Aegilops (Poaceae) in Italy: taxonomy, geographical distribution, ecology, vulnerability and conservation.

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Aegilops (Poaceae) in Italy: taxonomy, geographicaldistribution, ecology, vulnerability and conservationEnrico V. Perrinoab, Robert P. Wagensommera & Pietro Medaglica Botanical Garden Museum, University of Bari, via E. Orabona 4, 70126 Bari, Italyb Agronomic Mediterranean Institute, via Ceglie 9, 70010 Valenzano (Bari), Italyc Department of Biological and Environmental Science and Technologies, University ofSalento, Via Monteroni 165, 73100 Lecce, ItalyPublished online: 14 May 2014.

To cite this article: Enrico V. Perrino, Robert P. Wagensommer & Pietro Medagli (2014) Aegilops (Poaceae) in Italy: taxonomy,geographical distribution, ecology, vulnerability and conservation, Systematics and Biodiversity, 12:3, 331-349, DOI:10.1080/14772000.2014.909543

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Research Article

Aegilops (Poaceae) in Italy: taxonomy, geographical distribution,ecology, vulnerability and conservation

ENRICO V. PERRINO1,2, ROBERT P. WAGENSOMMER1 & PIETRO MEDAGLI3

1Botanical Garden Museum, University of Bari, via E. Orabona 4, 70126 Bari, Italy2Agronomic Mediterranean Institute, via Ceglie 9, 70010 Valenzano (Bari), Italy3Department of Biological and Environmental Science and Technologies, University of Salento, Via Monteroni 165, 73100 Lecce, Italy

(Received 26 November 2013; revised 18 March 2014; accepted 20 March 2014)

Aegilops L. includes wild species from which, over millennia, man has cultivated forms of Triticum L. Ten species ofAegilops occur in Italy. Three species are allochthonous and eight out of ten species are recorded in the Apulia region. Fiveout of the ten species have been included in Red Lists. Each taxon is presented and discussed, citing old and new sites ofoccurrence, by examining specimens from many different herbaria, and describing their ecology and habitats, according tothe Directive 92/43 EEC. A new taxonomic key, for the identification of all Aegilops species growing in Italy, is provided.The occurrence of Aegilops caudata L., A. peregrina (Hack. in J. Fraser) Maire & Weiller and A. speltoides Tausch in Italyis doubtful.

Key words: Aegilops, ecology, identification key, Italy, Red List

IntroductionAegilops L. has been intensively studied because of its

close relationship with cultivated wheats. The green revo-

lution caused wheat genetic erosion and for this reason

wheat improvement programmes required the use of

germplasm collections with the greatest diversity. Landra-

ces and wild relatives attracted increasing attention,

because they can provide characters related to adaptation

(Hu et al., 2012). This, in turn, led to the need for a com-

prehensive collection of species of Aegilops s.l., espe-

cially for those at risk of extinction (Hammer, 1980).

Assembling this worldwide collection of germplasm

resulted in a re-evaluation of the relationship between

Aegilops and Triticum, which is a subject of intense

debate. The separate status of the two genera is advocated

mainly for practical reasons, for which the correct names

under both genera are usually given by most of the authors

(van Slageren, 1993).

Linnaeus (1753) enumerated five species: four of these

still belong to Aegilops with A. ovata L., A. caudata L.

and A. triuncialis L. still with their original names, while

A. squarrosa L. refers actually to A. tauschii Coss.

(Bowden, 1966). The fifth species, A. incurva L., is now

referred to Parapholis incurva (L.) C.E. Hubbard. Today,

after much work (Jaubert & Spach, 1850�51; Boissier,

1884; Zhukovsky, 1928; Eig, 1929; Hammer, 1980; van

Slageren, 1994) the number of recognized species is 23

for Aegilops and one for Amblyopyrum.

The phylogeny and monophyly of Aegilops, on a

genetic basis, has also been discussed widely in several

papers (Kimber & Zhao, 1983; Hsiao et al., 1995, Kellogg

& Appels, 1995; Kellogg et al., 1996; Seberg & Frederik-

sen, 2001; Petersen & Seberg, 2002; Vakhitov et al.,

2003; Mason-Gamer, 2005; Petersen et al., 2006;

Alnaddaf et al., 2012). The common conclusion based on

phylogenetic analysis is that Aegilops would be stable if

A. speltoides Tausch, with B genome, was removed from

A. and transferred to the genus Sitopsis (Jaub. & Spach)�A. L€ove (L€ove, 1984; Petersen et al., 2006). On this basis,

Aegilops may be considered a monophyletic genus, which

is not to be included in Triticum (Mason-Gamer &

Kellogg, 1996; Sallares & Brown, 2004; Bordbar et al.,

2011). The last revision of van Slageren (1994) reported

the correct naming and typification of 27 accepted taxa of

Aegilops and two taxa of Amblyopyrum.

Excluding the USA and China, the genus Aegilops

occurs from about 10� W to 82� E and about 24� S to

47� N (van Slageren, 1994). The altitude distribution isCorrespondence to: Enrico V. Perrino. E-mail: [email protected] & [email protected]

ISSN 1477-2000 print / 1478-0933 online

� The Trustees of the Natural History Museum, London 2014. All Rights Reserved.

http://dx.doi.org/10.1080/14772000.2014.909543

Systematics and Biodiversity (2014), 12(3): 331�349

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from �400 m b.s.l. (Dead Sea area) to 2700 m a.s.l., with

a great variation among species. The genus may be char-

acterized as a Mediterranean-western Asian element

(Hegde et al., 2002), and it occurs in the following

regions: Central and Mediteranean Europe, Southern

Ukraine, the Crimea and the Ciscaucasus and Transcauca-

sus; for Africa, only in the north of Sahara and in the

Canary Islands; and for Asia (Western and Central part)

in the region bordered by the deserts of the Arabian penin-

sula in the south and by the Tian Shan mountains and the

Himalayas in the east. The centre of diversity is the Fertile

Crescent region of Palestine, Lebanon, South-Eastern Tur-

key, Syria, Northern Iraq, North-Western Iran, with abun-

dant occurrence also in Transcaucasus. None of the

species appears to be endangered, not even Aegilops shar-

onensis, which only occurs on the coastal plain of Pales-

tine and in southern Lebanon (Kimber & Feldman, 1987).

Probably only the European Aegilops uniaristata Vis. and

both varieties of Aegilops comosa Sibth. & Sm. might

become endangered as a result of their limited distribu-

tion, grazing and tourism.

The geographical distribution was outlined in detail by

Eig (1936). Since then, there has been a great deal of col-

lection activity for herbarium specimens and, more

recently, for threatened germplasm. According to the

most updated overview of the current distribution of the

taxa by van Slageren (1993) and others (Miller, 1987; van

Slageren, 1994; Feldman et al., 1995; Heywood &

Zohary, 1995), nine species of Aegilops are native to

southern Europe, which includes Italy. These nine species

are more genetically distant from Aegilops species consid-

ered genetically closer to Triticum L., but yet their gene

pools can be used in wheat breeding.

In summary, the genus Aegilops belongs to the tribe

Triticeae, subfamily Pooideae and family Poaceae

(Cenkci et al., 2008; Kilian et al., 2011), which is one of

the most intensively investigated group of plants (Miller,

1987). On a world scale, Aegilops includes 23 wild annual

species, 11 of which are diploids and 12 allopolyploids

(Hammer, 1980; Kilian et al., 2011). The revision of the

genus Aegilops with regards to its genome and taxonomy

results in a total of 27 specific and intraspecific taxa (van

Slageren, 1994; �Cerepanov, 1996). In the flora of Europe,

Aegilops is represented by 11 species (Tutin &

Humphries, 1980), which become 13 (A. kotschyi Boiss.

and A. peregrina (Hack.) Maire & Weiller) by adding

those reported by van Slageren (1994). Italy is the most

important continental country with 10 of the 13 European

species (A. biuncialis Vis., A. caudata L., A. cylindrica

Host, A. geniculata Roth, A. neglecta Req., A. peregrina

(Hack.) Maire & Weiller, A. speltoides Tausch, A.

triuncialis L., A. uniaristata Vis. and A. ventricosa

Tausch) (Pignatti, 1982; Conti et al., 2005), and six of

them (A. biuncialis, A. cylindrica, A. neglecta, A. triuncia-

lis, A. uniaristata and A. ventricosa) are included in the

IUCN categories. Recently A. fragilis Parl. has been cor-

rectly included with A. ventricosa (Conti et al., 2005),

since all characters of A. fragilis described by Pignatti

(1982) fall within the description of A. ventricosa (van

Slageren, 1994).

