibious, eitherhemicryptophytic* or annual.• Stem hollow and grooved, at first briefly prostrate and radi-cant, then upright, erect or floating, capable of growing up to 40 cmlong in water.• Leaves alternate, of two types: basal leaves submerged, two-to four- pinnatisect* and completely divided into long, slenderstrips; emergent leaves simply pinnatisect with toothed seg-ments.• 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 thanthe 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 pedicelswhich are very swollen below.

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


Distribution/range

Apium crassipes is considered to be a Tyrrheno-Sicilo-Numidianendemic species50.

FranceThe species only occurs in Corsica.

Other Mediterranean countriesOccurs in Italy (Sardinia, Pontins marshes and Sicily292) and inNorth Africa (Tunisia and eastern Algeria). Casper & Krausch63

include the Nile Delta in the range.

Habitat

1. “Mediterranean temporary pool” habitat

General description

In Corsica and Sardinia, Apium crassipes is an important con-stituent in terms of the biomass* of the flooded phase of manytemporary pools. In southern Corsica, A. crassipes, together withvarious floating species (Ranunculus peltatus, R. ophioglossi-folius, Illecebrum verticillatum, Baldellia ranunculoides), is abun-dant in the temporary pools of Padulellu, Tre Padule de Suartoneand Padulu219, 221, 284. In Tunisia, A. crassipes is associated with Isoetes velata, Myosotissicula, Illecebrum verticillatum, Eryngium barrelieri (E. pusillum),Ranunculus ophioglossifolius298.

Habitats Directive“Mediterranean Temporary Pools” (code 3170).

CORINE Biotopes22.34 Mediterraneo-Atlantic amphibious communities (Isoetalia).

22.341 Short Mediterranean amphibious swards (Isoetion).22.3412 Aquatic Mediterranean Quillwort swards: com-munities containing Isoetes velata in water bodies ofvariable depth.

Phytosociology*In its aquatic form, Apium crassipes belongs to formations of thesubmerged phase of the Isoetion, within communities of Isoetesvelata. Some authors include the hydrophyte* formations inwhich A. crassipes is found within the communities of the Potamo-getonetea pectinati class284.

2. Other habitats

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

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

Apium crassipes1 cm

10


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

Phytosociology*In Algeria, around the Mafragh, Tonga and Oubeira lakes, Géhuet al.156, for seasonally flooded meadows resulting from aggrada-tion, created the association with Paspalum distichum and Apiumcrassipes which they named Helosciadi crassipes-Paspaletum dis-tichi and which they included in the class Agrostietea stoloniferae,the order Plantaginetalia majoris and the alliance Paspalo-Agrostion.

Ecology


Life formHydrophyte*, either hemicryptophytic* (perennial) or therophytic*(annual).

ReproductionAerial.

FloweringIn Corsica, from mid-March to late April.

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

Description of achenesLength: c. 1.25 to 1.5 mm.

Phenological cycle


Assessment of populations

In Corsica, probably over 20.

Conservation status

In CorsicaThe populations fluctuate from one year to another, but appearto be stable in the medium term at the Padulellu, Tre Padule andPadulu pools. On the other hand, as a result of the cessation ofcattle grazing several decades ago, the Arasu, Mura dell’Undaand Muratellu pools now have small populations. The populationof the Chevanu temporary pool, which is very bare and where thesubstrate is very compacted, is heading for extinction.

Legal status of the species/level of protection

Conservation problems/threats

Natural factorsIn Corsica, the likely cessation of extensive cattle grazing in thenear future threatens to cause a severe fall in numbers at almostall 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, Ericascoparia, Myrtus communis, Pistacia lentiscus, etc.).


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

Recommendations

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

Bibliography

Anonymous, 199912 ; Bissardon & Guibal, 199739 ; Briquet & deLitardière, 193850 ; Casper & Krausch, 198163 ; Géhu et al., 1994156 ;Grime, 1979163 ; Lorenzoni & Paradis, 1997219, 2000221 ; Paradis etal., 2002284 ; Pignatti, 1982292 ; Pottier-Alapetite, 1952298.

Authors: Paradis G. & M. L. Pozzo di Borgo Collaborators: Grillas P. & N. Yavercovski


GerminationGermination

Vegetative growth

Flowering

Fruiting

��

Corsica

IUCN 1997Bern ConventionHabitats DirectiveNational and regional protectionNational Red Data Books and Red Lists

-----

Environmental conditions

HydrologyFlooding in winter, even of short duration, appears to be essen-tial for germination or the reactivation of buds and for thegrowth of young plants.

SubstrateLoose substrate, quite thin, (less than 10 cm deep), of varyingparticle size (sand, silt or clay) and acid.

Interspecific competitionA heliophilous* (light-demanding) plant, which does not thrive inthe shade of tall species.

Impact of perturbations• A low level of disturbance (by grazing, trampling), whichcauses a limited degree of substrate compaction, appears to bebeneficial.• Heavy, frequent disturbance, for example due to all-terrainvehicles, that causes severe substrate compaction, are very detri-mental (Chevanu pool, in Corsica).

11

Plant species

ANGIOSPERMS

ASTERACEAE

Main synonymsNone

French name: Armoise de Molinier

SubspeciesNone

Description/identification criteria • Sub-woody plant, very aromatic, with the same scent (likecamphor) as Artemisia arborescens, 30 to 60 cm tall.• Stem branching above the neck. Leaves bluish-green, finelyhispid, especially on the lower sides. Petiole equal in length tothe leaf, which is triangular in outline, bipinnatifid* with thread-like lobes 1-1.5 mm wide.• Capitula fairly small, ovoid (2.5 x 2.5 mm), with 10-15 floretsarranged in a dense spike. Inner involucral bracts lanceolate-spatulate. The achenes (0.8-1 x 0.2-0.3 mm) are greyish with fiveill-defined angles, smooth, slightly longitudinally grooved, andcut off somewhat obliquely at the apex*.

Similar species The genus Artemisia is difficult. Historically, A. molinieri has beenconfused with A. alba Turra31, which belongs to another section.A. molinieri belongs to the Artemisia section which is charac-terised by its capitula having a glabrous receptacle and beingallogamous* with the outer florets female and the disk floretshermaphrodite, all fertile. Within this section, A. molinieri is closeto the species with green leaves and with the petiole lackingbasal auricles*. Two of these species may be found in France:• Southernwood, (A. abrotanum L.), a cultivated wormwood whoseorigins are disputed, and which is distinguished from A. molinieriby its lemon scent, larger globose capitula (3-4 x 3-4 mm) with25-30 florets arranged in a leafy panicle, and its oval innerinvolucral bracts.• A. chamaemelifolia Vill. of dry grassland in the mountains ofsouthern Europe and beyond, from Spain to Iran. This wormwoodis distinguished from A. molinieri by its sessile paired caulineleaves with the lower lobes clasping the stem, and its slightlylarger capitula: 2.5-3 mm.

A. molinieri, A. chamaemelifolia and some populations of A. abro-tanum have 2n = 18.

All the wormwoods are found in dry habitats (rock, steppes, rub-ble etc.), apart from A. molinieri which is the only wetland worm-wood.

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


Distribution/range

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

Habitat

General descriptionLarge natural depressions of karstic* origin, flooded irregularlydepending on the year: flooding varying from more than sixmonths to completely absent.


CORINE Biotope22-342: Tall Mediterranean amphibious swards (Preslion cervi-nae Br.-Bl. 1931), with a type specific to this habitat: theArtemisia molinieri community245, 307.

Phytosociology* Class: Isoeto-Nanojuncetea

Order: IsoetaliaAlliance: Preslion cervinae

Association*: Artemisia molinieri community307

Companion species At Gavoty and Redon:• Schoenoplectus lacustris, Scirpus maritimus, Phalaris arundi-nacea, Alisma lanceolatum, Eleocharis palustris, Oenanthe globu-losa, Rorippa sylvestris, Ranunculus ophioglossifolius, Sisymbrellaaspera, Veronica anagalloides.• Annual amphibious companion species of high natural-heritagevalue (for France), subject to competition from the wormwood

Artemisia molinieri Quézel, Barbero & Loisel

Artemisia molinieri

1 cm

12


and favoured by the opening-up of the wormwood mat (by tram-pling associated with grazing, or cultivation during dry years):Lythrum tribracteatum, Damasonium polyspermum, Heliotropiumsupinum, Schoenoplectus supinus, Chenopodium urbicum, Crypsisschoenoides, etc.

Ecology


Life formArtemisia molinieri is a competitive perennial species (chamaeo-phyte*), which forms dense species-poor stands. The few gaps,deriving from past disturbance, permit the development of adiverse annual flora.

Biological cycle Vegetative growth begins in February with the appearance of thefirst green leaves. Its reproductive cycle takes place in summer(from mid-July to the end of August). The seeds begin to bemature from mid-September. The last green leaves wither inOctober/November.The plant produces runners, also perhaps layers, and it also readilypropagates vegetatively. Couteaux & Pons93 considered that vege-tative reproduction was the plant’s only means of propagation. The plant is anemphilous382 but the pollen is very poorly dispersedbecause the stamens remain contained within the corolla of thehermaphrodite florets*93. In addition, the pollen has a very low via-bility (10 to 30% of pollen viable depending on the population)382.However, the plant produces abundant achenes, which have nopappus* but are very light and are without doubt wind-distributed.Their production is irregular among individuals and stations307, andgermination rates are extremely low382. This latter trait must berelated to the irregularity of pollen production in a taxon whoseorigin probably derives from hybridisation*93, or to the demandingecological conditions with which the wormwood is confronted(grazing, flooding)382. This is no doubt compensated for by the largenumber 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.

SubstrateColluvial silts of anthropogenic origin (erstwhile erosion of culti-vated catchment areas)93. Meso-eutrophic* substrate.

Interspecific competitionArtemisia molinieri is heliophilous*, and the establishment ofwoody plants (especially Fraxinus angustifolia) during dry years,or of helophytes* during wet years, is probably harmful to it (nodata).

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


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 thou-sand aerial shoots).The cultivation of part of Lake Redon, in 2001, destroyed a signi-ficant part of the station*. The Artemisia molinieri population hassince proved to be healthy, many germinations having been seenduring the subsequent years. There are several other depressions, which seem capable of havingsupported the wormwood in the past307, within a radius of a fewkilometres from Gavoty and from Redon. Drainage operationsand intensive cultivation could have wiped it out here. At threeof these depressions, on fallow land, the recent discovery of smallpopulations of A. molinieri (from one to under ten stalks) raisesthe question of whether these may be relict or alternatively pio-neer populations. Moutte & Triat261 found wormwood pollen inthe sediment at a fossile lake northwest of Brignoles. They hypo-thesised that there was formerly a population of A. molinieri atthis lake, which is several kilometres away from the currentlyknown populations. However, Couteaux & Pons93 questioned thishypothesis.


IUCN 1997

Bern Convention

Habitats DirectiveNational and regional protection

National Red Data Books and Red Lists

“endangered” (E)399

-

-France, PACA regional list:Decree of 09/05/1994/JournalOfficiel 26/07/1994 France, national Red DataBook: priority species274


Leafing and vegetative growth

Flowering

Fruiting


Anthropogenic factors The deforestation and subdivision of catchment areas leads tothe infilling of depressions and changes in water quality. The culti-vation of pools in dry years probably constitutes a risk factor.


HydrologyThe 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 orabsent. • A maximum water depth of 20 to 70 cm, with the optimumapparently around 40 cm.

ReproductionFlowering, which is late, is aerial.

13

Plant species

Natural factorsAt Lake Gavoty, woody plants (ash, elm) are becoming establishedas a result of natural succession. This may eventually transformthe depression into woodland, not susceptible to flooding andincompatible with the survival of a healthy Artemisia molinieripopulation.

Risks relating to populationsThe risk of extinction at any site is very low in both of the pop-ulations known for a long time, which are very large (see vol. I,chapter 3, box 27).


Current measures • Management plan in progress in the context of the productionof the objectives document for Natura 2000* sites, including thetwo main pools with Artemisia molinieri.• Land acquisitions completed and in progress at Redon, in theframework of LIFE “Temporary Pools”. • Collection and ex-situ preservation of seeds by the Con-servatoire Botanique National Méditerranéen de Porquerolles, andprecautionary cultivation of the species on Conservatoire landsince 1995.

Recommendations• Research into the role of disturbance in the dynamics and con-servation of formations containing Artemisia molinieri and ofother priority habitats (Lythrion tribracteati and Heleochloion)subject to competition from the wormwood.

• Taxonomic study of A. molinieri, whose status as a species andas an endemic has been questioned. This type of study may onlybe 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 Ulmusminor and Blackthorn Prunus spinosa) around the pools, at theedge of the flooded zones. This plays a vital role as a naturalwater filter before it discharges into the depression and as a trapfor fine materials which could cause the pools to silt up.

Bibliography

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., 1964-1980386 ; Walter & Gilett, 1998399.

Authors: Michaud HCollaborators: Rombaut D. & N. Yavercovski

14


Crypsis schoenoides (L.) Lam.

ANGIOSPERMS

POACEAE

Main synonymsHeleochloa schoenoides (L.) Host ex Roem.

French names: Crypside ovoïde, Crypside faux choinItalian name: Brignolo ovatoEnglish name: Swamp Pricklegrass

SubspeciesNone

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, slightlyswollen sheaths*, ligule divided into hairs, blade 1 to 3 cm longand 3 to 4 mm wide, at an angle to the stem.• Inflorescence a cylindrical spike, 5 to 8 mm in diameter and 1to 3 cm long, comprising many spikelets each consisting of oneflower with three stamens.

Similar species Young stages of Crypsis aculeata and Cynodon dactylon, but atthe 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 tinycapitulum buried among the bracts*, with two stamens per flower.


Distribution/range

The taxon is considered to be Paleo-subtropical292.

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

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

SpainScattered throughout the country; most frequent in the centre177.

MoroccoTingitan peninsula, northwest Morocco, Middle Atlas.

AlgeriaTell region.

Habitat

General descriptionCrypsis schoenoides may be found in summer and early autumn,on the dried-out clay-silt edges of mesotrophic* ponds, on thebanks of a few hydro-electric reservoirs (Corsica), and in tempo-rary pools (in France, in Morocco in the dayas of the plains andlow mountains etc.). It sometimes occurs on the silty edges ofwatercourses (Corsica, Algeria), in ditches, and also in poachedpastures subject to flooding.Its optimal living conditions are clay-silt substrates which undergoan alternation of a lengthy flooded phase and a drying-out phaseduring summer (sometimes up to the beginning of autumn). The maximum water depth may vary between a few tens of cen-timetres (pools or ditches) to several metres (hydro-agriculturaldams). This species will tolerate a slightly saline substrate.


CORINE Biotopes22.343 Mediterranean halo-nitrophilous swards (Heleochloion). “Post-aestival slightly halophilic and nitrophilous vegetation oftemporarily flooded land, with Crypsis schoenoides, C. aculeata,C. alopecuroides and Centaurium spicatum”.

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

Crypsis schoenoides (formerly named Heleochloa schoenoides) ischaracteristic of Heleochloion formations.

In Corsica and Spain Class: Isoeto-Nanojuncetea

Order: NanocyperetaliaAlliance: Heleochloion

Associations* and communities given in the literature:- Crypsio (aculeatae)-Heleochloetum schoenoidis

Oberdorfer 1952 (southern mainland Europe, Corsica).

Crypsis schoenoides

1 cm

15

- Heleochloo schoenoidis-Fimbristyletum biumbel-lattae Br.-Bl. & Rivas Goday in Rivas Goday 1956corr.54 (Spain).

- Heliotropio supini-Heleochloetum schoenoidis RivasGoday 1955 (Spain, Corsica).

- Lythro flexuosi-Heleochloetum schoenoidis RivasMartinez 1966 (Spain).

- Crypsio schoenoidis-Cyperetum micheliani MartinezParras et al., 1988 (Spain, Corsica).

- Chenopodio chenopodioidis-Crypsidetum schoenoidisParadis & Lorenzoni 1994 (Corsica).

- Echinochloo cruris-galli-Crypsidetum schoenoidisParadis & Lorenzoni 1994 (Corsica).

- Community with Crypsis schoenoides and Corri-giola litoralis (in Paradis & Lorenzoni277) (Corsica).

- Community with Crypsis schoenoides and Cotulacoronopifolia (in Paradis & Lorenzoni277) (Corsica).

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

In Morocco Crypsis schoenoides is most often found with Corrigiola littoralis,Heliotropium supinum and Hypericum tomentosum.

Notewhereas the Isoetion communities appear in spring and are olig-otrophic*, the Helochloion communities appear in summer-autumnand are meso-eutrophic*.

Ecology


Life formTherophyte* (annual species).

ReproductionAerial.

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 floodingand by animal grazing during the summer and early autumn.Stress results from the drying-out of the soil during the summerand early autumn, which is variable depending on the topogra-phy and on edaphic* characteristics.

CaryopsesLength: 0.5 mm maximum.

Biological cycle • At the end of spring or the beginning of summer: germinationof caryopses followed by seedling development during the dryperiod (date variable, depending on climate and hydrology), on asubstrate which is still damp or under a very shallow layer ofwater (less than 10 cm).

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


HydrologyAlternation between a long period of flooding in winter-springand summer drying appears to be essential.

SubstrateMeso-eutrophic* substrate with fine particles (clay and silt), richin organic debris and in nutrients, particularly nitrates. It issometimes oligotrophic*, in the dayas* of Cork-Oak woods inMorocco. It may be slightly brackish at the surface (often moresaline deeper down), which explains the presence of Tamarix spp.close to many coastal Crypsis schoenoides stations. There is usu-ally a fairly significant degree of substrate compaction.

Interspecific competitionAs a light-demanding (heliophilous*) species, Crypsis schoenoidesdoes not readily tolerate shading by tall species such as Phrag-mites australis, Typha latifolia, T. angustifolia, various Scirpus orPaspalum distichum, hence its restriction to bare areas.

Impact of perturbationsThe potential for the establishment and survival of Heleochloionplant communities depends on denudation, generally due to humanactivity:• Deliberate cutting of helophytes*, to create shooting areas forexample.• Grazing and trampling by cattle: trampling by cattle causes afairly high degree of substrate compaction, which prevents tallhelophytes from establishing themselves and does not hamper thegermination and growth of the therophytes of the Heleochloion.• etc.



In CorsicaThis species occurs around ten sites:


Growth

Flowering

Fruiting

Germination

Corsica


Vegetative growth

Flowering and fruiting

Dispersal of caryopses

Germination

Morocco

Plant species

16


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

In mainland FranceIt occurs at more than twenty stations: • Pyrénées-Orientales: Villeneuve-la-Raho reservoir and alluvialplain of the Agly. • Aude and Hérault: Capestang, Poilhès, Ouveillan, Estagnolponds and elsewhere in the lower valley of the Aude. Sesquiersmarsh at Mèze, pools of Notre-Dame-de l’Agenouillade at Agde,pools and ditches at Roque-Haute, Saint-Martin-de-Londres andSaint-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-du-Valat, Verdier marsh).• Var: Lake Redon at Flassans, Badelune basins at Cannet-des-Maures, pools at Grimaud and Hyères, ditches at Roquebrune-sur-Argens. No recent records from Fréjus.

In MoroccoMore than twenty stations are known from the Tangier, Larache-Casablanca and Middle Atlas regions.

Conservation status

In FranceThe population at Tanchiccia (Corsica) appears to be decreasingin comparison with previous observations by Paradis281. On theother hand, all the other populations on the island are either sta-ble or increasing.In mainland France, populations are large and sometimes in veryhigh numbers in the region of Capestang, Ouveillan, Poilhès, etc.The species appears to have decreased on the coast in the Pyrénées-Orientales and has disappeared from the Alpes-Maritimes coast332.It is surviving well elsewhere: the number of new sites largelycompensates for unconfirmed sites mentioned in earlier records.It seems to have appeared in the Camargue and in the floodplainof the Rhône between Arles and Tarascon as a result of thedesalination which followed the increase of ricefields256.

In MoroccoThe populations appear to be fairly stable despite the increasingpressures on the sites.



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

Natural factorsThis plant is favoured by sheep or cattle grazing (France, Morocco)as well as by certain types of management for hunting, such asthe creation of pools in reedbeds.

Risks relating to populations In mainland France, the isolation of the Var populations and ofthe very small inland pools in Languedoc-Roussillon constitutesa potential threat.


Current measures • In Corsica: at present there are no management measures inplace, due to the absence of control over land ownership at thesites.• In mainland France: the objectives document for the Natura2000* site pools of the Centre Var takes this species into account,but no conservation measures are known for the other sites. TheConservatoire Botanique Méditerranéen de Porquerolles collectsseeds and preserves them (ex-situ).• In Morocco: the species is not protected and does not benefitfrom any management measures.

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

Bibliography

Anonymous, 199912 ; Bissardon & Guibal, 199739 ; Braun-Blanquet et al., 195249 ; Brullo & Minissale, 199854 ; Grime,1979163 ; Loisel, 1976215 ; Maire, 1952-1987230 ; Martinez Parraset 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 Lorcaet al., 1988286 ; Pignatti, 1982292 ; Rivas Goday, 1964318, 1970319 ;Rivas-Martínez et al., 2001320 ; Salanon et al., 1994332 ; Valdés etal., 2002391.

Authors: Paradis G. & M. L. Pozzo-di-BorgoCollaborators: Grillas P., H. Michaud, L. Rhazi & N. Yavercovski

IUCN 1997

Bern Convention



-

-

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

17

Plant species

Damasonium polyspermum Coss.

ANGIOSPERMS

ALISMATACEAE

Main synonymsD. stellatum Thuill. var. polyspermum (Coss.) Loret & BarrandonD. stellatum Thuill. var. polyspermum (Coss.) P. Fourn. D. alismaMill. 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 grainesSpanish names: Saeta de agua, Cola de Golondrina, Almea,AzumbarItalian name: Mestolaccia sicilianaMoroccan name: Mizmar er raaïEnglish names: Starfruit, Thrumwort

SubspeciesNone

Description/identification criteria • Small annual plant with the leaves all basal and arranged in arosette.• First submerged leaves linear and not persisting, subsequentlyreplaced by floating leaves with long petioles and narrow lanceo-late blades, attenuated or more or less truncated at the base,with faint transverse venation.• Leaves produced during the non-flooded phase similar to thefloating leaves but with a shorter petiole and a thicker blade.• Inflorescence most often with one, sometimes several super-posed umbelliform whorls. Flowers hermaphrodite with three per-sistent green sepals and three deciduous coloured petals. Petals2 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 withpink 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 withtransverse ridges (dimensions: 0.9-1.2 mm x 0.5-0.7 mm), in eachmature carpel.

Similar species Damasonium alisma Mill. has only two large seeds (1.7-2.5 mmx 0.8-1.2 mm) per carpel when the fruit is ripe. According to someauthors, additional distinguishing characters include the plant’soften larger size and more robust build, leaves rounded to cor-date* at the base with oblique transverse veins visible against thelight, smaller petals (4.2 ± 0.5 mm x 3.7 ± 0.5 mm) lacking pinkcolour at the tips, and shorter anthers (0.75 ± 0.006 mm v. 0.82± 0.12 mm). However, the relevance of these characters needs tobe confirmed, especially in relation to the influence of the envi-ronment (depth of water, length of submersion, etc.).

The two Damasonium are not always separated by botanists. Accor-ding to Rich & Nicholls-Vuille316, they are also separable on thebasis of their largely different geographical ranges and by theirchromosome numbers (D. alisma 2n = 28, tetraploid; D. polysper-mum 2n = 14, diploid*).

Baldellia ranunculoides (L.) Parl. is distinguished, at all stages ofdevelopment, by the scent of coriander that is released when theleaves are crushed. During flowering or fruiting, the arrangementof the many obovate carpels (≈ 2 mm) in a globose head removesany doubt.

Alisma spp.: The three Alisma species included in the French floraare usually far more robust than Damasonium polyspermum. Atflowering or fruiting they are easily distinguished by their manycarpels, which are blunt oval shaped, very tightly packed andwhorled in a single row. Unlike Damasonium, these are all peren-nial* species.


Distribution/range

Western Mediterranean species.

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

Other Mediterranean countriesNorth Africa: in freshwater dayas*, especially in mountains, inCyrenaica, in the High Plateaux of northwestern Algeria and inthe Middle Atlas in Morocco.Italy: southwest Sicily in the coastal zone292.Edges of ponds and pools in the Iberian Peninsula, in central andsouthern Spain as well as Portugal, Greece and Syria.

Habitat

General descriptionLand temporarily flooded by fresh or slightly brackish water inthe meso-Mediterranean zone: pools, edges of ponds and lagoons.

Damasonium polyspermum

1 cm

18



CORINE Biotopes22.32 Euro-Siberian dwarf annual amphibious swards(Cyperetalia fusci).22.34 Mediterraneo-Atlantic amphibious communities (Isoetalia).

22.341 Short Mediterranean amphibious swards (IsoetionBr.-Bl. 1931).

Phytosociology*• Euro-Siberian dwarf annual amphibious swards (Cyperetaliafusci): - Lythrion tribracteati (e.g. the Lythrum tribracteatum-Teucriumaristatum association*215, 252.- Elatino triandrae-Eleocharition ovatae (e.g. Elatinetummacropodae).• Southern amphibious communities (Isoetalia): - Short Mediterranean amphibious swards (Isoetion).• Damasonium polyspermum sometimes overlaps into formationsappearing later in the year and having a more distinctly eutro-phic* or even halophilic* character: Heleochloion schoenoidis.

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

Companion species The species most often associated with Damasonium polysper-mum are: Lythrum tribracteatum at all the French localities, within addition, depending on the site, L. hyssopifolium, Pulicaria vul-garis, P. sicula, Myosurus minimus s.l., Juncus bufonius s.l., J. tenageia and Herniaria glabra.

Ecology


Life formAnnual species (therophyte*).

ReproductionAerial.

Seeds (see above § Description)The lifespan of seeds in the sediment is not known, but observa-tions of the dynamics of natural populations indicate that it islong.

Biological cycleSeed germination and growth of leaves during the flooded phase(end of winter, beginning of spring). Flowering in spring (fromApril to June) up to the first dry periods, but may be prolongedor repeated in autumn if the substrate is sufficiently wet. Ripe-ning of fruits and dispersal of seeds during the dry phase (sum-mer) and before the autumn rains which mark the end of thecycle.


Flowering

Germination and vegetative growth

Seed dispersal

Formation and ripening of fruits

Southern France


HydrologyThe germination of seeds, which is very variable from one year toanother, requires immersion in water (a few centimetres) or water-logged soil in spring, and little vegetation cover (strong insolation).The plant can remain in the vegetative stage under very shallowwater, with characteristic floating leaves, but must emerge quicklyfor flowering and fruiting. The size and density of individuals andthe length of their life cycles exhibit spatial variability within thepool, in relation to hydrological gradients (duration of flooding,dates of flooding and drying out) that are themselves dependenton topographical gradients.

SubstratePlant tolerant of a wide range of soils and of slightly saline con-ditions.

Interspecific competitionA very light-demanding (heliophilous*) species, and thereforevery sensitive to any plant cover at the germination stage.

Impact of perturbationsDisturbance which helps to control competing vegetation is bene-ficial, whereas disturbance resulting in the burial of seedsimpede germination (trampling and digging by livestock or wildanimals, ploughing, overdeepening or dredging of the pool, etc.)(see Vol. 1, box 38).



FranceThe species currently occurs at 10 sites: • Hérault: Vendres/Sauvian (Malhol de l’eau), Agde (Notre-Damede 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), Saint-Antonin-sur-Bayon (Plateau du Cengle).• Var: Gonfaron (Lake Bonne Cougne), Flassans-sur-Issole (LakeRedon), Besse-sur-Issole (Lake Gavoty).• Absent from Corsica.

ItalyOnly 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 thecountry (between 50 and 100 sites), and three provinces in thesouth of the country (about twenty sites).

19

Plant species

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

MoroccoThe species is not considered to be rare133.

NoteAs the two species of Damasonium have not always been sepa-rated by botanists, some data remain uncertain: for example oldpublications list D. alisma sensu lato from the closed depressionsof the Roussillon plain (between Têt and Tech)351, in the coastalpools at Argelès-sur-Mer155 and in the Montmajour marsh(Arles/Fontvieille) (Jacquemin 1848 in Molinier257). In the FrenchMediterranean region, D. alisma is only known with certaintyfrom the extreme south of Larzac30 and appears therefore to beabsent from the meso-Mediterranean zone.

Conservation status

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


Natural factors As the species is an annual, it does not compete well with colo-nial perennial species (Artemisia molinieri, Mentha cervina,Isoetes spp. Eleocharis palustris, etc.), this being partly compensatedfor by its abundant seedbank. In many depressions, the appearanceof Damasonium is irregular from one year to another37.

Risks relating to populations This risk is low in the absence of any major perturbations, due tothe fecundity of the plant and the longevity of the seeds in thesediment.


Current measures The Lanau pool (Bouches-du-Rhône) was acquired by the CEEP in1998.The only management measures known at present are:• Monitoring of the population of the Lanau pool in the Crau, inrelation to grazing management.• A management plan in progress for the pools of the Centre Var,in the context of drawing up the Objectives Documents forNatura 2000* sites.• The ex-situ preservation of seeds by the ConservatoireBotanique National Méditerranéen de Porquerolles.

Recommendations• Promotion or maintenance of grazing, which is favourable tothe growth of the species, as is any other means whereby peren-nial vegetation communities are opened up.• Promotion of the acquisition and management of Damasoniumsites, which are in addition often rich in other rare species.

Bibliography

Bernard, 199730 ; Bigot, 195536, 37 ; Braun-Blanquet, 193548 ; Contiet 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.

IUCN 1997Bern ConventionHabitats DirectiveNational and regional protection


---

- France, national list: Decree of 20/01/1982/Journal Officiel13/05/1982, amended by Decreeof 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


Anthropogenic factors Several populations have disappeared following the infilling ordrainage of the depressions which supported them, in connectionwith agricultural developments (Vendres plateau, pools and pondsof the Costière Nîmoise) or with urban development (Rigaudpools at Agde).

20


Elatine brochonii Clavaud

ANGIOSPERMS

ELATINACEAE

Main synonymsE. hydropiper L. var. pedunculata (Moris) FioriE. hexandra (Lapierre) DC. subsp. brochonii (Clavaud) P. Fourn.

French name: Elatine de Brochon

SubspeciesNone

Description/identification criteria76, 96, 138, 381

• Herbaceous plant, amphibious, short (2-7 cm) with a relativelylong and significant root system.• Leaves opposite, spatulate, more or less petiolate, with asmooth slender stalk, prostrate, much-branched and rooting atthe nodes.• Flowers hermaphrodite, actinomorphic*, small (not more than5 mm), sessile, solitary in the leaf axils, with three free sepalspersisting in fruit scarcely longer than the fruit, and three freepetals, white with pink veins. Androecium of six stamensarranged on two verticils.• Fruit: spherical, slightly flattened capsule* with three valves.Seeds many, small, reticulate, straight or slightly arcuate, withlongitudinal ribs, transversely grooved.• Plant brownish-coloured at the end of the cycle.

Similar species • Elatine hexandra has pedicellate flowers (sessile in E. bro-chonii).• 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.


Distribution/range

A western Mediterranean species.

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

Other Mediterranean countriesSpain76, Morocco303, Algeria292, 386.

Habitat

General description In France, Elatine brochonii is found on the edges of temporarypools in southwest France on siliceous substrates with irregularflooding between years. In Corsica, this species occurs at a sin-gle site (Padulellu pool) in a depression that cuts into a graniticridge (105 m altitude) covered with low matorral*. Flooding anddrying out, which are irregular from one year to another, takeplace through rainwater and evaporation. The maximum waterlevel recorded in 1996 (a very wet year) was 80 cm219.

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


CORINE Biotopes22.34 Southern amphibious communities (Isoetalia).

22.341 Short Mediterranean amphibious swards (Isoetion).

Phytosociology*Elatine brochonii is found together with Juncus pygmaeus, J. bufo-nius, Lythrum hyssopifolia, Illecebrum verticillatum, Exaculumpusillum and Isoetes velata, which means that it is included intwo alliances, the Isoetion velatae and the Cicendion219: Class: Isoeto-Nanojuncetea

Order: IsoetaliaAlliance: Isoetion velatae Br.-Bl. 1931Alliance: Cicendion filiformis Rivas Goday (1961, 1964)Br-Bl. 1967.

Ecology


Life formTherophyte* (annual).

Adaptive strategyStress-tolerant-ruderals (S-R)163.

ReproductionAerial.

Elatine brochonii

0.25 cm

21

Plant species

SeedsSeeds are abundant (33 to 50 seeds per capsule), long-lived* inthe sediment, small (length: 0.63 mm, width: 0.27 mm), andweigh less than 1.4 mg.

Biological cycleAmphibious plant with a very short life cycle. It begins its bio-logical cycle under water or just after emergence and ends it outof water. Its appearance, irregular from one year to another andon varying dates, has been observed in southwest France346.In Corsica germination takes place at the end of spring (May-June), only during wet years. Flowering occurs from May to July,fruiting in June and July, and the seeds are dispersed in July andAugust.In Morocco, Elatine brochonii germinates in February-March and inApril forms flower buds which very quickly develop into flowers.Fruiting takes place from April to June with the dispersal of seeds313.



In mainland France, 13 stations were formerly known in thesouthwest, of which only one appears to have survived274; thereis one site in Corsica (Padulellu pool219), and at least eight inMorocco133, 346.

Conservation status

The interannual variability of the species causes difficulty inevaluating the status of the populations. However, it no longerappears to be present at most of its mainland French sites, andthe Corsican population remains vulnerable. In Morocco the pop-ulations appear to be better preserved, but are subject to rapidlyincreasing human pressures, especially around Rabat. The highsurvival rate of the seed stocks* in the soil gives grounds for hop-ing that populations may still exist in the dormant state.



Anthropogenic factors Digging out and removal of sediments, frequent vehicle move-ments creating tracks through the pool (Mamora pool in Morocco),or the formation of a sandy alluvial fan partly covering the Elatinestation (Padulellu pool in Corsica).

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

Risks relating to populationsDespite its large seedbank, which enables populations to persist,risks of extinction may arise in the future, especially at the sitessouth of the Mediterranean where the species may be consideredto be at the southern limit of its range. The high frequency of dryyears in these countries (Morocco) could result in the long termin a decline in the populations of Elatine brochonii. The low num-ber of sites increases the risk of extinction.


Flowering

Fruiting

Seed dispersal

Germination

Morocco


Flowering

Germination

Seed dispersal

Fruiting

Corsica



---

- France, national list: Decree of20/01/1982/Journal Officiel13/05/1982, amended by theDecree of 31/08/1995/JournalOfficiel 17/10/1995 - France, national Red DataBook274: priority species - Spain, national Red List: “vulnerable”13

- Morocco: “very rare” (RR)133, 181, 381


HydrologySeed germination requires the soil to be flooded (a few centime-tres of water) or saturated during March, and intense insolation(bare soil). The plant may remain in the vegetative stage undervery shallow water, but must emerge quickly for flowering andfruiting314.

SubstrateAcid, oligotrophic* at Moroccan stations, mesotrophic* inCorsica.

Interspecific competitionA very light-demanding species (heliophilous*), Elatine brochoniiis very sensitive to any plant cover at the germination stage.

Impact of perturbationsPerturbations which help to control competing vegetation arefavourable to Elatine brochonii, while those resulting in burial ofthe seeds will impede germination (87% germination rate forseeds on the surface compared with only 13% and 6% for seedsburied in sand to depths of 2 mm and 5 mm respectively). Thisburial may be caused in the field by the passage or digging ofwild and domestic animals, sedimentation (Padulellu) and themovements of vehicles.

22



Current measures None

Recommendations• In France and in Morocco systematic monitoring of the knownpopulations is recommended, as well as an assessment of theenvironment, the condition of the populations (including seed-banks), and the possible opportunities for their restoration. • At the Padulellu pool (Corsica), it is a matter of urgency thatthe process of sedimentation be halted (sand originating fromthe erosion of the track running next to the pool) and, to thisend, that the belt of Myrtle maquis be reinstated; the driving andparking of vehicles in the pool should also be prohibited. • In Morocco, the continuance of grazing appears to be benefi-cial in controlling the amount of competing vegetation.

