Upperi^w/w

nebouxii excisa,

Isabela Island

(Galapagos Islands.

Lower iChelo-

noidis nigra spp,

Santa Cruz Island,

C. Darwin Re-

search Center.

Photos by

Francesco, Toscano

Napoli, Italy

The Galapagos Islands. The Galapagos Islands (official name Archipielago de Colon) are

an archipelago of volcanic islands distributed on either side of the Equator in the Pacific

Ocean, 926 km West of continental Ecuador, which they politically belong to. The

Archipelago consists of 1 8 main islands, 3 smaller islands and 107 rocks and islets, with the

surrounding waters being a National Park (since 1959) and a Biological Marine Reserve

(since 1 986). In 1 978, Unesco recognised the islands as a World Heritage Site and, in 1 985, as

a Biosphere Reserve including, later (in 2001), the Marine Reserve. In September 1835 the

Galapagos Islands were reached by the English naturalist Charles Darwin who aboard the

"Beagle" spent more than a month on the exploration of this archipelago. He conducted

numerous obseiwations on geological, botanical and zoological features of these islands

subsequently merged into some of the basics of his theory on the evolution of the species. In

particular, it was obvious to C. Darwin as geographical isolation and adaptation to new

environments were crucial in the processes of speciation. These islands, all of volcanic

origin, were uninhabited in the beginning; subsequently, the various life forms mostly

coming from the South American continent underwent a first process of speciation in the

occupied territories. In many following steps, these species colonised other islands and, from

time to time, adapting to new environments and exploiting the numerous trophic niches,

further diverged. Al! this resulted in the presence of many taxa related to each other but

distinct by form and function which occupied different islands. Darwin, for example,

observed that the species of finches of the Galapagos Islands showed the beak with a different

shape and function as a result of the natural selection. The differentiation within this group

led to the final result of 13 species organized into three separate genera. Another example of

this evolutionary radiation are the "Galapagos giant tortoises", belonging to a single species

(Chelonoidis nigra group) divided into several subspecies depending on the island they

occurrin, nearly all considered as endangered taxa.

Michele Bellavista, Via C. De Grossis, 7 -90135 Palermo, Italy

email: michele.bellavista@gmail.com

Biodiversity Journal, 2012, 3 (4): 263-264

Introduction

The importance of insularity and biodiversity

Roberto Poggi

Museo Civico di Storia Naturale “Giacomo Doria”, via Brigata Liguria, 9 - 16121 Genova, Italy; e-mail; rpoggi(a)comune.genova.it

Proceedings of the International Congress “Insularity and Biodiversity”, May 11 *-13*, 2012 - Palermo (Italy)

The importance of biodiversity is now widely

recognized in the scientific field, but too often

ignored or underestimated by those who, by law,

should defend the territory and its environmental

components, for which we can only applaud any

initiative that wants to emphasize the value of

biological diversity and the need for its careful

protection.

The Congress held in Palermo in May 2012 not

only wanted to examine several examples of bio-

diversity, but also put them in connection with the

study of insularity (Figs. 1, 2), an argument which

has always been beloved to systematists, ecolo-

gists and biogeographers, since each island, big or

small, has the fundamental advantage of being per-

fectly delimited within its boundaries, which

makes it an excellent model for both field analysis

and theoretical interpretations.

The contributions have emphasized some

aspects of island populations (especially those

more closely Mediterranean) from a zoological as

much as botanical point of view, proving once

again, despite the limitation of means that charac-

terizes the current historical moment, the level and

the ability of researchers, either "professional" or

"amateurs", as well as the crucial importance of a

Figure 1. Western Camere islet, North-Eastern Sardinia, Figure 2. Figarolo islet, Aranci Gulf, North-Eastern Sar-

opposite “Costa Smeralda”, 11. IV. 1986, photo R. Poggi. dinia,ll.IX.1987, photo R. Poggi.

264

Roberto Poggi

careful and proper conservation of naturalistic col-

lections, essential reference point for any kind of

investigation that requires dynamic comparisons

between the present and the past.

Finally, relations have also highlighted the

need for the protection of our environmental heri-

tage, too often subjected to aggression dictated by

a momentary individual profit and ignored, or at

least underestimated, by some political, admini-

strative and management structures that, instead,

should be the Paladins of its protection, hoping to

pass on to the future generations the Earth in a

state at least similar to that in which our generation

received it in temporary usufruct.

Biodiversity Journal, 2012, 3 (4): 265-266

Preface

Flora and insular diversity

Francesco M. Raimondo

Orto Botanico and Herbarium Mediterraneum, University of Palermo, Italy; e-mail: franeeseo.raimondo@unipa.it

Proceedings of the U‘ International Congress “Insularity and Biodiversity”, May 11*-13*, 2012 - Palermo (Italy)

As it is well known, the flora of a eontinental

region as that of an island represents the results

of interaetion between aetual environmental fae-

tors and other ones of historieal nature. Therefore,

the flora of any geographieal region eonsists of

the whole of plant speeies that evolved there to-

gether with those that gradually arrived from

other regions through natural veetors.

Indeed, insularity is a eondition eireumseribed

to a ten'estrial spatial eontext, basieally subjeet to

isolation with respeet to another equal, similar,

eontext or else a quite different, being separated

through any barrier. This eondition physieally

eonforms with the geographieal unit defined by

the term “island” (latin insula): the barrier is ge-

nerally the sea or however by water; but it eould

also be represented by quite different environ-

mental system. From a biologieal point of view,

aeeording to Greater (2001) island is an insulating

barrier; mountains surrounded by extensive

plains, bogs and lakes eireumseribed to arid

zones, woody topsoils within the steppe are even

islands. In faet, the flora of the Mediterranean Re-

gion takes its riehness and peeuliarity from the

huge fragmented habitats and isolation pheno-

mena, not always generated by orographieal or

aquatie barriers. The eontributions presented in

the symposium “Flora and insular diversity”

whose proeeedings are presented here, almost

refer to the system "island" in its usual meaning

of territory surrounded by the sea.

The role played by the insular faetor in the ge-

nomie proeesses of plant diversity has been very

earefully followed by the seholars, taxonomists

and phyto-geographers in partieular. On this re-

gard, the Mediterranean biogeographieal region,

whose flora is among the riehest in eomparison

with its geographieal range, whieh is marked by

quite peeuliar events of both elimatie and geolo-

gieal nature and by a huge number of eontinental

island systems, has represented an important la-

boratory for biologieal researehes and interpreta-

tions. Indeed, many studies on this field are

definitely referred to the Mediterranean area.

Among these, I just remember the numerous eon-

tributions by Werner Greuter who in many oeea-

sions treated this topie with speeial regard to

Crete and the Aegean islands, where he earried

many personal surveys (Greuter, 1970, 1972,

1975a, 1975b, 1979, 1980, 1991,1995, 2001).

As remarked by several seholars, insularity is

at the same time a geographieal barrier and an im-

portant faetor in the reproduetive isolation, at

least as the spatial separation is eoneerned. The

reproduetive isolation, aeeording to Gerola

(1995), as it obstruets genes to freely migrate

among populations belonging to the same speeies,

supports eeotypes to assert themselves and then

allowing do new speeies form when they likely

eould not establish themselves without eontinue

interbreeding, by gene migrations from eaeh eeo-

type - or population - to other. These are, there-

266

Francesco M. Raimondo

fore, outside impediments, as defined by George

L. Stebbins, the Ameriean botanist and genetieist,

father of the plant biosistematies, otherwise insu-

perable barriers affeeting both pollination proees-

ses and the dispersal of speeies. Therefore, islands

are at the same time eonservative and ereative for

the plant life.

This statement is eonfirmed by the eontribu-

tions at the symposium “Flora and insular diver-

sity”. These are preminently dealing with topies

eoneerning the plant diversity in the Mediterra-

nean islands. There is also ineluded an additional

eontribution whieh brings into foeus the eonse-

quenees of insularity on the plant evolution under

another both geographieal and elimatie range. In

partieular, the system whieh is flood-lit is the Ar-

ehipelago of Soqotra, in the Indian Oeean, a small

orographieally diversified area, strategieally pla-

eed in front of the Afriean Horn, westwards, and

the Arabian Peninsula eastwards, respeetively.

In eonelusion, the eontributions here presented

eonfirm the rule: wherever plants are loeated, is-

lands are both ereative and eonservative eentre of

diversity, a on the whole eontribute supporting the

biologieal evolution of the Earth.

REFERENCES

Gerola F.M., 1995. Biologia e diversita dei vegetal!.

UTET, Torino.

Greuter W., 1970. Zur Palaogeographie und Floren-

gesehichte der sudlichen Agais. Feddes Report. 81:

233-242.

Greuter W., 1972. The reliet element of the flora of

Crete and its evolutionary signifieance. In: Valentine

D.H. (Ed.), Taxonomy, phytogeography and evolution.

London & New York, pp. 161-177.

Greuter W., 1975a. Die Insel Kreta - eine pflanzen-

geographische Skizze. - Veroff. Geobot. Inst. ETH Stif-

tung Riibel Ziirieh 55: 141-197.

Greuter W., 1975b. Historical phytogeography of the

southern half of the Aegean area. In: Jordanov D., Bon-

dev L, Kozuharov S., Kuzmanov B., Palamarev E. & Vel-

cev V. (Eds.), Problems of Balkan flora and vegetation.

Proceedings of the first international symposium on Bal-

kan flora and vegetation, Varna, June 7-14, 1973. Sofija,

pp. 17-21.

Greuter W., 1979. The origin and evolution of island

floras as exemplified by the Aegean archipelago. In:

Bramwell D. (Ed.), Plants and island. London & New

York, pp. 87-106.

Greuter W., 1980. The endemic flora of Crete and the

significance of its protection. In: Antipas B. (Ed.), Prak-

tika synedriou prostasias panidas-hloridas-biotopon.

Athenai 11-13 Oktobriou 1979, pp. 91-97.

Greuter W., 1991. Botanical diversity, endemism, ra-

rity, and extinction in the Mediterranean area: an analysis

based on the published volumes of Med-Checklist. Bo-

tanica Chronica, 10: 63-79.

Greuter W., 1995. Origin and peculiarities of Medi-

terranean island floras. In Quezel P. (Ed.), Connaissance

et conservation de la flore des lies de la Mediterranee.

Ecologia Mediterranea, 2 1 : 1-10.

Greuter W., 2001. Diversity of Mediterranean island

floras. Bocconea, 13: 55-64.

Biodiversity Journal, 2012, 3 (4): 267-272

Dream islands and island dreams

Alessandro Minelli

Department of Biology, University of Padova, Via Ugo Bassi 58 B, 135131 Padova, Italy; email: alessandro.minelli@unipd.it

ABSTRACT The contribution to the development of biogeography and evolutionary biology offered by

investigations on insular floras and faunas is briefly reviewed. Implications of the dynamic

nature of insular biota for faunistic and floristic research are stressed.

KEY WORDS allopatric speciation; chronogeonemy; dispersal; island biota; vicariance.

Received 12.05.2012; accepted 03.09.2012; printed 30.12.2012

Proceedings of the International Congress “Insularity and Biodiversity”, May 11*-13*, 2012 - Palermo (Italy)

EXPLORING ISLANDS

Ever since Darwin visited the Galapagos Islands

in 1835, the study of insular biotas has offered fac-

tual evidence of critical importance for the develop-

ment of evolutionary biology (e.g., Grant, 1998),

but also unparalleled contributions to biogeography

(e.g., Carlquist, 1966). This is especially true of the

oceanic islands, with the Hawaiian archipelago

eventually becoming a cherished treasure trove for

naturalists and population geneticists alike, ready

to spend their lives studying the archipelago’s hun-

dreds of Drosophila Fallen, 1823 species, or to do-

cument distribution and interrelationships of the

amazingly diverse tree-dwelling snails of the genus

Achatinella Swainson, 1828 (Cook & Kondo,

1960), or to reconstruct the evolutionary history of

the endemic, nectar-feeding drepanid birds. A jump

across the Pacific brings us to Moorea, and to the

local Partula Ferussac, 1821 snails, another group

of land gastropods that features among the most po-

pular taxa in evolutionary biology since Crampton’s

(1916, 1925, 1932) classical investigations.

Several monographic studies of insular biota are

by now classics of evolutionary biology and/or bio-

geography. A very short, selective list includes Al-

fred Russell Wallace’s Island Life (1880), Henry

Brougham Guppy’s (1906) monograph on plant di-

spersal across the Pacific, the books on the Galapa-

gos’ Darwin finches by David Lambert Lack (1945,

1947) and Peter R. Grant (1986), and Sherwin

Carlquist’s comprehensive books (1965, 1974,

1 980) on the origin and adaptations of insular plants

and animals.

Naturalists’ fascination with insular biota helps

understanding the prominent role long acknowled-

ged in biogeography to long-distance dispersal in

explaining distributions.

Dispersalist interpretations have been extensi-

vely challenged in the last few decades, since the

advent of the vicariance models of a new biogeo-

graphy rooted in cladistics, beginning with Brun-

din’s (1966) long essay on transantarctic

relationships; for classic reference texts on vica-

riance biogeography, see Nelson & Platnick (1981),

Nelson & Rosen (1981) and Humphries & Parenti

(1986); for a comparative perspective on biogeo-

graphic models based on vicariance vs. dispersal,

see Morrone & Crisci (1995).

Vicariance models have their obvious merits

when applied to continental biotas. However, when

investigating the origin of insular faunas and floras,

the traditional interpretations based on dispersal still

deserve consideration. Still of interest are a few

268

Alessandro Minelli

classical works on land molluscs of the Pacific area,

sueh as Vagvolgyi (1975), as well as others devoted

to land plants (e.g., Fosberg, 1948, 1956, 1963) or

inseets (e.g., Zimmermann, 1948; Gressitt, 1956,

1961; Gressitt & Yoshimoto, 1963).

CONCEPTUAL MODELS

Sinee Darwin (1859), the study of insular biota

has offered preeious examples of the evolutionary

seenario eurrently known as allopatrie speeiation

(Mayr, 1942; White, 1978; Coyne & Orr, 2004;

Grant & Grant, 2008), but has also stimulated the

formulation of other important eoneepts in evolu-

tionary biology, as the founder effeet (Mayr, 1942;

see also Barton & Charlesworth, 1984) or, more re-

eently, the perhaps less popular notion of taxon

eyele (Wilson, 1959, 1961; see also Rieklefs, 1970;

Whittaker, 1998; Rieklefs & Bermingham, 1999).

As defined by Rieklefs & Bermingham (2002),

“taxon eyeles are sequential phases of expansion and

eontraetion of the ranges of speeies, assoeiated ge-

nerally with shifts in eeologieal distribution. The im-

portant eontribution of the taxon eyele to

biogeographieal analysis is its emphasis on evolutio-

nary and eeologieal interaetions among eolonizing

and resident speeies, whieh influenee their extinetion

dynamies and establish patterns of geographieal di-

stribution. Taxon eyeles were inferred originally

from the distribution of speeies aeross island arehi-

pelagos, where a eorrelation was noted between gaps

in island oeeupaney and the degree of phenotypie dif-

ferentiation. This pattern implied that phases of eo-

lonization were followed by range eontraetion.”

This eoneept offers a bridge between evolutio-

nary biology and biogeography. Within the latter,

however, the single most important eontribution of-

fered by the study of insular biota has been the de-

velopment of the theoretieal models of insular

biogeography (Me Arthur & Wilson, 1963, 1967;

Wilson, 1969), by whieh the speeies eomposition

of insular biota is explained in terms of eombined

dynamies of eolonization and extinetion.

EXPERIMENTAL MODELS

The quantitative nature of MaeArthur and Wil-

son’s models invited soon to test them experimentally.

A first test was performed on a set of very small

and low islands off the eoast of Florida. Here, the

existing fauna was earefiilly inventoried, then it was

eompletely eliminated by fumigation, thus setting

time to zero before starting to reeord (re)eoloniza-

tion and survival/extinetion dynamies (Simberloff,

1969; Simberloff & Wilson, 1969, 1970; Wilson &

Simberloff, 1969). It is fair to remember, however,

that Nature had already offered a broadly similar

opportunity on a grand seale, when the island of

Krakatau was the theatre of the famous dramatie ex-

plosion (1883) that annihilated most if not all of its

original biota. Unfortunately, the loeal fauna and

flora had not been inventoried before the explosion.

However, subsequent reeolonization was studied

over about one eentury and has been eventually do-

eumented and diseussed in important monographs

(Doeters van Leeuwen, 1936; Dammerman, 1948;

Whittaker et al., 1989; Thornton, 1996).

ISLAND AS METAPHORE

In geography, an island is a pieee of land sur-

rounded by water. In eeology, however, there are

many units that deserve be ealled islands beeause

of the sharp boundaries than delimit them in respeet

to a radieally different surrounding landseape,

where the inhabitants of the ‘eeologieal island’

eould not survive permanently. Thus, islands are

bare mountaintops separated from the nearest

mountaintops by green and perhaps forested val-

leys. Eeologieal islands are erater lakes fully lae-

king eonneetion to other water bodies. To some

extent, eaves are also a kind of islands, although

their aetual limits do not eoineide, as a rule, with

the spaees potentially aeeessible to exploration by

humans, beeause of more or less extensive intersti-

tial spaees that may represent inhabitable eorridors

for the subterranean fauna.

Diseussing the mammalian faunas of mountain-

tops, Brown (1971) eharaeterized these as true re-

liets, rather than biota materializing equilibria

between rates of eolonization and extinetion, as pre-

dieted by MaeArthur and Wilson’s model of insular

biogeography. In the same years, however, the po-

tential usefulness of this model in deseribing the

biota of eeologieal islands was demonstrated by Cul-

ver (1970) for subterranean aquatie arthropods and

by Vuilleumier (1970) for the birds eommunities in-

habiting the isolated paramos of the Northern Andes.

Dream islands and island dreams

269

The ‘splendid isolation’ (to use the words

adopted by Simpson (1980) in reeonstrueting the

history of long separation and reeent temporary in-

terehange between the North Ameriean and South

Ameriean faunas) of several faunistieally rieh

lakes has been long studied, revealing striet paral-

lels with the evolution of the faunas (and, to a

more limited extent, the floras) of larger islands

sueh as New Caledonia (Chazeau 1993; Moral,

1993) or Madagasear (Goodman & Benstead,

2003). Let’s just mention Lake Baikal (Kozhov,

1 963), with its unique diversity ranging over fishes,

snails, amphipods, planarians and other groups,

and the great Afriean lakes (Vietoria, Malawi, Tan-

ganyika), whose huge eohorts of eiehlid fishes

have offered some of the most extensively investi-

gated histories in speeiation (see Goldsehmidt,

1996 for a popular aeeount on Lake Vietoria

eiehlids, under the signifieant title of Darwin’s

dreampond). Similarly, extensive karstie areas sueh

as those aeeompanying the Southern margin of the

Alps (in turn, part of a more extensive system ran-

ging from the Pyrenees to the mountains of Anato-

lia) hosts a huge number of strietly loealized

speeies of subterranean arthropods (espeeially ea-

rabids and eholevid beetles among the terrestrial

forms, and amphipods among the aquatie ones)

eomparable to the multiplieity of single-island en-

demies found e.g. in the Hawaiian arehipelago.

One more example of ‘insular’ fragmentation of a

lineage in a eontinental setting is offered from sin-

gle-mountaintop endemies sueh as the numerous

eamaenid snails distributed on mountain ranges of

South-Eastern Australia (Smith, 1984).

A very peeuliar way to use the metaphor of the

eeologieal island is its applieation to host-parasite

relationships (Brooks, 1979), where parasites (or

parasitoids) are deseribed as potential eolonizers of

‘islands’ represented by the available host speeies,

on eaeh of whieh the eolonizer may eventually sur-

vive, more or less permanently, or go rapidly extiet.

WHY ARE ISLAND A NATURALIST’S

DREAM MODELS?

The most remote oeeanie islands are at the same

time a eherished subjeet of everyman’s dreams, and

exemplary loeations where to study speeiation and

dispersal.

However, if on the emotional level the attraeti-

veness of islands is fundamentally dependent on the

adventurous journey we must undertake before put-

ting our foot on them, an island’s attraetiveness for

the biogeographer or the evolutionary biologist lies

foremost in its eleareut physieal boundaries.

Narrow straits between seleeted islands have al-

lowed biogeographers to fix the borderline between

the Oriental and the Australian Regions, although

without unanimity of eonsent - the ehoiee being of-

fered ineluding Wallaee’s line, based on mammals

and birds, and plaeed between Bali and Lombok

(Wallaee, 1876), and the more eastward traeed lines

of Weber and Lydekker (ef Mayr, 1944; van Stee-

nis, 1950; Whitmore, 1981).

Having preeise boundaries means also that we

ean preeisely determine an island’s distanee from

other islands, or from the nearest eontinent - basie

parameters in MaeArthur and Wilson’s model.

Good for biology - in so far as using number means

seientifie, modem, safe.

FROM DREAM TO REAL LIFE

Remote islands, however, are diffieult targets

for potential eolonizers. With inereasing distanee

from a potential souree (a eontinent, or another is-

land), the proeess of eolonization goes on progres-

sively slower, thus offering limited ehanee to

eompensate for extinetion. On the other hand, ex-

tinetion is always happening, espeeially on small

islands, where it may be diffieult for many animals

and plants to maintain a population of viable size.

The problem may beeome espeeially aeute for spe-

eies that have developed the prima faeie paradoxi-

eal syndrome of insular gigantism. Among

animals, this is exemplified by the huge tortoises

of Aldabra and the Galapagos. More numerous

examples are offered by plants, where several ge-

nera represented on eontinents by small herbaeeous

speeies have evolved giant, woody representatives

on several arehipelagos. There are many examples

among the Asteraeeae (Hemsley, 1885; Carlquist,

1974, 1980), e.g. Bidens Linnaeus, 1753 with giant

speeies in Southern Polynesia, a phenomenon also

found in some Senecio Linnaeus, 1753 of New

Zealand, and some Centaurea Linnaeus, 1753 of

the Canary Islands, not to mention the extraordi-

nary Hawaiian eomposites traditionally elassified

270

Alessandro Minelli

in the genus Wilkesia A. Gray, 1852 mid Argyroxi-

phium DC, 1836 (Keek, 1936; Carlquist, 1980).

Further example are provided e.g. by Echium Lin-

naeus, 1753 (Boraginaeeae) and Euphorbia Lin-

naeus, 1753 (Euphorbiaeeae) in Madeira, the

Canary Islands and the Cape Verde Islands.

Additionally, an island’s very existenee is also

to some extent preearious. The Hawaiian arehipe-

lago provides a dramatie example of the regular

eyele through whieh individual voleanie islands

first emerge from the sea, then grow through repea-

ted eruptions but eventually lose eonneetion to the

deep magmatie ehannels and start being demoli-

shed, redueing to small emerged roeks to be even-

tually eaneelled from the world’s list of islands.

There is no need to offer detail about the deva-

stating eonsequenees on insular biotas, of oeeanie

islands espeeially, of human temporary or pemia-

nent settlement, with the aeeompanying introdue-

tion of exotie animals and plants. With the arrival

of man, the faunistie and floristie history of islands

turns soon into a history of extinetions. Islands fea-

ture very prominently indeed in all surveys on plant

and espeeially animal extinetions in historieal times

(for a summary, see Balouet, 1990; worth reading

is still Greenway ’s (1956) old, but exemplary mo-

nograph on the extinet birds of the world), as well

as in the major works on introdueed speeies (for

exemple. Long’s (1981, 2003) monographs, on in-

trodueed birds and mammals, respeetively). Aleo-

ver et al. (1998) ealeulated that the arrival of

humans is direetly or indireetly responsible for the

extinetion of at least 27% of autoehthonous mam-

mal speeies that evolved on the world's oeeanie and

oeeanie-like islands, a pereentage rising to 35%

when flying mammals are exeluded.

As a eonsequenee of their intrinsie fragility, in-

sular biota deserve speeial priorities in eonservation

planning (Diamond, 1975).

There are interesting lessons to be learned from

these dramatie stories at global level - lessons, in-

deed, that should inform the biogeographer in

his/her daily work at doeumenting and interpreting

speeies distributions. The eore lesson is, that loeal

speeies eomposition may ehange at a paee we ean-

not ignore. The list of all plant or bird speeies re-

eorded for a given area over one eentury may not

eorrespond to that area’s aetual flora or avifauna at

any time. You ean not (or should not) imagine

an eeologieal network involving all those spe-

eies - they have never been all together in the area

you are investigating, and quite probably will never

be in the future. Geographie preeision in reeording

findings is eertainly preeious, but it may be inade-

quate, if the spatial information is not aeeompanied

by the reeord’s date. Distribution maps ignoring

time of reeorded presenees must be replaeed by

ehronogeonemies (Brandmayr et al., 2006). The

amazing turnover in the beetle fauna reeorded over

just fifteen years by Owen (1991) in her garden in

Cambridge may well serve as a warning against the

eeologieal implausibility of total speeies lists for an

island, be it geographieal or eeologieal, aeeumulated

through deeades of observations.

REFERENCES

Alcover J.A., Sans A. & Palmer M., 1998. The extent of

extinctions of mammals on islands. Journal of Bio-

geography, 25: 913-918.

Balouet, J.-C., 1990. Extinct species of the world. Les-

sons for our future. Letts, London, 192 pp.

Barton N.H. & Charlesworth B., 1984. Genetic revolu-

tions, founder effects, and speciation. Annual Review

of Ecology and Systematics, 15: 133-164.

Brandmayr R, Casale A., Puzzo F. & Scalercio S., 2006.

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Biodiversity Journal, 2012, 3 (4): 273-280

Species richness in isolated environments: a consideration on

the effect of time

Genuario Belmonte

Laboratory of Zoogeography and Fauna, DiSTeBA, University of the Salento - 73100 Leeee, Italy

ABSTRACT The widely aeeepted MacArthur & Wilson model of island biogeography proposes a number

of species that, after an initial growth, stabilize on each oceanic island at an equilibrium point.

This species number depends on the available space, the vicinity of mainland, and the habitat

diversification, thus being directly correlated with space characteristics. This space based

model, however, does not explain some astonishingly evidences of species richness. The story

length (the age) of each environment, possibly associated to the stability of conditions, should

offer a better interpretation of species richness in each situation. Lakes, as water islands, more

than land islands, have been considered in the present review as evidences of such an affection

of the time on the species richness. The high species richness in the most ancient lakes is pro-

bably completely due to the genetic drift which produces diversification within each population

possibly without any dependence from variability of conditions and habitats.

KEY WORDS Ancient lakes; Evolutionary time; Island biodiversity; species richness.

Received 12.05.2012; accepted 20.08.2012; printed 30.12.2012

Proceedings of the L‘ International Congress “Insularity and Biodiversity”, May 11*-13*, 2012 - Palermo (Italy)

INTRODUCTION

The ecological frame: space and unstable

conditions as responsible of species richness

Hutchinson (1959) justified the embarrassing

abundanee of speeies on the earth with the exi-

stenee of trophie nets. Aeeording to this proposal,

small sized speeies, eharaeterized by short life

span whieh does not eneompass many develop-

mental stages, ean eoneentrate themselves upon

narrower trophie speeializations, thus living toge-

ther in high numbers in the same site, without

eompetition. Complexity of trophie nets was in-

tended as higher when speeialization oeeurs, sueh

a speeialization being the possible result of evolu-

tion under stable environmental eonditions.

This assumption indireetly gave a role to the

time: is the environmental stability (intended as

time passed under invariant eonditions) whieh al-

lows the existenee of eomplex trophie nets.

But this implieit admission was never evident; in

faet, Hutehinson did not develop it in the future.

Only 2 years later the same seientist (Hutehinson,

1961) was eonvineed that the stability (homogeneity)

of the water environment eould not explain the high

biodiversity into the plankton, for whieh the seaso-

nality (time instability) eould be the aetual reason.

In aeeordanee with the prineiple of eompetitive

exelusion (Cause, 1934), only variation allows the

eo-existenee of eeologieally similar speeies (= eon-

stant eonditions lead to the affirmation of only one

subjeet in eaeh eeologieal role). In the water of the

pelagie habitat, mieroalgae of phytoplankton do not

affirm themselves beeause in a variable spaee, but

probably beeause in a variable time, so avoiding the

eompetitive exelusion. The time was a determining

faetor, but the variability, and not stability, was the

eause affeeting the speeies riehness. Probably also

due to this double, eontradietory time implieation

in the justifieation of speeies riehness, sueeessive

eeologists did not take eare of the time role.

274

Genuario Belmonte

To tell the truth, the role of the evolutionary time

in the biological diversification (enrichment) of the

planet is probably given as obvious and, avoiding

to discuss it, scientists wanted to demonstrate how

many and what causes were involved in addition.

Sepkowsky (1984) demonstrated that the num-

ber of taxa has a positive, although interrupted,

trend in the whole earth story. On the other hand,

however, Gould (1989) pointed out that after the

Cambrian explosion, only extinctions occurred on

the earth, regarding phyla, and the biodiversity

growth was relative to exaggerated speciation in-

side one or few phyla. Along the evolutionary time,

the bio-richness at level of phylum was considered

as impoverished notwithstanding the evident

growth of the species number.

Coming back to the Cause principle, just to cer-

tify that time for ecologists is not the same entity of

that for evolutionists, it implicitly admits an impo-

verishment of the species number with time (the

ecological exclusion destroys locally the diversifi-

cation if conditions became constant).

After Hutchinson, in ’60s of the XX century, the

scene has been monopolized by the model of Ma-

cArthur and Wilson (1963; 1967) where the species

number possibly existing on an island is linked to

the available space (island size) and to the distance

from the continent (both problems of space). The

role of time in that model is limited just to what

would be necessary to reach the equilibrium point

(to replenish the available space). The model is true

only for newly formed islands, e.g. for the oceanic

ones. What could change in an oceanic island at its

biological equilibrium, when new arrivals equal the

extinctions, is the type of species present, not their

number. A number of studies tending to demon-

strate the validity of this rule on a general scale

were realized in the successive 20 years considering

mountain tips, lakes, marine life around the islands,

as isolated biota subjected to the same rule.

The consideration of temporary ponds as islands

added also a time point of view to the problem. Ac-

cording Ebert & Balko (1987) the length of perio-

dical existences of temporary ponds (the so called

hydroperiod) affects the number of species possibly

present in each pond independently from the basin

size. The Authors proposed a time based re-inter-

pretation of the MacArthur & Wilson space based

model with the area dimension corresponding to pe-

riods of existence of the pond, and distances from

continent corresponding to periods of water ab-

sence. The time they considered, however, was an

ecological factor, because related to something as

the seasonal absence/presence cycles, and not use-

ful for the realization of evolutionary modifications.

In a book, Rosenzweig (1995) describes as in

the periphery of the geographic distribution of large

populations (where the pangamy is not real) small

populations differentiate, giving new species with

time. The process, intrinsic to the population dyna-

mics, is completely disconnected from the environ-

mental diversification and it occurs also in

homogeneous habitats.

Notwithstanding this evolutionary (timely) ap-

proach to the problem of the species richness, Ro-

senzweig officially adhered to the hypothesis of

Terborgh (1973) who gave to the land extension the

responsibility for the geographic distribution of spe-

cies on the whole earth. In fact, even if is only the

time which can allow the realization of genetic dif-

ferences and their affirmation, for Rosenzweig this

can be possible only for large populations, hence

for those populations which occupy large spaces.

More extended areas contain a larger number of

rare species because each lonely individual in each

site is connected to other scattered individuals di-

stributed in a larger space. On the contrary, a lonely

individual on an island (a delimited space) cannot

maintain such a connection with the rest of its own

population and the species will extinguish with his

death on that island.

Terborgh (1973) and Rosenzweig (1995) did not

affirm explicitly what this can represent for the spe-

cies number on each isolated system, but it is easy

to deduce that when a continental island is formed

(e.g. with the sea level uplift) the number of species

can only diminish (for the extinction of the rare

ones) from that moment onward, until a new situa-

tion of equilibrium. This is just the opposite case of

the species richness on oceanic islands (it grows

until the equilibrium).

The hypothesis of Rosenzweig, however, admits

a role of the time on the variability of the species

number, absolutely independent from the environ-

mental variations. Species richness spontaneously

evolves towards a growth due to a mechanism in-

trinsic to the population dynamics. In addition, and

in opposition with the Hutchinson’s thought, the

Rosenzweig ’s hypothesis is funded on the sympa-

tric/parapatric speciation.

Species richness in isolated environments: a consideration on the effect of time

275

Figure 1. Hypothesis of the speeies number growth in a delimited area (island). A, the area is empty. B, one speeies esta-

blishes sueeessfully on the area, oeeupying as mueh as possible spaee, with a high number of individuals. C, aeeording

the Rosenzweig's Hypothesis, periphery subpopulations undergo genetie drifts due to the impossibility that pangamy is

realized. This situation will give more speeies than the equilibrium situation due to the model of MaeArthur and Wilson,

without neither immigration, nor environment variability.

This evolutionist position appeared in a eontext

of great sueeess for Eeology whieh aeeepted wi-

thout problems the Connell (1978) theory of the in-

termediate disturbanee, evident deseent of the

eontinuous instability of Hutehinson, as the main

responsible of speeies riehness and/or eeologieal di-

versity in eaeh geographie site.

Henee Rosenzweig opposes his evolutionary

theory (there are more speeies in large areas where

large populations ean exist, and this is true also

under eonstant eonditions) to the eeologieal one of

Connell (there are more speeies where the environ-

ment is variable and subjeeted to intermediate di-

sturbanee). One thesis affinns that speeies riehness

is independent by environmental eonditions, al-

though favoured in mature and stable situations; the

other thesis affirms that speeies ric hn ess is favoured

by variations whieh maintain in a state of immatu-

rity the system (Fig. 1).

Isolated environments as witnesses of the

role of time and stability on species richness

Lyneh (1988) summarized the refugia hypothe-

sis from whieh the present biodiversity eould de-

rive. During a fragmentation period, speeies were

separated in many and not eomiuunieating popula-

tions whieh had the time to aeeumulate genetie no-

velties with the meehanism of the genetie drift (in

eondition of allopatry). The same interpretation

(fragmentation of the system leading to isolation of

many populations) has been proposed also for the

floek speeiation of Ciehlidae fish in the Afiriean rift

lakes (Sturmbauer, 1998).

What has been always distant from the eeologi-

eal interpretations is just the island situation and the

existenee of aneient lakes (water islands). These last

evidently do not follow the MaeArthur & Wilson

rules, and the eeologieally based predietions on the

number of speeies they should host are constantly

disobeyed. The most aneient lakes today existing

on the earth are all eharaeterised by an elevated

number of speeies, eoupled with an elevated ende-

mism. They eonflrm so simply the dependenee of

speeies number from the time, an uneonsidered ele-

ment from the island biogeography model of Ma-

eArthur & Wilson.

It remains to be diseussed if the numerieal

growth of speeies ean be attributed to the eondition

variability or to the stability in the time of the lake’s

lifespan. Lakes as Bajkal, Tanganika, Malawi, Vie-

toria, Biwa, Ohrid, seem to show a speeies riehness

direetly proportional to their own age, more than to

their geographie position, their size, and/or episodes

of fragmentation whieh eould be sueeeeded during

their existenee. The dependenee of the speeies num-

ber from the time (on an evolutionary seale) is evi-

dent beeause sites with high speeies number (the

276

Genuario Belmonte

mentioned lakes) show also an endemism grade di-

reetly eorrelated with their presumed age. Conse-

quently, speeies present in sueh environments did

not arrived there from elsewhere, beeause they for-

med in situ aeeording to a meehanism (speeiation)

whieh required time and, aeeording to the Rosen-

zweig hypothesis, they simply need large initial po-

pulations.

The evolutionary phenomenon of speeiation pro-

duees endemism, and needs, in lakes, probably more

than 10,000 years beeause postglaeial lakes (formed

from 10,000 years ago) in the boreal hemisphere

have not endemies among their speeies. The neees-

sary time for evolution is, on the other hand, eom-

pletely eompatible with voleanie islands of the

Mae Arthur & Wilson model. For example, Galapa-

gos or Hawaii, well known for the studies on the

presenee of endemies, have ages of some million

years. Howarth (1990) admitted the time as main re-

sponsible of the high speeies riehness of Hawaiian

Drosophilidae (at least 1000 speeies reeognized),

giving importanee not only to the biota fragmenta-

tion (the Hawaiian arehipelago) but, more, to the

presenee of similar eonditions on all the arehipelago

islands, during million years of story.

Briggs (1995) reports as the marine fauna ri-

ehness from the two islands Saint Helen and Aseen-

sion, notwithstanding they should be eolonized

aeeording to the MaeArthur & Wilson model,

shows numbers eompatible with the different ages

of the two islands, and not with their distanee from

the eontinent, or their size. Notwithstanding Aseen-

sion is eloser than St Helen to the Afriean eontinent

(1500 Km against 1850 Km), it has less fish spe-

eies. If we look at the position of the two islands re-

latively to the middle Atlantie ridge, we diseover

that Aseension is eloser to this fraeture line, henee

it is younger (it is the last whieh has been formed,

being the elosest to the ridge).

In the marine dominion also other environments

add data to the time-speeies model. One of these si-

tuation is the eontinental slope between 500 and

2000 m depth. Some pioneer data of benthonolo-

gists have astonished the seientifie eommunity sho-

wing the diversifieation of sueh an environment.

Here not only speeies are numerous (862 speeies on

a total sampled area of only 2 1 m^), but they some-

times reaeh maxima of possible abundanee, with no

speeies more abundant than another (Grassle,

1991). Also here the reason was searehed among

eeologieal eues and the intermediate disturbanee of

Connel (1978) was eonsidered as the only possible

eause. Nobody has never found what is the inter-

mediate disturbanee whieh determines sueh a situa-

tion, but the eontinental slope has been eonsidered

as inevitably interested by turbid falls of the just

strength, extension, and frequeney, to authorize

sueh a speeies diversifieation.

Continental slopes are, however, the most an-

eient marine habitats of the planet, and depths

below 1000 m are those where temperature is inva-

riably fixed around 3 °C, where the light is eomple-

tely absent, differenees of pressures are relatively

minimum, and probably also turbulenee is weak.

Due to the eontinental drift whieh is opening the

North Atlantie Oeean, the slope studied by Grassle

is opening from about 190 million years. While, ho-

wever, surfaee eoastal areas suffer the variability of

seasons and elimates, the eited slopes (1000 - 2000

m depth) represent the zones of maximum age

where present eonditions has been realized first

(and they are possibly remained eonstant).

The time seems the most responsible of speeies

ric hn ess aeeording the Rosenzweig’s hypothesis of

spontaneous formation of speeies from a large ini-

tial population, and the strongest allianee for this

results eomes from the stability of eonditions whieh

should stay there. This is eompletely opposite to the

eeologieal monopoly of reasons to explain speeies

riehness (spaee or time variability, intermediate di-

sturbanee, fraetioning of habitats, and so on) (Fig.

2). Unfortunately, the short time seale (1-50 years)

whieh our studies ean use to establish a rule, has not

efiieaey from the evolutionary point of view.

The area of an environment, or its variability,

ean sure be the eause for a temporary enriehment

of speeies, eoming from elsewhere. But all the bio-

geography ean be better understood if interpreted

in terms of environment ages. Latitudinal and alti-

tudinal gradients are strietly eorrelated with the age

of environments, due to the faet that eold environ-

ments (at high latitudes and/or high altitudes) have

obligatorily less years than the warm ones, due to

the last iee expansion during the last glaeial era.

The polar eaps retraetion on high latitudes, and

the rising on high altitudes of the mountain glaeiers,

are the eertifieation of the young age of nowadays

periglaeial isolated environments. On a short time

seale, the effeets of elimate variations, of the human

aetivities, and of eeologieal sueeessions, are all mo-

Species richness in isolated environments: a consideration on the effect of time

211

0 20 40 60 80 100 120 140 160 180 200 220 240 260 280

Figure 2. A map of Atlantic Ocean with colours indicating the different ages of its bottoms (see the reported colour scale from

0 to 280 million years). The yellow square indicates the area from where Grassle (1991) described astonishing high species

richness. Four arm s star indicates the position of Ascension island. Five arm s star indicates the position of St.Helen island.

dem effects, and field of Ecology, which subtract

attention to the evolutionary (long time scale) field.

Islands (isolated environments) of millions

years of age can offer the possibility to easily eva-

luate the correlation of species number with the sy-

stem age. Such a challenge has been accepted by

Limnologists more than land Island Biogeogra-

phers. The ancient lakes allow us to explain the spe-

cies richness avoiding to search a solution into the

dispersal/immigration and to point on the in situ

speciation.

The case of lake Aral

The Aral Sea (today reduced at 14% of its past

extension) had a surface about 20 times that of lake

Ohrid, in the Balkans. This notwithstanding, and

contrarily to the rule which wants the area correla-

ted with the species number, Aral has an evident

low species richness (maximum 310 species accor-

ding to Aladin, 1995) if compared with the "small"

Ohrid (about 1200 according to Albrecht & Wilke,

2008). Reading from more sources it is possible to

advance the hypothesis that the Aral did not exist

in ancient historical times, and its story has only

some centuries.

The expert Arab geographers of the middle age

did not indicate a great lake in the position of the

present Aral, and the lake has been reported on

maps only from Russians after the conquer of those

steppe lands after 1620. The history tells about the

Sultan of Khiwa who turned the Oxus river into the

desert to avoid it could be used by the Czar's sol-

diers to individuate the city and invade it. Oxus is

very possibly what now is called Amu Daria, the

river southern tributary of the Aral; and it possibly

278

Genuario Belmonte

arrived into the Caspian Sea, from where the Czar

soldiers easily eould reaeh Khiwa. This is probably

only a hypothesis to be confirmed, but the present

retraction of the Aral leaved dried lake bottoms

where in 2004 an ancient city with architectural

remnants of the XIII-XIV century has been reco-

gnized. Hence it is now evident that the Aral Sea,

as we remember it, was a young environment and,

although exaggeratedly large, the species it hosted

were only those which arrived from elsewhere, ac-

cording the process described by MacArthur & Wil-

son, and no one of them had the time to undergo the

genetic process of the multiple speciation.

Dodson (1991; 1992) confirmed the ecological

rule area/species number on the crustacean species

abundance in lakes of the northern hemisphere. But

this Author considered mostly postglacial lakes,

hence those (as the majority of boreal lakes) formed

in the post-glacial era, hence with a comparable age

(20,000 - 10,000 years).

In the study of 32 European lakes (Dodson,

1991) the regression line on the area/species log

plot, shows an evident scattering distribution of

points around the x value (area) of about 1x10^ km^

where it is easy to recognize the Finstertaler (a

dam), the Port Bielh (at 2700 m above the sea

level), and the Latnjajaure (near the Arctic polar cir-

cle), as the youngest lakes and at the lowest y (log

species) values; a different hypothesis could be at-

tempted with lakes arranged according an age/spe-

cies plot (Fig. 3).

The growth of species number with time is ob-

vious at the start of the story of each island (and

lake). The Krakatau "son" (an island remnant from

the explosion of 1 883) has been considered as re-

plenished of species after not more than 35 years.

Simberloff & Wilson (1970) demonstrated the re-

storation of the insect number in experimentally de-

fauned islets after only 2 years. Also a study on

dams (lakes of a young, known, age) in southern

Italy (Alfonso et al., 2010) confirmed that the spe-

cies number in the plankton grows until 50 years

from the lake birth, but no more. The time necessary

to the biological replenishment of newly birth is-

lands is relatively short but, however, it is only the

first part of the story. Successively to the reaching

of the equilibrium, the number of species stop to

grow only apparently. This number grows with a

slower rhythm, over a longer time, and only great

age differences among islands can unveil this time

effect. Aral, in Kirghiz language, means Island. The

species richness of this large island of water was

just that which has been possible to recruit from

water environments geographically around.

According MacArthur & Wilson they reached

the maximum number possible at an equilibrium

point between new immigrants and local extincts;

the large area allowed the presence of demographi-

Figure 3. The Dodson’s correlation Area/Species as reported from 32 European lakes. Lakes of about 1x10^ Km^ are vi-

sibly scattered around the regression line. Fin is the Finstertaler dam (less than 100 years old), PBi is Port Bielh, an

alpine lake at 2700 m above the sea level (less than 6000 years old), Lat is the Latnjajaure (near the Arctic polar circle,

less than 6000 years old)

Species richness in isolated environments: a consideration on the effect of time

279

cally enormous populations, but the lake age (400-

500 years) was too short to allow the system to

enter the phase of the speeiation even for single

groups (the speeies floek) as Rosenzweig suggested

as possible. The ease of the Island=Aral also sug-

gests that, differently from what proposed by Hut-

ehinson (1959) at least for long time intervals

(million years) the speeies riehness is probably di-

reetly eorrelated with the stability of the system (in

the ease of Aral its existenee). The speeiation ae-

eording the model of Rosenzweig, in addition,

needs not variability of eonditions. The Age/Speeies

eurve, henee, has to be eorreeted into a sum of spe-

eies eurves aeting eaeh for a different time duration,

and nothing ean be said on the existenee of a limit

to sueh a number (Fig. 4).

Aeeording to the reeent model of Hubbell (2001)

on a neutral theory of speeies geographie distribu-

tion, it exists a speeies number, typieal for eaeh en-

vironment, whieh depends not simply upon the

MaeArthur & Wilson model, but also on speeiation

and evolution. This number, however, obeys to a sort

of zero-sum game where the number of speeies ean-

not exeeed the earrying eapaeity of the system. If the

earrying eapaeity is a biomass value, the number of

speeies ean grow in eombination with a deerease of

the number of individuals per population, and/or a

deerease of the size for each species.

But this model deserves an appropriate set of

data to be completely accepted.

CONCLUSIONS

The time has undoubtedly an effect on the spe-

cies number which stay in an environment. This ef-

fect, however, is not simply that indicated by

MaeArthur & Wilson on oceanic islands, due to the

immigration/extinction rate of species. The role of

the time on the species richness starts to be reco-

gnizable only on isolated environments where con-

taminations from neighbors are limited or null, and

after many years to allow the evolution to do its job.

This action is so slow that it cannot be recogni-

zed in terms of ecological times, and even a limit

(a maximum number of species) is probably diffi-

cult to be found. Time, however, elegantly explains

the disobeying faunal situation of St. Helen and

Ascension in the middle of the Atlantic, the species

richness of ancient lakes (or the species poorness of

Aral), the latitudinal gradient of the species number,

the species diminution with the altitude, the astoni-

shing species richness of benthic fauna on the con-

tinental slope of the north Atlantic Ocean.

The temptation to re-describe each pattern and

geographic situation is great. In Italy, for example.

Figure 4. The species number in each island grows according different phases: (black) early colonization, arrival of pioneer

species (r-strategists) which sustain the successive immigration (Ecological Succession); (purple) immigration from nei-

ghbors (MaeArthur and Wilson model) up to a constant number (the equilibrium point) of species (k strategists) where

new arrivals equal extinctions; (red) speeiation and species flock appearences. The species number grows for intrinsic,

evolutionary reasons and independently from the ecological ones.

280

Genuario Belmonte

a series of faunal geographie provinees have been

reeognized on the eeologieal basis of the elimate.

If we look at the lakes, however, we ean note as

the Padano-veneta provinee has large lakes at low

altitudes, formed at the beginning of the iee retrae-

tion (about 40,000 years ago), whereas the Alpine

provinee hosts small lakes over the tree altitudinal

line, henee iee free from only 5,000-7,000 years.

Along the Italian peninsula (Apennine pro-

vinee) are eoneentrated several Voleanie water ba-

sins whieh have ages of hundred thousand years,

and at the extreme south, and on islands (insular

provinees), the most of the lakes have been reali-

zed only reeently with dam buildings (ages lower

than 100 years). In this brief ease the biogeogra-

phie subdivision of a territory eould be based not

on eeology, but on environment age, and it propo-

ses the age of environments as an alternative key

in the understanding speeies distribution, other

than the speeies riehness.

In any ease is probably time to start with a

deeper eonsideration of the age effeet in the bio-

geographie framework.

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MacArthur R.H. & Wilson E.O., 1963. An equilibrium

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MaeArthur R.H. & Wilson E.O., 1967. The theory of is-

land biogeography. Princeton University Press, New

Jersey. 224 pp.

Rosenzweig M.L., 1995. Species Diversity in Space and

Time. Cambridge University Press. 436 pp.

Sepkowsky J.J.Jr., 1984. Akinetic model of Phanerozoic

taxonomie diversity. III. Post-Paleozoic families and

mass extinetions. Paleobiology, 10: 246-267.

Simberloff D.S. & Wilson E.O.,1970. Experimental zoo-

geography of island. A two year record of coloniza-

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Sturmbauer C., 1998. Explosive Speeiation in ciehlid

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Biodiversity Journal, 2012, 3 (4): 281-296

Relict species and the challenges for conservation: the emble-

matic case of Zelkova sicula Di Pasquale, Garfi et Quezel and

the efforts to save it from extinction

Giuseppe Garfi' & Stephane Buord^

'Istituto di Genetica Vegetale, Consiglio Nazionale delle Ricerche, corso Calatafimi 414 - 90129 Palermo, Italy; email:

giuseppe.garfi@igv.enr.it

^Conservatoire Botanique National de Brest, 52, allee du Bot - 29200 Brest, Franee; email: s.buord@ebnbrest.eom

ABSTRACT The foreseen rapid climate changes are thought to represent a major threat for many plant spe-

cies. Owing to that, in the Mediterranean Basin, one the most important area worldwide for

biodiversity, a number of taxa are at risk of extinction, especially the endemics and the rarest

ones. Within them a particular significance is recognised to the pre-glacial relicts, that are rather

conspicuous in the area and represent a valuable heritage for their biogeographical relevance

and evolutionary history. Zelkova sicula is one of the most prominent plants within this flora.

To date, it is a very rare species, and due to a number of threatening factors it is on the brink

of extinction. In the present paper, we discuss the main topics related to its conservation criti-

calities and the ongoing integrated strategies to save it from extinction. Particular emphasis is

addressed to assisted colonisation, a “last resort” conservation approach consisting in the esta-

blishment of pilot-planting out of the species native range.

KEY WORDS Assisted colonisation; clonality; Life+ EC programme; Sicily; threatened plants

Received 11.05.2012; accepted 20.12.2012; printed 30.12.2012.

Proceedings of the P' International Congress “Insularity and Biodiversity”, May 11*-13*, 2012 - Palermo (Italy)

INTRODUCTION

In the Mediterranean area a number of rare and

threatened island plants have been reeently the ob-

jeet of a eampaign of awareness for their eonserva-

tion (Montmollin & Strahm, 2005). The eampaign

was promoted by lUCN in order to mobilize eon-

servation praetitioners, deeision makers and the ge-

neral publie to take aetions needed to save from

extinetion the most endangered speeies. Among the

fifty speeies eonsidered, as mueh as ten were selee-

ted from Sieily and the surrounding islets, so highli-

ghting the notable signifieanee of this region as a

major biodiversity hotspot within the whole Medi-

terranen Basin (Quezel & Medail, 2003).

Aeeording to the lUCN Red List Categories and

Criteria (lUCN, 2001), nine out of the ten Sieilian

speeies therein ineluded are quoted as Critieally En-

dangered (CR) mainly due to their extremely re-

strieted area and/or the low number of surviving

individuals. Zelkova sicula Di Pasquale, Garfi et

Quezel is one of the most renowned and prominent

taxa owed to both its biogeographieal relevanee and

evolutionary history.

First deseribed as a new speeies to the seienee

in 1991 (Di Pasquale et al., 1992), Z. sicula is a gla-

eial reliet tree, taking part in the eonspieuous eon-

tingent of Sieilian reliets plants - e.g. Abies

nebrodensis (Lojae.) Mattel, Bupleurum dianthifo-

lium Guss., Cytisus aeolicus Guss. ex Lindl., Erica

282

Giuseppe Garf'i & Stephane Buord

sicula Guss., Petagnaea gussonei (Sprengel) Rau-

schert, Pseudoscabiosa limonifolia (Vahl) Devesa,

Thymus nitidus (Guss.) Jalas, etc. - that notably con-

tribute to the world evolutionary inheritance, and

remarkably significant for conservation of plant di-

versity (Petit et al., 2005).

In the present paper we outline an overview

about the special meaning of biodiversity in insular

environments, with some remarks on relict species

and their vulnerability particularly in the Mediter-

ranean. Within this issue, the case of Z. sicula, as

an outstanding representative of a relict and most

threatened flora, will be discussed especially with

respect to the main topics related to its conservation

criticalities and the ongoing strategies to take it

away from the brink of extinction.

BIODIVERSITY, RELICT SPECIES AND

THREATS IN INSULAR ENVIRONMENTS

Biodiversity hotspots: the preponderance of

insular environments and their vulnerability

Conservation International identified 34 priority

zones for conservation. These hotspots are concer-

ning only 2.3% of earth’s area but gather more than

50% of plant and 45% of animal species of the pla-

net. They all are threatened by human activities. For

the most part, these hotspots are made, totally or

partially, of islands and archipelagos, such as New

Zealand, East Melanesia, Polynesia, Micronesia,

Japan, Philippines, Indonesia, Sri Lanka, Madaga-

scar and Indian Ocean islands, Caribbean islands,

Mediterranean Basin islands, etc..

Insular species are witnesses of a unique evo-

lutionary history. The endemism rate in insular en-

vironments is much higher than in continents. For

example, the Hawaiian archipelago counts 90% of

endemic species, whereas 50% of vascular plants

are endemic to Mauritius and 93% to Madagascar,

which contains as much as 8000 endemics. Given

these endemism rates, insular biodiversity repre-

sents a very important part of world biodiversity,

which has no link with the small area covered by

these islands.

Planetary insular environments are particularly

fragile. Since the sixteenth century and European

expansion, the impact of human activities (agricul-

ture, urbanisation, introduction of invasive species.

etc.) has caused a lot of damages on insular biodi-

versity. Currently, islands figure amongst the most

threatened territories on the earth. In this way, 2017

out of over 7000 plant species of the Caribbean hot-

spot are threatened or even disappeared. Similarly,

among 3334 Polynesian and Micronesian species,

926 are currently threatened (Brooks et al., 2002).

The recession of natural insular habitats is even

more informative. The original forest which cove-

red Mauritius currently represents just 5% of the

total area of the island. In Madagascar, human oc-

cupation led to a diminution of 90% of the original

forest in only 200 years.

Peculiarity and viability of the Mediterra-

nean Basin flora

The five biggest Mediterranean biomes, i.e. Me-

diterranean Basin, California, Mediterranean Chile,

South Africa and Southwest Australia, count 50, 48,

50, 68 and 75% of endemic species respectively

(Medail & Quezel, 1997). With 5000 islands and

small islets, the Mediterranean Basin is one of the

biggest insular sets in the world. There are more

than 25000 superior plants according to Quezel

(1985) and about 30000 species and subspecies ac-

cording to Greater (1991).

This biodiversity is primarily due to the particu-

lar climatic conditions, habitat heterogeneity and

different origins of the flora, which can be divided

into three main biogeographic groups (Quezel

1985, 1995): a native species group, a southern af-

finity species group and a non-Mediterranean Ho-

larctic-Eurasiatic group.

In addition, geological, palaeogeographical and

historical factors have helped to create highly di-

verse environments and the insular mountain or iso-

lated edaphic systems generally appear to be major

endemic centres (Gomez-Campo et al., 1984).

Comparing the vascular flora of each of the above

mentioned Mediterranean biomes, the circum-Me-

diterranean one shows the highest taxonomic ri-

chness although the degree of endemism varies

from 50% (Quezel, 1985) to 59% (Greuter, 1991).

Relict species and insularity in the Mediterranean

Regarding young islands, the isolation, the poor

number of pioneer elements, the presence of unoc-

Relict species and the challenges for conservation: the emblematic case of Zelkova sicula

283

cupied ecological niches and a natural selection

lower than in the continent are major ways of spe-

ciation. More insular refuges are likened to arks

protecting their biota from extinction during hostile

events (Terborgh, 1992).

In the Mediterranean, small islets have served

as natural laboratories of plant evolution while large

islands have contributed to the conservation of mid-

dle Tertiary flora with a high degree of endemism

(Greater, 1995). In this context, the high rate of en-

demism results from the very complicated natural

(i.e. tectonic, geologic and climatic) history since

the middle Tertiary (Quezel, 1985; 1995). Due to

the moderate direct impact of the Quaternary gla-

ciations, especially the Wiirm (Debrandt-Passard,

1986), several zones have acted as refuges (Hewitt,

1999; 2000). Thus, more favourable themial condi-

tions and plenty of conservative microhabitats

(cliffs, nunataks, etc.) have helped to preserve nu-

merous elements from a Tertiary palaeoflora (Me-

dail & Verlaque, 1997).

Accordingly, we can distinguish two kinds of

endemics: palaeo-endemics, i.e. ancient vestiges of

taxa that were once widespread, and neo-endemics,

that have evolved only recently. The presence of

paleo-endemic trees is also remarkable, particularly

on larger islands, and includes fir species Abies ne-

brodensis and^f. cepholonica Loudon in Sicily and

Cephalonia respectively, Zelkova sicula in Sicily

and Z. abelicea (Lam.) Boiss. in Crete, Quercus al-

nifolia Poech and Cedrus brevifolia (Hook.f.)

A.Henry in Cyprus (Barbero et al., 1995).

Relict plants, especially trees, as having been

able to survive impressive environmental changes

over millions of years, are of great concern for their

valuable contribution in improving understanding

of past and recent biogeographical and evolutionary

processes (Kozlowski et al., 201 1). Moreover, in the

frame of forecasted climate changes, some elements

could have a role in the future establishment of

novel ecosystems with new species combinations,

thanks to their aptitude in niching strategy, as re-

ported for several relict taxa in the forests of Geor-

gia (Transcaucasia) (Denk et al., 2001). On the

other side, human-induced future climate scenarios

could reveal quite catastrophic for many of these

elements, acting as the “last nail in the coffin for

ancient plants” (Connor, 2009). Hence, any effort

should be made to improve the perspective of their

conservation.

Mediterranean plants and conservation issues

Conservation strategies represent a crucial issue

in the Mediterranean biome because this area,

which represents only 2% of the world’s surface,

houses 20% of the world’s total floristic richness

(Medail & Quezel, 1997). Many of these Mediter-

ranean insular species are currently threatened.

Despite the long history of botany in the Medi-

terranean, there is a lack of species distribution and

abundance maps which hinders effective conserva-

tion and management of local botanical heritage.

The assessment of the Mediterranean Islands Flora

(Delanoe et al., 1996) suggests that a significant

proportion of the islands flora is under threat.

Crete, followed by the Balearics, has the highest

percentage (11%) of plants endangered at a global

level. The Maltese flora has the highest percentage

(28%) of plants threatened at the local level reflec-

ting the pressure on the island’s habitat. The publi-

cation of the lUCN Top 50 Mediterranean Island

Plants (Montmollin & Strahm, 2005), albeit not an

exhaustive check-list, has the merit to draw the ge-

neral public’s attention to some of the most endan-

gered plant species, stressing particular situations

and conservation needs.

In the Mediterranean, the destruction of habitats

due to a long and ancient human presence led to

their extreme rarefaction and a drastic diminution of

their populations. The low altitude zones are most

affected, particularly coastal areas, rocky grasslands

and damp ecosystems, but the risks now extend

throughout all sectors due to the increase of human

activities. As mentioned in Olivier et al. (1995), en-

demics regression in Mediterranean area is primarily

due to the destruction of favourable habitats due to

urban development, roads, etc. and by direct human

impact (agricultural and forestry exploitations, wil-

dfires, trampling). The biotic threats (concurrence

of exotic species, competition with woody species),

together with biological weakness (e.g. low repro-

ductive efficiency, genetic erosion, ineffective di-

spersal) are traditionally considered less important,

although recent research on plant conservation ge-

netics acknowledges to the latter a much higher si-

gnificance (Kramer & Havens, 2009).

To face with these major threats, numerous pro-

tective actions must be taken immediately through

appropriate management of indigenous populations

and scientific studies undertaken to analyze popu-

284

Giuseppe Garf'i & Stephane Buord

lation viability. Among these native species, the re-

lict species such as Z sicula are priorities for con-

servation because they represent millions years of

evolution.

ZELKOVA SICULA: FROM FOSSILS TO A

LIVING PLANT

Until 1991 the genus Zelkova Spach was known

to Sicily only through fossil records (Beguinot,

1929) and nobody before that time would have su-

spected the existence of living trees in the island.

At present-day Z. sicula (Fig. 1) is a very rare spe-

cies exclusive from South-Eastern Sicily, belonging

to a genus which became extinct in the whole con-

tinental Europe during the Quaternary Glacial Age.

Up to very recent times the Sicilian species was

known to consist worldwide of only a single popu-

lation (hereinafter referred as ZSl) of about 250

trees, included in a small area less than 0.4 hectares

within the Bosco Pisano, in the Iblei Mts. At the end

of 2009 a second population (hereinafter ZS2) was

Figure 1. Branchlet of Zelkova sicula.

unexpectedly discovered in the same mountainous

massif (Fig. 2) (Garfi et al., 2011). As the former

one, it includes a few hundreds stems, once again

distributed over a limited area of about 0.5 hectares,

in quite similar environmental conditions. In the

conservation perspective, such extraordinary occur-

rence surely contributed to reduce the level of threat

for this species, but at the moment it was not

enough to allow a reduction of risk level adopting

lUCN categories.

Included among the family of Ulmaceae, the

genus Zelkova is currently represented by six spe-

cies, whose worldwide distribution is quite frag-

mented: three taxa, Z. serrata (Thunb.) Makino, Z.

schneideriana Handel-Mazzetti and Z. sinica C. K.

Schneid., are widely spread in the lush forests of

Eastern Asia, and one, Z. carpinifolia (Pall.) Dippel,

is common in Transcaucasia, whereas the last two,

Z. sicula and Z. abelicea, are narrow endemics to

the Mediterranean islands of Sicily and Crete, re-

spectively (Denk & Grimm, 2005).

In the ecological point of view, some differences

may be outlined among all taxa. Z. sicula is the only

species relatively able to face Mediterranean sum-

mer drought stress. It grows in the thermo-(-meso)

Mediterranean bioclimate, within sclerophyllous

sparse communities dominated by Quercus suber

L., Olea europaea var. sylvestris (Miller) Brot and

Q. virgiliana (Ten.) Ten.; its biogeographically clo-

sest relative Z. abelicea lives in areas with fresh mi-

croclimate and good water balance (N-facing

slopes, small valleys, dolines), at elevations bet-

ween 850 and 1,800 m a. s.L, within supra- and oro-

Mediterranean mixed discontinuous woody stands

with Acer sempervirens L., Quercus coccifera L.

and occasionally Cupressus sempervirens L. (Bar-

bero & Quezel, 1980; Egli, 1997). Finally, the four

Asian species mostly thrive under a humid and

warm-temperate climate, where rainfall usually ex-

ceeds 1000 mm/yr and can rise up to 2000-2500

mm/yr (cf. Garfi et al., 2011).

Additional aspects distinguish the Sicilian spe-

cies. Both populations are currently restricted to

gullies bottom and streamsides. Furthermore, in

contrast to its relatives, usually represented by me-

dium to tall trees, Z. sicula in population ZSl cur-

rently e xhi bits an explicit shrubby growth form

(maximum height 2.5 m), with many plants stunted

and a general poor conservation; in population

ZS2, which can enjoy of a bit more favorable water

Relict species and the challenges for conservation: the emblematic case of Zelkova sicula

285

Figure 2. Current distribution map of Zelkova sicula (see text for abbreviations).

supply, the overall eonditions look mueh better in

terms of health and vigour, either at tree as well as

at population level, with many plants that ean attain

the height of 5-6 m (Fig. 3) (Garfi et al., 2011). In-

deed, a few individuals eultivated ex-situ in less

eonstraining environmental eonditions (no water

stress, milder temperature, partial shadowing) re-

vealed a growth potential of veritable trees. In ae-

eordanee with that it is suggested that in the past the

speeies was probably more widespread in fresher

and more humid environments.

Sueh an assumption is quite eonsistent with the

data issuing from palaeobotanieal investigations,

showing Zelkova remains within floristie assem-

blages dominated by deeiduous broadleaved trees,

from several European loealities up to the Late

Pleistoeene (e.g. Follieri et al., 1986; Garfi, 1996;

De Paola et al., 1997). Aetually, the genus Zelkova

belongs to a hygro-mesothermie floristie unit

whieh was very eommon in the luxuriant Tertiary

forests throughout the Northern Hemisphere. In

Eurasia, elimate ehanges oeeurred sinee the late

Plioeene/early Pleistoeene with the onset of the lee

Age, eaused the progressive rarefaetion from nor-

thern to southern latitudes of the less eold-resistant

elements. The Italian peninsula, among others, pla-

yed in this regard a major role of refuge allowing

the persistenee at Valle di Castiglione (Central

Italy) of the last Zelkova remnants until 31000

years B.P. After that time, aeeording to Follieri et

al. (1986), the genus Zelkova beeame definitively

extinet from the whole eontinental Europe. Only in

the two Mediterranean Islands of Crete and Sieily,

where glaeiations effeets were less severe, the two

endemies Z. abelicea and Z. sicula survived until

the present-day (Quezel et al., 1993), tolerating

more or less effieiently the summer drought stress

of the eurrent Mediterranean elimate.

Aeeording to reeent investigations (Finesehi et

al., 2002; Christe et al., in press), Z. sicula is gene-

tieally depauperated and seems to have had a hi-

story of severe isolation. It is suspeeted to be a

speeies of hybrid origin and its parents are sugge-

sted to be elose to Z. abelicea and Z. carpinifolia

aneestral speeies. This is rather eonsistent with re-

286

Giuseppe Garf'i & Stephane Buord

suits of Nakagawa et al. (1998), which point out

that leaf fossils, formerly referred by palaeobota-

nists to as Z. carpinifolia, most likely might instead

be attributed to Z. sicula. Whatever the ease, it

seems quite alike that sueh harsh and eomplex vi-

eissitudes eould be invoked as responsible of mu-

tations resulting in the eurrent triploid earyotype of

Z. sicula, whieh differs in this regard, too, from all

its diploid relatives (Garfi, 1997a). This is very rare

within the angiosperms and further emphasizes the

great eoneem of this speeies.

In the whole, the aforementioned issues finally

highlight the extreme importanee of this taxon for

both studies of eonservation biology and the reeon-

struetion of historieal biogeography of the genus

Zelkova (Kozlowski & Gatzfeld, in press).

WHY IS Z. SICULA SO VULNERABLE?

CURRENT CONSERVATION AND THREATS

At present a variety of eauses threaten a satisfae-

tory eonservation or, what is more, the survival it-

self of Z. sicula. Some faetors are strietly depending

on its biology, others on different exogenous agents.

A general review ean be summarized as follows.

Unsuccessful sexual regeneration

The fruetifieation of Z. sicula is quite irregular

(Garfi, 1997a), as already known for its Cretan re-

lative Z. abelicea (Egli, 1997). In the Sieilian spe-

eies it seems usually eoineiding with the rainiest

winter years and has been most often observed on

the same, very few trees. Moreover, aeeording to

eurrent knowledge (Garfi, 1997b) seeds seem ste-

rile due to its triploid eonditions. Therefore, at pre-

sent-day regeneration is exelusively depending on

vegetative meehanisms sueh as root suekering and

layering (Fig. 4).

Consequently, in both populations most trees are

very probably of elonal origin. This situation, in ad-

dition to the long geographie isolation, most likely

involved severe deeline in gene flow and a rapid de-

erease of intra-speeifie genetie variability (Finesehi

et al., 2002; 2004; Christe et al., in press). The su-

speeted predominanee of elonality eould have eon-

trasting implieations: i) on the one side it ean

represent the meehanism that assured the perpetua-

tion of a sueeessful genotype differentiated in re-

sponse to the hardiness of a ehanging environment

throughout the geologie times, and then the survival

of the speeies until present-day; ii) on the other side,

sinee elonal reeruitment implies absenee of gene re-

eombination and then strong genetie impoveri-

shment, it hugely diminishes populations’ viability,

raising vulnerability to any external adversity. In the

same time, elonality adds a further element of ex-

eeptionality to sueh an emblematie speeies. If future

investigations would reveal that both two eurrent

disjunet stands are entirely elonal populations, this

eould mean that, as for Lomatia tasmanica (Fineh

et al., 1998), eaeh of them is a very old organism,

even aged many thousand years, therefore beeo-

ming among the oldest living plant individuals (!)

known to date.

In any ease, the laek of fiinetional seeds strikin-

gly enhanees the risk of extinetion of the speeies as

it surely involves major diffieulties in propagating

it, both for in-situ and ex-situ eonservation aetions.

Designing efifieient protoeols for in-vitro and in-vivo

vegetative multiplieation is to be eonsidered an ir-

replaeeable goal for any efforts of aetive and effi-

eient eonservation. In this regard, propedeutie

investigations about the residual genetie diversity

need to be earried out in order to eonserve as mueh

as possible intra-speeifie variability.

Anthropic disturbance and habitat degradation

Aeeording to eurrent knowledge (Garfi et al.,

2011), Z. sicula is supposed to be a speeies thriving

in typieal forest habitat. Heavy human pressure

(grazing, wildfires, past silvieultural over-exploi-

tation), in addition to the foreseen elimate deterio-

ration are responsible of its eurrent habitat

degradation, as involving failure in natural regene-

ration of forest speeies and the further impoveri-

shment in forest eomposition and strueture.

Wildfires represent a major hazard in the whole

forest area. In the past large patehes of forest eover

have been destroyed or severely damaged by wil-

dfires, whereas very reeently fire injuries have

even direetly eoneemed the population ZS2. Their

origin is usually to be related to human aetivities.

As in many Mediterranean areas, sinee aneient

times fire has represented a kind of very “primi-

tive”, but extremely deleterious method of land ma-

nagement and in the last years it has beeome also a

means of soeial elaim. The general eonsequenees

eould be the simplifieation of the eeosystem and

Relict species and the challenges for conservation: the emblematic case of Zelkova sicula

287

Figure 3. Population of Zelkova sicula from site Contrada Ciranna (ZS2) with trees 5-6 m high. Figure 4. Vegetative pro-

pagation through root suekers. Figure 5. Withered leaves following summer water stress in August 2011. Figure 6. Massive

proliferation of epieormic short shoots due to browsing (a) and detail of a deeapitated tip (b). Figure 7. Selerophillous speeies

diffusion at site Boseo Pisano (ZSl) after grazing exelusion. Figure 8. Fencing and irrigation plant built at site Bosco Pisano

(ZSl) within actions of the LIFE project Zelkov@zione.

288

Giuseppe Garf'i & Stephane Buord

landscape mosaic, with a decrease of ecosystem

functions. The habitats suffering the greatest risk

are those directly related to Z sicula (i.e. 9330 -

Quercus suber forests, and 6220 - Pseudo-steppe

with grasses and annuals of the Thero-Brachypo-

dietea), but the other habitats could be also concer-

ned at various extent. Reiterated events can involve

the definitive loss of most vulnerable species and

the selection of fire-resistant taxa. Notwithstanding

its great ability of resprouting after an injury, Z. si-

cula for its rarity and punctiform distribution could

easily disappear as a consequence of repeated wil-

dfires at short intervals.

As far as concerns grazing, historical data te-

stify that it has been one of the main types of local

land use. Its pressure became heavier since the last

sixty-years period, after land abandonment and the

migratory flows abroad of local population that

made possible the settlement of transhumant gra-

ziers originating from the mountainous area of

Northern Sicily (Garfi & Di Pasquale, 1988; Di Pa-

squale & Garfi, 1989).

In contrast to fire, in some particular cases (e.g.

for the habitat 6220 - Pseudo-steppe with grasses

and annuals of the Thero-Brachypodietea) grazing

is considered a promoting factor for plant richness.

However, it must be kept in mind that in the area of

interest it has been in the time a formidable cause

of forest ecosystem degradation. Such an issue must

be carefully evaluated in the conservation planning

perspective, especially in a region like Sicily where

forest cover has been destroyed on large areas since

the antiquity. The great worry is that in the next fu-

ture the unregulated way this practice is currently

carried out can notably enhance the destruction of

the last remnants of “natural” forest patches of this

part of Sicily. In the specific case of Z. sicula, brow-

sing is proved responsible of severe growth sup-

pression (Fig. 5), flowering inhibition and even

death of trees, already hardly weakened by a con-

straining environment (Garfi, 1997b).