Considering the practical importance of Aegilops, the

aim of the present work was to update and harmonize

the knowledge of species occurring in Italy, about which

the authors found gaps in geographical distribution, taxon-

omy and, to some extent, in ecology, genetic erosion,

extinction and conservation status.

Materials and methodsIdentification and nomenclature was performed following

Pignatti (1982) and using the monograph of van Slageren

(1994). The geographical distribution and altitudinal range

was updated by: (i) checking the sites of the old official

herbarium specimens; (ii) indications provided by Pignatti

(1982), van Slageren (1994) and Conti et al. (2005);

(iii) other local and national studies; (iv) grey (unpub-

lished) literature; and (v) new data provided by the team.

The results refer to 254 herbarium specimens deposited in

27 herbaria, 223 of which were directly observed and con-

sulted on-line (Appendices 1 and 2, see supplementary

material, which is available on the Supplementary tab of

the article’s Informaworld page at http://dx.doi.org/

10.1080/14772000.2014.909543), and 31 collected by the

authors during 2009�2013, in Apulia, Basilicata and

Sicily. The new herbarium specimens (A. biuncialis, A.

geniculata, A. neglecta, A. triuncialis, A. uniaristata and

A. ventricosa) were deposited in the Herbarium of the

Botanical Garden Museum of University of Bari (BI). The

field observations related to spring and summer during

2008�2013, for the taxa reported in the Red List, gave

deeper knowledge on: (i) ecology (dissemination, fertil-

ity); (ii) phenology (flowering, fructification); (iii) threat

category, according to the criterion B of the IUCN (2012;

2013) and/or in alternative those by Conti et al. (1997);

and (iv) habitat, according to Directive 92/43 EEC (Euro-

pean Commission DG Environment, 2007; Biondi et al.,

2009). A new dichotomous key for identification of all

taxa of Aegilops found in Italy was developed, using as a

model, the monograph of van Slageren (1994).

ResultsA key of Aegilops for Italy was developed. We also

describe the occurrence of the 10 species of Aegilops with

short notes on altitudinal range of their distribution,

phenology, fertility, plant community, priority for in situ

and ex situ conservation, threats, strategies, actions

and norms for conservation. The geographical distribution

of all 10 species is reported, species by species, indica-

ting whenever possible localities, citations and the

332 E. V. Perrino et al.

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reference of the herbarium specimens (Appendix 2,

online supplemental material which is available from

the article’s Taylor & Francis Online page at http://dx.

doi.org/10.1080/14772000.2014.909543). The occurrence

of the species in each administrative region is shown

in Fig. 1, while the geographical distribution from the

data reported on the examined specimens is shown

in Fig. 2.

Fig. 1. Occurrence of the species of genus Aegilops in the administrative regions of Italy.

Aegilops in Italy 333

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Fig. 2. Geographical distribution of the species of genus Aegilops in Italy, from the data reported on the consulted specimens(see Appendix 2).

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Key to the species of Aegilops in Italy

1 Spike ovoid or subovoid, long (without awn)

1�4 cm..........................................................................2

� Spike cylindrical, subcylindrical or moniliform, long

up to 15 cm...................................................................6

2 Glume with (3�)4�5 awns, caryopsis 4�6 mm long,

spike with (2�)3�4 spikelets, the upper one sterile, the

lowest 1�3 subventricose, widest at or below the mid-

dle, constricted above; the glume apex with (3�) 4�5

awns (rarely even only 2), shorter towards the apex of

the spike: 2�4.5 cm at the basis, decreasing to

1�3.5 cm at the apex.................................A. geniculata

� Glume with 1�2(�3) awns, caryopsis 5�8 mm

long...............................................................................3

3 Glume with 1 awn, spike subovoid (rarely monili-

form).........................................................A. uniaristata

� Glume with 2�3 awns, spike ovoid...............................4

4 Glumes with veins unequal in width, sunk into the

glume surface, unequally spaced; caryopsis adherent to

lemma and palea............(doubtful in Italy) A. peregrina

� Glumes with veins equal in width, � parallel, protrud-

ing from the surface, equally spaced; caryopsis free

from lemmaand palea...................................................5

5 Spikelets 2�3 all fertile, rudimentary spikelets 1(�2),

awn almost rough........................................A. biuncialis

� Spikelets 3�6, the upper 1�3 sterile, rudimentary spi-

kelets (2�)3, awn almost smooth.................A. neglecta

6 Spike subcylindrical or distinctly moniliform, glume

(sometimes only the apical ones) with awn 1�6 cm

long................................................................................7

� Spike narrowly cylindrical, glumes 1�few mm long,

without awns (except in A. caudata) or only those of

apical spikelet with 1 awn............................................8

7 Spike subcylindrical, glumes with 2�3 awns, rudimen-

tary spikelet (2�)3, caryopsis free.............A. triuncialis

� Spike distinctly moniliform, glumes (sometimes only

the apical ones) with only 1 awn, rudimentary spikelet

absent (rarely 1�2), caryopsis adherent...A. ventricosa

8 Spike 3�10(�12) cm long, glume of apical spikelet

with awn 3�12 cm long................................................9

� Spike 7�15 cm long, glumes of all spikelet without

awns............................................................................10

9 Spike 5�8(�12) cm long excluding awns, glume of

apical spikelet with 3�6 cm awn with 2 teeth at the

base (sometimes not developed), lemma of apical

spikelet with 4�8 cm awn, apical glume and lemma

awns always shorter than length of entire spike, rudi-

mentary spikelet 1�2.................................A. cylindrica

� Spike 3�12 cm long excluding awns, glume of apical

spikelet with 4.5�12 cm diverging awn (thus longer

than the entire spike) without lateral teeth, lemma of

apical spikelet with a short awn, rudimentary spikelet

1�3..................................(doubtful in Italy) A. caudata

10 Only lemma of the 2 lower florets of the apical spike-

let with (equally) long (4.5�10 cm) awns, spikes

7�15 cm long........................................................

.............(doubtful in Italy) A. speltoides var. speltoides

� Lemmas of all lateral as well as apical fertile florets

with awns, increasing in length from 2.5 cm in basal

spikelets up to 10 cm, spikes 7�10 cm long .................

................(doubtful in Italy) A. speltoides var. ligustica

Distribution in Italy

Old official herbarium specimens in Italy were very useful

for discovering new growing sites of Aegilops species.

Their occurrence and distribution previously described by

Pignatti (1982) and Conti et al. (2005, 2007) were

updated. In the following list the exclamation marks (!)

indicate allochthonous species in the described region.

The new map produced for the Italian geographical distri-

bution of species of Aegilops (Fig. 1) shows that Apulia,

with eight species (A. biuncialis, A. cylindrica!, A. genicu-

lata, A. neglecta, A. peregrina!, A. triuncialis, A. uniaris-

tata and A. ventricosa) is the region with the highest

number of Aegilops species. This is followed by Basili-

cata with seven species (A. biuncialis, A. cylindrica!, A.

geniculata, A. neglecta, A. triuncialis, A. uniaristata, A.

ventricosa), while the poorest regions have only two spe-

cies, such as Valle d’Aosta (A. cylindrica!, A. genicu-

lata), Molise (A. geniculata, A. neglecta) and Marche (A.

geniculata, A. triuncialis?). Aegilops geniculata, A.

neglecta and A. triuncialis, showed similar regional distri-

butions and are the most widespread species in Italy.

From a conservation point of view A. biuncialis and A.

uniaristata need special attention. In fact, the former can

be found in only two regions (Apulia and Basilicata),

while the latter is present in five regions (Apulia,

Basilicata, Campania, Tuscany and Veneto).