Bibliography

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, 1931-1934181 ; Lesouëf & Richard, 1995210 ; Lorenzoni & Paradis, 1997219 ;Maire, 1926228, 1932229, 1952-1987230 ; Médail et al., 1996243 ; Olivieret al., 1995274 ; Pignatti, 1982292 ; Quézel & Santa, 1962-1963303 ;Rhazi et al,. 2001313, 314 ; Rivas-Goday, 1970319 ; Schotsman & Bos-serdet, 1966346 ; Schotsman, 1985347 ; Titolet & Rhazi, 1999381 ;Tutin et al., 1964-1980386 ; Vanden Berghen392, 1966 ; Vivant,1960396.

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

23

Plant species

Eryngium pusillum L.

ANGIOSPERMS

APIACEAE

Main synonymsE. barrelieri Boiss.

French name: Panicaut nain de Barrelier Italian name: Calcatreppola di Barrelier

SubspeciesNone

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 arosette of leaves at its base and ending in a capitulum (inflores-cence). The stem may branch and give rise to new units.• The young rosette leaves are elongated and not prickly. As theyage they become stiff, with thin spines along their edges. • There may be very many capitula on a single clump. Each carriesseveral very prickly bracts. The flowers are very small and bluish.They produce tiny achenes (indehiscent dry fruits)220, 274, 292.

Similar species None


Distribution/range

Southern Mediterranean species

FranceOnly in southern Corsica

Other Mediterranean countriesSardinia, Sicily, southern Italy, Tunisia, Algeria, Syria, Morocco274, 292.

Habitat

General description The Musella depression is a poljé* (karstic* depression) whichpasses through very different ecophases* over the course of ayear, whose two extremes are a winter flooded ecophase and avery dry summer ecophase.Flooding, which is due mainly to the rising water table in theunderlying limestone, occurs in winter during wet years and mayin exceptional cases last from November to April (e.g. in 1996and 2001). Drying out begins in March (in April in very wetyears), 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).

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

In spring 22.34 Mediterraneo-Atlantic amphibious communities.

Phytosociology*In summer and early autumn: Mentha pulegium and Eryngiumpusillum meadows, classifiable within the Trifolio-Cynodontionalliance (order Plantaginetalia majoris156 and class Molinio-Arrhenatheretea), which does not appear to correspond to anycategories in CORINE Biotopes.

Ecology


Life formPerennial species, hemicryptophyte*. It is formed of sympodial*units. Rosettes of leaves develop at the base of the units, whichare monocarpic (only flower once). The different units togetherform a low clump220.

GerminationAquatic.

ReproductionAerial for flowering220.

Survival strategy of the clumps Stress-tolerant (S) according to the terminology of Grime163

Biological cycleGermination in November if the site is flooded, dispersal ofyoung plants during the flooded period (hydrochory), vegetativegrowth from November to May, flowering end of May and June,fruiting in June and July, dispersal of achenes for a very short dis-tance from the parent plant and dispersal of the seedlings byfloatation (hydrochory)220. During and especially at the end ofsummer the leaves wilt and disappear, while new leaves appearwith the first rain of September-October.

Eryngium pusillum

1 cm

24



HydrologyThe alternation of a flooded ecophase (from the end of autumnto the beginning of spring) and a dry ecophase is necessary. Inthe absence of winter flooding the seeds do not germinate inspring. An absence of germination for several years is, however,not a handicap for this species, as the large amount of germina-tion during wet years, plus vegetative* reproduction, enable thepopulation to rebuild itself.

SubstrateMesotrophic*.

Interspecific competitionA strictly heliophilous* species, negatively affected by the shadefrom taller vegetation.

Impact of perturbationsA prickly species once the site dries out, not (or very little)grazed, but highly stimulated by grazing cattle, which restrictcompeting species and create bare areas which are favourablefor germination.



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

Conservation status

In Corsica, the Musella population is increasing, slowly but surely220,but it has disappeared from the two Corsican stations at Vix andVico274.



Anthropogenic factors At Musella, changes in the substrate have been observed as aresult of ploughing and sowing. In addition, the cessation of cat-tle grazing by the site manager in 2002 and its replacement withgrazing by a flock of sheep poses the risk, for the future, of alter-ing the ecological conditions at the site.At Vix, development works destroyed the population and itshabitat274.

Natural factorsAt Musella, the Eryngium pusillum population should not experi-ence any problems in the short term. However, the developmentof woody plants (Blackthorn, Ulmus minor and Rubus ulmifoliuscould be detrimental in the medium term (shading).


Current measures • Satisfactory at the Musella site up to 2001 as a result of theregular extensive cattle grazing. Moreover, the Musella pool ispart of a Natura 2000* site.• Seed collection and ex-situ storage by the ConservatoireBotanique National Méditerranéen de Porquerolles, and precau-tionary cultivation of the species on Conservatoire land since1993.

RecommendationsAs shading has a detrimental effect on the species, surveillanceand control of the spread of woody plants is advisable (scrubclearing, maintenance of grazing). Accordingly, keeping a flock ofsheep at Musella for part of the year is strongly recommended.

Bibliography

Anonymous, 199912 ; Bissardon & Guibal, 199739 ; Fennane & IbnTattou, 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.

Author: Paradis G.


Flowering

Vegetative growthVegetative growth

Fruiting and dispersal of achenes

Germinationand dispersalof seedlings

��



---

- France, national list: Decreeof 20/01/1982/Journal Officiel13/05/1982, amended by theDecree of 31/08/1995/JournalOfficiel 17/10/1995- France, national Red DataBook : priority species274

- Morocco: “very rare” (RR)133, 181

25

Plant species

Illecebrum verticillatum L.

ANGIOSPERMS

CARYOPHYLLACEAE

Main synonymsCorrigiola verticillata (L.) Kuntze

French name: Illécèbre verticilléItalian name: Corriggiola verticillataPortuguese name: AranhoesMoroccan name: Souifa diel el maEnglish name: Coral-necklace

SubspeciesNone

Description/identification criteria • Annual herb with thread-like stem, often radicant, quadran-gular, glabrous, pinkish. • Leaves opposite (can appear whorled due to the developmentof 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 ster-ile stamens. Pistil with small ovoid ovary, with very short styleand two stigmas.• Fruit: achene with membranous pericarp, retained within thepersistent calyx. • Seed sub-elliptical, shining brown, about 1 mm long.

The species exists in two very different forms, one terrestrial andone aquatic63:• The terrestrial form develops in spring when the pool has driedout. The stems are short (about 5 to 20 cm) and prostrate, withvery short internodes. The flowers are very abundant.• The aquatic form develops in winter and spring. The stems areupright and unranked and may be up to 60 or 70 cm long, theinternodes are very long and the submerged leaves are very fine.In March and April the upright stems develop floating branchesat the surface of the water which bear a few inflorescences andresemble the terrestrial form, i.e. short internodes and more obo-vate leaves. The aquatic form produces very few flowers com-pared with the terrestrial 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 Corrigiolalittoralis, but this species has long basal leaves (up to 6 cm) inrosettes.


Distribution/range63, 174, 292, 348

This species is “widely distributed in Europe from southwestSpain to Sweden and Poland, in a band extending 200 to 400 kminland from the Atlantic seaboard”348. It is rarer in theMediterranean region.

FranceWestern France and Corsica.

Other Mediterranean countriesWestern 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 andMorocco).

Non-Mediterranean regionAzores, 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 (aquaticform).

I. verticillatum is usually associated here with Apium crassipes,Ranunculus peltatus, R. ophioglossifolius, Myriophyllum alterni-florum, Baldellia ranunculoides, Eryngium barrelieri, Isoetesvelata, etc.72, 219, 221, 298, 314

I. verticillatum also occurs in its terrestrial form at the edges ofpools, in small transitory rain pools within Mediterraneanmaquis, and in cupular pools* (for example in Corsica at Evisa).Here it forms communities in association with Anagallis parvi-flora, Cicendia filiformis, Exaculum pusillum, Kickxia cirrhosa,Radiola linoides, Silene laeta, Solenopsis laurentia etc.54

Illecebrum verticillatum

1 cm

26



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 ofvariable depth.

Phytosociology*In its aquatic form Illecebrum verticillatum is a member of thesubmerged phase of the Isoetion formations within the Isoetesvelata communities. Some authors place the hydrophyte* forma-tions to which it belongs in the communities of the Potamoge-tonetea pectinati class221, 284.

The communities and associations* which include I. verticillatumand which are situated in places undergoing brief periods offlooding have received more attention from phytosociologists*.Thus, according Brullo & Minissale54, the phytosociological affini-ties of such communities and associations are as follows: Class: Isoeto-Nanojuncetea

Order: Isoetalia- Alliance Cicendio-Solenopsion laurentiaeVarious associations, in which I. verticillatum is aspecies which ranks highly in terms of occurrence - Alliance Agrostion pourretii

Association Illecebro-Agrostietum pourretii

2. Other habitats

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

Ecology


Life formTherophyte (annual).

ReproductionAerial.

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

Adaptive strategy (sensu Grime163)Stress-tolerant-ruderal (SR).

Description of seedsLength approx. 1 mm.

Biological cycleIn the deep parts of temporary pools: • Germination during winter (from December).

• 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 ofautumn-winter.


Germination

Corsica


Morocco

Germination

Vegetative growth

Flowering

Fruiting

GerminationGermination

Vegetative growth

�

� �

�

Flowering

Fruiting


HydrologyWinter flooding or saturation of the sediment, even for a briefperiod, appears to be necessary for the germination of Illecebrumseeds. This germination, which takes place rapidly as soon as thefirst rain falls, enables the species to appear in the field almostevery year314.

SubstrateLoose, very thin substrate (less than 3 cm deep) acid, siliceous,sandy or gritty and oligotrophic*.

Interspecific competitionThis strictly heliophilous* species does not tolerate cover fromother species, hence its confinement to substrates that are verythin and poor in absorbable mineral salts. In very turbid pools(Mamora, Morocco) it only grows in abundance after the waterhas fallen.

Impact of perturbationsDisturbance resulting in the opening up of the vegetation isclearly favourable (e.g. trampling).



In CorsicaThe species probably occurs at more than 20 sites.

In MoroccoThere are over thirty stations distributed between the Rif, north-ern Atlantic Morocco, and mid-Atlantic Morocco. These sites arelocated on siliceous soils of the plains and low mountains sub-ject to semi-arid, sub-humid and humid Mediterranean biocli-mates*134.

27

Plant species

In SpainThe species is present at 76 sites scattered through some twentyprovinces177.

Conservation status

In Corsica, Spain, Portugal and Morocco the species is notthreatened, appearing to be stable despite more or less wideinterannual fluctuations depending on the site.



Anthropogenic factors

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

In Morocco Urban development and the filling-in of pools are the mostimportant threats. Thus some sites in the Mamora forest havedisappeared following the construction of the Rabat-Fez motor-way. Some other sites close to large towns (Casablanca, Rabat,Benslimane, Bouznika, etc.) are also threatened by urban devel-opment in the medium term.

Natural factors

In CorsicaThere is a risk that the likely discontinuation of extensive cattlegrazing in the near future will result in a severe decline at sev-eral sites due to the spread of tall herbaceous plants (Schoenus

nigricans, Oenanthe globulosa etc.) and woody species (Phillyreaangustifolia, Erica scoparia, Myrtus communis, Pistacia lentiscusetc.). This is true of the temporary pools of southern Corsica, wherethe Illecebrum verticillatum population sizes are very small (Muradell’Unda, Arasu, Muratellu pools), probably due to the cessationof cattle grazing several years ago which has led to the establish-ment of a heavy biomass of herbaceous plants.

In Morocco Practically all the sites are grazed, allowing the vegetation to bekept open and preventing any possible spread by the surroundingwoody plants.


Current measures At present, in Corsica as well as in Morocco, there are no man-agement measures, due to the lack of control over land owner-ship at the sites.

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

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 BorgoCollaborators: Grillas P., L. Rhazi & N. Yavercovski


-----

28


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

LYCOPODIOPHYTES300

ISOETACEAE

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.

French names: (1) Isoète de Durieu, (2) Isoète épineuxItalian names: (1) Calamaria di Durieu, (2) Calamaria istriceEnglish name: (2) Land Quillwort

SubspeciesNone


• Terrestrial perennial plants with a very short stem forming akind of underground bulb bearing roots, and many fronds (“leaves”)arranged in rosettes at ground level. The fronds of Isoetes durieiare decurved towards the ground, while those of I. histrix arenarrow and tough.• Bulb: usually firmly embedded, ringed with blackish spiny scalescorresponding to the sclerified* remains of the bases of the pre-ceding 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 pitcontaining 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) andmicrosporangia containing microspores (mâles).

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


Distribution/range

(1) Isoetes durieiWestern Mediterranean species (distribution map in Quézel306).

FranceOn the mainland, it is common at Les Maures and Estérel, fairlyrare elsewhere (Biot Massif, between Béziers and Montpellier,southern catchment of the Caroux, southern foothills of the Mon-tagne Noire, Albères, the Roussillon plain, valleys of the Gardon

rivers in the southern Cévennes); in Corsica, it is fairly commonin the coastal zone and at low altitudes up to 300 m (but hasbeen found exceptionally at 1,000 m).

Other Mediterranean countriesPortugal, 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.

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

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

Isoetes duriei

1 cm

megaspore

frond base

0.25 cm

0.35 mm

Isoetes histrix

1 cm

megaspore

frond base

0.35 mm

0.25 cm

29

Plant species

Other Mediterranean countriesCentral and western Iberian Peninsula (Spain, and all the Por-tuguese provinces), mainland of Italy, Sardinia, Sicily, Macedonia,Greece, Crete, Malta, Croatia, Morocco, Algeria, Tunisia, MiddleEast (Syria, Lebanon) and Turkey.

Habitat

General description The most suitable habitats are short Mediterranean swards, wetor waterlogged in winter and the beginning of spring and verydry in summer. The substrate is usually siliceous (silts and sands)and relatively rich in organic matter. At Les Maures and theEstérel, Isoetes duriei also grows equally well in soils with littleorganic matter where, however, it grows to a smaller size than inaccumulation 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 wetthan those defined for I. duriei, and form a stage transitional tothe annual swards of drier biotopes (with Tuberaria guttata andAnthoxanthum ovatum).

Companion species (1) I. duriei• In Corsica: Ophioglossum lusitanicum, various Serapias, Airacapillaris, Bellis annua, Cicendia filiformis, Radiola linoides, Linumbienne, Anagallis arvensis subsp. parviflora, Lythrum hyssopifolia,Juncus bufonius, J. tenageja, J. capitatus, Ranunculus revelieri andLotus conimbricensis.• In mainland France21, 22, 215: the same species, with in additionIsolepis cernua (=Scirpus savii), Nasturtium asperum, Agrostispourretii, Airopsis globosa, Veronica acinifolia, Juncus pygmaeus,J. fasciculatus and Mentha pulegium.

(2) I. histrix• Species common to the various sites: Radiola linoides, Linumbienne, Anagallis arvensis subsp. parviflora, Serapias lingua, Juncusbufonius, Isolepis cernua (= Scirpus savii), Bellis annua.• At the former sites of the littoral Maures, where the specieshas not recently been seen: Allium chamaemoly, Romulea colum-nae and Isoetes duriei22.• In Morocco: Poa annua, Filago (= Logfia) gallica, Rumex buce-phalophorus 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 vegetationoften invades the short swards where Isoetes histrix occurs, andit 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: formationswith Isoetes histrix, I. duriei, of ephemeral aquatic habi-tats.

Phytosociology*Owing to the difference in ecology between Isoetes velata (ofperiodically flooded habitats) on the one hand, and I. duriei andI. histrix (of non-flooded habitats) on the other, de Foucault98

split the Isoetion of Braun-Blanquet48 and Rivas-Goday306, 319 intotwo alliances:The Antinorio agrostideae-Isoetion velatae for topographicallylow-lying areas, periodically flooded.The Ophioglosso lusitanici-Isoetion histricis for topographicallyhigher areas, not flooded.The Quillwort communities of non-flooded habitats are thereforeincluded in the Ophioglosso lusitanici-Isoetion histricis alliance,order Isoetalia velatae and class Isoetea velatae98, 194.

In the Isoetion s.l. (Isoetalia), the habitat of these two speciescorresponds to the following communities:

(1) I. duriei• Isoetetum duriei association*, defined in Languedoc48 and alsodescribed from Spain319.• Association with I. duriei and Juncus capitatus described fromCorsica233.• Association with I. duriei and Nasturtium asperum described atLes Maures21.

(2) I. histrix• In France, in the Serapion alliance, Aubert & Loisel 1971: sub-association with I. histrix of the Serapio-Oenanthetum, describedby 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. histrixand Radiola linoides, described by Chevassut & Quézel72.

Ecology


Life formPerennial species: bulb-forming geophyte*292.

Adaptive strategy163

Stress tolerant-ruderal (S-R).

ReproductionAerial 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 autumnrains21 and in winter when the substrate is very wet; ripening ofspores in spring, followed by rapid withering of the fronds at theend of spring, when the habitat is drying out rapidly.

30


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



(1) I. duriei• France (mainland and Corsica): large number of sites (seeabove, § Distribution), but number of stations within the sitesimpossible to estimate accurately, given the very scattered natureof 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 coastof the peninsula)137, 292.

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

Conservation status

(1) I. durieiThe populations appear to be stable.

(2) I. histrixThe populations appear to be stable in Corsica and Morocco asthey are fairly resistant to anthropogenic and natural threats.They have declined in mainland France, in Roussillon (extinctiondue to the destruction of the Saint-Estève pool) as well as at LesMaures (not found recently in the Saint-Tropez region, or at thecapes of Lardier, Benat, and Taillat, but recently rediscovered atHyères, at La Londe-les-Maures and Le Cannet-des-Maures). InMalta, the species has not been seen again since its discovery atthe end of the 19th century344.

Legal status/level of protection


France

Maturation of spores

Withering of fronds

and vegetative growth Germination

��


France


Withering of fronds

vegetative growth


Morocco


Withering of fronds


��Germination and

��



---

France (1) and (2), national list:Decree of 20/01/1982/JournalOfficiel 13/05/1982, amended byDecree of 31/08/1995/JournalOfficiel 17/10/1995 - Balearics (2): “endangered”(EN)331

- Malta (2): “presumed extinct”(Ex ?)344

Isoetes duriei

Isoetes histrix


HydrologyA very wet substrate is necessary at the beginning of the cyclebut these plants do not like to be submerged for long.

Substrate(1) not (or very slightly) oligotrophic* in Corsica, oligotrophic on themainland (Permian sandstones at the Plaine des Maures, rhyo-lites at the Colle du Rouet).(2) more or less oligotrophic (usually sand-silt).

Interspecific competitionLow growing species, light-demanding (heliophilous*) and conse-quently not very tolerant of competition from taller plants.

Impact of perturbationsThe main perturbations are the “ploughing” of the substrate bywild boars, and trampling (by cattle and people). They are bene-ficial as they limit the amount of cover and litter produced by tallerplants.

TemperatureIt appears to govern the distribution of Isoetes histrix at LesMaures, where it is confined to very warm coastal stations. It isusually replaced by I. duriei away from the coast, in the Mauresplain and massif in particular22.

31

Plant species



FranceIn Corsica, development projects (such as construction works) area potential threat to some coastal sites.In Provence, in Les Maures, urbanisation and tourist develop-ments (golf courses) have destroyed a large number of Isoetesduriei stations*. Similarly, development along the coastline hasdestroyed all the coastal I. histrix stations (region of Saint-Tropez,Le Lavandou, Les Bormettes), and constitutes a threat to the otherstations.

MoroccoAlong with urban development, the greatest threat to the I. histrixpopulations is the infilling of pools with debris.

Natural factors• In the absence of disturbance, the closing-up of the habitat dueto the spread of taller plants (Schoenus nigricans, Scirpus holo-schoenus, Dittrichia viscosa and especially various maquis species)could constitute a threat for several sites in Corsica. This is equallytrue at Les Maures where the cessation of grazing is favouringthe growth of perennial herbaceous species (Paspalum dilatatum,Juncus conglomeratus, Scirpus holoschoenus), and even maquisspecies. This is not the case at the Moroccan sites, where grazing iscommonly practised.• The frequency of dry years does not affect the Isoetes histrixpopulations, which sustain themselves by their spores as much asby their bulbiform rhizome.

Risks relating to populations(1) I. durieiNo extinctions currently known in Corsica or on the mainland.(2) I. histrixNo extinctions currently known in Corsica. On the mainland, theinherent risks to populations are low, but possible, due to thelimited numbers and the very marked isolation of each of thepopulations, which are not very vigorous.

Management measures

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

At Les Maures:• The clearing of firebreaks (provided that it is sensitive, i.e. thatit does not upset the microtopography), and especially theirgrazing, are beneficial.• A Natura 2000* area, in the process of being established in thePlaine des Maures, provides the opportunity to take the require-ments of management for the conservation of I. duriei intoaccount, as does a Réserve Naturelle project that includes themajority of the stations.• The LIFE “Temporary Pools” project has enabled experimentaltesting of the management of plant communities with I. duriei tobe carried out, as well as the purchase by the CEEP of land wherethis species grows.

Recommendations

FranceIn Corsica, retention of extensive cattle grazing at the siteswhere it is still present, and its reintroduction at sites where it isno longer practised.In mainland France:• Retention of horse grazing (one Isoetes histrix station) andpedestrian traffic (two I. histrix stations); avoidance of destruc-tive activities (mountain biking,* scrambling, fly-tipping).• Reintroduction of extensive grazing, beneficial for the conser-vation of I. duriei, at Les Maures.

MoroccoAvoidance of repeated cultivation of the catchment areas near-est to I. histrix sites, as the colonisation of the habitat by arableweeds may have a negative effect (competition).

Bibliography

Anonymous, 199912 ; Barbero, 196521, 196722 ; Bissardon &Guibal, 199739 ; Braun-Blanquet, 193548 ; Castroviejo, 1986-200164 ; Chevassut & Quézel, 195672 ; Cirujano et al., 199279 ;Danton & Baffray 199596 ; Foggi & Grigioni, 1999137 ; Foucaultde, 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 BorgoCollaborators: Catard A., H. Michaud, L. Rhazi, D. Titolet &N. Yavercovski

32


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

LYCOPODIOPHYTES300

ISOETACEAE

Main synonyms(1) I. delilei Rothm, (2) I. variabilis Le Grand

French names: (1) Isoète sétacé, Isoète grêle,(2) Isoète à voile, Isoète voilé Italian names: (1) Calamaria setacea, (2) Calamaria velataMoroccan name: (2) Lehyet-el-rebbEnglish name: (1) and (2) Quillwort (genus name)

Subspecies: (2) Isoetes velata A. Braun subsp. velataI. velata A. Braun subsp. intermedia (Trabut) Maire & WeillerI. velata A. Braun subsp. tegulensis Batt. & TrabutI. velata A. Braun subsp. perralderianaI. velata A. Braun subsp. tenuissima (Boreau) O. Bolos & Vigo


• Perennial amphibious plants: aquatic at least in winter andearly spring, then terrestrial.• Habit is an upright tuft when submerged. Stem very short,forming a kind of bulb which bears roots and many slender, frag-ile 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 fer-tile (sporophylls*): each has a depression at its base on the inte-rior (upper) side, occupied by a large sporangium. The sporangiaof I. setacea are bare, those of I. velata are partly or completelycovered by a well-developed veil.• Outer fronds bear macrosporangia (bare or covered with a veilaccording to species), containing macrospores. The inner frondsbear microsporangia (bare or covered depending on the species),containing microspores, which are very abundant.• Main difference between the two species: a very obvious veilover the sporangia of I. velata, while this veil is absent in I. setacea.

Similar species • The more terrestrial Quillworts (Isoetes histrix and I. duriei)during the dry phase have their bulbs surrounded by old, persis-tent sclerified* frond bases; these scales are absent in I. setaceaand I. velata whose bulbs, at least in winter and spring, have awhitish appearance299.• The subspecies I. velata subsp tenuissima is endemic to centralFrance, so there is no risk of confusion with subspecies velata.


Distribution/range

(1) Isoetes setaceaWestern Mediterranean species (distribution map in Quézel306).

FranceExtremely rare plant, occurring in: • The Hérault: temporary pools in the Plateau de Roque-Hauteand the Plaine de Béziers (Grand-Bois).• The Pyrénées-Orientales: Torremila pool and the Plateau deRodès (Fenouillèdes).

Note: Quézel’s map306 wrongly includes Corsica for I. setacea.

Other Mediterranean countriesSpain (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).

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

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

Isoetes setacea

Isoetes velata

megaspore

frond base

frond base

megaspore

1 cm

1 cm

0.25 mm

0.2 cm

0.25 mm

0.2 cm

33

Plant species

Other Mediterranean countries• Spain (in more than half of the country, mainly in the west andon the island of Minorca), Portugal (in almost the whole of thecountry), mainland Italy, Sicily, Sardinia (subspecies tegulensis)86

and Greece.• Maghreb (Morocco, Algeria, Tunisia, Libya). The subspeciesintermedia and tegulensis occur in Tunisia, Algeria and Morocco,subspecies perralderiana in Algeria. The subspecies velata occursin all the countries of the species’ Mediterranean range.

Habitat

General description

(1) I. setaceaAt Roque-Haute, I. setacea lives in pools hollowed out in theQuaternary basalt.

(2) I. velataHabitats 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 veryfine sediment with a very low content of organic matter (sand,silt and clay). Most of the sites are on siliceous rock, but the tem-porary pool of Padulu (Bonifacio, Corsica), has a calcareous sub-soil. The floodwater is oligotrophic* and its depth varies betweensites and between years: approximately 10 to 60 cm.

Companion species

(1) I. setaceaBraun-Blanquet48 described an Isoetetum setacei for these pools,with the following companion species: Juncus pygmaeus, Myosotissicula, Lythrum borysthenicum, Cicendia pusilla, Lotus angustis-simus, Lythrum thymifolium and Marsilea strigosa.

(2) I. velataSpecies associated with Isoetes velata are varied, according tothe depth of water (zonation) the time of year (phenophase) andalso the substrate21, 22, 219, 221, 284, 302: • Aquatic plants: Tolypella glomerata (Charophytes), Callitrichetruncata, Ranunculus peltatus s.l., R. ophiogossifolius, Myrio-phyllum alterniflorum, Illecebrum verticillatum, Pilularia minuta,Apium crassipes, Alopecurus bulbosus, Baldellia ranunculoides,Littorella uniflora, Glyceria fluitans, etc.• Other species, more or less associated with flooding: Crassulavaillantii, Lythrum borysthenicum, Juncus pygmaeus, Myosotissicula, Laurentia michelii, Ranunculus sardous, Scirpus setaceus,Agrostis pourretii, Bellis annua, Elatine brochonii, Corrigiola lit-toralis, Hypericum tomentosum, etc.



22.341 Short Mediterranean amphibious swards (Isoetion).22.3412 Aquatic Mediterranean Quillwort swards: com-munities including Isoetes setacea or I. velata of waterbodies of variable depth.

Phytosociology*

(1) I. setacea• For France, Braun-Blanquet48 described the Isoetetum setacei(see above). • For Spain, Rivas Goday319 considered I. setacea to be a charac-teristic species of the Cicendion alliance (order Isoetalia).

(2) I. velataVarious communities and associations* with I. velata have beendistinguished 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.

NoteOwing to the wide ecological differences between, on the onehand, I. setacea and I. velata living in periodically flooded habi-tats, and on the other hand I. duriei and I. histrix living in non-flooded habitats, de Foucault98 split the Isoetion ofBraun-Blanquet48, Malcuit (for I. velata)233 and Rivas-Goday319

into two alliances306, 308: • The Antinorio agrostideae-Isoetion velatae for low topograph-ical levels, periodically flooded.• The Ophioglosso lusitanici-Isoetion histricis for higher topo-graphical levels, not flooded.The I. setacea and I. velata communities are thus included in theAntinorio agrostideae-Isoetion velatae alliance, order Isoetaliavelatae, class Isoetea velatae.Julve194 accepted this classification.

Ecology


Life form• Perennial species. However, annual populations of Isoetesvelata have been identified in the cupular pools of the Colle duRouet Massif (Var); they have a very low level of reproductivesuccess295.• (1) hydrophyte*292 and geophyte*48.• (2) hydrophyte292, geophyte48 and sometimes therophyte*295

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

ReproductionMaturation of spores: (1) aquatic (2) aerial.

Description of sporesThe megaspores of the two species are fairly similar. Those of I. setacea (0.4 to 0.9 mm) have a few inconspicuous tuberclesand those of I. velata (0.4 to 0.5 mm) have more or less numer-ous tubercles on their various sides. The microspores are echinatewhen mature.

34


Biological cycle Growth of fronds in the water during winter (from the firstautumn rain on the mainland for Isoetes velata); maturation ofspores in spring (1), or after the water has receded (2); rapiddeath of fronds at the beginning of summer, when the habitat isno longer flooded and dries out severely.

Isoetes setacea

(2) I. velata

France• In Corsica, about twenty sites.• On the mainland, a single station in the Roussillon, with theremaining populations confined to the Var: - In the Plaine des Maures, three small recently discovered sta-tions.- In the Plaine de Palayson, the Catchéou pool and a few tem-porary streams.- In the Estérel, about fifty small cupular pools* on the rhyoliteescarpments of the Colle du Rouet (distributed between aboutfifteen stations), and the Barres de Roussiveau (a few stations).

MoroccoMore than 50 stations for Isoetes velata.

Spain and PortugalThe species is scattered throughout the Iberian Peninsula (apartfrom the northern and western provinces), and on the island ofMinorca64.

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

Conservation status

(1) In mainland France a more or less slow decline of the popu-lations of Isoetes setacea and of the Isoetion plant associationsis taking place. In particular, the populations of the pools atSaint-Estève (Pyrénées-Orientales), Grammont (Hérault), andRedessan (Gard) have disappeared.

(2) In Corsica as well as in Morocco, at sites with loose, thin, evenovergrazed substrates, the populations of I. velata are either sta-ble or expanding. On the French mainland the populations in theRoussillon (Saint-Estève) and the Gard (Redessan, Pazac) havedecreased, while they appear to be stable at Les Maures andEstérel.




Withering of fronds

and vegetative growth Germination

��



Withering of fronds

and vegetative growthGermination

��


National Red Data Booksand Red Lists

---

- France (1) and (2), national list:Decree of 20/01/1982/JournalOfficiel 13/05/1982, amended byDecree 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

Isoetes velata


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

SubstrateUsually non-calcareous, oligotrophic and thin.

Interspecific competitionLight-demanding species (heliophilous) and therefore intolerantof competition.

Impact of perturbationsFavourable, by opening up of the habitat.



(1) I. setacea

FranceI. setacea occurs in about 100 pools in the Roque-Haute NatureReserve, in one pool in the Béziers plain, one at Torremila, sev-eral pools and ditches at Rodès.

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

35

Plant species



(1) I. setaceaAt Roque-Haute, the discontinuation of sheep grazing, accumu-lation of organic matter and significant shading are having aneffect on the growth of I. setacea (reduction in weight of bulbsand 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).

(2) I. velata• In France: the spread of agriculture has destroyed some I. velata stations in the Costière Nîmoise, as well as the principalstation in the Pyrénées-Orientales (Saint-Estève pool). In LesMaures and Estérel, urbanisation and developments affecting thenature of the soil (plantations, golf, improvement of grazing inthe maquis by overseeding and fertilisation) have caused areduction in the number of pools capable of supporting thisspecies; the Saint-Raphaël and Fréjus stations, among others,have disappeared. Some small artificial hill reservoirs are appar-ently becoming suitable for its colonisation.

• In Morocco, the problems are mainly associated with infilling,extraction of sediment and drainage of pools.

Natural factorsIn mainland France and Corsica (but not in Morocco, where graz-ing still takes place) the closing up of the habitat and the accu-mulation of litter, resulting from the increase of tall plant species(Schoenus nigricans, Scirpus holoschoenus, Dittrichia viscosa andvarious maquis species), is deleterious to these species, as isinfilling by eroded sediments (Plaine des Maures and Colle duRouet).

Risks relating to populations(1) No short-term risk of extinction.(2) In Corsica, several small stations are threatened with disap-pearance as a result of the closing up of the habitat (pools ofMuratellu, Arasu and Mura dell’Unda). The recently rediscoveredRoussillon population is very vulnerable due to its small size.


Current measures (1) I. setaceaThe Roque-Haute site has legal “Réserve Naturelle” (NatureReserve) status, which is not being implemented at present dueto the impossibility of gaining access to the station (no controlover usage of the site). The I. setacea sites have been the subjectof a survey addressing water levels, the dynamics of woody veg-etation and clonal* species315.

(2) I. velataIn France and Morocco, the opening up of the tallest vegetationthrough grazing by cattle (Tre Padule Nature Reserve, Capandulapools in Corsica, Plaine de Palayson on the mainland) and bysheep (Padulu site in Corsica) is beneficial.In France, in the Var, further measures are in progress:

• Diagnosis of management problems and implementation of aprotocol for monitoring the cupular pools at the Colle du Rouetas part of the LIFE “Temporary Pools” project. • Classification in the “ecological series” by the ONF* of theCatchéou pools (Plaine de Palayson), as a prelude to enabling thenatural-heritage issues associated with these pools to be inte-grated into forestry management.• Establishment of Natura 2000* areas at the Var sites, provid-ing the opportunity to include the protection of Quillwort in theconservation objectives for these sites.

Recommendations(1) I. setacea• Ensuring that the pools are not kept permanently flooded.• Reintroduction of grazing in the pools with helophytes*, and inthe pools with woody plants (after clearing). • Removal of litter from the pools to allow the spore stocks togerminate satisfactorily.• Evaluation of the dynamics of the vegetation following theintroduction of grazing (and any modifications of management).

(2) I. velata• In France:- In Corsica and on the mainland, maintenance of cattle (andsheep) grazing at the sites where it still takes place and its rein-troduction at sites where it is no longer practised, while avoidingintensive practices (overseeding, fertilisation), and followingclearing operations at sites which have been heavily colonised bywoody vegetation (Corsica).- At the Colle du Rouet, implementation of monitoring at thecupular pools in the context of contractual management whichis being put in place by the ONF, the commune and the CEEP, and,in the case of pools which are silting up, planning for minorintervention (removal of litter). • In Morocco:Discouraging any modification of the hydrology of the pools(drainage, infilling) as well as extraction of sediment, whichdestroys the seedbank.

Bibliography

Anonymous, 199912 ; Barbero, 196521, 196722 ; Bissardon &Guibal, 199739 ; Braun-Blanquet, 193548 ; Castroviejo, 1986-200164 ; 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 BorgoCollaborators: Catard A., H. Michaud, L. Rhazi, M. Rhazi &N. Yavercovski

36


Littorella uniflora (L.) Ascherson

ANGIOSPERMS

PLANTAGINACEAE

Main synonymsL. lacustris L.

French name: Littorelle à une fleurItalian name: LittorellaEnglish name: One-Flowered Shoreweed

SubspeciesNone


Perennial herbaceous plant, hydrophytic* or amphibious, withtwo morphological types: a sterile submerged (aquatic) form anda 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 finelypointed at the tip, with air vessels, and measuring 3 to 15 cm longand 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 leavesthat are reduced to scales. At their tips, the stolons form newrosettes of leaves and take root in the soil by forming adventi-tious roots. Hence the stolons enable the plant to propagate itselfvegetatively to a considerable degree under the water.

Terrestrial formThis results form the transformation of the aquatic form after thewater has dropped. It takes the form of dense rosettes of leavesbut does not produce stolons. It produces flowers (the species ismonoecious). 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 withmany stomata321. The upper surface, especially of the internalleaves, has a longitudinal groove, while the lower surface issemi-cylindrical. The male flowers are borne at the tips of fairly long peduncleswith the female flowers at their bases. The male flowers are small(sepals 4 mm long and stamens with filaments 10-20 mm), thescarcely visible female flowers are protected by foliar sheathsand surrounded by long woolly hairs. The flowering of each plantis usually rapid and concentrated over one to two weeks321.Flowering follows the emergence of the substrate from thewater.Fruit dry, indehiscent, with one seed (achene), 2 mm in length.

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


Distribution/range63, 292

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

FranceA species of the Atlantic seaboard, but also occurs in Corsica. Onelocality, not recently confirmed, in the Crau (Etang du Luquier,Bouches-du-Rhône).

Other Mediterranean countriesMorocco in the Rif region, Sardinia (where it is extremely rareand reaches the southern limit of its European range), northernItalian provinces (in a non-Mediterranean climate), Spain andPortugal.

Habitat

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

Littorella uniflora

1 cm

37

Plant species

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

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


22.341 Short Mediterranean amphibious swards (Isoetion).22.3412 Aquatic Mediterranean Quillwort swards: com-munities containing Isoetes velata in water bodies of variabledepth (where I. velata has a much earlier phenology thanLittorella uniflora).