Summer water stress

Due to its incomplete adaptation to the Medi-

terranean climate seasonality, since its discovering

Z. sicula has suffered periodical summer drought

stress (Fig. 6), and several evidences indicate simi-

lar occurrences also in past times (Garfi et al.,

2002). Water stress cause moderate to severe injury

to the peripheral parts of the crown. Premature se-

nescence and shedding of leaves have been most

frequently noticed, whereas in case of abrupt dehy-

dration leaves withered and died but remained atta-

ched. Sometimes, severe increasing water stress

also leads to the death of twigs and branchlets, or

even of the entire stem. The plants can recovery in

the following season, but repeated episodes of stress

along more than one single year involve the death

of trees (Garfi et al., 2002). For instance, in summer

2007 the entire population ZS 1 suffered very severe

water stress and demographic follow-up enabled to

assess that almost 20% of trees have died.

Furthermore, recent trends in climate changes at

global scale depict scenarios characterised by an in-

crease in summer drought, both in terms of less

rainfall amount and duration and raise of tempera-

ture. Major detrimental effects can then be expected

for the conservation of the species in the nature

(Allen et al., 2010; Borghetti et al., 2012).

Potential interspecific competition

Some years after its discovery, the population

ZSl was fenced in order to protect it against brow-

sing. During the years following the exclusion of

grazing disturbance, Z. sicula experienced a rather

remarkable growth increase. But in the same time

progressive succession processes were observed

in its habitat: vegetation communities’ patterns

began to change and woody species cover became

more and more relevant. Xerophilous taxa, such

as Calicotome infesta (C. Presl) Guss., Pyrus spi-

nosa Forssk., Phillyrea latifolia L., Sarcopoterium

spinosum (L.) Spach and some trees like Quercus

suber, Q. virgiliana and Celtis australis L. began

to spread (Fig. 7).

On the one side such positive dynamic trend is

expected to improve in the long period the global

habitat stability and ecosystem functions. On the

other side, such processes are believed to be able to

trigger in the short period inter-specific competition

at the expense of Z. sicula, which surely cannot ex-

press a comparable adaptive and growth ability of

typical Mediterranean xerophilic species.

This could entail problems of decrease or even

survival of the species of concern, so that a perio-

dical monitoring of both populations should be re-

quired in order to assess the demographic trends

and population resilience and get information for

conservation management.

Relict species and the challenges for conservation: the emblematic case of Zelkova sicula

289

Improper use of plants or parts of them

Considering the peeuliarities of Z. sicula (e.g.

its aneient origin, its troubled history throughout the

geologieal periods, its rarity) sinee its diseovery a

speeial interest has arisen about it from a rather va-

riegated multitude of people, such as scientists, en-

viromuentalists, common tourists, plants collectors,

bonsai producers, nurserymen, etc.. Small to large

groups of people, usually not controlled by any of-

ficial guide or surveillance, during their visits scam-

per everywhere in the area causing soil subsidence,

overall injuries to vegetation and even involuntary

trampling of small Zelkova plants. Moreover, the

last four categories of visitors are the most dange-

rous since they often gather parts of plants as sou-

venir and, especially the bonsai lovers, even uproot

young plantlets for their collections.

Although not a priority cause, like for Z. abeli-

cea from Crete (Egli, 1997), this kind of distur-

bance/misuse could reveal detrimental for

conservation since responsible for reduction of po-

pulations’ size and, given its rarity, significantly in-

crease the risk of extinction.

AN INTEGRATED STRATEGY FOR THE

CONSERVATION OF Z. SICULA

As previously outlined the entire genus Zelkova

is a taxon of special concern. Accordingly, in recent

years it has deserved a particular interest within the

scientific community and conservationists (BGCI,

2010; Kozlowski et al., 2011). This resulted in a

global conservation plan, set out into three main

complementary goals (Kozlowski & Gratzfeld, in

press): i) conservation, comprising elaboration of

an action plan, proposing concrete recovery and/or

reintroduction measures, etc.; ii) basic and applied

research, including molecular phylogeny, phylogeo-

graphy, population genetics, population structure,

genetic comparison of wild populations with ex-situ

collections, etc.; iii) public awareness and outreach,

involving development of travelling exhibitions, and

organisation of national and international conferen-

ces and seminars to exchange knowledge and share

conservation expertise. In parallel, a specific project

entirely addressed to the conservation of the Sicilian

species was presented by a composite partnership

(i.e. the Sicilian Department of Environment - DRA,

the Sicilian Regional Authority of Public Forests -

AFDRS, the Institute of Plant Genetics of the Italian

National Research Council - IGV-CNR, the National

Botanic Conservatory of Brest - France - CBNB,

and Fegambiente, an Italian environmental associa-

tion) within the frame of the FIFE+ EC programme,

specially conceived for the conservation of nature

and biodiversity in the EU territory.

The project, named “Zelkov@zione - Urgent ac-

tions to rescue Zelkova sicula from extinction”

(http://www.zelkovazione.eu/) was funded in 2011

and, given its evident relevance with the internatio-

nal initiative, a sharing relationship was soon esta-

blished with it. The project aims to ensure the

survival of Z. sicula through in-situ and ex-situ con-

servation integrated actions to be carried out along

almost five years. It entails four comprehensive to-

pics all related each other (Table 1): i) knowledge

and monitoring, ii) active conservation, iii) exper-

tise and communication, and iv) education/aware-

ness. Some short remarks can help catching the

fundamentals of the project strategy.

Collecting information for conservation planning

Albeit a conspicuous literature already exists

(cf. reference list), much still need to be understood

both in terms of basic knowledge and issues to ad-

dress conservation. The viability of Z. sicula is a

preliminary information required to correctly plan

conservation policy. In this regard the first step to

assess the current status of the target species invol-

ves the inventory and mapping of all discrete trees

(i.e. single/multiple trees or shoots apparently ori-

ginating from spatially distinct stumps), in addition

to the implementation of a georeferenced database

including biometric data (stem height and diame-

ter), pheno logical features (e.g. flowering trees,

fruiting), individual vigour, past damages (water

stress, biotic disturbances), microsite characteristics

and spatial distribution patterns. Periodical monito-

ring will allow appraising the demographic trends

of the target species and the forest stand dynamics

with the aim to prevent inter-specific competition

phenomena following disturbance suppression.

Genetic investigations are propaedeutic to any

actions of multiplication, in order to detect residual

genetic variability, if existing, and allow conserva-

tion of the most diverse genotypes. Vegetative pro-

pagation through in-vivo and in-vitro techniques is

an immediate successive step.

290

Giuseppe Garf'i & Stephane Buord

Type of actions

Involved activities

Expected results

Knowledge/

Monitoring

1. Updated inventory of both ZSl and ZS2 popula-

tions, ineluding the position of eaeh tree trough sub-

metrie GPS deviee, the implementation of a

georefereneed database with biometrie, biologieal

and miero-topographie attributes

2. Monitoring of both the target speeies demographie

trends and the forest eommunity dynamies in eurrent

and future populations

S.Genetie investigations

4. Definition of an effieient in-vitro and in-vivo ve-

getative propagation protoeol for the target speeies

•Improvement of knowledge about the biology and the

eeology of the target speeies

•Evaluate the viability of Zelkova populations and pre-

vent any inter-speeifie eompetition phenomena follo-

wing disturbanee elimination

•Evaluation of the residual genetie diversity in order to

preserve the as highest as possible rate of variability

•Contribute to a more effieient eonservation planning

Active

conservation

1. Building an effieient system of feneing

2. Setting up of an emergeney irrigation system

3. Massive produetion of multiplieation material of

Z. sicula and native forest speeies

4.1n-situ plantation of the target speeies and establi-

shment of 5 new populations, 2 within the eurrent

habitat and 3 in supposed more favourable bioelima-

tie eonditions

5. Plantation of native forest speeies on about a 10 ha

area ineluding the eurrent site of ZS 1

6. Ex-situ eultivation of at least 200 trees from both

ZSl and ZS2, at the CCG and CBNB

7. Prompting formal proeedures to release a regional

administrative Aet for the legal proteetion of Z si-

cula and the SCI Boseo Pisano

8. Starting of formal proeedures for the inelusion of

Z sicula in the list of priority speeies of the Habitat

Direetive

•Improving the proteetion in the nature against disturbanee

from grazing or improper eolleeting of plant material

•Attenuation of summer water stress episodes for the po-

pulation ZS 1

•Reinforeement of the eurrent populations of Z sicula

•Inerease the number of population in the nature and test

the feasibility of uneommon eonservation approaeh as

“assisted eolonization”

•Habitat enforeement/rehabilitation in order to ereate a

more suitable eeologieal environment, through reeove-

ring the as highest as possible eeosystem funetionality

under eontrolled situation and in reasonably fast times

•Seeuring through ex-situ eonservation the as highest as

possible genetie diversity in publie eonservation eentres

•Reeognition of the status of proteeted speeies at re-

gional/national level and starting of the proeedure for

its aekowledgement as priority speeies (sensu Habitat

Direetive)

•Implementation of a standard normative proeedure for

proteetion of all threatened speeies in publie/private lands

Expertise/

Communication

1 .Implementation of operative conservation plan-

ning according to the rules proposed in the Ma-

nagement Plan “Monti Iblei”

Z.Drawing of a grazing management plan for the

Boseo Pisano to attenuate the pressure on the con-

cerned habitat according to the rules proposed in

the Management Plan “Monti Iblei”

3. Planning of special operative procedures for

wildfires prevention

4. Activation of training activities for local people

(e.g courses for naturalistic guides) to improve

knowledge and introduce young people to the la-

bour market

•Applying site-dedicated concrete conservation mea-

sures according to the Management Plan “Monti

Iblei”

•Involving the competent local Forest Authority to

improve surveillance and protection of the species

and habitat through active policy

•Realisation of a communication network among pu-

blic bodies, stakeholders, schools, etc., in order to

share and monitor experience and results

Education/

Awareness

1. Establishing a permanent round-table among

partnership, public authorities, no-profit organi-

sations and local stakeholders to monitor the pro-

cedures of LIFE+ project.

2. Carrying out of campaigns of awareness and di-

vulgation on the specific project and the general

problem of conservation of biodiversity

3. Realisation of a dedicated website to inform and

divulgate objectives, procedures, monitoring,

feed-backs and results

•Removing/reducing principal economic/social type

threats for the species and habitat by involving au-

thorities, organizations and stakeholders

•Involving local people in efforts to improve local

economy on green tourism

•Improving the didactic value of the CCG, with be-

neficial feed-back on the territory in regards to public

awareness about the global problem of biodiversity

conservation

•Prompting local awareness and information about

the problems of loss of biodiversity

Table 1. Main activities foreseen in the LIFE+ project “Zelkov@zione”.

Relict species and the challenges for conservation: the emblematic case of Zelkova sicula

291

The active actions: habitat rehabilitation, as-

sisted colonisation and legal protection

Active conservation mainly includes a number

of concrete measures directly in the field, in addition

to traditional ex-situ conservation. Outright actions

as fencing against grazing/misuse disturbances and

the setting up of an emergency irrigation system

(Fig. 8), can contribute to eliminate or significantly

reduce some of the most flagrant threats.

More indirect interventions will concern the pre-

sent habitat. As formerly mentioned, the optimal ha-

bitat of Z. sicula is assumed to be typical forest,

with complex floristic composition and structure.

In contrast, the current habitat is heavily degraded

due to long-lasting human pressure. Therefore the

recovery of ecosystem functions involving actions

of reinforcement and rehabilitation needs to be pur-

sued in order to (re-)create a more suitable ecologi-

cal environment for the target species. The action

will be exclusively based on the employ of native

forest species, produced from multiplication mate-

rial collected in the same forest area. Moreover, it

will be performed according to the modem princi-

ples of naturalistic silviculture and water saving

(minimize the impact of preparatory works and con-

serve as soon as possible the existing vegetation

cover, employ native shmb and tree species, use of

hydrogels, i.e. polyacrylates of very high molecular

weights binding water up to 400 times their mass,

that can prolong the survival of trees under water

stress up to 300%).

Particular attention will be also paid to the plan-

ting patterns, by favouring the creation of hedge-

rows and the random distribution of various size

plant groups in the aims to obtain a variety of habi-

tats and fauna feeding sources. Restoration will also

involve the removal of exotic tree groups {Eucalyp-

tus spp. and Cupressus spp.) introduced near ZSl

population during past afforestations.

At the population level, new plantings in the

current sites are aimed to enlarge and reinforce the

current stands. But one of the greatest efforts of the

project consists in the introduction and establi-

shment of novel populations in new sites, three of

which selected in areas supposed to be more suita-

ble for the species. In fact, according to results of

recent investigations (Garfi et al., 2002; 2011), pa-

laeoecological data (e.g. Beguinot A., 1929; Follieri

et al., 1986; De Paola et al., 1997) and personal ob-

servations on cultivated trees, a more humid envi-

ronment (e.g. supra-Mediterranean or montane fo-

rest habitats dominated by Fagus, Acer, Carpinus,

Taxus, deciduous-type Quercus etc.) (Fig. 9) is in-

ferred to better match with the ecological require-

ments of Z. sicula.

A similar approach, in which an endangered

species is introduced outside of its historically

known native range, is quite uncommon in actions

of plant rescue. Actually just in the last few years

a lively debate has arisen on this subjects (for a re-

view see Brooker et al., 2011), especially fostered

by the increasing recognition of the likely inability

of many species to cope with rapid climate war-

ming (Thomas, 2011). Many terms have been pro-

posed to indicate this approach of moving species

at risk from their current locations to those areas

expected to be suitable for their growth under fu-

ture climate change scenarios, but the most com-

mon used is “assisted colonisation” (Hunter, 2007;

Brooker et al., 2011).

Although some conservationists (Ricciardi &

Simberloff, 2008) remain very critic in this regard,

assisted colonisation is invoked as the “last resort”

when other conservation strategies have been pro-

ved to be ineffective or are highly prone to fail.

This is specially true for narrow endemics confi-

ned to very specialised habitat (isolated mountain,

single lake, unique geo-pedologic substratum) that

are surrounded by environments fundamentally

unsuitable for them which became insurmountable

barriers (Thomas 2011; Brooker et al., 2001). Ge-

netic patterns are as much as detrimental, since ge-

netic erosion can limit short-term resilience,

evolutionary potential for adaptation, and long-

term survival of plant species in the face of rapid

environmental change (Kramer & Havens, 2009).

It is extremely paradigmatic the example of Tor-

reya taxifolia Am., from Florida, that since the fif-

ties of the last century suffered an inexplicable

dieback up to the present number of about 500

trees (Barlow & Martin, 2004), therefore having

been the object of intensive planting outside its

original range.

In the case of Z. sicula we can account several

reasons to refer to assisted colonisation, especially

when related to future climate model of rapid war-

ming (Allen et al., 2010; Borghetti et al., 2012). Fir-

stly, we must consider its propagation ability, that

even at the genus level is not quite performing. For

292

Giuseppe Garf'i & Stephane Buord

Figure 9. View of the Boseo Tassita (Nebrodi Mts.) as an example of possible (re-)introduetion area oiZelkova sicula

aeeording to assisted eolonisation

instance, it has been suggested that the extant di-

stribution of Z. carpinifolia is probably also due to

the rather inefifieient dispersal meehanisms that may

have hindered its expansion from Pleistoeene refu-

gia to Holoeene alluvial plains in the Colehie lo-

wlands (Denk et ah, 2001). The situation of the

Sieilian speeies is even more dramatie, given its

ineffeetiveness to regenerate through seeds that

makes its dispersal impossible over long distanees.

In addition, its eurrent area is extremely isolated

and fragmented, being separated from supposed

more suitable habitats (e.g. the Nebrodi Mountain

Range, Northern Sieily) by vast lowlands. Moreo-

ver, as mentioned by some papers (Jaekson &

Hobbs, 2009; Brooker et al., 2011), palaeoeeologi-

eal data ean provide important information on past

oeeurrenee of the speeies and/or the plant eommu-

nity it took part to, then eneouraging our transloea-

tion projeets. This should be rather innovative for

the Mediterranean area, and in ease of sueeess, sueh

an aetion would eonfirm that our assumptions are

sound and show that an apparently ‘unnatural’ in-

tervention might even deeisively eontribute to save

a speeies from extinetion.

Traditional living ex-situ eolleetion is also eon-

templated. At least 200 trees from both popula-

tions will be eultivated at the CCG and the CBNB,

in order to seeure the speeies and eonserve the as

highest as possible genetie diversity, aeeording to

the reeommendations of Kozlowski et al. (2011).

The aetuation of legal proteetion is the last but

not the least issue of eonerete aetions. At present in-

deed Z. sicula does not yet enjoy for any legal/for-

mal measure of safeguard. Therefore different

aetions are addressed to both the sites and the spe-

eies. Formal proeedures are prompted for the reeo-

gnition of SCI ITA090022 “Boseo Pisano” (site of

ZSl) as a Speeial Area of Conservation (SAC)

sensu Habitat Direetive 92/43, and to enlarge the

perimeter of SCI ITA090024 “Cozzo Ogliastri” in

order to inelude the site ZS2, situated in its elose

proximity. In parallel, as the target speeies is not

Relict species and the challenges for conservation: the emblematic case of Zelkova sicula

293

comprised either in the Annex II or IV of the Habi-

tat Direetive, the Italian Ministry of Environment

will be requested through the DRA to inelude it in

the list of “priority” speeies (in danger of disappea-

ring, needing partieularly striet proteetion).

At regional/national level the main goal is to

obtain also the aeknowledgement of the status of

proteeted speeies through the adoption by the DRA

of a normative doeument (Couneilor’s Deeree)

eontaining appropriate measures that will ensure

the legal proteetion of the target speeies and its ha-

bitat. This will represent the first step to serve for

the implementation of a standard normative proee-

dure for proteetion of all threatened speeies in pu-

blie/private areas.

Capitalizing knowledge and expertise and

developing public awareness

Knowledge and expertise as well as aetions in

the field eannot disregard the need of divulgation

of the results in order to inerease publie awareness.

The implieated and ongoing expertise will eneom-

pass as key outeome the implementation of an ope-

rative eonservation planning for the areas of

eoneem aeeording to the rules proposed in the Ma-

nagement Plan “Monti Iblei” (ARTA, 2009), al-

ready eompiled within the Regional Management

Plans of SCI and ZSP. This will direetly lead to the

drawing of a site-dedieated management plan for

grazing and the wildfires prevention, aiming to at-

tenuate the pressure on the habitat and reduee the

major anthropogenie threats. Aeeordingly, the eom-

petent loeal Forest Authority will be involved in im-

proving surveillanee and proteetion of the speeies

and habitat through an aetive poliey.

The establishment of a permanent eommuniea-

tion network among eonservationists, publie bodies,

stakeholders, proteeted areas managers, ete. is a

eomplementary goal in order to share and monitor

experienee and results. Additionally, the aetuation

of training aetivities are foreseen for loeal people

(e.g. eourses for naturalistie guides) to improve

knowledge and stimulate the dissemination of an

environmental eonseiousness.

Dedieated eampaign of awareness and divulga-

tion on both the speeifie projeet issues and the ge-

neral problem of biodiversity eonservation will be

promoted through media, edueational programmes

in primary and seeondary sehools at loeal and re-

gional level, organisation of periodie meetings and

workshop on the aims and advaneement of the pro-

jeet in publie venues. The events will allow to eon-

tinuously evaluate the “state of the art” and to

illustrate the final results.

The organization of a generalist network will aim

at sharing the projeet aetions among publie authori-

ties, no-profit institutions and loeal stakeholders in-

volved in the eonservation issues of eritieally

endangered speeies. Finally, the realisation of a pro-

jeet-dedieated website will serve to inform and dis-

seminate as mueh as possible objeetives, proeedures,

monitoring, feedbaeks and results of the projeet.

CONCLUSIONS

In the frame of the foreseen rapid elimate ehan-

ges at world level, and in the Mediterranean in par-

tieular, many plant speeies unable to eope with

unsuitable environment are doomed to beeome ex-

tinet. Insular endemies and reliet taxa are the most

threatened, espeeially those suffering for habitat de-

struetion and biologieal weakness (e.g. low repro-

duetive effieieney, genetie erosion, ineffeetive

dispersal). In this regard Z. sicula represents a quite

paradigmatie eonservation ease, whieh must faee to

very ehallenging tasks.

As a general rule, priorities for biodiversity eon-

servation management should be of two kinds: i)

priorities for extensive areas where high biologieal

diversity has aeeumulated over long periods of geo-

logieal time and ii) priorities for distinetive areas

with high endemism. Beeause of the small extent

and high eoneentration of unique taxa, the risk of

rapid and irreversible loss of biodiversity is high

and important eeosystem flinetions may also rapidly

be lost. Biogeography has an important role to play

in this proeess of eonservation, but mueh more kno-

wledge is required about the habitat preferenees of

speeies and their response to habitat fragmentation

resulting from inereasing development pressures

and elimate ehange.

Fandseape eeology will play an inereasingly im-

portant role, providing the spatial eontext within

whieh to seleet eeologieally important sites for pro-

teetion, for monitoring ehange and for identifying

sites for habitat restoration and reliet speeies (re-)in-

troduetion. Among all, one of the most intriguing

and ehallenging strategy surely must be referred to

294

Giuseppe Garf'i & Stephane Buord

the translocation of species in presumed more sui-

table habitats, becoming the “last resort” to face the

deleterious impact of forecasted climate deteriora-

tion. Pilot (re-)introductions attempts worth to be

carried out and results could notably contribute to

the present debate concerning the application of as-

sisted colonization, especially for relict species, as

a valuable conservation tool for dealing with the

threats of a changing world.

ACKNOWLEDGMENTS

Authors are grateful to all colleagues of the par-

tnership of project Zelkov@azione, with special re-

gards to Matilde Fiore and Giandomenico

Maniscalco (from DRA), Francesco Carimi (from

CNR-IGV), Giancarlo Perrotta (from DRAFD), Sal-

vatore Livreri Console and Nicola Corona (from Le-

gambiente), Dominique Dherve and Catherine

Gautier (from CBNB), and the respective administra-

tive and technical staff for their invaluable contribu-

tion during all steps of the project implementation.

The project “Zelkov@zione - Urgent actions to re-

scue Zelkova sicula from extinction’Vas co-funded

by the EC LIFE+ Programme, Grant Agreement n.

LIFE 10 NAT/IT/000237.

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Biodiversity Journal, 2012, 3 (4): 297-310

Impact of alien insect pests on Sardinian landscape and

culture

Roberto A. Pantaleoni'’^*, Carlo Cesaroni',C. Simone Cossu', Salvatore Deliperi^ Leonarda Fadda', Xenia

Fois', Andrea Lentini^ Achille Loi^ Laura Loru', Alessandro Molinu', M. Tiziana Nuvoli^ Wilson Ramassini^

Antonio Sassu', Giuseppe Serra', Marcello Verdinelli'

'Istituto per lo Studio degli Ecosistemi, Consiglio Nazionale delle Ricerche (ISE-CNR), traversa la Crucca 3, Regione Baldinca,

07100 Ei Punti SS, Italy; e-mail: r.pantaleoni@ise.enr.it

^Sezione di Patologia Vegetale ed Entomologia, Dipartimento di Agraria, Universita degli Studi di Sassari, via Enrieo De Nieola,

07100 Sassari SS, Italy; e-mail: pantaleo@uniss.it

* Corresponding author

ABSTRACT Geologically Sardinia is a raft which, for just under thirty million years, has been crossing

the western Mediterranean, swaying like a pendulum from the Iberian to the Italian Peninsula.

An island so large and distant from the other lands, except for its “sister” Corsica, has inevitably

developed an autochthonous flora and fauna over such a long period of time. Organisms from

other Mediterranean regions have added to this original contingent. These new arrivals were

not randomly distributed over time but grouped into at least three great waves. The oldest two

correspond with the Messinian salinity crisis about 7 million years ago and with the ice age,

when, in both periods, Sardinia was linked to or near other lands due to a fall in sea level. The

third, still in progress, is linked to human activity. Man has travelled since ancient times and

for many centuries introduced allochthonous species to Sardinia which radically modified the

native flora and fauna, but always at a very slow and almost unnoticeable rate.

The use of sailing or rowing boats, with their low speeds, hindered the transport of living

organisms from one place to another. The use of the steam boat, introduced around 1 840 but

widely diffuse around 1870-1880, opened the doors to more frequent arrivals and also to orga-

nisms from the American Continent. This technical innovation had an influence over the whole

world economy, with its well-known grain crisis, and coincided in Sardinia with the arrival of

Roman dairymen, producers of pecorino cheese and the beginning of the expansion of sheep

farming which would continue uninterrupted until the present day. In this period of sudden so-

cial and environmental change, an insect was introduced which would turn out to be probably

the most economically devastating agricultural pest in Europe: the Grape Phylloxera. The vi-

neyard and wine business collapsed first in France then in Italy. The Phylloxera arrived in Sar-

dinia in 1883 and wine production crashed a very short time later and only resumed after the

distribution of American vine rootstock at the beginning of the 20th Century. From then, vine

cultivation in Europe was modified with the essential use of this rootstock.

Since then methods of transport have increased enormously in number and speed. The num-

ber of allochthonous and invasive species has increased proportionally: some of them along

with exotic plants which are cultivated on the island, others following man in his activities.

Often these new pests attack and destroy ornamental plants which have become part of the

Sardinian landscape, causing it to change; just as often their presence requires methods of pest

management which are different from the traditional methods on specific crops; finally in at

least one case (the Asian tiger mosquito) they pose a threat to our health.

KEY WORDS invasions; dispersion by man; Mediterranean; allochthonous species; history.

Received 12.05.2012; accepted 10.09.2012; printed 30.12.2012

Proceedings of the F‘ International Congress “Insularity and Biodiversity”, May 11*-13*, 2012 - Palermo (Italy)

298

R. A. Pantaleoni et alii

INTRODUCTION

Sardinia is in the centre of the Western Mediter-

ranean, one of the most bio-geographically complex

and interesting areas of the Palearctic region. Its na-

tural history is closely linked today with the history

of man. Before he arrived, the rhythms of evolution

and change in fauna and landscape were synchro-

nised with geological rhythms.

Long periods of standstill were interrupted by

some “catastrophic” moments of radical change

(Baccetti, 1964; 1983). After the arrival of man,

who represents the last of these natural catastro-

phes, the speed of change was completely modified.

The rate of arrival of new species grew exponen-

tially and has probably reached its maximum levels

in these years.

NATURAL INVADERS

Geologically, Sardinia is a raft which, for just

under thirty million years, has been crossing the we-

stern Mediterranean, swaying like a pendulum from

the Iberian to the Italian Peninsula. Despite its long

never-ending journey the Sardinian terrane still

hosts some descendants of the original organisms

which were present when it broke away from the

continent. Among these animals, a good number of

small invertebrates which live in caves and in the

soil like the cave ground beetle Sardaphaenops su-

pramontanus Cerruti & Henrot, 1956 (Coleoptera

Carabidae) (Fig. 1), stand out.

Organisms from other Mediterranean regions

have added to this original contingent. These new

arrivals were not randomly distributed over time but

grouped into at least three great waves. The oldest

one corresponds with the Messinian salinity crisis

about 7 million years ago. This was the time when

the Mediterranean became isolated from the Atlantic

and dried up through evaporation. So Sardinia

found itself connected to Africa and the northern re-

gions. In this period many of the amphibians and

reptiles still present on the island arrived, e.g. the

endemic Sardinian Brook Newt Euproctus platyce-

phalus (Gravenhorst, 1829) (Urodela Salamandri-

dae) (Carranza & Amat, 2005), and many insects

including the lovely orthopteran Pamphagus sar-

deus (Herrich-Schaeffer, 1840) (Orthoptera Pam-

phagidae) (Fig. 2).

The second wave of arrivals came about du-

ring the sea-regressions of the ice age, in particu-

lar during the sea-regression named “Cassia” by

Italian paleontologists, which occurred between

one million and eight hundred thousand years

ago. In this period animals coming from cold fau-

nas in the north reached Sardinia. The most fa-

mous among the vertebrates is the Prolagus

sardus (Wagner, 1832), a species of rabbit (Lago-

morpha Prolagidae) extinct at the end of the XVII

century (Smith, 2008).

Among the insects there are some species

which are now limited to the peaks of the Gennar-

gentu mountains such as the Winter Moth Opero-

phtera brumata (Linnaeus, 1758) (Lepidoptera

Geometridae) (Figs. 3, 4) (Hartig, 1976; Cao,

2005). The third wave of arrivals which is still un-

finished has only one cause: man. Man caused at

first a real mass extinction among vertebrate fauna,

mammals were almost completely substituted by

species introduced by man. Paleontologists have

little doubt in claiming that boar, mouflon, deer,

fallow deer and other common animals were

brought here by man (Vigne, 1992).

HISTORICAL PERSPECTIVE

In light of this information, the Nuragic cargo

ships with a whole collection of domestic animals

could be interpreted in a more realistic and less al-

legorical sense than archaeologists have allowed

until now. An impressive ship is the one called “Ve-

tulonia” (Fig. 7) in which we can recognise pigs,

rams, a buffalo and a cow, dogs and maybe a cat

(Depalmas, 2005). In other cases there are birds

which were probably chickens. One species of tor-

toise (perhaps two), once used as food, was also

certainly introduced (Corti et al., 1999; van der

Kuyl et al., 2002), as well as the Fat (or Edible)

Dormouse, Glis glis (Linnaeus, 1766) (Rodentia

Gliridae) (Massed, 2005).

We know little or nothing about the arrival of

insects and other invertebrates with man. Almost

certainly he arrived with his retinue of synanthro-

pic insects such as cockroaches (Figs. 5, 6) or lice

(Robinson, 1996). But he surely also introduced

other species in the ballast of ancient ships or with

the transport of wood. Among these could have

been termites.

Impact of alien insect pests on Sardinian landscape and culture

299

Figures 1-7. Some old invaders. Fig. 1. Sardaphaenops supramontanus (Coleoptera Carabidae), a eave-dwelling inseet be-

longing to the original eontingent of Sardinian Fauna. Fig. 2. Pamphagus sardeus (Orthoptera Pamphagidae), a lovely or-

thopteran belonging to the oldest wave of “invaders” eorresponds to the time of Messinian salinity erisis about 7 million

years ago. Figs. 3, 4. The Winter Moth Operophtera brumata (Lepidoptera Geometridae), a female (3) and a male (4), today

limited in Sardinia to the peaks of the Gennargentu mountains, belonging to the second wave of arrivals during the sea-re-

gressions of the ice age. Figs. 5, 6. The Oriental Cockroach Blatta orientalis (Blattaria Blattidae), an immature female (5)

and a brachypterous male (6), belonging to the organisms dispersed by the ancient man. Fig. 7. Bronze Nuragic cargo ship

with a whole collection of domestic animals found in Vetulonia, Tuscany, and deposited at the Archaeological Museum of

Florence. Photos by G. M. Delitala (1), M. Romano (2), D. Morel (3), P. Mazzei (4), A. Giannotti (5), and S. Guermandi (6).

Figure 7 redesigned from original Leoncini’s draw in Falchi (1887) by Montelius (1924).

In the same way it is possible that the Human

Flea, Pulex irritans Linnaeus, 1758 (Siphonaptera

Pulieidae) was among the synanthropie inseets in-

trodueed. Many years later, this would serve as a

vehiele for the plague whieh would sweep aeross

Sardinia several times (Cau & Pozzi, 2003). What is

eertain is that, apart from the plague, in some thou-

sands of years of trade and travel with the widest

range of sailing ships, the inerease in the number of

speeies introdueed has remained relatively low.

300

R. A. Pantaleoni et alii

So the Phoenicians, the Punics, the Romans, the

Arabs, the Pisans, the Genoans, the Spanish and all

the others who travelled around the Mediterranean

introduced tens of species to Sardinia which were

entirely indifferent to man. They did not alter his

crops, attack his farms, or threaten his health. Al-

though there is no certain and documented proof of

this phenomenon, the distribution of some orga-

nisms seems only to be justified by anthropochory.

One of the most interesting cases is the ground bee-

tle Dicheirotrichus punicus Bedel, 1 899 (Coleop-

tera Carabidae), found only in Cagliari and in a few

stations in North Africa and other Mediterranean is-

lands including Carthage (Gridelli, 1944).

In the first 350 years after America was disco-

vered, along with the numerous crops imported

from that continent (potatoes, tomatoes, com,

beans, but also prickly pears, peppers and tobacco)

only three insect pests arrived in Europe (Corte,

1991): the Angoumois Grain Moth, Sitotroga ce-

realella (Olivier, 1789) (Lepidoptera Gelechiidae),

the Woolly Apple Aphid, Eriosoma lanigerum

(Hausmann, 1802) (Sternorrhyncha Aphididae),

and the Greenhouse Thrip, Heliothrips haemorrhoi-

dalis (Bouche, 1833) (Thysanoptera Thripidae).

Things began to change halfway through the

1 8* century with the introduction of the first steam

ships. The American continent suddenly came

much closer, just as the Far East and Australia

came closer too. Improved shipping connections,

with the birth and growth of the railway, had an in-

fluence over the entire world economy. The arrival

of low cost wheat from America caused a crisis in

Europe, forcing whole nations to change their

crops. The same crisis caused an enormous flow of

immigrants with millions of people who went in

the opposite direction seeking their fortune on the

other side of the Atlantic.

Sardinia too was the object of epochal changes

in the last 20 years of the 1800s (eighteen hun-

dreds). Firstly, through heavy exploitation of fore-

stry, with wide indiscriminate deforestation. At the

same time the expansion of sheep farming began,

which would continue unintermpted until the pre-

sent day with the arrival of Roman dairymen, pro-

ducers of pecorino cheese. In this period of sudden

social and environmental change, an insect was in-

troduced which would turn out to be probably the

most economically devastating agricultural pest in

Europe and Sardinia: the Grape Phylloxera, known

at the time as Phylloxera vastatrix Planchon, 1 868,

and nowadays as Daktulosphaira vitifoliae (Fitch,

1855) (Sternorrhyncha Phylloxeridae) (Figs. 8-12).

The Grape Phylloxera induces the formation of

leaf and root galls on American Vitis species. The

severe viticultural impact of Phylloxera became

evident when it was imported into Europe after the

1860s. It devastated the European grape, Vitis vini-

fera, vineyards first in France, then spread across

the continent, and finally around the world. The vi-

neyard and wine business collapsed first in France

then in Italy (Crovetti & Rossi, 1987). The Phyllo-

xera arrived in Sardinia in 1883 and wine produc-

tion crashed a very short time later and only

resumed after the introduction of the practice to

graft the susceptible European vine species on

rootstook of resistant American vine species at the

beginning of the 20* century. From then, vine cul-

tivation in Europe was modified with the essential

use of this rootstock (Cau, 1999). Between the com-

mercial introduction of the steam ship and the end

of the second World War the scene did not drasti-

cally change. Arrivals came in succession at a high

rate especially from the American continent, with

landings at various European ports and successive

diffusion all over the continent.

In Sardinia, the situation is only slightly diffe-

rent. Only few “invaders” never arrived, like the

Colorado Potato Beetle, Leptinotarsa deeemlineata

(Say, 1824) (Coleoptera Chrysomelidae) (Fig. 14),

a very problematic Solanaceae pest, that was intro-

duced to Europe in 1897 where it established itself

from 1 920 and came later to Italy in 1 944 (Melis,

1950). In 1946, the island was hit by the most vio-

lent locust invasion of all time. Wartime, with the

abandonment of the countryside, had created sui-

table conditions for the development of this curse.

It was treated mostly with arsenical and organo-

phosphate insecticides (Pantaleoni et al., 2004).

But two enlightened entomologists, Guido Paoli

and Francesco Boselli, proposed a biological con-

trol programme with the introduction of three na-

tural enemies of the locusts present in Italy and

absent in Sardinia: two bee-flies (Diptera Bomby-

liidae) and the beetle Mylabris variabilis (Pallas,

1781) (Coleoptera Meloidae) (Paoli & Boselli,

1947) (Fig. 15). The latter in particular, set loose

in only 22 locations, has been present all over the

island for years (Boselli, 1954; Crovetti, 1966).

For once, a new welcome guest!

Impact of alien insect pests on Sardinian landscape and culture

301

With the end of the tragie events of the Seeond

World War, a new means of transport began to de-

velop: the aeroplane. Even more eargo and people

started to eross oeeans and eontinents. Journey spe-

eds inereased and the number of organisms tran-

sported from one eountry to another inereased at

the same rate (Pellizzari & Dalla Monta, 1997).

The most important points of entry were no longer

ports but airports. In Italy, the North East beeame

one of the most frequent points of entry due to the

presenee of a higher eoneentration of Ameriean mi-

litary bases.

In Veneto for example, the Syeamore Laee Bug

Corythucha ciliata (Say, 1 832) (Heteroptera Tingi-

dae) arrived for the first time in Europe in 1966

(Servadei, 1 966) and the Aeanaloniid Planthopper

Acanalonia conica (Say, 1830) (Fulgoromorpha

Aeanaloniidae) in 2004 (D’Urso & Uliana, 2004;

2006) and many of the speeies will be mentioned

further on sueh as the Leaf-footed Conifer Seed

Bug, the Citrus Flatid Planthopper, and the Asian

Tiger Mosquito.

TODAY

New invasions of inseet pests in Sardinia have

aeeelerated dramatieally over the last few years.

Some partieularly harmful speeies arrived between

1995 and 2000, many after 2000. Often these new

pests attaek and destroy ornamental plants whieh

have beeome part of the Sardinian landseape, eau-

sing it to ehange; just as often their presenee requi-

res methods of pest management whieh are

different from the traditional methods on speeifie

erops; finally in at least one ease (the Asian Tiger

Mosquito) they pose a threat to our health. The most

signifieant examples are listed.

Figures 8-12. Daktulosphaira vitifoliae (Stemorrhyncha Phylloxeridae) (= Phylloxera vastatrix Planchon, 1868). Fig. 8.

Opening of the gall on the upper surfaee of the leaf showing the eggs of the galleeoles. Figs. 9, 10. Galleeole, dorsal (9) and

ventral (10) views. Fig. 1 1 . Leaf-galls proliferate on the underside of the leaf Fig. 12. Seetion of a gall eontaining eggs and

galleeoles. Photos by C. Cesaroni/ISE CNR Sassari.

302

R. A. Pantaleoni et alii

Some invaders attaek ornamental plants. The

Geranium Bronze Butterfly, Cacyreus marshalli

(Butler, 1898) (Lepidoptera Lyeaenidae) (Fig. 13),

is a native of southern Afriea. It was aeeidentally

introdueed into the Balearie Island of Mallorea,

Spain, probably in 1987, and sinee then it has

spread to the other Balearie Islands (Menorea and

Ibiza) and other eountries in Southern Europe (Eit-

sehberger & Stamer, 1990; Raynor, 1990; Sarto i

Monteys, 1992). The Geranium Bronze oeeurs on

eultivated geranium {Geranium and Pelargonium)

speeies in Europe and ean pass through five to six

generations per year in Mediterranean loeations

(Favilli & Manganelli, 2006). Almost all eultivated

geranium varieties are at risk and plants ean be

eompletely destroyed. In Sardinia it is already

found almost everywhere (Contini et al., 2005).

The Citrus Flatid Planthopper Metcalfa prui-

nosa (Say, 1830) (Fulgoromorpha Flatidae) origi-

nates from South and Central Ameriea, and is

widespread from Quebee to Brazil. In our eountry

it was spotted for the first time 25 years ago in Tre-

viso (Veneto) (Zangheri & Donadini, 1980) and

sinee then it has spread almost all over the eountry

(Pantaleoni, 1988). The Flatid has been reported on

a long list of plants, ineluding many forest trees, or-

ehard and eitrus trees, grape and other vines, nume-

rous shrubs, and some herbs. Despite its name it is

not frequently found on Citrus plants in Sardinia.

The Flatid ordinarily does very little damage to

plants but its presenee is very evident, being revea-

led by the long, eurled filaments of waxy exudate.

This woolly material often obseures the nymph pro-

dueing it (Lueehi, 2000).

The Red Palm Weevil, Rhynehophorus ferrugi-

neus (Olivier, 1790) (Coleoptera Cureulionidae)

(Figs. 17-19) originating in Southern Asia and Me-

lanesia, has been advaneing westwards very rapi-

dly sinee the mid 1980s. It reaehed the eastern

region of Saudi Arabia in 1985. Then it was diseo-

vered in Egypt in 1992. In 1994, it was eaptured in

the south of Spain and in 1999 was found in Mid-

dle East. The Red Palm Weevil is a large reddish

brown beetle about 3 em long. Usually the damage

eaused by the larvae is visible only long after infe-

station, and by the time the first symptoms of the

attaek appear, they are so serious that they gene-

rally result in the death of the tree. In the Mediter-

ranean area the main palm eoneemed is Phoenix

eanariensis, the most eommon ornamental speeies,

but it eould attaek some other palms (Malumphy

& Moran, 2007). In Sardinia, the Red Palm Weevil

was diseovered in 2007 in Barisardo (Central West

Sardinia) on Phoenix eanariensis plants imported

from plant nurseries in the Campania region. It has

reeently been found also in the urban green areas

of Pula (South Sardinia) on palms whieh eame

from Sieily and it is expeeted that it will spread

throughout the whole region.

Also pine forests have been affeeted by the ar-

rival of new pests. In an area of the South West of

Sardinia from 2006, the Pine Proeessionary Moth,

Traumatoeampa pityoeampa (Denis & Sehiffer-

rniiller, 1776) (Lepidoptera Notodontidae), has

been found (Lueiano et al., 2007). It is eonsidered

among the most important limiting faetors for both

the growth and survival of pine forests in Southern

Europe and Mediterranean eountries (Laurent-

Hervouet, 1986). In reeent years, the speeies has

shown a tendeney to expand its range to upper la-

titude and elevation and large outbreak areas have

been observed in regions where the pest was ab-

sent or rarely reeorded (Battisti et al., 2005; 2006).

As a eonsequenee, also in Sardinia, the speeies

will foreseeably have a strong soeio-eeonomieal

impaet also due to its eaterpillars that have tiny

sharp barbed hairs and a toxin whieh ean eause ir-

ritation and allergie reaetions in people and ani-

mals (Lamy, 1990).

The Nearetie Leaf-footed Conifer Seed Bug,

Leptoglossus oceidentalis Heidemann, 1910 (Hete-

roptera Coreidae), is eonsidered a severe pest for eo-

nifer seed orehards, and it sometimes eauses serious

alarm when large numbers of adults suddenly invade

houses looking for overwintering sites (Bernardi-

nelli & Zandigiaeomo, 2001). It is a big inseet, the

adults are 9-18 mm long. This inseet was never re-

eorded for the European fauna, but in 1999 it was

first eolleeted near Vieenza (Veneto) (Taylor et al.,

2001). Up to now several speeimens have been ob-

served in different loealities of Italy ineluding Sar-

dinia (Vieidomini & Pignataro, 2007).

We eannot ignore the Euealyptus either. On this

originally Australian tree, the Yellow Euealypt Lon-

gieom, Phoraeantha reeurva Newman, 1 840 (Co-

leoptera Cerambyeidae), has reeently arrived (Cillo

et al., 2006). A wood feeder beetle whieh is slowly

ousting the Common Euealypt Longieom, Phora-

Impact of alien insect pests on Sardinian landscape and culture

303

Figures 13-19. Many kinds of invaders. Fig. 13. The Geranium Bronze Butterfly, Cacyreus marshalli (Lepidoptera Lyeaenidae),

a reeent “urban” invader from South Afriea: two old speeimens drinking from the soil moisture. Fig. 14. Colorado Potato

Beetle, Leptinotarsa decemlineata (Coleoptera Chrysomelidae), an old invader of Europe whieh never arrived in Sardinia.

Fig. 15. Mylabris variabilis (Coleoptera Meloidae), a loeust enemy introdueed into Sardinia from Italy. Fig. 16. Aedes albo-

pictus (Diptera Culieidae), an invader withpublie health implieations: adult female. Figs. 17-19. Rhynchophorus ferrugineus

(Coleoptera Cureulionidae), an Asiatie invader that attaeks palms: damage (17) and male adult (18, 19). Photos by M. Tomasi

(13), E. Musumeei (14), B. de Ruvo (15), S. Deliperi/University of Sassari (17), C. Cesaroni/ISE CNR Sassari (16, 18, 19).

cantha semipunctata (Fabricius, 1775), Australian

too, and present in Europe for about thirty years

(Tassi, 1970; Cavaleaselle & Contini, 1973). A leaf-

galling wasp, Ophelimus maskelli (Ashmead, 1900)

(Flymenoptera Eulophidae) (Figs. 20, 21), oeeur-

ring on Eucalyptus in Australia, has been reeorded

reeently in Europe (Protasov et al., 2007) and Sar-

dinia; this speeies was apparently not assoeiated

304

R. A. Pantaleoni et alii

with heavy damage to Eucalyptus trees, but the

adults appear in sueh large numbers that they di-

sturb people. There have been problems espeeially

in eampsites found in Eucalyptus woods. One of

its parasitoids Closterocerus sp. (Hymenoptera Eu-

lophidae) (Fig. 22) has reeently been introdueed to

Italy (Rizzo et al., 2006) and also to the island. In

2010 the invasive Red Gum Lerp Psyllid, Glyca-

spis brimblecombei Moore, 1964 (Stemorrhyneha

Psyllidae) (Fig. 23), has been reeorded for the first

time in Italy (Faudonia & Garonna, 2010; Peris-Fe-

lipo et al., 2011) and the year after in Sardinia

(OEPP/EPPO, 2011).

In 2012 its speeifie ^diXdisiioid Psyllaephagus bli-

teus Riek, 1962 (Hymenoptera Eneyrtidae) has

been deteeted in Sardinia and Sieily (Floris, pers.

eom.) probably aeeidentally introdueed together

with the psyllid.

Figures 20-23. Insects on Eucalyptus. Figs. 20, 21. Ophelimus maskelli (Hymenoptera Eulophidae), a gall wasp: female

(20) and old leaf galls (21). Fig. 22. Closterocerus sp. (Hymenoptera Eulophidae), a parasitoid of the previous. Fig. 23. Red

Gum Lerp Psyllid, Glyeaspis brimbleeombei (Stemorrhyneha Psyllidae), the most recent invader on Euealyptus in Sardinia:

adults with eggs. Photos by C. Cesaroni/ISE CNR Sassari (20-22), V. Risoldi (23).

Impact of alien insect pests on Sardinian landscape and culture

305

Figures 24-28. The Oriental Chestnut Gall Wasp, Dryocosmus kuriphilus (Hymenoptera Cynipidae), a parthenogenetie

speeies attaeking ehestnuts that induees gall formation on shoot tips, leaves and eatkins: galls (Fig. 24), seetion of a gall

eontaining the larva (Fig. 25), magnifieation of the larva head (Fig. 26), pupa inside the gall (Fig. 27), the speeifie pa-

rasitoid Torymus sinensis (Hymenoptera Torymidae) parasitizing a gall (Fig. 28). Photos by M.Verdinelli (24-26), and

M. Fara (27, 28)/allISE CNR Sassari.

306

R. A. Pantaleoni et alii

The Oriental Chestnut Gall Wasp, Dryocosmus

kuriphilus Ydisumdiisu, 1951 (Hymenoptera Cynipi-

dae) (Figs. 24-27), native to China but introdueed

some time ago in Japan, Korea, Nepal (Abe et al.,

2007) and eastern parts of the United States (Rie-

ske, 2007), has been diseovered in our eountry, in

2002 in Piemonte (Brussino et al., 2002), then in

other Appenine regions. The speeies was found in

the spring of 2007, even in Sardinia in Barbagia di

Belvi (Nuoro). It might have been introdueed bet-

ween 2003 and 2005 on some nursery plants whieh

eame from Piemonte (Pantaleoni et al., 2007). The

gall wasp attaeks both the European ehestnut and

Euro- Japanese hybrids. The development of shoots

is extremely limited and fruetifieation is redueed.

Chestnut produetion ean reeord losses of 50-70%

(OEPP/EPPO, 2005).

This adversity is therefore extremely serious

and eould have a negative influenee on the loeal

agro-forestry eeonomy of the Barbagia di Belvi

area whieh is elosely linked to ehestnut eultiva-

tion. In order to mitigate gall wasp aetivity, in

2009, it has been introdueed its speeifie parasitoid

Torymus sinensis Kamijo, 1982 (Hymenoptera To-

rymidae) (Fig. 28).

The Asian Tiger Mosquito, Aedes albopictus

(Skuse, 1894) (Diptera Culieidae) (Fig. 16), is a

veetor speeies, whieh has spread from its original

areas in Asia to the rest of the world through ship-

ments of used tires (Eritja et al., 2005). The Asian

Tiger Mosquito was deteeted in Cagliari, in the

South of Sardinia, in 1994-95, but the prompt in-

tervention of the loeal publie health ageney aehie-

ved the eradieation of the introdueed population

(Romi, 1995; Romi et al., 1999). In autumn 2006,

the Asian Tiger Mosquito was deteeted again on

the island. It is present in two important port eities,

again Cagliari and Olbia in the North-East (Con-

tini, 2007). In the latter loeation, the Tiger Mo-

squito seems to have reaehed a high population

density in eontrast to the south of the region where

the elimatie eonditions are not so favourable for

this speeies (Cristo et al., 2006). In 2007 an out-

break of Chikungunya fever, earried by Tiger Mo-

squito, took plaee in North Italy (Rezza et al.,

2007). No predietion ean be made about spread and

persistenee of the virus in Italy. Nevertheless Italy

is at risk of infeetion with arboviruses, sueh as den-

gue virus and West Nile virus, whieh have serious

effeets on publie health.

There are still a lot of inseets to mention that are

harmful to erops. We will eite only four speeies as

an example. The Western Flower Thrip, Frankli-

niella occidentalis (Pergande, 1895) (Thysanoptera

Thripidae), is an important pest inseet in agrieulture.

This speeies of thrip is native to North Ameriea but

has spread to other eontinents via transport of infe-

sted plant material (Lueiano & Piga, 1988).

The Woolly Whitefly, Aleurothrixus floccosus

(Masked, 1895) (Sternorrhyneha Aleyrodidae), is

almost eertainly native to South Ameriea. In the

Mediterranean region it was introdueed into the

Canary Islands in 1959, then Spain and Franee, be-

fore invading mainland Italy in 1970. Now it is

kept well under eontrol by several natural enemies,

some of whieh are imported (Delrio et al., 1982;

Ortu & Ibba, 1985).

The Citrus Leafminer, Phyllocnistis citrella

Stainton, 1856 (Lepidoptera Graeillariidae) (Figs.

29-31), is a small leafmining moth that is a poten-

tially serious pest of Citrus native to Asia (Ortu et

al., 1995; 2002). The Tomato Borer Tuta absoluta

(Meyriek, 1917) (Lepidoptera Geleehiidae) is one of

the most reeent arrivals reported in Europe. This

inseet, whieh is espeeially harmful to the tomato,

was found between the winter of 2006 and the au-

tumn of 2008 first in Spain, then in Algeria, Mo-

roeeo and Corsiea, and finally in Sardinia, where

it proves to be already widespread on tomatoes

and aubergines both in the field and in the green-

house (Viggiani et al., 2009). The eontrol of all of

these speeies has required (or will require) the use

of new management teehniques by farmers.

CONCLUSIONS

Man is a tremendous re-mixer of biodiversity,

voluntarily and involuntarily. The situation in Sar-

dinia is not very different from the other regions of

Italy. Its insularity ean either defend the island from

new arrivals or make new arrivals more problema-

tie. Our new guests ean turn out to be of no impor-

tanee to human eeonomy or they eould deeply

affeet it. Sometimes the problems are so serious that

they entail real ehanges in habits, traditions and me-

thods like for example the ease of the Grape Phyl-

loxera and maybe in the future, the Oriental

Chestnut Gall Wasp.

Impact of alien insect pests on Sardinian landscape and culture

307

Figures 29-31. The Citrus Leafminer, Phyllocnistis citrella (Lepidoptera Graeillariidae), a small moth from Asia. Fig. 29. Ma-

gnifieation of the larva. Figs. 30, 31. Mine and larva on the underside of the leaf Photos by C. Cesaroni/ISE CNR Sassari.

In other words, cultural changes. Other times

they could even lead to deeper changes. What will

happen to the palm-lined city streets, balconies flo-

wering with geraniums, pine- woods? A great deal

will depend on our ability to manage these pro-

blems.

ACKNOWLEDGMENTS

We express our deep gratitude to the friends

Giuseppe M. Delitala, Bruno de Ruvo, Marcella

Fara, Antonio Giannotti, Stefano Guermandi, Paolo

Mazzei, Daniel Morel, Enzo Musumeci, Vittorio

Risoldi, Marcello Romano, Mirko Tomasi for pro-

viding their marvellous photos. We also thank Drs.

Stefania Bagella and Anna Depalmas of Sassari

University for useful information on the Nuragic

cargo ship “Vetulonia”.

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Biodiversity Journal, 2012, 3 (4): 311-330

Lampione, a paradigmatic case of Mediterranean island bio

diversity

Pietro Lo Cascio' & Salvatore Pasta^

'Associazione Nesos, via Vittorio Emanuele 24, 98055 Lipari, Messina, Italy; e-mail: ploeaseio@nesos.org

^CNR - Istituto di Genetiea Vegetale, UOS Palermo, Corso Calatafimi 414, 90129 Palermo, Italy; e-mail: salvatore.pasta@igv.enr.it

ABSTRACT The papers aims at underlining the “unespeeted” value of Lampione’s biological heritage, as

well as the fragility of its ecosystem. Despite its very little size, this islet harbours a very rich

pool of plant and animal species of high biological and/or conservation interest. Special at-

tention is paid to the biogeographic meaning of local endemics, on local extinction and tur-

nover processes, on some ecological or biological patterns which contribute to the

distinctiveness of local biota. However, further investigations are needed in order to complete

the list of animals and to monitor the demographic trends of all species. In particular, it is ne-

cessary to assess if local seagull colony may represent a major threat for local diversity.

KEY WORDS island biogeography; conservation biology; rate of endemism; extinction; micro-insularity.

Received 12.05.2012; accepted 23.09.2012; printed 30.12.2012

Proceedings of the International Congress “Insularity and Biodiversity”, May 11*-13*, 2012 - Palermo (Italy)

INTRODUCTION

In I960, the German botanist J. Kohimeyer has

entitled a short note “Lampione, an unspoilt island

of the Mediterranean” (Kohimeyer, 1960a). This

definition, even if only partially true, is rather re-

presentative of the feeling that this islet may tran-

smit to its oeeasional visitors, espeeially if they

are naturalists.

The extremely isolated geographieal position,

the low profile that impedes to pereeive Lampione

(Fig. 1) on the horizon from long distanee, the diffi-

eulty of landing due to the frequent adverse sea eon-

ditions and to old health laws whieh forbade to visit

the islet unless the travelers underwent a long qua-

rantine, and espeeially the oeeurrenee of just faint

traees of an aneient human presenee, might do to re-

gard this plaee as a natural refuge where loeal biota

has not undergone drastie anthropie disturbanee that,

e.g. in the neighboring island of Lampedusa, has

strongly affeeted the present natural landseape.

Despite this apparent integrity, however, the

maintenanee of its biodiversity and the risk of a fast

environmental degradation seem to be regulated by

fragile equilibriums: in faet, a remarkable biologi-

eal value for Lampione has been highlighted

through two eenturies of seientifie exploration star-

ted with G. Gussone, who landed there in August

1828 (Gussone, 1832-1834; 1839), and whieh has

been eontinued by a number of botanists and zoo-

logists, in partieular around the mid-twentieth een-

tury, when the Pelagian Islands were studied in the

framework of a biogeographieal researeh projeet

eoordinated by E. Zavattari (Zavattari et al., 1961).

These investigations provided a rather exten-

sive knowledge on the loeal floristie and faunal

eommunities (Di Maria di Monterosato, 1892; Gi-

glioli, 1886; 1907; Mertens, 1926; Di Martino,

1958; 1961; Bernard, 1958; Kohimeyer, 1960a;

1960b; Gridelli, 1961; Lanza & Bruzzone, 1961;

Catanzaro, 1968; Moltoni, 1970; lapiehino &

Massa, 1989; Beekmann, 1992; Bartolo & Brullo,

312

Pietro Lo Cascio & Salvatore Pasta

1993; Baccetti et al., 1995; Mei, 1995; Cianfanelli,

2002; La Manila et al., 2002; Pasta, 2002b; 2002c;

Sferlazzo, 2003; Goggi, 2004; Lo Cascio, 2004),

leading to the description of new endemic taxa (cf.

Gussone, 1832-1834; Arcangeli, 1955; Canzoneri,

1972; Di Marco et al., 2002; Brullo et al., 2009;

Leo & Lo Cascio, in press); but have also allowed

to understand the degree of complexity that, in

particular, characterizes the relationships between

local plant communities and seagulls, extinction

rate, resources sharing, ecological adaptations,

etc., under harsh micro-insular constraints (cf.

Pasta, 2002b; Carretero et al., 2010; Lo Cascio,

2010; Lo Cascio & Massa, 2010).

In the present paper, an overview of the main

biogeographical and ecological traits of that ex-

treme example of Mediterranean insularity repre-

sented by the islet is given, based both on the

analysis of the available literature and on updated

information, obtained during a 10-year field work

carried out at Lampione. ABBREVIATIONS.

PLC = P. Lo Cascio; SP = S. Pasta.

GEOGRAPHICAL AND HISTORICAL

SETTING

Lampione (Fig. 1) (35°33’00” N - 12°19’11”E

Greenwich) is the smallest of the Pelagian Islands

(Channel of Sicily), with a surface of 0.021 km^.

750 m of coastal perimeter, and a maximum eleva-

tion of 36 m a.s.l. The islet is located 17 km off the

W coast of Lampedusa Island, and its morphology

is characterized by the occurrence of a flat top

which slopes gradually to the East, while the we-

stern side is a vertical cliff. Lampione is entirely

calcareous, with dolomitized carbonates composed

by associated wackestone and packestone referred

to the “Halk al-Menzel Formation” (Tunisian off-

shore, 46-34 Myrs BP: Bonnefous & Bismuth,

1982; Grasso et al., 1985). Thus, it belongs to the

African plate and its definitive isolation from North

Africa (as well as to Lampedusa) only dates back

to the last eustatic sea event (i.e. 18 Kyrs BP).

Except for few meteorological data collected

during a physical-astronomical expedition based on

the islet in 1971 (Cappatelli & Righini, 1972), no

information is available about its climate, although

the latter should not differ significantly from that of

Lampedusa, with an average annual rainfall and

temperature, respectively, of 320 mm and 19 °C

(see Pasta, 2002a and references therein). In parti-

cular, during the xeric season (from early April to

late October), rainfall results generally lesser than

35 mm and average monthly temperature ranges

from 18.7 to 26.1 °C (Vittorini, 1973). The islet is

now uninhabitated, but an early human presence,

probably only seasonal, is evidenced by the ruins

of some buildings, which have been referred to the

late Roman age (Smyth, 1824; Ashby & Litt, 1912).

Figure 1. The Islet of Lampione, Pelagian Arehipelago (Sieilian Channel, Mediterranean Sea).

Lampione, a paradigmatic case of Mediterranean island biodiversity

313

TAXA

SP & PLC

AIZOACEAE

Mesembryanthemum nodiflorum L.

AMARYLLIDACEAE

Allium commutatum Guss.

Pancratium sp.

APIACEAE

Daucus rupestris Guss.

ASPARAGACEAE

Asparagus horridus L.

Bellevalia pelagica C. Bmllo, Bmllo & Pasta

ASTERACEAE

Senecio leucanthemifolius Poir. s.l.

CAPPARACEAE

Capparis orientalis Veill.

CHENOPODIACEAE

Atriplex halimus L.

Halimione portulacoides (L.) Aellen

Arthrocnemum macrostachyum (Moric.) Moris

Sarcocornia fruticosa (L.) A. J. Scott

CONVOLVULACEAE

Convolvulus lineatus L.

Convolvulus siculus L.

CUSCUTACEAE

Cuscuta epithymum (L.) L.

EUPHORBIACEAE

Mercurialis annua L.

FABACEAE

Melilotus indicus All.

Melilotus sulcatus Desf.

Medicago truncatula Gaertn.

Lotus edulis L.

FRANKENIACEAE

Frankenia laevis L.

MALVACEAE

Malva veneta (Mill.) Soldano, Banfi & Galasso

OROBANCHACEAE

Orobanche amethystea Thuill.

Orobanche pubescens Dum.-Urv.

PAPAVERACEAE

Fumaria cfr. bastardii Boreau

314

Pietro Lo Cascio & Salvatore Pasta

PLUMBAGINACEAE

Limonium albidum Guss.

POACEAE

Catapodium rigidum (L.) C.E. Hubb. subsp. rigidum

Dactylis glomerata L. subsp. hispanica (Roth) Nyman

Trachynia distachya (L.) Link

Hordeum leporinum Link

Parapholis incurva (L.) C.E. Hubb.

SOLANACEAE

Lycium intricatum Boiss.

Table 1. Diachronic list of the vascular flora of Lampione. Families and species are listed in alphabetical order. G: Gussone

(1828); DM: Di Martino (from 1955 to 1958); K: Kohlmeyer (1959); SP & PLC (from 2001 to 2010). 1) Before its descrip-

tion, Bellevalia pelagica has been recorded as Muscari comosum and Bellevalia sp., respectively, by Di Martino (1961) and

Kohlmeyer (1960b); 2) according to Domina et al. (2011), locality not confirmed for this species.

Finally, during the 20th century an automatized li-

ghthouse was built by the Italian Navy.

BIOLOGICAL DIVERSITY

Present knowledge on plant and animal di-

versity and abundance

An up-to-date list of the species of vascular

plants recorded for Lampione is given in Table 1 .

Furthermore, Kohlmeyer (1960b) quoted for the

islet two lichens, Collema sp. and Roccella fucoides

(Neck.) Vain., both identified by F. Mattick, while

an unidentified mushroom belonging to the genus

Psalliota has been recently collected by one of us

(SP). Reliable data on the population size of plant

species are known just for the endemics Bellevalia

pelagica C. Brullo, Brullo & Pasta (Fig. 11), which

occurs with about 60 individuals (Brullo et al.,

2009), and Limonium albidum Guss., whose consi-

stence may be estimated in 20-30 individuals.

Other species also are represented by few (e.g.

Lycium intricatum Boiss.) or even by a single indi-

vidual (e.g. Asparagus horridus L.) (Sferlazzo,

2003). Available information on invertebrates

(Table 2) is yet partial: in fact, for some faunal

groups (Arachnida Acarida, Chilopoda, Insecta

Diptera, Insecta Hymenoptera except Fomiicidae),

even if occurring at Lampione, no records are given

in literature as well as no specimens were collected

or studied during recent samplings; similarly, there

are no data on the consistence of local vertebrate

populations. For the islet. La Mantia et al. (2002)

have listed 39 species of birds, mostly migrants, but

also including 4 breeding species: Cory’s shearwa-

ter, Calonectris diomedea (Scopoli, 1769) (Fig. 7),

Storm petrel, Hydrobates pelagicus (Linnaeus,

1758), Yellow-legged gull, Larus michahellis (Nau-

mann, 1 840), and Eleonora’s falcon, Falco eleono-

rae Gene, 1839 (Fig. 6). Among them.

Yellow-legged gull is the largely dominant species

in the Lampione ecosystem, with a colony of about

250 nesting pairs; Cory’s shearwater and Eleonora’s

falcon occur respectively with about 50 and 5 pairs,

while local consistence of Stonu petrel is uncertain,

but probably less than 10 pairs (La Mantia et al.,

2002; PLC, unpublished data). Apart from an old

record of the occurrence of the Monk seal (Smyth,

1 824), the only terrestrial vertebrates are the Ocel-

lated skink, Chalcides ocellatus (Forsskal,

1775)(Fig. 13), and the Maltese wall lizard, Podar-

cis jilfolensis (Bedriaga, 1876) (Mertens, 1926;

Lanza & Bruzzone, 1961) (Fig. 5).

Both species are represented on the islet by large

populations: using standard methods, Lo Cascio et

al. (2006) estimated for Maltese wall lizard a den-

sity of 7,500-8,000 individuals/ha (i.e. 15,000-

16,800 individuals on the whole islet), while from

field observations the ratio of apparent abundance

between this species and Ocellated skink was 3 : 1

approximatively (Carretero et al., 2010), estimating

for this latter a probable consistence of about 5,000

individuals.

Lampione, a paradigmatic case of Mediterranean island biodiversity

315

TAXA

REMARKS

GASTROPODA

PULMONATA

Clausiliidae

Lampedusa lopadusae subsp. nodulosa

(Monterosato, 1892)

this taxon has been negleeted by authors

after its deseription (see Cianfanelli,2002

and referenees therein), but should be

eonsidered an endemie subspeeies (Li-

berto et al., 2012; Nordsieek, s.d.)

Ellobiidae

Ovatella myosotis (Drapamaud, 1801)

Enidae

Chondrula pupa (L., 1758)

Helicidae

Cantareus apertus (Bom, 1778)

Eobania vermiculata (O.F. Muller, 1774)

Theba pisana (O.F. Muller, 1774)

Hygromiidae

Caracollina lenticula (Miehaud, 1831)

Cernuella virgata (Da Costa, 1778)

Trochoidea afF. cumiae (Caleara, 1847)

the loeal population is extremely differen-

tiated from those of Lampedusa and pro-

bably belongs to an endemie, undeseribed

speeies (Cianfanelli, 2002)

Sphincterochilidae

Sphincterochila candidissima

(Drapamaud, 1801)

now extinet (Cianfanelli, 2002)

ARACHNIDA

PSEUDOSCORPIONES

Olpiidae

Calocheiridius olivieri (Simon, 1879)

new record

Olpium pallipes (Lueas, 1849)

new record

ARACHNIDA ARANEAE

Dysderidae

Dysdera sp.

new record

Hahniidae

Hahnia sp.

new record

Gnaphosidae

genus and speeies unidentified

new record

Palpimanidae

Palpimanus gibbulus Dufour, 1 820

new record

Prodidomidae

Prodidomus amaranthinus (Lueas, 1846)

new and first record for Italian fauna

Salticidae

Euophrys sp.

new record

MALACOSTRACA

ISOPODA

Armadillidiidae

Armadillidium hirtum subsp. pelagicum

Areangeli, 1955

uncertain taxonomie status, according to

Camso & Lombardo (1995), who have not

seen the type material of this endemic sub-

species

316

Pietro Lo Cascio & Salvatore Pasta

TAXA

REMARKS

INSECTA ZYGENTOMA

Lepismatidae

Ctenolepisma ciliata (Dufour, 1831)

INSECTA EMBIOPTERA

Embiidae

Embia ramburi Rimsky-Korsakow, 1905

recorded by Kohlmeyer (1960b) from only

one specimen provisionally identified by K.

Friederichs; however, other specimens re-

cently collected and now under study show

strong morphological differences from E.

ramburi (PEC, unpubl. data)

INSECTA ORTHOPTERA

Acrididae

CalUptamus barbarus (Costa, 1836)

INSECTA

STERNORRHYNCHA

Aphididae

Dysaphis crataegi (Kaltenbach, 1 843)

INSECTA HETEROPTERA

Pyrrhocoridae

Scantius aegyptius (L., 1758)

INSECTA COLEOPTERA

Anobiidae

Ptinus obesus Lucas, 1847

new record

Apionidae

Malvapion malvae (R, 1775)

Carabidae

Cerambycidae

Syntomus fuscomaculatus (Motschulsky,

1844)

Parmena algirica Laporte de Castelnau,

1840

it has been previously referred by Kohlme

yer (1960b) and Sama (1988) to P. pube-

scens; some specimens have been reared

from small branches of Malva veneta

Coccinellidae

Tytthaspis sp.

new record

Curculionidae

Amaurorhinus bewickianus (Wollaston, 1860)

Dermestidae

Otiorhynchus poggii Di Marco, Osella &

Zuppa, 2002

Thorictus sp.

collected specimens are still under study

and probably belong to a N-African spe-

cies; new record

Melyridae

Aplocnemus pectinatus (Kiister, 1849)

new record

Melolonthidae

Geotrogus vorax Marseul, 1878

Mordellidae

Mordellistena oranensis Pic, 1900

unique record for Italy (Goggi, 2004)

Tenebrionidae

Catomus sp.

collected specimens are still under study

and probably belong to a N-African spe-

cies; new record

Lampione, a paradigmatic case of Mediterranean island biodiversity

317

TAXA

REMARKS

Tenebrionidae

Eutagenia aegyptiaca tunisea

Normand,1936

Glabrasida puncticollis moltonii

(Canzoneri, 1972)

Machlopsis doderoi Gridelli, 1930

new record

Opatrum validum rottembergi Canzoneri, 1972

Tentyria n. sp. Leo & Lo Caseio, in press

the loeal population belongs to a new spe-

eies; previously it has been referred to T.

sommieri (Canzoneri, 1972; Goggi, 2004)

INSECTA LEPIDOPTERA

Geleehiidae

Pexicopia malvella (Hiibner, 1 805)

Hesperiidae

Carcharodus sp.

INSECTA HYMENOPTERA

Formieidae

Tetramorium sp.

reeorded as T punicum by Bernard (1958)

but surely misidentified, aeeording to Mei

(1995)

Table 2. List of invertebrates of Lampione. Among orders and suborders, families and speeies have been listed in alpha-

betieal order. New reeords (in bold) for Pseudoseorpiones, Araneae and Coleoptera are given on the basis of speeimens

identified, respeetively, by G. Gardini, P. Pantini and PLC; the material is kept in their eolleetions.

PLANT SPECIES

REPRODUCTIVE STRATEGY

DISPERSAL STRATEGY

Allium commutatum

entomogamy

baroehory

Arthrocnemum macrostachyum

anemogamy

hydroehory

Asparagus horridus

entomogamy

endozooehory

Atriplex halimus

auto/ entomogamy

anemoehory

Bellevalia pelagica

entomogamy

baroehory

Capparis orientalis

entomogamy

endozooehory

Convolvulus siculus

entomogamy

baroehory

Frankenia laevis

entomogamy

baroehory

Fumaria ef bastardii

autogamy

baroehory

Limonium albidum

entomogamy

epizoo/baroehory

Lycium intricatum

entomogamy

endozooehory

Malva veneta

entomogamy

baroehory

Melilotus sulcatus

entomogamy

epizooehory

Mercurialis annua

entomo/ anemogamy

myrmeeoehory

Mesembryanthemum nodiflorum

entomogamy

baroehory

Pancratium sp.

entomogamy

baroehory/hydroehory

Table 3. Reproduetive and dispersal strategies of the vaseular plants oeeurring at Lampione.

318

Pietro Lo Cascio & Salvatore Pasta

Conservation status

Despite the very small geographieal range and

size of their populations, neither of the two endemie

plant speeies of Lampione is proteeted by national

and regional legislation. More in detail, three spe-

eies figured within the regional red lists eoneeming

Italian vaseular plants eompiled by Conti et al.

(1997). Following lUCN risk assessment, Daucus

rupestris (apparently extinet on the islet) is eonsi-

dered “EN” (= endangered), while to Limonium al-

bidum and Lycium intricatum the risk eategory

“LR” (= Low risk) was assigned.

Among them, only Daucus rupestris still figures

within the updated red list of Raimondo et al.

(2011). For the reeently deseribed Bellevalia pela-

gica, Brullo et al. (2009) have proposed the lUCN

risk eategory CR (= “Critieally endangered”) B2ab

(ii,v); C2a (ii). Also the oeeurring endemie inverte-

brates (see below) are not proteeted by the existing

legislation. In eontrast, Maltese wall lizard and

Oeellated skink are ineluded in the Annex 4 of the

EU Direetive 92/43 “Habitat” and in the Annex 2

of the Bern Convention.

The loeal avifauna also has great importanee in

eonservation terms: Storm petrel, Cory’s shearwater

and Eleonora’s faleon are listed in the Annex 1 of

the EU Direetive 09/147 and in the Annex 2 of Bern

Convention; the latter two are also elassified as

SPEC2 (speeies whose breeding population is

mainly eoneentrated in Europe, with unfavourable

eonservation status) aeeording to BirdLife Interna-

tional (2004).

BIOGEOGRAPHIC ALAND ECOLOGICAL

TRAITS

Endemism and high biological value

The reeently eensused flora (Table 1) eompri-

ses 16 speeies, whieh inelude Limonium albidum

and Bellevalia pelagica, both exelusive endemies

of the islet; therefore, the rate of endemism is

equal to 12.5%, whieh results relatively high in

eomparison to that known for the eireum-Sieilian

islands (ef. Pasta, 1997; Mazzola et al., 2002; Boe-

ehieri & liriti, 2011).

However, remain still unelear both the identity

and the taxonomie status of the loeal population of

Pancratium, previously reeorded by Kohlmeyer

(1960b) and Di Martino (1961) as E maritimum L.