Altitudinal range

In Italy, the altitudinal range of Aegilops is a maximum of

about 1000 m a.s.l., thus covering only about one-third of

the total altitudinal range of the genus, which is from

�400 m b.s.l. (Dead Sea area) to 2700 m a.s.l. In particu-

lar, A. geniculata ranges from sea level up to about

1000 m a.s.l.; A. ventricosa only for Umbria is reported at

more than 850 m a.s.l.; while all other species range from

0 up to 600 m of altitude. Aegilops uniaristata is an excep-

tion, since in the past it was never reported at more than

300 m a.s.l. (Pignatti, 1982); however, recently it was

found up to about 500 m a.s.l. Unfortunately, the herbar-

ium specimens of A. speltoides and A. caudata (Flora

Aegilops in Italy 335

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Italica Herbarium of Bertoloni) referred to their presence

near Genoa, but did not report any data about altitude.

Phenology and fertility

The phenologic observations indicate that the blooming

starts in April (instead of May as the rest of Europe and

Asia), continuing all through May (more rarely until

June), in some colder countries of Eurasia the blooming

may end in July. The ripening of caryopsises generally

occurs in June (more rarely in July). From a conservation,

competition and/or survival point of view, it is of consid-

erable importance to stress that most species have a low

number of fertile spikelets and A. biuncialis and A. uniar-

istata represent the extreme cases, carrying only 2�3 fer-

tile spikelets, each of them bearing only one caryopsis.

For these species, the survival of populations is strictly

bound to the few seeds produced by each individual and

to their percentage of germination. Once the caryopsis is

ripened, the spikelet falls on the ground. The seed remains

generally within the spikelet (barochory dissemination).

The penetration of spikelets in the soil is favoured, after

appropriate hydration of spikelet tissues, by the ability to

rotate and twist of the long awns. This mechanism (hygro-

nastic) occurs only in A. biuncialis, A. geniculata, A.

neglecta, A. uniaristata and A. triuncialis. Sometimes part

of the seeds are dispersed to the ground and are ‘captured’

and then transported to anthills by ants (myrmecochory

dissemination), or the spikelets anchor with their awns to

the hairs of livestock, especially sheep (ectozoochory dis-

semination). Naturally, these mechanisms do not allow

the seeds to disperse very far from the point where the

spikelet dropped. The low percentage of seeds produced

(especially by A. biuncialis and A. uniaristata), and the

low efficiency of the dissemination system, explain the

poor ability of these species to expand their area of distri-

bution and the difficulties that they may meet in increas-

ing the number of individuals of the population. For these

populations, it is therefore necessary to adopt specific

safeguards, starting from in situ conservation.

Plant communities

Species of Aegilops are present in different plant commu-

nities, often part of habitats of Directive 92/43 EEC:

(i) they characterize peculiar annual meadows of Brachy-

podietalia distachyi Rivas-Mart�ınez 1978 order (subtype

3 of priority habitat 6220�); (ii) they are sporadic of peren-nial termoxerophilous grasslands of the Lygeo-Stipetea

Rivas-Mart�ınez 1978 and Poetea bulbosae Rivas Goday

& Rivas-Mart�ınez in Rivas-Mart�ınez 1978 classes (sub-

type 1 and 2 of the same priority habitat 6220�), (iii) theyare sporadic of perennial mesic grasslands of Festuco-

Brometea Br.-Bl. et R. Tx. ex Klika et Hadac 1944 class

(habitats 62A0, 6210), and (iv) they are sporadic of the

subnitrophilous vegetation of Stellarietea mediae class.

Detailed information for each species is reported hereaf-

ter, in alphabetical order.

Distribution at the regional level, herbarium

specimens, and other information for each

species

Aegilops biuncialis Vis.

Fl. Dalmat. 3: 344, 1952 (Figs 3 & 9)

Fig. 3. A. biuncialis (Gravina di Palagianello, Palagianello, Taranto, Apulia, 1.05.2013, E.V. Perrino).

336 E. V. Perrino et al.

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Fig. 4. A. geniculata (Platani, Agrigento, Sicily, 3.05.2012, E.V.Perrino).

Fig. 5. A. neglecta (Chiancarete, Sasso di Castalda, Potenza,Basilicata, 7.06.2013, E.V. Perrino).

Fig. 6. A. triuncialis (Masseria Semeraro, Mottola, Taranto,Apulia, 29.04.2010, E.V. Perrino).

Fig. 7. A. uniaristata (Gravina di Palagianello, Palagianello,Taranto, Apulia, 1.05.2013, E.V. Perrino).

Fig. 8. A. ventricosa (Bosco Santantuono, Mottola, Taranto,Apulia, 14.06.2012, E.V. Perrino).

Aegilops in Italy 337

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Homotypic synonyms. A. geniculata Roth subsp. biun-

cialis (Vis.) Zangh., Fl. Ital. 1: 979, 1976, comb. inval.; A.

s ovata L. var. biuncialis (Vis.) Fiori & Paoletti, Fl. Italia

1: 109, 1896; A. ovata L. subsp. biuncialis (Vis.) Emb. &

Maire, Cat. Pl. Maroc 4: 947, 1941; Triticum biunciale

(Vis.) K. Richt., Pl. Eur. 1: 128, 1890, nom. illeg.

Heterotypic synonyms. A. biaristata Lojac., Fl. Sicul. 3:

370, 1909; A. biuncialis Vis. var. archipelagica Eig,

Repert. Spec. Nov. Regni Veg. Beih. 55: 137, 1929; A.

biuncialis Vis. var. velutina Zhuk., Trudy Prikl. Bot. 18:

483, 1928; A. connata Steud., Syn. Pl. Glumac. 1: 356,

1854; A. intermedia Steud., Syn. Pl. Glumac. 1: 354,

1854; A. lorentii Hochst. Flora 28: 25, 1845; Aegilops

macrochaeta Shuttlew. & Huet ex Duval-Jouve, Bull.

Soc. Bot. France 16: 384, 1869; A. macrochaeta Shuttlew.

& Huet ex Duval-Jouve subsp. pontica Degen, Magyar

Bot. Lapok 30: 111, 1931; A. ovata L. var. latearistata

Lange, Pugillus Pl. Hispan. 1: 56, 1860; A. pontica

(Degen) Waleff, Fl. Nar. Republ. Bulgariya 1: 473, 1963;

A. triaristata Willd. var. trispiculata Hack. ex Trab., Fl.

Alger 107, 1884; A. trispiculata (Hack. ex Trab.) Hack.

ex Batt. & Trab., Fl. Alg�erie Monocot.: 241, 1895.

A Mediterranean-western Asiatic element. The flower-

ing period extends from April to June. It grows from sea

level up to 850 m a.s.l., where it characterizes annual

meadows on calcareous and clay-marly soils, which are

included in the priority habitat ‘Pseudo-steppe with

grasses and annuals of the Thero-Brachypodietea’ (sub-

type 3 of 6220�). For over a century, the Italian botanists

thought that the only known station of A. biuncialis was

that of Gravina di Leucaspide, in Apulia (Groves, 1887),

later confirmed by Curti (1986). It was a wrong assump-

tion, because the species is also reported for Basilicata,

Campania, Tuscany and Veneto, as shown by the herbar-

ium specimens. According to more recent findings, other

Apulian stations are those of Alimini Lakes (Lecce)

(Mele et al., 2006), Gravina di Palagianello (Taranto)

(Carruggio et al., 2008) whose annual meadows are local-

ized in the clearing of Quercus trojanaWebb wood, Capi-

tolo (Monopoli, Bari) (Perrino & Signorile, 2009), where

the species occurs as a small population, with few individ-

uals, Vernole (Lecce) (Medagli et al., 2014), Murge della

Morsara and Masseria Cingavalle (Santeramo in Colle,

Bari), Monte Sant’Elia and Masseria Semeraro (Mottola,

Taranto), Gravina di Pentimelle (Castellaneta, Taranto)

(Perrino & Wagensommer, 2013a) and Portico del Ladro

(Massafra, Taranto). In Campania, other stations to be

included are those of Monte Pruno (Roscigno, Salerno),

which is the second regional station located at about

850 m a.s.l. (Del Vico et al., 2007). The new recent sta-

tions and the checking of old official herbarium specimens

have increased the Italian geographical distribution area

of the species, and applying the criterion B (IUCN, 2012),

the threat category was lowered, from Critically

Endangered (CR) (Conti et al., 1997) to Vulnerable

[VU B1ab(i,ii,iii,iv)þ2ab(i,ii,iii,iv)] (Perrino & Wagen-

sommer, 2013a).