Phytosociology*Littorella uniflora is a companion species in the Isoetes velata com-munities (Corsica). Elsewhere it is characteristic of formations ofthe class Littorelletea in Spain79 and in northwest Europe whereit is also a component of submerged plant associations* of theclass Potamogetonetea pectinati47, 82,127.

Ecology


Life formPerennial species: hemicryptophytic* hydrophyte*.

ReproductionAerial.

FloweringIn Corsica from mid-May to the end of June, and in April-May inMorocco.

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

Biological cycleAt the beginning of spring, under water 10 to 30 cm deep:• The achenes germinate and give rise to new individuals whichsend out stolons that produce rosettes (submerged form).• The rhizomes break dormancy and produce short stolons giv-ing rise to new rosettes (submerged form). At the end of spring, when the pool has dried out or in very shal-low water (less than 10 cm):• Transformation of rosettes from the submerged form to theterrestrial form (beginning of May).• Flowering of plants that have emerged or are under very shal-low water (end of May and June). In summer: • Fruiting (June and beginning of July). • Transition of achenes and short rhizomes into the dormantstate.


HydrologyWinter flooding is essential for germination and for the activa-tion of the buds on the short rhizomes to produce stolons. Emer-gence from the water is essential for flowering117, 127.

SubstrateSubstrate acid, loose, sometimes pisolithic, fairly thin (less than10 cm deep), of variable particle size (silt, sand, gravel), without(or with little) organic debris and oligo-mesotrophic.

Interspecific competitionA heliophilous* plant, intolerant of cover from tall species (Eleo-charis palustris, Phragmites australis, various Scirpus).

Impact of perturbationsLow-intensity disturbance (grazing, trampling) that causes asmall amount of substrate compaction is beneficial127.Drainage, by altering the hydrology of the pools, may have an effecton the germination of the achenes and sexual reproductive suc-cess.

LightLight-demanding (heliophilous plant), which could explain itsabsence from turbid pools in Morocco.



• France:- In Corsica, only 6 sites (see above).- In non-Mediterranean France: the species has a scattered dis-tribution in the western, central, eastern and extreme northernregions 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 throughthe northwestern half of the Iberian Peninsula (Medina, pers.com.). • Morocco: the species has been recorded only in the sub-humidto humid mountainous region of the Rif (Issaguene dayet).


Morocco


Transformation aquatic� terrestrial form

Germination and development of stolons

Corsica

Fruiting

Flowering


Fruiting

Flowering

38


Conservation status

• In Mediterranean France: in Corsica the populations fluctuatefrom one year to another, but appear to be stable in the mediumterm at the Tre Padule pools. There is a lack of data from theFrasselli pools. The four plants recorded by Lorenzoni225 at theMura dell’Unda pool (north of Porto-Vecchio) have not beenfound again (looked for in 2001, 2002 and 2003). Gamisans145

reported its occurrence at the Lac de Ninu (1,800 m altitude), butthe species could not be found there during a recent study of thelake. In the Crau, where the species was collected in 1954 at theEtang du Luquier257, it has not been seen again and could havedisappeared (changes in water quality, invasion of the banks byPaspalum distichum). • In non-Mediterranean France, it is considered to be scatteredand vulnerable96. • In Italy, the species is decreasing in the north of the country86. • In Morocco, the populations appear to be stable in themedium term, despite wide interannual fluctuations (linked withvariable rainfall).



Anthropogenic factors In Corsica, up until now, these kinds of threats appear to beabsent.

In Morocco and Spain, drainage is the most important threat, aswell as overgrazing in Morocco. In northern Italy, increasingeutrophication* of the water is resulting in the decline of thespecies86, 292.

Natural factors In Corsica, the thorough drying out at the end of spring and insummer and the shallowness and nutrient-poor status of theloose substrate, as well as extensive cattle grazing at the six poolswith L. uniflora, are sufficient to prevent the spread of morecompetitive species (such as Eleocharis palustris).


Current measures Corsica: the Tre Padule site is protected by its legal “RéserveNaturelle” status. Morocco: no management measures.

RecommendationsCorsica: 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 BorgoCollaborators: Grillas P., H. Michaud, L. Rhazi & N. Yavercovski



---

France, national list: Decree of 20/01/1982/JournalOfficiel 13/05/1982, amended byDecree of 31/08/1995/JournalOfficiel 17/10/1995 - Morocco: “very rare” (RR)133, 230

- Italy, national Red Data Book: “vul-nerable”86, 181

- Spain, regional lists (Medina, pers.com.)

39

Plant species

ANGIOSPERMS

LYTHRACEAE

Main synonyms(1) Peplis erecta Moris, P. hispidula Durieu, P. nummulariaefoliaJord., L. biflorum (DC.) J. Gay.(2) L. Salzmannii Jord., L. dibracteatum Guss., L. hyssopifolium L. subsp. salzmanni (Jordan) BonnierL. hyssopifolia L. var. pseudo-bibracteatum Tallon

French names: (1) Péplis dressé; (2) Salicaire à trois bractéesSpanish name: (2) Alheli silvestre177

Italian names: (1) Salcerella a foglie ovali; (2) Salcerella con duebratteeEnglish names: (1) Loosestrife (genus name); (2) ThreebractLoosestrife

SubspeciesNone

Description/identification criteria (1) Lythrum borysthenicum • 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 cmin height.• Flowers solitary, subsessile in the leaf axils.• Calyx expanded at the base, in a short tube, almost as wide aslong, without prominent ribs and longer than the capsule, withfive or six short teeth, blunt, equal or almost equal.• Petals crimson, five or six in number, or absent. • Stamens six, enclosed in each flower.

(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 longby 0.1 - 0.3 cm wide), sessile, alternate.• Flowers small, solitary in the axils of the branch leaves, sub-sessile, crimson; five or six petals, equalling approximately halfthe length of the calyx tube.• Calyx narrowed at the base, very elongated into a tube, muchlonger than wide (0.5 - 0.6 cm in length), with prominent ribsand 8 to 12 very short, triangular, unequal, blunt teeth, and twolinear bracts, green, equal in length to the calyx and resemblingleaves, or alternatively very short and scarious.• Stamens five to six. Style enclosed.• Fruit: cylindrical capsule* approximately equal to the calyx.Seeds yellow, slightly constricted.

Similar species (1) Lythrum borysthenicum may be confused with L. (Peplis) por-tula, but this species has its leaves clearly petiolate and spatu-late, a prostrate stem, petals liliaceous, and calyx shorter thanthe capsule.(2) L. tribracteatum may be confused with L. thymifolia and L. hyssopifolia, but in both these species, the colouration is glau-cous, the habit is upright, the external calyx teeth are longerthan the internal, the calyx is tubular and the stem unbranchedor very little branched.

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

Lythrum borysthenicum

1.5 mm

Lythrum tribacteatum

1 cm

1.25 mm

1 cm

calyx

calyx

40



Distribution/range

(1) Lythrum borysthenicumSub-Mediterranean species.

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

Other Mediterranean countriesPortugal: absent from the north of the country.Spain: western half, to Gerona and Valencia; absent from a largearea of Galicia and Asturias64.Italy: Tuscany, Lazio, Sardinia, Sicily292.Morocco: Atlantic plains and mountains of low to medium alti-tude, in a sub-humid, semi-arid Mediterranean bioclimate*.Also present in Greece, Cyprus, Turkey, Algeria, Tunisia, Libya, Israel.

(2) Lythrum tribracteatumMediterranean species.

FranceAtlantic 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 Mont-pellierais, Etang de Capelle and Costière nîmoise, Petite-Camargue.• 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.

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.

Habitat

General description (1) L. borysthenicum: temporary pools on a non-calcareous sub-

strate (sandstone, granite, quartzite, schist, basalt, etc.), floodedin winter and very dry in summer, with oligotrophic water, butalso edges of marshes and pools, or slow-flowing shallow rivulets(in Les Maures).

(2) L. tribracteatum: temporary pools in plains, low and mediumaltitude mountains in sub-humid and humid Mediterranean bio-climates. In western France, L. tribracteatum also occurs in brackishgrasslands near the coast in depressions with bare wet soil, andplaces that are trampled by livestock96.


CORINE Biotopes22.34 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 (Nanocype-retalia) (France and Spain): (2)

Phytosociology*

MoroccoBoth 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. borys-thenicum is characteristic, along with Ranunculus revelieri, of anIsoetion association which is endemic* to this region. In Languedoc-Roussillon, it occurs in Isoetes setacea formations (Isoetetumsetacei Br.-Bl.)(2) L. tribracteatum is often associated with Damasoniumpolyspermum as a characteristic species of the Lythrion tri-bracteati alliance (order Nanocyperetalia flavescentis)153, 215. It alsooccurs in rice fields in the Camargue, and fairly often togetherwith Crypsis on the edges of ponds in Languedoc. It is also charac-teristic of the Elatinetum macropodae Br.-Bl. (1931), formerlydescribed from Agde. On the central Atlantic coast, it is characteristic of a Juncohybridi-Lythretum tribracteati which occurs in sub-halophilic362

marshes.

CorsicaL. borysthenicum is associated with Isoetes velata and Juncuspygmaeus in the Tre Padule de Suartone pools221, with Elatinebrochonii and Juncus pygmaeus in the Padulellu pool219, with L. hyssopifolia and Cotula coronopifolia in the coastal temporarypool at Tour d’Olmeto Point282.

Spain319

L. tribracteatum is characteristic of the Lythrion tribracteatialliance, as well as the Lythro-Heleochloetum schoenoidis asso-ciation Rivas Martinez 1966, within the Heleochloion alliance.

Ecology


Life formTherophyte (annual species); scapigerous*.

ReproductionAerial.

Biological cycleIn Morocco and Corsica, Lythrum borysthenicum and L. tribractea-tum begin to germinate in February if the site is flooded; flowering

41

Plant species

Impact of perturbationsIt is favourable when it restricts competing plants.



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

FranceL. borysthenicum is very rare in Languedoc-Roussillon, commonerbut local in the Provence-Alpes-Côte-d’Azur Region (Var), fairlycommon at the Corsican pools. L. tribracteatum is rare through-out the country, including the Mediterranean region (local in Lan-guedoc, very rare in the PACA Region and absent from Corsica).

AlgeriaL. borysthenicum is a rare species, while L. tribracteatum is fairlycommon303.

SpainThese species are not considered to be rare. L. borysthenicum iscommon in the western half of the Iberian Peninsula, and L. tri-bracteatum has a wide range over the whole peninsula (27provinces in Spain, six in Portugal)64.

Conservation status

MoroccoThe populations are fairly stable but there is the risk of a declinein the short or medium term as a result of increasing threats.

France(1) Lythrum borysthenicum• In the Mediterranean region of the French mainland, L. borys-thenicum has decreased more than L. tribracteatum. Its status isprecarious in Languedoc-Roussillon, where a number of popula-tions have already disappeared (Petite Camargue, Costière Nîmoise,Grammont, Agde, Lamoure, Saint-Estève pool), while others arein jeopardy (Fenouillèdes, plateaux of Vendres and Torremila). Inthe siliceous parts of Provence the species is surviving well; how-ever, it has disappeared from the Giens peninsula.• In Corsica, at one of the temporary pools (Chevanu pool), usedas a car park from May to September, a very severe decline in thepopulation 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 deGrand-Lieu (Loire-Atlantique)67.

(2) Lythrum tribracteatum• L. tribracteatum is surviving well in the coastal Languedoc andin the lower floodplain of the Aude. It has disappeared from theedges of the Etang de Berre, but about ten localities haverecently been discovered in the Var, which means that its statusis less precarious than is indicated in the Red Data Book274. Somelocalities are, however, threatened by urban development (Agde)or the restructuration of wetlands (Vendres). The very small station



Mainland France

Fruiting and seed dispersal

Flowering



Morocco


Flowering


Jan. Feb. March April May June July Aug. Sept. Oct. Nov. Dec..


Mainland France


Flowering



Morocco


Flowering

Lythrum tribracteatum

lasts from the end of April to July for L. borysthenicum and from thebeginning of May to July for L. tribracteatum. Fruiting begins at theend 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 June-July for L. borysthenicum and in May-June for L. tribracteatum.In Corsica, L. borysthenicum has the following cycle: germinationin February and March, growth in March and April, flowering andfruiting in May and the beginning of June, death of the plantwhen the habitat has dried out, in June.

Lythrum borysthenicum


HydrologyThese two amphibious pool species are often situated in morehydromorphic (low-lying) positions than L. thymifolium and L. thesioides. The germination of the seeds requires the soil to besaturated with water (flooding is not essential). At the vegetativestage the plant can tolerate shallow water (<4-9 cm) but it can-not survive for long under greater depths of water.

Substrate(1) L. borysthenicum grows on usually acid substrates (siliceoussoils).(2) L. tribracteatum occurs on siliceous or calcareous substrates(sand, sand-silt, silt or clay-silt soils). In mainland France, thesoil is often silt-clay, rich in active limestone* and sometimessalty (Camargue, coastal pools of Languedoc, west-centralFrance153, 362.

Interspecific competitionMajor growth of tall plants may have negative effects on theseheliophilous* species as a result of shading. This is the case inmainland France where grazing has been completely discontin-ued at the majority of the sites.

42


in the Hautes-Alpes is vulnerable. In the Aveyron, this Lythrumhas recently been observed as a adventive plant in the valley ofthe Tarn, but the other stations in this département have notrecently been seen.• In the Atlantic area, the populations of L. tribracteatum, for-merly fairly numerous, appear currently to be reduced to a smallnumber of stations*274, 362.

Spain and Portugal L. borysthenicum may be increasing due to its ability to coloniseartificial new habitats (Medina, pers. com.).



Anthropogenic factors The discontinuation of traditional stock rearing, conversion toarable land, urbanisation, infilling with debris, the use of poolsthat are dry or drying out (e.g. as car parks), and drainage, are thecauses of the declines and the vulnerability of the populations ofboth these species.

Natural factorsIn Morocco, at sites with a semi-arid bioclimate, a high fre-quency of dry years, which prevent the renewal of the seedstocks, is a potential threat for both these taxa. In France, themain threat for both species comes mainly from the closing-upof the habitat due to the spread of woody plants.

Risks relating to populationsThe fairly small sizes of the currently existing populations renderthem vulnerable.


Current measures

MoroccoThere are no management measures.

FranceLythrum tribracteatum benefits from the legally protected statusof some sites (Nature Reserves of Roque-Haute in the Héraultand Saint-Denis-du-Payré in western France), and L. borysthe-nicum from that of the Tre Padule de Suartone Nature Reserve(southern Corsica).L. borysthenicum also benefits at the Plaine des Maures (Var) fromexperimental management measures, in the context of the LIFE“Temporary Pools” Project.L. tribracteatum is the subject of annual monitoring at the Lanaupool (Bouches-du-Rhône), and its seeds are preserved ex-situ bythe Conservatoire Botanique National Méditerranéen de Porque-rolles. Finally, several sites (Roque-Haute, Les Maures plain andmassif, Notre-Dame de l’Agenouillade at Agde, etc.) are includedin 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 bymaintaining the plant cover via clearing.• Implementing legal protection measures for the species andthe sites. • Raising the level of awareness among the managers involved.

Bibliography

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. RhaziCollaborators: Paradis G. & M. L. Pozzo di Borgo



---

France: (1) regional list for Languedoc-Roussillon: Decree of 29/10/1997/Journal Officiel 16/01/1998 (2) national list: Decree of20/01/1982/Journal Officiel13/05/1982, amended by Decree of31/081995/Journal Officiel17/10/1995 - (2) France, national Red DataBook: priority species274

43

Plant species

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

ANGIOSPERMS

LYTHRACEAE

Main synonyms(1) L. hyssopifolia L. subsp. thymifolium (L.) Bonnier & LayensL. thymifolia L.(2) L. geminiflorum Bertol.L. thesioides M. Bieb. subsp. geminiflorum (Bertol.) Rouy & E.G.CamusL. purpurascens Châtenier

French names: (1) Lythrum à feuilles de thym, Salicaire à feuillesde thym; (2) Lythrum faux-Thésium, Salicaire faux thésionSpanish names: (1) Salicaria-menor (catalan)Portuguese names: (1) Salicaria-menorItalian names: (1) Salcerella con foglie di timo; (2) Salcerella afiori appaiatiEnglish names: (1) Thymeleaf Loosestrife; (2) Loosestrife (nameof genus)

SubspeciesNone

Description/identification criteria96, 139, 292, 386, 394

(1) Lythrum thymifolium• Annual plant, glaucous, more or less scabrid*, from 3-10 cm inheight, with stem glabrous, erect, spindly, very leafy and not verymuch branched.• Leaves linear, sessileµ, alternate, small (0.5-0.9 cm long x 0.1-0.2 cm wide), narrow, close together, very finely toothed on theedges.• Flowers very small, solitary in the axils of the leaves on thebranches, with short pedicels and with two linear bracts at thebase of the calyx that are variable in size from one individual toanother, sometimes within a single population234, 386.• Calyx elongated, tubular, short (0.20-0.25 cm long), bearingeight very unequal teeth at the tip: four long (0.5-1 mm) andpapillose, alternating with four very short and membranous.• Corolla with four pink petals scarcely extending beyond thecalyx teeth. • Two to four stamens per flower.• Fruit: capsule*, equal to or less than the length of the calyxtube.

(2) Lythrum thesioides• Very similar, though a little taller (up to about forty centime-tres).• Flowers very small, grouped in pairs or threes in the axils of allthe leaves (solitary in L. thymifolium). • Calyx short, bell-shaped, with 8-12 teeth, and corolla rose-purple, with 4-6 petals with darker central veining (in L. thymi-folium the calyx is cylindrical with eight teeth, and the corollahas four uniformly pink petals).

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

Lythrum thymifolium

1 mm

1 cm

Lythrum thesioides

1.5 mm

1 cm

calyx

calyx

44


hexamerous* and pedicellate. The six petals clearly extendbeyond the calyx, which has 12 unequal teeth at the tip (six 0.5-1 mm long, not very papillose, alternating with six short andmembranous). 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, onthe coast or inland. It has a prostrate habit, a stem that is slightlywoody at the base and very branching, petals purplish pink withserrated elongated limbs, and narrow spatulate leaves slightlycurled at the edges.


Distribution/range

(1) L. thymifoliumMediterranean 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.

(2) L. thesioides Sub-pontic species.

FranceLower Rhône valley: Pierrelatte (Drôme), Caderousse and Orange(Vaucluse), Tresques, La Capelle-et-Masmolène, Meynes, andJonquières-Saint-Vincent (Gard).

Other Mediterranean countriesTwo old records from Italy in the floodplain of the Pau.

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

Habitat

General description (1) Habitats suitable for L. thymifolium include pools that aretemporarily flooded or simply saturated in winter, and very dry insummer, and also the edges of streams, marshes and ponds. Thesubstrate is usually siliceous (sandstones, schists, basalts, etc.),the floodwater oligotrophic* and of variable depth between sitesand years.(2) L. thesioides is also associated with habitats that are flood-prone or very wet during the winter, such as the banks of natu-ral freshwater ponds or the alluvial terraces of the Rhône or itstributaries. On the edges of ponds, L. thesioides occurs in the outerband 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 prob-ably formerly at Jonquières, in the Gard).


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 (Nano-cyperetalia)

Phytosociology* and companion species

(1) Lythrum thymifoliumIt belongs to the class Isoeto-Nanojuncetea and to the Cicendionfiliformis 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) asperumBarbero.

There are a large number of associated species: Juncus pygmaeus,J. capitatus (especially in Languedoc), J. bufonius, L. borys-thenicum, L. hyssopifolia, Isoetes velata, I. duriei (ecologicallymore similar, in France, to L. thymifolium than to I. velata), Laurentiamichellii.In the Crau (Lanau pool), the species is associated with species ofthe Lythrion tribracteati alliance.

(2) Lythrum thesioidesVery few facts are known about the habitat of this species, whichhas been observed as much in cultivated land in flood-prone areasas on the shores of temporary ponds. At Jonquières-Saint-Vincent192

as well as at La Capelle-et-Masmolène249, L. thesioides was abun-dantly accompanied by L. tribracteatum. At this station there alsooccurred Potentilla supina, formerly present at Jonquières. At Meyne,L. thesioides was accompanied by several Centaurium (C. pul-chellum, C. spicatum and C. tenuiflorum), Blackstonia serotina,Deschampsia media, Brachypodium phoenicoides, Phleum pratensesubsp. serotinum, L. hyssopifolium etc.359. The habitat of L. the-sioides thus appears to be quite different from that of L. thymi-folium and belongs to the Lythrion tribracteati alliance or eventhe Deschampsion mediae.

Ecology


Life formTherophytes (annual species).

ReproductionAerial.

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 outersurface, applanate-concave on the inner. Integument covered invery short appressed hairs when dry192.

45

Plant species

Biological cycle(1) In Morocco, germination at the beginning of February if thesite is flooded, vegetative growth from February to the end ofMarch, flowering in April-May, fruiting end of May and June, andseed dispersal a short distance from the parent plant. In Franceflowering 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 inJune (one record). The phenology* of this species is thereforelater than for L. thymifolium, and closer to L. tribracteatum forexample. The stamens have anthers which extend to the samelevel as the stigma, which raises the a priori possibility of auto-gamy*. This mode of reproduction is all the more probable as theflowers often lack petals. The capsule, which barely extends out-side the calyx, opens at the tip to form four very short valves.Seed dispersal is consequently probably limited. The seedsundoubtedly remain viable for a long time, as the plant may notappear every year and may disappear for many years betweenappearances at a given place.


HydrologyFor seed germination it is only necessary for the soil to be satu-rated with water. Flooding is not essential. At the vegetativestage the plant can tolerate shallow water (< 5-7 cm).

SubstrateSandy to sandy-silty, even silty for L. thesioides. Substrate aboveall non-calcareous (silica, basalt) for L. thymifolium.

Interspecific competitionDeep shade from tall plants may be unfavourable for these helio-philous* species, especially at sites which are not grazed (France).

Impact of perturbationsFavourable in that the habitat is opened up.



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

Caux-Fontès-Pézenas and Agde, Quaternary siliceous plains ofBéziers and Montpellier in the Hérault, Costière Nîmoise inthe Gard, Plateau du Coiron in the Ardèche, the Issoire area in thePuy-de-Dôme, Upaix in the Hautes-Alpes, the Lanau pool in the Bouches-du-Rhône, Plateau de l’Evenos , the Plaine and Massifdes Maures, Colle du Rouet and Bois de Palayson, Estérel ofFréjus in the Var and the Biot Massif in the Alpes Maritimes246, 274.• In Spain and Portugal: on the Balearic Islands and distributedover a large part of the peninsula, with the exception of theCordillera Cantabrica64.• In Italy: in the three regions of Puglia, Basilicata and Lazio,and in Sardinia292.• In Morocco: pools in the Atlantic plains (Benslimane) and poolsin the Atlas Mountains.

(2) L. thesioidesIn France the plant has only been found 14 times at seven local-ities since it was discovered by A. Jordan in August 1841 at theEtang de Jonquières. The Etang de la Capelle is the only site atpresent (1998).

Conservation status

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

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




Morocco

Fruiting

Flowering

Seed dispersal



France

Fruiting

Flowering

Seed dispersal

?

?



-(2): strictly protected (annexe I)

-(1) and (2) France, national list:Decree of 20/01/1982/JournalOfficiel 13/05/1982, amended byDecree of 31/08/1995/JournalOfficiel 17/10/19951) and (2) France, national RedData Book: priority species274

(1) Morocco: “rare” (R)133, 181, 381

(2) Italy, national Red Data Book:“extinct”86

Lythrum thymifolium

46



Anthropogenic factors Urban development, as well as drainage of pools and ponds, theirfilling in and their conversion to agriculture, constitute the greatestthreats for both species.

Natural factorsIn Morocco, successions of dry years prevent the renewal of theL. thymifolium seedbank*, which could pose a problem for somepopulations in the long term. In France the closing-up of thehabitat, resulting from the steady increase of woody plants, con-stitutes a threat for this species.

Risks relating to populations• (1) In France, the L. thymifolium stations are vulnerable andsubject to the risk of local extinction, especially as the species isat the northern limit of its range. • (2) The same is true of L. thesioides whose very insecure sta-tus in France is perhaps partly mitigated by the longevity of theseedbank in the soil. However, the very severe decline in thespecies throughout its entire range exposes the single currentlyconfirmed population to particular risk.


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

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

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

Bibliography

Anonymous, 199912 ; Aubert & Loisel, 197118 ; Cabanès, 190358 ;Castroviejo, 1986-200164 ; Charrel, 191370 ; Châtenier, 192271 ;Conti et al., 199286 ; Danton & Baffray, 199596 ; Fennane & IbnTattou, 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. RhaziCollaborators: Antonetti P., L. Garraud, J. Molina & G. Paradis

47

Plant species

Marsilea strigosa Willd.

POLYPODIOPHYTES300

MARSILEACEAE

Main synonymsM. 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)

SubspeciesNone

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 across at the apex of the petiole (rachis), like a Four Leaves Cloverleaf.• Two types of fronds: - The first with a floating blade, glabrous, borne on a long peti-ole, 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 intwo rows along the rhizome.

Similar species Water Shamrock (Marsilea quadrifolia L.; syn.: M. quadrifoliata(L.) L.) has always glabrous leaflets and sporocarps with shortpedicels, 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 therisk of confusion. In Spain, M. strigosa can be confused with M. quadrifolia as wellas with an endemic species of the Iberian Peninsula, M. batardae (Medina, pers. com.). The latter species has the rhi-zomes 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 anddistinctly pedicellate sporocarps, grouped in twos or threes


Distribution/range160, 386

Marsilea strigosa is a western Mediterranean species; it alsooccurs in some localities to the north of the Caspian Sea. It ispresent in France, Italy, Sardinia, Spain, the Balearics, Algeria,Morocco, Egypt, Romania and Ukraine (Crimea). Its existence inPortugal has been disproved.

FranceIt is only found today in three localities: • Hérault: pools in the Roque-Haute plateau and a wet depres-sion in the wine-growing plain of Vendres. • Pyrénées-Orientales: alluvial terraces of the Têt, in the uncul-tivated land of Torremila.

Other Mediterranean countries

Italy: south of the peninsula (several localities in the province ofPuglia, 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 andMinorca64.

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).

Egypt: Nile Valley and Delta46.

Habitat

General descriptionMarsilea strigosa appears to occupy a relatively wide range ofnatural habitats and substrates, and to be part of several phyto-sociological units, some of which remain to be specified2.

Marsilea strigosa

1 cm

48


The species grows mainly in temporary pools with nutrient-poorand poorly mineralised water (Medina, pers. com.). It is foundwithin amphibious vegetation which are submerged for a fairlylong time (in France at Roque-Haute, or in Spain), dominated byIsoetes setacea or Mentha cervina, or in Crassula vaillantii-Lythrum borysthenicum formations2, 251, 319.In Morocco, in the Atlantic plains, it grows in Isoetes velata,Ranunculus peltatus and Myriophyllum alterniflorum forma-tions312.It also occurs in regularly flooded former vineyards, cleared ofvines on siliceous gravel* (Vendres plateau and Plaine du Roussillon,France), often in furrows in which the water collects.




Phytosociology*

FranceMarsilea strigosa is considered a characteristic species of theIsoetetum setaceae association* Br.-Bl. (1931) 1935215. On aban-doned farmland, the formations have yet to be incorporated intophytosociological units2. These are most likely Isoetion formationsheavily disturbed by viticulture.

SpainThe species is characteristic of the Sisymbrello-Preslietum asso-ciation (Preslion cervinae alliance319).

Companion species

France • In temporary pools (Roque-Haute and Torremila): Isoetessetacea, Pilularia minuta, Mentha cervina, Lythrum borys-thenicum, L. thymifolium, Eleocharis palustris, Myosurus brevis-capus, Juncus pygmaeus, Pulicaria vulgaris and Polygonumromanum subsp. gallicum49 (Grillas & Tan Ham, pers. com. CEN-LR,pers. com. ).• On abandoned farmland, an impoverished form of this com-munity can be found (disappearance of Isoetes, Mentha cervina,Pilularia minuta and Eleocharis palustris) augmented by ruderalspecies (Conyza spp., Dittrichia viscosa, etc.)2, 246.

MoroccoIsoetes velata, Ranunculus peltatus Lythrum borysthenicum,Myriophyllum alterniflorum, Eleocharis palustris and Baldelliaranunculoides312.

Ecology


Life formSpecies considered to be perennial* (radicant hydrophyte*, produc-ing stolons, traditionally considered as a geophyte* with a rhi-zome), but which appears to behave as an annual (therophyte*)395.

SporocarpsSuborbicular, tightly packed, sessile* or with very short pedicels(2 to 2.5 mm) covered in articulated hairs300, with a diameter of3 to 5 mm. These are highly efficient survival organs due to theircapacity to survive several decades, or even over a century ofdrought. 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 cycleThe sporocarps*, often saturated with water from the autumn,open in late winter and early spring, releasing the microsporan-gia and macrosporangia. Fertilisation occurs over the followingdays. The first, filiform, fronds appear within a few days, followedby fronds with two leaflets, then with four leaflets. They appearunder a thin layer of water, during the dry phase of the pool, inwinter in Morocco (January-February) and in spring in France (inMarch-April at Roque-Haute).

When the soil is waterlogged, vegetative growth is prolific. Therhizome continues to grow, emitting a large number of fronds(dense tufts) as well as stolons which take root, enabling newrhizomes to develop. The formation of sporocarps and spores hasbeen observed in March-April in Italy86 and from April to June inFrance.

When the pool dries out completely, the plant acquires its ter-restrial habit. The fronds wither and disappear towards the endof the summer in Morocco (Rhazi L., pers. com.) and up until theautumn in France. The plant then only survives in the form ofsporocarps arranged in two tight rows on either side of the dried-out rhizome.


Emergence of fronds and vegetative growth

Withering of fronds

Formation of sporocarps?

France


HydrologyThe habitat of Marsilea strigosa is characterised by a fairly longinundation period, between five and eight months: from De-cember to May-July in France (Rhazi M., pers. com.); from Decem-ber to March in North Africa312, and by a maximum water depthof around 40 to 50 cm. In experimental conditions (not published),the opening of the sporocarps and fertilisation are observedunder a thin layer of water, or even on saturated ground.

SubstrateUsually non-calcareous (basaltic plateau at Roque-Haute, siliceousgravels at Vendres and at Saint-Estève).

Interspecific competitionThis species is sensitive to competition by perennial grasses andto the shade of woody plants (heliophilous* species).

49

Plant species

Impact of perturbationsThe rhizomes of Marsilea do not appear to be sought by WildBoar, though they can be dug up by chance by these animals.They are resistant to the turning-over of the top layer of soil invineyards.



FranceVery rare species with only three sites known. At Roque-Haute,over an ensemble of 200 pools, Marsilea appears to be dissemi-nate in small populations in 10 to 26 pools depending on inter-annual variations in rainfall162, 246. On Torremila, the species ispresent in small scattered populations in three humid areas ofuncultivated land and one temporary pool.

ItalyConsidered as very rare (a few localities).

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

MoroccoMany localities. The species, more common than M. minuta, isnot considered to be rare or threatened133.

AlgeriaRare species303.

Conservation status

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

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

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




FranceThese are the main threats. The habitat of the Vendres andTorremila stations has been degraded and partially destroyed bydrainage and cultivation. It has returned to a wild state, wherethe species survives, though it risks being put under intensivecultivation again in the short term66. Only one intact pool sur-vives among the vineyards of Torremila. At Roque-Haute, despitethe “Réserve Naturelle (Nature Reserve)” status, the absence ofcontrol over land ownership and usage no longer permits man-agement or monitoring of the site, and seriously compromisesthe conservation of the Marsilea populations.

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

Natural factors

Vegetation dynamicsAt Roque-Haute, following the abandonment of grazing severaldecades ago, the spread of woody plants and invasion by Scirpusare threatening the habitat of Isoetes setacea315, which is alsothat of Marsilea251. At Torremila, the species is subject to compe-tition from perennial grasses (mainly Deschampsia media),encouraged by the natural dynamics of the uncultivated land.

Variability of hydrological conditionsThe interannual fluctuations in flooding cause fluctuations in theabundance of Marsilea without jeopardising the survival of thepopulations.

Risks relating to populationsIts very reduced capacity for dispersal results in a poor aptitudefor the colonisation of new sites, and significant isolation of pop-ulations, both geographically and genetically395.

IUCN 1997Bern Convention



-Strictly protected species (Annexe I)

Annexes II and IV - France, national list: Decree of 20/01/1982/ Journal Officiel13/05/1982, amended by Decree of31/08/1995/ Journal Officiel17/10/1995- France, national Red Data Book: priority species.- Italy, national Red Data Book: “vulnerable”86

- Spain, regional lists (Medina, pers. com.)- Balearics: “vulnerable”331

50



Current measures

FranceThe Roque-Haute site is protected by its legal “Réserve Naturelle”status. Marsilea was the subject of regular monitoring here from1992 to 2001. At Vendres, the site is included in a Natura 2000* area, and theConservatoire des Espaces Naturels du Languedoc-Roussillon (CEN-LR) has acquired land to favour the restoration of the hydrologicalregime.At Torremila, the Conservatoire has drawn up the Objectives Docu-ment (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 thesetting-up of a LIFE project for the conservation of Alpine andMediterranean flora of EEC interest at the European scale(France, Spain, Italy).Conservation ex-situ: sporocarps have been collected by theConservatoire Botanique National Méditerranéen de Porquerolles.Cultivation trials successfully carried out in January 2004, par-ticularly within the framework of a project for the restoration ofthe Vendres site, show the viability of harvested sporocarps.

SpainOnly a few localities are included in Natura 2000 areas. Some arethe subject of particular programmes (Province of Valencia,Balearics) (Medina, pers. com) such as the creation of two micro-reserves in the lagoons of Sinarcas (Province of Valencia) and theproduction of sporocarps ex-situ from fragments of rhizomeswith a view to the reintroduction of the species179. The species isclassified as “threatened” on the regional lists of the regionswhere it is present (Medina, pers. com.).

ItalyThe species is cultivated in the botanical gardens at Padua86.

Recommendations (for France)

ViticultureBan intensive viticulture on all sites, as it is totally unfavourableto the survival of Marsilea populations through the impact ofvarious agricultural practices (drainage, burying of sporocarps,herbicides). On the other hand, outside of the temporary-poolhabitat, extensive viticulture (no herbicides, light cultivation of

the soil surface) is not incompatible with the conservation ofMarsilea populations, provided it permits flooding from autumnto spring.

Competition from perennials and woody plants (closing-up of thehabitat)• In the Roque-Haute Nature Reserve: implement the manage-ment plan and more generally restore a concerted managementwith a natural-heritage objective: reintroduce grazing, preferablyby sheep and goats, together with an appropriate monitoringprogramme.• At Torremila: promote management of perennial grasses bywater-weed cutting and grazing.

Isolation of populations395

Genetic isolation has implications for conservation strategies: toproperly 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 usesRestore control over land ownership and/or usage, an importantprerequisite for the implementation of conservation manage-ment 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 ; Braun-Blanquet et al., 195249 ; Castroviejo, 1986-200164 ; CEN-LR,200166 ; Colas et al., 199683 ; Conti et al., 199286 ; BernConvention,197987 ; Danton & Baffray, 199596 ; Directive92/43/CEE, 1992119 ; Fennane & Ibn Tattou, 1998133 ; Fennane etal., 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

51

Plant species

Mentha cervina L.

ANGIOSPERMS

LAMIACEAE

Main synonymsPreslia cervina (L.) Fresen.

French name: Menthe des cerfs Spanish names: Menta de burro, Poleo de ciervoMoroccan name: Fliyou

SubspeciesNone

Description/identification criteria274, 292, 386

• Perennial plant with rhizome very close to the surface42,recognisable by its very musky mentholated scent, its uprighthabit, 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 smallerleaves borne on very short branches), and above all, by the char-acteristic palmate shape of the bracteoles* surrounding thewhorls of flowers, which evoke the antlers of a deer, and to whichthe 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 orroughly toothed.• Flowers hermaphrodite, forming compact whorls, widelyspaced on the upper half of the stem. Calyx tubular and villousat the throat, with four triangular teeth terminating in a finepoint. Corolla twice as long as the calyx, with two equal lips, pinkor white. Floral bracts identical to the leaves, but larger.