In faet, although De Castro et al. (2012) did not find

any genetie differenees from the “typieal” sea daf-

fodil, aeeording to numerous eeologieal, morpho-

logieal and biologieal evidenees (SP, unpublished

data), it eould belong to a different speeies, not ru-

ling out a priori that it may be another endemie ele-

ment of the islet flora.

In this ease, the rate of endemism should in-

erease to 18.7%, reaehing an outstanding level for

sueh a tiny Mediterranean islet. Morphological (Co-

lombo & Trapani, 1992) and caryological (Brullo

et al., 1995) data suggest that L. albidum is closely

related to L. lopadusanum, which occurs on the

other Pelagic islands (Brullo, 1980). They both be-

long to a quite isolated group of diploid Limonium

species, such as L. panormitanum (Tod.) Pignatti

(NW Sicily), L. hyblaeum Brullo (Egadi Archipe-

lago and SE Sicily) and L. mazarae Pignatti (SW

Sicily) (Brullo & Pavone, 1981; Trapani et al.,

1997). Interestingly, a closely related species, Li-

monium cyprium (Meikle) Hand & Buttler, is ende-

mic of the northern coasts of Cyprus (Hand, 2003).

Due to its striking resemblance, it was first descri-

bed as a subspecies of Limonium albidum (Meikle,

1983), then considered to fall within its variability

(Greuter et al., 1 989). The recently described B. pe-

lagica, instead, seems to be closely related to other

narrow endemics of the Bellevalia romana subsec-

tion, like the N African B. dolichophylla Brullo &

Minissale, from Cap Bon (NE Tunisia), and B. ga-

litensis Bocchieri & Mossa, from La Galite Island

(off the N coast of Tunisia) (Brullo et al., 2009).

Also the group of exclusive endemic inverte-

brate taxa is rather rich: it surely includes the snail

Lampedusa lopadusae subsp. nodulosa (Montero-

sato, 1 892), and the beetles Otiorhynchus poggii Di

Marco, Osella & Zuppa, 2002, Glabrasida puncti-

collis subsp. moltonii (Canzoneri, 1972) (Fig. 12),

Opatrum validum subsp. rottembergi Canzoneri,

1972, and Tentyria n. sp. (Leo & Lo Cascio, in

press). L. nodulosa is closely related to another Pe-

lagian endemic, the nominal subspecies L. lopadu-

sae (Calcara, 1846), which inhabits Lampedusa,

and belongs to a group of species with mostly E

Mediterranean distribution which includes other in-

sular endemics in the Maltese Archipelago (Giusti

et al., 1995; Liberto et al., 2012; Nordsieck, s.d.).

The weevil O. poggi is morphologically compara-

Lampione, a paradigmatic case of Mediterranean island biodiversity

319

ble with the Sicilian species belonging to the group

of O. cribricollis Gyllenhal, 1834 (Di Marco et al.,

2002). The new Tentyria Latreille, 1802 shows a

remarkable affinity with some N African congene-

rics (Leo & Lo Cascio, in press), while the two en-

demic subspecies of O. validum and G. puncticollis

belong to a W Mediterranean and a N African com-

plex of geographical taxa, respectively (Aliquo &

Soldati, 2010). According to Caruso & Lombardo

(1995), further investigations are needed to clarify

the taxonomic status of the Isopod Armadillidium

hirtum pelagicum Arcangeli, 1955, while the

endemic fauna could include also an undescribed

snail of the genus Trochoidea Brown, 1 827 which

so far has been referred to T. cumiae (Calcara,

1 847), occurring in the nearby Lampedusa Island

(cf. Cianfanelli, 2002).

Among the different faunal groups, Coleoptera

Tenebrionidae are characterized by the highest level

of endemism, equal to 50% of the specific and in-

fraspecific taxa; besides, they include Machlopsis

doderoi Gridelli, 1930, an endemic species with N

African affinity which inhabits also Lampedusa and

the nearby Conigli Islet. Furthermore, present data

show as Lampione represents the only Italian loca-

lity known for Prodidomus amaranthinus (Lucas,

1846), a spider distributed in the Mediterranean

area (cf Platnick, 2009), and for Mordellistena ora-

nensis Pic, 1 900, a N African Mordellidae (Goggi,

2004). The same is highly probable for the beetles

belonging to genera Thorictus Germar, 1834 and

Catomus Allard, 1876, recently collected on the

islet and whose identification is still in progress, as

well as for the webspinner recorded as Embia ram-

buri Rimsky-Korsakow, 1905 by Kohlmeyer

(1960b), which surely belongs to an unidentified

(probably N African) species (PLC, unpublished

data). Further investigations will allow to assess the

identity of an ant erroneously referred to Tetramo-

rium punicum (Smith, 1861) by Bernard (1958; cf.

Mei, 1995) and still unidentified; like other Pela-

gian populations, it must be closely related to N

African ones (Sanetra et al., 1999).

If the close faunal relationship between Lam-

pione and North Africa, as well as in the case of

Lampedusa, is easily explained in light of paleogeo-

graphic data, the overall biological value of the islet

is further enhanced by the occurrence of two N

African beetles, the melolonthid Geotrogus vorax

Marseul, 1878 and the tenebrionid Eutagenia ae-

gyptiaca subsp. tunisea ^orvcmnd, 1936, for which

the Pelagian Archipelago is the unique Italian area

where they have been recorded (cf. Baraud, 1985;

Aliquo & Soldati, 2010). The only endemic verte-

brate is the Maltese wall lizard, here represented

by the subspecies laurentiimuelleri (Fejervary,

1 924) which exclusively inhabits this islet, Linosa

Island, and was recently introduced at Lampedusa

(Lo Cascio & Corti, 2008). Despite the remarkable

geographical distance between the Pelagian and the

Maltese Archipelago from these island, the popu-

lations are genetically very little differentiated

from each other, and this suggests a relatively re-

cent colonization of Linosa and Lampione by this

species (Scalera et al., 2004).

Dispersability

The analysis of the dispersal modes of plants

provides an unexpected result for an islet where

seabirds seem to represent the main ecological con-

straint for vegetation composition, structure and

dynamics (Table 3).

In fact, endozoochory and epizoochory are

equal, respectively, to 18.7% and 12.5%, while the

prevailing dispersal is barochory (56.2%), not inclu-

ding in this category a species characterized by

mixed strategies (Limonium albidum). In the same

way, other long- (hydrochory and anemochory) and

short-distance dispersal modes (myrmecochory) are

less represented in the islet’s flora, all equal to 6.2%.

Finally, it is noteworthy that within Coleoptera

are largely prevailing wingless or brachypterous

species, for an amount of about 60% of the whole

fauna. This order includes all the four endemic in-

sects exclusively known for the islet, and all them

are also unable to fly.

Extinctions and turn-over

In Table 1 are summarized the results of the flo-

ristic surveys carried out at Lampione during the

early 19th century (by G. Gussone), half of the 20th

century (by A. Di Martino and J. Kohlmeyer) and

by us during the last decade.

Whereas Gussone ’s visit took place in full sum-

mer (August), so that some dormant species might

therefore have escaped to his observations, the

320

Pietro Lo Cascio & Salvatore Pasta

number of taxa (10) he found sounds reliable. Mo-

reover, the same Gussone (1839) writes: “le piante

fanerogame di Lampione non oltrepassano le 20

speeie” [the flowering plants of Lampione did not

surpass 20 speeies]. The eomparison between the

data provided by Di Martino (1961) and Kohlmeyer

(1960b), who have reeorded 26 and 19 taxa respee-

tively, and those eolleeted during the most reeent

samplings, earried about half eentury later, shows

that during this period several extinetions have oe-

eurred: in addition to one species (Fumaria cf. ba-

stardii Boreau) that could have colonized the islet

just recently, only 15 out of the 32 previously re-

corded taxa still form part of the present floristic as-

semblage; noteworthy, none of the 5 species

belonging to family Poaceae results confiimed, as

well as Apiaceae, Asteraceae, Cuscutaceae and Oro-

banchaceae are no longer represented in the islet

flora. This loss of biodiversity may be the result of

the increasing disturbance due to local seagull co-

lony (Pasta, 2002b).

The massive presence of gulls, in fact, produces

strong changes in habitat structure, particularly in

small islands where ecosystems are extremely vul-

nerable (cf Vidal et al., 1998). Seagull activity de-

termines direct and indirect effects mainly on plant

communities (Figs. 8, 9), both for i) physical distur-

bance and damages due to nesting, trampling, etc.;

ii) chemical and physical alterations of the soil, due

to the sensitive input of nutrients and organic matter

which in turn triggers nitrification and eutrophica-

tion processes (Anderson & Polis, 1999; Garcia et

al., 2002). Unfortunately, the trend of the local sea-

gull population during the last half century is un-

known, and few indirect information are given by

Moltoni (1970), which in April 1967 has seen “di-

verse coppie nidifrcanti” [several breeding pairs]:

this observation seems rather simplistic compared

to the current noteworthy size of the colony, which

includes about 250 pairs (PLC & SP, unpublished

data), thus it can be assumed that over the last few

decades there has been a significant increase.

Also the apparent population decline and spatial

segregation of the colony of Eleonora’s falcon,

more than the area occupied by gulls, could be other

indirect evidence of this trend: in August 1882, Gi-

glioli (1907) had found 12 pairs of this species ne-

sting on the open spaces of the top plateau of the

islet, now massively occupied by the seagull co-

lony, while the recently censused 5 pairs are occur-

Figure 2. Log species-log area curves for Coleoptera, Ga-

stropoda and vascular plants of Lampione, Conigli and Lam-

pedusa. Data sources: Goggi (2004) and present paper, for

Coleoptera; Cianfanelli (2002), for Gastropoda; Pasta (2001;

2002b), for vascular plants.

Figure 3. Life-form spectrum of the Lampione flora. Ch:

chamaephytes; G: geophytes; NP: nano-phanerophytes; T:

therophytes.

$1 jt}lapMseu$ ctmnmut

lading liablls

Figure 4. Frequency of trophic categories among the Cole-

optera of Lampione.

Lampione, a paradigmatic case of Mediterranean island biodiversity

321

ring exclusively in the western cliff, less frequented

by other birds (PLC & SP, pers. observ.). Anyway,

contrary to what occurs in other micro-insular en-

vironments, where seagull disturbance has determi-

ned the entry of ruderal and/or nitrophilous plant

species (cf Bocchieri, 1990; Vidal et ah, 2000; Cal-

darella et al., 2010; Lo Cascio & Pasta, 2011), at

Lampione it has caused local extinctions but not a

true turnover process.

Although seagulls might have favoured the ex-

pansion of some omithocoprophilous species al-

ready occurring in the islet, such as Malva veneta

(Mill.) Soldano, Banfi & Galasso, no data are avai-

lable to assess the possible changements affecting

the spatial distribution and the floristic composition

of local vegetation. Considering the extinction rate

within life-fonu groups (sensu Raunkiser, 1934),

Pasta (2002b) highlighted that Lampione is charac-

terized by a higher mean value (50.0) compared to

Lampedusa (25.3) and Conigli Islet (31.0).

It has involved especially hemicryptophytes

(100.0) and therophytes (76.5), while other groups

result less (nano-phanerophytes: 16.7) or not affec-

ted (0 value for both geophytes and chamaephytes)

by extinctions. Finally, there are no comparable data

for faunal inventories, and in all likelihood some in-

vertebrates might have been neglected or not seen

during previous samplings. The only documented

extinction concerns the snail Sphincterochila can-

didissima (Drapamaud, 1801), for which just shells

without living animals were found on the islet

(Cianfanelli, 2002).

Species poverty

Islands typically have fewer species per unit

area than mainland, and intra-archipelago species-

area curves are steeper the smaller is the surface of

each island (Rosenzweig, 1995; Whittaker, 1998).

Three groups (Coleoptera, Gastropoda and vascular

plants) offer a good example of this insular trait. In

fact, comparing Lampione with Lampedusa and

Conigli Islet, whose surface is respectively 20.20

and 0.044 km^, speciesjQg-areajQg correlation resul-

ted highly significant for beetles (P = 0.011) and

snails (P = 0.037), while for plants no significant

con*elation occurs (P = 0.274) (Fig. 2). This result

suggests that plant richness may be influenced by

other geographical features, and primarily by the di-

stance from the main pool source: in fact, Conigli

Islet, located few meters off the S coast of Lampe-

dusa, harbours 78 species vs. 16 censused on the

farthest Lampione.

Micro-insular and local disharmony

Using the term “disharmony”, island biogeogra-

phers indicated the different balance of species

compared to equivalent patches of mainland. In

fact, islands are disharmonic as their biotas depend

only from the dispersive portion of the mainland

pool, but this fact must be distinguished from sim-

ple impoverishment, as it should not be merely a

random subset of a potential mainland pool that is

missing (Whittaker, 1998).

Concerning Lampione, which has a continental

origin and whose definitive isolation occurred ra-

ther recently, disharmony should not be related to

dispersal ability of propagules, while it could reflect

other constraints (e.g. climatic features, soil com-

position and structure, etc.), mostly still unclear,

which seem to have acted as selective forces in the

assemblage of its unbalanced ecosystem. Life-form

spectrum of plant comiuunity results dishamionic,

due to the exceptionally low number of therophytes

(representing 60% of the whole Pelagian vascular

flora, see Mazzola et al., 2002) and, contrariwise,

to the abundance of nano-phanerophytes (Fig. 3).

Anyway, the latter is a rather common pattern in

other circum-Sicilian islets (Pasta, 1997), while the

unusually low representation of the annual species

depends from the above-mentioned loss of plant di-

versity which occurred during the last 50 years.

Furthermore, it should be noted the absence of

some life-form groups (hemicryptophytes, phane-

rophytes) which instead are found in the plant com-

munities of other tiny islets, such as Conigli (Pasta,

2002b). A certain degree of disharmony characte-

rizes also the invertebrate fauna. For instance, Te-

nebrionidae are equal to 35% of the whole

coleopteran species, and to 14% of the invertebra-

tes occurring at Lampione. To better understand

this fact, it should be considered that this family

represents only 11% of the coleopteran fauna in the

near Lampedusa Island.

The over-representation of darkling beetles is

assumed to be a typical trait of micro-insular envi-

ronments indeed: at Alboran, an islet of 0.071 km^

322

Pietro Lo Cascio & Salvatore Pasta

of surface that lies in the middle of the homony-

mous sea ehannel between S Spain and N Mo-

roeeo, this group is equal to 60% of the loeal

eoleopteran assemblage, whieh ineludes 10 spe-

eies, and about 16% of the whole terrestrial inver-

tebrates (Aguirre, 2006).

On the eontrary, the low number of Gastropoda

reeorded on this islet (1 vs. 9 oeeurring at Lam-

pione: ef Aguirre, 2006; Cianfanelli, 2002) depends

from the geologieal origin of the islets, beeause

snails are generally less abundant on voleanie base-

poor outerops sueh as those of Alboran.

Also, an over-representation eoneems the tro-

phie groups of Coleoptera, where detritivores

(whieh inelude Ptinus obesus Lueas, 1847, Thoric-

tus sp. and all the Tenebrionidae) are extremely

abundant if compared to other groups whieh usually

are dominant in Mediterranean environments (such

as phytophagous, including anthophagous and rhy-

zophagous species) (Fig. 4). However, the latter

eould be explained by the abundanee of debris that,

in addition to the litter produeed by the biologieal

eyele of plants, at Lampione is due to the presenee

of a large gull eolony.

Another disharmonie trait of the islet biota is re-

presented by the eomposition of loeal herpetofauna,

whieh ineludes two Saurians belonging to Laeerti-

dae and Seineidae but no Gekkonidae. This faet stri-

kes attention, beeause Gekkonidae i) are extremely

eommon in the xerie environments of other Pela-

gian Islands, and ii) are generally more able than

other Reptiles to survive in very small insular areas

(Corti et al., 2006; Lo Caseio & Corti, 2008).

Considering that Lampione harbours the likely

autoehthonous Chalcides ocellatus, it is difficult to

explain the absence of Tarentola mauritanica (L.,

1758), a species widely distributed on the nearby

areas (Lampedusa, Conigli and North Afriea) whieh

were repeatedly eonneeted to this islet during the

last marine regressions, unless we suppose that the

present herpetofaunal disharmony hides superve-

ning events that may have eaused a loeal extinetion.

Ecological or evolutionary responses?

Islands are eommonly indieated as both evolu-

tionary and eeologieal laboratories. Short- and long-

term ehanges oeeurring in life history of island

speeies may be eombined under the term of “island

Figure 5. A male of Podarcis filfolensis elimbing on Malva ve-

neta in seareh of food. Figures 6, 7. An overview on the biodi-

versity of Lampione: a young Falco eleonorae, Oetober 2005

(Fig. 6.) Calonectris diomedea in the nest, June 2005 (Fig. 7).

Lampione, a paradigmatic case of Mediterranean island biodiversity

323

phenomenons”, or “island rule” in its wider signi-

fieance (Femandez-Palaeios, 2010). However, it is

not always elear whether proeesses and meeha-

nisms of insular adaptation refleet eeologieal or

evolutionary time-seales. For instanee, if further

researeh will eonfirm the identity of the Pancra-

tium sp. found at Lampione as P. maritimum, as

previously reported by Di Martino (1961) and Ko-

hlmeyer (1960b), its eeology would represent an

uniqueness in the eontext of the Mediterranean po-

pulations of the speeies.

In faet. Sea daffodil is a stress-tolerant and

psammophilous pioneer typieal of the embryonie

sand dunes, but in this islet it oeeupies a sharply dif-

ferent niehe, adopting a new primary strategy

(sensu Grime, 2001) and aeting as ruderal-nitrophi-

lous and lithophilous speeies. An unusual pupation

strategy is aeted by the endemie Glabrasida pun-

cticollis moltonii, whose nymphs develop inside eo-

eoons. This is very likely an exeeptional behaviour

among the speeies belonging to this genus, but re-

mains under debate if it represents a distinetive trait

of the Lampione population life-history in evolu-

tionary tenns or, as more reliable, a peeuliar (or sea-

sonal) adaptation to speeial environmental eonditions

(e.g. prolonged and strong drought period, soil sear-

eity, ete.) (Lo Caseio & Massa, 2010).

Finally, lizard populations at Lampione show

eeologieal traits typieally related to miero-insula-

rity, sueh as i) high population density, and ii) ele-

vated levels of intra- and interspeeifie eompetition,

measured as tail autotomy, eannibalism and preda-

tion rates (Lo Caseio et al., 2006; Carretero et al.,

2010). In other tiny Mediterranean islets has been

observed that the oeeurrenee of large eolonies of

seabirds is often elosely related to that of high li-

zards’ densities (Pafilis et al., 2009).

Gulls do not prey generally on lizards, while the

latters appear to profit from gull presenee in diffe-

rent ways: Moltoni (1970) reeorded as oeeasionally

kestrels, Falco tinnunculus Linnaeus, 1758, preyed

on lizards, but most of the year gull aggression di-

seourages lizard predators near the islet; and, more

importantly, it is well known as gulls subsidize islet

eeosystems by importing nutrients in form of

guano, eareasses, fish seraps, ete. (ef. Anderson &

Polis, 1998), thus supporting dense lizard popula-

tions (Markwell & Daugherty, 2002; Barrett et al.,

2005; Pafilis et al., 2009). In this regard, a very si-

gnifieant episode was reported by Moltoni (1970)

as follows: “vidi una lueertola, la quale per la sete

ehe aveva, leeeava i liquid! ehe useivano da un uovo

nel quale il pieeolo aveva gia rotto il guseio” [I saw

a lizard, whieh for the thirst that had, was lieking

the fluid eoming out from an egg already broken by

a hatehling (of gull)]. Both Maltese wall lizard and

Oeellated skink at Lampione also show high rates

of tail autotomy or injuried tails (Lo Caseio et al.,

2006; PLC & SP, unpublished data), whieh seem to

be related likely to high intra- and interspeeifie ag-

gression than the oeeurrenee of predation pressure.

This behaviour is eonfirmed by the eases of

eannibalism reported for the Maltese wall lizard

and predation on the latter by Oeellated skink, re-

fleeting the moderate diet overlap due to the eon-

vergenee in trophie strategies between the two

speeies (Carretero et al., 2010). In faet, Chalcides

ocellatus preys upon medium- or large-sized bee-

tles and inseet larvae, while the diet of P. filfolensis

is mainly based on ants and smaller preys not eon-

sumed by the skink, but both speeies share a remar-

kable eonsumption of vegetal matter. While partial

herbivorism is relatively eommon among insular

laeertid lizards (ef. Perez-Mellado & Corti, 1993),

it is absolutely unknown for eontinental popula-

tions of Oeellated skink whereas it has been found

just within insular ones (Lo Caseio et al., 2008);

for this speeies, there is also a trend for inereasing

the degree of herbivory with isolation and island

surfaee deerease, as eonfirmed by the very high

proportion (about 50%) of vegetal matter reeorded

in the Lampione diet (Carretero et al., 2010).

Therefore, in this ease evolutionary history, ra-

ther than resouree partitioning, seems responsible

for the moderate trophie overlaps found and even

may explain why both speeies eoexist under the se-

vere eonditions of Lampione.

Is there an adequate pollination network?

Although a speeifie study on loeal pollinators

was never earried out, during ten years of field work

on the islet a number of empirieal observations on

plant-animal relationships have been gathered by

the authors, espeeially as regards pollination me-

ehanisms. The main bulk of loeal flora eonsists of

entomogamous plants, equal to 75% of the oeeur-

ring speeies, while self- and wind-pollinated plants

are both equal to 12.5% (Table 3).

324

Pietro Lo Cascio & Salvatore Pasta

Figures 8-13. An overview on the biodiversity of Lampione. Fig. 8. The flat top as it appears in July. Fig. 9. The same in May,

during the breeding season of Larus michahellis. Fig. 10. Parmena algirica under stones. Fig. 1 1 . Ablooming Bellevalia pelagica.

Fig. 12. Glabrasida puncticollis moltonii inside its pupal eooeon (photo by B. Massa). Fig. 13. Chalcides ocellatus.

Lampione, a paradigmatic case of Mediterranean island biodiversity

325

Efficient pollinators, such as Diptera Syrphidae

or Hymenoptera Apoidea, have been rarely seen as

flower visitors, perhaps because permanent popu-

lations are lacking on the islet, that they can reach

occasionally thank to their high dispersal ability: for

instance, just one unidentified Hymenoptera (pro-

bably an Halictidae bee) has been found on flowers

of Bellevalia pelagica (SP, unpublished data).

Conversely, Diptera Calliphoridae and Musci-

dae were found with remarkable frequency on flo-

wers and are very abundant at Lampione, as these

flies depend from avian wastes for their larval

growth, but are generally considered less efficient

pollinators (Kwak & Bekker, 2006; Perez-Banon et

ah, 2008). Therefore, it is rather intuitive that, simi-

larly to other insular ecosystems (cf. Olesen et al.,

2010), the pollination network of this islet results

extremely simplified, and further investigations

may clarify to what extent lack of adequate polli-

nators or small pollinating fauna could affect the re-

productive biology of the local plant community.

Anyway, it can be preliminarily assumed that

small population sizes, together with the shortness

of blooming period, might expose to a greater risk

some species (e.g. B. pelagica) under these severe

constraints. If pollination plays a significant role in

the maintenance of genetic variability and fitness

of plants, and pollinator scarcity may lead in some

cases to local extinctions (Barrett, 1996), it cannot

be excluded that this factor could have contributed

to some extent to the loss of plant diversity occurred

during the last fifty years. Finally, it remains unclear

the role of Maltese wall lizard as potential pollen

vector, especially on large-sized individuals of

plants such as those of the biennal Malva veneta,

where the lizards were frequently observed clim-

bing, apparently in search of flies and other insects

(PLC & SP, unpublished data).

Lizards could play an important role also as “vi-

cariant” pollinators of Allium commutatum and

Pancratium sp., as already observed in some islets

of Balearic Archipelago for the relatives am-

peloprasum L. and Pancratium maritimum L. by

Perez-Mellado et al. (2000). In particular. Pancra-

tium sp. has morphological traits that favour polli-

nation by hawkmoths (“sphingophyly” sensu

Manning & Snijman, 2002), but its local population

blooms just 1-2 weeks around mid-September,

when the weather is often characterized by rather

unsteady and windy conditions.

These conditions can result less favourable for

“optimal” pollinators like Macroglossum stellata-

rum L., 1758 (Eisikowitch & Galil, 1971) than for

lizards, which show a strong propensity for the con-

sumption of vegetal matter.

CONCLUDING REMARKS: HOW FRAGILE

AND KNOWN IS LAMPIONE ECOSYSTEM?

With its unsteady plants species’ assemblage

and its extraordinary concentration of “classical”

examples of micro-insularity, apparently far from

human disturbance but actually suffering some un-

decifered form of degradation, Lampione represents

a paradigmatic example of Mediterranean islets

realm, which an increasing attention has been paid

to in the recent past (Delanoe et al., 1996).

Some twenty years ago Greater (1991) argued

that extinction occurred very rarely within Mediter-

ranean basin. On the other hand, the available data

on the evolution of the plant assemblages of the cir-

cum-Sicilian islands (Pasta, 1997) suggest that this

statement is not totally correct. In fact, many spe-

cies, also endemic ones, have disappeared in last

decades. Also some of the most noteworthy plants

of Lampione underwent strong rarefaction (Limo-

nium albidum) or even local extinction {Daucus ru-

pestris) within last 50 years. As a matter of fact, our

fragmentary knowledge on the history of local plant

and animal populations does not enable to disentan-

gle biological crisis from normal turn-over proces-

ses (Diamond, 1976), so that it is hard to forecast

the future of the biological heritage of Lampione.

Nevertheless, the sharp changes which recently

affected both plant species number and the distri-

bution and cover rate of local vegetation, suggest

the importance of regular monitoring activities, as

recommended by the Management Plan concerning

the SCI ITA040002 “Isole di Lampedusa e Lam-

pione” (La Mantia et al., 2009).

As recently pointed out by Dominguez Lozano

et al. (2003), in order to develop a coherent and ef-

fective plant conservation strategy at least three fac-

tors should be taken into account: the

environmental range of each species, its level of

geographic rarity (both on the local and the whole

distribution range level) and its rate of threat (resul-

ting from two opposite patterns: anthropogenic in-

teraction and level of protection).

326

Pietro Lo Cascio & Salvatore Pasta

Although it is often difficult and dangerous to

make generalisations about the influence of biolo-

gical variables on rarity and threat, in our opinion

future field investigations should be concern the

biology (life-cycle, reproduction and dispersal vec-

tors, etc.) of Lampione’s “botanical highlights” and

to better evaluate the environmental effect of local

seagull colony.

Further studies are also needed in order to im-

prove the faunal knowledge and to monitor the de-

mographic trends of some species of conservation

interest, such as the marine birds and the Eleonora’s

falcon. Finally, the identity of some invertebrates

that are currently identified only at generic rank, as

well as the taxonomic status of other faunal ele-

ments previously attributed or probably belonging

to endemic forms, need to be investigated in order

to understand properly the biogeographical impor-

tance and meaning of local biological heritage.

ACKNOWLEDGEMENTS

We wish to express our gratitude to Giusi Nico-

lini, Giuseppe Sorrentino and all the staff of the Na-

ture Reserve “Isola di Lampedusa” and the Marine

Protected Area “Isole Pelagic” for their unvaluable

logistic support; to Simone Cianfanelli, Claudia

Corti, Flavia Grita, Tommaso La Mantia, Bruno

Massa and Damiano Sferlazzo, for their help during

field work; to Giulio Gardini and Paolo Pantini,

who have identified, respectively, Pseudoscorpio-

nes and Araneae collected at Lampione. Informa-

tion on Gastropoda provided by Ignazio Sparacio

was very much appreciated.

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Biodiversity Journal, 2012, 3 (4): 331-336

Vascular flora evolution in the Soqotra Archipelago (Indian

Ocean)

Gianniantonio Domina'*, Giuseppe Bazan' & Francesco Maria Raimondo^

'Dipartimento di Scienze Agrarie e Forestall, Universita di Palermo, via Arehirafi 38 - 90123 Palermo, Italy

^Dipartimento di Seienze e Teenologie Biologiehe, Chimiehe e Farmaeeutiehe, Universita degli Studi di Palermo, via Arehirafi 38

90123 Palermo, Italy

* Corresponding author: gianniantonio.domina@unipa.it

ABSTRACT The main floristic and vegetational features of the Soqotra Arehipelago are outlined. The theo-

ries of vieariance and dispersal are eommented with the support of examples suggesting the

idea that both are complementary in the establishment and evolution of the flora of Soqotra.

Finally the relation of alien vs natural elements of the flora is analyzed.

KEY WORDS Biogeography; Soqotra; alien plants.

Received 11.05.2012; accepted 7.12.2012; printed 30.12.2012

Proceedings of the International Congress “Insularity and Biodiversity”, May 11*-13*, 2012 - Palermo (Italy)

INTRODUCTION

The Soqotra Archipelago is placed in the nor-

thern part of the Indian Ocean between 12° 06’-

I2°42’N and 52°03’-54°32’E and consists of four

islands. Soqotra, the main one, rises up to 1500 m

a.s.L, 240 Km from the African Coast and more

than 400 from the Arabian peninsula. It is of conti-

nental origin in fact it was joined to the Arabian

plate not less than 15 myr ago. The stratigraphy is

characterized by an igneous and metamorphic ba-

sement complex that crops out in the Haghier

mountains and is overlaid by a plateau ranging from

300 to 900 m composed of Cretaceous and Tertiary

limestone. On the coastal plains, quaternary and re-

cent deposits of marine and fluvial origin overlie

the older limestone (Beydou & Bichan, 1970).

The existing literature on the vegetation of So-

qotra is rather heterogeneous. Recent attempts to

classify the plant communities and analyse their di-

stribution pattern can be found in Krai & Pavlis

(2006), Kiirschner et al. (2006) and De Sanctis et

al. (2012). The latter recognizes a geo-altitudinal

gradient divided in four main vegetation belts:

Semi-arid Haghier mountain from 1000 to 1550 m

a.s.L, Arid limestone hills and plateaux from 400 to

1000 m a.s.L, a transition zone from 200 to 400 m

a. si. and the Arid coastal plain from the sea level to

200 m of altitude (Fig. 1). These host eight woody

vegetation types, seven scrubland communities, six

main herbaceous vegetation types and seven groups

of halophytic vegetation concentrated in the arid

coastal plain. Due to its high levels of biodiversity

Soqotra has been designated a UNESCO World He-

ritage Site, UNESCO Man and Biosphere Reserve,

WWF Global 200 Ecoregion and Plantlife Interna-

tional Centre of Plant Diversity.

MATERIALS AND METHODS

This contribution is the result of personal obser-

vations made in the island during the 2007 expedi-

tion by the members of the Department of Botany

of the University of Palermo, exchanges of views

with Ahmed Adeeb, Ahmed Eissa Ali Afraar, Fahmi

332

G. Domina, G. Bazan & RM. Raimondo

NE SE

Figure 1 . Section of the island of Soqotra along the dominant direction of the monsoon with the vegetation belts recognized

by De Sanctis et al. (2012) (redrawn), a: Arid coastal plain; b: Transition zone; c: Arid limestone hills and plateaux; d: Semi-

arid Haghier mountain.

Abdullah Mohamed Ba Ashwan from Soqotra and

Lisa M. Banfield from Edinburgh hosted in 2009

and 2010 in the Botanieal Garden of Palermo as far

as from relevant literature.

RESULTS AND DISCUSSION

The Soqotra Arehipelago ineludes 837 vaseular

plants, 310 (37%) of whieh are endemie (Kilian &

Hein, 2006; Domina & Raimondo, 2009; Raimondo

& Domina, 2009). These speeies belong to 433 ge-

nera ineluded in 1 14 families. The ferns eonsist of 30

speeies and there is a single gymnospemi {Ephedra

foliata Boiss. ex C.A.Mey.). The family of Poaeeae

is the riehest one (90 speeies), followed by Fabaeeae

(70) and Asteraeeae, Euphorbiaeeae, Apoeynaeeae,

Aeanthaeeae and Rubiaeeae with over 20 speeies

eaeh. Together these families total more than 40% of

the flora (Miller & Morris, 2004). There are 15 ende-

mie genera: 12 ineluding only one speeies: Angkalan-

thus (Aeanthaeeae), Nirarathamnos and Oreofraga

(Apiaeeae), Duvaliandra and Socotrella (Aselepiada-

eeae) Hemicrambe, Lachnocapsa and Nesocrambe

(Brassieaeeae), Haya (Caryophyllaeeae), Dendrosi-

cyos (Cueurbitaeeae) Placopoda and Tamridaea (Ru-

biaeeae); two with 2 speeies eaeh: Trichocalyx

(Aeanthaeeae) (Fig. 2) and Rughidia (Apiaeeae); one

with 3 speeies: Ballochia (Aeanthaeeae).

Soqotra falls in an arid area that has experieneed

rainfall fluetuations over the last 150,000 years that

eaused widespread desertifieation in the region. The

varied topography of the island and the moisture

brought by oeean monsoons offered wet refugia for

many plants that have otherwise been unable to sur-

vive to arid periods that affeeted the area (Miller &

Morris, 2004). Aeeording to Takfrtajan (1986) So-

qotra belongs to the Eritreo-Arabian subregion

whieh ineludes South Arabia, Somalia, Ethiopia,

Kenya and Tanzania and is eharaeterised by high

levels of generie endemism (but not usually fami-

lies unless mono-generie) and very high levels of

speeies endemism. White (1983) ineludes the So-

qotra Arehipelago in the Somalai-Masai Regional

Centre of Endemism and eonsiders it as a loeal een-

tre of endemism due to its exelusive endemie s.

The flora of Soqotra Arehipelago shows elose re-

lations with the floras of the neighbouring Southern

Arabia and NE Afriea. Soqotra and NE Somalia

share a high number of near-endemies, e.g. Dira-

chma soqotrana Sehweinf. ex Balf.f, Caralluma so-

qotrana (Balf.f.) N.E.Br. and Erythroseris amabilis

(Balf.f.) N. Kilian & Gemeinholzer. Further biogeo-

graphie relations are with: NW Afriea, Maearonesia,

SW Afriea, S Asia, Madagasear and N Ameriea.

The speeies of Soqotra shared with the Mediter-

ranean are also spread in Tropieal Afriea, C and SW

Asia e.g. Dactyloctanium aegyptiurn (L.) Willd.,

Hyparrenia hirta (L.) Stapf (Poaeeae), Juncus bu-

fonius L. (Juneaeeae), SUene apetala Willd. (Caryo-

phyllaeeae), Erodium cicutarum (L.) (Geraniaeeae),

Galium setaceum Lamk. and Valantia hispida L.

(Rubiaeeae) or are weeds arrived in the last eentu-

ries, e.g. Anagallis arvensis L. (Primulaeeae). Si-

milarities eoneeming the Mediterranean- Soqotran

geographieal disjunetion refer to genera with ally

speeies: Cleome, Teucrium, Dipcadi, Lepturus, ete.

(Balfour, 1898) and are often related to problematie

Vascular flora evolution in the Soqotra Archipelago (Indian Ocean)

333

taxa, e.g. Convolvulus siculus L. (Convolvulaceae)

and Valerianella affinis Balf.f. (Valerianaceae) col-

lected only in the 1 9* Century but not seen recen-

tly. Strongest relationships were observed for

Highlands of Northern Somalia where 40 % of the

genera are in common with the Mediterranean re-

gion and several taxa could represent real disjun-

ctions (Fici, 1991).

Soqotra as all the oceanic islands attracted seve-

ral botanists and biogegraphers that applied traditio-

nal theories to explain the evolution of its flora. A

review of these theories is reported in Miller & Mor-

ris (2004) and Banfield et al. (2011). The main ones

concern Vicariance: Tethyan fragments, Dry Pleisto-

cene Corridor, Boreotropical Origin, and Dispersal:

Colonisation, Adaptive Radiation. Both vicariance

and dispersal look like complementary in the esta-

blishment and evolution of the flora of Soqotra.

Vicariance

Vicariance implies barriers that restrict gene flow,

the isolated populations evolve separately and be-

come unlike enough to turn different species. This is

suggested by disjunctions and anagenesys. The “Te-

thyan fragments” theory, supported by Bramwell

(1976, 1985), Marrero et al. (1998), Miller et al.

(2002), Andrus et al. (2004), Rodrigues-Sanchez &

Arroyo (2008), etc., provides for a widespread vege-

tation around the Tethys Ocean, the continental Drift

(Separation of populations) followed by aridification

that caused widespread extinctions and left of relict

populations. This theory is elucidated in Soqotra by

endemic taxa that have unclear affinities/relation-

ships with other more widespread ones, and are the-

refore assumed to be ancient relicts, e.g. Punica

protopunica Balf.f. (Punicaceae) and Dendrosicyos

soqotranus Balf.f. (Cucurbitaceae).

Other plants with a distribution split on both

sides of the African continent that could have been

generated by this phenomenon, are the arborescent

Dracaena (Dracaenaceae) with Dracaena draco L.

in Morocco and Macaronesia, D. tamaranae Mar-

rero et al. in Gran Canaria, D. serrulata Baker in

the Arabian peninsula, D. cinnabari Balf.f in So-

qotra, D. ombet Heuglin ex Kotschy et Peyr. and D.

schizantha Sinet in E Africa; and Campylanthus

(Scrophulariaceae) with 1 5 species that occur scat-

tered in Soqotra, Canaries, Cape Verde Islands, NE

Africa and Arabian peninsula.

Some taxa have distributions that support the

hypothesis of a dry corridor during the Pleistocene

(1.8 mya), linking SW Africa, Soqotra, Somalia,

Ethiopia, and Arabia. Plants showing this distribu-

tion occur within: Wellstedia (6 species: Soqotra,

Somalia, Ethiopia, South Africa, Namibia), Grade-

ria (four species occurring in Soqotra, South Africa,

Tanzania), Camptoloma (three species: Canary Is-

lands, Somalia, Arabia, Namibia, Soqotra), Chori-

sochora (three species: Soqotra, South Africa).

Some taxa have widely disjunct distributions

that stretch to the New World (America). During the

Tertiary Period (65-2 myr), a warm, wet climate and

tropical woodland vegetation covered large areas of

N America, Europe and Asia (Eurasia). The little

barriers to dispersal facilitated the spread of widely

diffused taxa. Now some of these taxa have become

separated because of: continental drift northwards,

glaciation in northern areas, increased aridity in

Northern Africa and the Middle East, severance of

the Bering Land Bridge. These phenomena caused

extinctions and barriers to dispersal and the popu-

lations remained isolated.

Plants showing a Boreotropical origin are:

Chapmannia (Fabaceae) (seven species: four in So-

qotra, one in Somalia and two in America), Tham-

nosma (Rutaceae) (six species: Soqotra, South

Africa, Somalia, Yemen and Oman).

Molecular analyses done on Echidnopsis (Apo-

cynaceae) proved that the four Soqotran endemics

sampled were monophyletic and have evolved from

a single colonising ancestor (Thiv & Meve, 2007).

Divergence time older than 35-17.6 my is required

to provide evidence for vicariance (Palaeo-ende-

mics). Closest relationship should be between So-

qotran and Arabian taxa. This was provided only for

Dendrosicyos soqotranus that has an estimated age

of 40 my that supports his vicariant history.

Dispersal

Dispersal implies the occurrence of different co-

lonization events that bring to adaptive radiation in

a particular cladogenesis (the splitting of a species

into two or more groups, which give rise to more new

species) that can occur when a taxon colonises a new

environments and rapidly evolves to fill available ni-

ches (Simpson, 1953). Adaptive radiation brings to

genera with many species, often lightly morphologi-

cally different, with high levels of endemism.

334

G. Domina, G. Bazan & RM. Raimondo

Examples in Soqotran floras are: Hibiscus (Mal-

vaeeae) with 15 speeies, 9 endemie; Heliotropium

(Boraginaeeae) with 17 speeies, 10 endemie; Heli-

chrysum with 13 speeies, 12 endemie, Boswellia

(Burseraeeae) and Pulicaria (Asteraeeae) with 7 spe-

eies, all endemie; Echidnopsis (Aselepiadaeeae) and

Hypericum (Clusiaeeae) with 5 speeies, all endemie.

However, eolonisation (or dispersal) was never

eonsidered the most important engine of differen-

tiation of the Soqotran flora due to many genera and

families with only one or two speeies that suggest

viearianee to have the biggest influenee. Diver-

genee time younger than 35-17.6 myr indieates di-

spersal events produeing neo-endemies. An

example is represented by Limonium guigliae Rai-

mondo et Domina andZ. paulayanum (Vierh.) J.R.

Edm. (Plumbaginaeeae) and their affinities with

other Limonium from Arabian peninsula (Rai-

mondo & Domina, 2009). Aeeording to Thiv et al.

(2006) three independent eolonization lineages

from the Eritreo-Arabian subregion of the Sudano-

Zambesian Region were revealed in Aerva (Ama-

ranthaeae) providing further support to

eolonization via dispersal, rather than a viearianee

origin of the island elements. The same applies to

Exacum (Gentianaeeae) (Yuan et al., 2005). Phy-

logenetie patterns indieate several dispersal events

from E Afriea and/or Arabia in several other taxa

sueh as Thamnosma (Rutaeeae), Echidnopsis

(Apoeynaeeae), Kleinia (Asteraeeae), Zygocarpum

(Fabaeeae), Polycarpaea (Caryophyllaeeae), Re-

seda (Resedaeeae), Camptoloma (Serophularia-

eeae), Pulicaria (Asteraeeae) in eontrast to more

rare affinities to Madagasear, the Masearenes, Sou-

thern Afriea, and tropieal Asia.

Aliens

Reeent ehanges in the flora that risk to have a

deep impaet both on endemie and non endemie ele-

ments are eaused by aliens. Invasive alien speeies

are widely reeognized as one of the major threats

to native biodiversity, partieularly on oeeanie is-

lands (Denslow et al., 2009; Caujape-Castells et al.,

2010; Kueffer et al., 2010). Naturalization of aliens

on islands is higher than in mainland environments

(Stohlgren et al., 2008).

A reeent survey on the alien flora of Soqotra

(Senan et al., 2012) reports 88 taxa: 17 of them natu-

ralized and four invasive: Argemone mexicana L.,

Colotropsis procera (Aiton) W.T. Aiton, Leucaena

leucocephala (Lam.) de Wi and Parkins onia aculeata

L.). The remaining are eultivated for food, forage,

medieine, or ornament but new naturalizations from

this pool are to be expeeted also eonsidering that se-

veral of them are ineluded in the Global Invasive Spe-

eies Database (http://www.issg.org/database).

Anyway these results are fortunately low if

eompared with other oeeanie islands. This eould be

due to the faet that in Soqotra agrieulture, whieh is

the main souree of alien speeies in oeeanie islands,

has always been limited to small traets of ground

near houses (Balfour, 1898); the same situation ean

be observed still nowadays. No large plantations

have been realized with the single exeeption of date

palm {Phoenix dactylifera L.) (Fig. 3) now natura-

lized and Afriean finger millet (Eleusine coracana

Gaertn.), now abandoned. Fruit, riee, vegetables

and khat (Catha edulis Forssk.) eonsumed in Soqo-

tra eome almost totally from outside. The airport,

opened in 2000, plays an important role in growth

of trade and transport faeilitations. Today there

seems to be little awareness of the problem of in-

vasive speeies, severe eontrols are done only on

plants and plant propagules that are exported out-

side the island; on the eontrary no seeurity or pest

eheek is done on the material that arrives.

CONCLUSIONS

Biogeography of oeeanie islands always attrae-

ted botanists attention. Soqotra represents an im-

portant ease study of an aneient island not

extremely isolated from the eontinents but with a

rieh and well differentiated flora that makes it an

exeellent world’s hotspot of biodiversity. Its flora

benefited of wet and dry refugia to survive geologie

and elimatie variations oeeurred in the area. A par-

tieular relevanee has the diversity in some tropieal

genera sueh as Boswellia and Commiphora that

have a high ethnobotanie interest. Its landseape is

eharaeterized by the representative oeeurrenee of

the endemie Adenium obesum subsp. sokotranum

(Vierh.) Lav. (Fig. 4). Both traditional theories and

moleeular evidenee tried to explain the origins and

modifieations of this flora using the vieariant as

well as the dispersal approaeh. But there are some

problematie taxa with unelear affinities and unelear

relationships with known relatives (e.g. Dendrosi-

Vascular fJoro evolution in the Soqotra Archipelago (Indian Ocean)

335

4 5

Figure 2. Trichocalyx obovatus Balf.f. at Flom Hil. Figure 3. Date plantation near Di Lishah, in the baekground the massif of Hag-

geher. Figure 4. Hilly shmbland c\macter\zed\yy Jatropha unicostata mdAdenium obaesum subsp. sokotranus. Figure 5. Centuries

old tree of tamarind at Zam Horn used as shelter for livestoek.

cyos soqotranus andPunica protopunica). However

the moleeular analysis eannot take into aeeount

wide-ranging extinetions that have been deseribed

for the region during aridifieation. More phytoge-

nies are therefore needed of more obvious relietual

speeies or those thought to be more aneient.

Considering the low impaet of residential and

agrieultural aetivities and the total absenee of indu-

strial ones, the most severe threat to this unique

flora is represented by the touristie development of

the island that, as happened in other parts of the

world, risks to provoke deep modifieations in the

environment damaging unequalled habitats. The

millenary balaneed relationship between man and

nature as underlined by the oeeurrenee of eenturies

old trees of tamarind (Tamarindus indica L.) (Fig.

5), used as meeting point for man and livestoek,

risks to be eonsiderably altered.

In order to attain an overall proteetion of both

human eulture and biodiversity of the island, the

loeal Administrations and the inter-govemmental

organizations need to elaborate a long time foreeast

planning addressed to an eeo-sustainable develop-

ment. In short time period a deeper eontrol on new

introduetions in order to prevent pests and new un-

desired alien taxa is indispensable.

ACKNOWLEDGEMENTS

Funding by Universita degli Studi di Palermo

(ex 60 %) and Prof. Attilio Carapezza for eritieal

reading of the text are gratefully aeknowledged. A

speeial thanks to Dr. Luigi Guiglia a true lover of

the island that guided us during our exploration.

336

G. Domina, G. Bazan & RM. Raimondo

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Biodiversity Journal, 2012, 3 (4): 337-342

Vascular flora evolution in the major Mediterranean islands

Gianniantonio Domina'*, Pasquale Marino^ Vivienne Spadaro^ & Francesco Maria Raimondo^

'Dipartimento di Scienze Agrarie e Forestall, Universita degli Studi di Palermo, via Arehirafi 38 - 90123 Palermo, Italy; e-mail:

gianniantonio . domina@unipa . it

^Dipartimento di Seienze e Teenologie Biologiehe, Chimiehe e Farmaeeutiehe, Universita degli Studi di Palermo, via Arehirafi 38

90123 Palermo, Italy

*Corresponding author

ABSTRACT Characteristics of Mediterranean island floras are analyzed with stress on endemic units. On

these bases the main relationships between the major Mediterranean areas and the inland ter-

ritories with the strongest floristic affinities are analyzed. Finally the role of aliens in Mediter-

ranean island floras and threats are discussed.

KEY WORDS Biogeography; Mediterranean; endemism; alien flora.

Received 11.05.2012; accepted 20.11.2012; printed 30.12.2012

Proceedings of the P' International Congress “Insularity and Biodiversity”, May 11*-13*, 2012 - Palermo (Italy)

INTRODUCTION

The Mediterranean is among the riehest regions

in the world for wild and eultivated speeies. Cir-

eum-Mediterranean eountries house about 25,000

speeies, almost one tenth of the world's vaseular

flora, the 63% of whieh are endemie (Greuter, 1991;

Medail & Quezel, 1999).

A peeuliarity of this area is the high amount of

species with a narrow range, many of which are local

endemics. In some sectors endemics are more than

20%. Among these there is Sicily and the other major

Mediterranean islands and island groups: Baleares,

Corse, Sardinia, Kriti, Cyprus with their mountain

areas. The main reason for this high endemism can

be searched in the pronounced habitat fragmentation

that characterises the Mediterranean area as a whole.

CHARACTERISTICS OF MEDITERRA-

NEAN ISLAND FLORAS

Mediterranean island flora is relatively well

known although each year new species, sometimes

completely unknown, are described (e.g. Ptiloste-

mon greuteri Raimondo et Domina. Fig.l). Tradi-

tionally the floras of large Mediterranean islands

are considered ancient, relatively poor in species,

rich in endemic taxa and particularly vulnerable

(Greuter, 1995).

Permanence in situ and prolonged evolutionary

standstill are salient characteristics of Mediterra-

nean island floras (Greuter, 1979). These have a re-

lictual nature and are at least ancient as the islands

themselves; for Cyprus, Crete and the Baleares

about 5-6 milion years, dating back to the post-Mes-

sinian transgression (Greuter, 1995). According to

the equilibrium theory of island biogeography

(MacArthur & Wilson, 1 967) the number of species

on an island is in a state of dynamic equilibrium;

diversity eventually stabilizes but turnover remains

high as species continuously colonize and go ex-

tinct. The flora of the Mediterranean was subject to

an impoverishment due to the climatic fluctuation

during Pliocene and a subsequent enrichment due

to immigration by long range dispersal (Quezel &

Medail, 2003). This is more manifest in the islands

where original species pool seems to become con-

338

G. Domina, P. Marino, V. Spadaro & F.M. Raimondo

siderably impoverished before immigration eould

balanee extinetion.

Indeed the open sea is a more impermeable bar-

rier for all kinds of land plants than is solid ground,

even when inhospitable. On the sea there is no

ehanee for small, ephemerous populations to get

established, no resting plaees for pollen-earrying in-

seets. Salt water will kill offmost swimming propa-

gules in a matter of hours or days, and those that

might survive will be deposited in saline habitats

hostile to most non-littoral speeies. Germination

loss after some days of saltwater exposure was ve-

rified for some Mediterranenan plants e.g. Lotus cy-

tisoides L. (Fabaeeae) and Plantago weldenii Rehb.

(Plantaginaeeae) (Potthoff, 1989), on the eontrary

to different Anfiatlantie plants that have a high re-

sistanee to saltwater (eg. Portulaca gr. oleracea L.)

(Danin et al., 1978).

Comparing island floras with the mainland ones

the ratio “speeies number/area size” does not give

Figure 1 . Ptilostemon greuteri in its natural habitat recently

found in the Inici Mt. around Castellammare del Golfo (Tra-

pani) few kilometres from the built-up area.

partieular differenee. But Mediterranean island flo-

ras are usually poor in endemies when eompared

with the relevant mainland areas (Greuter, 2001).

ENDEMISM

Number and rates of endemies in a given terri-

tory depend on the territory’s size and is not rela-

ted to its insularity. On the major Mediterranenan

island systems, around 1 0% of the speeies are en-

demie and sometimes eonfined to a single island.

This rate is lower also than in Oeeanie islands

(Table 1). In addition, in eontinental islands spe-

eies are often quite loealized and have a small

number of individuals, e.g. Abies nebrodensis

(Lojae.) Mattel in Sieily.

Adaptive radiation (Darwin, 1 854) in whieh na-

tural seleetion drives divergenee of an aneestral

speeies into deseendants that are better able to ex-

ploit eeologieal opportunity (Dobzhansky, 1948)

has no applieability in Mediterranean islands.

Whenever you look elosely at examples of varia-

ble, polymorphie groups you are likely to find mo-

saie patterns of geographieal viearianee rather than

sympatrie niehe differentiation eoneomitant with

speeiation, as is thought to be eharaeteristie of ge-

nuine adaptive radiation. An example of is the vi-

earious distribution of Anchusella variegata (L.)

Bigazzi, E.Nardi et Selvi in Italy and Northern

Greeee and Anchusella cretica (Mill.) Bigazzi,

E.Nardi et Selvi in the Peloponnesus. Geographie

viearianee is elear also within the apomietie/sexual

eomplex of Limonium. This phenomenon started

around 6 mya, at the same time as one of the most

dramatie ehanges that affeeted the Mediterranean

basin. The number of speeies of Limonium in the

major Islands varies aeeording to a deeresing gra-

dient from west to east: 57 in the Baleares, 27 in

Corse, 51 in Sardinia, 44 in Sieily, 19 in Kriti and

9 in Cyprus (Domina, 2011) (Fig. 2). Therefore the

West Mediterranean is being identified among the

main eentres of differentiation of the genus.

The islands split off from the mainland (eherso-

genous ones), like all Mediterranean ones of an ap-

preeiable size, already earried their own, fully

adapted and diversified flora when they beeame in-

sular. They are eonservative systems. Ideally, the

flora of eaeh island ean be eonsidered to be a reflee-

tion, perhaps impoverished but otherwise little

Vascular flora evolution in the major Mediterranean islands

339

Island or Archipelago

Area Km^

No. of

species

species/

Area

No. of

endemics

endemics/

Area

%of

endemics

OCEANIC

Juan Ferdinandez Island, Chile

147

1.58

118

1.27

Galapagos Islands, Ecuador

7844

543

0.07

229

0.03

Mauritius

1865

800

0.43

280

0.15

Rodrigues, Mauritius

104

145

1.39

0.46

Madeira, Portugal

769

750

0.98

129

0.17

Canary Islands, Spain

7273

2000

0.27

569

0.08

Ascension, UK

0.27

0.12

Soqotra, Yemen

3799

825

0.22

307

0.08

MEDITERRANEAN

Baleares, Spain

4996

1500

0.30

121

0.02

Corse, France

8748

2781

0.32

240

0.03

Sardinia, Italy

24090

2407

0.10

238

0.01

Sicily, Italy

25708

2939

0.11

407

0.02

Kriti

8258

1971

0.24

189

0.02

Cyprus

9253

1940

0.21

121

0.01

Table 1 . Comparison between Oeeanie islands and the main Mediterranean islands in number of speeies and endemies in

relation with their area.

Island or archipelago

% AUens in the flora

Area Km^

Altitudinal range

No. Total taxa / area

CORSE

16.4

8748

2710

0.318

SARDINIA

6.4

24090

1834

0.100

CYPRUS

10.5

9253

1953

0.230

SICILY

8.4

25708

3323

0.125

KRITI

7.3

8258

2456

0.239

BALEARES

15.1

4996

1450

0.346

Table 2. Comparison between the pereentage of aliens in the flora and geographieal features of the major Mediterranean islands.

340

G. Domina, P. Marino, V. Spadaro & F.M. Raimondo

changed, of the plant cover it was bearing at the

time when it beeame insular (Greuter, 2001).

For the same reason the endemies shared among

the floras of the main Mediterranean islands are

very poor. Sicily shares 65% of its flora with Sar-

dinia and Corsiea but only few endemies, e.g. Carex

panormitana Guss., Dianthus arrostii C. Presl and

Genista aetnensis (Raf. ex Biv.) DC.

Figure 2. Occurrence of Limonium species in the major islands of the Mediterranean, it is evident a decreasing gradient

from west to east. Figure 3. Floristic affinities between the major Mediterranean islands and the relevant inland territories.

Vascular fJoro evolution in the major Mediterranean islands

341

The Sardinian flora is better represented in Cor-

siea, where 84% of its flora oeeurs (Boeehieri et al.,

2006); strong similarities with the Balearie Islands,

as well as with eoastal Spain and Franee (Fig. 3) are

also elear. This pattern is expeeted beeause the is-

lands have a similar geologieal history and they,

partieularly Sardinia, were eloser to the Balearie Is-

lands and the Pyrenees in the Mioeene, e.g. So/ei-

rolia soleirolii (Req.) Dandy, Urtieaeeae.

Corsiea shows the strongest floristie affinity

with Sardinia, when we eompared endemies shared

between W Mediterranean islands. A partieularly

eonspieuous affinity also exists with the adjaeent

mainland of Franee (inel. Massif Central), NE

Spain and W Italy (e.g. Bupleurum stellatum L.

Apiaeeae). Crete shows the strongest floristie affi-

nity with Greeee (e.g. Potentilla speciosa Willd.

Rosaeeae) but few similarities with Anatolia and E

Mediterranean.

Cyprus shows the strongest floristie affinity with

Anatolia (e.g. Onosma mite Boiss. et Heldr., Bora-

ginaeeae) and few similarities with East Mediterra-

nean and North Afriean territories.

HUMAN IMPACT

In the last eenturies man has beeome the most

important veetor for neweomers surpassing the

other animals, birds, drift and wind; in this way for-

mer isolation barriers were broken (Domina &

Mazzola, 2011). By the trade of goods, plants and

animals Man has, aeeidentally or purposely, intro-

dueed large quantities of alien propagules into sui-

tably eleared, new manmade habitats, or at least

into a degraded, unbalaneed environment. This

eould have eliminated vulnerable old relie speeies.

But if these losses did happen, it must have been

long before reliable botanieal reeords were made

(Greuter, 2001). Looking at the damage that al-

ready oeeurred in a flora ean be assessed its vulne-

rability. Rather than by new speeies introduetion

most damages to floras are to be imputed to habitat

loss; reeent examples are the endemies Lysimachia

minoricensis J.J.Rodr. (Primulaeeae) extinet from

Menorea and Adenostyles nebrodensis (Wagenitz

et I. Mull.) Greuter (Asteraeeae) praetieally extinet

from Sieily.

Vulnerability of a flora is evideneed by its ope-

ning to aliens. New introduetions of plant diaspores

is diffieult in the Mediterranean beeause of niehe

pre-emption of a resident flora that has long been

well adapted to existing habitat eonditions (Rune-

mark, 1969). This is overeome when habitats have

been pre-eleaned by human aetivities. So sinee an-

eient times the Mediterranean islands have been af-

feeted by more intense human pressure, therefore

they host higher pereentage of alien than the main-

land. The most important relations in large islands

have been observed between aliens oeeurrenee and

floristie density. In faet the aliens oeeurrenee in

eaeh island is related to extension and variety of ha-

bitats aetually suseeptible to invasions (Domina &

Mazzola, 2011).

WHERE ARE WE GOING

Large Mediterranean islands eonserve mid-

Tertiary floras rieh in endemies. These plants are

well fitted to their natural environment and seve-

ral of them, in spite of their rarity, eontinue to sur-

vive if their habitats are not manipulated.

Touristie and urban development are seriously da-

maging unique biota and their plants. Researeh on

Mediterranean plant diversity, biology, demogra-

phy and distribution is to be intensified to better

know how we ean proteet plant admitting a wise

human development aware of natural resourees.

Awareness of eonservation problems is arising

throughout the Mediterranean, and if eoordinated

efforts are done, we are still in time to prevent

heavy losses and to leave this rieh and valuable

plant heritage to future generations.

ACKNOWLEDGEMENTS

Funding by Universita degli Studi di Palermo

(ex 60 %) is gratefully aeknowledged.

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the Voyage of H.M.S. Beagle Round the World, under

the Command of Capt. FitzRoy, R.N, 2nd ed. John

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tbase - the information resource for Euro-Mediterra-

nean plant diversity. Published on the Internet

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grafiche sulla presenza di specie aliene nella flora

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269-276.

Greuter W., 1979. The origin and evolution of island flo-

ras as exemplified by the Aegean archipelago. In:

Bramwell D. (ed.). Plants and island. London & New

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Greuter W., 2001. Diversity of Mediterranean island flo-

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Mediterranean Basin: Setting global conservation

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von Diasporen kiistenbewohnender Pflanzen der Sii-

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Quezel P. & Medail F., 2003. Ecologie et biogeographie

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90-129.

Biodiversity Journal, 2012, 3 (4): 343-362

The vascular flora of Algerian andTunisian small islands: if not

biodiversity hotspots, at least biodiversity hotchpotchs?

Errol Vela' & Daniel Pavon^

'Universite Montpellier-2, U.M.R. AMAP (Botanique et Bioinformatique de I’Arehiteeture des Plantes), TA A51/PS2 - 34398

Montpellier eedex 5, Franee; e-mail: errol.vela@eirad.fr

^Aix -Marseille Universite, U.M.R. IMBE 7263 (Institut Mediterraneen de Biodiversite et d’Eeologie, UMR), Batiment Villemin,

Europole de I'Arbois - BP 80, 13545 Aix-en-Provenee eedex 04, Franee; e-mail: daniel.pavon@imbe.fr

ABSTRACT Algerian and Tunisian coasts host more than one hundred small islands and islets, but are

still poorly known. We have compiled recently published and unpublished data from “PIM

initiative” (Mediterranean small island initiative) and other kind of expeditions. For each

small island or archipelago we seek to establish the membership to or relationship with the

regional hotspots of the Mediterranean basin and the important plant areas (IPA) of Algeria

and Tunisia, thanks to species-area relationships and biogeographical analyses. Nowadays,

25 small islands are considered as botanically well-known and can be analysed. Species-area

relationship follows a classical linear regression model while some islands are less rich than

predicted and other ones are more rich. These richest islands can be assessed as IPA following

criterion B. Some of them have been yet assessed as IPA following criterion A, especially

because of presence of local or regional endemism. Each main archipelago shows biogeo-

graphical links not only with neighbour continental coasts, but also with northern coasts or

big islands from the western Mediterranean, especially the Tyrrhenian complex. “Grand Ca-

vallo” and “Petit Cavallo” islands are highlighted here as the 23rd IPA from Algeria. As bio-

diversity hotchpotch, each small island or archipelago should play a significant role in the

conservation programs although some of them are still unexplored and a deeper taxonomical

knowledge is necessary in the north- African context.

KEY WORDS Important Plant Areas; endemism; PIM initiative; protected areas; species-area relationship.

Received 11.05.2012; accepted 28.11.2012; printed 30.12.2012

Proceedings of the U‘ International Congress “Insularity and Biodiversity”, May 11*-13*, 2012 - Palermo (Italy)

INTRODUCTION

Mediterranean Basin is famous for its thousands

of islands (more than 15,000), most of them small

islands or roeky islets. Eastern Mediterranean

shows more islands and islets than the Western side,

nevertheless the latter ineludes the biggest islands

like Sieily, Sardinia and Corsiea. Following PIM iti-

niative (http://www.initiative-pim.org/) around 1100

small islands are eonsidered in the Western Medi-

terranean, and 1 67 of them are loeated in front of

Moroeean, Algerian or Tunisian eoasts, islands be-

yond Gibraltar strait and beyond the Sieilian ehan-

nel exeluded (Fig. 1). Around one fifth are isolated

islands, the others are grouped in arehipelago eontai-

ning from 2 to 16 islands. All are “small islands”, ge-

nerally uninhabited, defined here as less than 1000

ha and with at least one vaseular plant (i.e. not totally

submerged by waves or tide). It ineludes “very small

islands”, defined by Panitsa et al. (2006) as islands

with area less than 50 heetares. Most of these small

islands from Northern Affiean eoasts of the Western

344

Errol Vela & Daniel Pavon

Figure 1. Screenshot centered on Algerian and Tunisan coasts after the PIM database (www.initiative-pim.org). Main ar-

chipelagos are circles spotted in red. Number of islets aggregates included within others circles are written in the circle.

Mediterranean belong to regional biodiversity hot-

spots like Betieo-Rifean and Kabylies-Numidia-

Kroumiria eomplexes (Medail & Quezel, 1997; Vela

& Benhouhou, 2007). Their biogeographieal position,

often near the putative mean glaeial reftigia areas

(Medail & Diadema, 2009), made them probably

good biodiversity refuges during glaeial ages, even

more beeause their area was eonsiderably inereased

by a lower sea level. Today yet they appear as site fa-

vourable for miero-speeiation (Greater, 1 995) and re-

fuges for some rare speeies and/or vulnerable biota.

But why do they give us that impression? At

least we ean say beeause small islands are unfavou-

rable for eultivated fields, urbanisation, industriali-

sation and other modem intensifieations, they look

as refuges for wilderness. Are they biodiversity hot-

spots (or part of them)? Are they key biodiversity

areas (at least for plants)? Are they useful for eon-

servation strategies? Answering this requests is dif-

fieult beeause of low knowledge of natural history

for most of them, indueing diffieulties to understand

their biogeography (Triantis et ah, 2008a), neglee-

ting assessments of areas with eonservation priori-

ties (Nikolic et ah, 2008) or erroneous eonelusion

following seientifie field surveys (Mifsud, 2011).

In the aim of highlighting the natural history of

Algerian-Tunisian small islands and their eonserva-

tion values, we analyse here a eompilation of pu-

blished and unpublished data mainly eolleeted by

authors and eollaborators during 2006-2012 PIM

expeditions eompleted by few eomplementary data.

MATERIALS AND METHODS

Historical knowledge

Historieal floristie investigations were very poor

until the 1990s, as only six islands or arehipelagos

were more or less investigated:

Habibas (NW- Algeria): Maire, Wilezek & Faure

in 1934 (ef. Maire & Wilezek, 1936).

Galite (NW-Tunisia): Chabrolin in 1931 (ef.

Chabrolin, 1933), Boeehieri & Mossa in 1983 (ef.

Boeehieri & Mossa, 1985).

Cani, Pilau, Plane (N-Tunisia): Pottier-Alapetite

before her death in 1971 and most probably before

Tunisian independenee in 1956 (ef Pottier-Alape-

tite, 1979-1981).

Zembra (NE-Tunisia): Doumet- Adamson in

1884 (ef. Doumet- Adamson, 1888); Letoumeux in

1887 (ef. Bonnet & Barratte, 1896), Barratte & Cos-

son in 1888 (ef. Bonnet & Barratte, 1896), Labbe

& Pottier-Alapetite in 1953 (ef. Labbe, 1954).

The PIM Initiative

The Mediterranean small islands Initiative (PIM

initiative; www.initiative-pim.org) wants to better

promote and aet for the eonerete management of

Mediterranean islands.

It partieipates in knowledge and proteetion of

these miero-island areas through implementation of

The vascular flora of Algerian and Tunisian small islands: if not biodiversity hotspots, at least biodiversity hotchpotchs? 345

Figure 2. “Petite Habiba” and unexplored satellite islets viewed from the main island “Grande Habiba”. Figure 3. Deser-

ted village in the sheltered bay and still inhabited semaphore on the top of the main island.

346

Errol Vela & Daniel Pavon

Figure 4. “Fauchelle” and “Galiton” in the baekground, viewed from the main island “La Galite”. Figure 5. “La Galite” in

the baekground, “Gallina” and “Pollastro” unexplored islets in the foreground, viewed from the top of “Gallo”.

The vascular flora of Algerian and Tunisian small islands: if not biodiversity hotspots, at least biodiversity hotchpotchs? 347

Figure 6. N-W slope of Zembra island with its ealeareous “Capo Grosso” in the baekground and “L’Antoreho” roeky islet on

the left. Figure 7. The unexplored “Cathedrale” islet viewed from Zembra main island. Figure 8. Global view of Zembra main

island with the “Cathedrale” small island on the left. Figure 9. “Les Pisans” island viewed from the Boulimate’s beaeh, Gou-

raya’s National Park, Bejaia (N Algeria). Figure 10. “Grand Cavallo” island viewed from El Aouana eity, Jijel (N Algeria).

348

Errol Vela & Daniel Pavon

concrete aetions on the field, promoting exehange

of knowledge and skills between managers, natura-

lists and seientists from the whole Mediterranean.

Sinee 2006, through the program named “Terra Co-

gnita” (i.e. to better know), the PIM team organizes

field expedition involving many experts. Thus, with

help of Algerian CNL (Commissariat National du

Littoral; http://www.matev.gov.dz/pdf/littoral/orga-

nigramme_du_enl.pdf) and Tunisian ARAL (Agenee

de Proteetion et dAmenagement du Littoral;

http://www.apal.nat.tn/), the main North- Afiiean is-

lands have been investigated:

*Reehgoun (NW-Algeria): 2006/05.

**Habibas arehipelago (NW-Algeria): 2006/05,

2007/10 (2 islands).

*Serijina (NE- Algeria) : 2008/05.

**Galite arehipelago (NW-Tunisia): 2008/05,

2008/11, 2009/07 (4 islands).

**Cani arehipelago (N-Tunisia): 2009/08 (3

islands).

Pilau (N-Tunisia): 2007/05.

Plane (N-Tunisia): 2007/05.

Zembra arehipelago (NE-Tunisia): 2007/06,

2008/05, 2009/07, 2012/06 (2 islands).

*never explored before !

**some islets never explored. . .

Complementary data

Thanks to reeent works on Jijel’s and Bejaia’s

eoasts in NE- Algeria (Bougaham, 2008; Hanifi-

Benhamiehe et al., 2011; Benhamiehe-Hanifi&

Moulai, 2012; Vela et al., 2012a), flora of seven

small islands from Kabylian eoasts is known for the

first time:

*E1 Eueh (NE-Alg.) : 2010/03-2010/06.

*Bejaia islands (NE-Algeria): 2010/03-2010/06;

2011/06-2011/07 (3 islands).

*E1 Aouana islands (NE-Algeria): 2009/03;

2009/06 (3 islands).

Finally, unpublished data are available for three

Algerian islet :

Vivier islet at Cap de Garde (NE-Algeria):

2002/05 ; 2003/04.

Cale Genoise islets at Cap Tends (NW-Algeria):

2012/09.

Thus, we’ve got now field inventories for the

Algerian-Tunisian main islands and arehipelago and

several of the numerous smaller ones.

IPA approach

Plantlife International (2004) proposed three

main eriteria for identifying important plant areas:

Criterion A (signifieant populations of threatened

speeies); Criterion B (exeeptionally rieh flora in

the eontext); Criterion C (outstanding example of

a habitat type of eonservation importanee). Be-

eause of missing information on habitat (types of

habitats, threatened habitats) and on global plant

riehness (no speeifie data on eommon speeies) in

Algeria and Tunisia, additionally to the great num-

ber of restrieted-range endemie speeies within the

whole Mediterranean hotspot, methodology to de-

teet Important Plant Areas were partially modified

by Yahi et al. (2012).

IPA were seleeted by presenee and/or riehness

of “trigger speeies” as: globally threatened speeies

[based on partial list from lUCN red lists as Walter

& Gillett (1998) and Gareia et al. (2010)], site-re-

strieted endemie speeies (< 100 km^), high number

of restrieted-range speeies (< 5000 km^) and natio-

nally threatened/rare speeies [following national

floras as Battandier (1888), Battandier & Trabut

(1895), Cuenod et al. (1954), Quezel & Santa

(1962-1963), Pettier- Alapetite (1979-1981) and

eurrent unpublished field data] Northern Algerian

and northern Tunisian territories, ineluding small

arehipelagos, were respeetively investigated by

Yahi & Benhouhou (2011), Yahi et al. (2012) and

Ghrabi Gammar (2011).

Generally, habitat diversity inerease with area,

then among sites with similar areas, speeies ric hn ess

inereases with habitat diversity (Kallimanis et al.,

2008), it is why the lattest one is frequently used as

extrapolation of the first one. On islands, speeies

number is a fiinetion of area and number of habitats

(Triantis et al., 2006), except below a threshold in-

dueing a so-ealled “small island effeef ’ (Lomolino

& Weiser, 2001; Triantis et al., 2006). Below this

small island effeet, direet efifeets of area are elimi-

nated and limited to indireet effeets as the role of

area on habitat diversity then the role of habitat di-

versity on speeies riehness (Triantis et al., 2006).

Finally, for very small islands (Panitsa et al.,

2006), plant speeifie riehness is related with other

parameters than area, like elevation (indueing itself

an inereasing of possible habitat diversity?), pre-

senee or absenee or grazing, and non-standard fae-

tors or stoehastie effeets.

The vascular flora of Algerian and Tunisian small islands: if not biodiversity hotspots, at least biodiversity hotchpotchs? 349

Nevertheless, as area remains the strongest de-

terminant of island speeies numbers (Kreft et ah,

2008), speeies-area relationships ean be explored in

order to show “exeeptionally rich flora in the con-

text” as an IPA indicator for islands and archipelagos

(“criterion B” sensu Plantlife International 2004).

Biogeographical analysis

Taxa present on each island and archipelago are

assessed from a biogeographical point of view. Re-

gional endemism and other remarkable taxa (e.g.

small islands specialised taxa) shared with nei-

ghbour areas are highlighted in order to identify

biogeographical links among small islands and bet-

ween small islands and continents.

RESULTS

Including PIM expeditions data, recent biblio-

graphy and unpublished data, we could say that

flora is “well known” for 25 small islands or islets,

14 in Algeria and 1 1 in Tunisia (Table 1).

Species-area relationship

From Table 1 data, we can draw a species-area

relationship for Algerian and Tunisian small islands

(Fig. 11). As richness is a logarithmic function of

area, richness is then a proportional function of log

(area). A linear regression limited to these small is-

lands (area between 0.4 and 732 ha) is a good ap-

proach to detect “rich” islands (richness higher than

prediction) and “poor” islands (richness lower than

prediction).