Aegilops caudata L.

Sp. Pl. 2: 1051, 1753

Homotypic synonyms. Orrhopygium caudatum (L.) �A.L€ove, Biol. Zentralbl. 101: 206, 1982; Triticum caudatum

(L.) Pers., Syn. Pl. 1: 110, 1805, nom. illeg.

Heterotypic synonyms. A. caudata L. subsp. dichasians

Zhuk., Bull. Appl. Bot., Gen. & Pl. Breeding 18: 512,

1928, nom. illeg.; A. caudata L. var. paucispiculigera O.

Schwarz, Repert. Spec. Nov. Regni Veg. 36: 68, 1934; A.

caudata L. var. polyathera Boiss., Fl. Orient. 5: 675,

1984; A. dichasians (Zhuk.) Humphries, Bot. J. Linn. Soc.

78: 236, 1979, comb. illeg.; Triticum dichasians (Zhuk.)

Bowden, Canad. J. Bot. 37: 667, 1959, comb. illeg.; Triti-

cum markgrafii Greuter, Boissiera 13: 172, 1967.

Eastern Mediterranean element occurring mainly in the

Aegean region and western Turkey. In Italy, it has not

been reported for more than a century. It was cited in

Liguria (Bertoloni, 1834) and in Sardinia at Pula

(Cagliari) (Fiori & Paoletti, 1896; Pignatti, 1982). In Sar-

dinia it was not recently reported (Viegi, 1993) and as far

Fig. 9. A. biuncialis (Gravina di Palagianello, Palagianello,Taranto, Apulia, 1.05.2013, 36252 - E.V. Perrino, BI).

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as the ligurian station is concerned the situation is con-

fused, since it is reported as erroneous (Conti et al., 2005)

or as A. cylindrica (Bertoloni, 1834), which according to

Van Slageren (1994) is A. caudata. In conclusion, consid-

ering that the citations are old and recently it was never

found in any of the two old stations, nor were new sites

discovered, the occurrence of the species in Italy should

be considered uncertain.

Aegilops cylindrica Host

Icon. Descr. Gram. Austriac. 2: 6, 1802

Homotypic synonyms. A. caudata L. subsp. cylindrica

(Host) Hegi, Ill. Fl. Mitt.-Eur. 1: 390, 1908, comb. illeg.;

A. caudata L. var. cylindrica (Host) Fiori, Nuov. Fl. Italia

1: 160, 1923, comb. illeg.; Cylindropyrum cylindricum

(Host) �A. L€ove, Biol. Zentralbl. 101: 207, 1982; Triticumcylindricum (Host) Cesati, Passerini & Gibelli, Comp. Fl.

Ital. 86, 1869.

Heterotypic synonyms. A. caudata L. var. hirsuta (Binz)

Hegi, Ill. Fl. Mitt.-Eur. 1: 390, 1908; A. cylindracea Kit.

(ex J�av) ex So�o, Syst. Geobot. Handb. Hung. Fl. 5: 354,1973, nom. illeg.; A. cylindrica Host var. albescens

Popova, Trudy Prikl. Bot. Selekts. 13: 461, 1923; A. cylin-

drica Host var. aristulata Zhuk. Tzvelev, Novosti Sist.

Vyssh. Rast. 10: 37, 1973; A. cylindrica Host var. brunnea

Popova, Trudy Prikl. Bot. Selekts. 13: 461, 1923; A. cylin-

drica Host var. ferruginea Popova, Trudy Prikl. Bot.

Selekts. 13: 461, 1923; A. cylindrica Host var. flavescens

Popova, Trudy Prikl. Bot. Selekts. 13: 461, 1923; A. cylin-

drica Host var. fuliginosa Popova, Trudy Prikl. Bot.

Selekts. 13: 461, 1923; A. cylindrica Host var. kastoria-

num Karat., Pl. Syst. Evol. 163: 19, 1989; A. cylindrica

Host var. multiaristata Jansen & Wacht., Ned. Kruidk.

Arch. 3: 138, 1931; A. cylindrica Host var. pauciaristata

Eig, Repert. Spec. Nov. Regni Veg. Beih. 55: 103, 1929;

A. cylindrica Host var. prokhanovii Tzevelev, Novosti

Sist. Vyssh. Rast. 10: 37, 1973; A. cylindrica Host var.

pubescens (Kloos) Jansen, Ned. Kruidk. Arch. 3: 66,

1914; A. cylindrica Host var. rubiginosa Popova, Trudy

Prikl. Bot. Selekts. 13: 461, 1923; A. cylindrica Host var.

rumelica Velen., Fl. Bulg. 627, 1891.

A widespread European and Mediterranean-western

Asiatic element. The Italian altitudinal range cannot be

defined because the locations shown in the ancient herbar-

ium specimens are too generic. For this reason the lower

altitudinal limit is the one reported by Pignatti (1982), for

which, probably, the maritime stations are those of Sardi-

nia and Liguria, while for the upper limit one cannot refer

to Mt. Muret (3062 m a.s.l.), since it is (highly) likely that

the sample of Alioth (G) was collected somewhere at a

lower level than the top of the mountain. In fact, the abso-

lute upper limit (except in the USA) does not exceed

2000 m (Van Slageren, 1994). The flowering period goes

from May to June (Pignatti, 1982). The herbarium speci-

mens indicate the occurrence of the species in the follow-

ing different regions: Friuli Venezia-Giulia, Liguria,

Piemonte, Sardinia Valle d’Aosta and Veneto. The species

is also indicated in Basilicata, Lombardia and Trentino

Alto Adige (Celesti Grapow et al., 2009). Fiori & Paoletti

(1896) reported the species as a widespread taxon in Apu-

lia, while Pignatti (1982) states that it is located only near

the town of Gravina in Puglia. About this point, further

research is needed. The species is marked as at Lower

Risk (LR) in Valle d’Aosta and Piemonte, and Critically

Endangered (CR) in Apulia (Conti et al., 1997). In Apulia

its occurrence is strongly questionable.

Aegilops geniculata Roth

Bot. Abh. Beobacht 45: 1787 (Figs 4 & 10)

Homotypic synonyms. Triticum ovatum (L.) Raspail,

Ann. Sci. Nat. 1: 435, 1825.

Heterotypic synonyms. A. altera Cam. ex Roth, Bot.

Abh. Beobacht. 45, 1787, nom. inval.; A. brachyatera

Pomel, Nouv. Mat. Fl. Atl. 2: 389, 1875; A. divaricata

Jord. & Fourr., Brev. Pl. Nov. 2: 129, 1868; A. echinus

Godr., M�em. Sect. Med. Acad. Sci. Montpellier 1: 456,

1853; A. erigens Jord. & Fourr., Brev. Pl. Nov. 2: 131,

Fig. 10. A. geniculata (Acqua Fitusa, Sicily, 2.05.2012, 36179 -E.V. Perrino, BI).

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1853; A. erratica Jord. & Fourr., Brev. Pl. Nov. 2: 130,

1890; A. fonsii Sennen, Butl. Inst. Catalana Hist. Nat. 20:

122, 1920; A. narbonensis Lobel ex Honck., Syn. Pl.

Germ. 1: 485, 1792, nom. inval.; A. ovata L. var. africana

Eig, Repert. Spec. Nov. Regni Veg. Beih. 55: 144, 1929;

A. ovata L. subsp. atlantica Eig, Repert. Spec. Nov. Regni

Veg. Beih. 55: 144, 1929; A. ovata L. subsp. eventricosa

Eig, Repert. Spec. Nov. Regni Veg. Beih. 55: 144, 1929;

A. ovata L. var. genuina Griseb., Spic. Fl. Rumel. 2: 425,

1846, nom. inval.; A. ovata L. subsp. gibberosa Zhuk.,

Trudy Prikl. Bot. 18: 471, 1928; A. ovata L. subsp. globu-

losa Zhuk., Trudy Prikl. Bot. 18: 473, 1928; A. ovata L.

subsp. hirsuta Eig, Repert. Spec. Nov. Regni Veg. Beih.