Similar species During and after flowering, no confusion is possible: it is the onlymint which has deeply lobed bracteoles and whose calyx has fourteeth and not five, hence its former inclusion in the genus Preslia.In the vegetative state, the narrow, sessile leaves are characteris-tic and differentiate it from the Pennyroyal (Mentha pulegium),frequently present in the same habitats but whose leaves arerounded with short petioles.Before flowering, the young shoots do not yet have their scent ofmint, and can be confused with young Hedge Hyssop (Gratiolaofficinalis) plants.


Distribution/range

Western Mediterranean species, found in six countries160, 386.

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

Other Mediterranean countriesSpain and Portugal: western half of the Iberian Peninsula, withsome localities in the eastern part AND in the Balearics (Medina,pers. com.).Italy: once found in the wetlands and temporary pools ofAbruzzo, but not seen recently292.Algeria: temporary pools of the high plateaux of the Algerois andthe Oranais, Geryville region303.Morocco: dayas* and edges of peat bogs of the Rif regions (RhaziL., pers. com.).

Habitat

General description Species of the thermo and meso-Mediterranean zone.

- In France, this mint occupies the long-flooded parts of tempo-rary pools and lavognes*, and a number of temporary watercourses of the Montpellierais garrigues (Molina, pers. com.). Itonce occurred in the ox-bows of the Rhône (breaches andlaunes* around Tarascon, Avignon, etc.), where hydraulic modifi-cations caused its disappearance. It forms, with the often peren-nial species associated with it, areas of tall-herb grassland ofseveral dm2 to several m2. - In North Africa (Morocco, Algeria), the species grows at an alti-tude where the rainfall is sufficient (above 1,000 m in the

Mentha cervina

1 cm

52


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


CORINE Biotopes22.34 Mediterraneo-Atlantic amphibious communities.

22.342 Tall Mediterranean amphibious swards (Preslion cer-vinae): vegetation of tall annuals of land covered by deepwater for long periods, with Mentha cervina.

Phytosociology*Class: Isoeto-Nanojuncetea

Order: IsoetaliaAlliance: Preslion cervinae

Associations*: • In France49, 215: - Preslietum cervinae: Mentha cervina-Eleocharis palustris asso-ciation, described from western Languedoc.- Preslio-Trigonelletum ornithopodioides: association describedfrom 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 bySisymbrella aspera, Veronica anagalloides, Cyperus Badius, Mar-silea strigosa, Eryngium galloides, Callitriche platycarpa, etc.- Preslio-Eryngietum corniculati: M. cervina is present here withEryngium corniculatus, I. velata, I. setacea, etc.

Companion species

France At Roque-Haute, the mint grows with Marsilea strigosa andIsoetes setacea, or with Eleocharis palustris in the deepest partsof the pools251.In the Montpellierais and Uzégeois garrigues the companionspecies are Eleocharis palustris, Sisymbrella aspera, Juncus articu-latus and Mentha pulegium.

MoroccoEleocharis palustris, Callitriche truncata and Alisma plantago-aquatica are present at its stations.

Ecology


Life formPerennial species with vegetative growth (suckers, rhizomes).Although considered by several authors as hemicryptophyte*, inFrance this species is more of a rhizomatous geophyte*, as it isnot visible before the beginning of the spring, and the youngshoots appear towards the month of April.

Biological cycleIn France, its vegetative growth occurs in the spring, and itsreproductive cycle in summer. At Roque-Haute, it flowers fromlate June to mid-August, fruits from late July to September andwithers in October.

In Morocco, its development is earlier: the shoots appear inMarch, flowering commences in May and fruiting in early sum-mer.


Emergence of shoots and vegetative growth

France

Fruiting

Flowering



Morocco

Fruiting

Flowering


HydrologyIn France, the habitat of this species is characterised by a longperiod of flooding, from the autumn to spring153 (six to eightmonths depending on the year), a depth of water of between 20 and70 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).

SubstrateVarious substrates, oligotrophic*274 or meso-eutrophic*98, onbasaltic (Roque-Haute), calcareous (garrigues), marl (Minervois),or granitic (Fenouillèdes) bedrock, or on alluvium (borders ofponds).

Interspecific competitionThis heliophilous* species is sensitive to shade. The water depthlimits invasion by woody species, but on the other hand createsconditions favourable to colonisation by fiercely competitivehelophytes* (reeds, Scirpus, Reedmace, sedges)153.



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

53

Plant species

Conservation status

FranceThe populations of this species seem to be undergoing significantdeclines over the whole of its range, except in the Hérault andthe Gard.• Gard: the species remains abundant in the Uzégeois-Bagnolais(plateaux of Méjannes-le-Clap, Lussan, etc.), and is surviving in theregion of Quissac (pools at Vibrac). It is still present in the marshesof Pujaut157, 246 and further north in a lavogne* in the eastern endof 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 theother populations are stable (temporary pools and streams of theMontpellierais garrigues, the Bittérois plain, and the plateau ofRoque-Haute, the Assignan pool in the Minervois).• Pyrénées-Orientales: the species has disappeared from theSaint-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 (Etangdes Aulnes), all the stations of the “Crau humide” (Marais duVigueirat, 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.

Other Mediterranean countries• In Italy, the Abruzzo populations are considered to have disap-peared292.• In Spain and Portugal, the species could be declining due to thedestruction of its habitat (Medina, pers. com.). • In Morocco, the populations appear to be stable.



Anthropogenic factors In France, habitat destruction is responsible for the disappearanceof 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 ofthe lavognes of the garrigues).• Game management of lavognes (sealing of the pool bed witha layer of concrete).• Extraction of materials: the Plateau de Rodès station is nowthreatened by the creation of a granite quarry246.

Natural factorsThe shade linked to invasion by woody species, as a result of theabandonment of grazing at some sites (France), is without doubtunfavourable for this species.

Risks relating to populationsIn North Africa and in France, with the exception perhaps of theMontpellierais and Uzégeois garrigues and the Roque-Haute pla-teau, a risk of extinction of populations exists in the medium tolong term, because of their small number, their distance from oneanother, and their reduced size.


Current measures274

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

Recommendations• Maintain a favourable hydrological regime• Maintain open vegetation by cutting or grazing. • Protect the habitat at the remaining stations (Rodès plateau inthe Pyrénées-Orientales; Etang de Pujaut, Quissac pools etc, inthe Gard). • Raise awareness among the rural public of the interest andrichness of the many lavognes of garrigue zones so they can bemaintained and prevent their beds from being concreted over.• Intensify prospecting in the areas where only a few popula-tions 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 theirhabitat.

Bibliography

Aizpuru et al., 20006 ; Anonymous, 199912 ; Bissardon & Guibal,199739 ; Bonnier, 199042 ; Braun-Blanquet et al., 195249 ; Conti etal., 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. YavercovskiCollaborators: Rhazi L., M. Rhazi, F. Ruchon F. & H. Souheil



---

France, PACA regional list:Decree of 9/05/1994/JournalOfficiel 26/07/1994 - France, national Red Data Book:priority species274

- Morocco: “very rare” (RR)181, 133

54


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 rioItalian name: Millefoglio d’acqua gracileEnglish names: Alternate-flowered Water Milfoil, Slender WaterMilfoil

SubspeciesNone

Description/identification criteria • Aquatic plant, herbaceous, slender (10 to -50 cm), rooted,glabrous, branching.• Leaves green, 6 to 30 mm in length, sometimes reddish, pin-nate hair-like. Submerged leaves whorled, aerial leaves situatedon the fertile branches and almost all alternate. • Bracts of upper flowers entire or toothed, shorter than theflowers.• 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 droopingspike, consisting of three to four female flowers in a whorl at thebase and five to eight male flowers in a spike at the tip (witheight stamens).• Ovary inferior, with four carpels, each one containing four loculiwith 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 pecti-nate and longer than the flowers. The spike is very elongated,terminating in leaves; the flowers are pink.


Distribution/range

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

France• On the mainland, the species is absent from theMediterranean region, and elsewhere occurs in streams andponds with a siliceous soil.• In Corsica, it is found in seven temporary pools in the southeast of the island: four pools in the Tre Padule de SuartoneNature Reserve and three pools, of artificial origin, on theFrasselli plateau.

Habitat

General descriptionIn Morocco and in Corsica, this species is found only in tempo-rary pools. In Morocco, it is present in the pools on the plains(region of Benslimane) and mountains in the Rif region (Outka andIssaguene), on siliceous soils (sandstone, quartzite) and in a semi-arid bioclimate*, sub-humid or humid. These pools are generallyfilled with rainwater in December-January and dry out in April-May.


CORINE Biotopes This species appears in the aquatic phase of oligotrophic* tem-porary pools. 22.34 Southern amphibious communities (Isoetalia).

Phytosociology*In Morocco, the species is characteristic of Myriophyllum alterni-florum-Ranunculus peltatus communities. The species associatedwith this community are Callitriche brutia, Glyceria fluitans,Chara vulgaris, Nitella opaca and Illecebrum verticillatum.In the pools of Corsica (pools with Isoetes velata), the associatedspecies are Ranunculus peltatus, Illecebrum verticillatum andApium crassipes, as well as Potamogeton pectinatus on theFrasselli pools (which indicates slight eutrophication linked tothe presence of livestock).

Myriophyllum alterniflorum

0.5 cm

55

Plant species

Ecology


Life formPerennial species in permanent waters (hydrophyte*) but with anannual cycle (therophyte*) in temporary Mediterranean pools.

Reproduction Aerial pollination. Flowering in March-April.

Description of seedsSeeds: four per fruit, small (length: 1.5 mm; width: 1 mm to 1.25mm).

Biological cycleThis begins under the water and often ends out of the water. In Morocco and Corsica, germination takes place in February-March, when the pool is under water, and flowering in March-April. Fruiting, which begins during the aquatic phase, takesplace in May-June, but the maturation of fruits and seeds, andtheir dispersal a few centimetres from the mother plant, occursin June-July after the drying-out of the pool. In addition to thissexual reproduction, the plant reproduces vegetatively during theaquatic phase, by division and regrowth.

Impact of perturbationsFavourable when they facilitate the opening-up of the habitat.

LightThe species is heliophilous*, which explains its absence in theturbid pools of the Cork-Oak forest of Mamora in Morocco andin deep pools (with greater depths of water).



• In Morocco, Myriophyllum alterniflorum has been found in 14 pools133, 230 in the Atlantic plains (Benslimane) and the moun-tainous region of the Rif. • In Algeria, the species is considered to be rare. There are fewsites with M. alterniflorum and these are confined to just tworegions, the Algiers region (coastal subregion) and the Kabyleregion (Greater and Lesser Kabyle and Numidia)303, 338.• In Mediterranean France, there are seven pools, with M. alter-niflorum, three of them artificial, and all of them in Corsica.• On the Iberian Peninsula and the Balearics, the species is widelydistributed (40 provinces in Spain, five in Portugal).

Conservation status

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



Anthropogenic factors Filling in, urbanisation, cultivation and the modification of thehydrology of pools (drainage) constitute the main threats for thistaxon. In Morocco, there is a significant risk of extinction of pop-ulations at some stations because of urbanisation and the filling-in of pools.

Natural factorsIn Morocco, the frequency of dry years limits the appearance ofthis species313 and reduces the chances of renewal of the seedbank *, which could pose problems in the long term.In Corsica, the closing-up of the habitat (invasion by woodyspecies or by helophytes) is a potential threat for the species ifgrazing is abandoned.


Germination

Fruiting

Flowering

Seed dispersal

Morocco and Corsica


HydrologyThe germination of seeds requires flooding or at least waterlog-ging of the soil. The plant is adapted to submerged conditions(divided leaves, etc.) and can withstand high water levels pro-vided there is sufficient light: tolerance to water depth increaseswith the clarity of the water (for example 50 cm in the pools ofBenslimane in Morocco or Padule Maggiore in Corsica). The plantforms its flowers under the water but they emerge above the sur-face of the water for opening and pollination (anemophily). Thewater is oligotrophic* to mesotrophic*.

SubstrateAcidic to neutral.

Interspecific competitionM. alterniflorum is abundant in the centre of pools, where herba-ceous species such as Ranunculus peltatus, Callitriche brutia, etc.grow, but it becomes less abundant or even absent in areas withhelophytes* (Scirpus maritimus, Eleocharis palustris), due to com-petitive exclusion.


----

- Morocco: “very rare” (RR)133, 181, 381

- Italy, national Red Data Book:“vulnerable”86

56



Current measures In Corsica, the populations of the Tre Padule de Suartone poolsbenefit 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 thevegetation (woody species and helophytes), and maintaining orintroducing grazing.• Implement legal measures for conservation of non-protectedsites.

Bibliography

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, 1962-1963303 ; Rhazi et al., 2001313 ; Rhazi, 2001312 ; Samraoui & deBelair, 1998338 ; Titolet & Rhazi, 1999381 ; Tutin et al., 1964-1980386.

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

57

Plant species

Nitella opaca (Bruzelius) Agardh

CHAROPHYCÉES

CHARACEAE

Main synonymsChara opaca Bruzelius 1824N. flexilis var. flexilis f. opacoides R.D. Wood 1962

French name: none English name: Dark stonewort

SubspeciesNone


• Dark green plants, small in size (10-30 cm), living entirely sub-merged. Thallus with whorled structure, entirely without cortica-tion.• Whorls composed of 6-8 primary rays (branchlets), dividedonce to form 2-3 secondary rays (dactyls). Dactyls always uni-cellular with a blunt tip. Gametangia* situated in the forks of theprimary rays.• Dioecious species, very fertile with a marked sexual dimorphism.• Male plants characterised by dense fertile heads with shortdactyls; antheridia* solitary and of large size (diameter 650-775 µm). • Female plants with longer primary and secondary rays, givingthem a more open appearance; fertile nodes carrying up to three,though usually two oogonia* 600-700 µm long and 500-600 µmwide. Oogonium bearing a coronule* composed of 2 x 5 tiny cells.Oospores laterally compressed, dark brown to black, c. 350-500 µmlong and 350-400 µm wide with 6-7 spiral ridges. In the freshstate, the oospores have characteristic “winged” extensions ofthe spiral ridges.- Incrustations of calcite forming more or lesse regularly spacedrings around yhe axis and branchlets, often occur at stations witha low water level. This feature, known as “banded phenomenon ofthe 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 insummer or autumn, not in spring.• N. capillaris (Krokeil) Groves and Bullock-Webster, which alsohas spring growth but is distinguished by gelatinous mucusaround the male and female gametangia90, and by its oosporeswhich are ornamented with papillae131, 357.

N. opaca is, with N. tenuissima, the only representative of thisgenus so far recognised in Languedoc-Roussillon and southeastFrance. It is easily distinguished from N. tenuissima which hasprimary branches with two to four forks (multiple dichotomy).


Distribution/range

FranceNitella opaca is mainly present in central and western France(Aquitaine andBritany)90. The numerous localities listed by Corillionall date from earlier than 1957 and even from the beginning ofthe 20th century. A single earlier mention of N. opaca concernssouthern France near Hyères in the Var (coll. Boulu 1851, deter-mined and then cited by Hy178). Today, there are only five locali-ties known in the south of France: Lanau pool (Bouches-du-Rhône), the temporary lakes of Bonne Cougne and Redon, (Var)358,some pools in the natural reserve of Roque-Haute (Hérault) andone 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 countriesRare 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 sedimentsof a fresh water locality close to the Mediterranean coast131.

Habitat

In the Mediterranean region, the species has been identified onlyin temporary freshwater habitats, characterised by flooding atthe end of the autumn and drying out in May/June. This type ofhabitat seems to be consistent both in southern France and thelocalities in North Africa.


Nitella opaca

male female

1 cm

Mod

ified

fro

m G

rove

s &

Bul

lock

-Web

ster

164

58



22.341 Short Mediterranean amphibious swards (Isoetion).22.342 Tall Mediterranean amphibious swards (Preslion cer-vinae).

Phytosociology*N. opaca is part of the aquatic phase of Preslion and Isoetioncommunities (Isoetalia).This species is also characteristic of the Nitelletum opacae asso-ciation, often in monospecific*90 populations or associated withSphaerochara (personal observation).

Companion species

At the Lanau pool in the Crau (Bouches-du-Rhône), N. opaca isassociated with Sphaerochara intricata355. These two species pre-cede the other hydrophytes* and decompose during the emer-gence of communities of amphibious annuals with Damasoniumpolyspermum, Lythrum tribracteatum, Mentha pulegium, etc.

At Lake Redon (Var), N. opaca is accompanied by Sphaerocharaprolifera. The growth of this species also early and precedes thatof Ranunculus ophioglossifolius, Baldellia ranunculoides and Buto-mus 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 Myrio-phyllum alterniflorum, Callitriche brutia, Glyceria fluitans, Charavulgaris and Illecebrum verticillatum312.

Ecology


Life formNitella opaca is an annual species.

Biological cycleThis is a vernal species, which grows while the water is cold.Thus, it appears very early in shallow habitats, while in westernFrance (Britany) and in northern Europe, it is found up until thesummer in permanent lakes and even down to a depth of 40 min Sweden90, 200. In the Mediterranean region, it is present fromFebruary onwards and fructifies abundantly from the end ofMarch to May. The plants then decompose entirely.

Environmental conditions90, 200

HydrologyN. opaca is a fresh water species of habitats with a pH* close toneutral (6.5-7.5). N. opaca can be found in both temporary andpermanent habitats and also adapts to weak currents. The mini-mum duration of submersion necessary to complete a full bio-logical cycle is estimated at 5-6 months.

SubstrateIndifferent to substrate.

Interspecific competitionBecause of its early appearance, N. opaca is not subjected tomuch competition from other hydrophytes.

Impact of perturbationsBecause it is abundantly fertile, the species is not very sensitive to“mechanical” disturbances, and even seems favoured by the tram-pling of livestock (Lanau pool) and wild boar (the longer lasting wetparts of Lake Bonne Cougne), which limit the terrestrial vegetation.

TemperatureGermination in cold water, fruiting between 12 and 18°C; N. opaca decays when the temperature of the water exceeds20°C (personal observation).

LightNot very light-demanding; however, sexual reproduction is reducedin deep lakes90.



FranceDistribution is concentrated in central and western France whereit nonetheless appears to be in decline. An estimate of the num-ber of sites in these regions is not currently possible. Five locali-ties are known in the French Mediterranean region.

Conservation status

FranceIn overall decline: more of 32 localities listed by91 for westernFrance, mainly Britany, had only been collected at the beginningof the 20th century.

Other European countriesUnevenly distributed in northern Europe; considered as fairly fre-quent, but nonetheless vulnerable, in Spain79, while Krause200 men-tions rare sites in central Spain. The species has been recordedrecently in the Balkan countries40.



Reproduction

vegetative growth

Decay

�� Germination and


-----

Figure established on the basis of monthly observations at the temporary lakes ofBonne Cougne and Redon in 2001358.

59

Plant species

There is currently no legal protection for charophytes in Franceand the Mediterranean countries, nor are they included in anyRed List of threatened species.


Anthropogenic factors In France:• The hydrological regime and the water quality of the LakeBonne 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 ownershipand usage (ban on access to the site by owners) is preventing themonitoring and the management of populations of Nitella opaca.


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

RecommendationsIt is essential to maintain the temporary nature of the hydrolog-ical regime with flooding during the cold season. In particular, inthe Mediterranean region, care must be taken to maintain thenatural alternation of flooding/drying out, which favours sexualreproduction and, because of this, enables the plant to spread tonew pools.

Populations of N. opaca are vulnerable due to the fragility oftheir habitat and it is thus important that the dynamics of thesepopulations 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 etal.,1986310 ; Rhazi, 2001312 ; Soulié-Märsche & Thiéry, 1998355 ;Soulié-Märsche & Vautier, 2004356 ; Soulie-Märsche, 1989357 ;Soulié-Märsche, 2003358 ; Wood, 1962403.

Author: Soulié-Märsche I.

60


POLYPODIOPHYTES300

OPHIOGLOSSACEAE

Main synonyms(1) O. ambiguum Cosson & GermO. polyphyllum auct. non A. Braun(2) O. vulgatum L. subsp. lusitanicum (L.) Bonnier & Layens

French names: (1) Ophioglosse des Açores, (2) Ophioglosse duPortugal Italian names: (1) Ophioglosso delle Azzorre, (2) Ophioglossolusitanico 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


• Perennial plants, delicate and unobtrusive from 5 to 10 cm (1),and 2 to 5 cm (2), present in the form of small, fairly localisedpopulations• Two types of fronds (“leaves”) very distinct, as in all theAdder’s-Tongues: a sterile type, with an extended blade, oval orlanceolate, sometimes lying flat but usually upright and curledover to form a furrow; the other fertile, reduced to a fruitingspike with a peduncle and bearing sporangia stacked up on topof 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 sterilefronds generally sheathing the fertile fronds), light green or yel-lowish-green in colour, and yellowing completely before wilting,after sporulation (production of spores).• Generally one single sterile frond per plant, but can grow inpairs in O. azoricum, in which the fronds are broader. • Spikes (or fertile fronds) do not invariably appear on everyplant.

Similar species 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 neartemporary pools, it is generally found in more eutrophic* habi-tats, undergrowth or damp grasslands.O. azoricum and O. lusitanicum are closer in size and share habi-tats that are at first sight similar, but are in fact different interms of hydrology. The easiest distinction criterion is theautumn-winter growth period for O. lusitanicum, spring for O. azoricum. In early spring the two species can be present at thesame time but their stages of development (colour, sporulation)are visibly out of step.Finally, note that the number of chromosomes differs betweenthe three species: O. lusitanicum is diploid* (2n=240 chromo-somes), O. vulgatum is tetraploid (2n=480) and O. azoricumhexaploid* (2n= 720), suggesting that this latter species is derivedfrom a cross between the other two.


Distribution/range

(1) Ophioglossum azoricumWestern European species present in France and around theMediterranean in Spain, Portugal, Sardinia and Italy160. It alsooccurs locally in Slovakia and in the Czech Republic, the Canariesand Madeira (Azores)300.

France• Atlantic coast.• Mediterranean region: O. azoricum is present in the Plaine desMaures and the Estérel (Var), in the garrigues of the Hérault, theCévennes in the Gard and Lozère, and in Les Fenouillèdes (Pyrénées-Orientales). In Corsica, this species is only found in rarespozzines* on the Tenda Massif and the Plateau du Coscione142.

(2) Ophioglossum lusitanicumMediterranean and Sub-Atlantic species present all around theMediterranean 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, theBiot Massif and Les Albères. Formerly reported from the marshesof Coustière in the Crau. Fairly common in Corsica at low altitude141.

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

Ophioglossum azoricum & Ophioglossum lusitanicum

1 cm

61

Plant species

Other Mediterranean countries• Europe: Portugal, Spain, Italy, Yugoslavia, Greece, Crete andCroatia.• North Africa: Algeria (in the Constantinois Tell), Tunisia andMorocco (Middle Atlas, Rif mountains, north and mid-Atlanticplains).• Turkey, Lebanon.

Habitat

General description

(1) Ophioglossum azoricumThe habitats favourable to this species are very wet in winter andspring but rarely flooded for a long period. These are generallydepressions, beds or edges of small streams and springtime tem-porary runnels on well-exposed hillsides on thin soils (Cévennesand Avants-Monts in Languedoc).Its habitat, in the Mediterranean region, is short swards, open orin a mosaic with the shrubby maquis vegetation, on thin, ratheracidic, soil. The habitats found in the Atlantic region are coastalswards, dune slacks, sandstone shelves or other types of rock sur-faces. On the mainland, the stations are not found above 700 min altitude but in Corsica they are found up to 1,500 m in peatybasins (pozzines), which remain damp in the summer.The substrate is not very thick, sandy and often acidic.

(2) Ophioglossum lusitanicumThe 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 themaquis and small basins in rocky outcrops. However, the speciesrarely mixes with the preceding species: it seems to seek stationswith even thinner substrates, and seems less demanding of awater supply. It is clearly more thermophilic and is even foundaway from the low-altitude coastal zone.In Morocco, O. lusitanicum is mainly found in the sandy or stonypastureland of the plains and low hills under a Mediterraneanmild semi-arid and sub-humid bioclimate. In Algeria, its presenceis limited to sandy soils.

(1) and (2): Ophioglossum lusitanicum profits from temporary run-nels resulting from autumn rains, which are generally more signi-ficant than spring rains, whereas the cumulative effect of thesupply of water in the depressions seems better adapted to thedemands of O. azoricum. On a typical profile through a tempo-rary pool on mainland France (Maures), O. lusitanicum will thusbe situated higher than O. azoricum. In Corsica, the two specieshave very different distributions: O. lusitanicum is found at lowaltitude, while O. azoricum is only present in some higher-altitudepozzines.

Directive Habitat “Mediterranean Temporary Pools” (code 3170).



Phytosociology*

Mediterranean regions• In a European list, Brullo & Minissale54 include Ophioglossumlusitanicum among the characteristic species of the allianceCicendio-Solenopsion laurentiae (within the order Isoetalia, classIsoeto-Nanojuncetea). In Spain, Rivas Goday319 has described anOphioglosso-Cicendietum filiformis association.• In crystalline Provence, Loisel215 described O. lusitanicum ascharacteristic of the association Isoeto-Nasturtietum.• In Languedoc, in the Avants-Monts, O. azoricum is noted as aparticipant in a hygrophytic variant of a Scillo-Ranunculetum palu-dosii98. In the Cévennes, O. azoricum forms part of Isoetion com-munities with Isoetes duriei which have not been described44, 195. • In Roussillon, O. azoricum is associated with various undescribedIsoetion communities212. O. lusitanicum is associated in tempo-rary runnels with I. duriei272.

Other regions• In western France, Ophioglossum azoricum is part of a north-Atlantic Ophioglosso azorici-Agrostietum caninae or a continentalthermo-Atlantic Ophioglosso azorici-Isoetetum histricis whereasO. lusitanicum is one of the characteristic species of a coastalthermo-Atlantic Ophioglosso lusitanici-Isoetetum histricis98. Inthe Natura 2000* Habitats Register, this latter community isdivided into two: Romuleo columnae-Isoetetum histricis and a Cha-maemelo nobilis-Isoetetum histricis153.

Ecology


Life formPerennial species persisting in the summer through a rhizome(geophyte* with rhizome).

Adaptive strategy163

Stress-tolerant (of summer drought).

ReproductionProduction of spores aerial, fusion of gametes in wet conditions.

Biological cycleGrowth of fronds with flooding (autumn or spring rains accordingto the species), maturation of spores in winter for Ophioglossumlusitanicum in late spring/early summer for O. azoricum; death ofthe leaves after sporulation, which coincides with the summerdrying-out period for O. azoricum. The phenology* depends onthe region. In the Mediterranean region, the cycles can differfrom one year to the next depending on the weather conditions.In Morocco, the biological cycle of O. lusitanicum also begins inautumn (November). It terminates in the spring (May) with thewithering of the fronds.

62



HydrologyGermination of spores requires a very wet soil, supplied regularlyduring this phase with oligotrophic* water. These plants with-stand occasional flooding.

SubstrateAcidic, fairly thin

Interspecific competitionHeliophilous* species, which does not thrive in the shade of tallerspecies.

Impact of perturbationsGenerally favorable, but overgrazing constitutes a mid-termthreat for Moroccan stations.



France• In mainland France, the two species seem very local; the num-ber of stations is not exactly known.• In Corsica, Ophioglossum lusitanicum is fairly frequent on thecoast, O. azoricum is local.

MoroccoTen or so stations* with O. lusitanicum have been censused, andthis number could increase with a more systematic explorationof the more inaccessible regions of the Middle Atlas and the Rif.

SpainO. 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 provincesin the centre and northeast64.

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

Conservation status

(1) O. azoricumThe number of stations is noticeably increasing in France, nodoubt 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 apriori vulnerable. The species has not been found again at Roque-Haute (Hérault).

(2) O. lusitanicumThe number of stations is in noticeable decline in France, partic-ularly in the Atlantic zone. In Morocco, the number of stations isfairly stable, although the populations are slightly increasing atthe mountainous stations. In Algeria, the species is considered tobe rare.


O. azoricum

O. lusitanicum



Withering of fronds


France



Withering of fronds

Germination andvegetative growth

France



---

- (1) France, national list:Decree of 20/01/1982/JournalOfficiel 13/05/1982, amendedby Decree of 31/08/1995/Journal Officiel 17/10/1995- (2) France, regional lists: • Languedoc Roussillon: Decreeof 29/10/1997/Journal Officiel16/10/1998• PACA: Decree of 9/05/1994/Journal Officiel 26/07/1994 - (1) France, national Red DataBook: priority species274


Anthropogenic factors The very specific requirements of the Adder’s-Tongues, in bothtrophic and hydrological terms, mean that these plants arespecies which are very sensitive to modifications in the balanceof their environment. The localised nature of the stations accen-tuates their vulnerability. Modification in the hydrology of thesites is a frequent degradation factor. As the habitats occupiedby these species are generally not very fertile for agriculture, themost immediate threat remains the development of sites, in par-ticular for all the coastal stations. The Adder’s-Tongues are oftenestablished on thin soils which are sensitive to erosion. Excessivenumbers of visitors can contribute to degradation at some sta-tions.

Natural factorsThe closing-up of the habitat by vegetation can be a limiting fac-tor for populations. However, the sites being generally not veryproductive (skeletal soils), the growth of the competitive vegeta-tion is slow: in particular, populations of Adder’s-Tongues on rockyshelves are in a virtually climax* situation and not vulnerable tothis factor.In addition, the competitive vegetation is often sensitive to per-turbations (grazing, scrub clearing, etc.).

63

Plant species


Current measures

France• The Conservatoire Botanique National Méditerranéen de Porque-rolles is conserving O. lusitanicum spores ex-situ.• In the Plaine des Maures: - Some areas supporting these species belong to the CEEP or theConservatoire 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 sitessupporting these species (Péguière pool, Les Aurèdes site) and asite has been acquired in the Bois de Rouquan by the CEEP.- The grazing management and scrub clearance implemented forthe Défense Forestière Contre les Incendies can be favourable forthese species.- Several stations are included in the Natura 2000* area.

MoroccoThere are no measures for the management of Adder’s-Tonguesites.

Recommendations• Management of the sites supporting Adder’s-Tongue popula-tions, through statutory measures guaranteeing the maintenanceof ecological conditions favourable to their survival, is the bestway of conserving the stations. In particular, the implementationof decrees for biotope protection, notably for certain coastalpopulations, is recommended.

• A more exact census of the stations of these inconspicuousplants is also recommended. In most favourable natural habitats,their presence is undoubtedly underestimated. Inclusion of thesites in various inventories (ZNIEFF*, Natura 2000, etc.) is essen-tial.• From time to time, management measures aiming to maintainan open habitat or re-establish a satisfactory hydrological regimecan 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

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

64


Pilularia minuta Durieu ex A. Braun

POLYPODIOPHYTES300

MARSILEACEAE

Main synonymsP. globulifera L. subsp. minuta (A. Braun) Bonnier & Layens

French name: Pilulaire délicateItalian name: Pilularia minoreEnglish name: Pillwort (name of genus)

SubspeciesNone


• Amphibious plant, perennating by means of its filiform creep-ing rhizome, just slightly buried under the surface of the ground.• Fronds (“leaves”) green, filiform, cylindrical, upright from 1 to3 cm in length and arranged singly along the rhizome, the intern-odes being around 1 cm. • Sporocarps* (“fruits”) globular, from around 0.75 to 1 mm indiameter, densely downy, with short pedicels (the pedicel isrecurved at the apex) and dark brown in maturity.

Similar species Seedlings of rushes, grasses and Isoetes. The distinctive charac-ters of Pilularia minuta are the filiform rhizome very slightly buriedin the soil, the globular sporocarps and the filiform leaves.


Distribution/range

Mediterranean species.

FranceHérault, Alpes-Maritimes and Corsica.

Other Mediterranean countriesPortugal, Spain (Minorca and probably in Andalusia andCatalonia)64, Italy (Sardinia, Sicily, Rome region), North Africa(Morocco and Algeria), Greece, Cyprus and Turkey.

Habitat

General descriptionPools with Pilularia minuta are mainly (mainland France, Corsica,Morocco), confined to low-altitude forests or matorrals* domi-nated, 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 situatedin degraded vegetation dominated by Woody Fleabane (Dittrichiaviscosa). The soil there is sandy to sandy-silty and the bedrocksandstone, quartzite, basalt or schist. In Morocco, floooding usu-ally occurs in December-January, solely by rainwater, and dryingout in April-May. At Roque-Haute (France) and in Corsica, floodingbegins in autumn (November) and ends between April and July,depending on the pools.



22.341 Short Mediterranean amphibious swards (Isoetion).22.3412 Aquatic Mediterranean Quillwort swards.

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

Phytosociology*• Traditional phytosociological system319: communities withPilularia minuta belonging to the alliance Isoetion Br.-Bl. 1931 ofthe order Isoetalia Br.-Bl. 1931 em. Rivas-Goday 1970 and theclass Isoeto-Nanojuncetea Br.-Bl. & R. Tx. 1943. Braun-Blanquet48 considers P. minuta to be one of the charac-teristic species of his Isoetetum setaceae or Isoetes setacea-Lythrum borysthenicum association (= Peplis hispidula) of Roque-Haute (Hérault).

• More recent phytosociological system98: communities with P. minuta belong to the Antinorio agrostideae-Isoetion velataealliance (Br.-Bl. 1931) of de Foucault 1988, of the order Isoetaliavelatae (Br.-Bl. 1931) of de Foucault 1988 and the class Isoeteavelatae (Br.-Bl. & R. Tx. 1943) of de Foucault 1988.

Ecology


Life formPerennial species: rhizomatous geophyte* or radicanthydrophyte*292.

Pilularia minuta

0.5 cm

65

Plant species

ReproductionAerial for the maturation of the spores.

Biological cycleGermination of spores in very damp or flooded conditions andgrowth of the fronds in the water in March-April, followed by theformation of sporocarps. In late spring, maturation of the sporo-carps and the spores after complete drying out; withering andrapid death of the fronds.

• In Greece, this species occurs in three coastal pools situatedon islands in the western and northern Aegean Sea290.• In mainland France, there are a few stations at the pools of theRoque-Haute Nature Reserve and the Béziers plateau (Hérault)45, 300.• In Corsica, around a dozen of stations have been confirmed orrecently 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

Populations vary greatly in size from one year to the next, butappear to be surviving at the various stations.• In mainland France it is, however, possible that P. minuta isvulnerable as it is at the northwestern limit of its range. Thespecies has, furthermore, disappeared from the pool at Biot,destroyed in 1975246. • In Algeria, the known stations in the Algiers area and nearOran97 appear to be threatened302, but recent information islacking.



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

Natural factorsThe closing-up of the habitat, following the growth of tall vege-tation, has a negative effect upon this very small species.In France, at Roque-Haute, the abandonment of grazing resultsin the colonisation of pools by woody species and helophytes.They impede the growth of this species, both directly by theirshade, and directly or indirectly through their litter (mechanicaland chemical actions). In Morocco, it is mainly the frequency of dry years which limitsthe appearance each year of this species314.



Morocco

Withering of fronds




France

Withering of fronds



HydrologyGermination of the spores requires the saturation or flooding ofthe substrate. The plant can remain in the vegetative stage underoligotrophic, very shallow water (<5 cm) and form sporocarps,but the complete drying-out of the habitat is essential for thematuration of the sporocarps96.

SubstrateNon-calcareous substrate at the Moroccan and French stations.

Interspecific competitionA heliophilous* species, Pilularia minuta germinates in the clear-est parts of the pool. The species is negatively affected by theshade of woody species, herbaceous perennials (Cistus monspe-liensis, C. salviifolius, Myrtus communis, Dittrichia viscosa, Ulmusminor, Fraxinus angustifolius, etc.) and helophytes (Scirpus mar-itimus, Eleocharis palustris, etc.), as well as by mats of charo-phytes (such as Tolypella glomerata in Corsica) in years with avery rainy winter (as in 2000-2001).

Impact of perturbationsThese are favourable (movements of livestock, wild boar, etc.) asthey reduce the biomass of larger plants and expose the soil.

Adaptive strategyAccording to Grime163, the species seems to have a stress-tolerantruderal (S-R) survival strategy



• 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).




-- Strictly protected species(Annexe I)

- - France, national list: Decree of20/01/1982/ Journal Officiel13/05/1982, amended by Decreeof 31/08/1995/Journal Officiel17/10/1995 - 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

66


Risks relating to populationsRisk of extinction of populations of Pilularia minuta in mainlandFrance, due to their reduced numbers and their isolation.