The best linear regression model (predicted

specific richness = 70.025 log area(ha) + 15.565)

obtain a R2 coefficient of determination higher as

0.6. We can also size small islands following the

residual between observed and predicted species

richness (Table 2). We can see fourteen islands

with positive residuals, i.e. with species richness

higher than predicted and thirteen islands with ne-

gative residuals, i.e. with species richness lower

than predicted. Among very “poor” islands, there

are very small islands as Plane and Grand-Cani (6

and 4 ha) and some bigger ones as Galiton or Re-

chgoun (30 and 26 ha). Among “rich” islands. La

Galite is the richest and also the biggest one (732 ha).

species-area relationship

400 —^

050 —

300

loglOarea (ha)

Figure 1 1 . Species-area relationship for plants in Algerian and Tunisian small islands, following linear regression model.

350

Errol Vela & Daniel Pavon

Island (wilaya, country)

Area (ha)

Species

source

Reehgoun (Ain Temouehent, DZ)

PIM (E. Vela, unpubl.)

Grande Habiba (Oran, DZ)

PIM (Delauge & Vela, 2007); E.Vela & A.

Saatkamp, unpubl.

Petite Habiba (Oran, DZ)

5.5

PIM (Delauge & Vela, 2007)

Gale Genoise W (Chief, DZ)

0.4

D. Amari & E. Vela, unpubl.

Gale Genoise E (Chief, DZ)

0.5

D. Amari & E. Vela, unpubl.

El Eueh (Bejaia, DZ)

1.5

Benhamiehe-Hanifi& Moulai, 2012

L'Ail (Bejaia, DZ)

0.4

Vela et al., 2012a

Pisans (Bejaia, DZ)

Benhamiehe-Hanifi& Moulai, 2012

Sahel (Bejaia, DZ)

0.5

Benhamiehe-Hanifi& Moulai, 2012

Grand Cavallo (Jijel, DZ)

Benhamiehe-Hanifi& Moulai, 2012

El Aouana (Jijel, DZ)

0.4

Benhamiehe-Hanifi& Moulai, 2012

Petit Cavallo (Jijel, DZ)

101

Benhamiehe-Hanifi& Moulai, 2012

Serigina (Skikda, DZ)

1.6

PIM (Vela, 2008)

Vivier (Annaba, DZ)

0.7

E. Vela & G. de Belair, unpubl.

Galite (Bizerte, TN)

732

340

PIM (D. Pavon et al., unpubl.)

Galiton (Bizerte, TN)

PIM (D. Pavon & M. Murraeeiole, unpubl.)

Fauehelle (Bizerte, TN)

13.6

PIM (D. Pavon & M. Murraeeiole, unpubl.)

Gallo (Bizerte, TN)

8.9

PIM (D. Pavon, unpubl.)

Grand Cani (Bizerte, TN)

PIM (M. Delaugerre et al., unpubl.)

Cani NE (Bizerte, TN)

PIM (M. Delaugerre et al., unpubl.)

Cani SW (Bizerte, TN)

PIM (M. Delaugerre et al., unpubl.)

Pilau (Bizerte, TN)

PIM (E. Vela, unpubl.)

Plane (Bizerte, TN)

PIM (E. Vela, unpubl.)

Zembra (Nabeul, TN)

389

255

PIM (E. Vela et al., unpubl.)

Zembretta (Nabeul, TN)

46 // 45

PIM (Serrano, 2008; Domina & Mokni, 2012)

Table 1. List of Algerian and Tunisian islands whieh flora is here eonsidered “well known”.

The vascular flora of Algerian and Tunisian small islands: if not biodiversity hotspots, at least biodiversity hotchpotchs? 351

islet

loglO(area)

observed

predicted

residual

Galite

2.8645

340

216.2

123.8

Zembra

2.5899

255

196.9

58.1

Petit Cavallo

0.4771

101

49.0

52.0

Sahel

-0.3010

-5.5

49.5

Pisans

15,6

36.4

El Aouana

-0.3979

-12.3

35.3

L'Ail

-0.3979

-12.3

33.3

Grand Cavallo

0.4771

49.0

33.0

El Euch

0.1760

27.9

32.1

Genoise-W

-0.3979

-12.3

19.3

Genoise-E

-0.3010

-5.5

12.5

Vivier

-0.1549

4.7

10.3

Serigina

0.2041

29.9

2.1

Cani-SW

15.6

-7.6

Cani-NE

15.6

-9.6

Zembretta

0.6989

64.5

-18.5

Grande Habiba

1.5440

123.7

-25.7

Pilau

0.6020

57.7

-32.7

Petite Habiba

0.7403

67.4

-43.4

Grand-Cani

0.6020

57.7

-46.7

Fauchelle

1.1335

94.9

-51.9

Plane

0.7781

70.1

-59.1

Rechgoun

1.4149

114.6

-60.6

Gallo

0.9493

82.0

-69.0

Galiton

1.4771

119

-73

Table 2. Algerian and Tunisian islands following overage or deficit in species richness compared to standard predicted by

linear regression.

352

Errol Vela & Daniel Pavon

Three of them, Sahel, Cavallo-S and L’Ail (the

smallest ones, 0.5 ha or less) are eonsidered rieh only

beeause their predieted riehness is negative as their

Log (area) is less than the origin value of the regres-

sion line. Exeluding this artefaet and the moderate

rieh Vivier and Serigina islands, we ean eonsider six

islands with a signifieantly high speeifie riehness

(Galite, Petit Cavallo, Zembra, Pisans, Grand Ca-

vallo, El Eueh).

Biogeographical affinities

Rechgoun and Habibas Archipelago (NW-

Algeria)

The Habibas Arehipelago has got a eontinental

origin but its roeks are mainly from voleanie origin,

it is without freshwater resourees and is almost

uninhabited. During the first half of the 20th een-

tury, a small village was built by fishermen on the

main island, now abandoned and in ruins exeept for

equipment storage. There is a lighthouse on the top

(105 m) of the main island (35 ha), eontinuously in-

habited by one keeper. Historieal flora is known

thanks to a previous study by Maire & Wilekzek

(1936). Vaseular flora is not very rieh (around 100

speeies for the largest island) regarding to the area

but remained relatively stable during 70 years in

speeies riehness and eomposition (Vela, 2013). It

presents numerous regional endemisms shared with

Afi-iean and/or European eontinent, eharaeterising

the Alboran sea biogeographieal area (Table 3).

Moreover, the “small island speeialisf ’ and We-

stern-Mediterranean Stachys brachyclada De Noe

is present on the main island while it is absent on

the small island and on the neighboring Reehgoun

island (Vela, 2013).

The island of Reehgoun eontain similar flora

but signifieantly poorer. Only three of the regional

endemies are present but rare: Anthemis chrysan-

tha, Fumaria munbyi and Sonchus tenerrimus

subsp. amicus (Vela, 2013).

Kabylian-Numidian small islands (N-Algeria)

In this area, small islands are very diverse, some

of them ealeareous (Pilot a Pail, ilot Sahel), the

other ones silieeous. Vaseular flora is sometimes re-

latively poor (Vivier, Serigina) but more often very

rieh (Petit Cavallo, Pisans, ete.). Only one regional

endemism was inventoried on ilot Sahel, Pancra-

tium foetidum Pomel var. saldense Batt., whieh is a

Kabylian rupieolous vieariant of the W-Algerian-

Moroeean P. foetidum yw. foetidum. Furthermore,

the “small island speeialisf’ and Central-Mediter-

ranean eommutatum Guss. is present on one

island, L’ilot a PAil at Boulimate (where it was first

mentioned for Algeria, ef. Vela et aP, 2012a).

La Galite Archipelago (NW-Tunisia)

This archipelago is very far from the continent

(40 km). It includes a relatively big island more or

less inhabited (La Galite, 732 ha) and other smaller

ones whose area is varying from 8 to 30 ha. The

main island, mainly granitic, hosts several inter-

mittent seeps. Vascular flora appears to be rich but

still misunknown for the largest island because of

the lack of intensive researches (a cumulative of

340 species including historical and actual field ob-

servations).

It presents local and regional endemisms with

Corso-Sardinian affinities and a lot of taxa exclu-

sive for Tunisia indeed Africa (Table 4). Belleva-

lia galitensis is considered like a local endemism

(Bocchieri & Mossa, 1991) and the study of the

local Limonium “cf. intricatum’’'’ will certainly

confirm the presence of another endemic for this

archipelago. Moreover, the “small island specia-

lisf’ and Central-Mediterranean eommuta-

tum Guss. is present on La Fauchelle, a peripheric

islet from the Galite Archipelago (Pavon & Vela,

2011). At least five taxa can be considered sho-

wing Corso-Sardinian affinities (during our sur-

veys we have not seen the two last taxa mentioned

by Bocchieri & Mossa, 1985):

•Limonium “cf. intrieatum’’' = sp. nov. ? (belon-

ging to L. articulatiim aggr.). This interesting and

understudied taxa was first mentioned on La Galite

by Bocchieri & Mossa (1985, sub 'Fimonium sp.”)

then partially highlighted by Pavon & Vela (2011).

Its strong affinity or else its identity with L. intri-

eatum described from the Bizerte coast by Brullo

& Erben (1989) clearly attests its belonging to the

L. artieulatum aggregate of micro-endemism from

Corso-Sardinian area.

•Brassiea insularis s.s. (non 5. atlantiea): in its

stricto sensu, this Corso-Sardinian endemism is sha-

red with African continent on Edough peninsula

The vascular flora of Algerian and Tunisian small islands: if not biodiversity hotspots, at least biodiversity hotchpotchs? 353

Taxa endemic from Alboran sea (s.l.)

Presence in

Europe

Presence in

NW-Africa

Abundance on

Habibas

Anthemis chrysantha J. Gay

SE-Spain

NW-Algeria

Arenaria cerastioides (Crantz) Maire van oranensis Batt.

NE-Mor./NW-Alg.

Asteriscus maritimus L. subsp. sericeus (Maire et Wilczek) Vela

NW-Algeria

Brassica spinescens Pomel

NW-Algeria

Fumaria munbyi Boiss. & Rent.

E-Spain

NW-Algeria

Lobularia maritima (L.) Desv. subsp. columbretensis R. Fern

E-Spain

Rostraria balansae (Coss.&Dur.) Holub

NE-Mor./NW-Alg.

Silene pseudoatocion Desf van oranensis Batt.

NE-Mor./NW-Alg.

Sonchus tenerrimus subsp. amicus (Faure, Maire & Wilczek) Vela

Chafarinas? (sub S. bour-

geaui Schultz Bip.?)

Spergularia pycnorrhiza Batt.

NW-Algeria

Table 3. Taxa endemic from the Alboran area present on Habibas archipelago (Vela 2013).

Taxa exclusive for Tunisia

(*indeed Africa ; **exclusive endemism)

Biogeography

Rarity on Ga-

lite archipelago

Source

Asplenium marinum L.

Sub-Atlant.

Muracciole et al. (2010)

Asplenium obovatum Viv. subsp. obovatum

Sub-Atlant.

Pavon& Vela (2011)

**Bellevalia galitensis Bocchieri & Mossa

N-Tunisia

Bocchieri & Mossa (1991);

Pavon & Vela (2011)

*Bituminaria morisiana (Pignatti & Metlesics)

Greuter

Sardinia

RR?

Bocchieri & Mossa (1991)

Brassica insularis Moris [excl. B. atlantica

(Coss.) G.E. Schulz]

Corsica, Sardinia,

NE- Algeria

Pavon & Vela (2011)

Cheilanthes maderensis Lowe

W-Medit . -Macarones .

Vela (unpubl.)

Diplotaxis viminea (L.) DC.

Europ.-Medit.

Pavon & Vela (2011)

Limonium cf. intricatum Brullo & Erben

[**sp. nov. ?]

N-Tunisia

Pavon & Vela (2011)

Ononis minutissima L.

Medit.

Pavon & Vela (2011)

*Serapias nurrica Corrias

Tyrrhen. islands

Vela et al. (2012c)

Table 4. Taxa exclusive for Tunisia registered on La Galite Archipelago.

354

Errol Vela & Daniel Pavon

(Yahi et al., 2012) and the La Galite Archipelago

(Chabrolin 1933; Pavon & Vela, 2011).

•Serapias nurrica: this Tyrrhenian taxa known

from Corsica, Sardinia, Menorca, Sicilia and Cala-

bria was recently discovered on La Calite island by

R. Ouni (Vela et al., 2012c).

•Hyoseris lucida L. subsp. taurina (Martinoli)

Peruzzi & Vangelisti is a poorly known Tyrrhenian

taxa that grows in coastal cliffs of Peninsular Ita-

lia, Sardinia, Sicily, Malta and La Calite (Boc-

chieri & Mossa, 1985; Brullo et al., 1997; Peruzzi &

Vangelisti, 2010) but also on NW-Tunisian and NE-

Algerian continental coasts (VCa, unpubl.).

•Bituminaria morisiana is a doubtful and ne-

glected taxa described from Sardinia and mentioned

on La Calite by Bocchieri & Mossa (1985) which

doesn’t appear in the Euro+Med database

(http://ww2.bgbm.org/EuroPlusMed/), in the Afri-

can Plant Database (Dobignard & Chatelain, 2010-

13) nor in the last Tunisian checklist (Le Floc’h et

al., 2010).

Bizerte’s coasts small islands (N-Tunisia)

In this area, small islands are relatively near

(Pilau, Plane) or very distant from the continent

(Cani Archipelago). Vascular flora is poor (between

6 and 25 species) and is exempted from regional en-

demism. Nevertheless, the “small island specialist”

and Central-Mediterranean Allium commutatum

Cuss, is present on two islands of the Cani Archi-

pelago and on Pilau island where it was first men-

tioned for Tunisia (cf. Le Floc’h et al., 2010).

Zembra Archipelago (NE-Tunisia)

This archipelago’s comprising a relatively large

island once inhabited (Zembra, 389 ha), a very

small one (Zembretta, 5 ha) and some rocky islets.

Floristic richness is high on the main island Zembra

(about 255 species). This archipelago hosts at least

one local endemism and shares eight regional en-

demism with Sicily and adjacent areas or with con-

tinental areas from Punic domain within or near

Cap Bon (Table 5). Furthermore, two Central-Ea-

stern Mediterranean species were recorded in

Zembra where they are exclusive for Tunisia, Sar-

copoterium spinosum (L.) Spach (Doumet-Adan-

son, 1888, Labbe, 1954) and Solenopsis minuta

(L.) C.Presl (Labbe 1954 sub "‘'Laurentia miche-

lif\ Domina & El Mokni, 2012).

Finally, the “small island specialist” and Cen-

tral-Mediterranean ^1 //mm commutatum Cuss, has

been recently discovered on the very small island

of Zembretta (cf. Domina & El Mokni, 2012).

Regional endemism

(*exclusive for Tunisia; **exclusive endemism)

Biogeography

Rarity on

Zembra arch.

Source

Allium cf lehmannii Lojac.

(= A. obtusiflorum DC.?)

Sicily, S-Italy? (Greece?)

Pavon & Vela (2011); Do-

mina & El Mokni (2012)

Bellevalia cf dolichophylla Brullo & Miniss.

NE-Tunisia

Domina & El Mokni

(2012)

Brassica atlantica (Coss.) O.E. Schulz

(non B. insularis Moris)

NE-Tunisia

Cosson (1883-1887)

Dianthus rupicola Biv. subsp. hermaeensis

(Coss.) 0. Bolos & Vigo

NE-Tunisia (sp: Sicily s.l.

and Mallorca)

Cosson (1882-1890);

Doumet-Adanson (1888)

*Filago lojaconoi (Brullo) Greuter

Sicily s.l. (Linosa,

Pantelleria)

Domina & El Mokni

(2012)

*Iberis semperflorens L.

S-Italy, Sicily

Cosson (1883-1887);

Doumet-Adanson (1888)

**Limonium zembrae Pignatti

NE-Tunisia

Pignattii (1982)

Sixalix farinosa (Cosson) Greuter & Burdet

N-Tunisia + NE -Algeria

Doumet-Adanson (1888);

Vela et al. (2012b)

Table 5. Narrow and regional endemism present on Zembra main island.

The vascular flora of Algerian and Tunisian small islands: if not biodiversity hotspots, at least biodiversity hotchpotchs? 355

Hotspot areas

At global level (Myers et al., 2000; Mittermeier

et al., 2004), biodiversity hotspots are delimited by

exeeptional botanieal and zoologieal riehness and

endemism rate on large areas as major mountain

ranges (Himalayas, Caueasus, Andes, ete.), major

tropieal islands and arehipelagos (Madagasear,

Sundaland, New-Caledonia, ete.) or major Medi-

terranean biomes (California, Cape region, Medi-

terranean Basin, ete.).

As part of the Mediterranean basin, small is-

lands like biggest ones are ineluding within the glo-

bal Mediterranean hotspot. At regional level

(Medail & Quezel, 1997; Vela & Benhouhou,

2007), plant biodiversity hotspot within the Medi-

terranean basin are delimited by semi-empirie data

as speeifie riehness for 10,000 km^ and

endemism/subendemism rate within areas as high

mountains (Atlas, Lebanon...), eollision ehains

(Betieo-Rifean are, Taurus, Kabylies. . .), big islands

(Tyrrhenian islands, Creta, Cyprus. . .) or oeeanie ar-

ehipelagos (Canaries and Madeira). Themselves,

small islands are not suffieient territories («

10,000 km^) to host high speeifie ric hn ess (> 2000

sp.) and endemism rate (> 10 %). But most of them

are loeated near to the eoast and eould be ineluded

within regional hotspots:

- Peripheral islets satellites from Corsiea, Sardi-

nia, Sieily and the Balearie ean be ineluded within

The Tyrrhenian Island hotspot area;

- Small islands near northern and southern eo-

asts of the Alboran sea (like Habibas arehipelago in

NW- Algeria) ean be ineluded within the Betieo-Ri-

fean hotspot area;

- Small islands near the Kabylian and Numidian

eoasts of NE- Algeria and NW-Tunisia (like Beja-

ia’s, JijeTs, Skikda’s and Annaba’s small islands)

ean be ineluded within the Kabylian-Numidian-

Kroumirian hotspot area;

Figures 12-14. Species exclusive for Tunisia, as example of the hotchpotch phenomenon. Fig. 12. Sarcopoterium spinosum

on Zembra island (N-E Tunisia). Fig. 13. Cheilanthes maderensis on La Galite is “new” for Tunisia.Fig. 14. Ononis minu-

tissima on La Galite (N-W Tunisia).

356

Errol Vela & Daniel Pavon

Figures 15-17. Irreplaceability characterised by “small islands specialist” species. Figure 15. Fumaria munbyi on Rechgoun

island, Ain Temouchent (N-W Algeria). Figure 16. Stachys brachyclada on Flabibas main island, Oran (N-W Algeria).

Figure 17. Allium commutatum on “Hot a FAil” near Boulimate’s beach, Bejaia, (N Algeria).

The vascular flora of Algerian and Tunisian small islands: if not biodiversity hotspots, at least biodiversity hotchpotchs? 357

Figures 18-20. Irreplaceability characterised by micro-speciation of narrow endemism. Figures 18, 19. Limonium cf in-

tricatum (possibly not described species) on La Galite island (N-W Tunisia). Figure 20. Limonium zembrae on Zembra

(N-E Tunisia).

- All the numerous small islands of Dalmatian

eoasts are de faeto part of the putative hotspot area

suggested by Nikolic et al. (2008);

- Most of the numerous small islands of Gre-

eian territories eould be more or less ineluded wi-

thin the related hotspot areas as Peloponnese,

Crete and Taurus.

Even if Galite and Zembra Arehipelagos have

got biogeographieal affinities with Corso-Sardinian

and Sieilian areas respeetively (ef. supra), it seems

imprudent to inelude them empirieally within the

Tyrrhenian hotspot area without further investiga-

tions. In order to explore riehness and endemism at

a lower level than the hotspot one, IPA methodo-

logy will be helpful.

IPA assessment in Algeria and Tunisia

In Algeria, Yahi & Benhouhou (2011) and Yahi

et al. (2012) have assigned 22 IPA whose only one

is insular, the Habibas Arehipelago. But several

islets loeated near the eoast eould be ineluded de

faeto in some littoral IPA (Traras mountains, Oran’s

hills. Cap Tenes, Chenoua mountain, Gouraya na-

tional park, Taza national park, Edough peninsula

and El Kala national park).

As an example, among the islets with well-

known flora, the three islands of Bejaia “Hot a

I’Ail”, “He des Pisans” and “Hot Sahel” (Benhami-

ehe-Hanifi& Moulai, 2012; Vela et al., 2012a), be-

eause of their geographieal proximity and

eeologieal similarity with eontinental eoasts make

them as part of the Gouraya’s IPA:

- Boulimate’s “Hot a TAil” hosts the main Al-

gerian population of Allium commutatum, a rare,

fragmented and range-edge speeies not known el-

sewhere in this IPA;

- “Hot Sahel” hosts the site-restrieted endemism

358

Errol Vela & Daniel Pavon

Pancratium foetidum var. saldense, the restricted-

range Algerian endemism Sedum multiceps and the

Algerian-Tunisian endemism Sedum pubes cens.

- “lies des Pisans” is one of the six richest small

islands regarding to the species-area relation. This

exceptional rich flora in the context met the island

with the criterion B and is an IPA indicator.

Other islands recently studied like El Aouana is-

lands “Grand Cavallo” and “Petit Cavallo” (Bouga-

ham, 2008; Hanifi-Benhamiche et al., 2011;

Benhamiche-Hanifi& Moulai’, 2012) are remarkable

following the presence of the restricted-range Alge-

rian endemism Aristolochia longa subsp.fontanesii

and Genista numidiea. Furthermore, with a high

specific richness (respectively 101 and 82 plant spe-

cies for an area of only 3 hectares each one), among

the richest islands studied here, these archipelago

could be assigned as an autonomous IPA.

In Tunisia, Ghrabi Gammar (2011) has assigned

24 IPA whose two are insular, the Galite Archipe-

lago and the Zembra Archipelago. A third IPA (Sidi

Ali el-Mekki), restricted to the littoral, is de facto

including the interesting island “He Pilau” which

hosts one of the three Tunisian populations of Al-

lium commutatum (Pavon & Vela, 2011).

23 24

Figures 21-24. Biogeographical affinities of Zembra (N-E Tunisia) as revealed by shared endemism. Figs. 21-22. Brassica

atlantica (AB. insularis s.s.?), endemism shared with Cap Bon peninsula (N-E Tunisia). Fig. 22. Allium ef lehmanii, ende-

mism shared with Sieily. Fig. 23. Dianthus rupicola subsp. hermaensis, endemism shared with Cap Bon (subspeeies) or

Sieily and Mallorea (speeies).

The vascular flora of Algerian and Tunisian small islands: if not biodiversity hotspots, at least biodiversity hotchpotchs? 359

DISCUSSION

Contrarily to the biggest islands of the whole

Mediterranean, small islands or arehipelago are not

eonsidered as regional hotspot themselves, eertainly

due to their very low area insuffieient to allow the

in situ speeiation (Lomolino & Weiser, 2001) and

keep them poor in exelusive endemism (Triantis et

ah, 2008b). Nevertheless they are sharing regional

endemism with neighbour areas like big islands or

eontinent. In eontrast their ean host speeialised spe-

eies absent from eontinent beeause of higher eom-

petition or they ean share speeies with far eountries

whieh are not present on the neighbouring areas.

Finally, eaeh islet, island or arehipelago host an

original eombination of native and exotie flora and

ean be assimilate as biodiversity hotehpotehs. As an

example, Sieilian Strait is a barrier to dispersal and

gene flow whieh may be erossed by natural or an-

thropogenie long-distanee dispersal overseas, assi-

sted by sea-level oseillations during the Pleistoeene

(Lo Presti & Oberprieler, 201 1). In the previous stu-

dies, examples of “signifieant populations of threa-

tened speeies” were satisfying eriterion A for IPA

Figures 25-27. Biogeographical affinities of Habibas (N-W Algeria) as revealed by shared endemism. Figure 25. Anthe-

mis chrysantha, endemism shared with Mureian eoasts (S-E Spain). Figure 26. Asteriscus maritimus subsp. sericeus. Fi-

gure 27. Brassica spinescens (right), endemism shared with Oran’s eoasts (N-W Algeria).

360

Errol Vela & Daniel Pavon

assessment. But fortunately, eriterion B can be te-

sted here because of a good knowledge of the spe-

cific richness on these small sites. The main ones

archipelago as Zembra (main island ~ 400 ha, 4 is-

lands) and La Galite (main island ~ 700 ha, 6 is-

lands) have been easily assessed as important plant

areas (IPA) following criterion A (Radford et al.,

2011). For the smaller ones, like Habibas Archipe-

lago (main island 35 ha, 4 islands), number of spe-

cies is relatively low but can contain some local

rarities like regional endemism (i.e. “restricted-

range species”), range edge, or specialised habitat

species and permitted to asses following criterion

A(Yahi et al., 2012).

Because they are most often devoid of local en-

demism (i.e. “site-restricted endemic species”) and

depleted in regional endemism, other “very small

island” (less than 50 hectares, often even less) were

not recognized as IPA following criterion A.

Nevertheless, thanks to our richness data by is-

land, we can now size island based on species-area

residuals. It follows that specific richness is abnor-

mally high (residual >30 species) for six islands.

The two main islands (La Galite; Zembra) belon-

ging to the two main archipelagos has been yet as-

sessed as IPA following criterion A (Radford et al.,

2011), and can now be re-affirmed with criterion B.

One very small islands (Pisans) belonging to a coa-

stal archipelago is considered here naturally inclu-

ded within the continental Gouraya’s IPA. Two

other very small islands (Grand-Cavallo and Petit-

Cavallo) forming a coastal archipelago near El Ao-

uana city should be proposed following criterion B

as a new IPA, ie the 23rd of Algeria (cf. Yahi &

Benhouhou 2011; 2012). Continental areas facing

the archipelago (Cap Noir, Grand Phare...) are

known for their floristic richness as hosting the rare

Silene sedoides Poir. and Limonium aff. minutum

(L.) Kuntze (Quezel & Pons, 1954) and probably

belong to this same IPA.

Finally, the last very small island classified here

as very “rich” (El Euch) is located near the coast bet-

ween Bejaia and Azzefoun but we need additional

investigation, particularly on the continental coasts,

in order to assess it as a probable IPA (or not?).

Then, with better knowledge of the flora and ve-

getation of each island or islet, managers can make

priorities for conservation and protection against in-

sidious threats like exotic invasive plants, introdu-

ced animals, local extinction, touristic activities and

many others. Nevertheless, deeper taxonomic inve-

stigations will be useful to complete biodiversity

comprehension while field survey are needed to un-

derstand impact of environmental change on biodi-

versity and ecosystems services.

If not exactly hotspots, but at least hotchpotchs

and/or key biodiversity areas for plants, north Afri-

can small islands appear to be good refugia for cur-

rent biodiversity conservation. They are until now

relatively difficult to access for local population and

show often better conservation status than continent

touristic/urbanistic areas. Furthermore they can host

not necessarily local endemism but at least specia-

lised species intolerant to competition against the

continental flora (e.g. Allium commutatum, Fuma-

ria munbyi, Stachys brachyclada, etc.).

Nevertheless, legal protection status as National

Park or Protected Marine Areas appears to be useftill

to maintain this level of conservation and to develop

long term management plan.

ACKNOWLEDGMENTS

Authors would like to thanks the organizers of

expeditions on Algerian and Tunisian small islands

(Conservatoire de I’Espace Littoral et des Rivages

Lacustres, France; Commissariat National du Lit-

toral, Algerie; Agence de Protection et d’ Amenage-

ment du Littoral, Tunisie). Authors are very grateful

also to F. Liberto (Cefalu, Italy).

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Biodiversity Journal, 2012, 3 (4): 363-368

Geophytes and evolution in the Sicilian Archipelago

Pietro Mazzola* Rosario Schicchi & Sebastiano Ciccarello

Dipartimento di Scienze agrarie e forestall, Universita degli Studi di Palermo, via Arehirafi 38 - 90123 Palermo, Italy

* Corresponding author: pietro.mazzola@unipa.it

ABSTRACT Geophytes oceurring in the Sicilian archipelago are examined with respect to their distribution

and evolution, and also taking into account correlations with the inner parts of this territory

and other regions in the central Mediterranean.

KEY WORDS Biogeography; geophytes; endemism; insularity.

Received 11.05.2012; accepted 09.12.2012; printed 30.12.2012

Proceedings of the International Congress “Insularity and Biodiversity”, May 11*-13*, 2012 - Palermo (Italy)

INTRODUCTION

According to Raimondo et al. (2010) the vascu-

lar flora of Sicily and surrounding islands is made

of 3252 taxa of speeifie and lower ranks, among

whieh Angiosperms are 3173, ineluding 2463 dieo-

tyledons and 710 monoeotyledons.

Several eomprehensive works on sueh plant he-

ritage have been earried out sinee the seventeenth

eentury (Boeeone, 1674; Cupani, 1696-1697; Ueria,

1789; Gussone, 1842-1845; Lojaeono Pojero, 1888-

1909; Giardina et al., 2007; Raimondo & Spadaro,

2009), together with a large number of other eontri-

butions (efr. Raimondo et al., 1982) mainly dealing

with the most important fields of traditional botani-

eal interest like floristies, taxonomy, history, ete.,

and, in addition, relationships with other Mediterra-

nean eountries (Lojaeono Pojero, 1888-1909), ge-

neral eharaeteristies of biology and ehorology (Di

Martino & Raimondo, 1979), phytogeographieal de-

limitation of the Sicilian floristic domain (“Dominio

siculo”) (Brullo et al., 1995), ete. More reeently Rai-

mondo & Spadaro (2011) analyzed eritieally this

flora and listed 502 endemies, 322 of whieh are ex-

elusive to the arehipelago. Owing to these studies,

Sieily is nowadays eonsidered among the better sur-

veyed in the Mediterranean basin (Raimondo, 1988;

Brullo et al., 1995) as far as flora, taxonomy and bio-

geography are eoneemed. The knowledge of this

plant riehness and diversity ean be still more impro-

ved by eorrelating endemism, karyology, insularity

and other heterogeneous faetors possibly suitable

(when taken together into aeeount) to elarify speeifie

problematie aspeets of biogeographieal nature.

Some remarks referring to geophytes are presented

here with respeet to their evolution.

MATERIALS AND METHODS

In order to mark eonneetions or isolation phases

in whieh the archipelago was involved, plants with

a geophyte life form have been seleeted sinee they

are perennial and their ranges apparently vary slo-

wly. The taxa taken in eonsideration belong to the

families Orehidaeeae and Liliaeeae s.L. They are in

part endemie to the whole arehipelago (ineluding

Malta, in some eases) and in part to small islands

364

P. Mazzola, R. Schicchi & S. Ciccarello

or to circumscribed areas of Sicily that were islands

in the past. These latter partly correspond to the flo-

ristie subdivision provided by Brullo et al. (1995).

Besides distribution outlines, for eaeh taxon,

when possible, karyologieal data, mostly dedueed

from literature, are also taken into aeeount. Nomen-

clature follows Giardina et al. (2007).

SICILY

The above quoted 322 exclusive endemies,

mark a signifieant evidenee of the whole Sicilian

archipelago insularity; while in the total of 502

there are also included other taxa whose ranges

show eontaets happened with other Mediterranean

areas. In addition to such general information,

from the analysis of single taxa some evidence on

their aetual insular endemic condition is deduced

and, besides, on the relevant evolution processes.

In particular, those endemies exclusive to Sieily

and there spread everywhere show their insular

condition.

Among these there are several orchids like

Ophrys oxyrrhynchos Tod., O. lunulata Pari, and

Orchis commutata Tod. (Fig. 1) which, occurring

throughout the region, is likely a good example of

a neo-endemie tetraploid (2n = 84) viearianee with

respeet to O. tridentata Seop., diploid (2n = 42),

whose range eovers Central and S-Europe up to the

region of Calabria (Mazzola, 1984) where is its sou-

thernmost distribution limit lies. O. tridentata is in-

deed missing in Sicily. Apart from some slight

differenees in size, these two taxa are almost indi-

stinguishable. O. lunulata (2n = 36) oecurs throu-

ghout Sieily, but it is more frequent in the extreme

SE, the Iblei Mountains (the Iblei District, accor-

ding to Brullo et al., 1995), where O. biancae (Tod.)

Maeeh. (2n = 36), O. caesiella P. Delforge, and se-

veral other more or less distinet taxa belonging to

the O. lutea and O. fusca groups oeeur frequently.

The oecurrenee of these geophytes, together

with a speeial terrestrial vertebrate palaeofauna, and

other loeal endemie or rare plant speeies (Brullo et

al., 1995) show that the Iblei Mountains were an is-

land during the Middle Pleistoeene (Guglielmo &

Marra, 2011). In that period, the rest of the present

Sieily was in part submerged, and in part, nor-

thwards, oeeupied by another large island, including

the mountainous ranges between Messina and Tra-

Figure 1. Orchis commutata, a tetraploid vicariant of O.

tridentata', distribution: Sicily.

pani. The western part of this palaeo-island is cha-

racterized by several other endemic geophytes.

Among these, O. pallida Raf. (Fig. 2) occurs bet-

ween the Madonie Mountains (here very rare) and

the mountains around and south of Palermo, espe-

eially in the Monte Busambra zone, and in the

Monte S. Calogero west of the town of Termini

Imerese, one of past eoastal islands having been in-

corporated in the northern Sicilian littoral (Rai-

mondo et al., 2001).

In western Sicily, Oncostema ceruleum (Raf.)

Speta occurs with a range similar to O. pallida, to-

gether with many other (not only) exelusive ende-

mies like Colchicum gussonei Lojae. (related to C.

cupanii Guss. and there included by Giardina et al.

(2007)), and several endemie Gagea speeies such

as G. busambarensis (Tineo) Park, G. longifolia

Lojae., G. sicula Lojae. G. lacaitae A. Terrace., G.

ramulosa A. Terrace, that virtually refer to the

above mentioned western side of the northern Mid-

dle Pleistoeene island, i.e. the Distriet Drepano-Pa-

normitano by Brullo et al. (1995). Gagea also

occurs on the Madonie Mountains with several spe-

Geophytes and evolution in the Sicilian Archipelago

365

Figure 2. Ophrys pallida, endemic to Sicily from the Ma-

donie Mountains to westwards.

Figure 3. Orchis brancifortii, endemic to Sicily, S-Calabria

and Sardinia.

cies as G. chrysantha A. Terrace., G. nebrodensis

(Tod. ex Guss) Nyman, G. granatelli (Pari.) Pari.

G. dubiaA. Terrracc., and with G. pratensis (Pers.)

Dumort., G. lutea (L.) Ker-Gawl. in the Nebrodi

and Peloritani Mountains (the northeastern district

together with the Madonie and the Etna Mountains,

(Brullo et ah, 1995), while G. trinervia (Viv.) Grea-

ter is found in South-Eastern Sicily.

About this incomplete list, it is to be noted that

most taxa have not been confirmed recently (Pe-

ruzzi & Tison, 2005). This, neither from the taxo-

nomical point of view, nor as far as distribution

aspects are concerned. It is, nevertheless, interesting

to state that there are at least two localities, on the

Busambra and surrounding mountains and on the

top of the Madonie, where individual morphologi-

cal diversity is so intensive that they could be con-

sidered as true centres of local variation. Regarding

the Madonie Mountains (the “Pizzo delle Case”, at

1850 m a.s.l.), such high variation has skilfully

been illustrated by the nineteenth naturalist France-

sco Mina Palumbo (2011). On the other hand, the

genus, being largely distributed, refers to the gene-

ral insularity of Sicily. As far as the Nebrodi and

Peloritani Mountains are concerned (in spite of

these areas are characterized by several noteworthy

local endemics like Petagnaea gussonei (Spreng.)

Rauschert, the unique representative of a monospe-

cific genus, besides Carduus rugulosus Guss., Cir-

sium vallis-demonii Lojac. etc. in the Nebrodi;

Centaurera seguenzae (Lacaita) Brullo, Centaurea

tauromenitana Guss., etc.), geophytes generally

mark repeated contacts with Italy through the sou-

thern Calabria. Some of these are the orchids Dac-

tylorhiza sambucina (L.) So6, Orchis morio L.,

Polygonatum gussonei Pari. (Convallariaceae), Fri-

tillaria messanensis Raf. (Liliaceae), Aristolochia

lutea Desf. (Aristolochiaceae) (2n = 8), A. clemati-

tis L. (2n = 14).

Indeed, as Nardi (1984) pointed out, the genus

Aristolochia can clarify some taxonomic and phy-

togeographical relationships between Sicily and

other surrounding regions. In particular, apart from

the two above mentioned species and A. sicula

Tineo (2n = 1 6), a taxonomically isolated endemic

to the Madonie, Nebrodi, Peloritani mountains and

366

P. Mazzola, R. Schicchi & S. Ciccarello

the Etna, A. navicularis Nardi (2n = 24), is distri-

buted in Sardinia, the Egadi Islands and in the nor-

thern Algeria and Tunisia; A. altissima Desf. marks

eontaets between SE Sieily and Algeria; A. clusii

Lojae. (2n =12) oeeurs in S-Italy, Sieily and Malta.

From a more general point of view, there are several

other geophytes showing similar eorrelations in the

Central Mediterranean.

Among these. Orchis brancifortii Biv. (Fig. 3)

oeeurs in Sardinia, Sieily, and probably in some eal-

eareous sites seattered in the southernmost Calabria.

O. longicornu (Poir.) oeeurs in Sieily, Sardinia and

Corsiea; Serapias nurrica Corrias, first deseribed

as endemie to Sardinia (Corrias, 1982) is presently

known oeeurring seattered in several loealities of

Sieily (Giardina et al., 2007), Calabria, Corsiea

(Griinanger, 2001) and Tunisia (Vela et al., 2012).

As regards other regions, in eomparison with O.

quadripunctata Cirillo ex Ten., O brancifortii ean

be eonsidered as an interesting ease of viearianee

relating to southeastern European taxa. Similar eor-

relation with SE European flora ean be found in se-

veral other taxa (Galanthus, for instanee).

CIRCUMSICILIAN ISLANDS

In addition to the above outlined relations, most

of the instanees eoneeming distribution and evolu-

tion of the endemie pattern are inside the Sieilian

arehipelago itself Here only some eases are taken

into aeeount.

The orehid genus Anacamptis Rieh., whose

wide range ineludes most of Europe and the Medi-

terranean, in Sieily is represented by A. pyramida-

lis (L.) Rieh. (2n = 36), wieh is widespread

generally on limestone ground. This speeies is also

found in the Maltese islands there bearing the te-

traploid 2n = 72 number. Furthermore in the same

islands another loeal diploid endemie speeies is

eonfined, A. urvilleana (Sommier) Caruana (2n =

36), being morphologieally, karyologieally and

phenologieally quite distinet from A. pyramidalis .

The oeeurrenee of these two isolated taxa has been

eonsidered as the result of eontaets happened bet-

ween Sieily and Malta and eonsequent evolution

proeesses that have been repeated at least two times

(Del Prete et al., 1991).

Another example in the family of Hyaeinthaeeae

refers to Oncostema, a genus with a eentral and W-

Mediterranean range, whieh in Sieily is represented

by 4 native speeies endemie to the Arehipelago

(Fig. 4) and 1 naturalized there {Oncostema peru-

vianum (L.) Speta, from SW Iberian peninsula).

The native taxa are: O. siculum (Tineo ex Guss.)

Speta (2n = 28) (Fig. 5) whieh oeeurs in several,

seattered loealities of NW and SE Sieily, and in

Malta where aneuploid elements have also been re-

eorded; O. hughii (Tineo ex Guss.) Speta (2n = 28)

(Fig. 6), whieh is eonfined in Egadi Island of Ma-

rettimo, west of Sieily; O. dimartinoi (Brullo et Pa-

vone) F. Conti et Soldano (Fig. 7), eonfined in the

islet of Lampedusa, loeated between Sieily and

Malta. Finally O. ceruleum (Raf.) Speta, distribu-

ted in the eentral and western part of Sieily. This

fragmented distribution at the speeies level gene-

rally agrees with some possible evolution happe-

ned in a rather far past. Therefore, some small areas

where O. siculum and O. ceruleum oeeurr ean pre-

sently be eonsidered as having had an insular status

like the eoastal more evident ones shown by Rai-

mondo et al. (2001).

Considering the eomplex surrounding Sieily, in

general eaeh island denotes its own insular eondi-

tion through some loeal representatives whose neo-

or palaeo- endemie nature refers to the voleanie or

ealeareous geologieal origins. Among these. Allium

is represented by several speeies loealized in very

small islets (efr. Giardina et al., 2007). In partieular,

in the Egadi islands, A. aethusanum Garbari and^.

franciniae Brullo et Pavone, are endemie to Favi-

gnana and Marettimo, respeetively.

In addition, A. lopadusanum Bartolo et al., is a

rare endemie to Lampedusa (Pelagie islands); A.

longispatum Lojae., of uneertain value, is loeated

in the past islet of Capo Catalfano (Raimondo et

al., 2001). Regarding other families showing simi-

lar distributions, in the Amarylidaeeae family,

Paneratium linosae Soldano et Conti is eonfined

in the Linosa islet (Pelagie islands); and among

the orehids, Serapias cossyrensis B. et H. Bau-

mann, related to S. eordigera L., is endemie to

Pantelleria (Pelagie islands). Finally, Ophrys sco-

lopax Cav. s.l. is cited owing to it occasionally oc-

curs in the islets of the Archipelago without any

constant localization. This is another characteristic

of geophytes, especially orchids, and their occur-

rence in the islands.

Geophytes and evolution in the Sicilian Archipelago

367

Figure 4. Distribution of Oncostema in the Sicilian Archipelago, showing its probable Pleistocenic origin. Figure

5. O. siculum, distribution Sicily and Malta; specimen from Corleone (NW-Sicily) cultivated in the Botanical

Garden of Palermo. Figure 6. O. hughii, confined in the Marettimo island. Figure 7. O. dimartinoi, confined in

the Lampedusa island.

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Biodiversity Journal, 2012, 3 (4): 369-374

Insular endemism in the Mediterranean vascular flora: the

case of the Aeolian Islands (Sicily, Italy)

Angelo Troia

Dipartimento di Biologia ambientale e Biodiversita, Universita degli Studi di Palermo, via Archirafi 38 -90123 Palermo, Italy; email:

angelo . troia@unipa . it

ABSTRACT The present paper briefly provides the state of the art of the knowledge on vascular plant

endemism in the oceanic (“thalassogenous”) Aeolian Archipelago (Sicily). Preliminary

analysis of distribution areas and review of recent literature on biosystematics of endemic

species revealed that: (a) Aeolian strictly endemic taxa are just 6, i.e. about the 0.7 % of

the local vascular flora; among them, just 4 can be considered (with doubt) derived from

in situ evolution, (b) The other 18 endemics are taxa occurring in wider areas, so they can-

not be generally considered “locally evolved” but relicts. This preliminary analysis con-

firms that not only continental (“chersogenous”) but all Mediterranean islands are primarily

conservative rather than evolutionary active systems.

KEY WORDS speciation; evolutionary refugia; dispersal; island biota.

Received 11.05.2012; accepted 18.12.2012; printed 30.12.2012

Proceedings of I International congress “Insularity and Biodiversity”, 11-13 May 2012, Palermo, Italy

INTRODUCTION

Islands and archipelagoes have long faseinated

biologists, primarily beeause of their unique and so-

mewhat unusual faunas and floras (Thomson,

2005). The emersion of the Aeolian Archipelago

(Fig. 1), a voleanie are separated from Sieily and

Italian Peninsula by sea ehannels of about 1,000-

2,000 m depth (Allen & Morelli, 1971), began

about 500,000 years b.p. (Calanehi et al., 2007).

Despite its relatively “young” age, but maybe

thanks to its continuing isolation during the Pleisto-

eene falls in sea level, a fair number of endemic

plant and animal species oceurs on one or more is-

lands (Lo Caseio & Sparaeio, 2010).

Endemism has multiple eauses and a diversity

of faetors can influence variation in range size. En-

demic plants are eommonly distinguished in two

main categories: relicts or newly formed. These two

eategories are eommonly referred to as palaeo-en-

demic and neo-endemic taxa, respectively (Favar-

ger & Contandriopoulos, 1961), or as Zohary

(1973) preferred, primary (aetive) and seeondary

(reliet) endemism. In the Mediterranean region, is-

lands are for the most part fragments of land that

have beeome isolated due to their separation from

eontinental areas; the few exeeptions inelude the is-

lands originated from submarine voleanie aetivity,

sueh as the Aeolian Islands. The fonuer are usually

known as “eontinental islands”, the latter “oeeanic

islands”, even if a different terminology was pro-

posed by Greater (1979): namely, “ehersogenous”

for the former, “thalassogenous” for the latter ones.

In thalassogenous islands, initially, “empty

spaee offers itself for a few neweomers to expand

and adapt. The chersogenous islands, to which all

Mediterranean islands of an appreeiable size be-

long, already earried their own, fully adapted and

diversified flora when they became insular. No

empty spaee could they offer, no speeial ehallenges

370

Angelo Troia

to new colonists, no new horizons (...). They are

conservative systems, remarkably well buffered

against the effects of climatic and evolutionary

change” (Greater, 2001). According to this view,

the endemic species, occurring nowadays on the

thalassogenous Aeolian Islands, are expected to be

derived from in situ evolutionary processes. Aim

of this paper is to briefly provide the state of the

art of the knowledge on vascular plant endemism

in this archipelago.

MATERIALS AND METHODS

The current available list of Aeolian vascular

plant endemic taxa (Mazzola et al., 2001; Lo Cascio

& Pasta, 2004) was verified, according to the more

recent available literature (Brullo & Minissale, 2002;

Peruzzi & Passalacqua, 2003; Cristofolini & Troia,

2006; Euro+Med, 2006-; Ferro, 2009; Peruzzi &

Vangelisti, 2009; Kadereit & Freitag, 2011; Hilpold

et al., 2011; Bacchetta et al., 2012), mainly to assess

the taxonomic status and the actual distribution area

of each taxon. Taxa of doubtful taxonomic value or

not endemics were excluded from the revised list.

The resulting strictly endemic taxa were tenta-

tively assigned to one of the two categories: pri-

mary (active) or secondary (relict) endemics, these

categories better defining the origin of the ende-

mism regardless of its age.

RESULTS

An up-to-date list of the endemic plant taxa occur-

ring in Aeolian Islands was achieved, distinguishing

the ones strictly endemic of the archipelago (Table 1)

from those endemic of wider areas (Table 2).

Anthemis aeolica Lojac., generally considered a

synonym of A. maritima L. (Pignatti, 1982; Giar-

dina et al., 2007) but preliminary accepted in some

recent works (Greater, 2006-2009; 2008), was in-

cluded in Table 1 . It was described as an annual

plant, occurring just in few small islets near the Pa-

narea island (Lojacono, 1902-1903; Lo Cascio &

Navarra, 2003). Considering that in annual Anthe-

mis “prevailing mode of speciation (...) is allopatric

speciation and local (founder effect) speciation is

usually encountered” (Oberprieler, 1998), and that

a local endemic species with a similar micro-insular

habitat occurs in Aegean area (Georgiou et al.,

2006), certainly this taxon of doubtful taxonomic

value deserves more attention and further studies.

DISCUSSION AND CONCLUSIONS

The oceanic island ecosystems offer great op-

portunities for the study of evolution and have for

a long time been recognized as natural laboratories

for studying evolutionary processes involved in di-

versification owing to their discrete geographical

nature and diversity of species and habitats (Emer-

son, 2002).

Preliminary analysis of distribution areas and re-

view of recent literature on biosystematics of ende-

mic Tyrrhenian taxa revealed that:

- Aeolian strictly (“narrow”) endemic taxa are

just 6 (Table 1); considering that Aeolian vascular

flora consists of approximately 900 taxa (Pasta,

1997; Privitera et al., 2008; unfortunately, a com-

plete published checklist of the Aeolian flora does

not exist), the strictly endemic taxa are just the 0.7

% of the local flora.

- Of these few strictly endemic taxa, just 4 can

be considered (with doubt) derived from in situ evo-

lutionary processes.

- Most of the “endemics” (sensu lato) are spe-

cies occurring in wider areas, or at least in another

place outside the archipelago (Table 2); this is a

clear clue to consider them secondary endemics.

Considering their biology and taxonomic/phyloge-

netic relationships (but also the young age of the is-

lands, and their relative proximity to other

landmasses), it is reasonable to think that - in gene-

ral - they are not new taxa, evolved in the studied

archipelago and then spread outside, but they colo-

nized these islands from other nearby populations;

this happened, in one case at least {Cytisus aeolicus.

Fig. 2, cf Conte et al., 1998), also for the nowadays

strictly endemic species.

- If all the endemics (24 taxa. Tables 1 and 2)

are considered, the percentage of plant endemism

in the archipelago becomes ca. 2.7 %, a low value

but comparable with others concerning Mediterra-

nean islands such as Malta (2.3 %), Balearic Islands

(6.5 %) (Medail & Quezel, 1997) or Egadi Islands

(5.9 %) and Pelagian Islands (6.1 %) (Mazzola et

al., 2001).

- As to Greater ’s (2001) view, mentioned in the

introduction, both palaeo- and neo- endemics occur

Insular endemism in the Mediterranean vascular flora: the case of the Aeolian Islands (Sicily, Italy)

371

in the thalassogenous/oceanic Aeolian Islands, pro-

bably according to random factors related to the

biology of the species and the geological history of

their habitat.

After this preliminary survey and waiting

for further investigations, some conclusions can

be made:

1) As already implicitly suggested by Greuter

(2001), not only chersogenous but all Mediterra-

nean islands are primarily conservative systems.

Verlaque et al. (1997) also observed that palaeo-

endemic species may be more prevalent in the en-

demic flora of Mediterranean islands compared to

continental areas. This observation suggests that

geographic isolation may be important not just be-

cause it promotes genetic differentiation and the

Figure 1. A view of the Aeolian arehipelago as seen from the East (from an airliner). Below, the Calabria (Italy). On the

left, after the Messina Straits, Sieily with its 3,330 m high Etna voleano. On the right, all the seven main Aeolian islands

ean be distinguished (photo A. Troia). Figure 2. Cytisus aeolicus Guss. It is a palaeoendemie small tree, whose wild po-

pulations oeeur only in the Aeolian archipelago. To stress its isolated taxonomic position, it was put in a monotypic

section (sect. Dendwcytisus) within its genus (see Cristofolini & Troia, 2006) (photo A. Troia). Figure 3. Strombolicchio,

near the island of Stromboli, is one of the several “minor” islets of the Archipelago. “Minor” is said with reference to

their size, but they often house very interesting animal and plant species. For example, Strombolicchio houses the only

Aeolian population of Eokochia saxicola (Guss.) Freitag & G. Kadereit, occurring also (and only) in the Campanian is-

lands of Capri and Ischia (here extinct) and in two recently discovered sites in mainland Campania, near Cape Palinuro

(Santangelo et al., 2012); photo A. Troia.

372

Angelo Troia

evolution of endemism, but also beeause it reduees

immigration of eommon speeies and thereby fa-

vours the persistenee of endemie speeies on islands

(Thomson, 2005).

2) Geographie distanees separating Aeolian Is-

lands from other nearby landmasses may be (and

may have been) too small to guarantee an efifeetive

“perfeef ’ isolation (ef. Troia et ah, 2012); this, eou-

Anthemis aeolica Lojac.

Primary endemic (?)

Centaurea aeolica Guss. ex Lojac. subsp. aeolica

Primary endemic (?) (*)

Cytisus aeolicus Guss.

Secondary endemic

Erysimum brulloi Ferro

Primary endemic (?)

Genista tyrrhena Vais, subsp. tyrrhena

Primary endemic (?) (*)

Silene hicesiae Bmllo & Signorello

Secondary endemic (?)

Table 1. Strictly endemic taxa of Aeolian Islands.

(*) The presence of the same species in two different archipelagoes (Aeolian and Pontine Islands) contradicts its in situ

evolution; assuming that the subspecific differences are significant, each subspecies (strictly endemic of a different archi-

pelago) can be considered locally evolved.

ENDEMIC TAXA OF AEOLIAN ISLANDS AND NEAR (N-E) SICILY

- Dianthus rupicola subsp. aeolicus (Lojac.) Bmllo & Minissale

- Limonium minutiflorum (Guss.) O. Kuntze

ENDEMIC TAXA OF AEOLIAN ISLANDS AND WESTERN SICILY

- Ranunculus spicatus subsp. rupestris (Guss.) Maire

- Seseli bocconi Guss. subsp. bocconi

ENDEMIC TAXA OF AEOLIAN ISLANDS AND TYRRHENIAN AREA

- Beilis margaritifolia Huter, Porta & Rigo

- Carlina hispanica subsp. globosa (Arcang.) Meusel & Kastner

- Carlina sicula Ten. subsp. sicula

- Eokochia saxicola (Guss.) Freitag & G. Kadereit (Fig. 3)

- Helichrysum litoreum Guss.

- Heliotropium suaveolens subsp. bocconei (Guss.) Brummitt

- Matthiola incana subsp. rupestris (Raf ) Nyman

- Micromeria graeea subsp. eonsentina (Ten.) Guinea

- Ranunculus pratensis C. Presl

ENDEMIC TAXA OF AEOLIAN ISLANDS AND CENTRAL- WE STERN MEDITERRANEAN

- Daucus carota subsp. rupestris (Guss.) Heywood

- Glandora rosmarinifolia (Ten.) D.C. Thomas

- Hyoseris lucida subsp. taurina (Martinoli) Peruzzi & Vangelisti

- Iberis semperflorens L.

- Silene turbinata Guss.

Table 2. Not-strictly endemic taxa of Aeolian Islands.

Insular endemism in the Mediterranean vascular flora: the case of the Aeolian Islands (Sicily, Italy)

373

pled with the relatively young age of the islands and

their voleanie instability, may have prevented loeal

(in situ) speeiation.

3) Further studies on Aeolian flora are needed.

ACKNOWLEDGEMENTS

I would like to thank Salvatore (“Salvo”) Pasta

for reading my manuseript in a short spaee of time,

with his great Aeolian experienee and usual proo-

freader’s eye.

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Biodiversity Journal, 2012, 3 (4): 375-384

Distribution and ecological patterns of orchids in Monte Pel-

legrino Reserve, Palermo (Sicily, Italy)

Vincenzo Bertolini'*,Anne Damon', Javier Valle Mora' &Angel Natanael Rojas Velazquez^

'Colegio de la Frontera Sur (ECOSUR). Carretera Aeropuerto Antiguo Km 2.5, Tapachula, Chiapas, Mexico; e-mail:

vbertolini@ecosur.mx

^Autonoma de San Luis Potosi, Facultad de Agronomia, Alvaro Obregon # 64 Colonia Centro, CP 78000, San Luis Potosi,

S. L. R, Mexico

*Corresponding author

ABSTRACT Despite heavy human impact through the ages, the Monte Pellegrino Reserve maintains an

interesting orchid flora currently estimated as 33 taxa, with 9 endemic elements, including

Ophrys lunulata Parltore (priority species for the European Commission) and O. sphegodes

panormitana (Tod.) Kreutz, which has its locus typicus in Monte Gallo, another reserve close

to Monte Pellegrino. The chorological types most represented in this study were Mediterra-

nean and Atlantic Mediterranean. The distribution of the orchids in this site was correlated to

various environmental parameters, depending upon species and chorological type. The ende-

mic species were the only chorotype that was positively correlated to two habitats that are re-

licts of the original Sicilian vegetation: clearings in deciduous thermophilic forest {Rhamno

alaterni-Quercetum ilicis subassociation terebinthi) and Ampelodesmetum mea-

dows (Helictotrico convoluti-Ampelodesmetum mauritanici). Prevailing wind direction and

seasonality were determining factors for orchid distribution. Identification of the correlation

patterns between chorological types and habitats could be useful for predicting species pre-

sence within areas with similar biogeographical characteristics, facilitate species mapping

and serve as a tool for the design and implementation of conservation strategies.

KEY WORDS orchids; chorotype; relict natural vegetation; Mediterranean ecosystems; priority species.

Received 11.05.2012; accepted 21.10.2012; printed 30.12.2012

Proceedings of the P' International Congress “Insularity and Biodiversity”, May 11*-13*, 2012 - Palermo (Italy)

INTRODUCTION

The Monte Pellegrino (Palermo, Sieily, Italy) is

a natural area whieh has been ineorporated into the

European eeologieal network Natura 2000, designa-

ted as a S.I.C. (Sito dTnteresse Comunitario: Site

of Community Interest) eode ITA 020014 Monte

Pellegrino, due to the presenee of rare and endemie

speeies and habitats, sueh as the priority speeies

Ophrys lunulata Pari. (Natura 2000 eode: 1905), the

speeies of eommunity interest Dianthus rupicola

Biv. (Caryophyllaeeae), (eode: 1468), as well as the

priority habitats, residual serub forest with Laurus

nobilis L. (Lauraeeae) (Natura 2000 habitat eode:

5230) and residual serub forest with Zyziphus lotus

L. (Lam.) (Rhamnaeeae), (eode: 5220) and the ha-

bitat of eommunity interest substeppe, or meadow,

with grasses and annual herbs Thero-Brachypodie-

tea (eode 6220) (Gianguzzi et al., 1995).

The Monte Pellegrino Reserve (38°1E42.60”N -

38°08'53.01”N and 13°21 '53.95”E - 13°19'58.82”E)

is partially surrounded by the eity of Palermo to-

wards the south and south-east, and by the sea in

the north and north-west. The perimeter of the re-

376

V. Bertolini.A. Damon, J.Valle Mora & A. N. Rojas VelAzquez

serve is eharaeterized by numerous steep slopes and

sea eliffs at altitudes of 100-150 m whieh prevent

aeeess (Quartarone et al., 1995). The reserve is si-

tuated on dolomitie limestone roek; the lithosoils

eontain 30 to 80% sand, 20% elay and a low humus

eontent with a mostly neutral pH (6.8-7. 5) (Monta-

nari, 1964; Raimondo & Venturella, 1996; Fierotti,

1997). The original vegetation of the area eonsists

of Mediterranean serub forests with key speeies:

Chamaerops humilis L. (Areeaeeae), Euphorbia ar-

borescens Salm-Dyek (Euphorbiaeeae), Pistacia

lentiscus L. (Anaeardaeeae), Olea europea L. (Olea-

eeae), Ceratonia siliqua L. (Fabaeeae), as well as

deeiduous thermophilie forest featuring Quercus ilex

L. (Fagaeeae) and lithophytie vegetation on exposed

roek faees (Quartarone et al., 1995; Raimondo &

Venturella, 1996; Raimondo et al., 1996a).

Territories that surround the Monte Pellegrino

Reserve have been eolonized by humans from pre-

historie times (Vigliardi, 1991) and natural resour-

ees have been exploited for eenturies, leading to the

almost total elimination of forest eover by the XIX

eentury, as witnessed by Goethe (1905). However,

from the beginning of the 1 9* eentury the gover-

nment has intervened with plans to reforest the hills

to reduee erosion although the speeies used for re-

forestation have not always been the most appro-

priate and have not taken into aeeount the

biogeographieal features of the site. Today, vast

areas planted to alien speeies ean be observed, in-

eluding Pinus halepensis Mill, and P pinea L. (Pi-

naeeae), Cupressus sempervirens L. (Cupressaeeae),

Eucalyptus globulus Labill. and E. rostrata

Sehleeht. (Myrtaeeae) as the main elements, but also

mchxdmg Acacia saligna (Labill.) H.L. Wendl. (Fa-

baeeae), Opuntia ficus-indica (L.) Mill. (Caetaeeae),

Jacaranda mimosifolia D.Don 1822 (Bignoniaeeae),

Koelreuteria paniculata Laxm. (Sapindaeeae), Schi-

nus mode L. (Anaeardiaeeae), amongst others (Sero-

fani, 1949). Other exotie speeies that have invaded

the site are Pennisetum setaceum (Forssk.) Chiov.

(Poaeeae) within the steppe grassland habitat, and

oeeasional populations of Ailanthus altissima (Mill.)

Swingle (Simaroubaeeae), Agave sp. (Asparaga-

eeae), Nicotiana glauca Graham (Solanaeeae), Aca-

cia karoo Hayne (Fabaeeae) and Parkinsonia

aculeata L. (Fabaeeae) (Gianguzzi et al., 1996). In

addition, Mazzola & Di Martino (1996) report the

use of exotie, ornamental speeies in the areas with

most human impaet. Although the history of the site

has elearly provoked extensive alteration to the na-

tural vegetation of Monte Pellegrino, Raimondo &

Venturella (1996) mentioned that of the 741 speeies

and hybrids that eonstitute the present day vaseular

flora, 716 are native in origin. Considering the eryp-

togamie flora, mosses aeeount for 76 of the present

taxa, liehens for 83 and fungi for 53 (Raimondo &

Venturella, 1996; Venturella, 1996). Extensive areas

of the territory are heavily affeeted by eattle gra-

zing, wood extraetion and burning, the dumping of

non-biodegradable residues and the eontamination

of the air with traffie fumes (Raimondo et al.,

1996a; Venturella, 1996).

The elimate of the area is Mediterranean, with

xerophytie and thermomediterranean sub seetions

(Gianguzzi et al., 1996). The sea, towards the north

and north west, gives rise to humid air eurrents

throughout the year, whieh mitigate and influenee

loeal mieroelimates (Raimondo & Venturella, 1996;

Raimondo et al., 1996b) making the north of the

area eooler and more humid as eompared to the

more xerophytie south; the average annual relative

humidity is 67%, with a minimum of 58% in Oeto-

ber and a maximum of 74% in Mareh (Albaria,

2011). In the north of the area, environmental eon-

ditions favour deeiduous thermophilie Rahmno ala-

terni-quercetum ilicis subassoeiatio pistacietosum

terebinthi, whieh in turn favor the presenee of Celtis

australis L. (Ulmaeeae), Fraxinus ornus L. (Olea-

eeae) and Rhus coriaria L. (Anaeardiaeeae) (Gian-

guzzi et al., 1996). Towards the south, the dominant

speeies of the xerophytie serub are Oleo sylvestris-

Euphorbietum dendroidis subassoeiation euphor-

bietosum bivouac and Chamaerops humilis.

Aeeording to G.I.R.O.S. (2009), Italy has about

200 taxonomie entities, for Orehidaeeae, of the total

of almost 700 registered for Europe and eireum-Me-

diterranean eountries, within whieh Sieily is one of

the regions of Italy with the highest ineidenee of or-

ehids, with 90 taxa, ineluding speeies and subspeeies

(Bartolo & Pulvirenti, 1997). The provinee of the

eity of Palermo has an extension of 5000 km^, with

60 taxa registered (Giardina, 2005), and ineludes the

Madonie National Park whieh plays a prineipal role

with extensive forested areas and environmental le-

gislation in aetion whieh guarantees the proteetion

of the biodiversity of this important natural lung,

with a total of 57 orehid taxa (Pueeia, 1995). The

first data for the Monte Pellegrino Reserve were eol-

leeted by the Sieilian botanist Filippo Parlatore

Distribution and ecological patterns of orchids in Monte Pellegrino Reserve, Palermo (Sicily, Italy)

377

(1858) who reported 9 taxa, whereas in more reeent

floristie and vegetation studies, Gianguzzi et al.

(1996) and Raimondo et al. (1996b) determined the

presenee of 28 orehid speeies in the reserve. Finally,

Bertolini & Giardina (2008) and Bertolini (2009a,

b), mentioned a total of 27 speeies and 4 natural hy-

brids, within 5 genera (Anacamptis, Barlia, Ophrys,

Orchis and Serapias). Additionally, Grasso (2009)

reported the diseovery of Ophrys incubacea Bianea

subsp. incubacea, and its hybrids, as well as a po-

pulation of Serapias lingua L., albino type (Berto-

lini, personal unpublished data).

Studies on the population dynamies of native

North Ameriean terrestrial orehid speeies have

shown that the probability that a seed will germinate

and that the plant will reaeh maturity depends upon

a multitude of biotie and abiotie faetors (Diez, 2007).

In the ease of the genus Goodyera (Diez, 2007), and

Australian terrestrial orehids (Perkins & MeGee,

1995) the seed has a greater probability of germina-

ting if it falls elose to an adult plant of the same spe-

eies, whieh will already be inoeulated with suitable

fungi, and where a low pH (aeid), and presenee of

organie matter favour proliferation of the fungi.

The presenee of grazing animals may be dama-

ging to orehid populations as the animals may eat

the aerial, reproduetive parts of the plants, or even

the whole plant (Hutehings, 1987). Human inter-

vention thus modifies natural habitats, artifieially

affeets the natural distribution and habitat preferen-

ees of the orehids and that of their mutualists also

(Bergman et al., 2006). To design appropriate eon-

servation strategies for terrestrial orehids, the first

step is to evaluate the distribution of eaeh speeies,

as related to elimate and habitat eharaeteristies

(Sanford, 1969) and detennine the identity and di-

stribution of mutualist organisms, sueh as myeor-

rhizal fungi and speeialized pollinators (Brys et al.,

2008). The distribution of hybrids and subsequent

proeess of speeialization, adaptation and speeiation

also depends upon the distribution of the pollinators

and the habitat eolonized by the orehids (Sehatz,

2006). Strategies may inelude ex situ eonservation,

as well as ex situ propagation and eultivation to

eontribute towards the restoration of in situ popu-

lations, of both the orehids and their symbionts

(Swarts et al., 2007). Due to the rarity of many of

the Mediterranean orehids, deseriptions of preferred

habitats and environments are limited, but the lite-

rature mentions wide distributions and plastieity as

typieal of these orehids (Pignatti, 1982; Del Prete

6 Tosi, 1988; Delforge, 2005; G.I.R.O.S., 2009),

sueh as in the ease of O. lunulata, a rare endemie

Sieilian speeies present in Monte Pellegrino and

found at 1000 m asl, in a variety of habitats sueh as

pasture, meadows, Meditteranean serub, forest elea-

rings and full sun. From the data available, we

sought to identify eeologieal patterns and eorrela-

tions between orehid ehorotypes and habitat type in

Monte Pellegrino, with emphasis on endemie, Me-

diterranean and Atlantie Mediterranean speeies, to

statistieally eonfirm empirieal observations. We

propose a model that eould be applied to eoastal

Mediterranean areas in general.

In this study we set out to analyze the spatial di-

stribution of terrestrial orehids within the Monte

Pellegrino Reserve to determine distribution pat-

terns and eorrelations with habitat type for eaeh or-

ehid ehorologieal type. Using the data generated in

Monte Pellegrino Reserve as a model, we sought to

test the hypothesis that habitat type within the eoa-

stal zone of Sieily eould be used to prediet and de-

signate priority areas for orehid eonservation.

These biogeographie relationships on a small

seale are revealing eomponents of orehid eeology

whieh eould then be used to design strategies for

orehid eonservation on a larger seale and for other

areas. This type of analysis eould also be applied to

studies eoneeming the impaet of habitat modifiea-

tion due to elimate ehange.

MATERIALS AND METHODS

The study was earned out during the period

2007-2009 with visits to the sites throughout the

year exeept for the months of July and August. The

whole of the Monte Pellegrino Reserve was map-

ped and deseribed before being divided into see-

tions, aeeording to observable, predominating

habitat eharaeteristies as defined by Gianguzzi et

al. (1996); inaeeessible areas were not ineluded.

A detailed survey of the orehid speeies present

was earried out in plots of approximately 20-30 ha

eaeh, 8 sueh plots were eompleted in the first year,

7 in the seeond year and 3 in the final year. Due to

the rarity and patehy distribution of the orehids we

did not use quadrants; instead, the populations were

diseovered using transeets, with a sampling effort of

4-6 hrs. per session (day), a total of 12 sessions per

378

V. Bertolini.A. Damon, J.Valle Mora & A. N. Rojas VelAzquez

month and an estimated linear distanee of 10-12 km,

eovering 4.8 heetares. Eaeh session eonsisted of

walking the transeet and loealizing orehids within

approximately 2 metres in all direetions. Onee the

orehid populations were loealized, termed “points

of interest” (PI), the site was sampled intensely. The

PI were then visited monthly to monitor flowering

and extraetion of individuals (Bertolini, 2009a).

PI eoordinates were reeorded with a GPS Ma-

gellan eXplorist XL and other data were reeorded

with a handheld Sony PEG-TJ35/U, ineluding date

of first register for eaeh PI, altitude, date of flowe-

ring, no. of individuals, habitat type, direetion of

light exposition, and general observations.

Speeies determination was earried out using the

deseriptions of G.I.R.O.S. (2009), as well as eon-

sultation with experts where neeessary. For ehoro-

logieal data we referred to Griinanger (2001).

The distribution of orehids was studied in the fol-

lowing habitat types, aeeording to the elassifieation

of Gianguzzi et al. (1996): clearings in deciduous

thermophilic forest (Cl.DTF) (Rhamno alaterni-

Quercetum ilicis subassociation pistacietosum tere-

binthi), clearings in areas reforested with pine

(Cl.RPF) (Pinus halepensis, P pined), clearings in

areas reforested with a mixture of Pinus and Euca-

lyptus (Cl.RPEF) {Eucalyptus camalduensis), xero-

phytic nitrophilic grassland (XNG) (Carlino

siculae-Foerulum communis with Asphodelus micro-

carpus), Ampelodesmetum meadows (MA) {Helic-

totrico convolute-Ampelodesmetum mauritanici),

transition between xerophytic nitrophilic grassland

and pine forest (XNG-PF), Mediterranean scrub

(MS) {Oleo-Euphorbietum dendroidis subassociation

euphorbietosum bivouac), eroded wasteland (EW)

and steppe grassland (SG) (Pennisetum setacei-Hy-

parrhenietum hirtae).

The geographical coordinates or “marks” (xj,yj,

combined into one unit) that define each orchid

plant or group of plants, effectively correspond to

a random distribution, as these points were disco-

vered at random, which creates a stochastic spatial

process. Therefore the present study contemplates

a random, spatial process with “marks”:

y = {( xi, mi), ( X2, m 2 ), ..., (x„, m,,)} ,x/GfVc R\ mi e M

where Xj are the PI registered and m^ are the cor-

responding “marks”. The analysis was carried out

using the software R, specifically Spatstat (Badde-

ley & Turner, 2005; Bivand et al., 2008; R Deve-

lopment Core Team, 2011). As a descriptive tool,

and preliminary analysis, the spatial data was plot-

ted graphically against each of the “marks” mentio-

ned previously. The correlation between orchid

species and habitat type was analyzed by construc-

ting a spatial process using the “marks” M, which

in this case are of type 1,2. The value of 1 refers to

habitat type e.g. “i”, and 2 refers to the orchid spe-

cies, e.g. “j”. The first step is to plot a graph of the

distribution pattern against the “marks”.

To verify whether there was a correlation bet-

ween the occurrence of the “marks” and the PI, we

determined the function of the “marks” correspon-

ding to a stationary process, which is a measure-

ment of the dependence between the “marks” of

two points in the process, with a distance between

them of r. This can be defined as follows

Pf (r) = E[fiMi,M 2 ]

E \f{M, M‘]

where Mj, M 2 are the “marks” of the two points

which are separated by a distance r; while M, M^

are processes independent of the marginal distribu-

tion of the “marks”.

Function f is understood in this case as

J" {m\, ind) 1 {/Ml = ni2}

as a function of the categorical “marks”. The

correlation function is not a measure of correlation

in the usual, strict statistical sense. A value of 1, in-

dicates no correlation; i.e. that the occurrence of the

“marks” is completely random.

We also applied a randomization test to the data,

by which we tested the hypothesis that the “marks”

are conditionally independent and have identical di-

stributions. The test was constructed considering

the coordinates as fixed points and using a process

of repeated sampling.

RESULTS AND DISCUSSION

Chorology and populations

As a result of the three years study in the region, we

registered 33 taxa and 404 PI, with an estimated

total of 3000 individuals classified into 6 genera

(Anacamptis, Barlia, Neotinea, Ophrys, Orchis and

Serapias), giving a total of 28 taxa and 5 natural hy-

Distribution and ecological patterns of orchids in Monte Pellegrino Reserve, Palermo (Sicily, Italy)

379

2500

2113

2000

1500

1000

500

I total PI

total plants

504

254

j ^

127

n 10,

^■9

298

48 i

# /

1400

1200

1000

800

600

400

200

208

tn 61 tn

™ 31 IS 22 “5“ 111' 3 “4" 2'

^ i*? <=■

1 / / --

^ ^ i

O O /

Figure 1. Percentage contribution of chorological types, within the orchid population of the Monte Pellegrino Reserve,

Sicily. Figure 2. Total number of points of interest (PI) and individual plants, per genus. Figure 3. Total number of points of

interest (PI) and individual plants, per species, within the mediterranean (MED) chorotype. Figure 4. Total number of points

of interest (PI) and individual plants per species/hybrid, within the Endemic (END) chorotype. Figure 5. Total number of

points of interest (PI) and individuals per species, within the Atlantic Mediterranean (A-MED) chorotype. Figure 6. Total

number of points of interest (PI) and individuals per species, within the West Mediterranean (W-MED) chorotype.