55: 144, 1929, nom. illeg.; A. ovata L. subsp. lanuginosa

Zhuk., Trudy Prikl. Bot. 18: 474, 1928; A. ovata L. subsp.

planiuscula Zhuk., Trudy Prikl. Bot. 18: 473, 1928; A.

ovata L. subsp. puberula Zhuk., Trudy Prikl. Bot. 18:

473, 1928; A. ovata L. subsp. umbonata Zhuk., Trudy

Prikl. Bot. 18: 72, 1928; A. ovata L. subsp. vernicosa

Zhuk., Trudy Prikl. Bot. 18: 473, 1928; A. parvula Jord.

& Fourr., Brev. Pl. Nov. 2: 131, 1868; A. peregrina Tab-

ern. ex Honck., Syn. Pl. Germ. 1: 485, 1792, nom. inval.;

A. procera Jord. & Fourr., Brev. Pl. Nov. 2: 129, 1868; A.

pubiglumis Jord. & Fourr., Brev. Pl. Nov. 2: 131, 1868; A.

sicula Jord. & Fourr., Brev. Pl. Nov. 2: 129, 1868; A.

veterum J. Bauhin ex Honck., Syn. Pl. Germ. 1: 485,

1792, nom. inval.; Festuca italica J. Gerard ex Honck.,

Syn. Pl. Germ. 1: 485, 1792, nom. inval

Stenomediterranean-Turanian element. Very common

in uncultivated habitats and roadsides, less frequent in

grasslands, garigues and scrublands. It is the most com-

mon species of the genus A., occurring in 19 out of the 20

Italian regions. Its upper altitudinal limit is reported in the

Vascular Flora of Latium (Anzalone et al., 2010): 1000 m

a.s.l. The absence in Trentino Alto Adige (Conti et al.,

2005) should be considered an anomalous datum since its

occurrence is documented in several works and herbarium

specimens of neighbouring regions with the same ecologi-

cal conditions. In Molise the species is reported by Conti

et al. (2005), but we found no specimens from this region

in the consulted herbaria. Typically, the species character-

izes aspects of sub-nitrophilous vegetation of Stellarietea

mediae R. T€uxen, Lohmeyer & Preising ex von Rochow

1951, although it is frequently a companion species in

many types of other plant communities, especially the

grasslands of Lygeo-Stipetea Rivas-Mart�ınez 1978 (habi-

tat 6220� of Directive 92/43 EEC) and Festuco-Brometea

Br.-Bl. & T€uxen ex Klika & Hada�c 1944 (habitat 6210

and 62A0 of Directive 92/43 EEC).

Aegilops neglecta Requien ex Bertoloni

Fl. Ital. 1: 787, 1834 (Figs 5 & 11)

Homotypic synonyms. Triticum neglectum (Req.ex Ber-

tol.) Greuter, Boissiera 13: 171, 1967. Heterotypic

synonyms. A. algeriensis Gand., €Osterr. Bot. Zeithschr.31: 81, 1881; A. calida Gand., €Osterr. Bot. Zeithschr. 31:81, 1881; A. campicola Gand., €Osterr. Bot. Zeithschr. 31:82, 1881; A. gussonii Link, Linnaea 17: 388, 1843, nom.

inval.; A. mesantha Gand., €Osterr. Bot. Zeithschr. 31: 82,1881; A. ovata Roth, Ann. Bot. (Usteri) 4: 41, 1793, nom.

illeg.; A. ovata var. triaristata (Willd.) Griseb., Spic. Fl.

Rumel. 2: 425, 1846; A. triaristata Willd., Sp. Pl. 4: 943,

1806, nom. illeg.; A. triaristata Willd. subsp. attenuata E.

Schiem., Ber. Deutsch. Bot. Ges. 47: 177, 1929; A. triaris-

tata Willd. var. brachychaeta Font Quer, Treb. Mus. Ci.

Nat. Barcelona, Bot. 5: 54, 1924; A. triaristata Willd.

subsp. contorta Zhuk., Trudy Prikl. Bot. 18: 479, 1928;

A. triaristata Willd. subsp. contracta Eig, Repert. Spec.

Nov. Regni Veg. Beih. 55: 141, 1929; A. triaristataWilld.

subsp. elongata E. Schiem., Ber. Deutsch. Bot. Ges. 47:

177, 1929; A. triaristata Willd. var. hirsuta Zhuk., Trudy

Prikl. Bot. 18: 480, 1928; A. triaristata Willd. subsp.

Fig. 11. A. neglecta (Bosco delle Pianelle, Martina Franca,Taranto, Apulia, 7.05.1984, 70166 - Brullo, Signorello, Spampi-nato, CAT).

340 E. V. Perrino et al.

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intermixta Zhuk., Trudy Prikl. Bot. 18: 479, 1928; A. tri-

aristata Willd. var. kabylica Battand. & Trab., Fl. Alg�erie1: 241, 1895; A. triaristata Willd. var. macrochaeta E.

Mey, Pflanzenbau 3: 303, 1927; A. triaristata Willd. var.

ochreata Zhuk., Trudy Prikl. Bot. 18: 480, 1928; A.

triaristata Willd. subsp. quadriaristata Eig, Repert. Spec.

Nov. Regni Veg. Beih. 55: 140, 1929; A. triaristata Willd.

subsp. recta Zhuk., Trudy Prikl. Bot. 18: 478, 1928; A. tri-

aristata Willd. var. trojana Eig, Bull. Soc. Bot. Gen�eve 19:2, 1928; A. virescens Jord. & Fourr., Brev. Pl. Nov. 2: 130,

1868; A. viridescens Gand., €Osterr. Bot. Zeithschr. 31: 81,1881; Triticum rectum (Zhuk.) Bowden, Canad. J. Genet.

Cytol. 8: 135, 1966; Triticum triaristatum (Bluff, Nees &

Schauer) Godr., Fl. France 3: 602, 1856, nom. illeg.

A Mediterranean-Western Asiatic and Circumboreal

element that probably exceeds 1000 m a.s.l. (locality:

Monte Paterno, Veneto) and is always very rare. The

period of flowering and fruiting extends usually from

April to June, but in Sicily goes from April to May

(Lojacono-Pojero, 1908�1909). The habitats in which it

grows are similar to those of A. geniculata, though

slightly less nitrophilous and anthropic, as roadsides,

annual meadows, grasslands, garigues, scrublands, eco-

tonal belts of cultivated areas, as vineyards and olive

groves, rarely in clearings of the woods. From a vegeta-

tion point of view it is not clear if the annual meadows

fall in habitat of Directive 92/43: ‘Pseudo-steppe with

grasses and annuals of the Thero-Brachypodietea’ (6220�

subtype 3). The herbarium specimens indicate the occur-

rence of the species in the following different regions:

Apulia, Basilicata, Calabria, Emilia Romagna, Friuli Ven-

ezia-Giulia, Latium, Liguria, Lombardia, where it was

documented also in the province of Cremona (Bonali

et al., 2006), Piemonte, Sardinia, Sicily, Tuscany, Umbria,

Veneto. In Molise it is reported in dry grasses (Lucchese,

1995). The species is not reported in Marche, Trentino

Alto Adige and Val d’Aosta, while it is doubtful in Cam-

pania and Abruzzo (Conti et al., 2005). It is morphologi-

cally similar to A. biuncialis, from which it differs for

having 3�6 spikelets, 1�3 of which are sterile (upper

ones) and 2�3 rudimentary (see identification key).

Aegilops peregrina (Hackel in J. Fraser) Maire &

Weiller Fl. Afrique Nord 3: 358, 1955 (Fig 12)

Homotypic synonyms. Aegilemma peregrina (Hack.) �A.L€ove, Feddes Repert. 95: 499, 1984; A. peregrina (Hack.)

Eig, Repert. Spec. Nov. Regni Veg. Beih. 55: 121, 1929,

nom. inval.; A. variabilis Eig var. peregrina (Hack.) Eig

& Feinbrun, Repert. Spec. Nov. Regni Veg. Beih. 55:

125, 1929; Triticum peregrinum Hackel, Ann. Scott. Nat.

Hist. 1907: 102, 1907.