Current measures

MoroccoNo current management measures.

France• Mainland France: at Roque-Haute, on certain sites with Pilu-laria minuta, the clearing of woody species and the removal oflitter to encourage Isoetes setacea were tested within the frame-work of the LIFE “Temporary Pools” project. However, the conser-vation of this population, situated on a Nature Reserve, is nowuncertain (ban on access, and unilateral management by owners).• Corsica: the creation in December 2000 of the Tre Padule deSuartone Nature Reserve (Bonifacio) should allow managementmeasures to be put in place which are favourable to this speciesin the four pools of the Reserve.• Collection and conservation ex-situ of the spores by the Con-servatoire 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 thehabitat to a certain extent. To attain this objective, the extensivegrazing of cattle should be be maintained or encouraged.

• Promote legal conservation measures for stations which arenot yet protected.• Implement the management plan at Roque-Haute, and moregenerally restore concerted management with natural-heritageobjectives on this Nature Reserve.

Bibliography

Anonymous, 199912 ; Anonymous, 200013 ; Bissardon & Guibal,199739 ; Boudrié et al. 199845 ; Braun-Blanquet, 193548 ; Castro-viejo, 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 ; Maire1952-1987230 ; 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

67

Plant species

ANGIOSPERMS

RANUNCULACEAE

Main synonyms(1) R. nodiflorus L. subsp. lateriflorus (DC.) P. Fourn.(2) R. nodiflorus L. subsp. nodiflorusR. lateriflorus DC. var. charbonelii Rouy & Foucaud

French names: (1) Renoncule à fleurs latérales(2) Renoncule à fleurs nodalesItalian name: (1) Ranuncolo a fiori sessiliSpanish name: (1) Ranunculo con flores laterales

SubspeciesThese two taxa are sometimes considered to be two subspeciesof Ranunculus nodiflorus.

Description/identification criteria • Annual herbaceous plants (5-20 cm) pale green with slender,fasciculate roots. • Species with erect fistulous* cylindrical stems, sometimes rad-icant at the lower nodes, dichotomously branching.• Basal leaves petiolate with elliptical or lanceolate-oval blades,sometimes floating and in that case with long petioles; middleand upper leaves with progressively shorter petioles, entire (orslightly 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 Ranun-culus lateriflorus, smaller (1.5 to 2.5 mm) in R. nodiflorus, soli-tary, sessile, (lower sometimes with short pedicels), and locatedin the forks of the stem or between two leaves on the unbran-ched parts of the main stem. Five pale yellow petals approxi-mately equal to the sepals.• Achenes not deciduous, with surfaces covered in tubercles,very compressed, 1.5 to 2 mm excluding the beak, which approx-imately equals the main body of the achene in R. lateriflorus andis 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, givenby some authors: in R. lateriflorus the petals are spoon-shaped,in R. nodiflorus they are flat (however, this criterion is difficult toapply objectively to very small petals which are quickly shed,especially as it is variable on a single flower); also, more over ofequal size, R. lateriflorus may be more robust than R. nodiflorus.

Similar species Other annual Ranunculus species with entire leaves and small totiny yellow flowers may grow in identical habitats (R. revelieri, R.longipes, R. ophioglossifolius). However, in reasonably well-developed individuals there can be no confusion with the twoabove species, which are the only ones with sessile flowers in theleaf axils and in the forks of the stem branches. The other specieshave pedunculate flowers.

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

Ranunculus lateriflorus

1 cm

1 cm

Ranunculus nodiflorus

68



Distribution/range

(1) Ranunculus lateriflorusPaleo-tropical species with a wide but everywhere patchy distri-bution, around the Mediterranean and east to central Asia andSiberia: 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 Regardin the south of the basaltic plateau of the Coiron (Ardèche).In Italy, three currently known localities: in Sicily, in the Apen-nines, and in the Abruzzo86.

(2) Ranunculus nodiflorusFranco-Iberian endemic.In France, the stations are scattered especially through the centreand west of France: Massif des Fenouillèdes (Pyrénées-Orientales),Massif Central in the southern Aveyron and on the basaltic pla-teaux (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 thecentre and the north.

Habitat

General descriptionIn France: non-calcareous areas that are temporarily flooded inwinter and dry out in spring (pools, ditches or nutrient-poor grass-land).

(1) Ranunculus lateriflorusIt grows in fairly deep pools and ditches, even in poor grasslandsprone to flooding. At latitudes further south than France, it isfound 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-siliceoussubstrate218.

(2) Ranunculus nodiflorusOutside the Mediterranean area, it shows a preference for flat-bottomed basins, very shallow, covered in a clay-gravely silt ofmaximum depth 3 to 4 cm and scattered around on plateauxwhere there is much exposed bare rock47, 106. In the rest of itsrange, this species is found in similar habitats to R. lateriflorus.In Corsica, the plant is rare141 and known only from the moun-tains in the south of the island, from about 900 to 1,600 m alti-tude: • In the pozzines* of the Massif de l’Incudine, in the Plateau duCoscione99, 143, 144.• A stream which dries up from June onwards in the Forêt deMarghèse in the Massif de l’Ospedale-Cagna107.

Several of the pools where these Ranunculus grow have partlyanthropogenic origins: granite quarries in the Pays Bigouden80,basalt quarries at Roque-Haute, stock watering ponds in theArdèche234.

CORINE BiotopeOutside the Mediterranean region (22-11 to 22-13) x 22-32 Oligotrophic to mesotrophic annualcommunities, topographically low-lying, of plain levels, withAtlantic affinities, of the class Isoeto-Juncetea.

Mediterranean region 22.34 Southern amphibious communities (Isoetalia).


Habitats Directive

Outside the Mediterranean region Community of EC interest “Oligotrophic to mesotrophic annualcommunities, acidophile, of medium plain to montane levels ataltitudes ranging from plains to mountainous, of the Isoeto-Juncetea” (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-Dama-sonium polyspermum community98. • Sicily: Ranunculo-Antinorietum insularis and Myosuro-Ranunculetum 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-Ranunculetumnodiflori106 (= Crassulo vaillantii-Ranunculetum nodiflori Abbayes1946) and Ranunculus lateriflorus var. charbonellii (= R. nodiflorus)-Sedum villosum community38.• Mediterranean France: undescribed communities, notably withIsoetes setacea, Les Fenouillèdes.• Corsica: Gamisans144, based on three releves from the pozzinesin the Plateau du Coscione, described the Ranunculeto-Juncetumbulbosi association, which should be renamed Ranunculo nodiflori-Juncetum bulbosi.

Companion species

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

69

Plant species

• High Atlas (Morocco): Sedum nevadense, Elatine macropoda,Ranunculus batrachioides, etc.• Iberian Peninsula: Sedum nevadense, Pulicaria paludosa, Ille-cebrum 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.

Ecology


Life formAnnuals (therophytes*).

Biological cycleIn France, both these Ranunculus germinate during the winter toproduce floating leaves. Flowering takes place in spring when thewater level is falling (April-June). The flowers quickly producefruits: flowering and fruiting are almost simultaneous. The plantsdie and wither from the beginning of summer.R. nodiflorus is preferentially autogamous and its achenes arevery buoyant196. This is probably also true of R. lateriflorus whichhas the same floral characteristics (very small petals, flowerswithout scent) and similar-sized achenes. The seed bank* ofcourse plays a vital role in both species196.

Ranunculus lateriflorus and R. nodiflorus

Interspecific competitionVery sensitive to competition from perennial species. In Brittanyfor example, simply root-stripping of the pool vegetationresulted in an increase from 47 R. nodiflorus plants in 1995 to3000 the following year at a single site80.



(1) Ranunculus nodiflorusIn France, this species occurs in the mountains of southern Cor-sica and a large population is known from Rodès in Les Fenouil-lèdes. At Montalba-le-Château, a station near the above and known inthe 19th century (Castanier in Herbier Oliver, Montpellier Uni-versity), has not been seen since. There are old reports from thePyrénées-Atlantique and the Hautes-Alpes. It has been mistakenlyreported from Savoie, the Alpes-Maritimes and the Var. Overall, in France, the species is declining due to the disappear-ance of pools (urban development) and the growth of vegetationfollowing the cessation of grazing274.

(2) Ranunculus lateriflorusThere are three remaining localities in France: Roque-Haute, Caux-Fontès-Pézenas and Coiron. It disappeared from the CostièreNîmoise in the late 1960s or early 1970s following land ownershipreorganisation which led to the filling-in of many of the small pools(laquets) which were scattered across this plateau (formerly at theLaquet de l’Oli at Beauvoisin, on Quaternary siliceous gravels).The populations of Aosta and Calabria (Italy) have not been seenrecently86.



Anthropogenic factors The depression with Ranunculus lateriflorus at Costière (France,Gard) was filled in during agricultural developments in the1960s-1970s. The population at the Plateau de Caux-Fontès-Pézenas survives only in a system of canals used to drain a naturaldepression that has been taken into cultivation. The R. nodifloruspopulation at Les Fenouillèdes is threatened by a quarrying pro-posal.


Growth

Flowering

Germination

Fruiting

Southern France


HydrologyTypically amphibious species that require a fairly long period ofwinter flooding in the spring, extending into March-April for Ranunculus nodiflorus in Brittany106 and to April-May for R. lat-eriflorus in Mediterranean France. In a relatively wet climate(Massif Armoricain, Paris Basin) the basins occupied by R. nodi-florus are shallow, allowing rapid warming in spring followed byearly drying out (des Abbayes, 1947; Bournérias et al., 2001). In aMediterranean climate, the habitats occupied by R. nodiflorus orR. lateriflorus may be under several tens of centimetres of waterand may dry out later.

SubstrateThese are both species of oligotrophic* pools, which attain theiroptimum on acid mineral substrates: sandy-gravely soil (pH 5.2-6.5) on the Roque-Haute basalts49, gravely-clayey silts (pH 5.6-5.8), very thin (optimum 1-2 cm), lying directly over rock in theMassif Armoricain106, granitic sands at Rodès (Fenouillèdes).



---

- (1) and (2) France, national list:Decree of 20/01/1982/Journal Officiel 13/05/1982,amended by Decree of31/08/1995/ Journal Officiel17/10/1995 - (1) and (2) France, national RedData Book: priority species274

- (1) Morocco: “rare” (R) 133, 134, 181

- (1) Italy, National Red Data Book:“vulnerable”86

70


Natural factors The natural succession of perennial formations suppresses thedevelopment of these Ranunculus; however, they appear to becapable of surviving in clearings, even very small ones, within densevegetation. In addition, the seed bank is of considerable size.

Risks relating to populationsLow risk of local extinction due to the size and longevity* of theseed bank.


Current measures

In FranceOnly the populations of Ranunculus lateriflorus at Roque-Hauteare located within a nature reserve which benefits from a man-agement plan. However, the management plan cannot put intopractice at this reserve owing to a dispute with the owners.The Conservatoire Botanique National Méditerranéen de Porque-rolles has made collections of R. nodiflorus seeds and is preservingthem ex situ.

In ItalyNo measures

RecommendationsWork towards the acquisition of land at the richest sites (Fenouil-lèdes, etc.) as well as conservation management by a competentorganisation.

Bibliography

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 ; Gami-sans, 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.

71

Plant species

Ranunculus revelieri Boreau

ANGIOSPERMS

RANUNCULACEAE

Main synonymsR. 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)

SubspeciesRanunculus 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 andare not accepted by all botanists141, 109.

Description/identification criteria • Small plant, 10 to 20-30 cm tall, with the stem upright, fistu-lous* (rarely in rodiei), branched and more or less hollow.• Basal leaves surrounding the stem, with long petioles and ovallanceolate blades. Stem leaves with shorter petioles and morenarrowly lanceolate.• Flowers in a pale yellow perianth, with long pedicels. Petalsglabrous, shorter than the sepals (very villous on the lower sur-face in revelieri and glabrous or slightly downy in rodiei).• Fruits: ovoid achenes, blunt, finely grainy, with a very shortbeak*96, 274, 386.

Similar species Ranunculus fontanus C. Presl and R. ophioglossifolius (Vill.)Boreau have the petals longer than the sepals96. R. ophioglossi-folius 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 givenrise to R. revelieri which is tetraploid (2n = 32). R. revelieri wouldthus be an apoendemic*, i.e. a polyploid* taxon no doubt derived,through evolution, from the diploid taxon R. ophioglossifolius85.


Distribution/range

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

France Ranunculus revelieri rodiei in the Var (Maures, Estérel) andR. revelieri revelieri in Corsica.

Other Mediterranean countriesR. revelieri revelieri in Sardinia (Italy)16, 85, 96, 292.

Habitat

General description In Les Maures, Ranunculus r. rodiei is found in the shallow partsof pools, but also in ditches and on the edges of temporaryrivulets. In the Plaine de Palayson, this species occurs in clay-siltpools situated in sandy depressions resulting from the erosion ofPermian sandstones246 (R. r. rod.).In Corsica, the R. r. revelieri stations are mainly on the edges oftemporary pools, as well as depressions and wet meadows16, 85, 108,

109, 222, 285, 292.




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

Ranunculus revelieri

1 cm

72


• In Corsica, R. revelieri is frequently located among the follow-ing perennials: Schoenus nigricans, Plantago lanceolata var. tim-bali, Oenanthe globulosa, Cynodon dactylon and Carex serrulata.Associated annuals are: Juncus pygmaeus, J. tenageia, Solenopsislaurentia, Cicendia filiformis, Lythrum hyssopifolium, Illecebrumverticillatum (non-submerged form) and Agrostis pourretii. Thistype of community should not be included in the Isoetion histri-cis but in the Cicendion.

Ecology


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

ReproductionAerial.

Seed characteristics96, 386

Length: about 1.5 mm; width: about 1 mm.

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

Interspecific competitionFor this heliophilous* species, according to Barbero21, competi-tion 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 detrimen-tal to this species, in particular on soils whose depth is greaterthan 20 cm.

Impact of perturbationsThere are probably few direct effects on populations (uprootingof individuals), but a positive indirect effect of irregular distur-bance of the substrate (stream dynamics, wild boar, etc) whichrestricts perennial species or improves the hydrological regime.



In France• 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 inParadis et al.285.

In ItalySeveral stations in northern Sardinia, in the Limbara mountains,near Macomer, at Molara, and on San Pietro island west ofCagliari (map in Arrigoni16, Conti et al.86).

Conservation status

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



Vegetative growth

Flowering

Germination Germination

Fruiting

France


HydrologyA hygrophytic* species found in shallow water (0-20 cm) butwhich can survive, in exceptional winter conditions, at depths ofup to 40 cm. Germination takes place under water or on wet soiland the plant can tolerate prolonged immersion of the substratefor several months per year, from mid-autumn to the beginningof spring153. Climatic instability causes interannual fluctuationsin the abundance of individuals. Ranunculus revelieri can alsogrow in slow-flowing streams and rivulets18, 153.

SubstrateIt usually grows on siliceous soils.

IUCN 1997399

Bern ConventionHabitats DirectiveNational and regional protection


- 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 of20/01/1982/Journal Officiel13/05/1982, amended by Decreeof 31/08/1995/Journal Officiel17/10/1995 - France, national Red DataBook: priority species274

- Italy, national Red Data Book:“threatened”86

73

Plant species


Anthropogenic factors “Cleaning up”, drainage or infilling of the habitat, urban devel-opment96, 274. In Provence, several stations have been destroyeddirectly or indirectly (disruption of hydrology) by tourism develop-ments (golf), transport infrastructure works and urban deve-lopment.

Natural factorsNatural vegetational succession may lead to the habitat closingup and to processes of aggradation by siltation. For example,biotopes suitable for the Lythrum borysthenicum and Ranunculusrevelieri association at the Péguière pool (Plaine des Maures) areprogressively changing into drier habitats, which are prone tocolonisation by other plant associations153, as a result of infillingwith sediments from the catchment area. More generally in LesMaures, the disappearance of grazing favours the developmentof 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, theactivities 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 populationsAt Ranunculus r. rodiei stations, the populations are not visiblevery year as their development depends closely on the pattern ofspring rainfall. Similarly, in Corsica, populations of R. r. revelieriare only visible in years when the winter-spring rainfall is suffi-cient to enable germination to take place (as was the case in2000-2001). However, their instability does not imperil the pop-ulations.


Current measures

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

Mainland France (Maures)• A Nature Reserve project at the centre of the plain should pro-tect 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 neces-sary requirements for the conservation of this species to be takeninto account. Currently, the grazing which has been reintroducedalong the firebreaks is favourable for it.• LIFE “Temporary Pools” has enabled experimental habitat man-agement methods to be tested for this species, as well as theacquisition by the CEEP of two sites where it is present (Bois deRouquan, Vallon de Sauronne).• Seeds have been collected and are preserved ex-situ by theConservatoire Botanique National Méditerranéen dePorquerolles.

Recommendations

In Corsica Managers should ensure that ground disturbance, due for exam-ple 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 oftaking the species into consideration during physical planning.- Control over management by acquisition or by contractualmeans.• 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

74


Genus Riccia L.

MARCHANTIOPSIDA

RICCIACEAE

Main synonymsNone still in use nowadays.

French name: RicciaEnglish names: Crystalwort, Riccia

Sous-genresTwo sub-genera occur in the Mediterranean area: - Riccia with a single section, Riccia.- Ricciella (A. Braun) Rchb. with two sections: Ricciella andSpongodes Nees.

Description/identification criteria • Small liverworts with gregarious thalli, forming either completerosettes or more or less connected linear lobes, from a few mil-limetres to a few centimetres in diameter, with a tough, compactor spongy consistency.• May be annual or perennial* species.• Lobes usually having the dorsal surface longitudinally groovedand swollen at the edges into more or less prominent tumidridges, varying widely in colour depending on the species and thegrowing conditions, from pure white to blackish-violet. Edges ofthe lobes sometimes with hairs, cilia or papillae and with scaleswhich often have a crimson tint.• In species with a compact thallus (subgenus Riccia), dorsal tis-sue consisting of uniseriate columns of almost contiguous verti-cal cells; in species with lacunose thalli (subgenus Riciella) dorsaltissue 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 tissueof the thallus, more or less projecting on the upper surface in theform of small upright protuberances.• Female gametangia (archegonia) also contained within thethallus and more or less protruding on the upper surface. Oncefertilisation has taken place, capsule development within thethallus tissue; at maturity, the capsule may project from theupper 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* amongthe spores. Spores coloured at maturity, turning from brown toblack and very often exhibiting polarity (the distal surface, whichis roughly hemispherical, contrasting with the proximal surfacewhich has three triangular facets). Distal surface (less often theproximal surface) usually provided with alveoli demarcated bywalls which may have tubercles in their angles (this spore orna-mentation is often very important for identifying species, henceit is always worthwhile to look for mature thalli in the field).- Vegetative* reproduction very rare.

Introduction to a few speciesThe genus Riccia includes over thirty species in the Mediterraneanarea, making it the most widely represented liverwort genus. Thespecies are often fairly difficult to identify in the field, althoughsome of them are easy to recognise with a little experience. Theyare not all strictly dependent on temporary pool biotopes but may

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


Distribution/range

FranceThe genus Riccia is particularly well represented in theMediterranean Midi but many species have a wider distribution.Various species with Mediterranean affinities are found alongthe Atlantic seaboard (Riccia crozalsii, R. michelii, etc.) or in theMassif Central (R. trichocarpa). Many taxa are rare and onlyoccur at a very limited number of stations. For example, theMassif des Maures, the Biot area and the Nature Reserve de Roque-Haute are sites that are extremely rich in liverworts of the genusRiccia.

Other Mediterranean countriesAll the countries around the Mediterranean are rich in Riccia,although many of them have a very uneven distribution at thisscale. For example, Riccia macrocarpa is much commoner in thewest of the Mediterranean Basin than in the east; R. sommieri isonly found in west of Mediterranean Basin; R. atromarginata ismuch less rarer in North Africa than in the north of the Medi-terranean Basin, etc.

Habitat

General descriptionSuitable habitats include, for example, short and pioneer grass-lands with many gaps in the vegetation cover, thoroughly wettedin winter and spring and drying out completely in summer. Thephysical characteristics of the substrate are important. Riccianeed a substrate which is firstly stable and relatively consoli-dated and secondly has a certain capacity to retain water. Sands,silts and clays (or mixed substrates) are the most suitable substratesin this respect.Biotopes with Riccia are often located close to temporary poolsbut not actually at the lowest topographical levels. Generally

Riccia sorocarpa

1 mm

75

Plant species

speaking, it may be said that the Riccia of temporary pools andstreams are linked with associations* of medium topographiclevels. They are concentrated mainly among populations that arekept damp by capillary action or that emerge from the waterearly in the year, for example Isoetes duriei populations. Higher plant species that are usually associated are mostly smallannuals (for example the dwarf Juncus, Radiola linoides, Lau-rentia michelii, Aira capillaris, Crassula vaillantii, Lythrum thymi-folium, Lotus angustissimus, etc.).

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

CORINE BiotopesThese communities with Riccia may be placed in: 22.34 “Southern amphibious communities” (Isoetalia), in particular,


Phytosociology*The relationships between liverworts and the vascular-plantcommunities are often complex. Liverworts and bryophytes* arein general very ephemeral and delicate plants whose populations(and thus the communities which they form) are subject to greatvariability in space and time. The communities are often patchyat the scale of a single station, where their appearance is fre-quently associated with openings in the herb layer. Some authors21, 319 have considered the Riccia to be characteris-tic of the overall Isoetion alliance. Barbéro21 and then Hébrard171

for their part emphasise the closeness of the sociological linkswhich join the communities with liverworts to the vascular-plantcommunities Isoetion, but they also point out that it is very dif-ficult to give an opinion on the true phytosociological affinitiesof the bryophytes. In addition, Hébrard171 mentions cases of veryclear extensions of bryological communities typical of tempo-rarily wet biotopes into Cistus formations, when the soil humidityis sufficiently high.

The Campylopetum introflexi Hébrard171 is an association whichunites the bryophytes of the rocky seepage surfaces of crystallineProvence, in which Riccia are considered to be companion species. The Riccietum crustatae is another circum-Mediterranean asso-ciation, well represented in Spain, characteristic of wet, saltysoils which dry out severely in summer326. However, most Ricciacannot tolerate the presence of salt in the substrate.It may be seen that the phytosociological characterisation ofbryological communities involving Riccia is only in its early stagesand that much descriptive work still needs to be carried out.

Ecology

Riccia are temporarily hygrophytic*, heliophilous*, thermophilicand terricolous species. However, the individual species’ ecologi-cal requirements show wide variations with respect to this gen-eral pattern. There are species for which the chemicalcharacteristics of the substrate are immaterial while others aredistinctly calcicolous or silicicolous.


Life formGenerally hepatico-therophytic. Annual or perennial species. Thespecies considered to be annual may also revive after the dryseason with the resumption of activity in the apical meristem orsometimes the whole thallus. It is possible to observe a resump-tion of activity of Riccia meristems that have been kept in aherbarium for over ten years, which is a good illustration of theirpowers of regeneration!

Adaptive strategy (sensu During126)• Type “a”, corresponding to “annual shuttle”: lifespan less thanone year, spores large and produced in fairly small numbers;major reproductive investment.• More rarely type “s”, corresponding to “short lived shuttlespecies” (maximum of a few years): short-lived perennials withhigh investment in reproduction and few fairly large spores.

ReproductionFertilisation requires a film of water on the surface of the thallior even the substrate; spore maturation takes place during thewet season and may continue into the beginning of the dry sea-son. Spore production is therefore aerial.

SporesThey are usually fairly large (of the order 100 µm) and heavy, andare 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 moreunpredictable. In most cases the spores are ornamented and thiscould facilitate their attachment to vectors, although this hypo-thesis has not been experimentally confirmed. Fragments of thallusor propagules* (very rare) may also be carried by animals.

Biological cycleRiccia are capable of carrying out the whole of their biologicalcycle very rapidly following the appearance of water in the sub-strate (rain, runoff) or merely on the thallus (condensation, noc-turnal dewfall, etc.), enabling them to avoid the driest period.They are therefore able to tolerate the stresses of the driestmonths to a certain degree through the suppression of anyphysiological activity combined with an extreme longitudinalfolding of the thallus, and by the production of spores that havevery tough walls and that retain their germinative capacity for avery 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 thewet season. Gamete production may take place very soon aftergermination (a few weeks for the fastest species), but may alsocontinue until the beginning of the dry season of the following

76


year. The spores are released by the decomposition of the oldparts of the thalli as growth continues at the tips of the lobes.The complete desiccation of the substrate and the thalli causesthe virtually complete disappearance of Riccia and the release ofall the spores produced; the latent phase then begins, with thesurvival in the substrate of the spores, which may enter into adormant state.

Conservation status

The conservation status of Riccia populations is intimately linkedwith the general health of the temporarily wet biotopes. Theseliverworts have undergone a major decline following the disap-pearance of many temporary pool biotopes.In Malta, R. melitensis was known from one locality, where it hasnever been seen again since its discovery344.



HydrologyA very wet substrate is necessary at the beginning of the cyclebut, in most species, flooding must not be too prolonged.

SubstrateMore or less compacted substrates are definitely the most suitable,whether they are sandy, clayey or silty. Terra Rossa*, more or lessmuddy alluvia, etc., also provide potential sites.

Interspecific competitionAll the species are pioneers and struggle to survive in the presenceof competition from other more colonist bryophytes and espe-cially perennial vascular plants, which will eliminate them fairlyreadily.

Impact of perturbationsThe main perturbations (“ploughing” of the substrate by wildboar, trampling by cattle and humans) may have a positive effect,due to the elimination of taller plants, the destruction of the lit-ter from these plants and the opening of the herb layer throughthe creation of bare patches (“tonsures”) which are very suitablefor Riccia.

TemperatureAll the species are thermophiles.

LightAll the species are heliophiles, but very slight shading whichmoderates the intensity of the solar radiation may be a positivefactor in the hottest regions of the Mediterranean Basin.



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


Fertilisation and maturation of spores

Release of spores


Latent spores and thalli


----

- Europe: Red Data Book forbryophytes128: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 forbryoflora (proposed)105:Riccia crustata R. ligula

Further species in this genus are locally rare to very rare, but arenot included on the protected lists as they are little known.


Anthropogenic factors All the threats with which temporary pool and stream biotopesare faced in the area around the Mediterranean consequentlyalso threaten populations of liverworts of the genus Riccia.Urbanisation, infilling with rubble, drainage and conversion toagriculture, etc. lead to the irreversible loss of “Ricciological”natural heritage.

Natural factorsThe absence of any factors that will disturb the vegetation, andthe closing-up of the natural habitat, will eventually cause thedisappearance of pioneer liverworts. The loss of the temporarycharacter of pools, through infilling or through disruption of thehydrological network, is very deleterious to Riccia as this allowsthe spread of common and much more competitive species ofhygrophytic bryophytes (usually pleurocarp* species). However,the presence of dormant spores in the substrate is a positive fac-tor in the re-establishment of populations following a reopeningof the habitat.

77

Plant species

Extinction risks relating to populationsMajor interchanges probably take place between neighbouring orwidely separated sites, but there are no experimental results tosupport this assumption.


Current measures The sites at Roque-Haute and the Massif and Plaine des Mauresare included within Natura 2000* sites (no. Fr 9101430 and no. Fr 9301622 respectively) which should allow particularattention to be paid to the conservation of the temporary poolsand their associated species.The Roque-Haute site also has legally protected “Réserve Natu-relle (Nature Reserve)” status, which is currently unenforced dueto the ban on access imposed on the manager by the owners.

Recommendations• 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).

• Promote the perpetuation of extensive agricultural practiceswhich create moderate disturbance, or management activitieshaving the same rationale. Extensive grazing may be favourableat stations where the vascular vegetation has developed toomuch. Inputs of excess organic matter should, however, be moni-tored.

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.

78


MARCHANTIOPSIDA

RIELLACEAE

Main synonymsDuriaea helicophylla Bory & Mont. (basionyme*)

French name: Riella à thalle hélicoïdeEnglish names: Liverwort, Riella (genus names)

SubspeciesNone

Description/identification criteria • Aromatic aquatic liverwort (scent fairly like coriander, some-times strong), measuring up to 3 cm high but often much smaller,annual, soft-tissued, formed from an upright thallus composed ofa main branch with a more or less undulate single-layer mem-branous wing (a single layer of cells) on one side and small lance-olate scales on the other. Wing more or less helicoid, very fragile,and usually reduced to barely recognisable fragments due to var-ious disruptions and to predation by phytophagous invertebrates.• Thallus fixed into the substrate by many rhizoids* growingfrom the base (plant appearing to be placed gently on the sub-strate).• Dioecious species whose male plants bear very small anthe-ridia*, arranged in a row on the edge of the membranous wing ofthe thallus. • Female plants at maturity bearing bottle-shaped capsules* atthe 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 toslightly granulous, bristling with many large “spines” (6 to 10 µmlong) which are truncated and widen at the tip. No elaters*among the spores.

The sporophyte (capsule), which is normally attached to thegametophyte*, may become quite independent of it (the thallus),when unfavourable conditions (drought, predation, etc.) result inits premature disappearance. The sporophyte then continues todevelop in a completely independent way, which is very unusual,if not unique, among bryophytes*.

The formation of single-layered propagules with indeterminategrowth, arising from the apical meristem, has been described.These propagules, in contrast to spores, do not have a dormantstage and are therefore capable of growing immediately aftertheir release.

Similar species Confusion is possible in the field with all the other species in thegenus. Identification can only be carried out by a thorough exami-nation using appropriate optical equipment. Sterile material ispractically impossible to identify to species. Mature sporophytesmust therefore be looked for in the field amidst the population.In addition, mixtures of species are relatively common.

The species of the subgenus Trabutiella (Riella affinis M. Howe &Underw. and R. cossoniana Trab.) are fairly easily distinguishedfrom R. helicophylla by their longitudinally winged involucre.R. notarisii (Mont.) Mont. is monoecious (often a difficult characterto 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), withthe 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 fromthe taxonomic point of view. Great confusion reigns in thenomenclature regarding various taxa*. A complete and world-wide review of the genus appears to be essential to clarify themany points which remain uncertain.


Distribution/range

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

Riella helicophylla (Bory & Mont.) Mont.

Riella helicophylla

20 µmmale

femalespore

0.4 cm

79

Plant species

• Station by the Etang de Thau (Marseillan), discovered in1966124, and very probably disappeared following the siting of ahuge refuse tip on the exact site.

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

Habitat

General descriptionThe ecological characteristics necessary for the development ofRiella 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 aretherefore of two types: • Temporary pools, lakes, estuaries etc. with saline to super-saline or brackish water (France, Spain, North Africa).• Possibly, temporary freshwater pools (France).

The muddy, sandy or clayey substrate is often base*-rich (pH* rangingfrom about 7 to 8.5).The depth of the water seems to be very important: often veryshallow (a few centimetres) and a maximum of 1 metre.

In Spain, Riella helicophylla has been collected from sites with awide 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 typicallygreater than 10g/l77, 78, 189. Continuous evaporation at some sitesleads to the formation of saline crusts on the surface. R. helico-phylla appears particularly to prefer waters containing sodiumchloride although it also colonises those with magnesium sul-phate. The waters in which it grows are subject to considerablevariations in salinity over a single year and also from one year toanother, depending on rainfall, runoff, evaporation, etc.

Companion species The species most frequently associated with Riella helicophyllaare other species of the genus Riella (in particular R. cossonianaand R. notarisii), aquatic vascular plants (Ruppia spp., Altheniaspp., Potamogeton spp.,) Characeae (Chara galioides, C. canescens,etc., Tolypella spp., Lamprothamnium spp.), rarely bryophytes(Leptodictyum riparium, Drepanocladus aduncus, etc.), GreenAlgae (Ulothrix spp., Enteromorpha spp., etc.), Cyanophyceae(Microcoleus, Lyngbya, etc.) and sulphur bacteria. When the sub-strate becomes completely dried out, the vegetation then changesits appearance, with the frequent arrival of a suite of more or lessxerophytic vascular and bryophyte species. Note that Riella onlyreally disappears when the substrate is completely dry.

Habitats DirectiveThe simplest approach is to include the Riella community undercode 3170, with the overall title “Mediterranean TemporaryPools” for the freshwater communities. On the other hand, thesubhalophilic and halophilic* communities could be includedunder code 1150, with the title “Lagoons”, or even code 1160,

“Large shallow inlets and bays”. Insofar as these codes did notoriginally take the associated bryophyte communities intoaccount, the affinities with one or other suggested code shouldbe explained in detail, so as to avoid any ambiguity or confusion.

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

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

Phytosociology*The Riella helicophylla communities are these days includedwithin the class Rielletea helicophyllae, order Rielletalia helico-phyllae, alliance Riellion helicophyllae, association* Rielletumhelicophyllae78. These vegetation types represent a transitorycover of species that are highly specialised for the colonisationof a very restrictive temporary biotope, at the stage where othercommunities, dominated by various groups of species (Charetea,Potametea, Ruppietea, Isoeto-Nanojuncetea, etc.), are absent orat an embryonic stage of development.

Ecology

It should be borne in mind that Riella helicophylla is a basi-philous*, halotolerant* species, capable of withstanding hyper-saline water; a heliophilous pioneer.


Life formHepatico-hydrotherophyte. Annual species (capable of living formore than one year in culture if it is permanently submerged).

Adaptive strategy (sensu During126)Type “a”, i.e. “annual shuttle” species (= lifespan less than oneyear, spores large and produced in fairly small quantities; majorreproductive investment).

ReproductionMaturation of spores aquatic then aerial; release aerial. Sporesexhibiting dormancy and capable of retaining their germinativecapacity for at least three months.

GerminationIt is at its maximum with low salt concentrations, whereas thethallus grows better at higher salinities.

Dispersal of sporesDispersal by water birds (waders, web-footed species), sometimesmigratory, by external or internal ornithochory*. This mode ofdispersal is assisted by the very large number of spines on thespores. anemochory* appears never to have been described.

Biological cycleGermination of the spores takes place after the first autumn rain,followed by the slow growth of the gametophyte during winterand the beginning of spring. The spores are released in aboutMay and constitute the resistive stage during the summer

80


drought. When the ecological conditions are favourable, it is pos-sible to see veritable population explosions of this species, whichthen excludes other species and attains coverage rates of closeto 100%.



Anthropogenic factors Urban development around towns such as Agde, Tunis or Algiersconstitutes a key threat. The filling in of pools with debris needsto be monitored, particularly at the Notre-Dame de l’Agenouilladesite (Hérault, France). The illegal dumping of rubbish and large-scale official tips are leading to the irreversible disappearance ofthe species.

Natural factorsNatural vegetational succession leads inexorably to the sponta-neous 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) whichare capable of carrying spores from one site to another. Thisreduction in the “accessibility” of some sites is clearly detrimen-tal to the survival of viable metapopulations*. In addition, a highfrequency of dry years may explain the species’ decrease or evenextinction at some of these stations. A minimum of wetness suf-fices to trigger the germination of dormant spores but a suffi-cient amount of water is absolutely necessary to complete thecycle and to produce spores for a new generation. Several dryyears may completely deplete a fairly small stock of spores.

Risks relating to populationsThe risk of extinction is high, given the low numbers at theNotre-Dame de l’Agenouillade station (Hérault, France). The iso-lation of the French populations compared with those of NorthAfrica and Spain, and their limited area, are a further risk factor.


Current measures No specific management measures appear to be taking place atsites supporting Riella helicophylla.In France, the Notre-Dame de l’Agenouillade site (Hérault), isincluded:• In the European LIFE “Temporary Pools” project, which hasenabled proceedings for obtaining control over land ownership tobe undertaken, hydrological monitoring to be carried out andprotocols for monitoring the pools’ vegetation to be drawn up.• In a Natura 2000* area (site Fr 9101416), which should enablethe environmental conditions favourable for the conservation ofthis species to be maintained.


Fertilisation and maturation of spores

Release of spores

vegetative growth

Spores dormant

��

Germination and


Hydrology• The optimal water depth ranges between a few centimetresand a few tens of centimetres.• The survival of the species is closely dependent on the annualflooding/drying regime. A disruption of the hydrological regime,a delay in the arrival of the rains or too low a level of annualrainfall may considerably restrict or even prevent the develop-ment of the species one or several years in succession (temporarydisappearances).• Water quality is important, especially salinity. Riella helico-phylla appears to be not at all obligatorily halotolerant* but,being a very uncompetitive species, it finds in salt water the con-ditions which allow it to exist without or almost without anycompetition.• The species can tolerate a certain degree of eutrophication* ofthe substrate and of the water.