380

V. Bertolini.A. Damon, J.Valle Mora & A. N. Rojas VelAzquez

brids (plus one unidentified hybrid, possibly deri-

ved from O. incubaced). The orehid speeies were

distributed between the ehorologieal types as fol-

lows: Mediterranean (MED) (31%), Endemie

(END) (27%) Atlantie Mediterranean (AT-MED)

(15%), West Mediterranean (W-MED) (12%), Eu-

romediterranean (EU-MED) (6%), Stenomediterra-

nean (S-MED) (6%) and East Mediterranean

(E-MED) (3%). A small number of plants (298) di-

stributed amongst 48 PI eould not be identified, as

flowering was not observed. In terms of individual

plants, again the ehorotype MED was the most

abundant (1794 individuals), followed by AT-MED

(567), W-MED (166) and END (141).

The E-MED, S-MED and EU-MED types are

the least represented ehorotypes (11, 52 and 7, re-

speetively) (Fig. 1). Analyzing the data for eonsi-

steney of ehorotypes, Ophrys is the genus that most

influenees the data with a total of 254 PI and 2111

individuals, as eompared to the genus Barlia whieh

is least influential with only two individuals in 2 PI

(Fig. 2). Within the MED ehorologieal type, Ophrys

bombyliflora Link has the most PI and the highest

number of individuals. Together with O. speculum

Link and O. lutea subsp. minor (Tod.) O. Daneseh

et E. Daneseh these are the most abundant orehid

speeies, while Neotinea lactea (Poir.) R.M. Bate-

man, Pridgeon et M.W Chase and Barlia rober-

tiana (Loisel.) Greater, are the rarest (Fig. 3).

Within the END type, Ophrys bertolonii subsp. ex-

planata (Lojae.) Soea and O. sphegodes subsp.

normitana (Tod.) Kreutz both have loeus typieus in

the Monte Gallo reserve (Parlatore, 1858), elose to

the study site, and have the highest number of indi-

viduals within this group. Ophrys lunulata, is a

priority speeies aeeording to the European Commis-

sion, and is very searee within the study area, with

only three individuals.

All the hybrids found, with the exeeption of

O.xsommieri E.G. Camus unidentified individuals

and others that have been reeognized as parental O.

incubacea subsp. incubacea (Bertolini, personal un-

published data), have been plaeed within the END

group, having at least one endemie parent. Hybrids

were infrequent, and limited to few individuals

(Fig. 4). Serapias lingua, Ophrys lutea subsp. lutea

Cav. and S. parviflora Pari, are among the orehid

speeies with highest frequeneies and number of in-

dividuals within the AT-MED group. In partieular,

S. lingua was found in 2 PI, one of whieh eontained

approximately 200 individuals and the other had

elose to 100 individuals of an albino variety. In both

eases the total area oeeupied was limited to a few

square meters probably due to the eapaeity of this

speeies to elone itself by extending stolons (Fig. 5).

Within the W-MED group, the most abundant spe-

eies were Ophrys tenthredinifera Willd., 1805 and

Anacamptis longicornu (Poir.) R.M. Bateman, Prid-

geon et M.W Chase, whereas the least frequent was

Ophrys lupercalis J. Devillers-Tersehuren et P. De-

villers with only two individuals in 2 PI (Fig. 6). The

E-MED group had 1 1 individuals of Ophrys lutea

subsp. phryganae (Devillers- Terseh. et Devillers)

Melki, 2000, in 2 PI.

Another infrequent speeies were Ophrys sphe-

godes subsp. sphegodes Mill. Huds., belonging to

the EU-MED group, eaeh found in only 1 PI. The

S-MED group was represented only by O. incuba-

cea subsp. incubacea, with 2 individuals plus some

hybrids.

Distribution

Analysis of the distribution of the different eho-

rologieal types, distinet groups, or sub-populations

ean be observed, in agreement with Perkins &

MeGee (1995) and Diez (2007), who stress that the

possibilities of seed gemiination are redueed with

inereasing distanee from an adult plant, whieh eould

explain the diserete patehes of plants observed em-

pirieally and graphieally in this study.

Orehids that remained unidentified and eould

not be assigned to a ehorologieal type appeared to

have a random distribution as eompared to identi-

fied individuals assigned to a ehorologieal type. In

partieular, the ehorologieal types END, MED, AT-

MED and W-MED showed elear patterns of eolo-

nization (Fig. 7). However, the limited number of

individuals assigned to the other ehorologieal types

was insuffieient for analysis.

The prevailing winds eould play an important

role in the distribution of orehids, by transporting

the seed and influeneing the behaviour of pollina-

tors. Observing the distribution patterns of the or-

ehids, areas towards the east (right of the maps) are

more densely populated. Orehid seed eapsules ma-

ture and dehisee during the dry season, from May

to September, and the prevailing winds at that time

blow towards the north-east (Albaria, 2011), favou-

ring population inerease and eolonization in that di-

Distribution and ecological patterns of orchids in Monte Pellegrino Reserve, Palermo (Sicily, Italy)

381

UNIDENTIFIED END.

^ »

■

E-MED.

AT-MED.

■ 7 *— ’ '

• t ”

»•

* ft

EE-MED. MED.

* ftftA«ftft

•r. f:

W-MED. S-MED.

EAST

NORTH

NORTH EAST

WEST

FULL SUN

SOUTH WEST

(CLIFF)

SOUTH

SOUTH/SOUTH

WEST

SOUTH EAST

£• = 2

— (JidoWKr)

{JWot-JUr)

(Jidot-JUr)

400

800

Figures 7-11. Distribution and Ecological Patterns of Orchids in Monte Pellegrino Reserve, Sicily. Fig. 7: Spatial distribution

of orchids by chorotype. Fig. 8: distribution of orchids within the 9 habitats that characterize the Monte Pellegrino Reserve.

Fig. 9: distribution of orchids according to light exposure. Fig. 10: Correlation function, Spatial dispersion and test of Ran-

dom Labeling, for the distribution of Endemic (END) chorotype orchids and the habitat Clearings in deciduous thermophilic

forest (Cl.DTF). Fig. 11: Correlation function. Spatial dispersion and test of Random Labeling, for Endemic (END) cho-

rotype orchids and the habitat Ampelodesmetum Meadows (MA), in the Monte Pellegrino Reserve, Sicily.

rection, but ultimately limited by the seafront. It is

possible that these populations had their origin in

areas that now eorrespond to the eity of Palermo

towards the south. Very few orehids were found

towards the western side of Monte Pellegrino Re-

serve, not only beeause of the winds, but also be-

eause of unfavourable environmental eonditions,

as the western perimeter of the reserve is highly

disturbed, with extensive grazing of goats whieh

are destruetive to the vegetation, and invasion by

steppe grasses (Pennisetum setacei-Hyparrhenie-

tum hirtae), whieh in eombination with other spe-

eies of Pennisetum, out-eompetes other plant

speeies.

During the months of January to April almost all

angiosperms found in the study site are in flower

and at this time the prevailing winds blow towards

the south-east, whieh may serve to bring pollinators

from the deeiduous, thermophilie forest towards the

north, whieh is the most speeies rieh habitat in the

area. In the ease of hybridization, almost all the in-

dividuals were found elose to at least one parent,

eonsidering the possibility that the winds might

have brought pollinaria-earrying bees from popula-

tions of other orehid speeies towards the north.

The distribution of the orehids of Monte Pelle-

grino Reserve varies depending on the habitat in

question, and Figure 8 demonstrates that Cl.DTF,

382

V. Bertolini.A. Damon, J.Valle Mora & A. N. Rojas VelAzquez

i p

Cl.RPF, Cl.RPEF, MS, XNG and MA are the pre-

ferred habitats in Monte Pellegrino. The areas refo-

rested to pine, and XNG species are found towards

the north of the study area and are traditionally

given over to cattle pasture. Cows cause less da-

mage to vegetation than sheeps and goats and de-

posit substantial amounts of manure which

propitiate excellent growing conditions for orchids,

with favourable pH and C/N ratio (Diez, 2007) which

is particularly important in areas where the soil is lea-

ched and eroded with exposed rocks. The variable

topography and vegetation cover of the study area

affects light exposure and creates unusual microcli-

matic conditions. The results of this study demon-

strate that exposure to the east, north and full sun are

related to higher numbers of orchids (Fig. 9). Along

the limits of the reserve, exposure towards the north

and east also benefits from cool, humid air streams

coming off the sea. The areas in full sun are the flat

tops of the hills reforested to pine, and with XNG

species largely dedicated to cattle pasture. For the

END chorotype, there was a positive correlation with

Cl.DTF, C1.RPF and Cl.RPEF, XNG, MA and MS.

The correlation between END species and Cl.DTF is

positive, with a high probability of finding another

individual within a radius of approximately 250 m

(Fig. 10), for Cl.RPF there is a high probability of

Figures 12-18. Distribution and Ecological Patterns of Orchids

in Monte Pellegrino Reserve, Sicily. Fig. 12: Correlation

function, Spatial dispersion and test of Random Labeling

for Endemic (END) chorotype orchids and the habitat Clea-

rings in areas reforested with pine (Cl.RPF). Fig. 13: Cor-

relation function, Spatial dispersion and test of Random

Labeling, for Endemic (END) chorotype orchids and the ha-

bitat Clearings in areas reforested with a mixture of Pinus

and Eucalyptus (Cl.RPEF). Fig. 14: Correlation Function,

Spatial dispersion and test of Random Labeling, for orchids

of the Endemic (END) chorotype and the habitat Substeppe

xerophytic nitrophilic grassland (XNG). Fig. 15: Correlation

Function, Spatial dispersion and test of Random Labeling,

for orchids of the Mediterranean (MED) chorotype and the

habitat Clearings in areas reforested with pine (Cl.RPF).

Fig. 16: Correlation function, spatial dispersion and test of

Random Labeling, for orchids of the Mediterranean (MED)

chorotype and the habitat Ampelodesmetum Meadows

(MA). Fig. 17: Correlation function. Spatial dispersion and

test of Random Labeling for orchids of the Atlantic Medi-

terranean (AT-MED) chorotype Clearings in areas reforested

with Pinus (Cl.RPF). Fig. 18. Correlation function. Spatial

dispersion and test of Random Labeling for orchids of the

Atlantic Mediterranean (AT-MED) chorotype and the habi-

tat Ampelodesmetum Meadows (MA).

Distribution and ecological patterns of orchids in Monte Pellegrino Reserve, Palermo (Sicily, Italy)

383

finding another individual within smaller distanees,

approximately less then 50 m (Fig. 12), in the ease

of Cl.RPEF the distanee is estimated at 100-350 m

(Fig 13) and finally, in XNG the distanee is estima-

ted at 350 m (Fig. 14). For the ease of MA habitats,

the positive eorrelation presents a random distribu-

tion. This may be due to the faet that the total area

is limited and fragmented, so the distribution pat-

tern may be aeting on a larger seale (Fig. 11).

The distribution of the END speeies is peeuliar,

but predietable, in that this is the only group with a

positive eorrelation with the habitat Cl.DTF, whieh

eonsists of residual fragments of the original forest

of the Monte Pellegrino area and it is logieal that

the endemie speeies would have an affinity for this

older, eonserved habitat where mutualist speeies are

more likely to be present. Similarly, the fraetion of

soil explored by the hypogeal parts of the orehids

benefit from relatively elevated quantities of orga-

nie matter derived from the proeesss of humifiea-

tion of the leaf litter from the native trees, implying

an elevated and speeifie eryptogamie flora and

fauna, of whieh rhizoetonious fungi are of partieular

interest. This situation would be benefieial for the

seeds of orehids, more than the situation oeeurring

in disturbed areas with alien speeies and different

dynamies of organie matter reeyeling.

None the less, the END ehorotype was also po-

sitively eorrelated to other habitat types, sueh as

XNG, due to the faet that some of the endemie spe-

eies were numerous, for example O. explanata, had

57 individuals (Fig. 5), whieh were also distributed

in these habitats. These speeies are probably less

demanding and eolonize nutrient poor habitats, un-

like endemies sueh as O. lunulata with only 3 indi-

viduals and all found in Cl.DTF. The MED

ehorotype is positively eorrelated to Cl.RPF, with a

high probability of finding another individual wi-

thin a radius of 50 m (Fig. 15). This is the ehorotype

with the widest distribution and ineludes the most

abundant speeies, O. bombiliflora, with a total of

1294 individuals (Fig. 4). No positive eorrelation

was found with MA (Fig. 16). The non-random di-

stribution assigned to the MED ehorotype is justi-

fied by the high number of individuals, in eontrast

to the small numbers of individuals for the speeies

in the END ehorotype, distributed within the same

habitat with a random distribution but positively

eorrelated to the habitat. The AT-MED ehorologieal

group displays a positive eorrelation with the

Cl.RPF with a high probability of finding another

individual within 100 m (Fig. 17). No sueh eorre-

lation was found with the MA (Fig. 18).

In eonelusion, the results shown are interesting for

eonservation purposes of Mediterranean orehids, be-

eause eontribute to understand distribution patterns

of Orehidaeeae family in eoastal environments. Mo-

reover, we ean apply this kind of knowledge to ana-

lyze other Natura2000 reserves that are elose and

similar to Monte Pellegrino, in terms of mieroelimate

and geographieal patterns as, for example, Monte

Gallo or Monte Catalfano in order to better under-

stand Orehidaeeae distribution at a larger seale.

ACKNOWLEDGEMENTS

We thank Dr. S.A. Giardina for taxonomieal de-

termination of the uneertain individuals.

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Biodiversity Journal, 2012, 3 (4): 385-396

Updated checklist of flora of the satellite islets surrounding

the Maltese Archipelago

Jeffrey Sciberras', Arnold Sciberras^* & Luca Pisani^

'24 'Camilleri Court' flat 5, il-Marlozz Str, Mellieha, Ghadira, Malta; email: wildalienplanet@gmail.com

^133 'Amest', Arcade Str, Paola, Malta; email: bioislets@gmail.com

^ 9, Milller Str, Sliema, Malta; email: luca.pisani@hotmail.com

*Corresponding author

ABSTRACT The present study provides an updated list of flora species encountered in multiple visits

carried out during 2010-2012 to the satellite islets surrounding the Maltese Archipelago.

KEY WORDS Flora; Satellite Islets; Maltese Archipelago.

Received 11.05.2012; accepted 18.11.2012; printed 30.12.2012

Proceedings of the P' International Congress “Insularity and Biodiversity”, May 11*-13*, 2012 - Palermo (Italy)

INTRODUCTION

Although the Flora of the Maltese islands has

been studied extensively from a taxonomie perspee-

tive, little is known regarding the distribution and

demographie status of most speeies. Moreover, re-

strieted or much less accessible areas have often

been poorly studied with the consequence that few

published records have been made available. Very

few works mention the minor satellite islets. John

Borg (1927) records 23 species of flowering plants

occurring on Selmunett Island. A study published

on Selmunett Island in 1983 by The Society for the

Study and Conservation of Nature (SSCN) (Lan-

franco, 1983) records 90 species and Sciberras &

Sciberras (2009) record 2 species for the site and

additional 7 species in a subsequent paper (Sciber-

ras & Sciberras, 2010). Camilleri (1990) includes

in a children’s article a preliminary list of the flora

of Tac-Cawl Rock. Cassar and Lanfranco provided

a preliminary list (unpublished) of plant species col-

lected by themselves on Haifa islet and Tac-(2awl

Rock, along with Stevens and Schembri.

A number of floral species are mentioned for se-

veral sites in the book on the natural environment of

the Maltese islands (Lanfranco, 2002); in 2007 one

of the authors (AS) along with Sdravko Lalov recor-

ded floral species for Fungus Rock (Sciberras, 2007;

Sciberras & Lalov, 2007; Sciberras, 2008). Recently,

Sciberras & Sciberras (2010) gave a detailed study

recording the majority of species as new records for

the respective locations, including the distributions

of various species found and a general description

of topography of each islet presented for the first

time. Other data for Tac-Cawl and Tal Haifa are pu-

blished by Mifsud (2011). Sciberras & Sciberras

(Unpublished Malta Environment Planning Autho-

rity MEPA report 2012) gave a detailed report of

present biodiversity on Fungus Rock.

The present work aims at bringing up to date the

latest records known of floral species observed for

the first time from these locations. These records

are based on unpublished or overlooked works of

the authors and personal communications. Most of

them were not present in Sciberras & Sciberras

(2010) work because proof or further identification

was required. To this day some are still undergoing

thorough investigation in order to study in detail

their taxonomy and distribution. Other species were

recently discovered. Some minor amendments to

the previous work were also made. This includes

386

J. SCIBERRAS, A. SCIBERRAS & L. PiSANI

corrections in taxonomy as well as species thought

not to be present but rediscovered after the 2010

work. The flora of Filfola and Manoel Island were

not included in this work because they are still

under study by the authors. Of the other islets, Fil-

flett, Fessej Rock, Ghallis Rocks, Ghemieri Rocks

Hnejja Rocks, Bear Rocks, Crocodile Rock, White

Rock of Marsalfom, Devil's End Rock and Xrobb

1-Ghagin Rock are constantly inundated and some-

times submerged by wave action during rough wea-

ther and therefore they do not support terrestrial

vegetation. Other boulders and stacks do exist but,

as far as in our knowledge, they are uninhabited.

MATERIALS AND METHODS

Multiple seasonal visits were carried out except

to those islets/rocks which require a legal permit to

visit as, for example. Fungus Rock, which in 2012

was visited once. Those were visited only in the

available restricted period. The location was gene-

rally divided in virtual transects to facilitate the

counting of species and individuals. Several species

were photographed for later identification perfor-

med by the authors and generally checked after-

wards with other local and foreign botanists.

Study area and flora

The Maltese archipelago consists of three main

islands, Malta, Gozo (Ghawdex) and Comino

(Kemmuna) together with a number of minor satel-

lite islands, islets and rocks. These smaller islets are

listed below in Table 1 and appear according to their

abbreviation letter in the Maltese archipelago maps

(Figs. 1-3). Following is the checklist of flora of the

Satellite Islets surrounding the Maltese Archipelago

(Table 2).

Figure 1. Map showing Malta and its satellite Islets/Roeks. Figure 2. Map showing Gozo and its satellite Islets/Roeks. Fi-

gure 3. Map shows Comino and its satellite Islets/Roeks (Maps by the authors).

updated checklist of flora of the satellite islets surrounding the Maltese Archipelago

387

ENGLISH NAME

MALTESE NAME

CODE

Malta's nearby islets

Malta (in Maltese)

Filfola

Filfla

Filflette

Il-Gebla ta’Xutu

Cheirolophus Roek

Il-Hagra tas-Sajjetta

Devil's End Roek

Il-Gebla tax-Xifer 1-Infem

Xrobb 1-Ghagin Roek

It-Taqtiegha

Manoel Island, but peninsula sinee

1750

Il-Gzira ta’ Manoel

Ghallis Roeks

Il-Gebla ta' Ghallis

Qawra Point or Ta’ Fra Ben islet

Il-Ponta/ Ras il- Qawra

(marked in map as “Red”

Islets/Roeks)

Selmunett Island/ St. Paul’s Island

Il-Gzira Ta’ San Pawl

Commons nearby islets

Kemmuna (in Maltese)

Old Battery' s Roek

Il-Gebla ta' taht il -Batterija

Lantern Point Roek

Il-Gebla Tal-Ponta 1-Irqiqa

Comino Cliff Faee Roek/ Pigeon

Rook

Il-Gebla ta' taht il-Mazz

Small Blue Lagoon Roek

Il-Hagra Ta’ Bejn il-Kmiemem iz-

Zghira

Large Blue Lagoon Roek

Il-Hagra Ta’ Bejn il-Kmiemem il-

Kbira

Ghemieri Roeks

L-Iskolli Ta’ 1-Ghemieri

(marked in map as “Green”

Islets/Roeks)

Com in otto

Kemmunett

Gozo's nearby islets

Ghawdex (in Maltese)

Barbaganni Roek

Il-Gebla tal-Barbaganni

Haifa Roek

Il-Gebla tal-Halfa

Hnejja roeks

Il-Gebel tal-Hnejja

Tac-Cawl Roek

Il-Gebla tac-Cawl

Fessej Roek

Il-Gebla tal-Fessej

Fungus Roek/ General’s islet

Il-Gebla tal-General

(marked in map as “Blue”

Islets/Roeks)

Croeodile Roek and Bear roeks (3

roeks in total)

White Roek-

Il-Gebla tal-Baqra u 1-Gebel tal-Or-

sijiet

Il-Gebla tal-Ghar Qawqla

Table 1 . List of satellite islets of the Maltese Islands surveyed in the present study.

388

J. SCIBERRAS, A. SCIBERRAS & L. PiSANI

SPECIES

SATELLITE ISLETS OF THE MALTESE ISLANDS

CHI JKLNMOPQSU

Aetheorrhiza bulbosa

Allium ampeloprasum

Allium commutatum

** * * *

Allium lojaconoi

*9 * *9

Allium melitense

*9 *

Aloe vera

Agave amerieana var. amerieana

Agave amerieana var. variegata

Agave sisalana

Ajuga iva

Ajuga pseudoiva

Anaeamptis eollina

Anaeamptis pyramidalis

* * *

Anaeamptis urvilleana

** ** *

Anagallis arvensis

* * ** *

Anthemis urvilleana

* ** ** * * *

Anthyllis hermanniae

subsp. melitensis

* *

Anthyllis vulneraria

** *

Arisarum vulgare

** ** ** ** ** **

Arthroenemum maerostaehyum

* * * * * *

Atraetylis gummifera

Arum italieum

Asparagus aphyllus

•k i: k k k k k k k

Asphodelus aestivus

k k k

Asteraceae sp.

Astragalus hamosus

* *

Avena sterilis

Beilis annua

updated checklist of flora of the satellite islets surrounding the Maltese Archipelago

389

SPECIES

SATELLITE ISLETS OF THE MALTESE ISLANDS

CHI JKLNMOPQSU

Beta maritima

Blackstonia perfoliata

*9 *

Bituminaria bituminosa

*9 *

Brachypodium retusum

Bromus madritensis

* * *

Bromus sp.

Borago ojficinalis

Catapodium marinum

Capparis orientalis

* **** ***

Carlina involucrata

** * *

Centaurea melitensis

Centaurium erythraea

Centaurium pulchellum

*9 *

Ceratonia siliqua

Cheirolophus crassifolius

Chamaerops humulis

Chenopodium murale

Chiliadenus bocconei

(Jasonia bocconei)

Cremnophyton lanfrancoi

Cichorium spinosum

** *

Crithmum maritimum

* * * * *

Crucianella rupestris

* * * ** * * *

Convolvulus althaeoides

Convolvulus lineatus

Convolvulus oleifolius

** * * * *

Coronilla scorpioides

Cuscuta epithymum

Cutandia maritima

390

J. SCIBERRAS, A. SCIBERRAS & L. PiSANI

SPECIES

C H

SATELLITE ISLETS OF THE MALTESE ISLANDS

I JKLNMOPQ

Cynara cardunculus

Cynodon dactylon

Cynomorium coccineum

Darniella melitensis

Dactylis hispanica

Daucus gingidium

** *

Daucus rupestris

Desmazeria marina

Desmazeria pignattii

Desmazeria rigida

Dittriehia graveolens

Dittriehia viseosa

Eeballium elaterium

Echium arenarium

Echium parviflorum

•

Erodium malacoides

* *

Euphorbia exigua

var. pycnophylla

Euphorbia dendroides

Euphorbia melitensis

Euphorbia peplus

Euphorbia pinea

Evax pygmaea

Fumaria officinalis

Fedia graciliflora

Ferula communis

* *

Ficus carica

Foeniculum vulgare

Frankenia hirsuta

updated checklist of flora of the satellite islets surrounding the Maltese Archipelago

391

SPECIES

SATELLITE ISLETS OF THE MALTESE ISLANDS

CHI JKLNMOPQSU

Frankenia pulverulenta

Galactites tomentosa

it it it it

Geranium rotundifolium

Gynandriris sisyrinchium

it it it

Halimione portulacoides

Hedypnois rhagadioloides

Hedysarum spinosum

Hedysarum coronarium

Helichrysum melitense

Hippocrepis biflora

itit itit

Hippocrepis multisiliquosa

Hypericum aegypticum

it it it it

Hypericum triquetrifolium

Hyoscyamus albus

Hyoseris frutescens

Hyoseris scabra

Iris pseudopumila

itit itit

Iris sicula

Limbarda crithmoides

ititititititititititititit

Lagurus ovatus

itit it

Lavatera arborea

(Malva dendromorpha)

* *

Leontodon tuberosus

Limonium melitense

it itit it it it it it it it

Limonium virgatum

it it it it it

Limonium zeraphae

Linaria pseudolaxiflora

*9 *

Linum strictum

itit it it

Linum trigynum

392

J. SCIBERRAS, A. SCIBERRAS & L. PiSANI

SPECIES

SATELLITE ISLETS OF THE MALTESE ISLANDS

CHI JKLNMOPQSU

Lobularia maritima

* * **

Lonicera implexa

Lotus cytisoides

* ** * * *

Lotus edulis

* *

Lotus ornithopodioides

Lygeum spartum

* * *

Matthiola incana

subsp. melitensis

* * *

Malva sp.

Malva parviflora

Medicago sp.

Medicago sp.2

MeUlotus sp.

Mercurialis annua

Mesembryanthemum nodiflorum

* ** * *

Muscari comosum

Narcissus serotinus

** ** ** *

Narcissus tazetta

* ** ** *

Nauplius aquaticus

(Astericus aquaticus)

Olea europaea

Ononis mitissima

Ononis ornithopodioides

** *

Ononis sieberi

Opuntia sp.

Opuntia ficus-indica

Opuntia stricta

Ophrys bombyliflora

Ophrys melitensis

Orchis coriophora

(Orchis fragrans)

updated checklist of flora of the satellite islets surrounding the Maltese Archipelago

393

SPECIES

SATELLITE ISLETS OF THE MALTESE ISLANDS

CHI JKLNMOPQSU

Ornithogalum arabicum

Ornithogalum narbonense

Orobanche sp.

** ** ** *

Orobanche cernua

Orobanche cf. densiflora

Orobanche ramosa

subsp. mutelii

Oxalis pes-caprae

Pancratium maritimum

Pallenis spinosa

* *

Parapholis filiformis

* ** ** *

Parapholis incurva

Parietaria cretica

Parietaria judaica

Periploca angustifolia

** * *

Phagnalon graecum

subsp. ginzbergeri

** * * *

Pistacia lentiscus

* * * *

Plantago afra

Plantago coronopus

* *

Plantago lagopus

* ** *

Prasium majus

** ** * *

Prospero autumnale

** ** ** ** *

Polypogon maritimus

subsp. subspathaceus

Reseda lutea

Rostraria cristata

Romulea columnae

Romulea ramiflora

Reichardia picroides

* *

Rhodalsine geniculata

394

J. SCIBERRAS, A. SCIBERRAS & L. PiSANI

SPECIES

SATELLITE ISLETS OF THE MALTESE ISLANDS

CHI JKLNMOPQSU

Rubia peregrina

Ruta chalepensis

** * **

Sagina apetala

Satureja microphylla

Scabiosa maritima

Scorpiurus muricatus

*9 * *

Scilla sicula

Sedum caeruleum

* *

Sedum caespitosum

Sedum litoreum

** ** * *

Sedum rubens

*9 **

Sedum sediforme

Smilax asp era

Seneeio bieolor

(Jacobaea maritima)

* * * *

Seneeio leueanthemifolius

* ** ** ** *

Sherardia arvensis

*9 **

Sideritis romana

Silene sedoides

* * * ** * * *

Spergularia sp.

Spergularia boeeonei

Spergularia marina

Sporobolus pungens

Solanum villosum

Sonehus tenerrimus

** * * *

Sonehus oleraeeus

** * * *

Suaeda vera

Teuerium frutieans

* * *

Theligonum eynoerambe

updated checklist of flora of the satellite islets surrounding the Maltese Archipelago

395

SPECIES

SATELLITE ISLETS OF THE MALTESE ISLANDS

CHI JKLNMOPQSU

Thymbra capitata

*7 * * * *7

Tordylium apulum

*7 **

Trachynia distachya

* *

Trifolium scabrum

"k kk

Trifolium stellatum

*7 *

Umbilicus horizontalis

Urginea pancration

k kk k k k kk

Urospermum picroides

Valantia hispida

* *

Valantia muralis

kk k kk k k

Total number of species for

each location observed

6 18 130 1 2 17 24 22 89 1 63 61 30

Table 2. Checklist of Flora of the Satellite Islets surrounding the Maltese Archipelago. Each letter represents the location, as in

Table 1 . ** = newly recorded species; * = species recorded in past works and observed by the authors; *? = species recorded

in past works but not observed by the authors.

RESULTS AND CONCLUSIONS

In this work a total of 205 species of plants were

recorded from 13 islets/rocks including Selmunett

having the highest variety with a total number of

130 species whilst Old Battery’s Rock and

Barbaganni Rock both containing the least variety

with I species. This is clearly due to the topography

and size of the sites. A total of 35 species are new

for Salmunett Islet's species list when compared to

Sciberras & Sciberras (2010), while 87 new species

were recorded considering the all islets/rocks.

It is rather unusual that conspicuous species

such as Scil/a sicula Tineo ex Guss. and Iris sicula

Tod. were not recorded before in previous writings.

Although Iris pseudopumila Tineo is rather com-

mon in the surrounding area such as in Selmun and

must be native to the island of Selmunett, Iris sicula

may have been recently introduced to the island as

it is popularly cultivated in many areas in Malta, be-

sides being planted in several private gardens and

parks such as Bahar ic -Caghaq; populations such

as those of Mellieha and those of Comino are

known to have been introduced in the late 1980's

and early 1990's . Related species were also planted

in Mgarr and Majjistral Park. In addition, other

conspicuous species like Pancratium maritimum L.,

Matthiola incana subsp. melitensis Brullo et al. and

Parietaria judaica L. went unnoticed until Sciber-

ras & Sciberras (2009; 2010) either due to their ra-

rity on site, season ideal for identification or where

they are located. Several other species especially

large alien species were also not recorded before

these writings. Also a very interesting observation

noted through this study is that the biodiversity of

these satellite islands tends to be somewhat diffe-

rent from the closest area on the mainland.

The main islands are often influenced and alte-

red by human activity, whilst these islets, being les-

ser known and often more difficult to access, have

remained virtually untouched. The biodiversity of

the islets depends on the specialization, adaptation.

396

J. SCIBERRAS, A. SCIBERRAS & L. PiSANI

(such as small, succulent or hairy leaves and growth

strueture) and natural seleetion of only the hardiest

speeies to survive in these extreme, unrelenting en-

vironments with limited land area and thin soils,

eonstant sea spray and the eontinuous exposure to

strong winds. The loeation of an islet or roek, toge-

ther with its topography and size plays an important

role in speeies diversity, number and distribution.

Speeies’ lists generally vary with different authors,

possibly due to a speeies being mistakenly identi-

fied or, in the time between different publieations,

populations may have gone extinet or overlooked.

Whilst on eertain sites very few or no new ob-

servations ean be made, others, espeeially those

more inaeeessible or restrieted by legislation, hold

a high potential for future diseoveries. Sehembri et

al. (1987) already listed some of the loeations men-

tioned in this work as loealities with eonservation

value even without giving any speeies for most

sites. We hope that the present work and future ones

to follow will aid in the proteetion and further raise

the value of their eonservation status for these sen-

sitive eeologieal gems.

ACKNOWLEDGEMENTS

The authors are indebted to Romario Seiberras,

Esther Seiberras, Matthew Borg Cardona, Simon

Sultana Harkins for their assistanee in field visits

and to Mario Gauei for his generous hospitality du-

ring Gozo field visits and assistanee in literature se-

areh. Edwin Lanfraneo is greatly aeknowledged for

eonfirming the identity of several speeies in this

work. Thanks also go to Mario Seiberras for provi-

ding numerous transport trips to various loeations

and staff members of Piseieulture Marine De Malta

Ltd for providing transportation several times to

Selmunett. MEPA is also aeknowledged for issuing

permits to visit the proteeted sites.

REFERENCES

Borg J., 1927. Descriptive Flora of the Maltese islands.

Government Printing Office, Malta.

Camilleri A., 1990. IL- Gebla Tac-Cawl. Il-Ballotra vol

1,5-6.

Lanfraneo E., 1983. The Flora of St. Paul’s Islands. Po-

tamon, 2: 23-31.

Lanfraneo S., 2002. L-Ambjent Natural! tal-Gzejjer Mal-

tin. Kullan Kulturali (45). Publikazzjoniet Indipen-

denza, Malta, 196 pp.

Mifsud S., 201 1 . A floristic survey on the Gozitan islets

of Tac-Cawl and Tal-Halfa in the Maltese islands.

MaltaWildPlants.com Online Publications (Ref:

MWPOP-OOl), Article Update Version 3.0(relea-

sed on 5-Dec-ll). URL: http://www.MaltaWil-

dPlants. com/pub l/index.php#W0 1

Sehembri P.J., Sehembri S., Lanfraneo E., Farrugia P. &

Sultana J., 1987. Localities with Conservation Value

in the Maltese Islands: Environment Division, Mini-

stry of Education, Malta.

Seiberras A., 2007. Lizards at Id-Dwejra. Dwejra Heritage

Park Gozo, Dwejra Management Board, p. 28-33.

Seiberras A. & Lalov S.V, 2007. Notes on the impact of

the black rat (Rattus rattus L.) on the flora and fauna

of Fungus Rock (Maltese Islands). The Central Me-

diterranean Naturalist, 4: 204-207.

Seiberras J., 2008. IL - Fejgel. Il-Ballottra issue 12. Na-

ture Trust Malta publications, 192 pp.

Seiberras J. & Seiberras A., 2009. Notes on the distri-

bution of Helichrysum melitense, Hyoseris frute-

scens and Matthiola incana melitensis in the

Maltese islands. The Central Mediterranean Natu-

ralist, 5: 28-34.

Seiberras J. & Seiberras A., 2010. Topography and

Flora of the Satellite islets surrounding the Maltese

Archipelago. The Central Mediterranean Naturalist,

5: 31-42.

Biodiversity Journal, 2012, 3 (4): 397-399

Flora of ‘‘U Briantinu”, a satellite stack of Panarea Island,

Aeolian Archipelago (Sicily, Italy)

Arnold Sciberras' &Jeffre)' Sciberras^

433 'Amest', Arcade Str, Paola, Malta; e-mail: bioislets@gmail.com

^24 'Camilleri Court' flat 5, il-Marlozz Str, Mellieha (Ghadira), Malta; e-mail: wildalienplanet@gmail.com

ABSTRACT This present study provides a list of flora species encountered during a visit to “U Brian-

tinu” one of the satellite stacks of Panarea Island in the Aeolian archipelago (Sicily, Italy).

KEY WORDS Flora; U Briantinu stack or Scoglio Brigantino; Aeolian Archipelago.

Received 11.05.2012; accepted 13.07.2012; printed 30.12.2012

Proceedings of the International Congress “Insularity and Biodiversity”, 11*-13* May 2012, Palermo, Italy

INTRODUCTION

The authors have been for the past decade or so

visiting small islands, islets and stacks around the

Mediterranean to collect data on the insular biodi-

versity of such sites also including the study of

these micro habitats and generally in comparison to

close by or satellite main islands. Such studies have

been intensely carried out in works like Sciberras

& Sciberras (2010). ‘U Briantinu Stack' or as com-

monly known by the locals as ‘Scoglio Brigantino'

is an interesting location which is quite inaccessible

to visitors. Its geomorphology is simply stunning in

being such a narrow vertical cliff face stack.

Slightly tilted to the south, thus facing north,

the stack has a length of approximately 32 me-

ters from west to east/east to west and a width no

more than a maximum of 5 meters from north to

south/south to north. It is estimated to be about

12-15 meters high from the sea level to the hi-

ghest part and 28 meters away from the closest

land excluding 2 rocks which do not support ter-

restrial life. It is situated in a pocket beach known

as ‘Gala Junco’.

Close by is a popular archeological site known

as the prehistoric village that dates back to the

Bronze Age (XIV - XIII century B.C.) No past lite-

rature is available for this stack at least regarding

that of natural history (P. Lo Cascio personal com-

munication). From a geological point of view, the

volcanic rock of the stack formed during the 4^^^ Tyr-

rhenian eruptive epoch, roughly 124,000 to 1 18,000

years ago. Composition of the rock is ‘andesitico -

dacitica calcalcalina’ and high in potassium (Calan-

chi et al., 2007).

MATERIALS AND METHODS

The authors visited the stack on 28.IX.2011.

They swam to the site and circulated the bottom of

the stack to take rough measurements of the stack

by 50 ft measuring tape. These measurements were

later compared with Google earth and thus the

rough measurements above were produced. Attempt

was made to climb the stack but the top of the stack

is practically inaccessible at least from sea or to

amateur climbers.

From every spot that climbing was succeeded,

numerous images were taken by a water proof ca-

mera (Fuji fine Fix Z33 wp) to photograph habitats

and wildlife. To have a better image of all the stack

the authors walked all the mainland coast of Pana-

rea island that circulate the stack and photographic

398

Arnold Sciberras & Jeffrey Sciberras

Figures. 1-3. “U Briantinu” a satellite staeks of Panarea Island (Aeolian Arehipelago, Sieily, Italy).

shots of all parts of the stack were taken with a

zoom of 560 mm by a camera (Canon powershot

SxlO) so identification was carried out both visual

and from images.

RESULTS

The following 9 species of macrophytic flora

were recorded:

Helichrysum litoreum Guss. (Compositae)

Centaurea aeolica Lojac. (Asteraceae)

Dianthus nipicola subsp. aeolicus (Lojac.) Brullo &

Miniss. (Caryophyllaceae)

Hyoseris taurina Martinoli (Asteraceae)

Dactylis glomerata L. (Poaceae)

Limbardia crithmoides (L.) Dumont (Asteraceae)

Erica arborea L. (Ericaceae)

Pistacia lends cus L. (Anacaridiaceae)

Daucus rupestris Guss. (Apiaceae)

Flora of“U Briantinu” a satellite stack of Panarea Island, Aeolian Archipelago (Sicily, Italy)

399

The dominant species on top of the stack seem

to be Helichrysum litoreum, Dactylis glomerata

and Centaurea aeolica, while dominant species on

cliff side are Limbardia crithmoides and Daucus

rupestris. Counting the specimens was virtually

impossible as such parts of site were only presen-

ted by images.

CONCLUSIONS

It is interesting to note that while the mentioned

species are dominant on the stack, the nearby

mainland area known as Milazzese was noted to

be dominated entirely by Cistus monsepeliensis L.,

(Cistacae) accompanied to a much lesser extent by

Pistacia lentiscus, Erica arborea and Calycotome

infesta Guss. (Fabaceae).

Although no terrestrial fauna survey took place

during visit a specimen of Lespisma sp. (Thysa-

nura Lespismatidae) was noted but could not be

collected for further identification. Same goes for

a specimen of Formicidae sp., and Lepidoptera

Colias crocea (Geofifoy, 1785), Pieris rapae (Lin-

naeus, 175), Pieris brassicae (Linnaeus, 1758)

(Pieridae) and Vanessa eardui (Linnaeus, 1758)

(Nymphalidae) were all noted feeding on Limbar-

dia erithmoides. The site was also checked for any

typical herpeto faunal activity but from the limita-

tion encountered during climbing it was not pos-

sible to speculate well the site; however the

authors were informed that no herpetofauna are

known from the stack (P. Lo Cascio personal com-

munication).

ACKNOWLEDGEMENTS

The authors are indebted to Pietro Lo Cascio

(Lipari, Italy) for his generous hospitality during

the Aeolian archipelago field visits and assistance

in literature search. The authors would also like to

thank Alan Deidun (Malta) for making such a trip

possible.

REFERENCES

Calanchi N., Lo Cascio R, Lucchi F., Rossi P.L. &

Tranne C.A., 2007. Guida ai Vulcani e alia Natura

delle Isole Eolie. LAC, Firenze, 467 pp.

Sciberras J. & Sciberras A., 2010. Topography and

Flora of the Satellite islets surrounding the Maltese

Archipelago. The Central Mediterranean Naturalist,

5: 31-42.

Biodiversity Journal, 2012, 3 (4): 401-406

Biodiversity and evolution of the dendroflora in the Medi

terranean

Pasquale Marino', Giuseppe Castellano' & Rosario Schicchi^

'Dipartimento di Scienze e Tecnologie Biologiche, Chimiche e Farmaceutiche, Universita degli Studi di Palermo, via Archirafi 38

90123 Palermo, Italy

^Dipartimento di Seienze Agrarie e Forestali, Universita degli Studi di Palermo, via Arehirafi 38 - 90123 Palermo, Italy

*Corresponding author:

ABSTRACT Sediment The main old representatives of the Mediterranean dendroflora, their origin and di-

stribution are treated. Relevant threats and strategy for in situ and ex situ eonservation are also

discussed here.

KEY WORDS Biogeography; dendroflora; endemism; insularity.

Received 11.05.2012; accepted 21.12.2012; printed 30.03.2013

Proceedings of the P' International Congress “Insularity and Biodiversity”, May 11*-13*, 2012 - Palermo (Italy)

INTRODUCTION

The Mediterranean basin is one of the 25 biodi-

versity hotspots identified at a global level to whieh

the storage is of essential importanee (Myers et al.,

2000). It is rieh in islands of all extensions, from

Sieily with its 25,700 square kilometres down to the

pebble size of those; other large islands are Sardinia

(24,090 sq km), Cyprus (9253 sq km). Corse (8748

sq km), Crete (8258 sq kiu), Baleares (4,996 sq

km), Malta (316 sq km). This basin represents the

remnant of the aneient Tethys Sea, a Mesozoie

oeean that underwent profound ehanges during the

Alpine orogeny, beginning in the Middle Cretaee-

ous, about 100 Ma (Gradstein & al., 2004), and ex-

tending to the late Mioeene, about 7 Ma.

Palaeoelimatie researehers studying the late Mio-

eene (Messinian, 7. 2-5. 3 Ma) have identified an

event of regional aridity, during whieh most of the

Mediterranean Sea beeame a marsh (Kovar-Eder et

al., 2008). Thereafter, during the Plioeene (5. 3-1. 8

Ma), the Mediterranean seasonality and the regional

eooling aeeentuated around 4.5, 3.6, 3.2, 2.8 and 2.4

Ma (Sue, 1984; Kovar-Eder et al., 2008; Jost et al..

2009). These elimatie oseillations, ending with the

Pleistoeene glaeiations, resulted in the eumulative

loss of several thermophilie speeies from the Euro-

pean eontinent, as well as in episodie expansions of

xerophytie eommunities (Pignatti, 1978; Sue, 1984;

Palamarev, 1989). Palaeoelimatie analysis suggests

the establishment of the eurrent Mediterranean eli-

mate seasonality, eharaeterized by two intra-annual

stress periods for plant growth, in summer and in

winter, during at least three elimatie erises dated to

3.2, 2.8 and 2.4 Ma (Sue, 1984; Fauquette et al.,

1999, 2007; Bmeh et al., 2006). By 10,000 BP eo-

niferous forests dominated by Pine and Juniper spe-

eies were oeeurring; by 5000 years BP deeiduous

trees of oak, elm, hornbeam beeeh ete., were beeo-

ming dominant (Willis & MeElwain, 2002).

THE MEDITERRANEAN DENDROFLORA:

A FOCUS ON ANCIENT WOODY SPECIES

Cupressaceae. This family appeared in the

Triassie (200 Ma) with the genus Wriddingtonia

that at now only in South Affiea oeeurs. During Eo-

402

P. Marino, G. Castellano & R. Schicchi

cene (55 Ma) Tetraclinis and Cupressus emerged in

the Mediterranean dendroflora (Palamarev, 1989).

Tetraclinis articulata (Vahl) Mast, is the only one

taxon of the Callitroideae subfamily that is spread

in the northern hemisphere, in Moroeeo, Southern

Spain, and Malta (Fig. 1). The genus Cupressus in-

eludes C. atlantica Gaussen and C. sempervirens L.

The former oeeurs in Moroeeo (High Atlas), the lat-

ter in Cyprus, Greeee and in the Balkan eoasts.

Pinaceae. This family, which appeared in the

Mesozoic, about 150 Ma, includes several genera

that are among the most important in the forestal

Mediterranean landscape: Abies, Cedrus dindPinus.

Nine species, one natural hybrid and several varie-

ties of Fir (Vidakovic, 1991) belong to the Mediter-

ranean dendroflora. Palaeoecological studies based

on fossil pollen and plant macrofossils show that

during the Pliocene (c. 5 Ma) the Mediterranean

Basin was covered by vast forest ecosystems, pre-

sumably including a common ancestor of the cur-

rent Mediterranean Firs (Pignatti, 1978; Meyen,

1987; Palamarev, 1989). From this common ance-

stor, migrations and subsequent population frag-

mentation led to smaller, isolated Fir forests around

the current Mediterranean Basin (Farjon & Ru-

shforth, 1989) (Fig. 2).

The genus Abies appears to have undergone si-

gnificant morphological differentiation that does

not necessarily imply reproductive isolation. Infact

long-term Mediterranean Basin dryness along a

south-eastern to north-western gradient may have

started a Miocene-Pliocene speciation sequence.

Pleistocene glacial cycles probably forced migra-

Figure 1. Formations with Tetraclinis articulata Mast., High

Atlas, Morocco.

tions leading to repeated contact between Fir spe-

cies in glacial refugia (Linares, 2011). In this con-

text is A. nebrodensis (Lojac.) Mattei, very rare with

only twenty-four mature individuals (Fig. 3).

Speciation of the genus Cedrus dates back about

58 Ma. Recent phytogeographic studies have revea-

led several sites of refuge in the Mediterranean

mountains during the Pleistocene (2 Ma) (Svenning

& Skov, 2005; Comes, 2004; Hellwig, 2004). Three

species belong to the Mediterranea dendroflora:

- Cedrus atlantica (Endl.) G. Manetti ex Car-

riere (Middle Atlas, Morocco) (Fig. 4)

- Cedrus libani A. Rich. (Lebanon and Turkey)

- Cedrus brevifolia Elwes & Henry (Cyprus)

The genus Pinus, the the richest in species

among all conifers, was already established in the

Cretaceous (145-66 Ma); in the Mediterranean it

includes:

- Pinus halepensis Mill, (widespread in the basin)

(Fig. 5)

- Pinus nigra Aiton

- subsp. nigra (Central Italy and Balkan area)

- subsp. calabrica (Loud.) A. E. Murray (Sicily,

Calabria, Corse) (Fig. 6)

- subsp. salzmannii (Dunal) Franco (Spain,

France)

- subsp. dalmatica (Vis.) Franco (Dalmatia)

- subsp. pallasiana Lamb. Holmboe (Romany,

Greece, Turkey)

- Pinus pinaster Aiton

- subsp. atlantica Villar (Atlantic coastlands of

Spain, France and Portugal)

- subsp. hamiltonii (Ten.) Villar. (Pantelleria and

Southern Spain)

- subsp. renoui (Morocco)

Angiosperms. Angiosperms, of prevalent Ter-

tiary origin, represent the largest group in the

world. In the Mediterranean basin they play a very

important role in forest and secondary shrub com-

munities showing high levels of specific diversity

(e.g. Acer, Quercus, Pyrus, Malus, Ulmus). Many

of these taxa are of remarkable interest. Among

these several very remarkable paleoendemics are

included, such as Zelkova sicula Di Pasquale,

Garfi et Quezel, which is confined in two very re-

stricted localities of Sicily (Marino & Spadaro,

2012) (Fig. 7).

Biodiversity and evolution of the dendroflora in the Mediterranean

403

Figure 2. Hypothetical post-glacial expansion of^. alba based on molecular data and fossil records, and present distribution

and diversity of the Mediterranean Abies species (Linares, 2011). 1) A. alba', 2) A. cilicica', 3) A. pinsapo', 4) A. numidica',

5) A. cephalonica', 6) A. bornmuelleriana', 1)A. nordmanniana', 8)^. equi-trojanv, 9) A. borisii-regis (A. • borisii-regis = A.

alba •A. cephalonica)', 10)^. nebrodensis', W) A. pinsapo var. maroccana', 12) A. pinsapo var. tazaotana.

THREATS

Fire, pasture and invasion of alien plants are

the main traits affeeting the Mediterranean den-

droflora (Table 1). Inelusion of eongenerie taxa to

the native ones in reforestation projeets is the most

serious threat of biologieal pollution and repre-

sents an important faetor in the genetie erosion

(e.g. Fraxinus excelsior subsp. siciliensis Ilardi et

Raimondo and Abies nebrodensis in Sieily)

(Sehieehi & Marino, 2011).

Exotie fauna is another faetor dangerous for the

wood renoval. A boar of Balkan and its eross in Si-

eily represent a very thi*eat for biodiversity eonser-

vation like in main proteeted areas.

From the phytopathologieal point of view the in-

trodution of alien speeies ean be identified as an im-

prtant souree of diseases for the native populations

(Sehieehi et al., 2008).

DISCUSSION

Mediterranean Islands possess aneient taxa da-

ting baek to Triassie (over 200 Ma) sinee Eoeene

(55 Ma); these are mainly Gymnosperms like

Abies, Cedrus, Cupressus, Finns and Tetraclinis.

This group suffers the highest risk, due the eeolo-

gieal eompetition and human threats. A relevant

example is represented by Abies nebrodensis that

in Sieily is loeated in a restrieted area dominated

by Fagus sylvatica and where several exotie firs

were introdueed in the past.

Cenozoie flora, also known as Terthiary flora,

represent for the Mediterranean area a biggest

group of woody speeies that dominate from the

level of sea to the high mountains. Acer, Ulmus,

Fagus, Quercus are the most important genera.

Other taxa are Pyrus, Mains, Sorbus that grow in

shrubby vegetation.

Preservation of Mediterranean dendroflora is of

vital importanee for the future of biodiversity.

Human aetivity has shaped these biologieal resour-

ees but tourism espeeially ean result in destruetive

deteriorattion. In addition, ehanging agrieultural po-

lieies, espeeially the EU ones, are likely to alterate

rural landseapes further.

Conservation strategies of speeifie habitats on a

regional seale is required to preserve biodiversity.

In the Mediterranean Islands, strategies for in situ

and ex situ eonservation are widely shared trough

eolleetion of seeds and plants. An important role,

improuving eonservation strategy, is played by seed

404

P. Marino, G. Castellano & R. Schicchi

Figure 3. Abies nebrodensis in the native area, Madonie

Mountains, Sieily.

Figure 4. A huge Cedrus atlantica tree. Middle Atlas,

Moroeeo.

Figure 5. Natural Reserve "Pino d' Aleppo" the only one

indigenous station of Pinus halepensis in Sieily.

Figure 6. A monumental tree of Pinus nigra subsp. cala-

brica in the southern slope of Etna voleano (Sieily).

Figure 7. Zelkova sicula in the loeus elassieus, Iblei

Mountains, Sieily.

Biodiversity and evolution of the dendroflora in the Mediterranean

405

ISLANDS

AREA (KM^)

MAIN ENDEMIC ENDANGE-

RED TAXA

THREATS

SICILY

25708

Abies nebrodensis

Zelkova sicula

Pasture, alien sp., fire

SARDINIA

24090

Pinus pinea

Fire, human aetivities

CYPRUS

9253

Cupressus sempervirens

Cedrus brevifolia

Fire, pasture

CORSE

8748

Pinus nigra subsp. calabrica

Fire

CRETE

8258

Zelkova abelieea

Fire, pasture

BALEARES

4996

Pinus pinaster subsp. renoui

Fire, pasture

MALTA

316

Tetraelinis artieulata

Fire, pasture

Table 1. Examples of Mediterranean dendroflora taxa and their main threats.

banks and Botanic gardens in the main Islands.

These living eolleetions eonsist eenturies of know-

how and expertise that now means they play a key

role in plant eonservation. Many of these aetivities

eontribute to ex situ eonservation, but botanie gar-

dens also play an important role in in situ eonser-

vation.

ACKNOWLEDGEMENTS

University of Palermo Funds are gratefully aek-

nowledged.

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Biodiversity Journal, 2012, 3 (4): 407-428

Annotated checklist of the birds from Pantelieria Island (Sici-

lian Channel, Italy): a summary of the most relevant data, with

new species for the site and for Italy

Andrea Corso'*,Verena Penna^, Marco Gustin^ Igor Maiorano'^ & Piero Ferrandes^

'Via Camastra, 10-96100 Siracusa, Italy; email: voloerrante@yalioo.it

^Via della Fossa, 15 -00186 Roma, Italy

^LIPU, Dipartimento Conservazione, Via Trento, 49 - 43 122 Parma, Italy

"'Viale Sanzio, 13 - 34128 Triste, Italy

^ Via Cossyra, 16-91017 Pantelieria, Trapani, Italy

Corresponding author

ABSTRACT The updated annoted clieeklist of all the bird species recorded at Pantelieria island (Trapani,

Sicily, Sicilian Channel) up to May 2012 along with data on the status of the birds are reported.

The total number of species recorded is 26 1 , 43 of which are new for the islands compared to

the previous checklist. During our study, very rare species for the Italian fauna were recorded

including Semi-collared Flycatcher, Citrine Wagtail, Steppe Eagle, Daurian Shrike, Desert

Wheatear, Trumpeter Finch, Black Wheatear and Rufous Bush-Chat. Detailed data and avai-

lable documentation are reported for the most relevant records.

KEY WORDS Pantelieria; birds; checklist; interesting species.

Received 12.05.2012; accepted 03.09.2012; printed 30.12.2012

Proceedings of the P' International Congress “Insularity and Biodiversity”, May 11*-13*, 2012 - Palermo (Italy)

INTRODUCTION

The information on the birds of the island of

Pantelieria (Sieilian Channel, Italy) are fragmen-

tary and outdated (Fosehi, 1968; Moltoni, 1971,

1973; Massa, 1985; lapiehino & Massa, 1989; Lo

Valvo et al., 1993; Allegri, 2000), exeept for sparse

reeords in a reeent work dealing with the Sieilian

avifauna (Corso, 2005) and in the Atlas of verte-

brates of Sieily (AA.VV., 2008). Like in all small

islands, the fauna of Pantelieria is highly subjeet to

ehanges, even relevant: the status of the speeies

breeding or migrating is variable annually, and at

the same time new speeies are likely to eolonize,

mostly when faeing with highly mobile groups

sueh birds. During studies on migrating raptors at

Pantelieria on behalf of LIPU, in the period 2004-

2012, extensive data on all migratory and breeding

bird speeies were also eolleeted (Gustin, 2005-

2009; Premuda et al., 2007; Corso & Gustin, in

press a). With regard to breeding speeies, an over-

view summary of the data has already been provi-

ded by Corso & Gustin (in press b). Many other

observations and eolleeted data will be used for the

preparation of additional eontributions: partieularly,

a work about Laughing Dove Streptopelia senega-

lensis at Pantelieria Island and another about the

“Birds of the Sicilian Channel Islands”.

In this paper, therefore, we wish to report a sum-

mary concerning primarily the migratory species ob-

served, with greater attention to the rare and irregular

ones, to vagrants or to the species simply not reported

previously at Pantelieria. Finally, we also report the

complete updated checklist for the island.

408

A. CoRso.V. Penna, M. Gustin, I. Maiorano & P. Ferrandes

MATERIALS AND METHODS

As a preliminary basis, all the available litera-

ture on birds reeorded on the island was studied,

known data served as a starting point for aequiring

new ones. Field observations were eondueted pri-

marily in spring, and exaetly over the period April

20^^ to May 20* of the years 2004-2012, on behalf

ofLIPU.

Moreover, in 2005, a visit was eondueted during

Oetober 15* to 22“^^. In 2008, observations were

made during the period August 20* to September

20*, again on behalf ofLIPU. In 2010-2012 obser-

vations were made by one of us (PF) throughout the

whole year (the reeords we have taken into aeeount

for this period are limited to those supported by

photographie evidenee). For the observations we

used binoeulars and spotting seopes lOx and 20-

60x. The song and ealls of many migratory and

breeding speeies were reeorded on a digital platform

with professional supports. For most of the rarest

speeies, photographie doeumentation is available. In

the ease of speeies reeorded in Italy less than ten

times, whieh have to be submitted to the COI (Com-

missione Omitologiea Italiana, i.e. the "Italian Or-

nithologieal Committee"), we report when the

reeord has been already ratified or if is still being

assessed (pending). With regard to seientifie no-

menelature and taxonomy, we referred to historieal

works sueh Vaurie (1959; 1965) and to the reeent

ehanges to the last eheeklist of birds of Italy by

CISO-COI (Fraeasso et al., 2009). For eaeh obser-

vation diseussed, we have indieated in braekets the

name of the author (or authors) of the reeord (in

ease this eoineides with one or more of the authors

of this paper, only the first letter for name and sur-

name are reported: eg. AC for Andrea Corso).

Finally, for what eoneerns the speeies whieh

breed on the island, we regarded as surely breeding

only the ones observed attending a nest, or feeding

juveniles, pairs with reeently fledged offspring, thus

following all the eriteria adopted in the Sieilian

Atlas of vertebrates (AA. W., 2008), the same was

for the possible and probable breeding speeies. All

photos are by A. Corso where not speeified.

ABBREVIATIONS. A = vagrant: a species for

which only few observations (records) are known,

the letter A is followed by the number of records

known in the case those are well documented (or in

the case these are less than 10); ad = adult; AB = A.

Belvisi; AC = A. Corso; B = breeding; det = deter-

mination (when a bird has been a posteriori identi-

fied by a different person from the observer or the

photographer); f = female; im = immature; ind = in-

dividual or individuals; juv = juvenile; m = male;

Mr = regular migrant; Mi = irregular migrant (often

due to lacking data); MV = M. Vigano; N = a new

species for the Island not reported earlier by the pre-

vious annoted list by Moltoni (1973); PF = P. Fer-

randes; SB = in the case the entire population

breeding in the site is sedentary (as opposite to mi-

gratory breeder); ssp = subspecies; S = Summer; VP

= V. Penna; W = wintering; Wp = a bird only par-

tially wintering in the site; ? = doubt related to the

status of some species, then in brackets, for exam-

ple (B irr), the status is uncertain and not supported

by sure proofs proving it behind doubt.; 2CY= 2"^

calendar year.

RESULTS

In total, 261 species are now known for Pan-

telleria, 63 of which nest on the island, 48 with cer-

tainty (1 extinct), 5 probably and 10 possibly

(Corso & Gustin, in press b). Compared to the

checklist by Moltoni (1973), 43 species are new to

the island.

The most interesting data about migratory spe-

cies or vagrant are briefly compared with those

available in the literature. For the rare species, the

status in Italy is also provided. The new relevant

data collected during the study are reported and di-

scussed below, species by species; for these, num-

ber of known records are reported, with date, site

of observation, no. of individuals, sex and age,

name of the observer/s. For vagrants, the number of

known records for Italy is also reported.

A complete list of all species observed to date

on the island is provided and the most relevant re-

cords are reported.

CHECKLIST OF THE BIRDS FROM PAN-

TELLERIA ISLAND

ANSERIFORMES

ANATIDAE

1.01590 Anser albifrons (Scopoli, 1769) A-1

(11/2011)N

Annotated checklist of the birds from Pantelleria Island (Sicilian Channel, Italy)

409

2.01610 Anser anser (Linnaeus, 1758) Mr (?)

3.01730 Tadorna tadorna (Linnaeus, 1758) Mr, Wp

Anas penelope (Linnaeus, 1758) Mr, Wp

5 .0X^20 Anas strepera (Linnaeus, 1758) Mr

6.01840^1^05' crecca (Linnaeus, 1758) Mr, Wp

7.01860^1/705 platyrhynchos (Linnaeus, 1758) Mr, Wp

8.01890^/705 acuta (Linnaeus, 1758) Mr

9.01910^/705 querquedula (Linnaeus, 1758) Mr

10.01940 ^/705 c/y//oo^o (Linnaeus, 1758) Mr

1 1 .01950 Marmaronetta angustirostris (Menetries,

1832) A- 1 (09/2008)N

12.02260 Oxyura leucocephala (Seopoli, 1769) A-1

(07/1954)

Aythya ferina (Linnaeus, 1758) Mr, Wp (?)

\A.f)2f)20 Aythya nyroca (GiildenstMt, 1770) MrN

15.02030 Aythya fuligula (Linnaeus, 1758) Mi N

16.02210 Mergus serrator (Linnaeus, 1758) Mi

(?), Wp (?)

17.02230 Mergus merganser (Linnaeus, 1758) A-1

(12/1972)

GALLIFOPIMES

PHASIANIDAE

1 8.03570 graeea (Meisner, 1804) SB extinet

19.03700 Coturnix eoturnix (Linnaeus, 1758) Mr, B

GAVIIFORMES

GAVIIDAE

20.62 00030 Gavia aretiea (Linnaeus, 1758) A

(11/1969)

PODICIPEDIFORMES

PODICIPEDIDAE

21.00070 Taehybaptus rufieollis (Pallas, 1764) Mr,

Wp(?)

22.00090 Podieeps eristatus (Linnaeus, 1758) Mr,

Wp(?)

23 .00120 Podieeps nigrieollis (C. L. Brehm, 1831)

Mr, Wp

PROCELLARIIFOPIMES

PROCELLARIIDAE

24.00360 Caloneetris diomedea (Seopoli, 1769)

Mr, B

25.00462 Puffinus yelkouan (Aeerbi, 1827) Mr, B

HYDROBATIDAE

26.00520 Hydrobates pelagieus (Linnaeus, 1758)

Mr (?), B (?)

PELECANIFORMES

PELECANIDAE

27 .OOSSO Peleeanus onoerotalus (Linnaeus, 1758) A

SULIDAE

2^.001X0 Morus bassanus (Linnaeus, 1758) Mr, W

PHALACROCORACIDAE

29.00720 Phalaeroeorax earbo (Linnaeus, 1758)

Mr, Wp

30.00800 Phalaeroeorax aristotelis (Linnaeus,

1761)AN

CICONIIFORMES

APO^EIDAE

3X .0X220 Ardea cinerea (LimvdiQus, 1758) Mr

32. 0X2A0 Ardea purpurea (LimtdiQus, 1766) Mr

33 .0X2X0 Casmerodius albus (hirmsiQus, 1758) Mi

34.01 1 90 Egretta garzetta (Linnaeus, 1766) Mr; Wp

35 .0X0^0 Ardeola ralloides (Seopoli, 1769) Mr

36.01110 5//6 ///c// 5 /6/5 (Linnaeus, 1758) MiN

37.01040 Nyeticorax nyetieorax (Linnaeus, 1758) Mr

38.00980 Ixobryehus minutus (Linnaeus, 1766) Mr

39.00950 Botaurus stellaris (Linnaeus, 1758) Mi N

CICONIIDAE

40.01310 Cieonia nigra (Linnaeus, 1758) Mr

41.01340 Cieonia cieonia (Linnaeus, 1758) Mr

THPIESKIORNITHIDAE

42.1360 Plegadis falcinellus (Linnaeus, 1766) Mi

A3. OX AAO Platalea leucorodia X^irmdiQus, 1758 Mi

PHOENICOPTERIFOPIMES

PHOENICOPTERIDAE

44.01470 Phoenicopterus roseus (Pallas, 1811) Mr

410

A. CoRso.V. Penna, M. Gustin, I. Maiorano & P. Ferrandes

FALCONIFORMES

PANDIONIDAE

45.03010 Pandion haliaetus (Linnaeus, 1758) Mr

ACCIPITRIDAE

46.02310 Pernis apivorus (Linnaeus, 1758) Mr, B

(r.?; 1 ep.)

47.02390 Milvus milvus (Linnaeus, 1758) Mi

48.02380 Milvus migrans (Boddaert, 1783) Mr

49.02470 Neophron percnopterus (Linnaeus, 1758)

50.02560 Circaetus gallicus (Gmelin, 1788) Mr (er-

ratie?)

51.02600 Circus aeruginosus (Linnaeus, 1758) Mr

52.02610 Circus cyaneus (Linnaeus, 1766) M r

53.02620 Circus macrourus (S. G. Gmelin, 1771) Mr

54.02630 Circus pygargus (Linnaeus, 1758) Mr

55.02730 Accipiter brevipes (Severtzov, 1850) A-1

(05/2012) N (pending COI ratifieation)

56.02690 Accipiternisus (Linnaeus, 1758) Mi, Bi ?

57.02870 Buteo buteo (Linnaeus, 1758) Mr, Wp, B

58.02880 Buteo rufinus (Cretzsehmar, 1827) ssp.

cirtensis (Mr, B); ssp. rufinus (A) N

59.02920 Aquila pomarina (C. L. Brehm, 1831)

Mr (?) N

60.02942 Aquila nipalensis (Hodgson, 1833) A-2

(05/2012) N

61.02950 Aquila heliaca (Savigny, 1809) A-1

(05/2010) N

62.02980 Aquila pennata (Gmelin, 1788) Mr, B

(SB?) N

FALCONIDAE

63.03030 Falco naumanni (Fleiseher, 1818) Mr

64.03040 Falco tinnunculus (Linnaeus, 1758) Mr, B

65.03070 Falco vespertinus (Linnaeus, 1766) Mr

66.03110 Falco eleonorae (Gene, 1839) Mr

67.03120 Falco concolor (Temminek, 1825) A-1

(05/2012) N (pending COI ratifieation)

68.03090 Falco columbarius (Linnaeus, 1758) Mi

69.03100 Falco subbuteo (Linnaeus, 1758) Mr, B

1 0.05\ AO Falco biarmicus (TQmmmc\s, 1825) ssp.

erlangeri A-1 ( 04/2006) N

71.03200 Falco peregrinus (Tunstall, 1771) Mr,

Wp, B; ssp. calidus Mr, Wi

GRUIFORMES

GRUIDAE

72.04330 Grus grus (Linnaeus, 1758) Mr

PGVLLIDAE

73.04070 Rallus aquaticus (Linnaeus, 1758) Mr,

Wp, B (i?)

74.04210 Crex crex (Linnaeus, 1758) Mi (?)

75.04100 Porzana parva (Seopoli, 1769) AN

76.04080 Porzana porzana (Linnaeus, 1766) A

77.04240 Gallinula chloropus (Linnaeus, 1758) Mr,

Wp, B (i?)

78.04290 Fulica atra Linnaeus, 1758 Mr, Wp

OTIDIDAE

79.04440 Chlamydotis undulata (Jaequin, 1784) A-1

(11/1911)

80.04460 Otis tarda (Linnaeus, 1758) A

81.04420 Tetrax tetrax (CixmsiQViS, 1758) A-1 (1967?)

CHARADRIIFORMES

HAEMATOPODIDAE

^2.04500 Haematopus ostralegus (Linnaeus, 1758) Mi

PIECURVIROSTRIDAE

^5.04550 Himantopus himantopus (Linnaeus, 1758) Mr

84.04560 Recurvirostra avosetta (Linnaeus, 1758) Mi

BURHINIDAE

85.04590 Burhinus oedicnemus (Linnaeus, 1758) Mr

GLAPIEOLIDAE

86.04650 Glareola pratincola (Linnaeus, 1766) M(i ?)

CHARADRIIDAE

87.04930 Vanellus vanellus (Linnaeus, 1758) Mr

88.04910 Vanellus gregarius (Pallas, 1771) (A-1,

09/1990) N

89.04850 Pluvialis apricaria (Linnaeus, 1758) Mr

90.04^60 Pluvialis squatarola (Linnaeus, 1758) Mr

91.04700 Charadrius hiaticula Linnaeus, 1758 Mr

92.04690 Charadrius dubius (Seopoli, 1786) Mr

Annotated checklist of the birds from Pantelleria Island (Sicilian Channel, Italy)

411

93.04770 Charadrius alexandrinus (Linnaeus, 1758)

Mr, Bi

94.04820 Charadrius morinellus (Linnaeus, 1758)

M(r ?)

SCOLOPACIDAE

95.05290 Scolopax rusticola (Linnaeus, 1758) Mr,

96.05180 Lymnocryptes minimus (Briinnieh, 1764)

97.05200 Gallinago media (Latham, 1787) A-2

98.05190 Gallinago gallinago (Linnaeus, 1758) Mr

99.05320 Limosa limosa (Linnaeus, 1758) Mi N

1 00.05340 lapponica (Linnaeus, 1758) AN

101.05380 Numenius phaeopus (Linnaeus, 1758)

M(r?)

\Q2. 05 A\0 Numenius arquata (Linnaeus, 1758) Mi

103.05450 Tringa erythropus (Pallas, 1764) Mr

104.05460 Tringa totanus (Linnaeus, 1758) Mr

105.05470 Tringa stagnatilis (Beehstein, 1803) Mi

106.05480 Tringa nebularia (Gunnerus, 1767) Mr

107.05530 Tringa ochropus (Linnaeus, 1758) Mr

108.05540 Tringa glareola (Linnaeus, 1758) Mr

109.05560 hypoleueos (Linnaeus, 1758) Mr

WO. 056X0 Arenaria interpres (Linnaeus, 1758) M(r?)

111.04960 Calidris canutus (Linnaeus, 1758) Mi

112.04970 Calidris alba (Pallas, 1764) Mr

113.05010 Calidris minuta (Leisler, 1812) Mr

114.05020 Calidris temminckii (Leisler, 1812) Mi

115.05090 Calidris ferruginea (Pontoppidan, 1763)

116.05120 Calidris alpina (Linnaeus, 1758) Mr

1 1 7.05 1 70 Philomaehus pugnax (Linnaeus, 1 758) Mr

1X^.05650 Phalaropus fulicarius (Linnaeus, 1758)

A-1 (08/1966)

STERCORARIIDAE

1 19.05690 Stercorarius skua (Briinnieh, 1764) Mi N

120.05660 Stereorarius pomarinus (Temminek,

1815) MiN

121.05670 Stereorarius parasiticus (Linnaeus,

1758) Mi

LAPaDAE

122.05880 Zarws' audouinii (Payraudeau, 1826) Mi

123.05910 Larus fuscus (Linnaeus, 1758) Mi

X2A. 05926 Larus michahellis (Naumann, 1840) SB,

W,Mr

X25 .05900 Larus canus (LXmvdiQus, 1758) A

126.05820 Chroicocephalus ridibundus (Linnaeus,

1766) Mr, Wp

127.05850 Chroicocephalus genei (Breme, 1840) Mi

128.05750 Larus melanocephalus (Temminek,

1820) Mr

129.05780 Hydrocoloeus minutus (Pallas, 1776) Mi

130.06020 tridactyla (Linnaeus, 1758) AN

STERNIDAE

131.06050 Gelochelidon nilotica (Gmelin, 1789) Mi

132.06060 Hydroprogne caspia (Pallas, 1770) Mi

133.061 10 Sterna sandvicensis (Latham, 1787) Mr,

134.06150 Sterna hirundo (Linnaeus, 1758) Mi

135.06240 Sternula albifrons (Pallas, 1764) Mi N

136.06260 Chlidonias hybrida (Pallas, 1811) MrN

137.06280 Chlidonias leucopterus (Temminek,

1815) MiN

138.06270 Chlidonias niger (Linnaeus, 1758) Mr

ALCIDAE

139.270 06540 Fratercula arctica (Linnaeus, 1758)

A-1 (1978)

COLUMBIFORMES

COLUMBIDAE

140.06650 Columba livia (van domestica) Gmelin,

1789 SB

141.06700 Columba palumbus (Linnaeus, 1758)

Mr, SB

142.06870 Streptopelia turtur (Linnaeus, 1758)

Mr, B

143.06840 Streptopelia decaocto (Frivaldszky,

1838) SB

144.06900 Streptopelia senegalensis (Linnaeus,

1766) SBN

CUCULIFOPIMES

CUCULIDAE

145.07160 Clamator glandarius (Linnaeus, 1758)

MiN

412

A. CoRso.V. Penna, M. Gustin, I. Maiorano & P. Ferrandes

146.07240 Cuculus canorus (Linnaeus, 1758) Mr,

B(r?)

STRIGIFORMES

TYTONIDAE

147.07350 Tyto alba (Scopoli, 1769) SB

STRIGIDAE

148.7390 Otus scops (Linnaeus, 1758) Mr, B(r?)

\A9 .^1 510 Athene noctua (Seopoli, 1769) M(i?),

(Bi?)