Heterotypic synonyms. A. variabilis (Maire & Weiller)

Eig, Repert. Spec. Nov. Regni Veg. Beih. 55: 21, 1929,

nom. illeg.; A. variabilis Eig subsp. cylindrostachys Eig &

Feinbrun, Repert. Spec. Nov. Regni Veg. Beih. 55: 125,

1929, nom. inval.; A. variabilis Eig var. intermedia Eig &

Feinbrun, Repert. Spec. Nov. Regni Veg. Beih. 55: 124,

1929, comb. illeg.; A. variabilis Eig var. latiuscula Eig &

Feinbrun, Repert. Spec. Nov. Regni Veg. Beih. 55: 124,

1929; A. variabilis Eig var. multiaristata (Eig & Feinbrun)

K. Hammer, Feddes Repert. 91: 237, 1980; A. variabilis

Eig var. mutica Eig & Feinbrun, Repert. Spec. Nov. Regni

Veg. Beih. 55: 124, 1929; A. variabilis Eig var. planispi-

cula Eig & Feinbrun, Repert. Spec. Nov. Regni Veg.

Beih. 55: 124, 1929; Triticum triaristatum (Willd.) Gren.

& Godr. var. brachyatera Hack., All. Bot. Zeitschr. 12:

33, 1915; Triticum variabile (Eig) Markgr., Repert. Spec.

Nov. Regni Veg. Beih. 30: 225, 1932.

This species includes two varieties: var. peregrina and

var. brachyathera (Boiss.) Maire & Weiller. The variety

peregrina is a Western Asiatic element occurring abun-

dantly only in Palestine, Western Jordan, Lebanon and

Western Syria, uncommon to rare elsewhere. The variety

brachyathera is more confined to coastal and lowland

areas, is less frequently encountered and not known in

Greece. In Italy these varieties are both reported only

from the province of Lecce, in Apulia (Conti et al., 2005).

The species was not found in the old stations after the

Fig. 12. A. peregrina (Gallipoli, Lecce, Apulia, 1881, H.Groves, FI).

Aegilops in Italy 341

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19th century and therefore in Italy its occurrence should

be considered highly doubtful.

Aegilops speltoides Tausch

Flora 20: 108, 1837 (Figs 13 & 14)

Homotypic synonyms. A. ligustica (Savign.) Coss. subsp.

speltoides (Tausch) Eig in Hannig & Winkler, Die Pflanze-

nareale 4: 46, 1936; Sitopsis speltoides (Tausch) �A. L€ove,Feddes Repert. 95: 491, 1984; Triticum speltoides (Tausch)

Asch. & Graebn., Mitteleur. Fl. 2: 711 1901.

Heterotypic synonyms. A. agropiroїdes Godr., M�em.

Sect. Med. Acad. Sci. Montpellier 1: 456, 1853; A.

aucheri Boiss., Diagn. Pl. Orient. 5: 74, 1844; A. aucheri

Boiss. var. schultzii N�abĕlek, Publ. Fac. Sci. Univ. Masar-

ykovy 105: 30, 1929; A. aucheri Boiss. var. striata Zhuk.,

Trudy Prikl. Bot. 18: 536, 1928; A. aucheri Boiss. var. vel-

lea Zhuk., Trudy Prikl. Bot. 18: 533, 1928; A. macrura

Jaub. & Spach, Ill. Pl. Orient. 4: 21, 1851; A. markgrafii

var. polyathera (Boiss.) K. Hammer, Feddes Repert. 91:

232 1980; A. singularis Steud., Syn. Pl. Glumac. 1: 354,

1854; A. speltoides Tausch var. macrostachys Eig, Bull.

Inst. Agric. Nat. Hist. Tel-Aviv 6: 73, 1927; Agropyron

tournefortii Savign., Flora 35: 569, 1847; Triticum

obtusatum Godr., M�em. Sect. Med. Acad. Sci. Montpellier

1: 454, 1853.

This species includes two varietes: var. speltoides and

var. (b) ligustica (Savign.) Fiori, both with an

Asian�Western distribution. The first one (var. spel-

toides) reaches South-Eastern Bulgaria. In Italy, both vari-

eties are reported as adventitious (Pignatti, 1982). The old

specimens of 1846 of Savignone (Savignone, 1847) are

the only Italian records (as reported by Fiori & Paoletti

1896 . . . in the hills of Genoa). In a situation like this, it

would be wise to conclude that the occurrence of the spe-

cies in Italy is doubtful. To reach a definitive conclusion,

further investigation on the exsiccata and new floristic

surveys on the territory are needed. This would be advis-

able also because A. speltoides is considered to be the

probable donor of the B genome to Triticum turgidum ssp.

durum (Desf.) Husn. (Petersen et al., 2006).

Aegilops triuncialis L.

Sp. Pl. 2: 1051, 1753 (Figs 6 & 15)

Homotypic synonyms. Aegilopodes triuncialis (L.) �A.L€ove, Biol. Zentralbl. 101: 207, 1982, comb. incorr.; A.triuncialis L. var. typica Zhuk., Bull. Appl. Bot., Gen. &

Fig. 13. A. speltoides var. ligustica (near Genoa, Liguria, 1846,s.n. - Savignone, BOLO).

Fig. 14. A. speltoides var. speltoides (near Genoa, Liguria, 1846,s.n. - Savignone, BOLO).

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Pl. Breeding 18: 500, 1928; Triticum triunciale (L.) Ras-

pail, Ann. Sci. Nat. (Paris) 5: 435, 1825. Heterotypic syn-

onyms: A. croatica Gand., €Osterr. Bot. Zeitschr. 31: 81,1881; A. elongata Lam., Fl. France 3: 632, 1778, nom.

illeg.; A. squarrosa L., Sp. Pl. 2: 1051, 1753; A. triaristata

Req. ex Bertol., Fl. Ital. 1: 789, 1834, nom. illeg.;

A. triuncialis L. var. albescens Popova, Trudy Prikl. Bot.

Selekts. 13: 476, 1923; A. triuncialis L. var. assyriaca

Eig, Bull. Soc. Bot. Gen�eve 19: 19, 1928; A. triuncialis L.var. biaristatum Arut. & Gandilyan, Biol. J. Armenia 43:

747�748, 1990, nom. inval.; A. triuncialis L. var. breviar-

istata Hack., Nuov. Giorn. Bot. Ital. 19: 212, 1887; A.

triuncialis L. var. brunnea Popova, Trudy Prikl. Bot.

Selekts. 13: 476, 1923; A. triuncialis L. subsp. caput-

medusae Zhuk., Trudy Prikl. Bot. 18: 499, 1928; A.

triuncialis L. var. constantinopolitana Eig, Repert. Spec.

Nov. Regni Veg. Beih. 55: 133, 1929; A. triuncialis L.

subsp. fascicularis Zhuk., Trudy Prikl. Bot. 18: 499,

1928; A. triuncialis L. var. ferruginea Popova, Trudy

Prikl. Bot. Selekts. 13: 476, 1923; A. triuncialis L. var. fla-

vescens Popova, Repert. Spec. Nov. Regni Veg. 13: 475,

1923; A. triuncialis L. var. glabrispica Eig, Bull. Inst.

Agric. Nat. Hist. Tel-Aviv 6: 71, 1927; A. triuncialis L.

var. muricata Zhuk., Trudy Prikl. Bot. 18: 500, 1928; A.

triuncialis L. var. nigroalbescens Popova, Trudy Prikl.

Bot. Selekts. 13: 476, 1923; A. triuncialis L. var. nigroar-

istata Flaksb., Trudy Prikl. Bot. Selekts. 13: 484, 1923; A.

triuncialis L. var. nigroferruginea Popova, Trudy Prikl.

Bot. Selekts. 13: 476, 1923; A. triuncialis L. var. nigrofla-

vescens Popova, Trudy Prikl. Bot. Selekts. 13: 475, 1923;

A. triuncialis L. var. nigrorubiginosa Popova, Trudy Prikl.

Bot. Selekts. 13: 476, 1923; A. triuncialis L. var. rubigi-

nosa Popova, Trudy Prikl. Bot. Selekts. 13: 475, 1923; A.

triuncialis L. var. triaristatum Arut. & Gandilyan, Biol. J.

Armenia 43: 747�748, 1990, nom. inval; Triticum

persicum (Boiss.) Aiton & Hemsl., Linn. Soc. London,

Bot. 3: 127, 1886, nom. illeg.; Triticum squarrosum (L.)