Impact of perturbationsPerturbations due to the weather (rainstorms, frosts, etc.), anthro-pogenic disruption (trampling, ploughing, etc.), or zoogenic dis-turbances (grazing, disturbance of the substrate, etc.) may befavourable to the extent that they restore pioneer surfaces, pro-vided they take place during the dry period (i.e. after the matu-ration of the spores).

Interspecific competitionNatural succession inevitably leads to the disappearance of pio-neer Riella helicophylla communities under increasing pressurefrom vascular plants and algae, whose increasing biomass* andground cover result in greater competition for light and otheressential resources. R. helicophylla does not survive in competi-tion with other plants.



A single site in France; many sites in Spain, distributed among 13provinces79; several sites in North Africa.

Populations appear to be stable or even increasing in Spain andthe Maghreb, but are highly threatened in France due to thepressure of urban development in the Agde area.




-Strictly protected species(Annexe I)Annexe IIMalta: protected species since 1993206

- Europe, Red Data Book forBryophytes128

“endangered” (E)- France, national Red Data Book(proposed)105

81

Plant species

Recommendations• A major survey exercise should be organised in order to locateany possible new population centres, especially in France.• Any management measures which have the aim of maintain-ing or restoring the pioneer character of the pools and their sur-roundings, such as controlled grazing or vegetation clearing, areto be encouraged, especially at sites where these activities havebeen discontinued.• Complete protection of the biotopes (control over land owner-ship, etc.) should be achieved in order to prevent any destructiveurban development projects. • Dumping of rubbish, infilling, and illegal tourism-relatedactivities should be strictly forbidden.

Bibliography

Aboucaya et al., 20022 ; Anonymous, 199912 ; Bissardon & Guibal,199739 ; Cirujano et al., 198877 ; Cirujano et al., 199279 ; Cirujanoet 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.

82


Teucrium aristatum Perez Lara

ANGIOSPERMS

LAMIACEAE

Main synonymsT. cravense Molinier & Tallon

French names: Germandrée de Crau, Germandrée aristéeEnglish name: Germander (genus name)

SubspeciesNone

Description/identification criteria • Small annual plant 10 to 20 cm, stem slender taproot.• Stem square in cross-section, upright, usually branchingalmost from the base; branches erect-spreading.• Leaves pinnatilobed*, with lobes broad, short and blunt, shedfrom the time of flowering.• Flowers geminate, with short peduncles. Calyx lobes erect,spreading, terminated by a beard. Corolla villous externally, laven-der blue with a dark-blue V-shaped mark on the inner base of themiddle anterior lobe; posterior lobes streaked with crimson inter-nally.• Fruits: each flower produces four achenes after fertilisation.

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


Distribution/range

Endemic species of Iberia and Provence.

FranceCrau d’Arles in the Bouches-du-Rhône.

Other Mediterranean countriesOnly in Spain, where it was formerly known (more than 100 yearsago) in three provinces in the southwest (Huelva, Cadiz andSeville)177 (Medina, pers. com.), and where it was rediscovered in2002 in the centre of the country (Morales, pers. com.).

Habitat

General description In France, this Germander occurs on the edges of a clayey basincreated in the Crau pudding stone, whose water is slightly cal-careous252.


CORINE Biotopes22.34 Southern amphibious communities (Isoetalia)

Phytosociology*In the Crau the vegetation of the Lanau pool is allied to a spe-cific association*215, 252: Order Nanocyperetalia

Alliance Lythrion tribracteatiAssociation: Lythrum tribracteatum-Teucrium arista-tum215, 252

At this site, the species also forms part of the Deschampsionmediae alliance.

Companion species 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


Life formTherophyte* (annual species).

Teucrium aristatum

1 cm

83

Plant species

ReproductionAerial: the floral organs as well as the fruits develop out of water.

SeedsAchenes oboval-obovate (2 mm x 1.1 mm), glandular, rounded,and with dense very short curved hairs at the tips.

Biological cycleGermination of seeds after the water has receded, on very wetsoil (in May in France). Flowering occurs at the end of spring(June) on a dry substrate, followed by the ripening of the fruitsand the dispersal of seeds during summer.

SpainThe species is very rare here, with a very limited range (in theprovinces of Cadiz, Seville and Huelva), but a few new stationswere discovered in 2002 in the centre of the country.



Anthropogenic factors The acquisition by the CEEP of the site supporting the onlyFrench population has enabled the pool to be saved from possi-ble threats of habitat destruction, and management favourableto this species to be introduced.

Natural factorsThis species is very sensitive to competition with perennial speciesthat produce dense cover (Carex divisa, Agrostis stolonifera, etc.).Towards the end of the 1990s, it had disappeared from a part ofthe Lanau pool where the cessation of grazing had facilitated amassive growth of perennial grasses, whereas it always main-tained itself in the grazed part of the pool.

Risks relating to populationsThe unique nature of the population (very isolated) entails a highpotential risk of extinction at the site.


Current measures

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


Flowering


Dispersal of seeds

? ?Fruiting and ripening of fruits

France


HydrologyThis Germander is above all a species of the edges of pools, whichneeds a brief period of submersion to develop. It is probable thatthe saturation of the substrate with water at a given period (endof April or May) triggers germination.

SubstrateIn France, silt-clay soils, poor in calcium and oligotrophic*.

Interspecific competitionVery sensitive to competition with perennial herbaceous plants.

Impact of perturbationsDisturbance (grazing, trampling) is favourable to this heliophilous*species since it controls the cover from perennial grasses. At thehigher parts of the Lanau pool, the Germander is most often seenon patches of bare soil: roosting places, cattle tracks, holes madeby hoofs, and former ditches, where it forms small dense popu-lations.

Conservation-Management


FranceA single site is known, Lanau pool in the Crau (Bouches-du-Rhône). Although it undergoes wide interannual fluctuations,this population has been seen regularly since its discovery in1946, and is not in danger at present. The temporary reductionof grazing at this site seems to have noticeably reduced the sizeof the visible population, which is in the process of re-establishingitself.

IUCN 1997

Bern ConventionHabitats DirectiveNational and regional protection


World: “rare” (R)- France: “endangered” (E)- Spain: “rare” (R)

- -

- France, national list: Decreeof 20/01/1982/Journal Officiel13/05/1982, amended byDecree of 31/08/1995/Journal Officiel 17/10/1995 - France, national Red DataBook: priority species274

84


SpainThe species and the sites which support it are not subject to anyspecific measures for protection or management (Medina, pers.com.)

Recommendations

France• Study the hydrological regime of the site and prevent any dis-ruption of it.• Maintain grazing (formerly sheep, nowadays cattle) which isfavourable to the development of annual species like Teucriumaristatum.• Study the population biology of T. aristatum, in particular thekey spring recruitment phase (ecology of germination) and theeffects of natural and anthropogenic perturbations.

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

85

Macrocrustaceans

Imnadia yeyetta Hertzog, 1935

Class: BRANCHIOPODS

Order: SPINICAUDATA

Family: LIMNADIIDAE

Main synonymsI. banatic Marincek & Petrov, 1982I. cristata Marincek, 1972I. panonica Marincek & Petrov, 1984I. voitestii Botnariuc & Orghidan, 194143

French name: noneEnglish name: Clam Shrimp (general name for Spinicaudata)

Subspecies/variabilityNo valid subspecies. Intraspecific variability more or less marked43.In Yugoslavia, Marincek239 and Marincek & Petrov237 have includedseveral species as being synonymous with I. yeyetta.

Description/identification criteria The Spinicaudata have a bivalve carapace that is generally stri-ated, laterally compressed and protecting the body. But unlikenumerous Spinicaudata such as Cyzicus, Leptestheria, etc., thebivalve carapace of Imnadia yeyetta is not striated over its entiresurface: 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 pairsof legs whereas the females have only 14; the males have legs P1and P2 prehensile, armed with “claspers”, enabling them to claspthe female by the edge of the carapace during mating; thefemales have the pairs P9 and P10 modified into adhesive cylin-der-like structures to agglutinate the eggs between their bodyand their carapace. The carapace, is yellowish to greenish whitein colour, slightly transparent, and measures a maximum of 11 to12 mm.The eyes are sessile.

Similar species The only other species of Spinicaudata possessing a carapacewhich is partially smooth around the umbo is a species of theLimnadiidae family, Limnadia lenticularis (L., 1761), found in theIle-de-France and eastern France. This species is Palaearctic: itsrange extends over central and northern Europe52. L. lenticularisis also present in Italy.


Distribution/range

I. yeyetta is a species mainly present in central Europe (Austria,Slovakia, Hungary, Serbia and Romania). In the Mediterraneanregion it is found in northern Greece and in the former Yugos-lavia52; in France, it is only found in the Bouches-du-Rhône.

Habitat

General description I. yeyetta colonises usually turbid temporary pools, in theCamargue Delta and the Crau. Its presence reflects the dispersalinto biotopes of resting eggs carried by migratory birds flyingbetween central and southern Europe, such as the Anatidae,which are particularly abundant in the region.


CORINE BiotopesLanau pool (Crau) and temporary pools in the Camargue: 22.32 Annual Mediterranean amphibious swards (Lythrion tri-bracteati).

Ecology


FeedingMicrophagous* species feeding by filtering water and sediments.Detrivorous* feeding regime.

BehaviourUsually benthic* species, living on the sediment or partly buriedin superficial sediments.

Description of eggsEggs ovoid, 150 to 190 µm in size, characterised by their spiralledsurface. Confusion is possible with the gyrogonites* (résistantforms) of charophytes, which are distinguished by the presenceof five spiral-shaped apical cells355.

Site and mechanisms of egg layingThe eggs, grouped together on the abdomen of the female, aredischarged into the water during each moult. Their distributionon sediments depends on the position of the female at the timeof the moult, the influence of vegetation and areas of open wateras well as the wind, in particular the mistral (northerly wind), whichcreates movements in shallow water through convection cells.The distribution of the eggs can thus be very variable dependingon the biotope.

Imnadia yeyetta

male

2 mm

egg (150-190 µm)

Mod

ified

fro

m D

efay

e et

al.10

2

86


LongevityLongevity is estimated at less than eight weeks when conditionsare favourable (no premature drying out). However, as for allbranchiopods, it is very variable depending on the temperature.

Biological cycleI. yeyetta is most often present in the spring but can also befound in autumn (November-December). A complete cycle wasstudied in the Camargue by Nourisson & Aguesse270. The femalescan lay up to 400 eggs at a time (the size of the clutch dependson the age of the female). This occurs shortly after mating. Thechitinous* carapace with two lobes appears rapidly, generally atthe metanauplius* stage. This first, or “heilophore”, bivalve stageoccurs three or four moults after the nauplius* stage. The mor-phology of the larval stages has been studied by Eder129. Unlikemost crustaceans, and in particular the branchiopods, the cara-pace is not discarded totally during successive moults, whichleads to the appearance of growth lines374.


Interspecific relationshipsI. yeyetta can be found with the anostracans Tanymastix stag-nalis, Branchipus schaefferi, Chirocephalus diaphanus and/or thenotostracan Triops cancriformis. For more data on the ecologicalconditions of coexistence of several species of branchiopods in atemporary pool in a Mediterranean climate, see Thiéry373.

Requirements for reproduction No data.

Environmental perturbationsNatural perturbations corresponding to the constraints linked tohabitats with fluctuating water levels.

Main natural mortality factorsNot well known. However, as for most branchiopods, the individ-uals develop in a cohort from the onset of flooding371, which lim-its coexistence with predators (larvae of Coleoptera and notonectidHeteroptera, for example) which only colonise the habitats severalweeks 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 tem-porary stillwater habitats of Provence (Thiéry, original data). Theimpact of birds is poorly known.



Number of sitesThe species is only present in France in the Camargue Delta andthe Plaine de la Crau (Lanau pool). In central Europe, I. yeyettahas a fairly wide distribution52.

PopulationsThe densities of the populations are quite low, with an irregularityin the hatching periods and thus of appearance.


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

Natural factorsNo data.

Risks relating to populationsIn the Lanau pool, the population appears to be fairly stable, giventhe state of the cyst bank355; no data on the state of the popula-tions in the central Camargue.


Current measures At two known sites, situated in Natura 2000* areas, the popula-tions appear to be stable, and their biotopes safe from anyhuman pressure. The Lanau pool was acquired by the CEEP in1998 and the temporary pools where this species lives in theCamargue benefit from the status of “Réserve Naturelle” (NatureReserve) (Tour du Valat estate).

RecommendationsGiven the current state of knowledge, no conservation measurecan be proposed. Only the maintenance of the biotopes in theircurrent 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


Hatching and growth of larvaeAdult stages (clutches)

Egg bank

��

IUCNBern ConventionHabitats DirectiveNational and regional protectionNational Red Data Books and Red Lists

-----

87

Macrocrustaceans

Class: BRANCHIOPODS

Order: ANOSTRACA

Family: LINDERIELLIDAE

Main synonymsNone

French name: None

Subspecies/variabilityNo subspecies.Very low morphological variability; low genetic variability (Thiéry,original data).

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

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 morphologyof the antennae: in the males, they are much more developed,biarticulate, and their pincer shape enables them to clasp thefemale during mating. The males possess two spiny ventral hemi-penises at the base of the abdomen (genital apparatus). Thefemales, with shorter antennae, have, on maturity, a heart-shaped brood pouch*, (bicoloured blue/brown) at the base of theabdomen, in which the eggs acquire their outer envelope.The individuals, creamy-greenish in colour and transparent, mea-sure a maximum of between 10 and 13 mm. The females areslightly larger than the males (morphological dimorphism).

Similar species For the non-specialist, all anostracans are very similarly morpho-logically: they can only be differentiated with an identificationkey102, 269. Linderiella massaliensis resembles Linderiella sp. ofSpain as well as L. africana which inhabits the temporary poolsof the Middle Atlas in Morocco.


Distribution/range

FranceSpecies endemic to southeastern France (Var).

BiogeographyLinderiella massaliensis is a vicariant* species of the Moroccanspecies L. africana Thiéry 1986370 which inhabits several dayas(temporary pools) of the Middle Atlas on the Azrou Plateau. Threeother vicariant species are so far known: two in California, L. occi-dentalis Dodds, 1923 and L. santarosae Thiéry & Fugate, 1994367,and one in Spain Linderiella sp.7. These species together consti-tute a case of adaptive radiation through fragmentation of thedistribution range of a “mother” species at the time of the openingof the Atlantic and the Thetys.

Habitat

General description Linderiella massaliensis is found in temporary pools which areflooded in autumn and winter, such as Lake Redon. Generallyspeaking, L. massaliensis occupies pools of karstic origin (poljés*,dolines), of varying depths. The waters are of low turbidity, poorlymineralised (temperature between 5 and 20°C, pH between 7.2and 8.4, C20 between 250 and 600 µS.cm-1) and rich in micro-crustaceans (more than 25 species of cladocerans and copepodsat Bonne Cougne).


CORINE Biotopes22.34 Southern amphibious communities (Preslion cervinae)22.32 Annual Mediterranean amphibious swards (Lythrion tribracteati)

Ecology


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

Description of eggsThe eggs are exclusively “resting eggs”, spherical and spiny. Theyhave a diameter of 240 to 260 µm.

LongevityOne generation (from hatching to the disappearance of the adults)can last, depending on the conditions, for a maximum of three tofour months.

Site and mechanisms of egg layingThe eggs are expelled in small jets by the females and fall to thebottom in a random fashion.

Linderiella massaliensis Thiéry & Champeau, 1988364

Linderiella massaliensis

egg (240-260 µm)

Crus

tace

an d

raw

ing

mod

ified

fro

m T

hiér

y &

Cham

peau

364

2 mm

male

88


Biological cycleAlthough the nauplius* stage only lasts a few hours, growth isslow as the adults are only present from February to March (bree-ding period) whereas the eggs hatch in autumn, from October toDecember. The species disappears from the pools at the end ofthe winter even if they remain flooded, generally until April-May.Little interannual variability in phenology. As for most anostra-cans, the population functions in cohorts (hatching once the poolhas flooded, synchronous growth of the individuals, death).


Interspecific relationshipsLinderiella massaliensis can live alone (Petit Laucien, Var) or befound with Lepidurus apus, Chirocephalus diaphanus (Lake Redon)and Branchipus schaefferi373.

Requirements for reproductionThe species requires an autumn and winter flooding regime whichcorresponds to its biological cycle (fresh water).

Environmental perturbationsEarly drying-out of the pool, in March for example, does not nega-tively affect the species as the adults are present from Februaryand have been able to begin breeding. The introduction of Pump-kinseed Sunfish (Lepomis gibbosus, a species introduced from theUSA) into Lake Bonne Cougne in 2001-2002 caused a collapse inthe 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 anostra-

can. For example, the species is not sensitive to freezing temper-atures (population present under the ice at Bonne Cougne inJanuary 2001, or a minimum water temperature of 1.8°C at PetitLaucien, in February 2003, Thiéry, original data). However, thespecies does not appear to tolerate water which is too warm.



Number of sitesThis endemic species has only been found in five temporary poolsin the Var (around Saint-Maximin and Brignoles) including LakeRedon, Lake Gavoty, and the Petit Laucien (type locality).

PopulationsRare species: the population densities are between 0.5 and 2 ind.L-1

at Bonne Cougne and Petit Laucien. The distribution, which isoften regular, can sometimes become aggregated (influence ofbeds of charophytes, Ranunculus, etc.). Two studies (McNutt &Thiéry and Meglecz & Thiéry) are ongoing, one to map the distri-bution of the cysts of L. massaliensis within Lake Bonne Cougne,the other to quantify the consequences, through the genetic bottle-neck* effect, of the excavation of sediments reducing the pool’segg bank (analysis of enzymatic polymorphism, levels of muta-tion and ‘genetic drift’*).


Anthropogenic factors The Lake Bonne Cougne (Centre Var) is seriously threatened byboth local activities (overdeepening, dyke construction, filling inwith spoil brought in from building work or hospital waste con-taining contaminants, etc.) and by a project for the creation of agolf course on the land above the depression. Although the plan-ning application deposited at the Var préfecture in Toulon wasinitially refused, it has just been approved. Generally speaking,the modifications of the water quality due to pumping from thekarstic water table, extraction of the run-off waters of the topo-graphical catchment area and to water running into the LakeBonne Cougne from the golf course above will jeopardise in theshort term (estimated at less than 10 years), the populations ofbranchiopods as well as those of planktonic crustaceans, clado-cerans, copepods, etc.377

Natural factorsIn the Lake Gavoty (Var), the development of the phytocenosis*considered to be natural, is in fact more likely to be influencedby the nutritional inputs provided by the catchment area (homeswithout a collective treatment system, livestock raising - llamas,goats, horses, etc.). The increase in nitrogen and phosporus levelsreduces the hatching rate of L. massaliensis, whose mechanismdepends on osmotic processes374. In addition, the covering of thesediments by a very dense vegetation bed reduces the mobility ofthe larvae and increases natural mortality.

Risks relating to populationsTwo populations are threatened (Bonne Cougne and Gavoty) outof the five known. If the possibilities for connectivity* are con-sidered for the five populations which are in a fragmented habi-tat within this metapopulation*, these threats represent a potentialloss of 40% of the genetic heritage entering into the gene flowsnecessary for the maintenance of the metapopulation.In 2002, an identical situation in California for the vicariantspecies L. santarosae367 resulted in the implementation of conser-vation measures. In contrast, no actions, or conservation mea-sures are planned in France.


Hatching and larval

Adult stages (egg laying)

Egg bank

��


-----

89

Macrocrustaceans


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

RecommendationsImperative protection of the sites in their current conditionwhere the species is present. Inclusion of the species on a RedList.

Bibliography

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.


90


Tanymastix stagnalis (Linnaeus, 1758)

Class: BRANCHIOPODS

Order: ANOSTRACA

Family: BRANCHIPODIDAE

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

French name: noneSpanish, Italian names: species is present in these two countriesbut there is no common name.English name: Fairy Shrimp (general name for Anostraca)

Subspecies/variabilityNo subspecies.Variability of adults: Margalef236 and Nourisson271 report aberrantT. stagnalis with hypertrophied antennae. The lenticular cysts canpresent some differences in diameter depending on their geo-graphical origin375. An aberrant morphology also exists, corre-sponding to a deformation of the lenticular shape376.

Description/identification criteria Body without carapace, eyes pedunculate, 11 pairs of phyl-lopods*, abdomen without appendages, and two cercopods, ver-milion in colour, forming the furca, at the end of the abdomen. The males are distinguished from the females by their prehensileantennae 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) issituated at the base of the abdomen. The female’s brood pouch*,also borne on the ventral side at the base of the abdomen, bearstwo widely spaced spines. The structure and development of thecyst envelopes have been studied by Garreau de Loubresse149, 150, 151.The males are a pale green colour and the females have a blue-brown to orange oviger*. Their maximum size varies from 6 to 20 mm.

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


Distribution/range

T. stagnalis is a Palaearctic species, present throughout theMediterranean region (Europe and North Africa) but it is veryunevenly distributed52.

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

Other Mediterranean countriesSpain7, Italy (including Sardinia )92, Macedonia and North Africa(Algeria)154, 339.

Habitat

General descriptionTanymastix stagnalis is present in habitats which dry out rapidlysuch as shallow pools (sometimes on rock shelves as in the Fon-tainebleau Forest), vegetated ditches or sansouires (Salicorniascrub) in the Camargue.

France• Roque-Haute Nature Reserve in the Hérault260.• Cupular pools of the Estérel Massif and the Colle du Rouet (seevol. 1, box 7). These basins of several dm2 of surface area, fill upfrom October to May, depending on the year. They are coveredwith an acidic sandy-silty layer (average pH 5.2) from 5 to 10 cmin depth363.• Oligo-brackish temporary pools on calcareous clayey-siltysubstrate in the Camargue (Tour du Valat estate).

ItalyTemporary clear water with sparse vegetation92.

Habitats Directive

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

Tanymastix stagnalis

egg (190-430 µm)

male

2 mm

91

Macrocrustaceans

Salicornia scrubMediterranean and Thermo-Atlantic halophilous scrubs (code 1420)

CORINE BiotopesT. stagnalis is present in Mediterranean temporary pools as wellas in other types of seasonally flooded habitat.

Temporary pools22.34 Mediterraneo-Atlantic amphibious communities.

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

Ecology


FeedingTanymastix stagnalis, like all the anostracans, is microphagous*and feeds by filtering the food particles in suspension in thewater (microplankton, micro-organisms and organic material),using its legs which are covered with bristles.

Description of eggsThe eggs, whose lenticular shape is specific to the genus Tany-mastix, are coppery brown in colour and measure between 190and 430 µm in diameter.

BehaviourThe adults react to variations in light intensity; behaviour involvingfleeing towards the pool bed, or even burying in the mud, havebeen observed when shade is cast over them.

LongevityThis varies according to the temperature and the season whenthe species appears in the habitat (from 30 days in summer toover 60 days in winter).

Site and mechanisms of egg layingThe eggs are laid in open water, where they usually float, andaccumulate towards the edges of the pool262.

Biological cycleTanymastix lives for a period of around three months at all sea-sons of the year, both in winter and in summer . It grows rapidlyand, depending on the temperature of the water, can reach sexualmaturity after seven to 40 days. The nauplius* stage only lasts afew hours.The biological cycle of T. stagnalis is very variable between sitesand from year to year. The cycle begins with the flooding of thepool and its duration depends on the temperature.


Interspecific relationshipsTanymastix stagnalis can be associated with the anostracans Chi-rocephalus diaphanus, Branchipus schaefferi, with the notostra-can Triops cancriformis and with Imnadia yeyetta (Spinicaudata).On the other hand, it is often the only species present in cupularpools, the other species having longer biological cycles.

Requirements for reproduction Pioneer species, colonising clear, fresh and poorly mineralised water.

Environmental perturbationsSpecies present in pools with dry sediments in the summer, with-out a groundwater capillary link. Any modification in the envi-ronment which can cause an increase in the mineral content(conductivity <200 µS.cm-1) reduces its hatching potential.

Main natural mortality factorsPerturbations in the habitat, especially the introduction of preda-tors (such as the fish Lepomis gibbosus, Gambusia affinis, etc.)are the main threats for T. stagnalis.



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

ItalyEleven localities, including six in Sardinia and four in the centreof the peninsula (see map in Mura263).

SpainUncommon species7, 236.

Conservation status

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


Hatching� larval� adults (egg laying) Hatching� larvae� adults

Egg bank

(egg laying)

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

92


Legal status of the species/level of protection control over land ownership and usage) and the pools of theColle du Rouet are included in a Natura 2000* site.

Recommendations It is vital that the small shallow biotopes be maintained in ahealthy condition.

Bibliography

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.



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Anthropogenic factors In general, the infilling of pools and the use of pesticides in therural environment threaten populations.


Current measures Some Italian pools containing Tanymastix are situated in protectedareas263. In France, the Roque-Haute site has a legal “RéserveNaturelle” status (currently ineffective because of the absence of

93

Macrocrustaceans

Class: BRANCHIOPODS

Order: NOTOSTRACA

Family: TRIOPSIDAE

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, 1776Apus productus Bosc, 1802; Apus apus (Linnaeus) Latreille, 1802Lepidurus productus (Bosc) Leach, 1819

French names: (1) Triops cancriforme (sometimes inaccuratelycalled apus – Apus is a bird genus), (2) LépidureEnglish name: Tadpole Shrimp (general name for Anostraca)

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

Description/identification criteria These crustaceans, with a dorsoventrally flattened body, have anoval, streamlined dorsal carapace covering the head, the thoraxand a part of the abdomen. They can reach a maximum size of 10to 11 cm, but the two cercopods* (furca), at the end of theabdomen, are almost as long as the body. The compound eyes are sessile* and dorsal. The phyllopods*, thefirst two pairs of which are locomotive, are more numerous inTriops (48 to 57 pairs) than in Lepidurus (35 to 48 pairs).The posterior abdominal segments are apodal (no legs): the lastfour to five segments in Lepidurus, the last four to seven seg-ments in female Triops and the last five to nine segments in maleTriops.Female Triops are also distinguished from the males by the struc-ture of the 11th pair of thoracic legs which are transformed intoa 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 Triopsbut less so in Lepidurus. The latter is characterised also by thepresence, on the final segment (the telson), of a caudal ramuswith toothed edges.The dorsal shield is greenish-brown in Lepidurus and brownish inTriops, in which the thoracic legs can be reddish or greenish.

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


Distribution/range

FranceTriops is present over the whole country (including Corsica) andis fairly frequent in the east and in the coastal regions, particu-larly the Mediterranean regions. Lepidurus appears to be absentfrom eastern France and Corsica, but is more frequent in the norththan Triops. Generally speaking, the two species are allopatric(separate ranges). The distribution of the two species seems clearlydetermined 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 drierregions than Lepidurus.These distributions may be erroneous, as the two species seem tohave been often confused. Furthermore, their presence at certainstations has not been confirmed since 1950. For further details,consult Nourisson & Thiéry269, Brtek & Thiéry52 and Defaye et al.102.

Other Mediterranean countriesBoth species are present throughout Europe, as well as in theMediterranean Basin for Lepidurus apus apus.• Spain: Lepidurus apus apus, Triops cancriformis simplex, T. c. mauretanicus7 (which replaces T. c. cancriformis in southernSpain).• Portugal: T. cancriformis.• Italy: L. apus (mainland Italy, Sicily, Sardinia ), T. cancriformis,and T. numidicus (= T. granarius), a species of African origin pre-sent in Calabria.

Triops cancriformis (Bosc, 1801) (1) & Lepidurus apus (Linné, 1758) (2)

Lepidurus apus

Triops cancriformis

Crus

tace

an d

raw

ings

mod

ified

fro

m A

lons

o7

egg (430-520 µm)

egg (340-400 µm)

1 cm

1 cm

94


• Malta: T. cancriformis205.• Croatia: L. apus.• Former Yugoslavia: L. apus, T. cancriformis238, 289.• Morocco: T. c. simplex, T. c. mauritanicus, L. a. lubbocki369, 371.• Israel: L. a. lubbocki, T. c. cancriformis (Thiéry, original data ).

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 intemporary watercourses with a weak current. Triops is also present in artificial habitats flooded in the summer,such as the ricefields in the Camargue297.


CORINE Biotopes

Mediterranean temporary pools(1) 22.32 Annual Mediterranean amphibious swards (Nanocype-retalia).(2) 22.34 Southern amphibious communities (Isoetalia).

Ricefields(1) 82.41 Ricefields.

Ecology


FeedingThey take a wide range of food and they can be considered aspredators, herbivores or detritivores. In turbid waters, they arescavengers: 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 digup the mud with the front part of their carapace.

Feeding can vary during the growth of the animal, which canpass from a strictly phytoplanktonophagous* feeding regime,when it measures less than 5 mm, to a herbivorous regime (in thebroadest senses); when it is larger than 1 cm, it also feeds onmicrocrustaceans. As an adult, it is capable of devouring weak-ened anostracans (usually Branchipus schaefferi), chironomid lar-vae, Oligochaete worms and tadpoles (Pelobates cultripes).

BehaviourThey usually swim close to the bottom, but can come up to thesurface, ventral side uppermost, if there is a lack of oxygen.

ReproductionMating is ventral-ventral in Lepidurus, whereas it appears to bedorso-ventral in Triops. Parthenogenetic reproduction is frequentin Triops, but when both sexes are present, sexual reproduction isthe rule. The sex ratio* varies according to latitude with anabsence of males beyond 50°N, a low percentage between 45°

and 50°N (northern and central Europe) and an equal sex ratiobelow 45°N (southern, western Europe and North Africa), withexceptions (mainland Italy, Sicily, etc.).The parthenogenetic females possess a hermaphroditic gland butthe spermatozoa formed degenerate and play no part in repro-duction.

Description of eggsThe 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 layingThe eggs (60 to 70 in Lepidurus) are deposited just before themoult, in clumps, on small pebbles where they remain during thedry period. In Lepidurus, the female can lay eggs on the stems ofaquatic vegetation or bury her eggs in the mud.

Biological cycleAfter hatching, the nauplius* larvae evolve rapidly into metanau-plius* larvae, with the dorsal shield already present.Lepidurus, which is present from January to June (most oftenfrom February to April), has a life expectancy of 4 to 6 months.Triops, which can be found in all seasons of the year in theflooded phase, from March to January (most often from May toOctober), has fairly rapid growth and can produce resting eggsfrom the 15th day, when it is still small in size.

Triops cancriformis


Requirements for reproduction Their cycle being relatively long, these notostracans require asubmersion period of three to five months.Triops, which can be found from March to January, is more ther-mophilic than Lepidurus, which is present from January to June.Triops cannot withstand temperatures higher than 35°C and thusfrom June to August is practically absent from habitats floodedin summer, such as the ricefields in the Camargue.


adult stages (egg laying)

Hatching and larval

Egg bank

��

stages

Temporary pool

Lepidurus apus Jan. Feb. March April May June July Aug. Sept. Oct. Nov. Dec.

adult stages (egg laying)

Hatching and larval

Egg bank

��

stages

95

Macrocrustaceans

Interspecific relationshipsTriops is usually found with Branchipus schaefferi and morerarely with Chirocephalus diaphanus. In the Camargue, it can alsolive alongside Tanymastix stagnalis, Imnadia yeyetta and Eoleptes-theria ticinensis. In ricefields, when its populations explode (densities reaching300 individuals per m2 have been measured in the Camargue297),it can become an important part of the diet of the Little EgretEgretta garzetta169.Lepidurus usually lives with Chirocephalus diaphanus but can, inthe Var, live alongside Linderiella massaliensis.

TerritoriesIn notostracans, there is a differential micro-distribution betweenthe sexes in the pool.

Main natural mortality factorsEarly 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 cer-tain populations (Camargue)169, 297.



In France, they are present at many stations: Triops is found notablyat the sites of Roque-Haute (Languedoc-Roussillon), Padulu (Cor-sica), in the Camargue and the Crau (Lanau pool), while Lepidurushas been found in the pools of the Var, Lake Redon, Lake Gavotyand Saint Maximin (PACA). A description of the distribution ofthe two species in Languedoc-Roussillon is given by Thiéry372.In Italy, these species are fairly widespread, in particular in thericefields.

Conservation status

In France, these two species are common but Triops seems tohave been often confused with Lepidurus and their presence atsome stations has not been confirmed since 1950.



Anthropogenic factors As Lepidurus can be found in the overflow pools of rivers (Loire,for example), physical planning of watercourses and their regu-lation in order to limit flooding can represent an indirect threatto their populations.Triops is sometimes considered to be a nuisance in ricefieldswhere it can be extremely abundant (it uproots young plants,eats their growing tips, and reduces photosynthesis activity bybringing sediments into suspension). Others estimate that theseperturbations are minor and that it acts as an extremely efficientbiological 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 pesticidesdestroys populations.


Current measures In France the pools of Roque-Haute and the Camargue are situa-ted in Nature Reserves (at Roque-Haute, however, conflicts withthe owners do not allow management of the reserve to take place).The lakes of the Centre Var, the Padulu pool in Corsica, and theLanau pool, acquired by the CEEP in 1998, are included withinNatura 2000* areas.

RecommendationsEnsure the conservation of the habitat (hydrological functioningand 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, 1978-1979199 ; Kuller & Gasith, 1996201 ; Lafranco, 1990205 ; Longhurst,1955217 ; Marincek & Petrov, 1992238 ; Moubayed, 1996259 ; Nou-risson & 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 IUCNBern ConventionHabitats DirectiveNational and regional protectionNational Red Data Books and Red Lists

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Malta: rare species (R)344

96


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


Like all the Zygoptera, these species have the eyes well separatedand the four wings of exactly the same shape. The family char-acteristics include the absence of any metallic reflections and theentirely hyaline (transparent) wings, which are petiolate (stalked atthe base). Both species have a mostly dark or black abdomen witha bronze tint and, in males, the eighth segment at least partlyblue (also the ninth in Ischnura pumilio). The pterostigma (darkmark on the wing) is scarcely longer than wide. In I. genei, thepterostigma is slightly larger on the forewings than on the hind-wings, while in I. pumilio it is almost twice as large on theforewings as on the hindwings. Only I. genei has a median tuber-cule on the prothorax* and this is clearly indented. Abdomen: 18 to 27 mm. Wings: 12 to 19 mm.

SubspeciesNone

Similar species Ischnura elegans (similar to I. genei in particular).


Distribution/range

FranceIschnura pumilio occurs only on the mainland (especially theRhône Valley, Pre-Alps, Champagne-Ardenne, Centre-Ouest, andVar), and I. genei is only found in Corsica122, 159.

Other Mediterranean countriesIschnura genei occurs only on the islands of the westernMediterranean (Sardinia, Sicily, Capri, Malta, Elba, etc.).I. pumilio is more widely distributed, along the whole of the coastof Mediterranean Europe and part of the North African andMiddle Eastern coasts95, 401.

HabitatBoth species are pioneer colonisers and are therefore attracted torecently created habitats, temporary or otherwise, usually withstill water. They may be found in brackish water, and sometimesin still areas of water associated with watercourses (oxbow lakes,etc.). They are usually seen below 800 m altitude, however somesites are known at up to 2,100 m (Ischnura pumilio)55, 95, 173.

I. pumilio does not like acid biotopes (Samraoui, unpublisheddata). It is also known to disappear from some sites after a fewyears, probably due to habitat succession68, 207.

Ecology


(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.), withephemeral populations323, while in Spain and northeast Algeriathe species appears in two waves, one in August, the other inSeptember/ 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 capa-ble of breeding six to ten days after emergence69 and the malespossibly even sooner207. This maturation is all the more rapid ifthe individual emerges late in the season, thereby maximising thechances of breeding207. After mating, the female goes alone to laythe 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 abovethe surface of the water and plunges her abdomen in so that shecan lay her eggs in the submerged part of the stem. She prefersto lay at the level of the leaf sheaths (rushes), where the growthof the plant tissues will completely cover the eggs; this appearsto be advantageous for their survival during dry conditions68. Theeggs can survive for at least two weeks in an emergent plant ifit has its base in the water. The eggs hatch quickly (after 15 daysat an ambient temperature of 25°C68, most often between 20 and30 days (Papazian, pers. com.). Rapid larval development enablesthe adults of the second generation to emerge before the pooldries out completely. In temporary habitats, this second genera-tion 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 andthe young larvae spend the winter in the diapause* stage.