\50. 01 610 Asia otus (Linnaeus, 1758) B (SB?), Mr,

151.07680 Asio flammeus (Pontoppidan, 1763)

M(i?) N

CAPRIMULGIFOPIMES

CAPRIMULGIDAE

152.07790 Caprimulgus rufieollis (Temminek,

1820) A- 1 (5/2008) N (pending COI ratifieation)

153.07780 Caprimulgus europaeus (Linnaeus,

1758) Mr,B

APODIFOPIMES

APODIDAE

154.07980 Apus melba (Linnaeus, 1758) Mr, B

155.07950 Apus apus (Linnaeus, 1758) Mr, B

156.07960 Apus pallidus (Shelley, 1870) Mr, B

CORACIIFORMES

ALCEDINIDAE

157.08310 Aleedo atthis (Linnaeus, 1758) Mr, Wp

MEROPIDAE

158.08400 Merops apiaster (Linnaeus, 1758) Mr, Bi

Coraeiidae

159.08410 Coraeias garrulus (Linnaeus, 1758) Mr

UPUPIDAE

160.08460 Upupa epops (Linnaeus, 1758) Mr, (B?)

PICIFORMES

PICIDAE

161.08480 Jynx torquilla (Linnaeus, 1758) Mr,

(Bi?)

PASSERIFOPIMES

ALAUDIDAE

162.09610 Melanoeorypha ealandra (Linnaeus,

1766) Mi

163.09680 Calandrella braehydaetyla (Leisler,

1814) Mr, B

164.09720 Galerida eristata (Linnaeus, 1758) Mi

\ 65. 091 40 Lullula arborea (Linnaeus, 1758) A

\66.09160 Alauda arvensisdrmRQus, 1758 Mr

HIRUNDINIDAE

167.09810 Riparia riparia (Linnaeus, 1758) M r

1 68.099 1 0 Ptyonoprogne rupestris (Seopoli, 1769) Mi

169.09920 Hirundo rustiea (Linnaeus, 1758) Mr, Bi

170.09950 Cecropis dauriea (Linnaeus, 1771) Mr

1 7 1 . 1 00 1 0 Deliehon urbieum (Linnaeus, 1758) Mr,

MOTACILLIDAE

172.10200 Motacilla alba (Linnaeus, 1758) Mr, Bi

173.10180 Motacilla citreola (Pallas, 1776) A-3

(Mi?) N

\1 4. \0\10 Motacilla flava (Linnaeus, 1758) Mr, (Bi?)

175.10190 Motacilla cinerea (Tunstall, 1771) Mr, Wp

\1 6. \0050 Anthus campestris (Linnaeus, 1758) Mr, B

111.10020 Anthus richardi (Vieillot, 1818) A-1

(10/1955)

11 S. 10090 Anthus trivialis (Linnaeus, 1758) Mr

119.10110 Anthus pratensis (Linnaeus, 1758) Mr, Wp

1^0.10120 Anthus cervinus (Pallas, 1811) Mr

1 8 1 . 1 0 1 40 Anthus spinoletta (Linnaeus, 1758) Mr, Wp

RULIDAE

182.13140 Rulus rulus (Linnaeus, 1758) Mr, Wp

183.13150 Rulus ignicapilla (Temminek, 1 820) Mr,

PRUNELLIDAE

184.10840 Prunella modularis (Linnaeus, 1758)

Mr, Wp

Annotated checklist of the birds from Pantelleria Island (Sicilian Channel, Italy)

413

Figure l.Anser albifrons, Pantelleria, 11.2011 (PF). Figure 2. Buteo rufinus cirtensis, adult, Pantelleria. Figure 3. Falco pere-

grinus ssp, f ad nesting at Bagno di Venere (Pantelleria). Figure 4, 5. Gallinago media, Bagno di Venere, 15.05.2011 (AC &

O. Janni). Figure 6. Streptopelia senegalensis, Pantelleria (MV). Figure 7. Sylvia melanochephala, m ad, Pantelleria, spring.

414

A. CoRso.V. Penna, M. Gustin, I. Maiorano & P. Ferrandes

TURDIDAE

185.10950 Cercotrichas galactotes (Temminck,

1820)A-4 (Mi?)

186.1 0990 Erithacus rubecula (Linnaeus, 1 758) W,

Mr, B(r.?)

187.11040 Ei/5'cm/a megarhynchos (Brehm, 1831)

Mr, (B?)

188.11060 Luscinia svecica (Linnaeus, 1758) Mi

189.11210 Phoenicurus ochruros (Gmelin, 1774)

Mr, W

190.11220 Phoenicurus phoenicurus (Linnaeus,

1758) Mr

191.11370 Saxicola rubetra (Linnaeus, 1758) Mr

192.11390 Saxicola torquatus (Linnaeus, 1766) Mr,

Wp, (B ?)

193.11580 Oenanthe leucura (Gmelin, 1789) A-1

(04/2009) N

194.11460 Oenanthe oenanthe (Linnaeus, 1758) Mr

195.1 1480 Oenanthe hispanica (Linnaeus, 1758) Mr

196.11490 Oenanthe deserti (Temminek, 1825) A-1

(12/2011) N

197.11440 Oenanthe isabellina (Temminek, 1829)

A-1 (03/2012) N

198.11 620 Monticola saxatilis (Linnaeus, 1 766) Mr

199.11660 Monticola solitarius (Linnaeus, 1758)

Mr, B

200.11860 Turdus torquatus (Linnaeus, 1758) A-2

201.11870 Turdus merula (Linnaeus, 1758) SB, Mr

202.11980 Turdus pilaris (Linnaeus, 1758) Mi

203.12010 Turdus iliacus (Linnaeus, 1766) Mi, Wi

204.12000 Turdus philomelos (Brehm, 1831) W, Mr

205.12020 Turdus viscivorus (Linnaeus, 1758)N

SB, Mr

SYLVIIDAE

206.12260 Cisticola juncidis (Rafinesque, 1810)

ssp. cisticola SB; juncidis Mi, (Bi ?)

201 .123W Locustella luscinioides (Savi, 1824) A-1

(5/2009) N

20^. \2A\0 Acrocephalus melanopogon (Temminek,

1823) AN

209. 12430 Acrocephalus schoenobaenus (Linnaeus,

1758) Mr

210.12510 Acrocephalus scirpaceus (Hermann,

1804) Mr

2\\ A2530 Acrocephalus arundinaceus (Linnaeus,

1758) Mr

212.12590 Hippolais icterina (Vieillot, 1817) Mr

213. 12600 Hippolais polyglotta (Vieillot, 1 8 1 7) Mi

214.13120 Phylloscopus trochilus (Linnaeus, 1758)

215.13110 Phylloscopus collybita (Vieillot, 1817)

Mr, W, (B ?)

216.13070 Phylloscopus bonelli (Vieillot, 1819)

M(r ?)

217.13080 Phylloscopus sibilatrix (Beehstein,

1793) r

218.1 3000 Phylloscopus inornatus (Blyth, 1 842) A-2

(04/1931; 05/2012)

219.12770 Sylvia atricapilla (Linnaeus, 1758) Mr,

W, (B?)

220.12760 Sylvia borin (Boddaert, 1783) Mr

221.12740 Sylvia curruca (Linnaeus, 1758) Mi

222.12750 Sylvia communis (Latham, 1787) Mr

223.12640 Sylvia conspicillata (Temminek, 1820)

Mr, B(i?)

224.12620 Sylvia undata (Boddaert, 1783) SB,

(Wp, Mi?)

225.12610 Sylvia sarda (Temminek, 1820) SB extinet

226.12650 Sylvia cantillans (Pallas, 1764) Mr, B ?

227.12670 Sylvia melanocephala (Gmelin, 1789)

SB, Wp, Mr

228.12690 Sylvia rueppelli (Temminek, 1823) A-1

(06/1970)

MUSCICAPIDAE

229.13350 Muscicapa striata (Pallas, 1764) Mr,

B(r?)

230 A3490 Ficedula hypoleuca (Pallas, 1764) Mr

231.1 3480 Ficedula albicollis (Temminek, 1815) Mr

232. 1 3470 Ficedula semitorquata (Homeyer, 1 885)

A-5 (Mi?) N

233.13430 Ficedula parva (Beehstein, 1794) A-1

(04/2008) N

222.10990 Erithacus rubecula (Linnaeus, 1758) W,

M reg, B (r?)

223.11040 Luscinia megarhynchos (C. L. Brehm,

1831) Mr, (B?)

224.1 1060 Luscinia svecica (Linnaeus, 1758) Mi

225.10950 Erythropygia galactotes (Temminek,

1820) A-4

226.11210 Phoenicurus ochruros (S. G. Gmelin,

1774) Mr, W

Annotated checklist of the birds from Pantelleria Island (Sicilian Channel, Italy)

415

Figures 8-13. Bagno di Venere (Pantelleria). Figure 8. Motacilla citreola, m, 16.5.2008, Figure 9. Motacilla sp., juv,

9.9.2008, showing mixed eharaeters of citreola and flava. Figure 10. Motacilla flava ssp. Figure \ \. Motacilla flava ssp,

m, '"xanthophrys” (likely feldeggxl). Figure 12. Motacilla flava feldegg, m ad. Figure 13. Motacilla flava cinereocapilla,

m ad, spring.

416

A. CoRso.V. Penna, M. Gustin, I. Maiorano & P. Ferrandes

227.11220 Phoenicurus phoenicurus (Linnaeus,

1758) Mr

228.11370 Saxicola rubetra (Linnaeus, 1758) Mr

229.11390 Saxicola torquatus (Linnaeus, 1766) Mr,

Wp, B?

230.11580 Oenanthe leucura (Gmelin, 1789) A-1

(04/2009) N

23 1 . 11460 Oenanthe oenanthe (Linnaeus, 1 758) Mr

232.11480 Oenanthe hispanica (Linnaeus, 1758)

233.11490 Oenanthe deserti (Temminek, 1825) A-1

(12/2011) N

234.11440 Oenanthe isabellina (Temminek, 1829)

A-1 (03/2012) N

PARIDAE

234.14622 Cyanistes teneriffae l.QSSon, 1831 - ssp.

ultramarinus SB

PIEMIZIDAE

235.14900 Remiz pendulinus (Linnaeus, 1758) A

(Mi?)

OPaOLIDAE

236.15080 Oriolus oriolus (Linnaeus, 1758) Mr, (B?)

LANIIDAE

237.15150 Lanius collurio (Linnaeus, 1758) M(i?)

238.15140 Lanius isabellinus (Hemprieh & Ehren-

berg, 1833) A-1 (11/2011) N (pending COI ra-

tifieation)

239.15190 Lanius minor (Gmelin, 1788) A-1

(11/2011) N

240.15230 Lanius senator (Linnaeus, 1758) Mr, B?

CORVIDAE

241.15673 Corvus cornix (Linnaeus, 1758) MiN

242.15720 Corvus corax (Linnaeus, 1758) Mi N

STURNIDAE

243.15820 Sturnus vulgaris (Linnaeus, 1758) W,

Mr, (B ?)

244. 15830 Sturnus unicolor (Temminek, 1 820) Mr,

PASSERIDAE

245.15910 Passer (domesticus) italiae (Vieillot,

1817) SBN

246. 15920 Passer hispaniolensis (Temminek, 1 820)

Mr, Br

247.15980 Passer montanus (Linnaeus, 1758) Bi?,

FRINGILLIDAE

248.16360 Fringilla coelebs (Linnaeus, 1758) W,

249. Fringilla (coelebs) spodiogenys Mi (Bi?) N

250.16380 Fringilla montifringilla (Linnaeus,

1758) Mi, Wi

251.16490 Carduelis chloris (Linnaeus, 1758) Mr,

Wp, B(i?)

252.16540 Carduelis spinus (Linnaeus, 1758) W, Mr

253.16530 Carduelis carduelis (Linnaeus, 1758)

SB, Mr, Wp

254.16600 Carduelis cannabina (Linnaeus, 1758)

SB, Mr, Wp

255.16400 Serinus serinus (Linnaeus, 1766) SB(i ?),

Mr, Wp

256.17170 Coccothraustes coccothraustes (Linna-

eus, 1758) W, Mr

257.16760 Bucanetes githagineus (Liehtenstein,

1823)A-5

EMBERIZIDAE

258. 1 8660 Emberiza hortulana (Linnaeus, 1 758) A

(Mi?)

259.18770 Emberiza sehoeniclus (Linnaeus, 1758)

MrN

260. \S500 Plectrophenax nivalis (Linnaeus, 1758)

A-1 (10/1970)

261.18820 Emberiza calandra Linnaeus, 1758 SB

(r?), Mr, W

RELEVANT RECORDS OF THE BIRDS

FROM PANTELLERIA ISLAND

ANATIDAE

Anser albifrons (Seopoli, 1769)

White-fronted Goose. New speeies for the is-

land. Throughout November 201 1 , a 1 st W bird, ap-

Annotated checklist of the birds from Pantelleria Island (Sicilian Channel, Italy)

417

Figures 14-20. Pantelleria. Figure 14. Oenanthe deserti, m, Arenella, 2.12.2011 (AB). Figure 15. Ficedula sp., m, Bagno di

Venere, 22/27.4.2009, showing eharaeters of the taxa speculigeraliberiae. Figures 16, 17. Lanius ef isabellinus,}uw, loealita

Suvaki, 9.11.2011 (AB). Figure 18. Lanius senator senator. Figure 19. Fieedula albieollis, m ad, Bagno di Venere. Figure

20. Hirundo rustiea.

418

A. CoRso.V. Penna, M. Gustin, I. Maiorano & P. Ferrandes

parently of the nominate ssp, was observed along

the north eoast of the island.

Marmaronetta angustirostris (Menetries, 1832)

Marbled Duek. Not previously reported, a single

reeord known eoneerning 1 ind observed on

15.09.2008 at Bagno di Venere (AC). This speeies,

onee a very rare vagrant in Italy, sinee 2000 begun

to nest with some pairs in southern Sieily (provin-

ees of Agrigento and Siraeusa) (Briehetti & Fra-

easso, 2003), probably as a eonsequenee of the

inerease of the Tunisian population (Corso, 2005;

Isenmann et al., 2005). The number of breeding

pairs in Sieily is estimated in 4-8 pairs (Corso,

2005; pers. obs.). Never reeorded in all the other

surrounding Sieilian islands.

Aythya nyroca (Giildenstadt, 1770)

Ferruginous Duek. Not reported previously, nu-

merous observations are known in reeent years,

both during pre-breeding and post-breeding movi-

ments, although usually referred to single individual

or small floeks (max 6). Speeies breeding in Sieily

with a population of national relevanee, among the

most relevant in Europe (Corso & Janni, 2001;

Corso, 2005).

PHALACROCORACIDAE

Phalacrocorax aristotelis (Linnaeus, 1761)

Shag. New speeies for the island. 1 im observed

on 5.09.2007 at Pantelleria harbour (Corso et al.,

2009a).

ACCIPITRIDAE

Milvus milvus (Linnaeus, 1758)

Red Kite. Not reported by previous authors, we

obtained at least 5 reeords eoneerning 8 birds, du-

ring the period 2009-2012 (Gustin, 2009, 2012; AC

& VP). Of these, 3 ind were observed together on

5.04.2011, along with 8 blaek kites (T. La Mantia,

pers. eomm.). The observation of this speeies eros-

sing the Sieilian Channel, not mentioned by Corso

(2005), is very interesting. In other small islands of

the Sieilian Channel, like Pelagie islands, the spe-

eies is surely rarer or even a vagrant (Corso et al.,

2009b).

Neophron percnopterus (Linnaeus, 1758)

Egyptian Vulture. During the survey on raptors

migration, 1 ind was observed in 2006 (Gustin,

2006) and a floek of 4 birds was observed from 5*

to 7* May 2012 (AC & VP; Gustin, 2012). The re-

produetion of this speeies begins in Mareh- April;

the observation of more than 2 ind together is rather

rare during the pre-breeding migration in the Siei-

lian Channel, something instead more frequently

observed during the post-breeding migration

(Corso, 2005).

Circaetus gallicus (Gmelin, 1788)

Short-toed Eagle. Already mentioned by Moltoni

(1973) with some observations reported. In Spring

2004 to 2012, during the raptors migration survey

earned out on behalf of LIPU, 20 ind in total were

eounted, with a maximum of 10 in 2004 (Corso &

Gustin, in press a; Gustin, 2005-2009; Gustin,

2012). During pre-breeding migration, most short-

toed eagles (mainly the experieneed adults) tend to

avoid erossing the Sieilian Channel, preferring to

migrate through the Iberian-Tyrrhenian flyway

(Agostini et al., 2002; 2009). It remains unelear, ho-

wever, if individuals were observed at Pantelleria in

aetive migration moving north, or if they were sim-

ply erratie birds arrived from Tunisia, where the spe-

eies breeds in good number (Isenmann et al., 2005).

Buteo rufinus cirtensis (Levaillant, 1849)

Atlas Long-legged Buzzard. Pantelleria is today

the only national site where the speeies is known to

breed (Corso & Gustin, in press b). Corso (2005)

reported the presenee of a maximum of 2 1 ind in

spring, most of them being immature in 2CY.

AA. VV. (2008) reported the possible breeding

of this taxon. Corso (2009) eonfirmed the breeding,

mentioning a ease of a mixed pair Buteo buteo

buteo X Buteo rufinus cirtensis in May 2008, whieh

fledged one juv, and reporting two pure breeding

pairs of cirtensis. Sinee 2004, observed regularly

on the island with 1-2 pairs, in 2007 and 2008 one

pair was observed attending a nest in a large eavity

on a eliff of the south/south-east slope of Mt

Grande. Observed up to 2 pairs simultaneously.

From 2004 to 2012 adults were observed in territo-

rial display, breeding eourtship, breeding behavior

Annotated checklist of the birds from Pantelleria Island (Sicilian Channel, Italy)

419

Figure 21 . Calandrella brachydactyla, Pantelleria, note the extensive rusty-orange plumage with least marked dark pattern,

eloser to the ssp. rubiginosa rather than to the nominate one. Figure 22. Calandrella brachydactyla rubiginosa, Linosa, Pe-

lagian Island (1. Maiorano). Figure 23. Ixobrychus minutus, Pantelleria. Figure 24. Nycticorax nycticorax, Pantelleria. Figure

25. Ciconia ciconia, Pantelleria.

420

A. CoRso.V. Penna, M. Gustin, I. Maiorano & P. Ferrandes

and territorial behavior, and several juveniles were

observed too. The presenee of birds whieh are dif-

fieult to identify, showing intermediate eharaeters

with Buteo buteo, as well as the presenee of typieal

cirtensis mating with bird eloser in appearanee to

Common Buzzard Buteo buteo makes the pieture

very eomplex and diffieult to define. In Europe, the

only other areas where this distinetive taxon is bree-

ding are southern Spain (Tarifa, Gibraltar) and Por-

tugal, where mixed pairs or presumed mixed pairs

and presumed hybrids have been observed during

the last ten years (Elorriaga & Munoz, 2010; L.

Palma, pers. eomm.).

Buteo buteo vulpinus (Gloger, 1833)

Steppe Buzzard. Not reported historieally (Mol-

toni, 1973), in reeent years, observed with some

birds during pre-breeding migration in Spring 2004-

2011 (Corso, 2005; Gustin, 2005-2009; Corso &

Gustin, in press a). During spring 2012, along with

the largest passage ever reeorded of Honey Buzzard

for Pantelleria, a reeord number of 8 ind was reeor-

ded (AC & VP; Gustin, 2012).

Aquila pomarina (Brehm, 1831)

Lesser Spotted Eagle. Not reported by Moltoni

(1973). Observed annually during the raptors pre-

breeding migration survey eondueted on behalf of

LIPU. In partieular : I ind observed in Spring 2005,

2006, 2009, 2010, 2011, while 2 (3) in Spring 2012

(AC & VP; Gustin, 2005-2009; Gustin, 2012; Corso

& Gustin, in press a). During the survey on raptor

migration at Cap Bon, Tunisia, the speeies is regu-

larly observed in spring, with a substantial number

of birds, therefore it is not unexpeeted that some

birds reaeh Pantelleria (Corso, 2005; Isenmann et

al., 2005).

Aquila nipalensis (Hodgson, 1833)

Steppe Eagle. New speeies for the island and

for the Sieilian Channel islands; 2 observed during

the survey on raptor migration on behalf of LIPU

in the Spring 2012: I juv (in 2CY) on 04.05.2012

and I ad on 5.05.2012 (AC & VP; Gustin, 2012).

The speeies is a rare vagrant in Italy with less than

20 reeords (Briehetti & Fraeasso, 2003; Corso,

2005; EBN Italia). At Cap Bon, Tunisia, observed

regularly with several dozen indd., therefore, the

absenee of reeords on Pantelleria was surprising

(Corso, 2005).

Aquila heliaca (Savigny, 1809)

Imperial Eagle. A single reeord, relating to 1 im

(2/3CY) observed on 5* and 6* May 2010, at Mt.

Grande (AC & VP; Corso & Gustin, in press a). The

speeies is an irregular migrant in Italy (Briehetti &

Fraeasso, 2003; EBN Italia). In spring 2010, eoin-

eiding with an unpreeedented influx into Italy of

speeies from the Balkans, 4 or 5 other imperial ea-

gles were observed, 3 of whieh in Sieily (Ruggieri

&Nieoli, 2011).

Aquila pennata (Gmelin, 1788)

Booted Eagle. Breeding reeorded for the first

time in Italy at Pantelleria by Corso & Gustin (in

press b), still pending approval by the COL A pair

nesting on the island with eertainty sinee at least

2004, the first year of our study. The same pair oe-

eupied the same territory from 2005 to 2012, but in

reeent years, the adult white morph female has di-

sappeared being replaeed by an adult dark morph.

Regularly seen in spring in territorial and mating

display, in eourtship display, and other breeding re-

lated behaviors. During summer to autumn, the

adult pair was observed aeeompanied by juvenile

birds, obviously reeently fledged, forming family

party (AC, PF). In the Spring 2012, another pair,

both sexes of the white morph, has oeeupied the is-

land, maintaining a different territory than the first

pair. In August-September 2012, one pair was ae-

eompanying two fresh juveniles (PF).

FALCONIDAE

Falco biarmicus (Temminek, 1825)

Lanner Faleon. 1 adult of the ssp. erlangeri

(Kleinsehmidt, 1901), typieal of North Afriea, ob-

served and photographed repeatedly while hunting

over the airport of Pantelleria during the period 1

to May 2006 (Gustin, 2006; R. Gildi, per.

eom.). Less than four historieal reeords are known

for Italy (Briehetti & Fraeasso, 2003; Corso, 2005)

and one reeent observation at Siraeusa in April 2012

of an adult bird (AC & B.J. Small). The reeord ob-

Annotated checklist of the birds from Pantelleria Island (Sicilian Channel, Italy)

421

Figure 26. Glareola pratincola, Pantelleria. Figure 27. Milvus milvus, juv, Dammusi di Ghirlanda, Pantelleria. Figure 28.

Cyanistes teneriffae ultramarinus, Pantelleria (1. Maiorano).

422

A. CoRso.V. Penna, M. Gustin, I. Maiorano & P. Ferrandes

tained in Pantelleria is certainly the most detailed

report and the best doeumented one (Corso, 2005).

SCOLOPACIDAE

Gallinago media (Latham, 1787)

Great Snipe. A single historieal reeord (Moltoni,

1973). We have seen this speeies on a single oeea-

sion: 1 ind found exhausted at Bagno di Venere on

15.05.2011 (AC & O. Janni). The bird was pieked

up, re -hydrated, fed and released with sueeess in the

same site of diseovery. During the same period

(April-May 2011) was reeorded one of the most si-

gnifieant migratory influx noted in Italy in reeent

deeades (LBN Italia).

COLUMBIDAE

Streptopelia senegalensis (Linnaeus, 1766)

Laughing Dove. From the first, a nest was found

with eggs in 2004, followed until hatehing (Corso,

2005) and the reeord aeeepted by the COI (Janni &

Fraeasso, 2009); to date the speeies has shown a si-

gnifieant and fast inerease up to approx. 62-70 pairs

estimated on 2011 (Corso & Gustin, in press b;

Corso et al., unpublished data). The population on

the island is eertainly attributable to the ssp. phoe-

nicophila (Hartert, 1916) typieal of North Afriea

(Corso, 2005).

TYTONIDAE

Tyto alba (Seopoli, 1769)

Bam Owl. The taxonomie position of the bree-

ding population in Pantelleria and at the Pelagie is-

lands is not elear. Corso (2005) believes that the ssp.

erlangeri (Selater, 1921) eould be involved due to

the proximity to the Afriean eoast from where likely

the eolonizers arrived. However, Vaurie (1965) does

not report Tunisia as breeding ground for the ssp.

erlangeri, moreover Isenmann et al. (2005) aseribe

to the nominate ssp the population found in Tunisia.

On the eontrary, the taxonomie group AERC-TAC

(the Taxonomieal Committee of the Assoeiation of

the European Rarities Committee) ineludes North

Afriea as distribution range of the ssp. erlangeri.

Therefore the taxonomie status of the barn owls

breeding on the islands of the Sieilian Channel re-

mains unelear and deserves further in depth study.

STRIGIDAE

Athene noctua (Seopoli, 1769)

Little Owl. Corso (2005) makes no mention of

the presenee in Sieily of other taxon than the nomi-

nate ssp. noctua (Seopoli, 1769). Briehetti & Fra-

easso (2003) report that Sieilian population may

show sometimes intermediate eharaeters with the

ssp. glaux (Savigny, 1 809), from the North Afriean

eoast. In Pantelleria, the Little Owl is frequently

observed, although not regularly, and is eonsidered

an irregular breeder (Moltoni, 1973; Corso, 2005;

Corso & Gustin, in press b), while rare are the ob-

servations for the Pelagie islands (new work in

preparation). Given the proximity of the Tunisian

eoastland, it is not exeluded that individuals found

on the islands in the Sieilian Channel ean belong,

at least in part, to the ssp. glaux (Savigny, 1809)

(or even saharae Kleinsehmidt, 1909). The few in-

dividuals seen in daylight, indeed, showed a very

pale plumage, sandy eolored, with quite broad pale

markings, making the pale pattern rather striking

(AC). Therefore, the little owls of Pantelleria and

the Pelagie islands, should be studied more tho-

roughly in the future, possibly obtaining genetie

samples, reeordings of voealizations and morpho-

metrie data.

CAPRIMULGIDAE

Caprimulgus ruficollis (Temminek, 1820)

Red-neeked Nightjar. Not reported in other stu-

dies, we report a reeent observation, probably re-

ferring to the ssp. desertorum (Erlager, 1899) from

North Afriea. Along the southern slope of Mt.

Grande, 1 ind heard singing on 18.05.2008 and 1

ind, presumably different, observed along the road

a few hundred meters away from the first sight

(AC; Ruggieri & Nieoli, 2009). This report is pen-

ding approval by the COI. However, repeated at-

tempts to reloeate the speeies, by using play-baek,

have not sueeeeded, to date (AC, O. Janni, M.

Robb). Given the high density of breeding pairs

found in the promontory of Cap Bon, Tunisia (Isen-

mann et al., 2005), loeated 73 km from Pantelleria,

and the similarity in the habitat between the two

areas, its oeeasional nesting or its arrival is not sur-

prising. Future more in depth researehes are there-

fore advisable. The reeord, if approved by the COI,

will be the 3^^^^ known for Italy. There is one further

Annotated checklist of the birds from Pantelleria Island (Sicilian Channel, Italy)

423

recent record of this species in Lampedusa, where

1 bird of the ssp. desertorum was ringed on

12.05.2010, recently accepted (Janni & Fracasso,

in prep.)

ALAUDIDAE

Calandrella brachydactyla (Leisler, 1814)

Short-toed Lark. Taxonomic status of the bree-

ding population of Pantelleria is unclear and deser-

ves further studies. Corso (2005) reported the

nominate subspecies only for the breeding popula-

tion of Sicily. For Pantelleria, Moltoni (1973) re-

ports the nominate subspecies only, probably on

account of the taxon found all over Italy and wi-

thout a proper plumage analysis of the breeding

birds. The breeding population of Pantelleria and

Pelagic, appears to show characters usually asso-

ciated to the ssp. rubiginosa (Fromholz, 1913). Al-

ready Cova (1969) noticed that many birds from

these Sicilian islands (with one bird described and

illustrated collected at Linosa in May) showed cha-

racters of the ssp. rubiginosa, warning therefore fu-

ture studies to establish their taxonomical status.

Indeed, according to direct personal observations of

the birds breeding in Pantelleria and Pelagic, the

whole plumage is conspicuously more rusty orange

or c inn amon buff brown, warmer and brighter com-

pared to the breeding population of Sicily and Italy,

with the crown almost unmarked orange, reddish or

rusty-colored, the mantle being rusty-rufous tinged

with the darker pattern just visible, often the strea-

king being barely darker and least marked (chiefly

on the crown where often is not visible in the field),

all characters reported to be typical of the taxon ru-

biginosa (Vaurie, 1959; Svensson, 1992). For the

coastal and Northern Tunisia, Isenmann et al.

(2005) reported the nominate subspecies. For Malta

some authors report the nominate subspecies as

breeding (Sultana & Gauci, 1982; Sultana et al.,

2011) while others include the area within the di-

stribution range of rubiginosa (Vaurie, 1959; Sven-

sson, 1992), finally Fenech (2010) in his recent

extensive work reports that both birds showing cha-

racters of nominate race as well as others looking

closer to rubiginosa are observable in Malta. Wha-

tever this taxon is valid or not is questionable (L.

Svensson, pers. com.), but surely further research

are deemed to clarify the taxonomic status of the

birds found in the islands of the Sicilian Channel; a

comparison of series of birds from all over Italy and

from the Sicilian Channel is in preparation.

MOTACILLIDAE

Motacilla citreola (Pallas, 1776)

Citrine Wagtail. Not reported previously. In re-

cent years was observed three times, so that with

wider and more frequent ornithological coverage of

the island, probably the species would result an ir-

regular migrant or a scarce regular migrant rather

than a rare vagrant. Specifically, we have the follo-

wing data, all backed up by videos, sound recor-

dings and photos: 1 m and 1 f Bagno di Venere,

16.05.2008 (AC); 1 m Bagno di Venere, 05.08.2010

(M. Robb, AC & IM). Also observed two presumed

hybrids Motacilla citreola x Motacilla flava ssp.,

showing characters intermediate between the two

species: 1 f Bagno di Venere, 16/05/2008; 1 juv,

Bagno di Venere, 09/09/2008 (both AC).

TURDIDAE

Cercothricas galactotes (Temminck, 1820)

Rufous Bush-Chat. Two historical records - 1 on

5.05.1971 and 1 on 02.05.1976 (Corso, 2005). Du-

ring this study, two more records were obtained in-

cluding one bird on 22.04.2009 and one on

23.04.2009, in two different sites of Pantelleria

(AC; Ruggieri & Nicoli, 2010).

Turdus viscivorus (Linnaeus, 1758)

Mistle Thrush. Not mentioned by Moltoni

(1973), reported by Corso (2005) as breeding in the

island and confirmed by Corso & Gustin (in press

b). It nests with some pairs since 2004, no other

breeding data are known for small satellite islands

of Italy (Brichetti & Fracasso, 2008). The breeding

population on the Sicilian mountains is considered

attributable to the nominate subspecies (Corso,

2005; Brichetti & Fracasso, 2008). The ssp. dei-

chleri Erlanger, 1897, typical of North Africa, is li-

mited in Italy to Sardinia (Brichetti & Fracasso,

2008). The subspecies of the breeding pairs on Pan-

telleria is unknown but probably they have coloni-

zed the island from the African coast, so they would

belong to the ssp. deichleri. Future studies are de-

sirable to clarify their taxonomic position.

424

A. CoRso.V. Penna, M. Gustin, I. Maiorano & P. Ferrandes

Turdus torquatus (Linnaeus, 1758)

Ring Ouzel. 1 juv, subspeeies not identified, ob-

served at Mt.Grande on 17.09.2008 (AC). Known

previously for a single historieal reeord of 1 bird

eaptured in Oetober 1964 (Moltoni, 1973).

SYLVIIDAE

Locustella luscinioides (Savi, 1824)

Savi’s Warbler. A single reeord is known for

Pantelleria island. 1 ind, singing on 18.05.2009 at

Bagno di Venere (AC). Reeords of this speeies in

Sieily are very searee and irregular, with less than

13 known data (Corso, 2005; lentile, pers. eom.),

many of whieh obtained in reeent years at Pelagie

islands (Corso, 2005; Corso et al, unpublished

data).

Phylloscopus inornatus (Blyth, 1842)

Yellow-browed Warbler. An historieal reeord re-

fers to 1 bird on 09.04.1931 (Moltoni, 1973). New

reeord: 1 at Pantelleria town in a garden with Ta-

merisks, on 17.5.2012 (AC). Speeies observed more

frequently in reeent years on the islands surroun-

ding Sieily (ehiefly at Linosa and Lampedusa) and

in general elsewhere in Italy, so that it eould now

be eonsidered a regular migrant albeit searee (Bri-

ehetti & Fraeasso, 2010). In Pantelleria, it is still a

very rare vagrant, surely due to the laeking of omi-

thologieal eoverage during the best period of oeeur-

renee. In faet, virtually all reeords obtained in Italy

are from Oetober-November, while Spring reeords

remain oeeasional.

Sylvia sarda (Temminek, 1820)

Marmora’s Warbler. As suggested already by

Corso (2005), VV. AA. (2008) and reported by

Corso & Custin (in press b), we eould eonfirm the

extinetion of this speeies in Pantelleria. Although it

has been assiduously sought, in all suitable breeding

habitat and also by the use of playbaek, the speeies

was never eontaeted in 2004-2012. As opposite, the

speeies was reeently found in several oeeasion in

Lampedusa (Pelagie islands), where it should be

elarified if it is simply a wintering/wondering bird

or if there is a little and loealized breeding popula-

tion (Corso et al., unpublished data).

MUSCICAPIDAE

Ficedula semitorquata (Homeyer, 1885)

Semi-eollared Flyeateher. First data for Pantel-

leria, 5 reeords of 5 to 7 birds. 1 f at Bagno di Ve-

nere, on 02.05.2004 (AC; Ruggieri, 2005); 1 f on

28.04.2008, same loeality (Ruggieri & Nieoli,

2009) ; 1 m photographed along the banks of Bagno

di Venere, on 11-12.05.2009 (Ruggieri & Nieoli,

2010) ; 2 m on 24.04.2010, same site; 2 m again on

27.04.2010 (or same birds of the previous reeord ?).

In reeent years, the speeies has been reeorded regu-

larly in Sieily and Italy, though always in a very

small number (Briehetti & Fraeasso, 2008).

Ficedula parva (Beehstein, 1794)

Red-breasted Flyeateher. Not reported previou-

sly, we know a reeent observation referred to 1 m

on 26.04.2008 at Bagno di Venere (Ruggieri & Ni-

eoli, 2009). This speeies is a regular autumn mi-

grant at the Pelagie islands (Briehetti & Fraeasso,

2008; Corso et al, unpublished data).

Oenanthe leucura (Cmelin, 1789)

Blaek Wheatear. First reeord for the island refers

to 1 m observed near Suvaki, on 24.04.2009 (Rug-

gieri & Nieoli, 2010). In Italy, where it was histori-

eally breeding, nowadays it is a rare vagrant

(Briehetti & Fraeasso, 2008).

Oenanthe deserti (Temminek, 1825)

Desert Wheatear. Not reported previously. One

reeent reeord: 1 m (1st W?), photographed at Are-

nella, on 02.12.2011 (A. Belvisi, det AC). In Sieily,

12 additional reeords are known to date (Corso,

2005; personal data).

Oenanthe isabellina (Temminek, 1829)

Isabelline Wheatear. Not reported in the litera-

ture. First observation known to us is relative to 1

bird photographed near the town of Pantelleria on

18.03.2012 (PF), in eorrespondenee with one of the

major influx of this speeies ever reeorded in Italy

(EBN Italia). The speeies is a regular migrant in ea-

stern Sieily and the Pelagie islands, mueh less fre-

quent in the northern area of the Sieilian Channel

Annotated checklist of the birds from Pantelleria Island (Sicilian Channel, Italy)

425

and in western Sieily (Corso, 2005). With more eo-

verage during the period of Mareh-early April, the

speeies would probably be more often reeorded also

in Pantelleria.

LANIIDAE

Lanius isabellinus (Hemprieh & Ehrenberg, 1833)

Isabelline (Daurian) Shrike. The first reeord for

Pantelleria regards 1 juv observed and photogra-

phed on 9. 1 1 .20 1 1 near Suvaki, probably belonging

to the taxon isabellinus (A. Belvisi, det AC). This

is the 1 1* reeord for Italy of the Lanius isabellinus

eomplex, six of whieh were observed in Sieily

(Corso, 2005; Briehetti & Fraeasso, 2011). The at-

tribution to a eertain taxon of the Lanius isabellinus

eomplex (sensu latu) is extremely diffieult (Pear-

son, 2000; Worfolk, 2000; van der Laan & CDNA,

2008; Panov, 2009; 2011; Pearson et al., 2012) and

a review of all the Italian reeords would be desira-

ble, even more so in light of the splitting of the taxa

phoenicuroides and isabellinus into separate speeies

as proposed by several authors (Pearson, 2000;

Worfolk, 2000; van der Laan & CDNA, 2008;

Panov, 2009; 2011; Briehetti & Fraeasso, 2011; Pe-

arson et al., 2012).

Lanius minor (Gmelin, 1788)

Lesser Grey Shrike. Not mentioned by Moltoni

(1973). We have obtained a reeent reeord referring

to 1 bird in 1st W plumage photographed on

17.11.2011 at Bagno di Venere (A. Belvisi). The

speeies is an extremely searee migrant through Si-

eilian Channel, having a more easterly migration

(Corso, 2005; Briehetti & Fraeasso, 2011).

CORVIDAE

Corvus cor nix (Linnaeus, 1758)

Hooded Crow. First reeord for Pantelleria rela-

ted to 2 birds observed on 16.05.2004 (Corso,

2005). Afterward, 1-2 ind regularly observed in

spring 2005 to 2012. The subspeeies of the birds

observed, as well as that of the birds of the Sieilian

population (Corso, 2005) is uneertain and would

deserve better studies.

Corvus corax (Linnaeus, 1758) (ssp.?)

Raven. First reeord for Pantelleria, referring to

1 bird observed at Seauri in May 2004 (Corso,

2005). Subsequently, observed in at least four other

oeeasions sinee 2006 with 1-2 ind (AB, AC, PF, R.

Gildi, P. Console). The subspeeies of the birds ob-

served on the island is doubtful, it is in faet uneer-

tain if they were birds eoming from Sieily, therefore

belonging to the ssp. corax or hispanus Hartert &

Kleinsehmidt, 1901 (aeeording to authors), or eo-

ming from Tunisia, in this ease attributable to tin-

gitanus (Irby, 1874) (Corso, 2005; Briehetti &

Fraeasso, 2011). This latter taxon, is now eonside-

red a separate speeies by some authors (Baker &

Omland, 2006). All ravens observed on the islands

of the Sieilian Channel, should therefore be exami-

ned earefully, and ideally, for a positive determina-

tion, a good sound reeording of the voiee should be

obtained (M. Robb, pers. eomm.).

STURNIDAE

Sturnus unicolor (Temminek, 1820)

Spotless Starling. Reported historieally in 1955

(Moltoni, 1973). In reeent years, I to 4 ind observed

regularly at Arenella and Grazia, from 2005 to 2012

(AC, VP, PF; E. Vigo, pers. eomm.). Breeding was

suspeeted in 2011 and eonfirmed for the first time

in summer 2012, when two pairs of adult aeeompa-

nied by 6 freshly fledged juveniles were photogra-

phed at Kazen, near Arenella (AC, PF).

FRINGILLIDAE

Fringilla (coelebs) spodiogenys (Bonaparte, 1841)

Afriean Chaffineh. 1 observed in Pantelleria on

1.06.1987 (lapiehino & Massa, 1989). In reeent

years some males observed and also heard singing

at Mt. Grande, into the woodland area. In May

2009, heard up to 2 or 3 m singing and observed I

f (Ruggieri & Nieoli, 2010). Heard at least 1 m in

2008 and 2010 but none in 201 1 and 2012. The po-

tential habitat and proximity to Tunisia, as well as

the repeated observations in suitable period at least

sinee 1987, with males singing in several different

days, suggest a possible breeding on the island,

though not regularly. Data regarding nests or reeen-

tly fledged juveniles should be obtained in the fu-

ture in order to eonfirm its reproduetion on the

island.

426

A. CoRso.V. Penna, M. Gustin, I. Maiorano & P. Ferrandes

Bucanetes githagineus (Lichtenstein, 1823)

Trumpeter Fineh. Reported by Moltoni (1973)

with 2 birds on 25.10.1970, 2 more on 02.05.1971

and 3 on 5.05.1971, while 1 or 2 birds on

07.05.1971. New data: 2 speeimens photographed

together at Arenella, on 19.11.2010 (PF).

CONCLUSIONS

Contrary to Moltoni (1973), we deeided to ex-

elude from the eheeklist of Pantelleria two speeies:

Great Tit {Pams major Linnaeus, 1758) and Wren

{Troglodytes troglodytes Linnaeus, 1758). For the

former, Moltoni (1973) reports the observation by

Steinfatt of 21 birds, on 12*^ April 1931. It is possi-

ble that this observation is due to eonfusion with

the North Afriean Blue Tit {Cyanistes teneriffae

Lesson, 1831), whieh at that time was not known

to be present on the island and whose appearanee

ean be similar to Great Tit if eonseious attention is

not paid (Moltoni, 1971). Further, no other reeords

of Great Tit are known for the islands of the Sieilian

Channel islands. To support reeords of Wren, no de-

tailed observation is reported, exeept a few general

information provided by some loeal hunters.

ACKNOWLEDGMENTS

A warm thank to LIPU, in partieular to Elena

D' Andrea and Claudio Celada, for finaneial support

for all studies performed in Pantelleria, during

spring 2004 to 2012, as part of the study for the pro-

teetion of migratory raptors in the eentral Mediter-

ranean. Thanks to other members of the group

MISC for their assistanee during field observations

and the eompany during all the "birding expedition"

on the islands of Sieilian Channel: Ottavio Janni,

Hans Larsson, Lueio Manisealeo and Miehele Vi-

gano. Thanks to Luke, Carol, and Lillo Antonioni

for hospitality and logistieal support on the island.

Thanks to Andrea Belvisi for his photos and the in-

teresting observations provided as well as to Enzio

Vigo, Andrea Nieoli, Maurizio Sighele, Franeeseo

Omaghi and many other observers who kindly re-

ported their observations. Thanks to Brian J. Small

(LIMOSA Birding Holidays) and Magnus Robb

(“The Sound Approaeh to Birding”).

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Biodiversity Journal, 2012, 3 (4): 429-444

Catalogue of malgasy genera of Pselaphinae Latreilie, 1 802

and first data on the diversity of Pselaphid beetles popula-

tion in the Amber Mountain National Park, Northern Ma-

dagascar (Coleoptera Staphylinidae)

Giorgio Sabella*, Fabio M. Viglianisi & Ettore Petralia

Department of Biologieal, Geologieal and Environmental Seienees - seetion of Animal Biology "M. La Greea" of University, via

Androne 81, - 95124 Catania, Italy; e-mails: fabiovgl@uniet.it; ettorepetralia@gmail.eom

Corresponding author: sabellag@uniet.it

ABSTRACT The catalogue of the genera of Pselaphinae Latreilie, 1802 from Madagascar is here presented.

For each genus is given the bibliographic reference relative to the original description, its sy-

nonyms and the number of total known species. A review of current knowledge about Psela-

phinae from the National Park of Amber Mountain (Northern Madagascar) shows that for this

area are at present known 16 genera (14 endemic to Madagascar and one of Amber Mountain)

and 23 species, all malgasy endemic, 19 of which are known only for the Amber Mountain

area. During faunistic researchs carried out in this district, from 18 to 31 March 2011, were

collected seven genera (Faronitopsis, Trissemus, Leichotrella, Ctnenistes, Acylotyrus, Ei-

chiella and Rhynchoclaviger) reported for the first time for Amber Mountain area. The rese-

arch confirms the high biodiversity of malgasy Pselaphid fauna and some of its characteristics

as the coexistence in the same area of more congeners species, sometimes very similar to each

other, contrary to what occurs in temperate regions. Besides the classic environment of the

soil, in Madagascar there are many species that live on herbaceous vegetation or on the bran-

ches of trees, in particular those belonging to the tribe of Brachyglutini and Ctenistini.

KEY WORDS Catalogue of genera; Pselaphinae; Madagascar; Amber Mountain; Biodiversity.

Received 12.05.2012; accepted 08.12.2012; printed 30.12.2012

Proceedings of the LUntemational Congress “Insularity and Biodiversity”, May 11*-13*, 2012 - Palermo (Italy)

INTRODUCTION

The Malgasy Pselaphinae fauna is relatively

well-studied due to the works of Raffray (1890;

1897; 1898; 1900; 1903; 1904); Wassmann (1891;

1893a; 1897); Fairmaire (1898a; 1898b); Jeannel

(1954a; 1954b; 1956a; 1959b; 1960a; 1960b;

1961b), Leleup (1969; 1972; 1976a; 1976b; 1977),

Cells (1969; 1970), Dajoz (1982); Coulon (1986;

1989), Hlavac (2005) and Hlavac & Banaf (2012).

At present for Madagasear are known 128 ge-

nera of subfamily Pselaphinae Latreilie, 1802

(where 105, representing about 82%, are Malagasy

endemies) for a total of 417 speeies, with 415 of

them (equal to 99.5%) endemie to Madagascar.

Below is presented an exhaustive picture of the

current knowledge about pselafidofauna from Ma-

dagascar, according to the classificatory scheme

proposed by Newton & Chandler (1989) as modi-

fied by Bouchard et al. (2011). For each taxonomic

group (supertribes and tribes) are indicated the

number of genera and species, specifying the num-

ber of taxa endemic to Madagascar. For each genus

is given the bibliographic reference relative to the

430

G. Sabella, RM. Viglianisi & E. Petralia

original description, its synonym and the number of

total known species; with E are highlighted the ge-

nera endemic to Madagascar.

Genera o/Pselaphinae from Madagascar

Supertribus Faronitae Reitter, 1882

3 genera and 15 species all endemic to Madagascar.

F aronites iQdimvQl, 1954a: 153 (11 spp.) E

Faronitopsis Jeannel, 1960a: 54 (1 sp.) E

Parafaronus Jeannel, 1960a: 54 (3 spp.) E

Supertribus Euplectitae Streubel, 1839

36 genera, 29 endemic to Madagascar; 147 species

all endemic to Madagascar

Tribus Bithynoplectini Schaufuss, 1890

12 genera, 10 endemic to Madagascar; 26 species

all endemic to Madagascar.

Anozethopsis Jeannel, 1956a: 6 (2 spp.) E

Apozethopsus Jeannel, 1954a: 159 (3 spp.) E

Basilewskyozethus Leleup, 1977: 76 (1 sp.) E

Decazethodes Coulon, 1989:232(1 sp.)E

Microzethinus 1954a: 165 (1 sp.) E

Microzethopsis Jeannel, 1960a: 64 (1 sp.) E

Nesiozethus Jeannel, 1954a: 164 (3 spp.) E

Petalozethopsis Jeannel, 1960b: 6 (1 sp.) E

Protozethopsus Jeannel, 1954a: 162

(13 spp., 10 spp. endemic to Madagascar)

Protozethus Jeannel 1954b: 62 (misspelling)

Metazethopsis Jeannel, 1960a: 68

Trizethopsis Dajoz, 1982: 482 (1 sp.) E

Zethinomorphus Jeannel, 1960a: 66 (1 sp.) E

Zethopsiola Jeannel, 1954b: 86

(67 spp., 1 sp. endemic to Madagascar)

Tribus Euplectini Streubel, 1839

14 genera, 12 endemic to Madagascar; 70 species

all endemic to Madagascar.

Afroplectus Jeannel, 1952a: 130

(231 spp., 2 spp. endemic to Madagascar)

Afwplectidius Jeannel 1952b: 200 (subgenus)

Afroplectodes Jeannel, 1952b: 202 (subgenus)

Afroplectaualax Jeannel, 1959b: 202 (subgenus)

Aminosimus Raffray, 1898a: 267 (12 spp.) E

Plectasymus Jeannel 1960a: 80 (subgenus)

Anotimus Dajoz, 1982: 484 (1 sp.) E

Asymoplectidius Jeannel, 1956a: 11 (1 sp.) E

Asymoplectus Raffray, 1897: 55

(54 spp., 9 spp. endemic to Madagascar)

Asymoplectodes Jeannel 1960a: 75 (subgenus)

Newton & Chandler, 1989: 19

NEW NAME for Humbertiella Jeannel, 1960b: 7

(preocc., not Saussure, 1869) (1 sp.) E

Neoplectidius Jeannel, 1960a: 84 (1 sp.) E

Neothesiastes Jeannel, 1960a: 73 (2 spp.) E

Nesiotoplectus Jeannel, 1954a: 175 (15 spp.) E (in-

cluding Comore islands)

Omotimiotes Jeannel, 1954a: 170 (1 sp.) E

Pachyeuplectus Jeannel, 1954a: 179 (1 sp.) E

Paraphiliopsis Jeannel, 1959a: 120; 1960a: 70

(5 spp.) E

Plectiastes Jeannel, 1960b: 9(1 sp.) E

Plectodytes Jeannel, 1956a: 8 (2 spp.) E

Pseudozibus Jeannel, 1956b: 365

(5 spp., 1 sp. endemic to Madagascar)

Aphiliopsis Besuehet, 1956: 369

Trimiophanes Jeannel, 1954a: 168 (16 spp.) E

Tribus Trichonychini Reitter, 1882

8 genera, 7 endemic to Madagascar; 5 1 species all

endemic to Madagascar.

Ambalavoa Dajoz, 1982: 487 (1 sp.) E

Ankavena Jeannel, 1954a: 192 (2 spp.) E

Apotectus Newton & Chandler, 1989: 28

NEW NAME for Autoplectus Raffray, 1883: 248

(preocc., not Balsamo-Crivelli, 1843) (7 spp., 6 en-

demic to Madagascar); known only from Madaga-

scar and Angola

Badensia Jeannel, 1954a: 193 (13 spp.) E

Imeriniella Jeannel, 1960b: 13 (3 spp.) E

Imerinella Jeannel, 1960a: 110 (objective syno-

nym of Imeriniella Jeannel, 1960)

Ranavala Raffray, 1898b: 224 (21 spp.) E

Masoala Jeannel, 1954a: 190 (subgenus)

Fanovana Jeannel, 1960a: 100 (subgenus)

Ranavalidius Jeannel, 1959b: 204 (3 spp.) E

Ranavalodes Jeannel, 1960a: 99 (2 spp.) E

Supertribus Batrisitae Reitter, 1882

9 genera, 8 endemic to Madagascar; 36 species, 35

endemic to Madagascar.

Tribus Batrisini Reitter, 1882

Batrischema Newton & Chandler 1989: 34

NEW NAME for Batriso schema Jeannel, 1960a:

123 (preocc., not Reitter, 1883) (1 sp.) E

Malgasy genera of Pselaphinae and Pselaphid beetles population in the Amber Mountain, Northern Madagascar 43 1

Batrisochorus Jeannel 1949a: 146

(16 spp., 1 sp. not endemic to Madagascar)

Batristellus Jeannel 1949b: 154 (subgenus)

Batrisodella Jeannel, 1954a: 243 (2 spp.) E

Batrisodites Jeannel, 1954a: 244 (19 spp.) E

Batrisolius Jeannel, 1956a: 25

Batrisomina Raffray, 1903: 316 (11 spp.) E

Batrixenus Newton & Chandler, 1989: 35

NEW NAME for Batrisoxenus Dajoz, 1982: 495

(preocc., not Leleup, 1971) (1 sp.) E

Franzorella Leleup, 1977: 82 (1 sp.) E

Jochmansiella Leleup, 1976b: 307 (1 sp.) E

Macrodelphus Leleup, 1977: 84 (1 sp.)E

Supertribus Goniaceritae Reitter, 1882

13 genera, 10 endemic to Madagascar; 88 species,

87 endemic to Madagascar.

Tribus Brachyglutini Raffray, 1904

10 genera, 7 endemic to Madagascar, 81 species, 80

endemic to Madagascar.

Baxyridius Jeannel, 1954a: 210 (2 spp.) E

Bryaxella Raffray, 1903: 319 (1 sp.) E

Leiochrotella Jeannel, 1954a: 231 (3 spp.) E

Leiochrotidius deannel, 1960a: 161 (1 sp.) E

Madabaxyris Dajoz, 1982: 501 (1 sp.)E

Rabyxis Raffray, 1890: 124 (43 spp.) E

(1 sp. also in Comore islands)

Pseudobaxyris Jeannel 1960a: 129 (subgenus)

Reichenbachella jQamiQl, 1949b: 80

(19 spp., 1 sp. not endemic to Madagascar)

Reichenbachia Leach, 1826: 451

(340 spp., 20 spp. endemic to Madagascar)

Dierobia Thomson, 1859: 54

Dicrobia Reitter, 1882: 474 (misspelling of Dierobia)

Reichenbachius Casey, 1906: 359

Trissemus Jeannel, 1949a: 95

(130 spp., 8 spp. endemic to Madagascar)

Corynecerus deanneX, 1949a: 111 (subgenus)

Trissemodes Jeannel, 1949b: 84 (subgenus)

Trissemidius Jeannel, 1952a: 182 (subgenus)

Apotrissemus iemmeX, 1954: 211 (subgenus)

Trissemites Jeannel, 1959a: 529 (subgenus)

Trissemellus Jeannel, 1959a: 529 (subgenus)

Trissemosus Jeannel, 1959a: 529 (subgenus)

Xenobryaxis Jeannel, 1954a: 234 (1 sp.) E

Tribus Iniocyphini Park, 1951

3 genera and 7 species all endemic to Madagascar.

Capnites Raffray, 1898: 245 (1 sp.) E

Sogaella Jeannel, 1960a: 125 (5 spp.) E

Leiochrotina Jeannel, 1959b: 208 (preocc., not

Westwood, 1883)

Trichopnites Dajoz, 1982: 500 (1 sp.) E

Supertribus Pselaphitae Latreille, 1802

28 genera, 19 endemic to Madagascar, 70 species

all endemic to Madagascar.

Tribus Ctenistini Blanchard, 1 845

7 genera, 4 endemic to Madagascar; 1 3 species all

endemic to Madagascar.

Chenniopsis Raffray, 1904: 338 (1 sp.) E

Ctenisophanes Jeannel, 1954a: 259 (4 spp.) E

Reichenbach, 1816: 75

(30 spp., 1 sp. endemic to Madagascar)

Dionyx Le Peletier & Serville, 1825: 221

Tecnesis Peyerimhoff, 1925: 59 (subgenus)

Leptoctenistes iearmeX, 1956c: 167 (subgenus)

Tectenis Jeannel, 1959a: 617 (misspelling of Tecnesis)

Tecnesites iedxmeX, 1961a: 451 (subgenus)

Desimia Reitter, 1882: 457

(23 spp., 3 spp. endemic to Madagascar)

Tetrads Sharp, 1874: 79

Desmia Leng, 1920: 132 (misspelling)

Desimiella Jeannel, 1949a: 200 (subgenus)

Xenodesimia Jeannel, 1959a: 632 (subgenus)

Enoptostomus Schaum, 1864: 528

(23 spp., 2 spp. endemic to Madagascar)

Hynneophorusl.Q\Qwp, 1972: 171 (1 sp.) E

Laphidioderomimus LoXeup, 1972: 167 (1 sp.) E

Tribus Odontalgini Jeannel, 1 949

3 genera, 2 endemic to Madagascar; 5 species all

endemic to Madagascar.

Algodontodes Jeannel, 1960a: 175 (1 sp.) E

Madontalgus Dajoz, 1982: 506 (1 sp.) E

Odontalgus Raffray, 1877: 286

(49 spp., 3 spp. endemic to Madagascar)

Herminiella Blattny, 1925: 211

Tribus Pachygastrodini Leleup, 1969

2 genera and 3 species all endemic to Madagascar.

Pachygastrodes Leleup 1969: 284 (1 sp.) E

432

G. Sabella, RM. Viglianisi & E. Petralia

Pachygastrodirius Leleup 1976a: 268 (2 spp.) E

Tribus Tyrini Reitter, 1882

8 genera, 6 endemie to Madagasear, 34 speeies all

endemie to Madagasear

Acylobythus Jeannel, 1960a: 178 (2 spp.) E

Acylopselaphus Raffray, 1883: 237 (11 spp.) E

Acylotyrus Jeannel, 1954a: 278 (13 spp.) E

Leptotyrus Jeannel, 1954a: 283

Centrophthalmosis Kaffray, 1904: 376

(27 spp., 1 sp. endemie to Madagasear)

Centrophthalmus Sehmidt Goebel, 1838: 7

(90 spp., 2 spp. endemie to Madagasear)

Camaldus Fairmaire 1863: 637

Franziotus Leleup, 1972: 175 (3 spp.) E

Nesiotyrodes Jeannel, 1954a: 285 (1 sp.) E

Vadoniotus Jeannel, 1954a: 287 (1 sp.) E

Tribus Arhytodini Raffray, 1890

5 genera and 12 speeies all endemie to Madagasear

Eichiella Leleup, 1976b: 311 (2 spp.) E

Holozodoides Dajoz, 1982: 503 (1 sp.) E

HolozodusVdiiYmaixQ, 1898b: 346

NEW NAME for Hologlyptus Fairmaire, 1898a:

338 (preoee., not Le Conte, 1866; Pomel, 1883)

(4 spp.) E

Pasiglyptus Berg, 1899: 79

Tetraglyptinus iQdixmQl, 1960a: 167 (1 sp.) E

Tetraglyptus Jeannel, 1956a: 46 (4 spp.) E

(ineluding Comore islands)

Tribus Pselaphini Latreille, 1802

3 genera, none endemie to Madagasear; 3 speeies

all endemie to Madagasear

Pselaphaulax^QiiiQX, 1909: 218

(61 spp., 1 sp. endemie to Madagasear)

Neopselaphaulax jQSLxmQl 1959a: 593

Pselaphidius Jeannel, 1951: 9

(15 spp., 1 sp. endemie to Madagasear)

PselaphoxysPjdiiixdiy, 1890: 139

(5 spp., 1 sp. endemie to Madagasear)

Supertribus Clavigeritae Leaeh, 1815

39 genera, 36 endemie to Madagasear; 62 speeies

all endemie to Madagasear

Tribus Clavigerini Leaeh, 1815

39 genera, 36 endemie to Madagasear; 62 speeies

all endemie to Madagasear

Andasibe Hlavac & Banaf, 2012: 60 (1 sp.) E

Ankarahitra Jeannel, 1954a: 308 (1 sp.) E

Antalaha Jeannel, 1954a: 318 (5 spp.) E

Antahala Dajoz, 1982: 521 (misspelling)

Apoderiger^dissxxmxxv, 1897: 263 (2 spp.) E

Articeronomus Raffray, 1898a: 268 (1 sp.) E

Articeropsisy^asxxvdixxxi, 1893a: 257

(4 spp., 2 spp. endemie to Madagasear) (also in Sri

Lanka)

Dimerometopus Celis, 1970: 245 (1 sp.) E

Fustigerodes Keittex, 1884: 168

(3 spp., 2 spp. endemie to Madagasear) (also in Bo-

livia?)

Commatoceropsis Raffray, 1890a: 167 (revised status)

Commatocerinus Wassmann, 1897: 260

Fustigeromimus Dajoz, 1982: 515 (1 sp.) E

Hadrophorus Fairmaire, 1898b: 342 (1 sp.) E

Xenofranzia Celis, 1969: 422

Madara Dajoz, 1982: 520 (1 sp.) E

Marofusiger T>a]oz, 1982: 517 (1 sp.)E

Merinia Newton & Chandler, 1989: 67

NEW NAME for /mm/taRaf&ay, 1897: 281 (preoee.,

not Ragonot, 1891) (2 spp.) E

Micrapoderiger iQaxmQl, 1960a: 202 (1 sp.) E

Microclaviger Wasmanxi, 1893: 108 (5 spp.) E

Nearticerodes Jeannel, 1954a: 299 (1 sp.) E

Neoceratopsis Jeannel, 1956a: 50 (1 sp.) E

Neocerus Wasmann, 1893a: 105 (1 sp.) E

Neocorynotus Jeannel, 1960a: 198 (1 sp.) E

Neofustigerinus Jeannel, 1960a: 194 (2 spp.) E

Novofustiger^assxxraxxxv, 1893b: 106(1 sp.) E

Paussiger^asxxmxxr, 1893a: 257 (1 sp.) E

Platycerodes Jeannel, 1960a: 193 (1 sp.) E

Pseudoradama Dajoz, 1982: 519 (1 sp.) E

Radama Raffray, 1883:230 (4 spp.) E

Radamellus Raffray, 1905: 456 (3 spp.) E

Radamides Wasmann, 1897: 261

(4 spp., 3 spp. endemie to Madagasear) E (also in

South Air iea)

Rhynchoclaviger Wasxxiarm, 1891: 4 (1 sp.) E

Rynchoclaviger Raffray, 1911: 175 (misspelling)

Semiclaviger Wasxaaxm, 1893b: 102 (1 sp.) E

Soalala Dajoz, 1982: 512 (1 sp.) E

Stenofustigerinus Celis, 1970: 241 (1 sp.) E

Malgasy genera of Pselaphinae and Pselaphid beetles population in the Amber Mountain, Northern Madagascar 433

Su/ifer Newton & Chandler, 1989: 66

NEW NAME for Fusifer Raffray, 1900: 524

(preocc., not Dendy, 1896) (2 spp.) E

Theocerus Raffray, 1897: 280 (2 spp.) E

Thysdariella Hlavac, 2005: 304 (1 sp.) E

Thysdariopsis Jeannel, 1960a: 211 (1 sp.) E

ThysdariusVdinmdiixe, 1904: 117

NEW NAME for Thysdrus Fairmaire, 1898b: 344

(preocc., not Stal, 1874) (2 spp.) E

Tricemtomerus deannel, 1960a: 195 (1 sp.) E

Trichomatosus Cells, 1970: 268 (1 sp.) E

Trymalius Fairmaire, 1898b: 345 (1 sp.) E

MATERIALS AND METHODS

Study area

The research was conducted within the Natio-

nal Park of Amber Mountain at the north side of

Madagascar, in the province of Antsiranana,

about 40 km southwest of the capital Diego Sua-

rez. The National Park was created in 1958 and it

is the first protected area in Madagascar, covering

about 185 square kilometers. Actually, the Amber

Mountain includes a complex of protected areas

which extend over 23,010 hectares, whose 18,200

hectares of the National Park and 4,810 hectares

of the Special Reserve Amber Forest (Fig. 1). A

track of about 30 km, opened by the French Fo-

reign Legion and today only partially practicable

with ofi-road car, allows to cross the National Park

along the east-west direction. From this track

many paths branch off, allowing to visit by wal-

king the entire area (Fig. 2).

The soil is volcanic and the area of the Natio-

nal Park culminates in the Amber Peak (1,475 m

above sea level) and, like an oasis, is located wi-

thin a region of dry savanna. The rain forest of the

park is lush; with their heavy rainfall (annual rates

of 3,585 mm of rain) it represents the water reser-

voir of the entire region and the rainiest area of

Madagascar. The old craters host numerous lakes

such as the “Lac de la Coupe Verte”, the “Petit

Lac” (Fig. 3), the “Grand Lac” and the “Lac Ma-

hery” near Sakharami (Fig. 4). The crest line is a

natural barrier that divides the relief of the Amber

Mountain in two very different areas. The western

side of the mountain is covered with a dense rain-

forest (Fig. 5), while its eastern side is characteri-

zed by a drier forest (Fig. 6). Due to the volcanic

nature of this basaltic massive, the Park is very

rich in streams and waterfalls which gush in many

localities, offering some surprising views such as

the Great Waterfall, the Little Waterfall and the

Sacred Waterfall (Fig. 7).

The list of collecting stations wit hin the Natio-

nal Park of Amber Mountain (Madagascar) is

shown below (Table 1). Are indicated the geogra-

phical coordinates (latitude and longitude), the

sampling technique used, the altitude, the date of

collection and the name of the sampler.

Sampling methods

Faunistic researchs were carried out on the

western side of the National Park of Amber

Mountain and focused on beetles especially with

regard to family Staphylinidae and the subfamily

Pselaphinae. The research, funded by the Univer-

sity of Catania and the Ministry of Scientific Re-

search of Germany, was supported in the field by

researchers of the University of Antananarivo (Fig.

8) and authorized by The National Association for

the Management of Protected Areas in Madagascar

(A.N.G.A.P.). As shelter was used the small fore-

stal barrack located in the Park. The entomological

samplings were carried out from 18 to 31 March

2011 in various environments (soil, litter, on her-

baceous, shrub and arboreal vegetation, in subcor-

ticicolous and sublapidicolous environment), by

sight or using the entomological net and sifting.

Images of the insects were obtained via a ste-

reoscopic microscope Leica MZ 205A (equipped

with the software auto-montage pro, Syncroscopy).

RESULTS

Current knowledge about Pselaphinae from

the National Park of Amber Mountain (Nor-

thern Madagascar)

On the basis of literature cited above, for the

study area are currently known 16 genera (14 en-

demic to Madagascar and one of Amber Mountain)

and 23 species, all malagasy endemic, 19 of which

are known only to the Amber Mountain area. He-

reinafter they are reported and briefly commented,

and with the letter E are indicated those strictly en-

demic of the district of Amber Mountain.

434

G. Sabella, RM. Viglianisi & E. Petralia

Amber Mmutam

NadoDsl Park **

Reserve speciale

de Andiiamena

MONZABIQUE

Indian

Ocean

• ^

^ /

^ Jbffrevill# ' ^

^ ■ T - j /i ^

/ P , *

- i

Amber ^Souht^in

^ National Park

a . ■ f

tL..

Antsalaka

^ ^ a'

•X

<-o

-X*

^ j'

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r' ^

Z' -I

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Figure 1. Madagascar: localization of the National Park of Amber Mountain.

Figure 2. Map of the National Park of Amber Mountain (from Google Earth, mod.).

Malgasy genera of Pselaphinae and Pselaphid beetles population in the Amber Mountain, Northern Madagascar 435

Northern Madagascar, Antsiranana province, Diana region, National Park Ambre Mountain

Mad 1 . Path between forestal home and Saered Waterfall.

12°31’39.27”S49°10’17.51”E, sifting of litter, 1,062

m 19/111/2011,0. Sabella.

Mad 2. Petit Lae. 12°32’10.04”S 49°10’33.59”E, sifting

of litter along banks, 1,070 m 20/IIE2011, G. Sabella.

Mad 3. Petit Lae. 12°32’12.73”S49°10’27.26”E, Sifting of

litter and rotten wood, 1,150 m 20/IIE2011, G. Sabella.

Mad 4. Temporary lake along Thousand Trees traek.

12°31’23.30”S 49°10’19.16”E, sifting of litter and

rotten wood, 1,040 m 21/III/2011, G. Sabella.

Mad 5. Temporary lake along Thousand Trees traek.

12°31’23”S 49°10’19”E, entomologieal net on her-

baeeous vegetation, 1,040 m 21/III/2011, G. Sabella.

Mad 6. 1 km after the Park entranee. 12°30’55”S

49°10’49”E, entomologieal net on herbaeeous vege-

tation, 990 m 22/III/2011, R. Ranaivosolo.

Mad 7. Big tree near forestal house. 12°3r38.13”S

49°10’17.83”E, sifting of litter, 1,060 m 22/III/2011,

G. Sabella.

Mad 8. Wetland along Thousand Trees traek. 12°3 1’ 16”S

49°10’24”E, entomologieal net on herbaeeous vege-

tation, 1,040 m 23/III/2011, G. Sabella.

Mad 9. Along Thousand Trees traek before wetland.

12°3r22”S 49°10’23”E, entomologieal net on her-

baeeous vegetation, 1,050 m 23/IIE2011, G. Sabella.

Mad 10. Along Thousand Trees traek before wetland.

12°31’16”S 49°10’23”E, sifting of litter and rotten

wood, 1,040 m 23/III/2011, G. Sabella.

Mad 1 1 . Along artifieial eanal near the wooden bridge.

12°3r27”S 49°10’21”E, entomologieal net on her-

baeeous vegetation, 1,050 m 23/III/2011, G. Sabella.

Mad 12. Saered Waterfall. 12°31’41”S 49°10’17”E, en-

tomologieal net on herbaeeous vegetation, 1,070 m

23/III/2011, G. Sabella.

Mad 13. Wetland along Thousand Trees traek.

12°3ri6”S 49°10’24”E, sifting of litter, 1,040 m

24/III/2011, G. Sabella.

Mad 14. Sommet traek 600 m after the eross Saered Water-

fall. 12°31’42”S 49°10’15”E, entomologieal net on

herbaeeous vegetation, 1,070 m 25/IIE201 1, G. Sabella.

Mad 15. Sommet traek, 1,700 m after the eross Saered

Waterfall. 12°31’59”S 49°10’09”E, sifting of litter

and rotten wood, 1,150 m 25/111/2011, G. Sabella.

Mad 16. Sommet traek, about 5 km after the eross Saered

Waterfall. 12°34’13”S 49°10’57”E, entomologieal

net on herbaeeous vegetation, 1,250 m 25/III/2011,

G. Sabella.

Mad 17. Big Caseade traek, about 1 km after the eross Sa-

ered Waterfall. 12°31’00”S 49°10’30”E, rotten

wood, 1,050 m 26/III/2011, G. Sabella.

Mad 18. Big Caseade traek, about 1,3 km after the eross

Saered Waterfall. 12°30’57”S 49°10’30”E, under

bark, 1,030 m 26/III/2011, G. Sabella.

Mad 19. Big Caseade traek, about 2,5 km after the eross

Saered Waterfall. 12°30’00”S 49°10’23”E, sifting of

litter and rotten wood, 850 m 26/III/20 1 1 , G. Sabella.

Mad 20. Big Caseade traek, about 2,8 km after the eross

Saered Waterfall. 12°30’04”S 49°10’23”E, entomo-

logieal net on herbaeeous vegetation and trees, 860 m

27/III/2011, R. Ranaivosolo.

Mad 21. Big Caseade traek, about 2,8 km after the eross

Saered Waterfall. 12°30’04”S 49°10’23”E, sifting of

litter, 860 m 27/III/2011, R. Ranaivosolo.

Mad 22. Big Caseade traek, about 1,3 km after the eross

Saered Waterfall. 12°30’57”S 49°10’30”E, sifting of

bark, 1,030 m 27/III/2011, G. Sabella.

Mad 23. Forestal house. 12°31’37”S 49°10’19”E, ento-

mologieal net on herbaeeous vegetation, night-time,

1,055 m 27/III/2011, G. Sabella & R. Ranaivosolo.

Mad 24. Thousand Trees traek, small waterfall.

12°31’23”S 49°10’16”E, sifting of litter, 1,055 m

28/III/2011, G. Sabella.

Mad 25. Sommet traek, 500 m after the eross Saered

Waterfall. 12°31’43”S 49°10’06”E, entomologieal

net on trees, 1,120 m 28/111/2011, G. Sabella & R.

Ranaivosolo.

Mad 26. Sommet traek, 250 m after the eross Saered Wa-

terfall. 12°31’42”S 49°10’23”E, sifting of litter and

rotten wood, 1,080 m 28/III/20 11, G. Sabella.

Mad 27a. Lae Mahery, Sakharami, around lae. 12°26’25”S

49°14’49”E, entomologieal net on trees (morning time),

370 m 29/III/2011, G. Sabella & R. Ranaivosolo.

Mad 27b. Lao Mahery, Sakharami, forest around lae.

12°26’13”S 49°14’47”E, sifting of litter and rotten

wood, 390 m 29/111/2011, G. Sabella.

Mad 27o. Lae Mahery, Sakharami, around lae.

12°26’25”S 49°14’49”E, entomologieal net on trees

(in the afternoon), 370 m 29/111/2011, G. Sabella &

R. Ranaivosolo.

Mad 27d. Lae Mahery, Sakharami, forest around lae.

12°26’31”S 49°14’49”E, sifting of litter and rotten

wood, 375 m 29/111/2011, G. Sabella.

Mad 28. Thousand Trees traek, small waterfall.

12°31’23”S 49°10’16”E, sifting of litter, 1,055 m

30/111/2011, R. Ranaivosolo.

Mad 29. Thousand Trees traek, between wetland and Big

Caseade traek. 12°31’04”S 49°10’21”E, entomolo-

gieal net on trees, 1,020 m 30/111/2011, G. Sabella &

R. Ranaivosolo.

Mad 30. Thousand Trees traek, between wetland and

Big Caseade traek. 12°30’53”S 49°10’18”E, sifting

of litter of epiphyte on wood, 980 m 30/111/2011, G.