Raspail, Ann. Sci. Nat. (Paris) 5: 435, 1825, nom. illeg.

A widespread Mediterranean-Western Asiatic Circum-

boreal element. Occurring in the most southern part of

Europe. In Italy, it is reported from sea level up to not more

than 600 m a.s.l., whose flowering goes from May to June

(Gismondi, 1950) in the central and northern part of the pen-

insula, in the southern part, as noted for Sicily, the flowering

period goes from April to June (Lojacono-Pojero,

1908�1909). The species has a discontinuous distribution

in Central and Northern Italy, occurring in the following

regions, as shown by the herbarium specimens: Apulia,

Calabria, Campania Friuli Venezia-Giulia, Liguria, Pie-

monte Sardinia, Sicily, Trentino Alto Adige, Tuscany, Ven-

eto. In Basilicata, the species is reported in province of

Matera by Medagli & Gambetta (2003) and by Silletti

(2010), sub Triticum triunciale (L.) Raspail (syn.). The spe-

cies is reported also from Latium (Anzalone et al., 2010)

while it is absent in Molise, Marche, Umbria, Val d’Aosta,

Lombardia, and has not been reported for a long time in

Abruzzo (Conti et al., 2005). It is difficult to explain these

gaps. Probably, the species escapes observation because it

constitutes very small populations. For instance, each of the

four apulian populations actually known, in the municipali-

ties of Crispiano, Palagianello and Mottola (province of

Taranto), is approximately 15/20 m2 wide. Only in Latium

it is reported in the Regional Red List with the status of

lower risk (LR) (Conti et al., 1997).

Aegilops uniaristata Vis.

Fl. Dalmat. 3: 345, 1852 (Figs 7 & 16)

Homotypic synonyms. A. uniaristata Steud., Syn. Pl.

Glumac. 1: 354, 1854, nom. illeg.; Chennapyrum uniaris-

tatum (Vis.) �A. L€ove, Biol. Zentralbl. 101: 207, 1982; Tri-ticum uniaristatum (Vis.) K. Richter, Pl. Eur. 1: 128,

1890.

Heterotypic synonyms. A. notarisii Clem., Mem. Acd.

Sci. Torin., Ser. 2: 335, 1857.

It is a Southeast-European species, occurring in Croatia,

Greece (islands included), Albania and Italy (Visiani,

1852; Hayek, 1932; Davis, 1986; van Slageren, 1994).

According to the recent works of the Vascular Flora of

Fig. 15. A. triuncialis (near Masseria Primicerio, Mottola,Taranto, Apulia, 1.05.2013, 36253 - E.V. Perrino, BI).

Aegilops in Italy 343

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Turkey (Semiz & Celik, 2005) the species does not occur

in Turkey. In Italy it is known only in Apulia and Basili-

cata. The new stations have increased the altitudinal limit,

by exceeding 500 m a.s.l., which, in Italy, remains the

lowest of all species of the genus. It grows on rocky, cal-

careous soils and red soils (terra rossa). The flowering

period extends from May to June (Fiori & Paoletti, 1896;

Pignatti, 1982), although in the maritime stations, as

observed in Apulia, it starts in April. In Apulia, the first

discovered station was that of Leucaspide (province of

Taranto) (Groves, 1887). Later it was reported also in the

South Eastern part of Murge (Taranto), in the Laterza

ravine and near Bosco delle Pianelle (Martina Franca),

between Spongano and Surano (province of Lecce)

(Bianco et al., 1989), Bosco di Rauccio and Santa Cesarea

Terme (province of Lecce) (Marchiori et al., 1993), Bosco

Difesa Grande of Gravina in Puglia (province of Bari)

(Forte, 2001), in olive groves between Maruggio and

Manduria (province of Taranto) (Caforio & Marchiori,

2006), in ancient olive groves between Fasano and Brin-

disi (province of Brindisi) (Perrino, 2011). The only data

for Basilicata are those from Bosco di Lucignano, near

Matera (Medagli & Gambetta, 2003). At national level

and applying the criteria A and B of the IUCN Protocol

(2013), the threat category of the species for Italy is

Vulnerable [VU B1ab(iii, v)þ2ab(iii, v)] (Perrino &

Wagensommer, 2012). Probably, A. uniaristata, a very

rare European taxon, might become more vulnerable

because of a combination of limited distribution and graz-

ing pressure. In addition, from an ecological and conser-

vation point of view, it is worth noting that, as for A.

biuncialis, it produces the fewest number of fertile spike-

lets of the genus, therefore the survival of (the few) popu-

lations depends on the rate of germination of the few

seeds produced by each individual. In particular, the Ital-

ian populations present no more than just 2�3 fertile spi-

kelets (Pignatti, 1982), even though van Slageren (1994)

reports 3�5 fertile spikelets. This can be explained by the

fact that van Slageren (1994) was referring to samples

covering the whole distribution range of the species. At

the same time, when present, A. uniaristata is an impor-

tant environmental component of vegetation that could be

included into ‘Pseudo-steppe with grasses and annuals of

the Thero-Brachypodietea’ (code �6220), priority habitat

of Directive Habitat 92/43 CEE (European Commission,

1992; European Commission DG Environment, 2007; San

Miguel, 2008). For these reasons, specific guidelines for

conservation were proposed within the project LifeþCENT.OLI.MED (see conclusions).

Aegilops ventricosa Tausch

Flora 20: 108, 1837 (Figs 8 & 17)

Homotypic synonyms. Gastropyrum ventricosum (Tausch)�A. L€ove, Biol. Zentralbl. 101: 208, 1982; Triticum ventrico-

sum (Tausch) Cesati, Passerini & Gibelli, Comp. Fl. Ital.

1: 86, 1869.

Heterotypic synonyms. A. fragilis Parl., Fl. Ital. 1: 515,

1850; A. squarrosa L. var. comosa Coss., Notes Pl. Crit.

68, 1850; A. squarrosa L. var. truncata Coss., Notes Pl.

Crit. 1: 68, 1850; A. subulata Pomel, Nouv. Mat. Fl.

Atlantique 2: 388, 1875; A. ventricosa Tausch var.

obscura Miczynski, Bull. Soc. Bot. France 76: 715, 1929;

Triticum fragile (Parl.) Cesati, Passerini & Gibelli, Comp.

Fl. Ital. 87 1869, nom. illeg.; Triticum subulatum (Pomel)

T. Durand & Schinz, Consp. Fl. Afric. 5: 939, 1894, nom.

illeg.

Western-Stenomediterranean species. In Italy, it is

very rare. Pignatti (1982) separated it from A. fragilis

Parl., on the basis of the reduced number of spikelets

(only 3�4), the small size of the awns of the lower

glumes, and the awn of terminal spikelet, which is

2 cm or more in length. However, these characters fall

under the description of A. ventricosa (Van Slageren,

1994). As a consequence, all the records of A. fragilis

relating to Apulia, Abruzzo and Sardinia (Fiori &

Paoletti, 1896; Pignatti, 1982), should be referred to A.

Fig. 16. A. uniaristata (Jazzo Pallanculo, Laterza, Taranto,Apulia, 19.06.2009, 36264 - E.V. Perrino, BI).

344 E. V. Perrino et al.

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ventricosa. The herbarium specimens testify the occur-

rence of the species in Apulia, Calabria, Campania,

Piemonte, Sardinia, Sicily and Umbria. Other regions

in which it occurs are Abruzzo, where it is reported

from alveo del Tordino, Teramo (Zodda, 1967;

Pignatti, 1982) and Chieti (Leporatti et al., 2001),

Basilicata, where it is reported for Ginestra and Serra

La Croce (Rionero in Vulture, Potenza) (Laghetti

et al., 1992) and in an oak wood, near Matera (Silletti,

2010), and Veneto, where it is reported from Monte

Baldo, as allochthonous species (Prosser et al., 2009).

At national level, applying the criterion B of the IUCN

Protocol (2013) to the new data of the Italian distribu-

tion of A. ventricosa, the threat category of the species

in Italy is Endangered [EN B2ab(i,ii,iii,iv)] (Perrino &

Wagensommer, 2013b). Previously the species has

been indicated as vulnerable (VU) in Tuscany and

Apulia, and as a Lower Risk (LR) in Abruzzo (Conti

et al., 1997).