Ischnura pumilio (Charpentier, 1825) (1) & Ischnura genei (Rambur, 1842) (2)

Ischnura pumilio

0.5 cm

male

97

Odonata

In the Mediterranean region, the short hatching time, rapid lar-val development, precocious emergence spread over a longperiod of time (advantageous in view of the interannual variabil-ity of the hydrological conditions), and the rapid maturation ofnewly emerged individuals enable two generations of the speciesto be produced per year. The rapid dispersal of individuals fol-lowing emergence (migratory instinct), together with the rapidturnover of generations, enable I. pumilio easily to colonise tem-porary 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 theend of September55.Regarding its ecology, one may refer partly to I. elegans, which isvery similar. Like I. pumilio and I. elegans, I. genei has two generationsper year, with rapid development of the pre-imaginal* stages5, 95.

Simplified phenological cycle of Ischnura pumilio


HydrologyBoth species lay their eggs in submerged stems. The eggs die ifthe plant is out of the water for more than about fifteen days (nosummer diapause at the egg stage), as do the larvae if the pooldries up too early.

VegetationThe sites used are rather sunny and the emergent aquatic vege-tation need not necessarily be very well developed173, 204. The plants used by Ischnura pumilio for egg laying have a grass-like form330 such as rushes (Juncus bulbosus, J. tenuis, J. subno-dulosus, J. inflexus, J. articulatus) or spike-rushes (Eleocharispalustris) 68,173. I. pumilio prefers to colonise water bodies withclayey beds and fairly shallow average depth (less than one metre).

CompetitionIschnura pumilio has a low level of tolerance of competition fromother dragonfly species. It is a requirement therefore that there arefew of these at the sites or at least that some areas are free fromcompetitors (notably I. elegans), or that the vegetation is well-devel-oped enough to allow I. pumilio to find refuge173. Accordingly, the adultsoften only occupy a limited territory within a suitable biotope330.



(1) Ischnura pumilio is less common than I. elegans. In the FrenchMediterranean region, it is known at the pools in the Plaine desMaures244, the Centre Var377, the Colle du Rouet, Porquerolles

(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 outof 50 stations visited as part of the dragonfly survey in Cor-sica159, 323, notably in the temporary pools at Tre Padule de Suartone.



Anthropogenic factors Interference with habitats that dry out in summer (drainage, fill-ing in), and lack of maintenance of pools.

Natural factorsOvergrowing of vegetation at water bodies, siltation and over-growing of wetlands; feeding competition with other dragonflies(such as Ischnura elegans), predation by spiders and natural mor-tality during heavy rain (very frail species).

Risks relating to populationsEarly and prolonged drought may result in the complete destruc-tion of all the larvae at a site. Only recolonisation will thenenable these Ischnura to return.


Current measures In Corsica, Tre Padule benefits from its legally protected “RéserveNaturelle (Nature Reserve)” status. The sites at the Plaine desMaures, the Centre Var and the Colle du Rouet are included inNatura 2000* areas.

RecommendationsThe aquatic vegetation should be maintained in an undevelopedcondition if the site is not much used by dragonflies, and evenmore so if there is strong competition. Vegetation managementshould favour small rushes or club-rushes, with sufficiently lowwater levels. In addition management should aim for the pool tobe flooded until spring (from November to April-May) to enablethe 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 etal., 2002377 ; Wendler & Nüss, 1994401.

Author: Gendre T. Collaborators: Jakob C., M. Papazian, B. Samraoui & N. Yavercovski


Winter larval diapause,followed by larval growth

Emergence of adults, egg laying and migration

Larval development, emergenceand dispersal to other pools

First generation (pool A)

Second generation

Egg laying (pool B)

Hatching of eggs

Larval diapause…

��

�

Temporary pool in southern France

IUCNBern ConventionHabitats DirectiveNational and regional protection


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Italy, regional protection(Tuscany): regional law n° 56of 06/04/2000

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98


Order: ODONATA

Suborder: ZYGOPTERA

Family: LESTIDAE

Main synonyms(1) Agrion nympha Hansemann, 1823Agrion barbara Fabricius, 1798Lestes barbara Selys, 1840(2) Lestes vestalis Rambur, 1842 Agrion virens Charpentier, 1825

French names: (1) Leste sauvage, (2) Leste verdoyantEnglish names: (1) Shy Emerald Damselfly, Southern EmeraldDamselfly, (2) Small Emerald Damselfly

SubspeciesL virens virens (in southern France)L. virens vestalis (in Italy)

Description/identification criteria (1) Lestes barbarus has a clearly bicoloured pterostigma (darkmark in the wing) when mature: brown in its basal half and yel-low-white in its apical half. The overall body colour is clearmetallic green at first, becoming more or less coppery, even dullbrown in old individuals. The lower hind part of the head, thecoxa, the metathorax, and the thoracic and abdominal lines arepale yellow. The male has short cerci*. The female has a roundedvulvar scale, untoothed, and ochre-coloured legs in most cases.Measurements of male: abdomen 26 to 35 mm; hind wings 19 to25 mm.Measurements of female: abdomen 26 to 33 mm; hind wings 21to 25 mm.

(2) Lestes virens has a uniformly pale brown pterostigma, edgedwith white veins (apart from in juveniles). The general bodycolour is brilliant bronze-green, the sides of the thorax and thetip of the abdomen are pale blue in mature males. The lower hindpart of the head is pale yellow, as in L. barbarus. The male hasvery short cerci. The female has a vulvar scale with a short, onlyslightly pointed tooth. L. virens is smaller than L. barbarus. Thesubspecies from southern France, L virens virens, shows a thinpale yellow line along the anterior part of the metathorax*.Measurements of male: abdomen 24 to 34 mm, hindwings 16 to24 mm.Measurements of female: abdomen 24 to 30 mm, hindwings 17to 23 mm95, 173, 322, 402.

Similar species Lestes barbarus is very similar to L. virens (especially at the juve-nile stage) and to its subspecies L. v. vestalis.


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 alti-tude. As a pioneer species, it appears more particularly to preferhabitats which dry out in summer (temporary pools) and brack-ish coastal marshes, where there are often large populations.(2) L. virens occurs in still bodies of fresh water, including tem-porary pools, colonised by vegetation, up to 1,000 m altitude.

Ecology


(1) In southern France, the adults of Lestes barbarus (known asimagos) are fairly common from the end of May (immature atthat time, they become mature in June) to the end of September(at which time they colonise new pools). In spring, after emer-gence, the immature imago leaves the water, to return when it issexually mature, usually in June-July. The period of maturationbetween flying away and returning to the pool is estimated to be15 days on average for Zygoptera89. In other regions maturation

Lestes barbarus (Fabricius, 1798) (1) & Lestes virens (Charpentier, 1825) (2)

Lestes barbarus

Lestes virens

males

1 cm

99

Odonata

may take place following a period of aestivation, as is the case inparticular of the populations of southern Italy and northernAlgeria338, 340, 390.The duration of the maturation period influences the egg-layingperiod: in southern France laying may take place from the end ofJune to September (Papazian, pers. com.), while in southeastAlgeria (in Numidia) the females become mature in Septemberand no breeding behaviour is seen before October (Samraoui,unpublished data). Mating (forming a heart shape) and the tandem flight which fol-lows, may last for between 30 and 160 minutes (for both species)and usually takes place during late morning387. After mating, thepartners continue their tandem flight and the females begin tolay eggs using their ovipositor (egg-laying organ). In both species,eggs are laid into the aerial parts of plant stems4: the female’sovipositor pierces the plant to insert the eggs, which are oftenlaid in groups in the same stem, with gaps of a few millimetresbetween them. The distances covered between stems range froma few centimetres to a few metres387. Often, the male breaksaway from the tandem formation after a certain time, and thefemale then continues to lay by herself. The total duration of egglaying may last for up to six hours, depending on the weatherconditions and diurnal rhythm.After a period of diapause* inside the plant, which varies accordingto the date of laying, the eggs hatch in the spring of the followingyear, and give birth to elongated larvae which develop fairlyquickly, taking about two months and moulting about ten times.These larvae are carnivorous, like all dragonfly larvae, and eattwice their own weight in food every day.Thanks to the diapause eggs (a primitive protective characteris-tic of the species) and to the short larval development period (sixto eight weeks) very synchronised emergence may take place inspring, with practically all the imagos emerging over a shorttime.The sex ratio* (calculated at a Camargue site) is 50% females forlarvae and 54.1% females for exuviae4. The adult dies later in theautumn. 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 thantwo and a half months4.

(2) The mature adults of Lestes virens may be seen from June toSeptember in southern France, and from May (occasionally endof April) to November in northeast Algeria (Samraoui, unpub-lished data). The female, who may or may not be accompanied bythe male, lays her eggs in pairs, in spring and summer, into plantstems. After a winter egg diapause (often preceded by a summerdiapause), larval development takes place in the following springas with L. barbarus, and lasts for eight to ten weeks402.In the North African part of the Mediterranean Basin, a proportionof Lestes also lays in summer, before the site becomes flooded;the diapause eggs then withstand the drought, then overwinterand do not hatch until the following spring340. In Numidia, this istrue of L. virens at lower altitudes (laying in June-July), while athigher altitudes breeding activity in this species does not beginuntil August (Samraoui, unpublished data).Both species (L. barbarus, L. virens) only have one generation peryear.

Phenological cycle of Lestes barbarus


Winter temperatures Lestes barbarus and L. virens are adapted to the Mediterraneanclimate due to their rapid life cycle and their temperaturerequirements95. However, in southern France, L. barbarus requiresa period of cold, of at least fifteen weeks below 10°C, to triggerthe hatching of all of the eggs4.

HydrologyAs far as is known for these two species, the eggs are laid mostoften in a dry environment; the pool need not be flooded at thetime of laying for the eggs to survive4, 340, since they are capableof 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 orsummer, and this is usually the case thanks to the rapidity of lar-val development.At the time of the emergence of the prolarva*, it sometimes hap-pens that the place where the eggs were laid is dry; the prolarva,which on hatching allows itself to drop off the stem, is thencapable of crawling to the water over a distance of a metre ormore (Papazian, pers. com.).

VegetationAccording to Utzeri et al.387, the plants preferred by Lestes virensvestalis for egg laying at a temporary pool in Italy (Rome), areCarex sp. and Juncus effusus (green or dry stems). L. barbarusshows a preference for the same species, as well as Juncus artic-ulatus, Polygonum hydropiper, Alisma plantago and Mentha sp.Samraoui (unpublished data) reports that in Numidia, L. barbarusprefers to lay in the stems of Juncus effusus.



Mainland FranceBoth species are known from the Alpes Maritimes at the Pic desCourmettes (Papazian, pers. com.), as well as in the Var: Plainedes Maures244, Colle du Rouet325, Joyeuse valley at Callas, lakes ofthe Centre Var377. Lestes barbarus has been observed at severalother localities in the Var (Palayson wood, Giens Peninsula)363 (Papa-zian, pers com.), as well as in the Bouches-du-Rhône in the Crauand in the Camargue363, and in the Hérault at Roque-Haute260.L. virens occurs in the Vaucluse along the Durance (Papazian,pers. com.)


Eggs in diapause

Larval development

Emergence of adults

Egg laying

Maturation of adults

Death of adults

��

Southern France

100


CorsicaLestes barbarus, a migratory and pioneer species, appears to becommoner than L. virens, with six stations out of the 50 visitedduring the survey of dragonflies in Corsica, compared with onlytwo stations for L. virens. Both species are known in particularfrom Tre Padule and Padule Maggiore159, 323.

Conservation status

No information.


Lestes barbarus and L. virens have no legal protection at the pre-sent time227.


Anthropogenic factors Drainage and infilling of the sites used.

Natural factorsSiltation and overgrowing of wetlands; feeding competition withother dragonflies (e.g. Sympetrum fonscolombii), predation bywater beetles, parasites, interannual variability of hydrologicalconditions (early drying out may result in the death of all larvae).


Current measures In France: • The Tre Padule de Suartone and Roque-Haute sites are legallyprotected due to their “Réserve Naturelle (Nature Reserve)” sta-tus (at Roque-Haute, the management of the site is compro-mised as there is no control over land ownership or the activitiesfor which the site is used).

• The lakes of the Centre Var, the Plaine des Maures, the Palaysonwood and the Colle du Rouet are included within Natura 2000*areas, as well as the Lanau pool, which was acquired in 1998 bythe CEEP.

Recommendations• Attempt to ensure, at temporary pools and fresh or brackishstill waters, that there are flooded conditions until spring (April-May), suitable for the development of the larvae.• Maintain and promote aquatic vegetation and bands of vege-tation around the edge of the pool, suitable for egg laying(rushes, sedges, water plantains, mints, bistorts).

Bibliography

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.

Authors: Jakob C. & J. FuselierCollaborators: Papazian M., B. Samraoui & N. Yavercovski


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101

Odonata

Order: ODONATA

Suborder: ANISOPTERA

Family: LIBELLULIDAE

Main synonyms(1) Libellula erythroneura Schneider, 1845Libellula insignis Brittinger, 1850(2) Libellula hybrida Rambur, 1842Libellula nudicollis Hagen, 1850Libellula meridionalis Sélys, 1841

French names: (1) Sympetrum de Fonscolombe, (2) Sympetrum méridionalEnglish names: (1) Red-veined Darter, (2) Southern Darter

Description/identification criteria Sympetrum have six to eight veins in the forewings. The bases ofthe hindwings have a yellow mark which is fairly well-developedin S. fonscolombii (but often absent in S. meridionale).(1) In S. fonscolombii, the legs are black with a yellow stripe onthe outer edge.Measurements of male: abdomen 22 to 29 mm, hindwings 26 to30 mm.Measurements of female: abdomen 22 to 28 mm, hindwings 26 to 31 mm.(2) S. meridionale has yellowish or brown thorax with practicallyno 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, hind-wings 20 to 30 mm95, 322, 401.

Similar species No confusion possible with Sympetrum fonscolombii, whereas S. meridionale may be confused with S. vulgatum ibericum andS. sinaiticum.


Distribution/range

France(1) S. fonscolombii is found especially in the Mediterranean area,including Corsica, and in the Rhône Valley; it is much rarer in therest of the country.(2) S. meridionale has the same range but appears to be com-moner in the rest of the country122, 401.

Other Mediterranean countriesThe whole of the Mediterranean region95.

Habitat

(1) Sympetrum fonscolombii prefers still, occasionally brackishwater (pools, ponds, gravel pits, marshes, boggy edges of lakes),from sea level up to 2,000 m (sometimes much higher for wan-dering individuals). It prefers wetlands that are very little vege-tated (or vegetated with helophytes* and hydrophytes*), wellexposed to the sun, and of very variable water depth.

(2) S. meridionale also prefers still water, more or less colonisedby vegetation, up to 1,800 m altitude95, 173.

Ecology


The eggs of Sympetrum often enter a diapause* phase after beinglaid, and do not hatch until they have spent the summer or win-ter in egg form89. However, in S. striolatum in Corsica it is alsopossible to observe larvae hatching out before the winter andthese spend the winter in the water of the pool (Papazian,unpublished data.) In both cases, this type of annual cycle, withthe eggs laid in autumn, corresponds to the flooding cycle ofMediterranean temporary pools340. S. meridionale and S. fons-colombii lay directly into the water (a highly evolved characteristic)but have partly retained the diapause at the egg stage.

Sympetrum fonscolombii (Sélys, 1840) (1) &Sympetrum meridionale (Sélys, 1841) (2)

Sympetrum fonscolombii

Sympetrum meridionale

1 cm

1 cm

males

102


(1) Adult S. fonscolombii show well-marked exploratory behaviourand undertake movements over great distances (migratoryspecies)95, 122, which explains their sudden appearance in more orless large numbers322. The imagos are usually seen from March(an early species) to December, mainly from March to June andfrom August to December in temporary habitats. The mass move-ments of various Sympetrum that are observed in the countriesaround the Mediterranean are explained as being flights betweenpools which dry out in summer and aestivation sites338.

In this species, there are at least two and even three generationsper year in the Mediterranean region: the first emerges in Marchand, following a period of rapid maturation, lays eggs whichhatch after very short incubation period; the larvae grow quicklybefore the water body dries up completely, and the emergence ofthe second generation takes place340 (Papazian, pers. com.). Thissecond generation lays at the end of summer or in autumn, assoon as the water returns. In southern France, either the eggs (ora proportion of the eggs) of the second generation hatch quicklyand the larvae spend the winter in diapause, or they do not hatchuntil the following spring, depending on biological and ecologi-cal 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 anautumn and winter growing phase which is prolonged due to thelow temperatures; in addition, if there is water available in sum-mer the second egg laying takes place earlier, and the speciesmanages to lay for a third time late in the season; this is also thecase in southern Europe in ricefields340.

Laying usually takes place in tandem (the male still clasping thefemale). In full flight, the female, who like all Libellulidae has noovipositor, taps the water surface with her abdomen: the eggsthus released into the water remain caught up on a support (anaquatic plant in most cases), before falling to the bottom. InFrance the larval stage of S. fonscolombii (first generation) isamong the shortest observed among Odonata in France (alongwith Hemianax ephippiger). In the Mediterranean region, possi-bly after a diapause due to the drying out of the pool or to win-try temperatures in the water, the eggs hatch in a short time(rapid incubation). Emergence is usually nocturnal, extendingfrom March in the Mediterranean region95, to November.

(2) In southern regions, adult S. meridionale may be seen fromthe beginning of June (a late species) to mid-October95. At poolsin northeast Algeria, Samraoui et al.341 have noted the presenceof adult S. meridionale around the pools in May-June (immatureimagos) and in September-October (mature).In temporary habitats, the adults have just enough time toemerge before the summer drought. They mature over the sum-mer. At this time in northeast Algeria340, they disperse to refuges(woodland) at high altitudes where they slowly develop sexualmaturity (imaginal diapause); adults may travel up to 35 km toaestivate at altitudes greater than 1,000 m.After the summer phase of maturation, the adults return, at theend of summer or in autumn, to the egg laying sites, which bynow are usually flooded, to breed. In southern France (Papazian,pers. com.), the eggs may either hatch rapidly, the larvae thenspending the winter in diapause, or delay hatching until the fol-lowing spring. On the other hand in northeast Algeria larval

development continues until the emergence of adults in the springof the following year; there is no embryonic diapause if water isavailable, but if it happens that the species lays in habitats thatare still dry, the eggs can then remain resistant to desiccationuntil the first rains340. Contrary to S. fonscolombii therefore, S. meridionale only has one generation per year in temporary habi-tats (southern France, North Africa)341 (Papazian, pers. com.). Theeggs (up to 550) are laid in places with shallow water, amongreeds and Carex, even in puddles.S. meridionale is a migratory species like S. fonscolombii. Thelifespan of the adults which survive to aestivation is among thelongest known among Anisoptera.

Simplified phenological cycle of Sympetrum fonscolombii


Hydrology• Larval development:Aguesse4 showed that laying does not always take place at timesand places that are favourable for the development of the larvae.Among Libellulidae such as Sympetrum, females are frequentlyseen laying eggs on the dry outer edge of a pool, which will notbe submerged at a suitable time, or on the water surface at apool which dries out a few weeks later. While larval developmentis jeopardized by the failure of the laying site to refill with wateror by early drying out of the pool, these laying strategies remainnonetheless effective overall thanks to egg diapause (faculta-tive), which takes place if the site is dry and which enables theeggs to survive the dry period. It should be noted that this groupof Odonata has the widest geographical range4.

• Adult diapause: For North Africa, Samraoui et al.341 described a prolonged, dia-paused adult phase in S. meridionale (3-4 months during thesummer dry period), when there is no reproductive activity. VonHagen (in Samraoui et al.341) described the same phenomenon inS. striolatum in southern Spain. Papazian (pers. com.) hasobserved it in Corsica and in southern France.

`


Diapause of eggsor larvae, followed by larval development

Emergence of adults, sexual maturation, followed by egg laying and migration

Larval development,emergence of adults, migration

First generation

Second generation

Aestivation and maturation of adults

Eggs laying and migration

��

Southern France

103

Odonata



Mainland FranceBoth species are fairly common near the Mediterranean, espe-cially 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 theColle du Rouet325, the Plaine de Palayson363, the oxidation pondsat 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-Hauteplateau3.

CorsicaSympetrum fonscolombii occurs at six out of the 50 stations vis-ited during the survey of the dragonflies of the island, and S.meridionale occurs at ten of these 50 stations323. Notably, bothare present at Padule Maggiore and Tre Padule.



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 aquaticvegetation is probably harmful to Sympetrum fonscolombii.(2) Male S. meridionale are often parasitized by mites (Acaruslibellulae De Geer) 322.


Current measures In France, some sites are wholly or partly legally protected bytheir Réserve Naturelle status (Roque-Haute, Camargue, Les TrePadule de Suartone); others are included in Natura 2000* areas(Colle du Rouet, Plaine des Maures, lakes of the Centre Var, Lanaupool), and among these some have been the subject of operationsof land ownership control in the context of the LIFE “TemporaryPools” project (some parts of the Plaine des Maures and theLanau pool).

Recommendations While Sympetrum fonscolombii is a common species in the FrenchMediterranean region, S. meridionale is much more confined tothe coastal fringe, and the survival of these populations there-fore depends above all on the conservation of coastal pools.

Bibliography

AGRN-RH, 20003 ; Aguesse, 19614 ; Aguesse, 19685 ; Anonymous,199711 ; Corbet, 199989 ; D’Aguilar & Dommanget, 199895 ;Dommanget, 1994122 ; Heidemann & Seidenbusch, 2002173 ; Médailet 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. FuselierCollaborators: Papazian M., B. Samraoui & N. Yavercovski


-----

104


Bufo calamita Laurenti 1768

AMPHIBIA

BUFONIDAE

Main synonymsNone

French names: Crapaud calamite, Crapaud des joncsSpanish name: Sapo corredorEnglish name: Natterjack Toad

SubspeciesNone

Description/identification criteria The length varies between 40 and 80 mm (sometimes up to 90 mmin the female). The build is sturdy with prominent paratoidglands* arranged in parallel, a horizontal pupil and pale greenish-yellow iris, and a greenish to brownish dorsal surface generallycrossed by a pale yellow mid-dorsal line125, 333. The breeding maledevelops brown nuptial pads on the toes. The species has a char-acteristic mode of locomotion: unlike most amphibians theNatterjack does not jump, but walks rapidly or runs. It is easy tospot thanks to its far-carrying song, which may be heard from upto a kilometre away.

Similar species The Green Toad (Bufo viridis) has no yellow dorsal line; it hashandsome green markings contrasting with its ash-grey back.The Common Toad (B. bufo) has a golden-red iris.


Distribution/range

France The Natterjack is found in all parts of France except for the highmountains (maximum 1,500 m in the Pyrenees) and Corsica. Thisspecies extends all the way into built-up areas in some towns. It isabundant in the south but its distribution becomes more discon-tinuous towards the north.

Other Mediterranean countries The Natterjack occurs in Spain (except for part of the northwestand the Pyrenees) and in Portugal168, but is absent from Italy.

Habitat

The Natterjack is typically an amphibian of open habitats, withshort vegetation and few trees. It prefers loose substrates, eithersandy (sandy back dunes) or stony (Plaine de la Crau, Plaine desMaures), degraded garrigue and maquis and also cool heathlands(Plateau de l’Aubrac). It adapts to man-made sites: parks andwasteland on the edges of built-up areas. Its aquatic habitat isfairly varied: live stock-watering ponds, flooded areas amongsand dunes, puddles, Mediterranean wadis, quarry lakes, etc.

These habitats are typified by often shallow water (a few tens ofcentimetres at most), an absence of predators, and sparse or evencompletely non-existent aquatic vegetation.

Ecology


This is a pioneer species, which is able rapidly to colonise recentlycreated habitats such as temporary pools. It prefers loose, sandysoils. In search of suitable habitats, Natterjacks may easily coverdistances of more than 2.5 km268. The males may live for sevenyears while females can sometimes live for 17 years125. Like almostall European amphibians, the Natterjack has terrestrial and aquaticphases. Breeding usually starts in March in the Southern Francewith the young toads leaving the water at the start of summer(June), extending into July further north. In the south, a secondbreeding season is sometimes observed at the end of summer,depending on the storms of August and September. Young frogsfrom this breeding season emerge at the end of September orbeginning of October. Males form choirs and sing in shallowwater to attract the females. After mating, the adults quicklyleave the pool, and the eggs, which are laid in strings, (2500-4000 at a time) hatch after a few days. Aquatic larval develop-ment is relatively brief (four weeks to three months). Their rapiddevelopment does not always protect the young Natterjacks fromfalling victim to early drying out184, 185. The juveniles are very smallat metamorphosis compared with other species, but grow quickly.Sexual maturity is generally attained when they are three yearsold. The tadpoles, which are completely black, may feed on algaeor other plants as well as detritus. They are sometimes canni-balistic268. Both adults and juveniles are predators of terrestrialinsects.

Bufo calamita

1 cm

105

Amphibians

Annual phenological cycle


HydrologyFlooding 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 notdisrupt the process of larval development and the year’s breed-ing success.

Substrate for spawningThis is one of the rare species capable of breeding in the com-plete absence of any aquatic vegetation, in concrete-lined ponds,rocky wadis or bare sandy areas.

Interspecific competitionThis species avoids temporary pools used by other species, espe-cially the Green Frog (Rana perezi) or the Common Toad; thedigestive tract of some species contain an alga which inhibits the larval growth of Natterjack Toad tadpoles191.

Impact of perturbationsThis species is highly adapted to unstable habitats and as a resultis very tolerant of perturbations in the aquatic environment. Itsshort larval development period, tolerance of high temperatures,ability to cover great distances to breed and production of a largenumber of eggs make it a species well-adapted to the risks ofdrying out and for the colonisation of new sites. On the otherhand, it needs open habitats for feeding and moving around, anddeclines when the habitat becomes wooded.

ShadeThe Natterjack prefers pools with strong insolation and rathershort terrestrial vegetation, the larvae being capable of with-standing high temperatures, up to 40°C for brief periods.



The Natterjack Toad is fairly abundant in the Mediterraneanregion, rarer and more sparsely distributed in northern Europe125.It occurs at about 29% of Réserves Naturelles de France land and10% of Réserves Naturelles Volontaires. However, if only thoseRéserves Naturelles located within its French range are included,it is present at about 48% of them135.



The Natterjack Toad is a relatively little-threatened species in theMediterranean region, aside from the impact of increasing amountsof woodland.

Anthropogenic factors The transformation of a shallow temporary pool into a deep oreven permanent pool poses problems for this species. Such a poolcan support greater numbers of invertebrate predators than ashallow 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 WesternSpadefoot (Pelobates cultripes), Parsley Frog (Pelodytes puncta-tus), Midwife Toad (Alytes obstetricans), Marbled Newt (Triturusmarmoratus), Great Crested Newt (T. cristatus), Palmate Newt (T. helveticus), Painted Frog (Discoglossus pictus) or Stripeless TreeFrog (Hyla meridionalis). This is the case for example at theValliguières pool and the Opoul pool, where temporal segregationfrom the other species allows it to breed successfully.The introduction of fish causes the decline and disappearance ofthis species.Finally, the degradation, destruction or modification of the ter-restrial habitat has harmful effects, by destroying the aestivationand hibernation sites which are essential for the survival of amphi-bians in the Mediterranean (stumps of trees or bushes, rocks,vegetation near the water bodies).

Natural factorsIf the aquatic and terrestrial habitats become overgrown throughnatural succession, this poses problems for pioneer species of openhabitats like the Natterjack. The increase of terrestrial plantcover, in particular, has been proved to be a factor in its decline:on a very small scale, plant cover provides shelter, but on a largerscale it may form an obstacle to moving around, and a bare habi-tat is greatly preferred (see the Plaine de la Crau, where the speciesreaches extremely high densities!).

Risks relating to populationsThe disappearance of shallow temporary water bodies and thedestruction of breeding sites increase the risk of local extinctionas a result of the increasing isolation of habitats.


Departure

Arrival

Spawning

Larval development then metamorphosis

Normal cycle

Exceptional cycle

Aestivation by adults then terrestrial phase and

hibernation

Spawning

Larval development

Departure, terrestrial phase andhibernation

�

�

Southern France



-Annexe IIAnnexe IV- France, national list: Decreeof 22/07/1993/Journal Officiel09/09/1993- 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

106



Current measures Recent scrub clearing around shallow pools such as those atRoque-Haute (terrestrial vegetation) is improving the quality ofthe habitat and migratory routes. The overdeepening of pools,frequently undertaken in the case of pools which have becomevery shallow through aggradation*, has a negative impact on thepopulation56, as does the creation of permanent pools next toshallow temporary pools35: both these measures favour other, com-peting species of amphibians.

RecommendationsIn most cases, the reinforcement of populations or the substitu-tion of recreated habitats for destroyed habitats are effective

methods and are often more prudent than deepening the pool.Population reinforcement may be achieved by creating a series oftemporary pools with a range of varying depths around a site27, 35.

Bibliography

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

Author: Jakob C.

107

Amphibians

Discoglossus sardus Tschudi, 1837

AMPHIBIA

DISCOGLOSSIDAE

Main synonymsNone

French name: Discoglosse sardeItalian name: Rana TirrenicaEnglish name: Tyrrhenian painted Frog

SubspeciesNone

Description/identification criteria This Painted Frog is a small amphibian of slender appearance witha strong resemblance to a frog of the genus Rana. The males,which are slightly larger than the females, may reach 5 cm inlength from snout to cloaca. The colour of the upperside is veryvariable, most often brownish but sometimes reddish, greyish oreven greenish, more or less heavily marked with irregular darkareas all over the body. A pale band runs between the eyes acrossthe top of the head. This pattern gives the animal the appearanceof a dead leaf, which makes it very cryptic on the ground and inthe water. The hind legs have pale and dark bands. The undersideis a rather uniform pearly cream colour. The pupil is shaped likea water droplet and the eardrum is not visible.Eggs are laid in tens at a time, separately or in the form of asheet 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 to4 mm gelatinous capsule.The tadpoles are small and dark-coloured, fairly similar to Bufotadpoles. Like all the Discoglossidae, they have a ventral spiracle*.

Similar species The Tyrrhenian Painted Frog forms a species pair with the Cor-sican Painted Frog, with which it was confused until 1984. Com-pared with this species the hind leg is shorter: if extended alongthe body, the heel usually reaches as far as between the eye andthe tip of the snout, but no further. The snout is more pointedand the part between the eye and the nostril is slightly sloping.


Distribution/range

This Painted Frog is restricted to the French and Italian Tyrrhenianislands: Sardinia and some of its islets, Corsica and the LavezziIslands, 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 long-standing, such that each of these populations has evolved distinc-tive and specific characteristics. In Corsica, it ranges over a goodpart of the island, from sea level up to at least 1300 m. At Port-Cros it is found in all the streams on the island, notably the Vallonde la Solitude and the Vallon Notre-Dame. It seems to be rare onthe Île du Levant where surveys are still required to ascertain itsprecise status.

HabitatThis species lives in a wide range of habitats, from low maquis tomature oak or pine woodland. It is undemanding in its choice ofbreeding sites: springs, coastal marshes, canals, brooks and streams,depressions cut off from the main channels of rivers, reservoirsand fountains. In particular it uses impermanent bodies of waterwhich in coastal regions are fed by unpredictable events (isolatedpuddles, temporary runnels, temporary pools, etc.). It is not uncom-mon for the eggs to fail to hatch, or to be dried out by evapora-tion or carried away by floods. This species can breed in relativelybrackish water and, in contrast with the Corsican Painted Frog,can adapt to disturbed or modified habitats (polluted streams onthe edges of villages, artificial reservoirs). Outside the breedingseason, the species may be observed under piles of leaves or undertrunks of fallen trees, several hundred metres from the nearestaquatic habitat.

Ecology


This species is active for a good part of the year, by day as wellas 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 followingheavy rain (two to three nights after the rain starts). The males’song is quiet, not very far-carrying and restricted to the breedingseasons. Males arrive at the breeding sites first and remain therelonger than the females. Mating is brief and the eggs are laid inbundles of 20 to 50. The small tadpoles hatch out after aboutthree days. These tadpoles grow rapidly and metamorphose afterfour to six weeks. Some, originating from eggs laid in autumn,grow much more slowly (20-25 weeks) during the winter. InCorsica as well as at Port-Cros, the eggs are laid over a very longperiod of time. Several generations of tadpoles coexist in a singleplace and it is common to see eggs, tadpoles of all sizes andjuvenile frogs all present simultaneously.At metamorphosis individuals are very small (about 8 mm) andresemble 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.

Discoglossus sardus

1 cm

108


Phenological cycle


HydrologyThis species is perfectly adapted to the unpredictability of the Medi-terranean hydrological regime (very extended breeding period,rapid development of larvae) reducing its vulnerability to climatichazards.

Aquatic habitatThis Painted Frog uses almost all types of aquatic habitat forbreeding, with the exception of flowing water and very large waterbodies. This species ranges widely over the countryside, withoutspecialising in any one type of habitat. Pools are therefore notessential for the survival of its populations, in contrast with otheramphibians.A slight degree of salinity is tolerated, as well as a high degreeof turbidity (muddy puddles). Organic pollution does not seem tohave any affect on breeding. In general, this species uses smalltemporary water bodies, preferably exposed to sunlight, althougha fairly large amount of plant cover may be tolerated.

Terrestrial habitatAlmost all kinds of habitat may be used: dense woodland, maquis,or partly built-up areas.



Most of the populations are large, except on some islets whichhave practically no fresh water such as the Lavezzi islets. Corsicaundoubtedly supports the largest numbers. In Sardinia this speciesseems to be less common and it is only known there from a smallnumber of sites. There are no quantitative data available for theTuscan archipelago, but according to Knoepffler (1962), the islandsof Giglio and Montecristo have very large populations due to thepresence of abundant fresh water.On Port-Cros, Knoepffler estimated the population size at around5000 adults at the end of the 1950s. Recent observations indi-cate that numbers may be lower at present. The introductionsome years ago of fish into the artificial reservoir where a highproportion of the island’s breeding adults were concentrated hadreduced the breeding capacity of the population. The eradicationof the fish in 2001 made the site once again available for thePainted Frogs, which bred there in numbers in 2002-2003.(Joyeux, pers. com.). On the Levant Islet, Knoepffler198 considered

the species to be common during the 1950s-60s, but threatenedby the deforestation of the island, the destruction of wateringplaces and the impact of the Viperine Snake on the larvae andthe adults. No surveys have been carried out since then to assessthe current status of this population. Most of the occupied sitesare situated on steep islands that are not very suitable for agri-culture and are well preserved overall.



Anthropogenic factors Owing to its lack of specialisation, the threats to this species arenot very serious. Dams interrupt the distribution of populationsand reduce the number of breeding sites; water pollution undoubt-edly affects the eggs and larvae. One threat which cannot beignored is that of forest fires, which as well as destroying adultshave the effect of profoundly altering the hydrological regime ofthe watercourses (more torrencial regime) and the nature of thebreeding sites (input of ash and sediment). Coastal populationsare no doubt affected by anti-mosquito treatments (Corsica) andby the spread of the Italian Pool Frog (Rana bergeri), introducedprobably recently into Corsica. The introduction of fish into closedwater bodies (small reservoirs, ponds) is recognised as constitutinga threat (for example the case of the Parc de Port-Cros). It isacknowledged that the introduction of alien competitive or preda-tory species in particular should be controlled (American crayfish,fish, frogs), as these will undoubtedly have catastrophic effectson this species.

Natural factorsNatural threats are few in number apart from the problem of fire,which may in some cases constitute a serious threat (small woodedislands such as Port-Cros). Reafforestation is known to constitutea threat to some populations by its effect on the breeding sites(shading of open water, and reduction of open water due to theaccumulation of litter).

Risks relating to populationsIt is necessary to maintain sufficient numbers to counter the riskof chance extinctions, which is particularly high in an island situa-tion.


Current measures The Tyrrhenian Painted Frog occurs at many sites in France andItaly which benefit from official protection either specifically for


Egg laying (sometimes) thenLarval growth and metamorphosis

Occasional cycle

Usual cycle

Extended egg laying

Larval growth and metamorphosis

Departure and terrestrial phase

��

Southern France



-Annexe IIAnnexes II and IV- France, national list: Decree of 22/07/1993/Journal Officiel09/09/1993- Italy, regional lists (Sardinia, Tuscany): regional law of 29/07/1998 - France, national Red DataBook: “rare”242

- Italy, national Red Data Book14

109

Amphibians

wildlife (Parc National de Port-Cros, Nature Reserves of Etang deBiguglia, Scandola and Bouches de Bonifacio in Corsica, ParcNaturel Régional de Corse, Tuscan Islands National Park) or as aresult of laws relating to land use (in France, sites belonging tothe Conservatoire de l’Espace Littoral et des Rivages Lacustresand 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 accountin the management plan, is recommended. The species is the sub-ject 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 populationswhich 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 offorest cover around the main breeding sites by selective stream-side felling. In addition, it would be worth reassessing the dry

stone dams along the watercourses, which were constructed inthe early years of the Park to hold back sediment: the piles ofleaves accumulated over the years have had the effect of blockingthe beds of the island’s streams, tending to cause the water torun underground, with consequent losses of the amount of openwater available for use by the aquatic fauna. The number ofbreeding sites has decreased significantly as a result.