Sabella.

Mad 3 1 . Thousand Trees traek, between wetland and Big

Caseade traek. 12°30’45”S 49°10’12”E, sifting of

bark, 1,000 m 30/111/2011, G. Sabella.

Mad 32. Sommet traek 600 m after the eross Saered Wa-

terfall. 12°3r42”S 49°10’15”E, entomologieal net

on trees, 1,070 m 31/111/2011, G. Sabella.

Mad 33. Sommet traek, 1.700 m after the eross Saered

Waterfall. 12°31’59”S 49°10’09”E, entomologieal

net on trees, 1,150 m 31/111/2011, G. Sabella.

Table 1. The list of eolleeting stations within the National Park of Amber Mountain (Madagasear)

436

G. Sabella, RM. Viglianisi & E. Petralia

Faronitae Reitter, 1 882

E Faronites curtipennis Leleup, 1977

Faronites curtipennis Leleup 1977: 72

Remarks. Endemic of Amber Mountain, known

for a single female collected by H. Franz in

22.V.1969 near the forestal station.

E Faronites jeanneli Leleup, 1977

Faronites jeanneli Leleup 1977: 72, fig. 1 (aedeagus)

Remarks. Endemic of Amber Mountain, known

for two male and two female collected by H. Franz

in 22.V.1969 at Samdramiana river, 800 meters

above sea level.

E Faronites robinsoni Jeannel, 1959

Faronites robinsoni Jeannel 1959b: 191, fig. 2 (aedea-

gus); Jeannel 1961b: 2

Remarks. Endemic of Amber Mountain, known

for a single female collected by A. Robinson in

XIL1958 at Les Roussettes by the soil washing,

1,100 meters above sea level.

Euplectitae Streubel, 1839

Bithynoplectini Schaufuss, 1890

Apozethopsus vadoni jQannQl, 1954

Apozethopsus vadoni Jeannel 1954a: 159, figs. 6 (habitus),

7 (palpus), 8 (aedeagus); Jeannel 1954b: 85, figs 39

(habitus), 40 (palpus), 41 (aedeagus); Jeannel 1960a:

61, figs. 22 (habitus), 23 (aedeagus); Coulon 1989:

133, figs. 99 (palpus), 100 (antenna), 101 (aedeagus)

Apozethopsus pauliani Jeannel 1954a: 159, fig. 9 (ae-

deagus); Jeannel 1954b: 85, fig. 42 (aedeagus); Je-

annel 1960a: 62

Remarks. Species known only for the Northea-

stern Madagascar (Andranofotsy) and Amber

Mountain, where 1 male have been collected by R.

Paulian in XII. 1948.

E Protozethopsus franzi Leleup, 1977

Protozethopsus franzi Leleup 1977: 75, fig. 2 (aedeagus)

Remarks. Endemic of Amber Mountain, known

for 2 male collected by H. Franz in 20.V.1969.

Euplectini Streubel, 1839

E Antinosintus ambreanus Jeannel, 1959

Aminosimus ambreanus Jeannel 1959b: 196; Jeannel

1961b: 5

Remarks. Endemic of Amber Mountain, known

for one female collected by sight by P. Remy in the

summer of 1957 on clayey ground.

E Asymoplectus ambreanus (Jeannel, 1954)

Biblioplectinus ambreanus Jeannel 1954a: 173, fig. 24

(aedeagus)

Asymopleetus (Asymoplectodes) ambreanus Jeannel

1960a: 79, fig. 46 (aedeagus)

Remarks. Endemic of Amber Mountain, known

for 5 specimens collected by R. Paulian in XII. 1948.

Asymoplectus remyi iQdiVmQl, 1959

Asymoplectus remyi Jeannel 1959b: 192, fig. 5 (aedeagus)

Asymoplectus (s. str.) remyi Jeannel 1961b: 3

Remarks. Species known from the Perinet Fo-

rest (Analamazoatra, Central Madagascar) and

Amber Mountain, where 6 specimens have been

collected near the Petit Lac in the soil by P. Remy

in the summer of 1957.

E Asymoplectus decoratus Jeannel, 1959

Asymoplectus decoratus Jeannel 1959b: 194, fig. 7 (ae-

deagus)

Asymoplectus {Asymoplectodes) decoratus Jeannel

1961b: 3

Remarks. Endemic of Amber Mountain, known

for one male collected by P. Remy in the summer

of 1957 near the Petit Lac in the soil.

E Nesiotoplectus megacephalus (Raffray, 1897)

Euplectus megacephalus Raffray 1897: 265

Nesiotoplectus megacephalus Jeannel 1954a: 176, figs

26 (habitus), 27 (aedeagus); Jeannel 1960a: 94, figs

66 (habitus), 61 (aedeagus); Jeannel 1961b: 9-10

Remarks. The species is known only for the en-

virons of Diego Suarez, where 3 specimens have

been collected by C. Allaud in 1893.

E Trimiophanes microphthalmus Jeannel, 1959

Trimiophanes microphthalmus Jeannel 1959b: 197, fig.

16 (aedeagus); Jeannel 1961b: 6 (microcephalus,

misspelling), 8

Remarks. Endemic of Amber Mountain, known

for 4 specimens collected by A. Robinson in

XII. 1958 at Les Roussettes by soil washing, 1,100

meters above sea level.

Malgasy genera of Pselaphinae and Pselaphid beetles population in the Amber Mountain, Northern Madagascar 437

7 8

Figures 3-6. Madagascar, Antsiranana province, National Park of Amber Mountain. Fig. 3. The Petit Lac. Fig. 4. The Lac

Mahery near Sakharami. Fig. 5. Rainforest on the western side of the Amber Mountain. Figure 6. Forest on the eastern side

of Amber Mountain. Fig. 7. National Park of Amber Mountain: the Sacred Waterfall (Photos R. Gerecke). Fig. 8. Researchers

of the University of Tuebingen (Germany) and Antananarivo (Madagascar) on the trail that leads to the entrance of the Na-

tional Park of Amber Mountain (Photo G. Sabella).

438

G. Sabella, RM. Viglianisi & E. Petralia

Trichonychini Reitter, 1882

E Badensia pauliani Jeannel, 1954

Badensia pauliani Jeannel 1954a: 193, figs 43 (habitus),

44 (aedeagus), 45 (metatibiae and metatarsi); Jeannel

1960a: 111-112, figs 100 (habitus), 101 (aedeagus)

Remarks. Endemic of Amber Mountain,

known for one specimen male collected by R. Pau-

lianin XIL1948.

E Ranavala ambreana Jeannel, 1959

Ranavala ambreana Jeannel 1959b: 207, fig. 23 (ae-

deagus)

Ranavala (s. str.) ambreana Jeannel 1961b: 12

Remarks. Endemic of Amber Mountain,

known for 5 specimens collected at Les Roussettes

in the soil by P. Remy in VII. 1957.

Ranavala crassiuscula Jeannel, 1959

Ranavala crassiuscula Jeannel 1959b: 206, fig. 24 (ae-

deagus); Jeannel 1960b: 13

Ranavala (s. str.) crassiuscula Jeannel 1961b: 12

Remarks. Species known for Marojejy Massif

(Sambava, Ambinanitelo) and the Amber Moun-

tain where it has been collected only one winged

female in the soil at Les Roussettes by P. Remy in

VII. 1957, 950 meters above sea level.

Batrisitae Reitter, 1882

Batrisini Reitter, 1882

E Batrisomina wewalkai Leleup, 1977

Batrisomina wewalkai Leleup 1977: 93

Remarks. Endemic of Amber Mountain,

known for one female collected by H. Franz near

the Petit Lac in 2 1 .V. 1 969, 1 ,400 m above sea level.

E Franzorella trifossulata Leleup, 1977

Franzorella trifossulata Leleup 1977: 83, figs 8 (habi-

tus), 9 (aedeagus), 10 (antenna), 11 (median leg), 12

(metatrochanter)

Remarks. Genus and species endemic of the

Amber Mountain, where have been collected two

males and one female by H. Franz in 20.V.1969

near the forestal station.

Goniaceritae Reitter, 1 882

Brachyglutini Raffray, 1904

E Rabyxis pauliani Jeannel, 1954

Rabyxis pauliani Jeannel 1954a: 198, fig. 47 (aedeagus)

Rabyxis (s. str.) pauliani Jeannel 1961b: 131, 134, fig.

132 (aedeagus)

Remarks. Endemic of Amber Mountain, known

for 5 specimens collected by R. Paulian in XII. 1 948.

E Rabyxis viduana (Raffray, 1 897)

Reichenbachia viduana Raffray 1897: 266

Rabyxis viduana Jeannel 1954a: 200, fig. 51 (aedeagus)

Rabyxis (s. str.) viduana Jeannel 1961b: 131, 138, fig.

139 (aedeagus)

Remarks. Species known for a single male col-

lected by C. Allaud in VI. 1893 at Diego Suarez.

E Reichenbachia auriculata Raffray, 1897

Reichenbachia auriculata Raffray 1897: 266; Jeannel

1960a: 151, 152

Reichenbachia (s. str.) auriculata Jeannel 1954a: 220,

225

Remarks. Species known for some specimens

collected by C. Allaud in VI. 1 893 at Diego Suarez.

Reichenbachia usitata Raffray, 1897

Reichenbachia usitata Raffray 1897: 267; Jeannel

1960a: 151, 153

Reichenbachia (s. str.) usitata Jeannel 1954a: 220, 228,

fig. 16 (aedeagus); Jeannel 1956a: 43

Remarks. Species widely distributed and sepa-

rated into several subspecies, whose taxonomic va-

lidity is to be verified. It is signaled for the Northern

Madagascar (Diego Suarez, 7 specimens C. Allaud

1893; Monte Tsaratanana, forest at 1,500 meters

above sea level, one male and one female R. Pau-

lian XL 1949), Eastern Madagascar (Ambodivo-

angy, Ankovana, Andranofotsy, by beating the

branches of trees) and Southern Madagascar (Be-

hara).

PsELAPHiTAE Latreillc, 1 802

Tyrini Reitter, 1882

E Acylopselaphus calcaratus Raffray, 1897

Acylopselaphus calcaratus Raffray 1897: 268; Raffray

1904: 330; Jeannel 1954a: 271, 275, figs 129 (an-

tenna), 130 (palpus), 131 (aedeagus); Jeannel

1960a: 180, 182, figs 220 (antenna), 221 (palpus),

222 (aedeagus)

Remarks. Species known for 1 male and 3 fe-

Malgasy genera of Pselaphinae and Pselaphid beetles population in the Amber Mountain, Northern Madagascar 439

Figure 9. Faronites sp. from Amber Mountain. Figure 10. Nesiotoplectus megacephalus Jeannel, 1954 from Lae Mahery,

Sakharami. Figure 11. Ranavala ambreana Jeannel, 1959 from Petit Lae. Figure 12. Ctnenistes vadoni Jeannel, 1954

from temporary lake along Thousand Trees traek.

440

G. Sabella, RM. Viglianisi & E. Petralia

males collected by C. Allaud in 1 893 at Diego Sua-

rez.

Clavigeritae Leach, 1815

Clavigerini Leach, 1815

E Articeronomus nitidus Raffray, 1 897

Articeronomus nitidus RafFray 1897: 269; Jeannel 1954a:

310; Jeannel 1960a: 202

Remarks. Species collected by C. Allaud at

Diego Suarez.

E Microclaviger alluaudi Raffray, 1897

Microclaviger alluaudi Raffray 1897: 270; Jeannel

1954a: 336, 337, fig. 200 (aedeagus); Jeannel 1960a:

212,213

Remarks. Two specimens of these species were

collected at Diego Suarez by C. Allaud in V. 1 893 in

a rotting tree trunk with Camponotus boisvini.

Preliminary data on Pselaphinae collected

in Amber Mountain from 18 to 31 March

2011

During the period of stay in Amber Mountain

159 specimens of Pselaphinae have been collec-

ted, whose study is currently being defined. At

present only the shunting at genera level has been

completed, while as regards the determination at

the specific level, with some exceptions, it will be

necessary the comparison with the typical mate-

rial. Hereinafter still are reported these prelimi-

nary data, indicating for each identified genus the

number of specimens and the number of collected

species.

Faronitae

F aronites iQdixmQl, 1954. 1 specimen and 1 species

(Fig. 9)

F aronitopsis iQmmQl, 1960. 1 specimen and 1 spe-

cies. Genus reported for the first time for the

Amber Mountain area.

Euplectitae

Bithynoplectini

Apozethopsus Jeannel, 1954. 2 specimens and 1

species.

Euplectini

Asymoplectus Raffray, 1897. 1 specimen and 1

species.

Trimiophanes Jeannel, 1954. 3 specimens and 3

species.

Nesiotoplectus megacephalus Jeannel, 1954

(Fig. 10)

Examined material. Lac Mahery, Sakharami,

12°26’25”S 49°14’49”E, 370 m, 12°26’31”S

49oi4’49”E, 375 m29/in/2011, G. Sabella, 1 male.

Ecological notes. Entomological net on trees.

Trichonychini

Ranavala ambreana Jeannel, 1959 (Fig. 11)

Examined Material. Petit Lac, 12°32’10.04”S

49°10’33.59”E, 1,070 m, 20/III/2011, G. Sa-

bella, 1 female; Petit Lac, 12°32’12.73”S

49°10’27.26”E, 1,150 m, 20/III/2011, G. Sa-

bella, 1 male and 1 female; temporary lake along

Thousand Trees track, 12°31’23.30”S

49°10’19.16”E, 1,040 m, 21/III/2011, G. Sa-

bella, 1 male and 2 female; along Thousand

Trees track before wetland, 12°3ri6”S

49°10’23”E, 1,040 m, 23/III/2011, G. Sabella,

1 female.

Ecological NOTES. Sifting of litter along banks, sif-

ting of litter and rotten wood.

Ranavala crassiuscula Jeannel, 1959

Examined material. Big Cascade track, about 2,5

km after the cross Sacred Waterfall, 12°30’00”S

49°10’23”E, 850 m, 26/III/2011, G. Sabella, 1

male and 1 female.

Ecological notes. Sifting of litter and rotten wood.

Ranavala Raffray, 1898. 3 specimens and 2 species.

Badensia Jeannel, 1954. 2 specimens and 1 species.

Batrisitae

Batrisini

Batrisomina Raffray, 1903. 1 specimen and 1

species.

Malgasy genera of Pselaphinae and Pselaphid beetles population in the Amber Mountain, Northern Madagascar 441

Goniaceritae

Brachyglutini

Reichenbachia Leach, 1826. 109 specimens and 6

species.

Rabyxis Raffray, 1890. 8 specimens and 3 species.

Trissemus Jeannel, 1949. 2 specimens and 1 spe-

cies. Genus reported for the first time for the

Amber Mountain area.

Leichotrella Jeannel, 1954. 1 specimen and 1 spe-

cies. Genus reported for the first time for the

Amber Mountain area.

PSELAPHITAE

Ctenistini

Ctnenistes vadoni Jeannel, 1954 (Fig. 12)

Ctnenistes vadoni Jeannel 1954a: 257, figs 103-104 (an-

tennae), 105 (adeagus); Jeannel 1956a: 49; Jeannel

1960a: 172.

Examined material. Temporary lake along Thou-

sand Trees track, 12°3r23”S 49°10’19”E

1,040 m, 21/III/2011 G. Sabella, 2 males; we-

tland along Thousand Trees track, 12°3ri6”S

49°10’24”E, 1,040 m, 23/III/2011 G. Sabella, 2

males and 3 females; forestal house, 12°3 1 ’37”S

49°10’19”E, 1,055 m, 27/III/2011, G. Sabella&

R. Ranaivosolo, 1 female.

Ecological notes. Entomological net on herbace-

ous vegetation. The genus is reported for the

first time for the Amber Mountain area. The spe-

cies is endemic to Madagascar where it is men-

tioned for numerous localities in the eastern

regions: Antakotako (Maroantsetra); beach of

Maroantsetra; Beanana (high valley Lokoho);

Perinet Forest; Fanovana (Moramanga).

Tyrini

Acylotyrus Jeannel, 1954. 2 specimens and 2

species. Genus reported for the first time for the

Amber Mountain area.

Arhytodini

Eichiellal.Q\Qwp, 1976. 1 specimen and 1 species.

Genus reported for the first time for the Amber

Mountain area.

Clavigeritae

Clavigerini

RhynchoclavigerWassmami, 1891. 1 specimen and

1 species. Genus reported for the first time for

the Amber Mountain area.

CONCLUSIONS

In conclusion, seven genera {Faronitopsis, Tris-

semus, Leichotrella, Ctnenistes, Acylotyrus, Ei-

chiella and Rhynchoclaviger) are reported for the

first time for Amber Mountain area. The research

confirms the high biodiversity of malgasy Pselaphid

fauna and some of its characteristics.

The number of genera and species endemic to

Pselaphinae of Madagascar, although already very

high, it is certainly expected to grow further, partly

because some areas of the Big Island are still

unexplored, and because even in areas that are re-

latively well known, such as the Amber Mountain,

focused research and in different periods of the

year allow to greatly increase the number of ge-

nera and species. In tropical regions and especially

in equatorial regions often coexist in the same area

more congeners species, sometimes very similar to

each other, contrary to what occurs in temperate re-

gions. In this regard, a significant example is re-

presented by the large number of species of

Faronites, Ranavala, Reichenbachia or Trimiopha-

nes collected in Amber Mountain.

Research confirms that for Pselaphid species of

Madagascar, with some exceptions, not ever occur

collections with a large number of specimens. In

fact, even in the literature, the species are known

for one or few specimens. This seems to be another

feature of Pselaphinae fauna from Madagascar. A

further consideration relates to the environment

where is possible to collect these beetles. Besides

the classic environment of the soil, in Madagascar,

as also known in the literature, there are many spe-

cies that live on herbaceous vegetation or on the

branches of trees, in particular those belonging to

the tribe of Brachyglutini and Ctenistini.

Finally, it is surprising that a relatively small

area such as the Amber Mountain guests a so rich

Pselaphid’s population in relation both to the num-

ber of genera and species and also to their rates of

endemicity, and richer, for example, than that of

Sicily.

442

G. Sabella, RM. Viglianisi & E. Petralia

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Biodiversity Journal, 2012, 3 (4): 445-458

State of the art on Neuropterida of Sicily and Malta

Rinaldo Nicoli Aldini', Agostino Letardi^ & Roberto A. Pantaleoni^'^*

'Istituto di Entomologia e Patologia vegetale, Facolta di Agraria, Universita Cattolica del Sacro Cuore, via Emilia Parmense 84,

29122 Piacenza, Italy; e-mail: rinaldo.mcoli@unicatt.it

^ENEA - C.R. CASACCIA UTAGRI ECO S.P. 046, via Anguillarese 301, 00123 S. Maria di Galeria Roma, Italy; e-mail:

agostino.letardi@enea.it

^Dipartimento di Agraria, Entomologia, Universita degli Studi, Via Enrico de Nicola, 07100 Sassari, Italy; e-mail:pantaleo@uniss.it

"^Istituto per lo Studio degli Ecosistemi, Consiglio Nazionale delle Ricerche (ISE-CNR), Traversa la Crucca 3, Regione Baldinca,

07100 Li Punti Sassari, Italy; e-mail: r.pantaleoni@ise.cnr.it

*Corresponding author

ABSTRACT Sicily, the largest Mediterranean island, is surrounded by many small islands (Aeolian Islands,

Ustica, Aegadian Islands, Pantelleria, Linosa, Lampedusa, Maltese Islands), some of whieh

forming archipelagoes. The authors, after a historieal sketeh of the researeh on Neuropterida

in Sicily (sensu lato), analyze the biodiversity of the area, highlighting the species richness and

providing an up-to-date check-list. The lack of knowledge on some of the most paradigmatic

communities of Neuropterida is discussed in relation to their various habitats. The distributional

patterns of Sicilian Neuropterida are interpreted in order to obtain a biogeographieal eharacte-

rization of the area. It is confirmed that the location of Sicily and its surrounding islands forms

a bridge between north and south and a door from the W Mediterranean region to the oriental

Mediterranean basin.

KEY WORDS Raphidioptera; Megaloptera; Neuroptera; faunistics; biogeography.

Received 11.05.2012; accepted 20.12.2012; printed 30.12.2012

Proceedings of the International Congress “Insularity and Biodiversity”, May 11*-13*, 2012 - Palermo (Italy)

INTRODUCTION

Sicily is the largest Mediterranean island. Its lo-

eation represents a bridge from north to south, bet-

ween the Italian Peninsula and the Tunisian Cape

Bon, and a door to the W Mediterranean region,

being in front of the Ionian Sea and the oriental Me-

diterranean basin. The main island is surrounded by

many small islands, some of whieh forming arehi-

pelagoes (Fig. 1). From an orographieal point of

view, along the northern eoast the Peloritani (1,300

m asl), Nebrodi (1,800 m asl), and Madonie (2,000

m asl) mountain ranges represent an extension of

the mainland Apennines. Voleanism is widespread.

Mount Etna, whieh dominates over the eastern

eoast at a height of 3,320 m, is highly aetive as well

as some other eones on the Aeolian Islands. Also

the islands of Ustiea, Linosa and Pantelleria are of

voleanie origin. The Maltese Islands eorrespond to

the high points of a shallow plateau between Sieily

and North Afriea and they are geologieally linked

with the Hyblaean Mountains (1,000 m asl) of SE

Sieily. Consequently, despite the faet that Malta be-

longs to a different State, the Maltese Arehipelago

and Sieily (with its surrounding small islands) to-

gether form a single geographie system.

Due to their loeation in the Mediterranean Sea,

Sieily and its surrounding islands have a partieu-

446

R. Nicoli Aldini, A. Letardi & R.A. Pantaleoni

Figure 1. Map showing the loeation of Sieily and the sur-

rounding islands (modified from a map by Norman Einsten,

Wikipedia).

larly rich flora and fauna and represent a crossroads

between various biogeographieal patterns. This is

also eonfirmed by the data on Neuropterida that has

been aeeumulating for some deeades. Here we in-

tend to review and summarize the eurrent kno-

wledge on Neuropterida, foeusing on some still

unresolved issues.

Notes. The upgrade ineludes data until April

2012. The main papers about Neuropterida from Si-

eily will be eited in the paragraph on historieal data.

In the text we use simply “Sieily” referring to the

whole geographie system eomprising mainland Si-

eily and its surrounding islands, ineluding the Mal-

tese arehipelago. Otherwise we speeify “mainland

Sieily” referring only to Sieily sensu strieto. We fol-

low Vigna Taglianti et al. (1992) in the names and

definitions of geographieal distribution patterns.

RESULTS AND DISCUSSION

History of the research on Neuropterida

The oldest mention of a neuropteran for Sieily

is in “La topografia di Palermo e de' suoi dintomi”

[The topography of Palermo and its surroundings]

(Seina, 1818). The abbot Domenieo Seina introdu-

eed oeeasional entomologieal observations in his

book, listing some inseets in a footnote in whieh he

mentions an unidentifiable snakefiy using the name

Raphidia ophiopsis. The seeond, mueh more impor-

tant reeord regards the deseription of a new, still

valid, speeies of owlfiy by the naturalist Bernardino

Angelini (1827). Despite mueh damage, the type

Ascalaphus siculus (today Libelloides siculus) is

still preserved at the Natural History Museum of

Verona (Fig. 2).

During the nineteenth eentury, the papers of

three authors were the fundamental steps to our

knowledge about Sieilian Neuropterida: two Ger-

Figure 2. The two syntypes oi Ascalaphus siculus AngAmi, 1827, preserved at the Civie Natural History Museum of Verona.

[Label under left speeimen: Ascalaphus \ siculus Angelini \ eotipo deeolorato \ dalla luee e dal tempo \ 937 det. Dott. F.

Capra. Label under right speeimen: Sieilia \ al tempio \ di Segesta. Photo by Daniele Zanini.

State of the art on Neuropterida of Sicily and Malta

447

man-speaking scholars who studied the material

collected by two traveling colleagues, and an eclec-

tic indigenous naturalist.

Wilhelm Gottlieb Schneider, an entomologist

and botanist from Silesia (Schroter, 1890), studied

the material collected by Philipp Christoph Zeller

(McLachlan, 1883), describing some new species

(Schneider, 1845). Hennann August Hagen, a fa-

mous entomologist and bibliophile from Prussia

who ended his career in the United States of Ame-

rica (Henshaw, 1894), published the list of Neurop-

terida collected by Jean-Baptiste Eugene Bellier de

la Chavignerie (Hagen, 1860). From Castelbuono

on the Madonie mountains, the physician Francesco

Mina-Palumbo wrote on Sicilian Neuroptera (in the

old sense, also comprising Odonata and other small

orders) (Mina-Palumbo, 1858; 1871).

The first mentions of Neuropterida from a small

island near Sicily, in this case Ustica, were due to

the teacher Giuseppe Riggio (1885; 1889) from Pa-

lermo. In the twentieth century, both the main Italian

specialists published a paper on Sicilian Neuropte-

rida: Felice Capra published a list of antiions collec-

ted in NE Sicily (Capra, 1934) and Maria Matilde

Principi published the results of the research on the

fauna of the Apennines (comprising the northern Si-

cilian mountains) conducted by the Natural History

Museum of Verona (Principi, 1966).

In the following decades, data accumulated

mainly in the form of scattered occasional records.

Nevertheless, some studies specifically dedicated

to Sicilian (or South-Italian) Neuropterida (Aspock

etal., 1980; Nicoli Aldini, 1983; Insomet al., 1986;

Pantaleoni, 1986; Duelli, 1994; Fo Valvo, 1994;

Pantaleoni & Fo Valvo, 1995; [Bernard!] lori et al.,

1995; Plant & Schembri, 1996; Fo Verde & Mon-

serrat, 1997; Nicoli Aldini & Baviera, 2001) appea-

red and the species known for Sicily rose from forty

to one hundred (Fig. 3).

Biodiversity by taxa: species richness

Almost half of the Neuropterida species found

in Italy, about a hundred (exactly 97 including the

doubtful ones), are currently known in Sicily

(Table l;Figs. 4-19).

1800 1850 1900 1950 2000

Figure 3. Cumulative progress of the number of Neuropterida speeies known for Sieily. Papers eiting more than five new

speeies for Sieily are highlighted.

448

R. Nicoli Aldini, A. Letardi & R.A. Pantaleoni

As the order Megaloptera is absent, they belong

to the orders Raphidioptera and Neuroptera.

Raphidioptera. Only two speeies of Raphidiop-

tera are eommon in Sieily, eaeh representing one of

the two families of the order, Raphidiidae and Ino-

eelliidae. The inoeelliid Fibla (Fibla) maclachlani

(Albarda, 1891) belongs to a typieal Mediterranean

genus. The W Mediterranean nominal subgenus

Fibla has three speeies: F {F.) hesperica Navas,

1915 from the Iberian Peninsula, F. (i^) peyerim-

hoffi Navas, 1919 from the Mediterranean areas of

North Afriea, and the Tyrrhenian F {F) maclachlani

from Corsiea, Sardinia and mainland Sieily. In the

E Mediterranean area there is another subgenus

with a speeies from Crete: Fibla (Reisserella) pasi-

phae (H. Aspoek et U. Aspoek, 1971).

The raphidiid Xanthostigma corsicum (Hagen,

1867) has a typieal Tyrrhenian distribution being

present in Corsiea, Sardinia and mainland Sieily,

even on small islands sueh as Capraia and Elba in

the Tusean Arehipelago, and on the Italian Penin-

sula, moreover on the “Sierra” of Bejar, mountains

of the Sistema Central, Spain (H. Aspoek et al.,

1991). In every geographieal area (e.g. Sieily, Cor-

sica and Sardinia, Spain, etc) this species shows

small but constant morphological differentiation.

A third species, the raphidiid Subilla confinis

(Stephens, 1 836), is known by a single unpublished

record from the Madonie mountains (label: Sicilia

(PA) Madonie / Piano Zucchi m 1300 / G. Sama leg.

// Ex larva / Acer / 17.V.84. specimens: 1 male & 2

females. Location: Natural History Museum of Ve-

rona. Identification: H. & U. Aspoek, 2002). This

species lives on tree canopies and has a wide distri-

bution from Central Europe to Japan through Sibe-

ria. S. confinis reaches the Mediterranean range

only through the Italian Peninsula, other similar Su-

billa inhabit the Iberian and Balkan peninsulas. Ho-

wever there is no report of this species along the

Italian peninsula between the Alpine and southern

populations.

Neuroptera. Seven families of Neuroptera are

known for Sicily: Nevrorthidae, Coniopterygidae,

Mantispidae, Chrysopidae, Hemerobiidae, Mymie-

leontidae, Ascalaphidae. The most speciose are the

green lacewings (Chrysopidae) with 25 species, fol-

lowed by antiions (Myrmeleontidae) with 23, du-

stywings (Coniopterygidae) with 20 and brown

lacewings (Hemerobiidae) with 18. Nevrorthidae,

Mantispidae and Ascalaphidae have respectively

one, three and four species.

The nevrorthid Nevrorthus iridipennis A. Costa,

1863 is confined in the range of the Peloritani

mountains (mainland Sicily) and in the facing

Aspromonte massif (Italian peninsula). Another

three species belong to the genus Nevrorthus: N. fal-

lax (Rambur, 1 842) from Corsica and Sardinia, N.

hannibal U. Aspoek et H. Aspoek, 1983 from Me-

diterranean areas of Algeria and Tunisia, N. apate-

lios H. Aspoek, U. Aspoek et Holzel, 1977 from the

Balkans reaching northwards to the foothills of the

eastern Italian Alps (Letardi et al., 2006).

The species of Coniopterygidae recorded for Si-

cily are approximately three quarters of those found

in Italy. This large number has been positively in-

fluenced by targeted, although occasional, research

carried out recently by some specialists (Lo Verde

& Monserrat, 1997). Actually, the small, delicate

and inconspicuous dustywings are rarely collected

by non neuropterists and they are very scarce in pu-

blic entomological collections. Consequently the in-

formation about this family’s habitat, ecology, and

species distribution is still poor. The list of Sicilian

species, with many widespread, European or Me-

diterranean taxa, is not particularly informative

(Table 1). Only Helicoconis hispanica Ohm, 1965

shows an interesting biogeographical pattern, being

previously known only for the south of Spain and

more recently for Sardinia (Lorn et al., 2011).

All three species of Italian Mantispidae are re-

corded for Sicily. Insects with a complex biology,

characterized by hypermetabolic development and

larvae evolving in spider egg-sacs, the W Palearctic

mantisflies show wide patterns of distribution. Re-

latively few species of Hemerobiidae are strictly

correlated to the Mediterranean climate, so the

number of species recorded for Sicily comes to only

40% of the whole Italian fauna. Among many spe-

cies which are widespread, or common in agro-eco-

systems, there are some with clearly defined

biogeographical patterns. Wesmaelius tjederi (Kim-

mins, 1963), W. ravus (Withycombe, 1923), Sym-

pherobius elegans (Stephens, 1836), nnd Micromus

paganus (Linnaeus, 1767), all found rarely or once,

have a northern distribution; Wesmaelius navasi

(Andreu, 1911) and Sympherobius fallax Navas,

1908 a southern one. Moreover W. navasi is the

only species recorded exclusively from Malta and

not from Sicily or other surrounding islands.

The knowledge about Chrysopidae is strongly

affected by unresolved taxonomical problems regar-

State of the art on Neuropterida of Sicily and Malta

449

Species list

Record reliability

and presence on

the islands

Distributional pattern

RAPHIDIIDAE

Subilia confinis (Stephens, 1836)

! S

Central-European

Xanthostigma corsicum (Hagen, 1867)

! S

W-Mediterranean

INOCELLllDAE

Fibla (Fibla) machlachlani (Albarda, 1891)

! S

Tyrrhenian

NEVRORTHIDAE

Nevrorthus iridipennis Costa, 1863

! S

Sicilian-Calabrian

CONIOPTERYGIDAE

Aleuropteryx loewii¥Adipk\e]L, 1894

! S

S-European

Aleuropteryx juniperi Ohm, 1968

! S,M

S-European

Helicoconis (Ohmopteryx) pseudolutea Ohm, 1965

! S

Turanian-Euro-Mediterranean

Helicoconis (Fontenellea) hispanica Ohm, 1965

! S

W-Mediterranean

Coniopteryx (Xeroconiopteryx) loipetsederi Aspoek, 1963

! S,M

Mediterranean

Coniopteryx (Coniopteryx) borealis Tjeder, 1930

! S

European

Coniopteryx (Coniopteryx) pygmaea Enderlein, 1906

! S

Euro-Siberian

Coniopteryx (Holoconiopteryx) haematica MeLaehlan, 1868

! S

Euro-Mediterranean

Coniopteryx (Holoconiopteryx) renate Rausch et Aspoek, 1977

! S

E-Mediterranean

Coniopteryx (Metaconiopteryx) arcuata Kis, 1965

! S

Mediterranean

Coniopteryx (Metaconiopteryx) esbenpeterseni Tjeder, 1930

! S

S-European

Coniopteryx (Metaconiopteryx) lentiae Aspoek et Aspoek, 1964

! S

S-European

Coniopteryx (Metaconiopteryx) tjederi Kimmins, 1934

! S

S-European

Parasemidalis fuscipennis (Reuter, 1894)

! S

Holarctic

Hemisemidalis pallida (Withy combe, 1924)

! S

Central-Asian Mediterranean

Conwentzia pineticola Enderlein, 1905

! S

Holarctic

Conwentzia psociformis (Curtis, 1834)

! S,M

Holarctic

Semidalis aleyrodiformis (Stephens, 1836)

! S

Palearctic

Semidalis pseudouncinata Meinander, 1963

! S

W-European

Semidalis vicina (Hagen, 1861)

! S,M

W-Mediterranean

MANTISPIDAE

Mantispa styriaca (Poda, 1761)

! S

Central-Asian Euro-Mediterranean

Mantispa perla (Pallas, 1772) (sensu Erichson, 1839)

! S

Central-Asian Mediterranean

Mantispa aphavexelte Aspoek et Aspoek, 1994

! S

Central-Asian Mediterranean

HEMEROBllDAE

Hemerobius humulinus Linnaeus, 1758

Holarctic

Hemerobius stigma Stephens, 1836

! S

Holarctic

Hemerobius handschini Tjeder, 1957

! S

S-European

Hemerobius micans Olivier, 1792

! S

European

Hemerobius gilvus Stein, 1 863

! S

S-European

Wesmaelius tjederi (Kimmins, 1963)

! S

S-European

Wesmaelius subnebulosus (Stephens, 1836)

! S,L

Palearctic

Wesmaelius ravus (Withycombe, 1923)

! S

Asian-European

Wesmaelius navasi (Andreu, 1911)

! M

Central-Asian Mediterranean

Sympherobius pygmaeus (Rambur, 1842)

! S,M

Turanian-Euro-Mediterranean

Sympherobius luqueti (Leraut, 1991)

! S,^

currently not definable

450

R. Nicoli Aldini, A. Letardi & R.A. Pantaleoni

Species list

Record reliability

and presence on

the islands

Distributional pattern

Sympherobius elegans (Stephens, 1836)

! S

Turanian-European

Sympherobius fallax Navas, 1908

! S,M

Afrotropieal-Mediterranean

Megalomus tineoides Rambur, 1842

! S

Mediterranean

Megalomus pyraloides Rambur, 1 842

! S

W-Mediterranean

Micromus variegatus (Fabrieius, 1793)

! S

Asian-European

Micromus angulatus (Stephens, 1836)

! S,M

Holaretie

Micromus paganus (Linnaeus, 1767)

Asian-European

CHRYSOPIDAE

Hypochrysa elegans (Burmeister, 1839)

! S

Central-European

Italochrysa italica (Rossi, 1790)

! S,M

Mediterranean

Nineta flava (Seopoli, 1763)

European

Nineta principiae Monserrat, 1980

! S

S-European

Chrysopa perla (Linnaeus, 1758) sensu Sehneider, 1851

?S,M

Asian-European

Chrysopa dorsalis BmmQistQY, 1839

! S

Turanian-European

Chrysopa formosa Brauer, 1850

! S, M, ^

Asian-European

Chrysopa viridana Sehneider, 1 845

! S

Turanian-Euro-Mediterranean

Chrysopa pallens (Rambur, 1838)

! S, M, ^

Palearetie

Pseudomallada flavifrons (Brauer, 1850)

! S, M, ^

Turanian-Euro-Mediterranean

Pseudomallada marianus (Navas, 1905)

! S

eurrently not definable

Pseudomallada inornatus (Navas, 1901)

! S

S-European

Pseudomallada sp. prope picteti (MeLaehlan, 1880)

! S,^

eurrently not definable

Pseudomallada prasinus (Burmeister, 1839)

! S

Palearetie

Pseudomallada zelleri (Sehneider, 1851)

! S

E-Mediterranean

Pseudomallada genei (Rambur, 1 842)

! S, M, L, P, M

Mediterranean

Pseudomallada venustus (Holzel, 1974)

! S,P

Mediterranean

Pseudomallada clathratus (Sehneider, 1845)

! S,M

E-Mediterranean

Cunctochrysa albolineata (Killington, 1935)

! S

Asian-European

Chrysoperla carnea (Stephens, 1836) s. 1.

? S, M, L, U

Chrysoperla agilis Henry, Brooks, Duelli et Johnson, 2003

! S

Mediterranean

Chrysoperla lucasina (Laeroix, 1912)

! S, M, L

Euro-Mediterranean

Chrysoperla mediterranea (Holzel, 1972)

! S,M

S-European

Chrysoperla pallida Henry, Brooks, Duelli et Johnson, 2002

! S

Euro-Mediterranean

Brinckochrysa chlorosoma (Navas, 1914)

! S,M

Afrotropieal-S-Mediterranean

MYRMELEONTIDAE

Palpares libelluloides (Linnaeus, 1764)

! S

Turanian-Mediterranean

Acanthaclisis occitaniea (Villers, 1789)

! S

Turanian-Euro-Mediterranean

Synclisis baetica (Rambur, 1 842)

! S,M

Turanian-Euro-Mediterranean

Myrmecaelurus trigrammus (Pallas, 1781)

! S, U, ^

Central-Asian Mediterranean

Myrmeleon formicarius Linnaeus, 1767

! S

Asian-European

Myrmeleon inconspicuus Rambur, 1842

! S

Turanian-Mediterranean

Myrmeleon hyalinus Olivier, 1811

! S, L, JEg, JE

Afrotropieal-Mediterranean

Myrmeleon punicanus Pantaleoni et Badano, 2012

! S,P

Sieilian

Euroleon nostras (Geoffroy in Foureroy, 1785)

! S

European

Dendroleon pantherinus (Fabrieius, 1787)

! S,M

Central-Asian European

State of the art on Neuropterida of Sicily and Malta

451

Record reliability

Species list and presence on Distributional pattern

the islands

Macronemurus appendiculatus (Latreille, 1 807)

Neuroleon arenarius (Navas, 1904)

Neuroleon egenus (Navas, 1915)

Neuroleon nemausiensis (Borkhausen, 1791)

Neuroleon microstenus (McLachlan, 1898)

Neuroleon ocreatus (Navas, 1904)

Distoleon tetragrammicus (Fabricius, 1798)

Distoleon annulatus (Klug, 1834)

Nemoleon poecilopterus (Stein, 1 863)

Creoleon griseus (Klug, 1834)

Creoleon lugdunensis (Villers, 1789)

Creoleon aegyptiacus (Rambur, 1 842)

Gymnocnemia variegata (Schneider, 1845)

ASCALAPHIDAE

Bubopsis agrionoides (Rambur, 1838)

Deleproctophylla australis (Fabricius, 1787)

Libelloides coccajus ([Denis et Schiffermuller], 1775)

Libelloides siculus (Angelini, 1 827)

!S, M, L,P,^g

Mediterranean

! S,M

Mediterranean

! S,M

Mediterranean

! S, M, JEg

Mediterranean

IS, Mg

E-Mediterranean

?! S

W-Mediterranean

! S

Turanian-European

!M, L,P,^,^g

Mediterranean

! S

Turanian-Mediterranean

! L

Afro trop. -Indian S-Mediterranean

! S, M, L, P, M

W-Mediterranean

! M,L

Turanian-Mediterranean

! S,M

Turanian-Mediterranean

! L

W-Mediterranean

! S,^

E-Mediterranean

! S

S-European

!S,^g

Sicilian

Table 1. Up-to-date Checklist of Sicilian Neuropterida. ! = reliable data, ?! = data to be confirmed, ? = uncertain data. S

= mainland Sicily, M = Maltese Islands, L = Lampedusa and/or Linosa, P = Pantelleria, M = ^olian Islands, U = Ustica,

^g = ^gadian Islands.

ding mainly the genera Chrysoperla and Pseudo-

mallada. The speeies of the former genus use aeou-

stie signals (eourting ealls) as a reproduetive barrier,

and exhibit extremely uniform habitus and genita-

lia. In Chrysoperla the reeent use of the eall patterns

as a diseriminating eharaeteristie was the starting

point for deteeting morphologieal distinguishing

features (Henry et ah, 2001). Instead, in Pseudo-

mallada the “songs”, though present, are not speei-

fie but probably aeeompanied by the emission of

pheromones. Therefore, without even bio-aeoustie

information as a guide, it is very diffieult to diseri-

minate between the different speeies. However pa-

tient work of eorrelation between subtle

morphologieal eharaeteristies, the bio-eeology and

the first data on DNA-taxonomy allows us to begin

to untangle this Gordian knot.

Chrysopidae is the most speeiose Neuroptera fa-

mily in Sieily, with half of the Italian speeies. Their

distribution patterns gravitate mainly in the Medi-

terranean. In the unique, very interesting, ease of

Brinckochrysa chlorosoma (Navas, 1914) mainland

Sieily represents the northern limit of distribution.

On the eontrary, it is more frequently a southern

limit of distribution, e.g.: Hypochrysa elegans (Bur-

meister, IS39), Nineta flava (Seopoli, 1763) andiV.

principiae Monserrat, 1980, Chrysopa perla (Lin-

naeus, 1758) sensu Sehneider, 1851, Pseudomal-

lada marianus (Navas, 1905), and Cunctochrysa

albolineata (Killington, 1935).

Like the Coniopterygidae, the speeies of Myr-

meleontidae reeorded for Sieily are approximately

three quarters of those found in Italy. However this

large number is not due to speeialized researeh but

to oeeasional eaptures over a long period by many

entomologists. Aetually, antiions are attraetive in-

seets, frequently eolleeted though always in low

numbers. Speeialized researeh has led us very re-

eently to the diseovery and deseription of a new

speeies of Myrmeleon (M. punicanus Pantaleoni et

Badano, 2012, known only on mainland Sieily and

Pantelleria) whereas a speeies of Creoleon is still

being studied. Despite the wide distribution of

many antiion speeies, Sieily seems to be effeetively

a erossroads for distributional patterns in this fa-

mily. There are two southern speeies reaehing only

452

R. Nicoli Aldini, A. Letardi & R.A. Pantaleoni

the island of Lampedusa near Afriea, Creoleon ae-

gyptiacus (Rambur, 1842) and C griseus (Klug,

1834), one speeies for whieh Sieily represents the

western limit of distribution, Nemoleon poecilop-

terus (Stein, 1863), three speeies for whieh Sieily

represents the southern limit of distribution rea-

ehing Etna, Myrmeleon formicarius Linnaeus,

1767, Euroleon nostras (Geoffroy in Foureroy,

1785) or the northern mountain range, Dendroleon

pantherinus (Fabrieius, 1787). Moreover the S Me-

diterranean antiion Distoleon annulatus (Klug,

1834) is reeorded for many small islands.

Among the speeies of Asealaphidae, Bubopsis

agrionoides (Rambur, 1838) is known only for

Lampedusa, having a W Mediterranean distribution

with searee and seattered reeords along the eoasts

of southern Franee, eastern Spain, Moroeeo and Tu-

nisia. On the eontrary the E Mediterranean Dele-

proctophylla australis (Fabrieius, 1787) is

widespread in Sieily. Owlflies are strong-flying in-

seets and the narrow (3 . 1 km) Strait of Messina

does not seem able to aet as a barrier against them.

However Sieily and Calabria, the two opposite re-

gions of the strait, share only one speeies of Libel-

loides whereas overall four speeies live in Calabria

and two in Sieily. The shared speeies is L. coccajus

([Denis et Sehiffermuller], 1775), the seeond Siei-

lian speeies is the endemie L. siculus (Angelini,

1 827). The last belongs to a group of Mediterranean

speeies (eonsidered subspeeies by some): L. icteri-

cus (Charpentier, 1 825) from W Mediterranean eon-

tinental eoasts, between south Franee and Tunisia;

the Tyrrenian L. corsicus (Rambur, 1 842) from Cor-

siea, Sardinia and Capraia, an island of the Tusean

Arehipelago; L. cyrenaicus H. Aspoek, Holzel et U.

Aspoek, 1976 from Cyrenaiea.

Biodiversity by habitat: the lack of knowledge

The following list of habitats does not derive

from a rigorous elassifieation of the Sieilian en-

vironment, but it is a simple frame of work inelu-

ding the most paradigmatie eommunities of

Neuropterida.

Freshwater. The only amphibiotie speeies of

Sieilian Neuropterida is the nevrorthid Nevrorthus

iridipennis. Nevertheless if the absenee of alderflies

(Megaloptera Sialidae) and osmylids (Neuroptera

Osmylidae) eould be plausible, the laek of reeords

regarding spongillaflies (Neuroptera Sisyridae) ap-

pears to be very questionable. In the W Mediterra-

nean, the speeies Sisyra iridipennis A. Costa, 1884,

is widespread ineluding islands smaller than Sieily,

sueh as the Balearies, and nearby Sardinia and Tu-

nisia. Speeialized researeh is required to resolve

these doubts.

Soil. In the Euro-Mediterranean region many

Neuropterida live in the soil, partieularly the larvae

of some Raphidioptera and almost all the larvae of

Myrmeleontiformia. But the former live among lit-

ter, under stones, among mosses and the latter on the

surfaee (Asealaphidae) or immediately below the

surfaee (other families), so they eould both be eon-

sidered epiedaphie. The only inhabitant of the soil

mineral layer (euedaphie speeies) are the larvae of

Dilaridae and, perhaps, of Berothidae. Both these fa-

milies are not reeorded from Sieily but the absenee

of Dilaridae does not seem plausible.

Wetlands and coasts. Neuropterida are searee

in wetlands exeluding the amphibiotie ones, but the

speeies living here are very interesting. The eoastal

environments are also the habitat of speeialized

Neuropterida, partieularly the Myrmeleontidae spe-

eies linked to loose sand. While no study was eon-

dueted in wetlands, some data on the eoasts is

available. Unfortunately wetlands and eoasts are

two extremely endangered environments in Sieily,

faeing destruetion through urban development and

reereational use.

Agro-ecosystems. Often the agro-eeosy stems in

the Mediterranean elimate are environments rieh in

inseet biodiversity. Previous studies in regions si-

milar to Sieily, regarding both the Chrysopidae, fa-

mily of agrieultural interest (Pantaleoni & Lepera,

1985; Pantaleoni & Curto, 1990), and the Myrme-

leontidae (Curto & Pantaleoni, 1987), notieed an

unexpeetedly high number of speeies. Unfortuna-

tely, in Sieily the eommunities of Neuropterida in

agrieultural eultivations were never studied, al-

though many of these sueh as vineyards, eitrus or-

ehards, olive groves, and the very loeal and

eharaeteristie manna ash groves or pistaehio or-

ehards appear very promising.

Northern mountain ridge. The northern

mountain ridge has the best preserved wildlife of

Sieily and is perhaps the most frequented area by

naturalists and entomologists. We have a lot of oe-

easional and seattered data on Neuropterida of

these mountains, but speeialized researeh would

produee some surprises.

State of the art on Neuropterida of Sicily and Malta

453

Figures 4-7. Sicilian habitats of some interesting Neuropterida communities. Fig. 4: freshwater, wetlands and coasts: the

mouth of the River Modione (Trapani), May 2008; Fig. 5: agro-ecosystems: Segesta (Trapani), April 2006; Fig. 6: Northern

mountain ridge: beech wood on Monte Mufara, Madonie (Palermo), April 2003; Fig. 7: small islands: Isola dei Conigli,

Lampedusa (Agrigento), September 2005. Photos by Marcello Romano.

454

R. Nicoli Aldini, A. Letardi & R.A. Pantaleoni

Mediterranean forests, woodlands, and

scrub. Mediterranean vegetation is the dominant

vegetation in Sieily and probably its wide diffusion

ereates less interest in entomologists. Indeed this

eommon environment would deserve more atten-

tion, as demonstrated by the reeent diseovery of a

new arboreal speeies of the genus Myrmeleon (Pan-

taleoni & Badano, 2012).

Hyblaean Mountains. The laek of researeh on

Neuropterida in the Hyblaean Mountains is one of

the biggest grey areas in our knowledge of these

inseets in Sieily.

In faet, the Hyblaean Foreland belongs to the

northern part of the Afriean Plate and it is of huge

biogeographieal interest; the reeent diseovery of a

new speeies of tree belonging to a reliet genus in

Figures 8-13. Sicilian Neuropterida. Fig. Xanthostigma corsicum female, Piano Battaglietta Madonie Palermo, 11.VL2006.

Fig. 9: Fibla maclachlani female, Paceco Trapani, 17.1V.2011. Fig. 10: Fibla maclachlani larva, L’Antennamare Peloritani

Messina, summer 2001. Fig. 11: Nevrorthus iridipennis adult, Valle dell’Alcantara Messina, 13.VL2002. Fig. 12: adult of

Coniopteryx sp., Catania, 30.1X.2007. Fig. 13: Mantispa perla adult. Piano Battaglia Madonie Palermo, 21.V1L2004. Photos

by Marcello Romano (8, 13), Luigi Barraco (9), Rinaldo Nicoli Aldini (10, 11) and Alessandro Strano (12).

State of the art on Neuropterida of Sicily and Malta

455

the Mediterranean range, Zelkova sicula Di Pa-

squale, Garfi et Quezel, 1992, is just one example

whieh is also well known by the general publie

(Garfi, 1996).

Small islands. Speeialized researeh was never

earried out on the small surrounding islands exeept

the Maltese Islands (Duelli, 1994; Plant & Sehem-

bri, 1996). In partieular, eurrent knowledge is

searee about Chrysopidae, Hemerobiidae and Co-

niopterygidae, slightly better about Myrmeleonti-

dae and Asealaphidae.

As far as we know, many studies appear very

promising: the Afriean speeies living in Lampe-

dusa and Linosa, the Neuropterida of the woods in

the Pantelleria mountains, the fauna of Aegadian

Islands and Ustiea.

Figures 14-19. Sicilian Neuropterida. Fig. 14: Sympherobius luqueti adult, Marausa Trapani, 30.VIIL2009. Fig. 15: Chrysopa

formosa adult, Nubia Trapani, 14.VI.2009. Fig. 16: Chrysopidae eggs on flower, Paceco Trapani, 2.VI.2007. Fig. 17: larva

of Pseudomallada sp. with its camouflage, Paceco Trapani, 20.VI.2009. Fig. 18: Myrmeleon hyalinus male, foce del Mo-

dione, Trapani, 1.VI.2007. Fig. 19: mating pair of Libelloides siculus, saline Trapani 16.V.2008. Photos by Luigi Barraco

(14, 15, 16, 17) and Marcello Romano (18, 19).

456

R. Nicoli Aldini, A. Letardi & R.A. Pantaleoni

Biodiversity by distributional pattern:

biogeography

It is generally believed that many Neuropterida

have wide distribution areas, however this is pro-

bably due primarily to the poor eurrent knowledge

on the taxonomy of some families, partieularly the

Chrysopidae. A paradigmatie example is that of

Pseudomallada prasinus (Bumieister, 1839), a spe-

eies whieh apparently has Palearetie distribution,

but is probably eomposed of many more loealized

sibling speeies. Unfortunately this situation seriou-

sly affeets the study of distributional patterns in

Neuropterida.

Anyway, as far as we know, also the list of Siei-

lian Neuropterida is rieh in speeies with a wide di-

stribution area (Table 1), but the eommonest

distributional patterns are the Mediterranean, in a

broad sense, and the S European ones. Obviously a

Mediterranean fauna is the normal baekground for

Sieily. The speeies with northern distributional pat-

terns are more interesting. The majority of (Cen-

tral-) European speeies, but sometimes also the

Asian-European or Euro-Siberian ones, follow the

beeeh woods (pure or mixed with eonifers) along

the Italian peninsula, reaehing the northern Sieilian

mountain ranges. The European beeeh woods are

the main habitat of some speeies of Neuropterida

sueh as, e.g., the green laeewing Hypochrysa ele-

gans (Burmeister, 1839) and the brown laeewings

Hemerobius micans Olivier, 1792 and Symphero-

bius elegans (Stephens, 1836).

The speeies with southern distributional patterns

are fewer. Among these there is the eommon Sym-

pherobius fallax Navas, 1908, but also Brincko-

chrysa chlorosoma (Navas, 1914), a very interesting

Chrysopidae, and two species of the African Creo-

leon recorded only for Lampedusa.

The position of Sicily in the W Mediterranean,

but at the border of the east side, favors the pre-

sence of both western and eastern Neuropterida on

the island. So we could find the E Mediterranean

taxa that colonize the Italian peninsula, such as the

laeewings Pseudomallada clathratus (Schneider,

1845) andE zelleri (Schneider, 1851) or the owlfiy

Deleproctophylla australis (Fabricius, 1787). On

the contrary we find also W Mediterranean species

like the dustywing Semidalis vicina (Hagen, 1861)

and the antiion Creoleon lugdunensis (Villers,

1789).

Finally, Sicily hosts some but few endemic or

subendemic (i.e. with small distribution area) Neu-

ropterida all belonging to W Mediterranean taxa.

CONCLUSIONS

The current state of knowledge on the Neurop-

terida of Sicily is satisfactory, especially if we com-

pare it to many other Italian regions. However, the

large number of records accumulated does not de-

rive from specialized research but from occasional,

sporadic sampling. Many environments have not

been adequately studied and some of these appear

to be particularly interesting. Certainly, much more

work will be necessary to fill in the many gaps left

in our knowledge. As far as Neuropterida are con-

cerned, Sicilian biogeography is nevertheless well

outlined and confimis that the island is a crossroads

for Central European, Afrotropical, E and W Medi-

terranean species which can be added to the main

Mediterranean and S European components.

ACKNOWLEDGMENTS

We are greatly indebted to some Sicilian natu-

ralists, particularly Marcello Romano (Capaci, Pa-

lermo), Luigi Barraco (Paceco, Trapani) and

Alessandro Strano (Catania), for their scientific

support and passionate interest; they have helped

us by providing many personal local observations,

comments, suggestions and some marvelous pho-

tos. Warm thanks also to Daniele Zanini (Verona)

who took the pictures of the types of Bernardino

Angelini.

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Annotated checklist of the dragonflies (Insecta Odonata)

of the islands of the Sicilian Channel, including the first re-

cords of Sympetrum sinaiticum Dumont, 1 977 and Pantala

flavescens (Fabricius, 1 798) for Italy

Andrea Corso'*, Ottavio Jan ni^ Maurizio Pavesi^, Michael Sammut^ Arnold Sciberras^ & Michele Vigano^

'MISC - Via Camastra, 10 - 96100 Siracusa, Italy; e-mail: voloerrante@yahoo.it

^Via G.G. D'Amore, n. 21 - 81016 Piedimonte Matese, Caserta, Italy

^Museo di Storia Naturale - Corso Venezia, 55 - 20121 Milano, Italy; e-mail: maurizio_pavesi@yahoo.com

"^11, Sqaq Rigu, Birkirkara, BKR 2131, Malta; e-mail: aquilarus@gmail.com

^24 'Camilleri Court' flat 5, il-Marlozz Str, Mellieha (Ghadira), Malta; e-mail: wildalienplanet@gmail.com

^ MISC - Via Ongetta, 5 - 21010 Germignaga, Varese, Italia; e-mail: mikivigano@yahoo.com

*Corresponding author

ABSTRACT In this paper we report data on the historieal and reeent status of all dragonfly species (In-

secta, Odonata) recorded for the Sicilian Channel islands: the Pelagic islands and Pantel-

leria, politically belonging to Italy, and Maltese Archipelago islands. The number of species

known for the former group of islands raises from 7 to 20. Of these, 2 are new for the Ita-

lian fauna, namely the Desert Darter Sympetrum sinaiticum, noticed through likely si-

ghtings starting from 2010 on Lampedusa, and confirmed through voucher specimens

collected in April 2012, and the Wandering Glider Pantala flavescens, first noticed in Oc-

tober 2012 on Lampedusa and Linosa; while Calopteryx sp. cf. haemorrhoidalis, Ischnura

genei, Aeshna mixta, Orthetrum nitidinerve, Orthetrum coerulescens anceps, Crocothemis

erythraea, Sympetrum striolatum, S. meridionale, Brachythemis impartita, Trithemis an-

nulata and T. kirbyi, already known for Italy, are new for the Italian islands of the Sicilian

channel. The Maltese fauna includes at present 18 recorded species; the previously reported

Trithemis arteriosa is to be deleted from the list, since the concerned specimen upon re-

examination proved to be T. annulata.

KEY WORDS Odonata; Sicilian Channel Islands; Sympetrum sinaiticum, Pantala flavescens; new for Italy.

Received 12.05.2012; accepted 22.12.2012; printed 30.12.2012

Proceedings of the L‘ International Congress “Insularity and Biodiversity”, 11*-13* May 2012, Palermo, Italy

INTRODUCTION

Because of local scarcity of aquatic biotopes, re-

sulting in low species richness, the dragonfly fauna

(Insecta, Odonata) of the Italian islands of the Sici-

lian Channel has received little attention. Indeed,

there are only three publications specifically dealing

with the Odonata of the Pelagic islands and Pantel-

leria (Consiglio, 1960; Lohmann, 1989; Pavesi &

Utzeri, 1995), while material from these islands is

also briefly discussed in other works (Carchini &

Di Domenico, 1992; Carfi & Terzani, 1993). Only

7 species are reported in the literature for the Italian

islands of the Sicilian Channel, including one only

known from historical records and considered lo-

cally extinct (Pavesi & Utzeri, 1995); of recently

460

A. CoRSO, O.Janni, M. Pavesi, M. Sammut, A. Sciberras & M.Vigano

reported ones, only 2 were found to breed, and of

these only 1 with a doubtless viable population.

This very low speeies riehness no doubt largely re-

sults from near-absenee of permanent or even tem-

porary water bodies, suitable for Odonata breeding

and larval development. Yet a lot of dragonfly spe-

eies are strong fliers, able to eover long distanees

as oeeasional vagrants or regular true migrants,

more so when supported by favourable winds, and

often also to breed, at least temporarily, even in ar-

tifieial, newly formed water bodies. Therefore the

little number of reeorded speeies is also due to li-

mited field work earried out over the years; on the

other hand, the number of aetually oeeurring spe-

eies may be subjeet to rapid ehanges, due to modi-

fieations of loeal eonditions, e.g. newly ereated

suitable biotopes. Other inseet groups attraeted

greater interest and have been the objeet of more

detailed studies, in parallel with the situation on

mainland Sieily (Corso, personal unpublished data),

whieh resulted in a elear, although limited, inerease

in the number of reeorded speeies.

The islands of the Maltese arehipelago, eom-

pared to Pantelleria and the Pelagie, despite of in-

tensive habitat destruetion still harbour many more

bodies of either fresh or braekish water, and loeal

Odonata reeeived a greater deal of interest. Lite-

rature about Malta and its satellite islands is varied

and extensive; in addition to historieal works (Val-

letta, 1949, 1957), an inereasing number of papers

in reeent years foeused on single speeies or on the

whole odonate fauna, ineluding diseussions of

their biology and loeal status (Ebejer et al., 2008;

Seiberras, 2008; 2011; Seiberras & Sammut, 2008;

2013; Seiberras et al., 2010; Gauei & Seiberras,

2010). This resulted in a more diverse known odo-

natologieal fauna, as far as eertainly or supposedly

breeding speeies are eoneemed: not less than 15

of the 1 8 speeies so far reeorded. Even so, further

additions are to be expeeted, sinee elimate ehanges

may result in formerly oeeasional or never found

speeies beeoming established, even with large po-

pulations. For example, two of the present-day

most abundant dragonflies, Orthetrum trinacria

and Trithemis annulata, were first reeorded only

after 2000.

From 2006 to 2012, we have visited the Pelagie

islands and Pantelleria with the primary purpose of

gathering omithologieal data, but we kept a high in-

terest in and have paid a great deal of attention to

other aspeets of islands biodiversity, ineluding Odo-

nata. This paper presents the results of our field

work, together with an overview of existing publi-

shed data. As a result of our field work, the number

of speeies reeorded for the Pelagie and Pantelleria

has more than doubled, with 20 speeies, ineluding

2 new for the Italian fauna.

ABBREVIATIONS. AC = A. Corso; AS = A.

Seiberras; COM = Comino; GOZ = Gozo; LIN = Li-

nosa; LMN = Lampione; LMP = Lampedusa;

MAL = Malta; MISC (Malati di Isolitudine alio Sta-

dio Cronieo) = a birding and nature group foeused

on islands of Mediterranean Basin; MP = M. Pavesi;

MS = M. Sammut; MV = M. Vigano; OJ = O. Janni;

PNT = Pantelleria.

MATERIALS AND METHODS

Taxonomy and nomenelature follow Dijkstra &

Lewington (2006) and Dijkstra & Kalkman (2012).

We eonsulted all the available literature on the

Odonata of our study area, and analysed both re-

eent and historieal works in order to note any po-

pulation trends (inereasing/deereasing/stable) and

have a baseline eheeklist against whieh to eompare

our findings.

Three of us (AC, OJ and MV) have visited Pan-

telleria and the Pelagie in spring (February-May),

summer (June -August), autumn (September-No-

vember), and a few winter visits in Deeember and

late January. More speeifieally, between April 2004

and November 2012 they have visited the Pelagie for

a total of almost 400 days, in the following periods:

April-May 2004, September-Oetober 2005, Mareh-

May 2006, September-Oetober 2006, Mareh-May

2007, September-Oetober 2007, January 2008,

Mareh-May 2008, June 2008, Oetober-November

2008, Mareh-July 2009, September-November 2009,

January 2010, Mareh-May 2010, Oetober-November

2010, February- April 2011, July- August 2011, Oe-

tober-November 2011, Oetober-November 2012. As

part of a LIPU-sponsored study on raptor migration

aeross the Mediterranean, they visited Pantelleria for

30-days periods in April-May 2005-2012 and Au-

gust-September 2008, during whieh they prospeeted

all potentially suitable habitats, and as general rule

tried to eover as mueh ground as possible, in order

to find both odonates breeding in loeal water bodies

and migrating individuals. Oeeasional field trips

Annotated checklist of the dragonflies (Insecta Odonata) of the islands of the Sicilian Channel

461

were also made by AS (April 2010 and September

2010) and MP (November 2012).

As for Maltese islands, two of the authors (AS,

MS) are resident on Malta, the largest one, and

were able to monitor the loeal odonate fauna all

over the last three deeades. All speeies but very few

have been photographed; voueher speeimens for

many have also been eolleeted and are now housed

in the private eolleetions of three of the authors

(AC, MP and AS). Eaeh speeimen eolleeted, whe-

ther mounted or in an envelope, is aeeompanied by

the following data: speeies (and subspeeies if ap-

plieable), sex, loeality (ineluding GPS eoordinates)

and date of eolleetion, number of observed indivi-

duals, eolleetor (legit) and responsible for the iden-

tifieation (det.). Determinations for eaeh reeord,

either based on material evidenee (voueher speei-

men/photo) or only on sighting, rely on the reeord’s

author himself, unless otherwise stated.

RESULTS AND DISCUSSION

Our study raises to 20 the number of odonate

speeies altogether reeorded for the Pelagie (20, of

whieh 2 tentatively) and Pantelleria (9, of whieh 1

with the only European large viable population),

vs. 7 previously reported in the literature (Table 1).

More exaetly, as for the Pelagie, reeorded speeies

riehness is as follows: Lampedusa (17, of whieh 2

tentatively), Linosa (13, of whieh 2 tentatively),

and Lampione (4). Two of the speeies reeorded in

the past for Pantelleria, one of whieh at that time

supposedly breeding, have never been found again

there during our survey (see below), while all the

speeies previously reeorded for the Pelagie were

repeatedly eonfimied.

It should be kept in mind that there is indeed

no reason for whieh any speeies found on one of

these islands would not to be expeeted, at least oe-

easionally, also in the others. All of them are rather

elose to eaeh other, therefore more or less equally

exposed to migratory influxes, aeeounting for most

odonate reeords, these islands being poor (Pantel-

leria) to almost or eompletely devoid of water bo-

dies (the Pelagie). Very few of these exist on

Lampedusa, where some speeies with evidenee or

presumption of breeding are reeorded, although

none is definitely known to have viable loeal po-

pulations; for instanee a fairly large, temporary

rain pool at Ponente, within the island’s nature re-

serve, a water-filled, shallow gravel pit at Albero

Sole, a temporary pool at the mouth of a mostly

dry ereek at Cala Puleino, or again man-made

small water reservoirs (see below, under Sympe-

trum fonscolombii). Newly built-up water eat-

ehments or reservoirs, to be kept filled with some

water throughout the year, where also aquatie ve-

getation may develop (e.g. within the Lampedusa

Island Natural Reserve), eould indeed result in

eonspieuous inerease of breeding speeies, with

some populations possibly beeoming established.

Some speeies may also, at least oeeasionally, breed

on Linosa, where no natural water bodies, yet here

and there some small reservoirs exist. Lampione

on the other hand is a small, not inhabited roek,

where not even the least water eatehment exists,

and therefore all Odonata there found are no doubt

migrants. Dragonflies oeeurrenee, sometimes mas-

sive, therefore results above all from migratory

events, mainly linked to strong winds from sou-

thern quadrants, namely “libeeeio” (from south-

west) and “seiroeeo” (from south-east). The

former above all seems to aeeount for most of the

influxes, beeause of mueh lesser distanee, between

the islands and the North- Afriean eoast, in south-

west direetion rather than in south-east one. Winds

from north-east (“greeale”) or north (“tramon-

tana”), eooler than southern ones, are usually not

eonsidered as souree of influxes, yet they may ae-

eount for oeeasional arrivals Ifom Sieily, ineluding

weak fliers sueh as zygopterans.

Differenees in number of reeorded speeies the-

refore result above all from different amount of ob-

servations. Lampedusa, the only one of the Pelagie

with an airport, is the easiest to reaeh, even with

stormy sea eonditions, while the other two ean be

reaehed only by ship. Exeeedingly low number of

speeies reeorded for Lampione partly relies on laek

of observations, only resulting from short visits,

beeause of non-existing loeal faeilities; on the

other hand, on its most limited extension, eompa-

red to Lampedusa and Linosa, therefore on its har-

dly aeting as rest site for migrant individuals.

Moreover, during strong winds periods, usually the

best situation to observe dragonflies on the Pela-

gie, reaehing the islet may prove impossible.

Pantelleria shows somewhat different eondi-

tions. Besides being by far the largest and highest

in altitude (836 m), it has a permanent saline lake.

462

A. CoRSO, O.Janni, M. Pavesi, M. Sammut, A. Sciberras & M.Vigano

SPECIES

PNT

LMP

LIN

LMN

STATUS

NOTES

ZYGOPTERA

CALOPTERYGIDAE

Calopteryx haemorrhoidalis

aeeidental

C. Virgo meridionalis

no reeords

aeeidental on Maltese islands

COENAGRIONIDAE

Ischnura fountaineae

breeding

Linosa reeords at reservoirs

Ischnura genei

breeding?

ANISOPTERA

AESHNIDAE

Aeshna mixta

oeeasional?

large migrant swarms regularly

observed in Sieily

Anax imperator

migrant, irregular

breeding

oviposition and exuviae reeorded on

Pantelleria

Anax parthenope

migrant, irregular

breeding?

oviposition reeorded on Pantelleria

Anax ephippiger

migrant, possibly

breeding

potentially suitable biotopes on

Pantelleria

LIBELLULIDAE

Orthetrum brunneum

no reeords

rare on Malta, breeding not eonfirmed

Orthetrum nitidinerve

vagrant

single reeord; oeeasional migration

reeorded on Malta

Orthetrum coerul. anceps

vagrant?

single reeord; common on Malta

Orthetrum chrysostigma

no reeords

records on Malta, status uncertain

Orthetrum eaneellatum

vagrant, formerly

breeding?

large population in the past on

Pantelleria, extinct

Orthetrum trinaeria

vagrant

migratory species, recently established

on Maltese islands

Crocothemis erythraea

oeeasional migrant,

irr. breeding?

oviposition recorded on Lampedusa,

breeding not confirmed

Sympetrum fonscolombii

migrant, irregular

breeding

large numbers regularly observed

Sympetrum striolatum

migrant, irregular

breeding?

oviposition recorded on Lampedusa,

breeding not confirmed

Sympetrum meridionale

vagrant

single record

Sympetrum sinaitieum

migrant

regular in winter-early spring, no

reproductive behaviour

Braehythemis impartita

oeeasional migrant

scattered records; established on

Italian major islands

Trithemis annulata

vagrant

recently established on Maltese islands

Trithemis kirbyi

vagrant

single record

Selysiothemis nigra

no reeords

breeding on Maltese islands

Pantala flavescens

vagrant

single cluster of records; migratory

species

Table 1. Odonata of the Pantelleria and Pelagie islands.

Annotated checklist of the dragonflies (Insecta Odonata) of the islands of the Sicilian Channel

463

SPECIES

MAE

GOZ

COM

STATUS

NOTES

ZYGOPTERA

CALOPTERYGIDAE

Calopteryx haemorrhoidalis

vagrant

loeal populations never reeorded,

despite potentially suitable habitats

C. Virgo meridionalis

vagrant

COENAGRIONIDAE

Ischnura fountaineae

no reeords

possibly overlooked

Ischnura genei

breeding

ANISOPTERA

AESHNIDAE

Aeshna mixta

oeeasional

Anax imperator

breeding

partly deelining, displaeed by

A. parthenope?

Anax parthenope

breeding

Anax ephippiger

migrant, irregular breeding

exuviae reeorded; no viable population

known

LIBELLULIDAE

Orthetrum brunneum

breeding?

rare, breeding not eonfirmed

Orthetrum nitidinerve

oeeasional

single eluster of reeords

Orthetrum coerul. anceps

breeding

Orthetrum chrysostigma

breeding?

seattered reeords, single emergenee

reeorded; possibly underestimated

Orthetrum eaneellatum

breeding

declining because of 0. trinaeria

settlement

Orthetrum trinaeria

breeding

recently established, abundant

Crocothemis erythraea

breeding

Sympetrum fonscolombii

breeding

strongly declining, locally disappeared,

overcome by O. trinaeria

Sympetrum striolatum

formerly breeding?

no records in last years, overcome by

O. trinaeria

Sympetrum meridionale

no record

Sympetrum sinaitieum

no reeords

possibly overlooked

Braehythemis impartita

no reeords

scattered records on Linosa,

established on Italian major islands

Trithemis annulata

breeding

recently established

Trithemis kirbyi

no reeords

Selysiothemis nigra

breeding

Pantala flavescens

no reeords

possibly overlooked

Table 2. Odonata of the Maltese islands.

464

A. CoRSO, O.Janni, M. Pavesi, M. Sammut, A. Sciberras & M.Vigano

with the only known viable population of Odonata

throughout these islands, and a fair number of man-

made water reservoirs and eatehments, where oeea-

sional breeding of some speeies was observed. The

lower number of reeorded speeies, eompared with

Lampedusa or even Linosa, ean therefore only result

from inadequate prospeeting, partieularly in some

periods (see above); little doubt exists that absenee

of observations during the appropriate periods ae-

tually aeeounts for non-existing loeal reeords of e.g.

Sympetrum sinaiticum or Pantala flavescens.

Two speeies reported for Pantelleria in the lite-

rature (Pavesi & Utzeri, 1995) were not eonfirmed

during our survey. Orthetrum cancellatum, reeorded

in 1875 in large numbers and probably breeding at

that time, was never found again and is very likely

to be loeally extinet. O. trinacria, reeorded in 1984

upon a single individual, was possibly never establi-

shed. Breeding was reeorded or supposed throu-

ghout different islands for some speeies, yet at

present a definitely viable population is only known

on Pantelleria: Ischnura fountaineae. Sympetrum si-

naiticum and Pantala flavescens are new for Italy;

Calopteryx sp. ef. haemorrhoidalis, Ischnura genei,

Aeshna mixta, Orthetrum nitidinerve, O. coerule-

scens anceps, Crocothemis erythraea, Sympetrum

striolatum, S. meridionale, Brachythemis impartita,

Trithemis annulata and T. kirbyi are first reeorded

for the Italian islands of the Sieilian Channel.