Priority for in situ and ex situ conservation

The criteria to decide priorities for in situ conservation of

the different species are different from those for ex situ con-

servation. For in situ conservation, the main criterion is the

risk that species run mainly in function of their presence,

abundance and geographical distribution. For ex situ conser-

vation, the main criterion is the presence of the species in

gene banks and botanical gardens, where they are kept

mainly for studies and use by breeders to improve cultivated

varieties of wheat (Bioversity International et al., 2009).

Based on sampling, geographical and environmental

representation, experts consider the priority as high,

medium, low or nil for species for which gene banks and

botanical gardens retain, respectively, with respect to the

number of samples. Aegilops geniculata is quite wide-

spread and well distributed, for which the priority for in

situ conservation is nil, while for the ex situ is high,

because there are no collections of this species in gene

banks. Aegilops neglecta, like A. geniculata, is wide-

spread and it is already well represented in gene banks;

therefore priority for ex situ and in situ conservation is

null. Aegilops biuncialis with regard to its presence in

nature has a different degree of risk for the different sites

and therefore the priority for in situ conservation is

medium, while it is nil for the ex situ, because gene banks

and botanical gardens already retain a considerable num-

ber of samples. Aegilops triuncialis is not as widespread

as the previous ones, but relies on a large number of sta-

tions and regions where it grows in Italy. It cannot be con-

sidered truly at risk, which is why the priority in situ is

medium, while it is low for ex situ, being already well rep-

resented in gene banks and botanical gardens. Aegilops

ventricosa, A. cylindrica and A. uniaristata are considered

species with a medium risk, for which the priority is

medium, both for in situ and ex situ conservation, since

they are reasonably represented in gene banks and botani-

cal gardens.

Aegilops caudata is in a completely different situation,

since we do not know for sure if it is really present in

Italy. Investigations are needed to determine whether it is

present or not. After that, if it is present, it is evident that

it needs immediate in situ conservation action, while the

priority for the ex situ conservation is low, because gene

banks and botanical gardens retain an appreciable number

of accessions. Even the presence of A. speltoides and A.

peregrina are doubtful for Italy, so is ex situ conservation

for the same consideration to be made for A. caudata to

be valid, but being well represented in gene banks the pri-

ority for their ex situ conservation is nil.

Threats

Applying the criteria proposed by IUCN (2012) to

Aegilops, excepting A. geniculata and A. neglecta since

both species are widespread and well distributed, the most

important threats are: (i) livestock farming and ranching �the over-grazing is potentially harmful, especially to popu-

lations of few individuals; (ii) increase in fire frequency/

intensity � fire, in the period before the dissemination, can

Fig. 17. A. ventricosa (near Torino, Piemonte, 2897 - G.F. Piot-taz, FI).

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be very dangerous, especially for species with low capacity

in seed setting; (iii) herbicides and pesticides � populations

growing in the most anthropized sites, such as margins of

field crops, roadsides, etc., are threatened by massive use

of pesticides and phytocides, and by high negative impact

on soil fertility, microflora, etc.; (iv) low densities � some

species depend on very small populations or very few indi-

viduals; (v) population fluctuations � fluctuation or genetic

drift of sub-populations can determine, even in a short

time, a drastic reduction of individuals; and (vi) restricted

range � this may be the case of A. uniaristata, A. biuncia-

lis and A. ventricosa, as far as the Italian populations of A.

species are concerned, it should be stressed that most of

the threats are mainly linked to the small ecological range

of their geographical distribution. Another important threat-

ening factor is the germination capacity, particularly if con-

sidered in relation to the low number of caryopsises

produced by each single plant of some Aegilops species,

especially in adverse environmental conditions (Perrino &

Wagensommer, 2012).

Strategies/actions and norms of conservation

The plant communities of A. biuncialis, A. triuncialis, A.

uniaristata and A. ventricosa identify annual meadows

that may fall within priority habitat of Directive 92/43

EEC ‘Pseudo-steppe with grasses and annuals of the

Thero-Brachypodietea’ (code 6220� - subtype 3 Brachy-

podietalia distachyi) (European Commission DG Envi-

ronment, 2007; San Miguel, 2008; Biondi et al., 2009); A.

geniculata and A. neglecta fall into subnitrophilous Stel-

larietea mediae class; in the case of the species whose

presence was no longer confirmed in Italy (A. speltoides,

A. peregrina and A. caudata) information about vegeta-

tion cannot be provided because it is unavailable.

For the threatened Aegilops species, as for any other

threatened taxa, ex situ conservation in seed banks is

highly recommended. The Germplasm Bank of the Botan-

ical Garden Museum of the University of Bari

(BG-MOBB) preserves seed samples collected from pop-

ulations of A. biuncialis and A. uniaristata growing in the

territory of Bari and Taranto (Apulia).

The small number of seed setting and dissemination

mechanisms of threatened A. species indicate their low

capacity to compete with other plant species, colonize

new areas and to enhance the number of individuals

within the same populations. For these reasons, it is essen-

tial to undertake in situ conservation activities and moni-

toring of the threatened populations at risk, in order to

conserve and protect threatened A. species in their natural

habitats (Kilian et al., 2011).

As far as A. uniaristata is concerned, within the lifeþCENT.OLI.MED. project, guidelines for in situ conserva-

tion were outlined (Perrino et al., 2014). Some of the

most significant are reported here: (i) collection of

caryopsises from plants growing in neighbouring popula-

tions with a high density of individuals; (ii) reinforcement

measures to enhance the number of individuals of the

threatened populations, in collaboration with gene banks,

and (iii) systematic monitoring of threatened populations,

with the aim to improve their in situ conservation.

As for the future, investigations should be made in the

following three directions: (i) checking if species of A. are

‘at risk’, have ‘always’ been there and if they were just

overlooked by previous botanists; (ii) verify if ‘recent’

environmental and climatic changes have created new

ecological niches; and (iii) find out if the species have

developed special adaptation and colonization capacity to

new ecological niches. Since none of these three cases

can be excluded more research on-field and in-lab should

be promoted in order to search for new sites and to check

the taxonomy of the specimens, whenever possible with

the help of molecular tools. Genetic variability correlated

with environmental conditions may show special adapta-

tion patterns. The results of these studies could throw

more light on the taxonomy of controversial species.

ConclusionsRecent reports, herbarium specimens and unpublished

data were used to update the geographical distribution of

Aegilops species in Italy. An emblematic case is that of A.

biuncialis, which for more than a century was reported

only in Apulia and now is known also in Basilicata,

Campania, Veneto and Tuscany. A total of five species

(A. biuncialis, A. cylindrica, A. triuncialis, A. uniaristata

and A. ventricosa) out of the 10 that occur (three of which

are doubtful) in Italy, are reported in the Italian Red List.

One of the threatened species, A. uniaristata, is at risk

also at Community level (van Slageren, 1994). The occur-

rence of A. speltoides, A. peregrina and A. caudata,

should now be considered highly doubtful, because they

have been documented by exsiccata dated before 1900

and recently not found.

Aegilops uniaristata, which grows in Italy only in

Apulia and Basilicata, shows 2�3 fertile spikelets

(Pignatti, 1982) and not 3�5 fertile spikelets per spike as

indicated by van Slageren (1994). Aegilops fragilis, on

the basis of the number of spikelets per spike and the

smaller size of the awns, falls within the description of

A. ventricosa and therefore, it cannot be considered an

autonomous species. If these observations are correct, it

means that we are dealing with the existence of two Italian

ecotypes of A. uniaristata and A. ventricosa.

AcknowledgementsThe authors wish to thank the curators of all cited herbaria

for facilitating the examination of herbarium specimens

346 E. V. Perrino et al.

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and for allowing the reproduction of images, namely: Her-

barium of Botanical Garden Museum of University of

Bari (BI), Herbarium of University of Bologna (BOLO),

Herbarium of Biological Department of University of Cat-

ania (CAT), Herbarium of Natural History Museum �Botanical Section ‘F. Parlatore’ of University of Florence

(FI). Thanks are also due to the reviewers for their recom-

mendations. Finally, the authors wish to thank Dr Jenny

Calabrese for her help in improving the English language

of the manuscript.

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