Bibliography

Anonymous, 200414 ; Capula & Corti, 199361 ; Cheylan &Massemin, 199973 ; Cheylan, 198375 ; Delaugerre & Cheylan1992103 ; Delaugerre 1999104 ; Gasc et al., 1997152 ; Knoepffler,1962197, 1973198 ; Maurin, 1994242 ; Pesme, 2001287 ; Salvidio etal., 1997335, 1998337, 1999336 ; Vandenbrouck, 1996393

Authors: Cheylan M. & M. Delaugerre

110


Pelobates cultripes (Cuvier, 1829)

AMPHIBIA

PELOBATIDAE

Main synonymsNone

French names: Pélobate cultripède, Crapaud à couteauxSpanish name: Sapo de espuelasPortuguese name: Sapo-de-unha-negraEnglish names: Western Spadefoot, Iberian Spadefoot Toad

SubspeciesNone

Description/identification criteria The Western Spadefoot is a fairly large amphibian (10-11 cm)with a squat appearance. The head is short and surmounted bylarge protuberant eyes. The eardrum is not visible. The skin issmooth, 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 canbe immediately recognised by its upright stance. There is littlesexual dimorphism, apart from the presence in the male of a yel-low lens-shaped protuberance on the forelegs. On the heel thereis a down-turned black spade used for digging in the soil. Thetadpoles, which are very large when fully-grown (up to 12 cm)are distinguished by a bulky oval body and a lead-grey to greenishcolouration, with fine golden markings. The caudal crest* is lance-olate shaped, pointed at the tip and practically unmarked. Whenthey leave the water the young measure on average 25 mm andweigh 1.5 to 2 g. The spawn takes the form of a long string, 12to 20 mm in diameter and 1.1 metres long on average. Spawnfrom different females is often mixed up, tangled together at theedge of a water body, usually caught up in vegetation 0 to 20 cmdeep. It may be distinguished from the spawn of “true” toads bythe irregular distribution of the eggs within the string and by thelarger number of eggs.

Similar species Impossible to confuse with other species, except in the case ofthe tadpoles.


Distribution/range

This Spadefoot is found in the Mediterranean regions of southwestEurope: Spain, south and southwest France, as well as in Portugal.In France the populations form two distinct nuclei: a Medi-terranean nucleus which extends eastwards to the Fréjus area inthe Var département and to the Ardèche in the Rhône Valley, andan Atlantic nucleus extending from the mouth of the Gironde tothe 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

Mediterranean arable land, up to 800 metres altitude in southernFrance and 1,700 m in the Iberian Peninsula. In Spain it is com-mon in the dehesa* country typical of southwest Iberia and aroundthe 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


Like most Mediterranean amphibians, the Western Spadefoot isactive especially in winter and spring, from November to May insouthern Spain, from March to June and then from the end ofAugust to November in France. Strictly nocturnal, they are espe-cially active during rain, whenever the temperature reaches atleast 8°C. In the daytime the animal remains buried in the soil ata depth of 6 to 20 cm, up to 40 cm or deeper in winter.

The breeding season varies depending on the local climatic con-ditions. It extends from October to February in the south of theIberian Peninsula and from the end of February to the beginningof May in central Spain. In southern France the breeding cycle ismarkedly bimodal with one more or less consistent breeding periodin autumn (September to November) and another in spring, usu-ally between the third week in February and the end of April,sometimes to the beginning of May. The onset of this event istriggered by continual rain over several days (20 mm), with an airtemperature of 9 to 16°C. The breeding adults arrive at thebreeding sites at dusk and remain under a few centimetres ofwater immediately next to the water’s edge. The largest malesarrive first, sometimes before the females, and remain for one toeight nights. The females usually stay for one to three nights atthe breeding place. During amplexus*, which lasts an average of72 hours, the male and the female both give a call which resem-bles the clucking of a chicken.

In Spain, the average number of eggs laid is 2318, while at a siteat the Ile d’Oléron it is 960 eggs. The strings of eggs are woundaround the vegetation and are laid in shallow water, usually lessthan 20 cm below the surface. The tadpoles feed mostly on plantmaterial, both algae and higher plants, but also detritus. Whenfood becomes scarce they may eat the tadpoles of other speciesor of their own species. Larval development usually takes three

Pelobates cultripes1 cm

111

Amphibians

and a half months for eggs laid in spring, up to eight months forautumn eggs. Metamorphosis takes place mostly in June and July.Some tadpoles do not metamorphose in summer and are still inthe water in autumn. In this case, they probably do not undergometamorphosis 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 maturityappears to develop at two years old. They have many aquaticpredators: dragonfly larvae, water beetles, water snakes, herons.

Phenological cycle


HydrologyThe length of the larval cycle constitutes a major constraint forthis species, particularly in view of the unpredictability of theMediterranean climate. Many breeding attempts are thereforedoomed to failure, whether due to the breeding site drying outearlier than usual or to delayed flooding. Ideally, the site shouldstart to fill with water at the end of October and remain floodeduntil the end of July.

Aquatic habitat It is usually characterised by fairly deep water, most often deeperthan 70 cm. The spawn is laid preferably on submerged vegeta-tion, very close to the water’s edge, in water containing not morethan 5 g of salt per litre (the tadpoles will tolerate slightly highersalinities, up to 10 g.l-1). The breeding sites are typically poor inaquatic vegetation: the highest densities of tadpoles are found insites with hydrophyte biovolumes of less than 10 kg/m3.

Terrestrial habitatThe presence of open areas appears to be a major factor. Otherwisethe 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 sandyareas, the presence of rabbits appears to be favourable for theWestern Spadefoot: it uses their burrows for shelter in the day-time. Grazing, especially by sheep, is also beneficial.



The Spadefoot is currently declining over the whole of its range.This decline is due to the destruction of its breeding sites or ter-restrial habitats, the widespread use of pesticides, and theincreasing 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 fourhistorically known sites still remain, but also in the Bouches-du-Rhône, the Vaucluse and all the Atlantic coastal départements.Most of the breeding sites hold fewer than 100 individuals, a fewtens being the norm. Six Conservatoire de l’Espace Littoral et desRivages lacustres sites and four Nature Reserves support thespecies.In Spain, the situation is a cause for concern in most provinces.The decline seems to have been particularly steep over the courseof recent decades. In many cases, the survival of the speciesdepends on the preservation of a single breeding site.



Anthropogenic factors Impacts of an anthropogenic nature include the destruction ofboth terrestrial habitats (conversion to arable, urban development)and aquatic habitats (infilling of pools), the introduction of com-petitive or predatory alien species (fish, American crayfish, MarshFrogs), the use of toxic chemicals (pesticides) and the destructionof the animals (road traffic). Coastal populations are especiallyaffected by tourism developments and by anti-mosquito treatments.Further inland, it is the abandonment of farmland and the spreadof woodland which are the two principal causes of decline.

Natural factorsThe main threats are due to weather conditions, either when thebreeding site dries out before the tadpoles have metamorphosed,or when a sudden dry period lowers the water level to below thestrings of spawn, which are usually laid at water level at the water’sedge. Some predators (notably Polecats) can have a major impacton the adults during the breeding season.


Arrival

Spawning

metamorphosis

Occasional cycle

Usual cycle

Larval development and

Arrival

Spawning

Larval development and metamorphosis

Departure, aestivation and terrestrial phase

��

Southern France



-Annexe IIAnnexe IV- France, national list: Decree of 22/07/1993/Journal Officiel09/09/1993- 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

112


Risks relating to populationsEssentially, it is necessary to maintain high enough numbers toovercome stochastic risks, as well as a dense enough network ofconnections between populations to ensure interchanges of indi-viduals and hence gene flow. Distances of less than a kilometrebetween sites would appear to be desirable.


Current measures In France, if some of this species’habitats are situated in pro-tected areas, the species is not the subject of specific conserva-tion measures, apart from the monitoring protocols set up at somesites in Charente-Maritime, in the Camargue (Tour du Valat estate),at the Valliguières site in the Gard département, in the Roque-Haute Nature Reserve in the Hérault, in the Parc du Lubéron inthe Vaucluse and at the Suze-la-Rousse site in the Drôme.

RecommendationsIt would be helpful, in the first place, to maintain the spatialcohesion of the breeding sites, if necessary by recreating suitablesites in recently abandoned areas. This could be attempted expe-rimentally at a few sites: the Giens peninsula and the Plaine dePalayson in the Var, the Vallée du Calavon in the Vaucluse, thePlateau d’Aumelas in the Hérault.In addition, it would be useful to carry out some activities at cer-tain breeding sites: providing information for owners and communes,

setting up management agreements, etc. As far as management isconcerned, the most important actions are the following: to con-trol the introduction of undesirable species (Louisiana Crayfish,fish, terrapins etc.), to ensure that the hydroperiod* is compatiblewith the species’ breeding cycle, to control plant cover over anarea of radius 100 to 200 m by grazing and/or scrub clearing, andto place shelters by the edges of the pools to promote the survivalof the newly metamorphosed young.

Bibliography

Alvarez et al., 19908 ; Busack & Zug, 197657 ; Cabral et al., 199959 ;Cei & Crespo, 197165 ; Cheylan & Poitevin, 199974 ; Conservatoirede 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.

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Amphibians

Triturus cristatus (Laurenti, 1768)

AMPHIBIA

SALAMANDRIDAE

Main synonymsNone

French name: Triton crêtéItalian name: Tritone crestatoEnglish names: Great crested Newt, Warty Newt

SubspeciesNone

Description/identification criteria Large newt (total length of adults: 19 cm), dark brown or almostblack in colour, finely spotted with white on the flanks. The under-side is yellow/orange, marked with black. The skin is very charac-teristically warty. During the breeding season, the male developsa high, spiky dorsal and caudal crest. This has a silvery streakalong 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, Tritu-rus blasii) is a hybrid between the Great Crested and MarbledNewts. It is distinguished from the Great Crested by its ventralsurface, which is greenish black above and yellowish with blackmarkings and fine white spots below.• The range of the Italian Crested Newt (T. carnifex) is almostcompletely separated from that of the Great Crested Newt. Its feetare longer, its body is more elongated and its upperside is paler,showing many black marks; the male’s crest is lower and lessdeeply indented.


Distribution/range

FranceThis species is found throughout the northern half of France,north of a line from Charentes to Savoie. In the southern half ofFrance it is currently known only from a few sites: one at Arles,Bouches-du-Rhône, in the Gard51, two in the Drôme and one inthe southern Ardèche.

Other countriesThe range extends from Great Britain in the west to the Ural Moun-tains in Russia and, north-south, from southern Scandinavia toAustria and northern Romania. This species is absent from Spainand Portugal.In the southern Mediterranean countries, it is replaced by twovicariant species formerly treated as subspecies of Triturus crista-tus:• T. carnifex (Laurenti, 1768) in Italy and the Balkans;• T. karelinii (Strauch, 1870) by the northwest coasts of theAegean Sea (Turkey, Greece).

Habitat

General descriptionThe Great Crested Newt is a species of lowlands which lives mainlyin fish-free permanent pools and/or temporary pools which dryout in summer. The three sites in the Mediterranean region take the form of:• a marlpit in a calcareous area covered with pine woods, gar-rigue 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-upareas and degraded (Trinquetaille, at Arles).

Ecology


In the Mediterranean region, the only information concerningthe species’ life cycle comes from the Etang de Valliguières. Atthis site the aquatic phase is variable in length between years,depending on the flooding period. In “normal” conditions, thenewts arrive at the pool in November/December and leave againaround May. The length of time spent in the water is about 80-90 days33. The adults feed mostly on aquatic and terrestrial insects19. Thelarvae feed on aquatic prey (especially microcrustaceans).

Breeding begins in April-May. The female lays 200 to 500 eggsper year248, placed one by one underneath the leaves of sub-merged plants which she then folds up with her hind feet (mainlyspecies with broad leaves such as Ranunculus gr. peltatus, Gly-ceria fluitans, etc.). Embryonic development takes from two tothree weeks depending on the water temperature. It is followedby a two- or three-month larval stage which ends with meta-morphosis, 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 damprefuge in the surrounding area, where they hide and feed. Aftertwo years, in optimal conditions, they attain sexual maturity(males usually one year earlier than females140, and return to thewater to breed. Immature T. cristatus tend to appear in the poollater than the adults. Adult survival is fairly high247: between 60and 75% depending on the year. The maximum lifespan in thewild is from 14 to 18 years.

Triturus cristatus1 cm

male

114


The Great Crested Newt is known for its fairly strong fidelity tobreeding sites, local variations in breeding success being com-pensated for by its long life expectancy. The newts do not appearto colonise new breeding sites more than 400 m away from theirnatal pool20. Juveniles may disperse for up to a kilometre15, 202.

During the terrestrial phase, the individuals hide underneathstones, under pieces of wood or in crevices in the ground, a fewhundred metres, or further, from the breeding sites. Fidelity toterrestrial sites appears to be linked to the different “scent com-plexes” which distinguish them, orientation being carried out bymeans of olfaction170.

Phenological cycle


HydrologySuccessful breeding depends on the presence of water up to theend of July, which is essential for the metamorphosis of the juve-niles.

Aquatic habitat The site must be relatively large and deep (>1 m) open to the sunand lacking fish. The presence of macrophytes* is essential foregg laying and for the development of an abundant invertebratefauna; however, vegetation should not cover more than 20% ofthe water surface (maximum tolerated by breeding individuals)88.

Terrestrial habitat The presence of wooded areas immediately adjacent to the siteappears to be necessary for the survival of adults (and newlyemerged young) during the terrestrial phase188. The presence ofnatural refuges (logpiles, piles of stones, stone walls) is also afavourable factor. Ditches and hedges provide biological corridorswhich facilitate dispersal.



Due to their isolation, the populations of Mediterranean south-ern France are highly endangered in the short term. The site atTrinquetaille (Bouches-du-Rhône) is situated within very degradedsemi-urban surroundings. The population at Valliguières (Gard) has

been subject to conservation measures for three years but its iso-lation, low numbers (about 200 individuals) and irregular breed-ing mean that its chances of survival in the medium to long termare questionable. The population at Peyreguil (Gard) is still poorlyknown. The presence of fish here constitutes a serious threat.



Anthropogenic factors The destruction of breeding sites (filling-in of pools), changes tothe farmed environment resulting from new management prac-tices, the isolation of different populations through habitatfragmentation203, and the introduction of competitive alien species(Marsh Frogs, fish, American crayfish) are the main causes ofdecline.

Natural factorsThe repeated drying-out of breeding sites may eventuallythreaten the survival of the population161; the effect of wild boarduring the emergence of the young newts also represents athreat.


Current measures The population of the Etang de Valliguières has been monitoredsince 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 floodingand thereby to encourage more regular breeding.• The creation of an artificial pool close to the existing site, inorder firstly to encourage more regular breeding and secondly tocreate a second breeding site with the aim of improving theprobability 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 speciesin the Mediterranean region.• Establish a network of potential breeding places in close prox-imity 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 maybe considered: installing electric fences around all or part of thesite during the period when the young are emerging, digging outa substitute pool, planting bramble bushes around the pool toincrease the number of refuges for the young newts.


Arrival and

Eggs laying

Larval growth

overwintering (aquatic phase)?

Larval metamorphosis

(juveniles, adults)

Departure, terrestrial phase and aestivation

��

Southern France



-Annexe IIAnnexes II and IV- France, national list: Decreeof 22/07/1993/Journal Officiel09/09/1993- France, national Red DataBook: “vulnerable”242

115

Amphibians

• Urgently introduce conservation measures at the Trinquetaille/Arles site in the Bouches du Rhône (translocation of individualsto a more suitable site could be considered).

Bibliography

Arntzen & Teunis, 199315 ; Avery, 196819 ; Baker & Halliday,199920 ; Besnard, 200233 ; Brogard et al., 199651 ; Cooke et al.,

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.

Authors: Lombardini K. & M. Cheylan

116


Triturus marmoratus Latreille 1800

AMPHIBIA

SALAMANDRIDAE

Main synonymsNone

French name: Triton marbréSpanish name: Triton jaspeadoPortuguese names: Tritão-marmorado, Tritão-verdeEnglish name: Marbled Newt

Subspecies T. m. pygmaeus in the Iberian Peninsula, recently considered to bea species in its own right.


Large newt (total adult length: 16 cm) black to brown in colourwith green marbling on the back. Females and juveniles have anorange-red line down the spine. In the breeding season, the maledevelops a smooth crest with vertical black and yellowish-whitebars. The underside is black, dotted with small white spots(orange-yellow with large black markings in the Great CrestedNewt).

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 markedwith black. Blasius’ Newt is a hybrid between T. marmoratus andT. cristatus. It may show morphological characters of both species,but it is completely absent from the Mediterranean region. ThePygmy Marbled Newt, an endemic species of southern Spain, hasa black underside like the Marbled Newt, but is smaller.


Distribution/range

FranceThis species occurs in the western half of the country, as farnorth as the Seine. The eastern limit of its range lies in the Som-mières area, in the Gard département.

Other Mediterranean countriesIberian Peninsula, north of the river Tagus, from the coast up to2,100 m in the central mountains.

Habitat

The Marbled Newt, during its aquatic phase, is found in smallbodies 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 grassyareas (Causses) and woodland.

Ecology

Biological characteristicsLike almost all the European amphibians the Marbled Newt hasa bimodal cycle, with a terrestrial phase and an aquatic phase. At all stages of development, they are carnivorous. The larvaefeed mostly on aquatic invertebrates405, the adults and juvenilesare 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 win-ter owing to the unfavourable condition of the terrestrial habitatin summer. In southern France the period for which they are inthe water extends from mid-November to mid-June, with con-siderable 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 slightlyvariable depending on interannual variations in the floodingperiod. Larval development takes a relatively long time (two tothree months140). A single female may take several weeks to layall her eggs. The eggs are laid one by one, in a fold in a leaf of anaquatic plant, with a preference for species with broad, flexibleleaves such as Mentha pulegium (Pennyroyal), Ranunculus spp. orCallitriche sp.9.

The terrestrial habitat in evidently very important for the aesti-vation of amphibians in the Mediterranean as the animals runthe risk of desiccation. Juveniles and adults hide under stones oramong the roots of bushes. They do not move far, travellingpartly underground during the summer drought.

Phenological cycle


Arrival and feeding

Egg laying

Larval development

Hibernation

Larval metamorphosis

(juveniles, adults)

(juveniles, adults)

Departure, then terrestrial phase and aestivation

��

Southern France

Triturus marmoratus1 cm

male

117

Amphibians

Environmental conditions183, 184, 405

HydrologyBreeding sites must be holding water by February at the latest toallow the eggs to be laid. A flooding period of five months is thennecessary for completion of larval development and successfulbreeding. Breeding failure in a given year does not constitute amajor problem for the population, unless it recurs too frequently.

Aquatic habitat The water should be of a reasonable depth (about 0.8 m) andprovide sufficient growth of aquatic vegetation to support thedevelopment of sufficient invertebrates for the feeding require-ments of the larvae and adults.

Terrestrial habitat It may be very varied in character (open heathland, garrigue, oakor pine woodland, arable or even built-up areas). Hedges and copsesaround the edge of breeding sites provide important terrestrialhabitats and migration routes135.



In Spain there are large populations which are relatively stable,apart from in the Ebro Valley and in north Catalonia where itappears to be more threatened. In France, its status gives no causefor concern although some local disappearances have beenrecorded, due mainly to the filling-in of pools or to land consoli-dation (destruction of hedges).It occurs at about 13% of Réserves Naturelles and 3% of RéservesNaturelles Volontaires. However, if only those Réserves Naturelleslocated within the newt’s French range are included, it occursin 42%182.



Anthropogenic factors Impacts of an anthropogenic nature include the destruction ofhabitats, 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 destructionof the animals (road traffic).

Natural factorsToo short a period of flooding (i.e. through silting up) may pre-vent the completion of larval development, which will jeopardisethe presence of the newt at a site.

Risks relating to populationsThe destruction of breeding sites and terrestrial habitats bringsthe risk of extinction by reducing the interconnections betweenpopulations.


Current measures No specific measures are currently being applied at the Roque-Haute Nature Reserve (Hérault, France). The effect of scrub clearingaround the pools (terrestrial vegetation) on the survival of adultsand their migration routes remains to be assessed.

Recommendations• Maintain the cohesion of populations at the local and regionalscales.• Control the introduction of fish and American crayfish intobreeding sites.• Maintain well-developed aquatic vegetation.• Do not burn the area around pools or the sites used as refugesin the terrestrial phase. Dry years, when breeding does not takeplace, do not present real problems for this long-lived species, aslong as the aestivation refuges enable good numbers of adults tosurvive. The maintenance of vegetation corridors around thepools 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 ; Zuider-wijk, 1997405.

Author: Jakob C.Collaborator: Cheylan M.



-Annexe IIIAnnexe IV- France, national list: Decree of 22/07/1993 Journal Officiel09/09/1993- Spain, national list293

- Portugal, national list59

- France, national Red DataBook: “vulnerable”242

- Portugal, national Red DataBook: “not threatened” (NT)59

- Spain, national Red DataBook: “low concern” (LC)293

118

Glossary

Actinomorphic: (= regular) describes a flower, or a group ofidentical 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 lime-stone ions thus released are capable of being absorbed by plants. Aggradation: Situation resulting from gradual infilling by accu-mulated material (soil, silt, sand, gravel, etc.).Allogamous: having a mode of sexual reproduction involvingcross-fertilization among plants.Amplexus: among amphibians, mating without internal fertilisa-tion. The eggs and sperm are emitted simultaneously and fertili-sation is external. Depending on the species, amplexus may belumbar (the male clasps the female around the lower part of theabdomen) or axillary (the male clasps the female under thearmpits).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, inalgae, mosses, liverworts and ferns. Apoendemic: describes a new plant taxon which has been pro-duced in a given region by polyploidisation (by becoming poly-ploid*) from diploid taxa* which are more or less widelydistributed 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 thebase of a leaf, petiole, ligule, etc.). Bases: all the exchangeable positive ions in the soil which raisethe pH*.Basionym: original name under which a species was firstdescribed. 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 varioustypes of continental environment, either at the scale of naturalregions of limited area or at the scale of stations occupying smallareas. Biomass: total mass of living material, animal and vegetable, ina defined biotope at a given time. Bipinnatifid: used of a pinnatifid* leaf whose divisions are them-selves pinnatifid.Bract: more or less modified leaf situated in the axil of a floweror an inflorescence.Bracteole: small bract situated in the axil of the ray of abranched inflorescence or in the axil of each of its constituentflowers.Brood pouch: see ovigerBryophytes: group comprising both mosses and liverworts.Capsule: in mosses and liverworts, designates a hollow dry-walledorgan of the sporophyte, in which the spores develop. Amonghigher plants, designates a dry dehiscent fruit (i.e. opening spon-taneously when ripe) containing more than one seed.Caudal crest: membrane on both sides of the tail in tadpoles (inthe Anura) and larvae (in Urodela) used for propulsion. In somespecies, this crest extends up the back to the back of the head(Tree Frog tadpoles for example).

Cercopods: (= cerci) in arthropods and crustaceans, appendageslocated on the last abdominal segment (usually two in number,one on each side).Chamaeophyte: (= chameophyte) low woody plant whose budsare 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 thethallus and by the highly complex structure of the reproductiveorgans (antheridia and oogonia).Chitinous: formed of chitin, a molecule involved in the structureof the arthropod cuticle.Climax: used of a plant community which has developed to asustainable state of equilibrium with the climatic and edaphicconditions of the environment, in the absence of human inter-vention.Clonal: used of an organism (animal or plant) derived from a sin-gle cell or a single individual, by asexual reproduction (known asvegetative propagation* in plants)Cohort: set of individuals which have experienced the sameevent at the same time (individuals born at the same time orbreeding at the same time in pools, for example).Connectivity: facilitation of the movement of individuals of aspecies between local sub-populations to form a single func-tional demographic unit.Cordate: describes a heart-shaped flat organ.Coronule: short apical cells which develop a crown shape at thetip of the oogonium* in the Characeae. Cupular pool: small temporary pool located in bowl created inrock by natural erosion (see volume 1, chapter 2a). Cyme: branched inflorescence in which each flower is located atthe end of a branch. Daya: a temporary pool in Morocco.Dehesa: Spanish name used to describe a landscape specific tocertain regions in the centre and south of Spain, consisting ofpasture or cereal cultivation scattered with evergreen oaks orCork Oaks, which are usually cut.Detrivore (or Détritivorous): feeding on dead organic material.Diapause: period during which metabolic activity and the devel-opment of an insect is suspended at a particular stage (egg, larva,nymph or adult), as a result of the action of internal or externalfactors. Diploid: describes an organism whose cell nuclei have a doubleset of chromosomes (2n). Echinulate: covered in small spines.Ecophase: during its life cycle, a species passes through differ-ent stages (egg, larva, juvenile, etc.). An ecophase corresponds toone of these stages having a different ecology from the otherparts of the cycle.Edaphic: refers to ecological factors linked with the soil as wellas to their relationships with plants. Elater: elongated cell attached to a spore, and capable of flexi-ble movements.Endemic: used of a species exclusively confined to a given bio-geographical area, often of limited extent.Espace naturel sensible (Sensitive Natural Areas): areas boughtby each département in France since 1985 “to preserve the qualityof 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 withmineral nutrients (phosphates, nitrates) often accompanied by a

119

Glossary

proliferation of algae and/or aquatic higher plants, leading tooxygen depletion of the deeper water as well as a build-up oforganic matter. Fistulous: hollow and cylinder-shaped. Gametangium: organ of the gametophyte* in which thegametes are formed.Gametophyte: in the life cycle of algae, mosses, liverworts andferns, designates the haploid* phase of the organism, which pro-duces gametes; in higher plants refers to the haploid organwhich produces the gametes.Genetic bottleneck: sudden decrease in the size of a populationassociated with a decrease in the total genetic variability.Genetic bottleneck: sudden decrease in the size of a populationassociated with a decrease in the total genetic variability. Genetic drift: within small populations, chance fluctuations ingene frequency with each generation, enabling rare genes (whichwould be eliminated by selective pressure within large popula-tions) to be continuously expressed.Geophyte: plant species which withstand the unfavourable sea-son thanks to the presence of bulbs, rhizomes or any other typeof underground reserve organ (see Figure). Gravel bed/Gravels: loose sedimentary formation consisting ofrock fragments. Gyrogonites: calcified female fructifications produced by charo-phytes, corresponding to fossil or living forms after dispersal.They are invariably made up of five cells in the form of left-handed spirals, joined at the tips.Halotolerant: tolerant of a certain amount of salt in the envi-ronment (water, soil, etc.).Haploid: organism or organ whose cell nuclei each contain aset of n chromosomes, comprising a single copy of each chro-mosome.Heliophilous: used of a plant which grows in conditions of strongsunlight.Helophyte: marsh plant whose budding parts, which enable it tosurvive during the bad season, are laid down in the sediment,while in the good season they develop an aerial structure whichextends above the water surface (Reed, for example).Hemicryptophyte: perennial herbaceous plant whose buddingparts, which enable it to survive during the bad season, remainon the surface of the soil, at the very base of the stems (or of thetuft 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 ofwhose constituent equivalent parts (petals, sepals, stamens, etc.)is six or a multiple of six. Hexaploid: said of an organism whose cell nuclei contain threepairs of each homologous chromosome.Hybridogenic: taxon derived from a cross between two or sev-eral different taxa usually different species or sub-species.Hydroperiod: period during the year when the pool containswater.Hydrophyte: plant that lives in an aquatic environment (WaterMilfoils, Water Lilies).Hydrophytic: refers to a hydrophyte* plant.Hygrophilous: organisms dependent on biotopes characterisedby high soil water content.Karstic: relating to karst (the whole of the surface and subter-ranean structure of a limestone massif, which results from thedissolution of rock and is characterised by a wide-ranging systemof underground water flow).

Laune (or lône): a local term meaning a former branch of awatercourse which can reconnect with the present watercoursewhen water levels are medium or high. Lavogne: local term meaning depressions dug out for wateringlivestock, in the Causses of the Cevennes and the Uzégeois andMontpellierais garrigues (France). Life form: five life forms (see Figure) were defined by Raunkiaerin his ecological classification of plants based on their strategyfor surviving through the unfavourable season. Each life form isdefined by the position of the buds in the architecture of theplant, 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 visibleto the naked eye.Matorral: word of Spanish origin, in areas with a Mediterraneanclimate usually applied to a bushy vegetation community, adaptedto drought and often dominated by evergreen species, often withsmall leaves.Meso-eutrophic: see Mesotrophic*.Mesotrophic: intermediate between eutrophic* and oligotrophic*. Metanauplius: larval stage following the nauplius* stage incrustaceans, characterised by the beginnings of body segmenta-tion. Several moults, over a few weeks or a few months, are thennecessary to attain the adult stage. Metapopulation: set of populations that are interconnected viamigration events (gene flow) and are subject to extinction andrecolonisation. This concept may be extended to include any setof populations developing in a more or less independent waywhich are, however, interconnected through rare instances ofmigration.Metathorax: posterior part of the thorax in insects, to which areattached the last pair of legs (and sometimes the second pair ofwings). Microphagous: feeding on small prey.Mucronate: terminating abruptly in a sharp point.Natura 2000: ongoing European policy, with the aim (throughthe implementation of the Habitats Directive119 and of theCouncil Directive 79/409/EEC of 2 April 1979 on the conservationof wild birds) of reconciling human activities and natural habi-tats within a network of sites having great ecological interest,some designated for their diversity of birds (Special ProtectionAreas) others for the rarity of their fauna, their flora and theirnatural habitats (Special Conservation Areas). Nauplius: first larval stage of crustaceans, followed by themetanauplius* 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. Itmay be external (carried on the body) and/or internal (absorptionand passage through the digestive tract). Pappus: plume of hairs crowning an achene.Parotoid glands: glands located at the back of the head amongsome species of amphibians (true toads, salamanders), producinga poisonous secretion as a deterrent against predators.Perennial: in botany, describes a plant species which lives formore than one year (as opposed to annual). Phenology: description of the various phases in the life cycle ofa species. Phytocenosis: collection of plant species having a homogenousappearance and colonising the same habitat (syn.: plant community).

120


Phytoplanktonophagous: feeding on phytoplankton.Phytosociology: see volume 1, chapter 2, box n° 4.Pinnatilobed: used of a divided leaf when the divisions cut intothe lamina for less than half its width. Pinnatipartite: used of a pinnate leaf when the divisions cut intothe lamina for more than half its width. Pinnatisect: used of a pinnate leaf when the divisions cut com-pletely 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 atthe ends of the gametophyte* stems.Polje: karstic* plain or subsidence valley, resulting from the coa-lescence of several karstic erosion systems (“ouvalas”) and coveredin decalcified clays. A polje is subject to more or less prolongedseasonal flooding which sometimes occurs suddenly, dependingon the underground karstic network. Polyploid: condition of an organism whose cells contain morethan two sets of homologous chromosomes, i.e. more than 2nchromosomes.Pozzines: high altitude swards dominated by Carex, growing ona peaty substrate and mostly dotted with pools (Corsica, Pyrénées,Sierra Nevada, etc.).Pre-imaginal stage: final larval stage before the adult stage, ininsects.

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, veryshort, 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 peb-bles bound together with a limestone or sandstone cement.Recruitment: the adding of new individuals to a population. Recruit-ment 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 ofcertain cells, in particular in bushes, trees and shrubs, making itrigid 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 orpeduncle. Zoological: microorganism attached to a support(stem, rock, etc.). Sex-ratio: ratio of the numbers of male and female individualsin a defined population.

Annual plants Perenialplants

**: Raunkiaer, C, 1934. The life form of plants. Oxford, Clarendon Press. 632 pages

Phanerophytes Chamaeophytes Hemicryptophyte CryptophytesGeophytes with bulb

CryptophytesGéophytes

with rhizome

Therophytes (surviving by seeds)

Figure. The main biological types, according to Raunkiaer**

Pistacialentiscus

Artemisia molinieri

Eryngium pusillum

Ophioglossum lusitanicum

Isoetes histrix

Ranunculusrevelieri

In perennial plants, position of the budssurviving the bad season

seeds

N. B

eck

121

Glossary

Sheath: a leaf base expanded to form a more or less cylindricalenvelope, split longitudinally or not and clasping the stem. Sorus: group of sporangia*, arranged in a distinctive pattern onthe lower surface of the fronds of some ferns. Spiracle: in tadpoles, opening which enables water to be ejectedduring respiration. In tadpoles of Painted Frogs, Midwife Toadsand Yellow-bellied Toads, it is located in a median position onthe underbelly. In other European amphibians it is situated on thesides of the abdomen.Sporocarp: in some ferns (Marsilea, Pilularia, etc.), a particulartype of sorus*, closed and with tough walls, enclosing the spores.Sporophyll: in ferns, specialised leaf or frond, bearing one ormore sporangia.Station (of plants): homogenous area of land where a plantspecies (or a particular plant community) is found.Stylopodium: small fleshy nectariferous disc which surmountsthe fruit in Apiaceae (= Umbellifers), bearing two separate styles.Sympodic: describes the parts of a plant resulting from sympo-dial growth, i.e. lengthening of the stem in a succession of seg-ments produced by lateral budding of the stems which haveflowered in the previous year.Terra Rossa: type of soil found in Mediterranean regions, derivedfrom the breakdown of limestones and characterised by an accu-mulation of ferric oxides.

Therophyte: synonym for an annual plant, a herbaceous plantwith a very short reproductive cycle, lasting a few months or incertain cases a few weeks, which survives the bad season in theform of seeds. (see Figure).Therophytic: describes a therophyte*.Trophic: everything relating to nutrition among plants and ani-mals.Vegetative reproduction: mode of reproduction of a plantspecies using vegetative organs (stolons, rhizomes, tubers, etc.).Vegetative stage: part of the life cycle of plants relating to ger-mination, growth and propagation (vegetative reproduction),excluding flowering and fruiting.Verticil (whorl): all the homologous organs arising from a stemat the same point and radiating out from the stem.Vicariant: used of animal or plant species that are taxonomicallyclosely related and which inhabit environments with similar eco-logical characteristics in different geographical regions. ZNIEFF: (Zone naturelle d’intérêt écologique floristique et fau-nistique) one of a list (begun in 1982 by the MEDD) of Frenchnatural areas, both terrestrial and marine, whose interest lieseither in the stability and the richness of the ecosystem or in thepresence of rare and threatened animals and plants.

122

Macrocrustacean inventory form

1. Essential data A) Source

Observer or code:Surname, first name:Adress:

Name of form compiler if different from observer:

Biblio. ref. or code:Author(s):Date:Title:

Periodical:Volume, number, page:

Coll. ref. or code:Where deposited:Inventory number:

Name of form compiler if different from observer:

or or

B) TaxonNumber of refe-rence list:

Genus Species Subspecies or variety

Taxon

or

determined by:

verified by:v.c.

C) PlaceCoordinates– in grades (Paris meridian)

Longitude: � E, �W

Latitude:

or coordinates– in degrees (Greenwich meridian)

Longitude: � E, �W

Latitude

Commune + locality:

INSEE code:

IGN map number:

or and and

° ‘ ‘’

° ‘ ‘’

Altitude:

At:

Name of pool or water body:

D) Date

hr 1 hr 2 day month year

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

2. Additional dataBiotopes:

� clay� sand � rock surface� clear water� turbid water� grassy ground� bare soil� algae� vegetation beds� ..............................

� Isoetes� Ranunculus� ....................

Method of collection:

� by hand� shrimp net� drag net� plankton net� bird stomach� amphibian stomach� fish stomach� ..............................

� salt marsh� salt water� fresh water

Depth of biotope:

Duration of flooding:

Colour on the living creature:

Systematic notes, associated fauna, biometry, etc.:

3. Additional information (in note form to aid validation)

Use reverse side of the form if necessary

metres

Please send to: Danielle DEFAYE, Muséum National d’Histoire Naturel, Laboratoire de Zoologie - Arthropodes (Crustacés), 61, rue Buffon, F-75005 Paris

123

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