The list of speeies reeorded on Maltese islands

at present remains at 1 8, the most reeent being Or-

thetrum chrysostigma; at least 15 are eertainly or

supposedly breeding (Table 2).

Here follows a systematie and eommented list

of the reeorded speeies.

Subordo ZYGOPTERA Selys, 1854

Family CALOPTERYGIDAE Selys, 1850

Calopteryx haemorrhoidalis (Vander Linden,

1825)

Seiberras & Sammut (2008; reported by Boudot

et al., 2009) aseribed to C. virgo (Linnaeus, 1758),

after some doubts about its possible belonging to

haemorrhoidalis, one speeimen, found dead in a

roeky pool at Marsaseala, Malta, eonsidering it as

possibly intermediate between ssp. meridionalis

Selys, 1873 and ss^.festiva Brulle, 1832; few fur-

ther reeords for Gozo exist, one of whieh was ehee-

ked by AS and proved identieal to Marsaseala spe-

eimen. Yet the latter, although badly damaged by

dermestid beetles (only three wings and small de-

bris of thorax still remain), from the pieture rather

resembles a C. haemorrhoidalis.

Wing length/width, eonsidered by Seiberras &

Sammut (2008) as diagnostie, proved an unreliable

feature, sinee eonsiderable variations may be obser-

ved even within a single population throughout the

season, depending on emergenee period: the earlier

the emergenee, the broader the wing, and also the

larger the individual size (Gallesi et al., in prepara-

tion). On the other hand, wing venation, dark blae-

kish instead of bright metallie blue, remnants of

thorax eutiele also dark blaek-bluish, basal elear

area of wings light tan instead of almost eolourless

hyaline, all argue for C. haemorrhoidalis and

against virgo s.l. However, having the speeimen

been found dead in a roeky pool, these eolour fea-

tures may also result from a post mortem alteration.

More reeently, Seiberras & Sammut (2013) dealt

with some additional eolleetion materials; besides

the above material, they also report and figure a

fairly preserved speeimen from Marsaseala, elearly

reeognizable as C. haemorrhoidalis (along with a

true meridionalis, see below). It is somewhat sur-

prising that no loeal population was ever notieed,

sinee in Maltese islands some flowing-water bodies

exist, very likely unsuitable for the highly rheophi-

lous and mierothermophilous C. v. meridionalis,

but potentially matehing C. haemorrhoidalis re-

quirements. Therefore the loeal oeeurrenee of C.

haemorrhoidalis is to be regarded as oeeasional,

eonsequent to either human aeeidental introduetion

(e.g. by ship) or to arrival of single straggler indi-

viduals, possible supported by natural events, sueh

as strong winds (Seiberras & Sammut, 2008) or

even floating debris drift. The likely origin areas are

Sieily or North Afriea; in both of them C. haemor-

rhoidalis is eommon and widespread, often with

large populations.

A single female on Linosa, 12.X.2009, not far

from Gala Mannarazza, near a small fountain in a

private garden (AC), was only briefly observed and

eould not be eaught; we eannot assign it to a definite

speeies, yet what eould be notieed argues for the

most likely C. haemorrhoidalis. This is the only

known reeord of a Calopterygidae for the Italian is-

lands of the Sieilian Channel, where no potentially

Annotated checklist of the dragonflies (Insecta Odonata) of the islands of the Sicilian Channel

465

suitable habitat for viable populations is known to

exist; small man-made fountains or springs, sueh as

the eoneemed one, may only prove temporary at-

traetive sites for straggler individuals.

Calopteryx virgo ssp. meridionalis Selys, 1873

Formerly reeorded for the Maltese islands (Sei-

berras & Sammut, 2008; again dealt with in Seiber-

ras & Sammut, 2013) upon poorly preserved

material, eonsidered as possibly intermediate bet-

ween C. virgo meridionalis and C. virgo festiva

Brulle, 1825. This reeord is herewith tentatively

aseribed to C. haemorrhoidalis (see above). Howe-

ver, the same authors (Seiberras & Sammut, 2013)

also found in a eolleetion another speeimen from

Dwejra (Gozo), elearly to be assigned to C. virgo

meridionalis. No other reeord is known, nor any

evidenee exists that the speeies ever oeeurred with

breeding populations in the Maltese islands. Sinee

virgo meridionalis is not known for Sieily (nor it

does oeeur in southern Italian mainland, replaeed

by forms supposedly referable to festiva), the likely

origin areas is only North Afriea, where the speeies

only oeeurs in few, seattered plaees.

Family COENAGRIONIDAE

Ischnura fountaineae Morton, 1905

The speeies in literature is often reported as 1.

fountainei; however, sinee it is dedieated to a

woman. Miss Margaret E. Fountaine, under provi-

sions of International Code of Zoologieal Nomen-

elature, 4th edition, art. 3 1 . 1 .2, the eorreet spelling

is fountaineae, as eorreetly reported e.g. in Dijkstra

& Lewington (2006) or in the website Fauna Euro-

paea.

Lohmann (1989) reported the first Italian (and

European) reeord for Pantelleria, at the voleanie

lake named Bagno dell’Aequa (the loeality name

used by the author and by subsequent ones) or

Speeehio di Venere, on the basis of sightings made

on 14 and 15.VIII.1984; subsequently larvae, exu-

viae and adults were seen and eolleeted by Pavesi,

Ratti, Carehini, Di Domenieo and others (Pavesi &

Utzeri, 1995). More preeisely, a very large number

of adults, partly emerging, and exuviae was reeor-

ded on 7.VIII.1985 (MP); no reproduetive beha-

viour was observed, possibly beeause of stormy

wind throughout the day, and no females with eo-

loration other than immature bright orange were no-

tieed. However, 1 female from Pantelleria with the

following data: VII. 1954, leg. E. Moltoni, with ma-

ture olive-greenish eoloration, was already housed,

without determination, in the Museo di Storia Na-

turale, Milano (Pavesi & Utzeri, 1995). Ragusa

(1875), about Odonata notieed on Pantelleria at the

lake, made no mention of any Ischnura', the speeies

may of eourse have been overlooked, yet it is well

possible that the speeies is a more reeent eoloniser,

not oeeurring there at that time. Besides supposed

ehanges in physieal eonditions (namely salinity),

the reported huge population of Orthetrum cancel-

latum may have prevented a small zygopteran from

beeoming established (see also below, under this

speeies). As for the Pelagie, a very teneral female

(MP det.) was found on Linosa, Monte Rosso,

19.IX.2010 (AS), where some Roman age small re-

servoirs (“gebbie”) exist. Further observations are

needed to assess whether small breeding popula-

tions of this speeies and the following one loeally

oeeur. Not reported for the Maltese islands, possibly

beeause overlooked.

From 2006 to 2012, we have monitored the

Speeehio di Venere population, the only one known

for Italy and Europe, in order to evaluate population

trends, identify aetual or potential threats and

launeh a genetie study (R. Ana Sanehez, unpubli-

shed). The population appears to be numerieally

stable, and although a slight deerease was noted in

2012, it may simply refleet later emergenee rather

than an aetual deeline; indeed, all our observations

were made in the months of April and May. Al-

though Pavesi & Utzeri (1995) report 1 male and 1

female eolleeted by Ratti on 2. V. 1984, we never ob-

served emergenee before 14.V. One of the said au-

thors (Utzeri) found no adults or larvae on

28.IV. 1991, so that they speeulated on possible

loeal repeated extinetions of the speeies beeause of

exeeedingly arid eonditions and therefore high sa-

line eoneentration in some years, followed by sub-

sequent reeolonisation; whieh may have aeeounted

for supposed absenee of the speeies in the said pe-

riod. The same authors, on the other hand, do not

exelude the eyele may be primarily univoltine, with

late mass emergenees, and partly (oeeasionally) se-

mivoltine, with some larvae delaying emergenee

until following year.

466

A. CoRSO, O.Janni, M. Pavesi, M. Sammut, A. Sciberras & M.Vigano

Despite intensive searehes, we never eould find

any larvae exeept for the days immediately preee-

ding emergenee. Considering the high numbers of

individuals oeeurring in the site, this very likely de-

pends on larvae behaviour, to remain hidden inside

thiek eane tufts until ready to emerge, and may have

been the real reason for the reported laek of fin-

dings. Considering observed emergenees starting

from May, and definitive eoloration of females

found in July, yet totally absent in the huge popula-

tion observed in August, there is no reason not to

suppose a bivoltine/multivoltine eyele, as usual in

Ischnura. On the other hand, this elusive larval

habit makes even more diffieult to monitor the sta-

tus of the Pantelleria population, under eurrent si-

tuation threatened with extinetion. Although the

Speeehio di Venere is a Nature Reserve, SPA (Spe-

eial Proteetion Area) and SCI (Site of Community

Importanee), it is in faet exposed to eontinuous and

very strong human pressure, fi-om both loeal people

and tourists swimming in the pond and walking

along the edges, so trampling the breeding area of

the speeies. Moreover, the eultivated fields imme-

diately adjaeent to the lake are regularly and heavily

subjeet to pestieides and fertilizers applieation. Ur-

gent aetions are strongly required for an effeetive,

not only virtual proteetion of this unique biotope;

the best preserved part of the area, delimited follo-

wing researehers’ indieations, should be elosed to

people’s transit and aeeess should be only allowed

for seientifie researeh, while the remaining eould

remain open for reereational purposes. Sueh mea-

sures should be of eourse strietly enforeed with ade-

quate patrol serviee by rangers (see also Pavesi &

Utzeri, 1995).

Ischnura genei (Rambur, 1842)

Reeorded for the Maltese islands already in past

literature (Ebejer et al., 2008; Boudot et al. 2009),

where it oeeurs with healthy populations on the is-

lands of Malta, Gozo and Comino. Never reeorded

for Pantelleria. As for the Pelagie, two speeimens,

male and female (MP det.), were eolleeted on Li-

nosa, Cala Mannarazza, 18.IX.2010 (AS). Although

the breeding is not eonfirmed (yet possible, see

above), the reeord is quite interesting, being an evi-

denee that migratory influxes from Sieily, mediated

by northern (“tramontana”) or north-eastern (“gre-

eale”) winds, may oeeasionally oeeur. /. genei, en-

demie of major (and some of the smaller) Italian is-

lands and Maltese arehipelago, does not oeeur in

North Affiea.

AS also observed two Isehnura individuals on

Lampedusa, Spiaggia dei Conigli, 16.IX.20 10, not

eolleeted, therefore not to be identified. They may

belong to either previous speeies, as well as to /. sa-

harensis AguQSSQ, 1958. Considering the proximity

with the Tunisian eoast, and the zoogeographie eom-

position of the Pelagie, loeal oeeurrenee, at least oe-

easional, of /. saharensis is not unlikely; the speeies

oeeurs with very large breeding populations e.g. on

Djerba island (AC, unpubl.). The above spottings,

as for number of speeies known for eaeh island (see

above), are treated as belonging to a single speeies.

Subordo ANISOPTERA Selys, 1854

Family AESHNIDAE Rambur, 1842

Aeshna mixta Latreille, 1805

It was not previously reeorded for the Pelagie or

Pantelleria (Pavesi & Utzeri, 1995). During our

study, we found it on Pantelleria, Punta Spadillo,

19.V.2011, 1 male (AC). This is the only reeord

known to us for the island. Single spotting were no-

tieed in 2010 on Lampedusa, Cala Calandra, 15. IX

and on Linosa, Monte Rosso, 1 9. IX (AS). In au-

tumn 2012, a rather relevant influx has been notieed

on the Pelagie, with notably up to 5 a day on Linosa

on last week of Oetober and first week of Novem-

ber, of both sexes (but mostly females), and single

individuals on Lampedusa by the end of Oetober

(AC, OJ, MV, I. Maiorano & G. Soldato). No si-

ghting was notieed on Lampedusa between 23. XI

and 29. XI (MP). In April and May, dozens and so-

metimes hundreds of immature individuals of this

speeies and of Aeshna ajjinis Vander Linden, 1 820

ean be seen migrating along the eastern eoast of Si-

eily (from Capo Peloro to Capo Passero), and many

are seen eoming in off the sea in the Syraeuse area,

probably arriving from the Sieilian Channel. The

paueity of reeords for Pantelleria and the Pelagie is

therefore rather puzzling and may only be due to

laek of observers during its peak migration period.

The present reeords eonfirm the speeies as highly

migratory and able to eover large distanees (Boudot

et al., 2009), therefore also to eolonise new areas or

to appear far out of range.

Annotated checklist of the dragonflies (Insecta Odonata) of the islands of the Sicilian Channel

467

Figures 1-6. Ischnura fountaineae, Specchio di Venere, Pantelleria (AC). Figure 1. Mature male. Figure 2. Immature

female, C-type. Figure 3. Mature male, display. Figure 4. Immature female, C-type, display. Figure 5. Mature female,

C-type. Figure 6. Mating pair (female C-type).

468

A. CoRSO, O.Janni, M. Pavesi, M. Sammut, A. Sciberras & M.Vigano

Recorded for the Maltese islands by Boudot et

al. (2009) upon a single specimen (Ebejer et al,

2008; Sciberras, 2008). During the spring of 2012

a total of six specimens were recorded for Malta,

Comino and Gozo (Sciberras & Sammut, 2013).

Anax imperator Leach, 1815

Pavesi & Utzeri (1995) report a single record of

1 female, collected at Mursia, Pantelleria, on

9. DC. 1994. During our study, we have repeatedly ob-

served it in May at various localities on Pantelleria:

Specchio di Venere, Punta Spadillo, Rukia, Rekhale,

Grazia, Scauri, Ghirlanda, Pian di Monastero. Both

sexes have been observed, with preponderance of

females. There are also records, documented with

photos, in VI-VII.2010 and 2011, and X.2011 (A.

Belvisi). At least some of these individuals may be

stragglers from Africa, or even Sicily. However,

breeding was photographically documented in 201 1,

when a few females were observed laying eggs and

exuviae were found in a water catchment pool in the

locality of Arenella (A. Belvisi). The species has

also been repeatedly observed, from April to Sep-

tember, on Lampedusa and Linosa (AC, OJ, MV, AS

& I. Maiorano), always in limited numbers compa-

red to the other two reported Anax species. Also

spotted on Lampione, 5.IV.2010 (AS).

In the Maltese islands it was always reported in

old literature as common to very common, deemed

to be the most abundant Aeshnidae; this possibly

resulted either from being the species in the past

much more common, or from confusion with the

sometimes similar yt. parthenope. Although it is in-

deed common and widespread, with several bree-

ding sites, as for abundance it is at present exceeded

by the latter. In most coastal water bodies, the for-

merly dominant A. imperator in recent years has

been progressively overcome by A. parthenope

(Sciberras, 2008). It can be observed searching for

prey in a variety of habitats, including cultivated

fields or urban areas, even far away from water. It

catches several insects, especially Diptera, but also

larger ones, including various Lepidoptera, e.g. La-

siocampa quercus (Linnaeus, 1758) and Vanessa

eardui (Linnaeus, 1758), and other Odonata. Some-

times it catches preys standing on vegetation or

ground (Sciberras, 2008), either lifting them or de-

vouring them on the spot; in Italy a female was ob-

served to hawk upon a motionless Orthetrum

caneellatum female perched on the ground, to block

it with legs over its wings and to feed upon it (MP,

unpubl.). At the same time this species also forms

an important food source for migrating birds, espe-

cially Hobby Falco subbuteo Linnaeus, 1758 and

European Bee-eater Merops apiaster Linnaeus,

1758 which are regularly seen catching it (MS, per-

sonal data).

Anax parthenope (Selys, 1839)

Recorded for Lampedusa (common in August,

mainly during southern winds periods) and for Pan-

telleria (a single record in 1994) by Pavesi & Utzeri

(1995) which consider it as regular and potentially

breeding, although with no evidence of the latter.

During our study it was repeatedly observed in

numbers, besides on Lampedusa, also on Linosa

and Lampione (AC, OJ, MV, AS & I. Maiorano),

e.g. on VII.2009 and 2010. On some days, in spring

(March to May), it arrives erratically by hundreds

from southern quadrants. This mass of individuals

forms an important food source for the breeding

Eleonora's Falcon Fa/co eleonorae Gene, 1839, and

for many other migratory birds (Corso, 2011), in-

cluding passerines (MISC, unpubl). At least attempt

of breeding was documented on Pantelleria at the

Specchio di Venere, where in 2009 it was observed

laying eggs. No reason however does appear, for

which water catchments, reported as oviposition

site for A. imperator (see above), could not be also

suitable for A. parthenope.

In the Maltese islands it is reported as common

(Ebejer et al., 2008), found by Sciberras (2008) to

be the commonest local aeshnid, generally domi-

nant in coastal areas, including a number of bioto-

pes formerly dominated by A. imperator. Mass

migrations have been reported, as well as small

numbers mixed with large migrating swarms of A.

ephippiger (Sciberras, l.c.). Definitely recorded for

Malta and Gozo. It is to note that records only based

on collection materials may lead to largely undere-

stimate its frequency, individuals being often excee-

dingly difficult to catch.

Anax ephippiger (Burmeister, 1839)

Recorded for Lampedusa only by Pavesi & Ut-

zeri (1995), with a single record in IV. 1987. The

same authors however suggested that it may have

Annotated checklist of the dragonflies (Insecta Odonata) of the islands of the Sicilian Channel

469

Figure 7. Anax imperator, mature male, found on the small pool at Arenella, Pantelleria, 6.X.2011, A. Belvisi. Figure 8.

Idem, egg-laying female, 1 .VIL201 1 . Figure 9. A. imperator, mature male, Speeehio di Venere, Pantelleria, 28.VI.201 1, A.

Belvisi. Figure 10. Exuvia of A. imperator, AveneWa, Pantelleria, 4.X.20 11, A. Belvisi. Figure 11. A. ephippiger, mature fe-

male, eaught at Speeehio di Venere, Pantelleria, 20.IV.20 11 during a passage of thousands through the Sieilian Channel

(AC). Figure 12. Idem, mature male.

470

A. CoRSO, O.Janni, M. Pavesi, M. Sammut, A. Sciberras & M.Vigano

been largely overlooked, and that possible breeding

in small natural or artifieial water eatehments, as

well as in the Bagno dell’Aequa, was to be verified,

eonsidering its ability to breed in temporary pools

in arid environments, beeause of very rapid larval

development. We have found it to be eommon du-

ring all our visits to Pantelleria and the Pelagie, in-

eluding Lampione (AC, OJ, MV & I. Maiorano).

On some spring days, espeeially in Mareh and

April, but oeeasionally from late February, thou-

sands of individuals ean be seen arriving off the sea

from North Afriea. Mueh like A. parthenope and

Sympetrum fonscolombii, they are an important

food souree for breeding Eleonora’s Faleon Falco

eleonorae and for many other speeies of migratory

birds (Corso, 2011), ineluding passerines (MISC,

unpubl.). For example, in Mareh- April 2011 an im-

pressive influx of thousands of individuals was re-

eorded in Sieily (Corso, personal data) and on

28.IV.20 11 almost two thousands individuals were

estimated at the Speeehio di Venere, with many spe-

eimens eolleeted (AC). Breeding was not doeumen-

ted; however, sinee the other two Anax were

definitely found (imperator) or supposed {parthe-

nope) to breed, the same habitats may prove suita-

ble also for A. ephippiger.

On Maltese islands the speeies is a regular mi-

grant from Mareh to Oetober, and breeding has re-

eently been eonfirmed on Malta (Seiberras, 2011).

Mueh like in Sieily, this author reports that the lar-

gest influx of migrants was noted in Mareh 2011,

with about 4,000 individuals estimated on

1 8.III.201 1 on the island of Gozo (Maltese arehipe-

lago).

Family LIBELLULIDAE Rambur, 1842

Orthetrum brunneum (Fonseolombe, 1837)

Not reeorded for the Pelagie or Pantelleria,

where on the other hand no suitable biotopes exist.

In the Maltese islands it is rare and only oeeurs in

some freshwater streams on Malta (Ebejer et al.,

2008).

Orthetrum nitidinerve (Selys, 1841)

A stream-dweller, on the Pelagie or Pantelleria

only reeorded upon a single sighting, on Lampedusa,

Cala Croee, 14.IX.20 10 (AS), obviously vagrant. As

for brunneum or eoerulescens aneeps, no suitable

habitat does exist anywhere on these islands.

Boudot et al. (2009) do not report it for the Mal-

tese islands. Only reeorded by Seiberras et al.

(2010), who report males and females for different

loealities on Malta, over a very short period only,

1 8-22.VII.2008, without any observed reproduetive

behaviour; never notieed sinee. Therefore the same

authors regard its oeeurrenee as resulting from mi-

gration not followed by sueeessful breeding, whieh

is very likely eorreet.

Orthetrum eoerulescens aneeps (Sehneider, 1845)

On the Pelagie or Pantelleria only onee obser-

ved, on Lampedusa, Punta Sottile, 14.IX.2010

(AS), most likely a vagrant. O. eoerulescens is re-

eorded for the Maltese islands by Boudot et al.

(2009), who refer to elinal populations, seemingly

intermediate between nominal subspeeies and coe-

rulescens aneeps (ef. aneeps IclinQ obsolescent)', yet

aeeording to Seiberras (2008; 2011), Ebejer et al.

(2008) and Gauei & Seiberras (2010), Maltese po-

pulations should be aseribed to ssp. aneeps. Their

taxonomie status and variability indeed deserve an

in-depth study. In faet, aeeording to Boudot et al.

(2009), mainland Sieily populations appear inter-

mediate between eoerulescens and aneeps. Mauer-

sberger (1994) and Dyatlova (2006) are of the same

opinion. On the other hand, studies earned out

throughout Sieily from 2006 to 2012, as for aeees-

sory genitalia have shown a number of intermediate

speeimens, together with others seemingly referable

to the nominal subspeeies, but none really matehing

ssp. aneeps as Sardinian or North Afriean animals

do (Corso, personal data). Two Maltese speeimens

eolleeted by AS, from the pietures were found (by

AC and MP) to be, one obviously a true aneeps, the

other a chrysostigma (see below). Unfortunately

part of Maltese Orthetrum material, sent for study

by AS to AC, never reaehed the latter and is defini-

tely lost. It is therefore at present rather diffieult to

know whether some other aneeps reeords aetually

refer to chrysostigma.

Orthetrum chrysostigma (Burmeister, 1839)

Not reeorded for the Pelagie or Pantelleria,

where nevertheless its oeeurrenee may me expeeted

Annotated checklist of the dragonflies (Insecta Odonata) of the islands of the Sicilian Channel

471

(Pavesi & Utzeri, 1995), because of its ability to

breed also in temporary pools in desertic environ-

ments, so presumably also in man-made water cat-

chments.

Boudot et al. (2009) do not report it for the Mal-

tese islands too. First records for the Maltese archi-

pelago, all from Malta, are reported by Gauci &

Sciberras (2010), a total of 4 females, from 2008 to

2010. One of these, found by Gauci at a man-made

small freshwater pond in the Ghadira Nature Re-

serve, a saline marshland, on 12.VI.2010, from the

photo is clearly newly emerged, no doubt locally,

and most likely in the very same spot, although exu-

via was not found. This record was also discussed

on Forum Natura Mediterraneo on 19-20.XII.20 12,

between Charles Gauci (“Selys”) and MP (“gom-

phus”). No other records exist; this may argue for

only temporary breeding, resulting from occasional

arrivals and followed by local extinction. However

a fully mature male collected by AS, first believed

to be coerulescens anceps, from the picture was re-

cognized by AC and MP separately as actually

being chrysostigma. It is to be stressed that light

stripes on the thorax, considered by the authors as

diagnostic, are obvious only in females and not

fully mature males when thorax is not yet covered

with pruinescence. Old males have dark, densely

blue-pruinose thorax, with obsolete, unconspicuous

stripes, therefore quite resembling anceps, and

when occurring in low numbers among large popu-

lations of the latter may remain overlooked and un-

noticed. As said above, because of partly lost

material, it is at present impossible to check some

of previous Maltese anceps records, maybe partly

to actually refer to chrysostigma.

Orthetrum cancellatum (Linnaeus, 1758)

Pavesi & Utzeri (1995) report a single historical

observation for Pantelleria by Ragusa (1875), refer-

ring to “hundreds” of individuals seen at the Spec-

chio di Venere, as well as to large numbers of dead

dragonfly larvae, supposedly belonging to this spe-

cies and to equally abundant Sympetrum fonscolom-

bii, on the bottom of the lake. The species was never

found again there, and the same authors presumed

it to be locally extinct, or at least to have become

extremely rare. Historical data about other groups

of insects lead to suppose at that time a much lesser

salinity of the lake, likely related to greater rainfall

amount. Increased salinity because of rainfall shor-

tage may have led to disappearance of O. cancella-

tum, providing in the same time a suitable, almost

predators-free habitat to the highly tolerant

Ischnura fountaineae (Pavesi & Utzeri, 1995). No

further record for the island exists, despite small

water catchments and reservoirs (see above, under

Anax imperator) being a well-known habitat of this

species. Not positively recorded for the Pelagic. On

20.XI.20 12, a supposed female of this species was

observed, but not caught or photographed, on Lam-

pedusa, Albero Sole, at a water-filled gravel pit

(AC). Few days later, on 23.XI.2012, quite close to

the previous spot, a large libellulid, tentatively re-

ferred to this species (possibly the very same indi-

vidual), was briefly observed at some distance,

perched on the ground of a small dirty road (MP).

It disappeared before a clear sighting was possible.

No other individual was seen.

In the Maltese islands it is recorded for Malta

and Gozo (Ebejer et al., 2008) as common and oc-

curring in any type of water bodies, including gar-

den ponds. However Sciberras (2008) reports it as

declining, possibly also because of progressive in-

vasion of biotopes by O. trinaeria. Some observa-

tions exist of O. cancellatum caught and devoured

by O. trinaeria.

Orthetrum trinaeria (Selys, 1841)

Pavesi & Utzeri (1995) report a single record for

Pantelleria, a sighting by Lohmann (1989) on 14-

15.V.1984, presumably a vagrant. The species was

collected on Lampedusa, Gala Croce and Spiaggia

dei Conigli, 14.X.2010 (AS), no doubt upon mi-

grant individuals. O. trinaeria is a strong flier, well

known as a migratory species (cfr. e.g. Fraser,

1936). No further records for any of the Italian is-

lands of the Sicilian Channel exist.

In Maltese archipelago large breeding popula-

tions occur on Malta and Gozo, first recorded in

2003 (Ebejer et al., 2008). Today the species is wi-

despread and common especially on Gozo, where

because of its highly aggressive territoriality and

predatory attitude, it has locally overcome most

other Odonata (Balzan, 2008; Sciberras, 2008;

Sammut, personal data), systematically chasing

away, when not hunting, any other dragonfly; cases

were reported of O. trinaeria preying upon O. can-

cellatum. Sympetrum fonscolombii, as well as the

472

A. CoRSO, O.Janni, M. Pavesi, M. Sammut, A. Sciberras & M.Vigano

more occasional S. striolatum, were reported to di-

sappear, following O. trinacria settlement (Sciber-

ras, I.C.). The only smaller dragonflies able to

coexist with large O. trinacria numbers seem to be

Crocothemis erythraea and Trithemis annulata, the

latter another recent coloniser, whose populations

had over last years a spectacular increase (Balzan,

I.C.; Sammut, personal data).

Crocothemis erythraea (Brulle, 1832)

This is one of the commonest and most wide-

spread odonates all over Italy, including Sicily

(Corso, personal data), also known for its migratory

attitude. It was not reported for Pantelleria or the

Pelagic by Pavesi & Utzeri (1995), who neverthe-

less anticipated possible future records. On the for-

mer island, man-made water reservoirs, such as

those mentioned as breeding sites fovAnax impera-

tor or Sympetrum fonscolombii, may well prove sui-

table for this species. In September 2005, it was

observed and photographed on Lampedusa for the

first time by M. Romano. In this island, we have

only seen it in July 2009, September 2010 and last

week of October 2012 (AC, AS). During the last si-

ghting, 26-28.X.2012, not less than 4 pairs were ob-

served mating and egg-laying in a small sewer

outlet in the old harbour of Lampedusa town (AC

& 1. Maiorano). No occurrence was noticed there

on 23-24.XL2012 (MP). Its occurrence on Lampe-

dusa is likely to be only occasional, due to the lack

of large enough, permanent water bodies; the actual

successful breeding in the above reported, heavily

polluted outlet remains in need of confirmation.

Also noticed on Linosa, Monte Nero and Cratere,

18.IX.2010 (AS).

Recorded for the Maltese islands by Ebejer et

al. (2008), Sciberras (2008), Boudot et al. (2009),

as the most abundant and widespread dragonfly all

over the Maltese archipelago, and one of the very

few small species surviving habitat invasion by Or-

thetrum trinacria.

Sympetrum fonscolombii (Selys, 1840)

Reported for Pantelleria, Lampedusa and Linosa

by Pavesi & Utzeri (1995), who mentioned nume-

rous records, including exuviae from Lampedusa,

Cala Pisana, and deemed it to be one of the com-

monest odonates on these islands, as well as the

possibly only species able to breed in man-made re-

servoirs on Lampedusa. Known for Pantelleria as

early as the 19th century (Ragusa, 1875). It is in-

deed quite common on all circum-Sicilian islands

(Terzani & Lo Cascio, 1997; Corso, personal data),

including those of the Sicilian Channel. Because of

its attitude to regularly migrate, it can very often be

seen in large numbers even on small islands com-

pletely devoid of any water body. We have regularly

observed egg-laying tandems on Pantelleria and

Lampedusa. On the former island, exuviae and dead

adults are often found in the Roman cisterns at the

San Marco acropolis (AC & OJ), and it can be seen

at various localities along the coast, especially near

the locality of Arenella, where a semi-permanent

rain-fed pool is present, but also quite far from

water, e.g. at Scauri, Punta Spadillo, Kamma, Ghir-

landa (AC & V. Penna). On Lampedusa we have

observed territorial behaviour, mating pairs and tan-

dems from June to November in various localities

(e.g. Albero Sole, Ponente, Cala Pulcino, Cala Pi-

sana, Cala Madonna, Cala Francese, Valle Imbria-

cola). Egg-laying has regularly been observed at

Ponente, in a fairly large, temporary rain pool in the

western clearing of the pine grove within the is-

land’s nature reserve (the latest recorded on

25.XI.2012, MP), and also in a water-filled, shallow

gravel pit at Albero Sole (see above, under Orthe-

trum cancellatum). As reported by Pavesi & Utzeri

(1995), large numbers of this species arrive, toge-

ther with Anax parthenope (Selys, 1839), during

summer (especially August), on southern winds; we

found this to happen not only in summer, but also

in spring (April-May) and autumn (September-Oc-

tober). At these times, hundreds and sometimes

thousands of individuals can be seen arriving off the

sea, as in many other European coastal sites (cfr.

Owen, 1958; Heyne, 1989). These influxes are

often accompanied by numerous migratory birds,

as noted for other species of migratory dragonflies

(Anderson, 2009). Indeed, it seems that this species,

together with Anax parthenope and A. ephippiger

(see above), is a key food source for many migra-

tory birds, especially raptors, as well as for breeding

Eleonora’s Falcons Falco eleonorae (Corso, 2011).

Numerous specimens, including late records (De-

cember), have been collected, held in AC and MP

collections. Pavesi & Utzeri (1995) report the si-

ghting of two unidentified Sympetrum on Pantelleria,

3. XII. 1992, tentatively attributed to fonscolombii as

Annotated checklist of the dragonflies (Insecta Odonata) of the islands of the Sicilian Channel

473

the most likely speeies. Although identifieation is

not eonfirmed, the late date indicate a winter acti-

vity for Sympetrum sp. already in the early 90’s, like

regularly observed nowadays in both Sicily and Si-

cilian channel islands (Corso, personal data).

In the Maltese islands formerly considered very

common and widespread, it has drastically declined

in recent years and locally disappeared, because of

predatory pressure from the increasingly abundant

Orthetrum trinacria. It is still quite common in

sites, mostly in coastal areas, not suitable to support

breeding populations of O. trinacria.

In nearly all specimens from the Pelagic exami-

ned by us, the yellow area at the base of hind wings

is highly reduced to hardly visible, on fore wings

almost always absent. Also body size is somewhat

diminutive. North African populations have an

identical pattern (Corso pers. obs.; C. Mancin pers.

com). However a series of specimens collected on

Ponente, 23.XI.2012 (MP) mostly have indeed on

hind wings the yellow area reduced, reaching up to

about mid-length between base of wing and base of

triangle, not invading the median space; yet in two

the yellow reaches up to the triangle base, with me-

dian space lightly tinged, and in another the yellow

encloses the entire triangle and also fore wings are

distinctly tinged at base. In Sicilian populations, the

extent of yellow area is generally larger, and the

same proved for Maltese ones, the yellow on hind

wings often reaching about the base of triangle,

upon an overview (MP) of photos, on the website

Forum Natura Mediterraneo, by Albert Floridia

(“xilpa”) and Charles Gauci (“Selys”).

Since large numbers of S.fonscolombii regularly

migrate across the Mediterranean, the existence of

genetically separate and morphologically different

populations on their north and south sides is highly

unlikely. It is to be stressed that most of the indivi-

duals occurring on the Pelagic or Pantelleria very

likely originate from Africa; although this species

is definitely recorded to breed e.g. on Lampedusa,

there is little doubt that no viable resident popula-

tion actually does exist, the locally emerged stocks

likely being inadequate in numbers to support a

such population, also because of local aquatic bio-

topes becoming completely dried-up all over sum-

mer; and on the other hand, locally emerged stocks

being totally overcome by large, ceaseless influx

from North Africa. The situation on Maltese islands

is quite different; given the greater distance from

North African coast, as well as the local existence

of suitable biotopes with large breeding popula-

tions, it can be supposed that migrant individuals

are a minority compared to whose locally emerged,

and that most of individuals found on Maltese is-

lands originate from locally breeding populations.

Influxes from Sicily, if any, are supposed to be nu-

merically insignificant.

Evidences exist that exposure of larvae to dif-

ferent average temperatures prior to emergence

may result both in reduced marking pattern and in

decreased size of adults. Waiting for a confirmation

from rearing experience, we speculate that occa-

sional individuals with more or less large yellow

pattern found on Lampedusa may result from local

breeding, while the majority, with reduced pattern,

may be of North African origin. The Maltese is-

lands do not have an obviously different climate,

when compared with the Pelagic, yet unlike the lat-

ter ones they harbour large viable populations (see

above), which may account for having Maltese in-

dividuals usually larger yellow pattern than those

from the Pelagic.

Sympetrum striolatum (Charpentier, 1840)

Not reported for the Pelagic or Pantelleria by Pa-

vesi & Utzeri (1995), nor by other authors, it ac-

tually occurs on all these islands (not yet reported

for Lampione), as well as on other circum- Sicilian

islands, although erratically and in much smaller

number than the previous species (Corso, personal

data). No spotting was noticed in autumn 2012 (AC,

MP). It occurs in spring (March-May) and again in

autumn (October-November). However, from seve-

ral specimens collected and held in AC collection,

all spring individuals, as well as for late winter to

early spring S. sinaiticum (see below) are quite aged

according to general appearance and worn-out

wings, which obviously indicates they have over-

wintered, either on the Pelagic or in North Africa.

Absence of February records, contrary to S. sinai-

ticum, may be simply coincidental and only result

from insufficient observations. S. striolatum may

breed at least on Lampedusa, at Ponente, where we

have occasionally seen pairs in tandem; yet no evi-

dence from exuviae or newly emerged individuals

does exist.

In the Maltese Islands already in the past it was

found to be scarce; only in a single instance, in

474

A. CoRSO, O.Janni, M. Pavesi, M. Sammut, A. Sciberras & M.Vigano

2007, a fairly good number of individuals was no-

ticed (Sciberras, 2008). Strongly declined, if not to-

tally disappeared in recent years (the last record on

18.VI.2009), most likely because of its breeding

sites being progressively invaded by Orthetrum tri-

nacria (Sciberras, 2008).

Sympetrum meridionale (Selys, 1841)

The hitherto only record for Sicilian Channel is-

lands is a male, caught and subsequently released

on Lampedusa, Cala Croce, 14.IX.2010 (AS), no

doubt a straggler. The species, widespread in North

Africa, is known to migrate over long distances.

Sympetrum sinaiticum Dumont, 1977

Not recorded previously for the islands of the

Sicilian Channel, or anywhere else in Italy. On mid-

March 2009, at various localities on Lampedusa,

we observed several unfamiliar Sympetrum indivi-

duals, quite pale, the wings without any trace of

basal yellow spot, obviously quite aged according

to general appearance and worn-out wings; particu-

larly male features, such as pattern of thorax and

abdominal segments, matched none of the locally

likely species we are familiar with, such as S. strio-

latum, S. meridionale or S. fonscolombii, conver-

sely these argued for the Saharan and Near-Eastern

Sympetrum sinaiticum Dumont, 1977. The females,

without close-up view or in-hand examination, re-

mained somewhat puzzling. An individual of the

same species was seen on 14.IX.2010 (AS). In Fe-

bruary 2011, several individuals, of which 4 males,

were observed again in various localities on Lam-

pedusa (AC); also on Linosa individuals, possibly

belonging to the same species, were observed later

in March 2011, yet no close enough to enable defi-

nite identification. Finally, on 12-15.IV.2012, of 6

males seen on Lampedusa at Albero Sole, 4 mature

to aged could be collected (K. Bacon leg.), enabling

definite identification by AC; now housed in AC (2)

and MP (2) collections. All these likely reached

Lampedusa from North Africa supported by sou-

thern winds.

In North Africa S. sinaiticum breeds in winter

(Jodicke, 2003; Boudot et al., 2009), after which

many individuals disperse, often reaching the coast

of Tunisia. For instance, numerous individuals were

seen on the island of Djerba (Gulf of Gabes) in Fe-

bruary 2009-2011 (AC, personal data). Emergences

take place in late spring, but the newly hatched in-

dividuals estivate, often travelling far from the

emergence site, and do not breed until late autumn

or early winter. Therefore during years with high

breeding success, some individuals disperse nor-

thwards, and under favourable conditions may

reach the southernmost islands of the Sicilian Chan-

nel, such as Lampedusa, more often in February-

April. Although a single record on September exists

(see above), it should be noted that throughout Oc-

tober/November 2012, when strong southern wind

periods resulted in outstanding records of North

African dragonflies (see also under Trithemis kir-

byi), not a single sighting was noticed (AC, MP). It

should also be stressed that only more or less aged

individuals were observed on Lampedusa; moreo-

ver no mating pairs or ovipositing tandems were

ever noticed, despite observations period falls into

reproductive season of the species. Therefore at pre-

sent no evidence exists even to suppose that the spe-

cies may breed on Lampedusa, despite its

seemingly quite regular occurrence from late winter

to early spring.

On the other hand, S. sinaiticum breeds in de-

sertic environments, in temporary, summer-dry

water bodies, and European populations, wide-

spread in Mediterranean Spain (Dijkstra & Lewin-

gton, 2006; the same authors suggest it may be

overlooked elsewhere), are reported to breed also

in concrete water reservoirs, where exuviae were

found. Therefore further investigations are needed,

in order to assess the actual status in the Pelagic, as

well as likely winter occurrence, or even possible

breeding, on Pantelleria. Never reported for Maltese

islands, possibly because overlooked.

Brachythemis impartita (Karsch, 1890)

Only recently (Dijkstra & Matushkina, 2009) re-

cognized as separate species, a long time confused

with the supposedly unmistakable B. leucosticta

(Burmeister, 1839). Therefore, all papers cited in

the following text refer to the species as to leuco-

sticta. The latter is not known to occur north of Sa-

hara (Dijkstra & Matushkina, 2009).

Previously unrecorded for the Pelagic or Pantel-

leria; on Linosa 1 male found on 20.X.2009 (AC),

and again spotted on Monte Vulcano, 7.IV.2010, 1

female according to entirely colourless wings (AS).

Annotated checklist of the dragonflies (Insecta Odonata) of the islands of the Sicilian Channel

475

Figures 13-20. Sympetrum sinaiticum, male, Lampedusa, Albero Sole, 15.IV.2012, the first eonfirmed reeord for Italy of this

species, widespread in North Africa; some distinctive features. Figures 13-14. Two specimens (MP coll.). Figure 15. Flead,

frontal view. Figure 16. Idem, ventral view. Figure 17. Thorax, lateral view. Figure 18. Accessory genitalia, lateral view.

Figure 19. Terminal appendages, lateral view. Figure 20. Idem, ventral view. Note highly reduced dark markings, pale wing

venation, hind wings totally devoid of hasal amher spot, peculiar accessory genitalia. Photos 13, 14, 17, 18 hy M. Zilioli.

Healthy populations of this species are found in cen-

tral and Northern Tunisia and throughout southern

Sicily (Galletti et al., 1987; Boudot et al., 2009;

Corso, personal data) and whole Sardinia. As such,

considering the ability of the species to breed also in

water reservoirs, colonisation of the islands of the Si-

cilian Channel would not be unexpected, as sugge-

sted already by Pavesi & Utzeri (1995). The Linosa

records came, the former on a rather late date, the lat-

ter on a quite early one, compared to the main flight

period of this species in mainland Sicily (Corso, per-

sonal data), arguing for a North African origin, where

the flight season extends into October (Dijkstra &

Lewington, 2006). Indeed, much of the Pelagic fauna

originates from North Africa rather than from Sicily,

both with regards to arthropods (cf Massa, 1995) and

birds (Corso, 2005). On the other hand, B. impartita

is a well-known migrant, since all Italian established

populations result from recent colonisation from

North Africa (Galletti et al., 1987; Pavesi & Utzeri,

1995), as well as South European ones; previous re-

cords for North Mediterranean countries were refer-

red to vagrant individuals not breeding in the area.

Not recorded for the Maltese islands.

476

A. CoRSO, O.Janni, M. Pavesi, M. Sammut, A. Sciberras & M.Vigano

Trithemis annulata (Palisot de Beavois, 1807)

Only reported for Pantelleria and the Pelagie

through the sighting of 2 males and 1 female on

Lampedusa, Gala Calandra, 14.IX.2010 (AS); ad-

ditional reeords are to be expeeted, given the reeent

eolonisation and very rapid population inerease in

the nearby Maltese islands, where it was first reeor-

ded in 2005. From 2007 forward, regular reeords

followed (Seiberras et al., 2007; Balzan, 2008; Ebe-

jer et al., 2008). In a very short time, the observa-

tions beeame more frequent and the population

inereased rapidly. The first exuviae were found on

Malta, at Chadwiek Lakes and the Chinese Gardens

in Santa Lueia. In reeent years, exuviae have also

been found at il-Qattara (Gozo). Today it is the se-

eond most abundant anisopteran species in the Mal-

tese islands, superseded only by Crocothemis

erythraea. It seems to be among the few small dra-

gonflies to thrive in territories dominated by Orthe-

trum trinacria (MS, personal data). T. annulata is

indeed extremely aggressive, also towards other

species; in Greece it was seen attacking and chasing

away considerably larger dragonflies such as O.

cancellatum or even Lindenia tetraphylla (Vander

Linden, 1825) (MP, unpubl.). Some cases are

known in which habitat colonisation by T. annulata

resulted in local decline of the formerly abundant

C. erythraea (see Balzan, 2008; MP, unpubl.).

Trithemis kirbyi Selys, 1891

Not previously reported for Pantelleria and the

Pelagie, nor for Maltese archipelago. On 28.X.2012

morning 1 male and 1 female were very well and

long observed at Capo Grecale, Lampedusa (AC &

R. Finati), yet they proved impossible to catch; al-

ready in the early afternoon of the same day they

were no longer to be seen, nor any further sighting

occurred later. This constitutes also the first record

for Sicily and the first Italian record outside Sardi-

nia, where it was recorded on 23.VL2003 at Oridda

stream, Villacidro (VS) (Holusa, 2008) but never

noticed since, despite active researches in recent

years (B. Kunz, pers. comm.).

The record for Lampedusa raises some doubts

about the actual status of T. kirbyi in Italy. Although

the species may breed also in water reservoirs, there

is little doubt that the above individuals did not re-

sult from local breeding on the island, nor else-

where on the Pelagie or Pantelleria. The above re-

cord shows that T. kirbyi, at least with favourable

conditions of southern winds, is able to migrate

over long distances. Indeed, the simultaneous oc-

currence on the Pelagie, during a period of sustai-

ned “libeccio” (south-western wind), of two

African, never locally recorded species such as T.

kirbyi and Pantala flaveseens may hardly be regar-

ded as coincidental. Such wind periods are also par-

ticularly fruitful for observation of several bird

species migrating across the Sicilian Channel; the

above-said one also resulted in several North Afri-

can bird species records in the Mediterranean basin

(Corso, unpublished data). The minimum distance

between southern Sardinia and Tunisia is about 1 80

km, no doubt within the reach of even a less strong

flier, when supported by wind. Given the complete

lack of further records, in subsequent years and de-

spite intensive researches, in the concerned locality

or elsewhere in Sardinia, the actual occurrence of

breeding Italian populations of T. kirbyi clearly

needs confirmation.

Selysiothemis nigra (Vander Linden, 1 825)

Not recorded for Pantelleria or the Pelagie.

Given the status and positive trend in the Maltese

islands, possible future records are to be expected.

The species is well known also for mass migrations

(Fraser, 1936); on the other hand, Compte-Sart

(1960) reports for Mallorca (Balearic islands) large

populations breeding in concrete water reservoirs.

First recorded for the Maltese islands by Valletta

(1957), with 2 specimens collected in 1952; no fur-

ther records until 1996, when a specimen was col-

lected (AS). In 2007, a single specimen was

collected in July, at Ramla Bay, Gozo, and 5 fema-

les were observed in August in a burnt field at Tas-

Sellun, Xaghra, Gozo. During the same period, a

permanent population was discovered in two artifi-

cial pools in a valley at Marfa, Malta. In 2008 two

further, rather large populations were found at L-

Ahrax and Ghadira, Malta, where from 19. VII to

22 .VIII the species was observed daily with a ma-

ximum of 15 observed at the same time. Although

an increased abundance of S. nigra in recent years

is likely, it is suspected that in the past this species

was simply overlooked and may have been more

regular than believed. This is primarily due to its

elusive behaviour; its tendency to fly very low.

Annotated checklist of the dragonflies (Insecta Odonata) of the islands of the Sicilian Channel

All

along with small size and unconspicuous colouring,

make it more difficult to detect than other Odonata,

even more when the species is unexpected and the-

refore not expressly searched for. Since 2009, it

has been regularly observed in small numbers

even in other areas, including Buskett and Dingli

(Malta), especially on August- September (MS).

On 17. VII. 20 11 e.g., not less than 13 individuals

were observed at Cirkewwa (AS).

Pantala flavescens (Fabricius, 1798)

First records for Italy, and among the very few

for Europe, on Lampedusa and Linosa, X.2012 (OJ,

MV, AC, I.Maiorano & G. Soldato), no doubt in the

same days also to be found on Pantelleria. No lon-

ger recorded on 23-25.XI.2012 (MP). Because of

their outstanding interest, they will be thoroughly

dealt with in a separate paper. Never recorded for

Maltese islands.

SPECIES ERRONEOUSLY RECORDED TO

DELETE FROM LOCAL FAUNAL REPORTS

Trithemis arteriosa (Burmeister, 1839)

Reported for Malta upon a single specimen, an

aged female, collected in 2002 (Ebejer et al., 2008).

However, after re-examination the specimen was

found to be T. annulata (Sciberras, 2008), determi-

nation now confirmed (AC, MP). Although its oc-

currence may be not unlikely, at present T. arteriosa

is to be deleted from the list of Maltese Odonata.

ACKNOWLEDGEMENTS

We would like to express our heartfelt thanks to

our longstanding travel and research companions

and island lovers from our MISC group: Hans Lar-

sson, Igor Maiorano, and Lucio Maniscalco. For

their help in the field and unpublished data, we

would like to thank Giacomo Assandri, Giovanni

Soldato, Cosmic Mancin, Kevin Bacon, Andrea

Belvisi, Piero Ferrandes, Rosa Ana Sanchez, Sonia

Ferreira, Jean-Pierre Boudot, Bemd Kunz, Sonke

Hardersen, Elisa Riservato, Verena Penna and Rai-

mondo Finati. Finally, we thank Marco Gustin,

Elena d’ Andrea, and LIPU, who funded most of AC

visits to Pantelleria and partially to Lampedusa pri-

marily for ornithological studies. Also AS greatly

thanks his family for their support: Esther, Romario

and Jeffrey Sciberras. MS thanks his many field

companions, but especially Joseph Grech, Christo-

pher Cachia Zammit, and Dominic Frendo and the

late John Azzopardi and for various discussions and

communications with Charles Gauci who shares a

high interest in Odonata. MP also thanks Charles

Gauci for discussions about Maltese Odonata on the

website Forum Natura Mediterraneo. Finally many

thanks to Michele Zilioli (Museo di Storia Naturale,

Milano) for the beautiful photos of Sympetrum si-

naiticum material.

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Biodiversity Journal, 2012, 3 (4): 479-486

Preliminary notice on the genus Carabus Linnaeus, 1 758 (Co-

leoptera Carabidae) of the islands of Peter the Great Gulf in

the far East of Russia, Primorski province, Vladivostok area

with description of a new subspecies

Ivan Rapuzzi

Via Cialla, 47 - 33040 Prepotto, Udine, Italy; e-mail: info@ronehidieialla.it

ABSTRACT In the present paper the Fauna of Carabus Linnaeus, 1758 of some islands of the Peter the

Great Gulf in the Far East of Russia (Vladivostok area) is investigated. After the study of large

series of Carabus specimens from the islands and the mainland a new subspecies, Carabus

(Morphocarabus) hummeli putyatini n. ssp., is described and figured; moreover, comparative

notes with the closest taxa are provided.

KEY WORDS Carabus-, new subspecies; Peter the Great Gulf; Vladivostok; Russia.

Received 12.05.2012; accepted 24.06.2012; printed 30.12.2012

Proceedings of the L' International Congress “Insularity and Biodiversity”, May 11*-13*, 2012 - Palermo (Italy)

INTRODUCTION

The Peter the Great Gulf is the largest gulf of the

Sea of Japan adjoining the eost of Primorski Krai

(Fig. 1). The Muravyov- Amursky Peninsula and a

ehain of outlying islands divide the gulf of about

6,000 km^ into the Amur Bay and the Ussuri Bay.

The eoastline of about 1,500 kilometres is indented

by many smaller bays. The islands inelude two ar-

ehipelagos separate from the mainland by the Ea-

stern Bosphorus: Rimsky-Korsakov Arehipelago

eonsists of six small islands (the largest are Askol’d

Island and Putyatin Island) and few roeky islets (ke-

kurs) whieh are uninhabited and very hard to reaeh.

Eugenie de Montijo Arehipelago is a group of five

large islands (Popov Island, Shkota Island, Reyneke

Island, Russky Island and Rikord), and a lot of small

islands, islets and roeks (Figs. 2-5). Despite the vi-

einity with the large town of Vladivostok the genus

Carabus Linnaeus, 1758 of the islands is still little

known while the Carabus Fauna from the mainland

is well known and investigated from many years.

The first notiee for the genus Carabus for the Is-

lands of Peter the Great Gulf was the paper dated

1932 by Semenov and Znojko with the deseription

of C. (Aulonocarabus) praedo Semenov & Znojko,

1932 now eonsidered a mere synonym of C. (Aulo-

nocarabus) careniger careniger Chaudoir, 1 863 by

different Autors (Shutze & Kleinfeld, 1999; Bre-

zina, 2003; Deuve, 2004; ). After that only very few

information are available by the literature: C. (Au-

lonocarabus) canaliculatus jankowskiellus Deuve,

1991 and C. (Morphocarabus) hummeli vladoby-

dovi Obydov, 2009. Other works on Carabus of

these regions have been earried out by different au-

thors: Kraatz (1878), Haury (1879), Semenov-Tian-

Shansky (1898; 1906), Breuning (1932-1936),

Deuve (1991; 1997; 2012), Imura (1993), Ivanovs

(1993), Shilenkov (1996), Obydov (2005; 2009),

Deuve & Li (2007), Kleinfeld & Reuter (2009).

Untill the XXI eentury the knowledge of the Ca-

rabus from these Islands was poor and laeunose. In

the last 20 years thanks to the investigations of Mr.

Andrey Plutenko (Russia) who visited part of the

480

Ivan Rapuzzi

islands (Reyneke, Shkota, Putyatin, and Askol’d) a

lot of new data became available and new taxa were

described (Rapuzzi, 2010): C (Megodontus) vietin-

ghoffi rugicolor Rapuzzi, 2010, C. (Coptolabrus)

smaragdinus robinzoni Rapuzzi, 2010 and C. (Au-

lonocarabus) gossarei mareschii Rapuzzi, 2010.

In the present paper the results of the recent re-

search in the area are provided. The examination of

some Carabus specimens from Putyatin Island have

identified a new taxon following described.

ABBREVIATION. RC = I. Rapuzzi collection.

RESULTS

Carabus (Morphocarabus) hummeli putyatini

n. ssp.

Examined material. Holotypus male (Fig. 6),

Russia, South Pimorskiy region, Putyatin Island,

50-200 m, 14/22.VIII.2008, A. Plutenko leg. The

holotypus is deposited in the author collection. Pa-

ratypes: 32 males and 32 females, same data as ho-

lotypus. The paratypes are deposited in Plutenko

and author collections.

Description of Holotypus male. Length inclu-

ding mandibles: 28.3 mm, maximum width of ely-

tra: 9.8 mm. Color black with metallic luster on

dorsum brownish-violet with margin of pronotum

copper and green, margin of elytra golden-green

and primary foveae violet, shiny. Femora reddish-

brown. Head moderately thickened. Frontal impres-

sions deep anteriorly, exceeding anterior margin of

eyes; vertex slightly convex, surface of the vertex

slightly punctured, posterior margins of eyes with

punctures deep and subrugose; very short neck.

Mandibles short with dorsal surface smooth. Palpi

thin and long, labial palp bi-setose. Eyes very sa-

lient. Antennae extending with 5 antennomeres be-

yond pronotal base. Pronotum not sinuate,

transverse, about 1.37 times as broad as long, sli-

ghtly convex ; sides of pronotum very narrow mar-

gined, not bent upwards; hind angles very short,

truncate, slightly protruding behind its base, incli-

ned downwards; surface of pronotum strongly pun-

ctured, coarsely distributed, more dense and

roughly at sides and base; basal depressions small

and deep, roughly punctured with metallic luster.

Elytra very elongate, oval, convex, maximum width

behind middle; shoulders rather narrow and roun-

ded; sculpture triploid homodyname, intervals uni-

formly convex, interrupted in the row in quite long

links, slightly punctured striae. Male aedeagus (Fig.

10) characteristic of the species but with the median

portion larger, apex shorter and less curved.

Variability. The length of the body ranges from

22 mm to 30 mm. Very few variable color: dorsum

with more or less intense brownish- violet and mar-

gins green or golden-green. Pronotum sometimes

slightly less transverse. Intervals sometimes more

interrupted.

Etimology. The given name is dedicated in

honor to the famous Russian diplomatic Admiral

Yevfimy Putyatin (1803-1883).

Comparative notes. C. (Morphocarabus) hum-

meli Fischer von Waldheim, 1 823 is a very variable

species with several subspecies described from the

Vladivostok area: C. hummeli smaragdulus Kraatz,

1878 (Figs. 7, 11), C. hummeli pusongensis Imura,

1993 (= C. hummeli gustavi Shilenkov, 1996) (Figs.

8, 12), C. hummeli tristieulus Kraatz, 1878 (Figs. 9,

13), and C. hummeli vladobydovi Obydov, 2009.

C. hummeli smaragdulus: the new subspecies

differs by stronger body, more transverse pronotum,

different color, uniformly and regular intervals, and

shape of aedeagus. C. hummeli smaragdulus is the

closest subspecies to C hummeli putyatini n. ssp.

C. hummeli pusongensis and C. hummeli tristi-

eulus: C. hummeli putyatini n. sp. is definitely dif-

ferent by the larger size, transverse pronotum, color

and shape of aedeagus. C. hummeli tristieulus is the

geographical closest subspecies on the mainland.

C. hummeli vladobydovi: the new subspecies

differs for the larger size and stronger body, more

transverse pronotum and intervals less interrupted

forming longer links. C. hummeli vladobydovi was

described from Popov Island in the Eugenie de

Montijo Archipelago and seems to be more related

with pusongensis /tristieulus group.

List of the Carabus from the investigated islands

of Peter the Great Gulf

I. REYNEKE ISLAND

1. C. (Parhomopterus) Mannerheim, 1827

2. C. (Carabus) granulatus telluris Bates, 1883

3. C. (Carabus) arvensis faldermanni Dejean, 1829

4. C. (Morphoearabus) venustus Morawitz, 1862

The genus Carabus of the islands of Peter the Great Gulf in the far East of Russia with description of a new subspecies 48 1

Figure 1. Peter the Great Gulf in the far East of Russia. Figure 2. Reyneke Island. Figures 3, 4. Putyatin Island. Figure 5.

Askol’d Island.

482

Ivan Rapuzzi

Figure 6. Carabus (Morphocarabus) hummeli putyatini n. ssp., holotypus and (Fig. 10) aedeagus: lateral (left) and frontal view

(right). Figure 7. C. (M.) hummeli smaragdulus, Russia, Primorskyi reg., Ozyomyi klutsch river, A. Plutenko leg. (RC) and

(Fig. 11) aedeagus, lateral view. Figure 8. C. (M.) hummeli pusongensis, Russia, Primorskyi reg., Nadezhdenskyi distr., Ana-

n’jevka riv., A. Plutenko leg. (RC) and (Fig. 12) aedeagus, lateral view. Figure 9. C. (M.) hummeli tristiculus, Russia, Primorskyi

reg., Shkotovka river, 10 km East from Novaya Moskva vil., A. Plutenko leg. (RC) and (Fig. 13) aedeagus, lateral view.

The genus Carabus of the islands of Peter the Great Gulf in the far East of Russia with description of a new subspecies 483

Figure 14. Carabus (Aulonocarabus) gossarei Haury, 1879. Russia, Primorskiy reg., Shkotovskiy distr., Anisimovka vill., Li-

vadiyskiy Mt. Rug., 1000 m (CR). Figure. 15. C. (Aulonocarabus) gossarei mareschii, holotypus, Askol’d Island (CR). Figure

16. C. (Aulonocarabus) canaliculatus jankowskiellus. Russia, Primorskiy reg., Maravyov, Bogataya riv., Okeanskiy khrebet,

150/200 m (CR). Figure 17. C. (Aulonocarabus) canaliculatus jankowskiellus, Reyneke island (CR). Figure 18. C. (Aulono-

carabus) careniger. Russia, Primorje, Kamenushka (CR). Figure 19. C. (Aulonocarabus) careniger, Askol’d Island (CR).

484

Ivan Rapuzzi

Figure 20. Carabus (Megodontus) vietinghoffii caesareus Semenov, 1906. Russia, Primorskiy reg., Zanadvorovka vill. env.

(CR). Figure 21. C. (Megodontus) vietinghoffii rugieolor, holotypus, Reyneke island (CR). Figure 22. C. (Coptolabrus) sma-

ragdinus mandschuricus Semenov, 1898. Russia, Primorskiy reg., Ertem env., Ozemyi klueh (CR). Figure 23. C. (Coptolabrus)

smaragdinus robinzoni, holotypus, Reyneke island (CR). Figure 24. C. (Aeoptolabrus) sehrencki Motsehulsky, 1860. Russia,

Far East, Chemye Mts, Sukhaya (CR). Figure 25. C. (Aeoptolabrus) sehreneki hauryi, Putyatina Island, 50/200 m (CR).

The genus Carabus of the islands of Peter the Great Gulf in the far East of Russia with description of a new subspecies 485

5. C. (Aulonocarabus) canaliculatus jankowskiellus

Deuve, 1991 (Figs. 16, 17)

6. C. (Hemicarabus) tuberculosus Dejean, 1829

7. C. (Megodontus) vietinghoffii rugicolor Kapuzzi,

201 1 (Figs. 21, 26, and ssp. caesareus Semenov,

1906 Fig. 20)

8. C. (Acoptolabrus) schrencki hauryi Gehin, 1885

9. C. (Coptolabrus) smaragdinus robinzoni Ra-

puzzi, 2011 (Figs. 23, 29)

II. SHKOTA ISLAND

1 . C. (Parhomopterus) billbergi Mannerheim, 1 827

2. C. (Carabus) granulatus telluris Bates, 1883

3. C. (Carabus) arvensis faldermanni Dejean, 1829

4. C. (Morphocarabus) venustus Morawitz, 1862

5. C (Aulonocarabus) canaliculatus jankowskiellus

Deuve, 1991

6. C. (Acoptolabrus) schrencki hauryi Gehin, 1885

(Fig. 27)

7. C. (Coptolabrus) smaragdinus efr. mandschuri-

cus Semenov, 1898

III. PUTYATIN ISLAND

1 . C. (Parhomopterus) billbergi Mannerheim, 1 827

2. C. (Carabus) granulatus telluris Bates, 1883

3. C. (Carabus) arvensis faldermanni Dejean, 1829

4. C. (Morphocarabus) venustus Morawitz, 1862

5. C. (Morphocarabus) hummeli putyatini n. ssp.

6. C. (Aulonocarabus) careniger Chaudoir, 1863

(=praedo Semenov & Znojko, 1933)

7. C. (Acoptolabrus) schrencki hauryi Gehin, 1885

(Figs. 25, 28)

IV. ASKOL’D ISLAND

1 . C. (Parhomopterus) billbergi Mannerheim, 1 827

2. C. (Carabus) granulatus telluris Bates, 1883

3. C. (Carabus) arvensis faldermanni Dejean, 1829

4. C. (Morphocarabus) venustus Morawitz, 1862

5. C. (Aulonocarabus) gossarei mareschii Rapuzzi,

2011 (Figs. 14, 15)

6. C. (Aulonocarabus) careniger Chaudoiv, 1863

(=j??raer/o Semenov & Znojko, 1933) (Figs. 18, 19)

Figure 26. Carabus (M.) vietinghoffii rugicolor from Reyneke Island. Figure 27. C. (A.) schrencki hauryi from Reyneke

Island. Figure 28. C. (A.) schrencki hauryi from Putyatin Island. Figure 29. C. (C.) smaragdinus robinzoni from Reyneke

Island.

486

Ivan Rapuzzi

7. C. (Acoptolabrus) schrencki hauryi Gehin, 1885

8. C. (Coptolabrus) smaragdinus cfr. mandschuri-

cus Semenov, 1898

V. POPOV ISLAND

1 . C. (Morphocarabus) hummeli vladobydovi Oby-

dov, 2009

CONCLUSIONS

The Carabus inhabiting the investigated islands

of Peter the Great Gulf are elosely related with the

Carabus inhabiting the mainland. Five Carabus

taxa of subspeeifie rank are endemie to single is-

lands: C. (Morphocarabus) hummeli vladobydovi,

C. (Morphocarabus) hummeli putyatini n. ssp., C.

(Aulonocarabus) gossarei mareschii, C. (Megodon-

tus) vietinghoffi rugicolor, C. (Coptolabrus) sma-

ragdinus robinzoni. All the other Carabus are

distributed on the mainland and islands.

Further investigation will be neeessary to better

understand the Carabus fauna of the region.

ACKNOWLEDGMENTS

I wish to thank Mr. Audrey Plutenko for provi-

ding me the very interesting Carabus used for the

present study.

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sammelten Carabus. Deutsche Entomologische Zeit-

schrift, 22: 241-253.

Obydov D., 2005. Faune des Carabus de Siberie & d’Ex-

treme-Orient russe - II. Neocarabi. Collection syste-

matique, Vol. 11. Association Magellanes, Andresy,

130 pp.

Obydov D., 2009. Faune des Carabus de Siberie et d’Ex-

treme-Orient russe - 1. Lipastromorphi. Collection sy-

stematique, Vol. 20. Association Magellanes,

Andresy, 148 pp.

Rapuzzi L, 2010. Descrizione di tre nuove sottospecie

di Carabus provenienti dalle isole al largo di Vladi-

vostok nell’Estremo Oriente russo (Coleoptera, Ca-

rabidae). Lambillionea, 110: 310-314.

Schutze H. & Kleinfeld F., 1999. Carabusformen Zen-

tral-Asiens und Sibiriens - Taxa, Systematik, Bi-

bliogra-phie - mit einem besonders ausfuhrlichen

Fun- dorteverzeichnis. 2. Auflage. Gottingen/Furth,

242 pp.

Semenov-Tian-Shansky A., 1898. Symbolae ad congni-

tionem generis Carabus (L.) A. Mor., II, Horae So-

cietatis Entomologicae Rossicae, 31: 316-541.

Semenov-Tian-Shansky A., 1906. Analecta coleopterolo-

gica, XIII. Revue Russe d’Entomologie, 3-4: 150-156.

Semenov-Tian-Shanskiy A. & Znojko D., 1932. Nouvel-

les donnees a E etude du genre Carabus. IV. Comptes

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Biodiversity Journal, 2012, 3 (4): 487-492

Forest-ecological aspects of the genus Allardius Ragusa, 1 898

(ColeopteraTenebrionidae) in Sicily and Sardinia

Michele Bellavista' & Ignazio Sparacio^

'via C. De Grossis, 7 - 90135 Palermo, Italy; e-mail: miehele.bellavista@gmail.eom

^via E. Notarbartolo, 54 int. 13 - 90145 Palermo, Italy; e-mail: isparaeio@inwind.it

ABSTRACT The genus Allardius Ragusa, 1898 (Coleoptera Tenebrionidae) includes only two species: Al-

lardius oculatus (Baudi di Selve, 1876) endemic to Sicily and A. sardiniensis (Allard, 1877)

endemic to Sardinia. They are infrequent species in nature with few reports in entomological

bibliography. The authors describe and illustrate the larvae and the biological aspects of Allar-

dius. In particular, it is highlighted the strong saproxylophagous activity of these beetles and

the importance of their role in the ecology of a forest in relation to the presence of "dead wood".

KEY WORDS Tenebrionidae; Allardius; larvae description; Deadwood; saproxylic diversity.

Received 12.05.2012; accepted 01.10.2012; printed 30.12.2012

Proceedings of the P' International Congress “Insularity and Biodiversity”, May 11*-13*, 2012 - Palermo (Italy)

INTRODUCTION

The genus Allardius Ragusa, 1898 (Coleoptera

Tenebrionidae, subfamily Tenebrioninae, tribe He-

lopini, subtribe Helopina) ineludes only two speeies

medium-sized, winged, with noetumal habits and

summer phenology, sylvieolous and eortieieolous

(i.e. living under the bark); they show an obvious

sexual dimorphism with males that appear smaller,

opaque on the dorsal surfaee, with median tibiae

strongly eurved; the females, larger and more ro-

bust, with dorsal surfaee shiny and median tibiae

slightly eurved. The geographieal distribution is li-

mited to Sieily for A. oculatus (Baudi di Selve,

1876) (Fig. 1) and to Sardinia for A. sardininensis

(Allard, 1877) (Fig. 2) (Luigioni, 1929; Porta, 1934;

Gardini, 1995; Aliquo et al., 2007; Lobl et al., 2008;

Aliquo & Soldati, 2010).

A. oculatus was deseribed for Sieily by Baudi di

Selve (1 876) on a single female speeimen eaught in

the Madonie (see also Ragusa, 1 898) sent to him by

the Sieilian entomologist E. Ragusa. Allard (1877)

believed that these two speeies, beeause of the par-

tieular shape of the pronotum and the length of the

metastemum, should belong to a new genus whieh

was later deseribed by Ragusa (1898) as a subgenus

of Helops Fabrieius, 1775 and dedieated to Allard.

A. oculatus is eonsidered a rare speeies with very

few speeimens reported so far, reeently signalized

in mixed forests and eork at medium altitudes in

two Sieilian loealities (Caronia, Messina provinee

and Palermo, Addaura), illustrated and deseribed by

Aliquo et al. (2007) and Aliquo & Soldati (2010).

A. sardiniensis seems limited to Southern Sar-

dinia, partieularly in the area of Monte Sette Fra-

telli, Cagliari (Aliquo et al., 2007). It is reported in

various web pages for Monte Sette Fratelli, Cagliari

(http ://www.naturamediterraneo . eom/ forum/ topie . a

sp?TOPIC_ID=60 136; http://www.entomologiita-

liani.net/publie/ forum/phpBB3/viewtopie.php?f= 1 7

8&t=3105) and Monte Serpeddi, Musui, 800 m

asl,Cagliari, Bureei (http://www.naturalphotode-

sign.it/taxonomy/gruppi-sistematiei-trattati/Regno-

Animalia/Phylum- Arthropoda/ Classe-Inseeta/ Ordi-

ne-Coleoptera/F am. -Tenebrionidae/ Allardius-sardi-

niensis-Allard- 1 877-Sardegna.html).

488

Michele Bellavista & Ignazio Sparacio

On the whole, the researeh carried out in recent

years on the island faunas has allowed us to disco-

ver several numerous populations of the two species

of Allardius and make interesting, preliminary ob-

servations on their biology and ecology.

MATERIALS AND METHODS

Two Sicilian populations of A. oculatus (Pa-

lernio, Addaura, 100 m above sea level; Trapani,

Castellammare del Golfo, Monte Inici, 400 m asl)

(Figs. 3-6) and one population of A. sardiniensis

(Cagliari, Monte Sette Fratelli, Campu Omu, 455

m asl) (Fig. 7) have been investigated. The environ-

ments of collection were similar to each other, being

very dense wooded areas with a predominance of

oak (Quercus ilex L.) in the dead wood of which the

larvae of Allardius were developing. Observations

were conducted in the field, in the places of disco-

very and, later, in laboratory where the dead wood

attacked by the larvae of Allardius was transferred

by special boxes. It was thus possible to observe

and record periodically the various stages of larval

development (Figs. 8, 9) and, in the summer period,

to study the behavior of the adults. All sites of col-

lection, along with the wood attacked by the larvae

and all stages of the life cycle were photographed

with a Canon EOS 400D. The study, which began

in February 2010, for the Sicilian populations, and

in April 2011 for the one of Sardinia is still ongoing.

The studied material is preserved in the collections

of the authors.

RESULTS

Allardius oculatus. The study began with the

identification and gathering of branches of Quercus

ilex attacked by the larvae of Allardius. The wood

was collected directly on the ground, in the thick of

the forest, between the wet foliage litter and debris

at the foot of oak trees. The branches, whose dia-

meter ranged from 10 to 105 mm, often displayed

on the bark traces more or less extensive of fungal

attacks or holes out of Allardius emerged in pre-

vious years. Even the consistency of the branches

was variable in relation to the larval attacks suffe-

red; indeed, the larvae occur in the same wood with

different generations and in different years, digging

numerous tunnels into the wood until its complete

degradation (Fig. 10). In our study it was possible

to observe simultaneously larvae in varying degrees

of development, at the moment up to three years.

The tunnels are nomially more or less straight and

parallel to the length of the branch attacked but

often, in more advanced stages of wood degradation,

tend to collide with one another and to have irregular

course. The larvae are very active, move quickly and

use the urogomphi (i.e. paired horns) to defend

themselves; they develop into the wood with very

large populations and in almost absolute monopoly;

in our study we found larvae of Stictoleptura cordi-

gera (Fussli, 1775) (Coleoptera Cerambycidae) and

Anthaxia (Cratomerus) hungarica (Scopoli, 1772)

(Buprestidae). Despite this crowding condition, lar-

vae did not show any form of aggression or canni-

balism behavior. Pupae (Fig. 11) occur at the end of

May- June and last about twenty days. Adult flicke-

ring starts in mid- June and continues in July. The

flicker holes are circular and show a diameter of 5-

8 mm. Adults observed in the laboratory are quite

active and move quickly on the surface of dead bran-

ches; they stop very quickly and tie up at the point

where they are at the slightest glimmer of artificial

light; in the rest position they often adhere to the

wood with the head flexed forward and antennae

straight forward slightly broaden and bent at the

apex. Male is smaller than female and during mating

(Fig. 12) is located on her back holding her with the

front and medium legs placed at the sides of the ab-

domen and with the antennae on the sides of the pro-

notum. On some occasions we also observed 2-3

males on the back of a female (Fig. 13). The female

is under the male with the antennae straight and lean

forward, parallel to the substrate.

Description of the larva. The description is

based on a larva 28 mm long, 3.1 mm wide, head

capsule 2.4 mm broad (Figs. 14-16). The body is

brownish-yellow with blackish mandibles; cuticle

sclerotized, with shiny and very rugose surface of

tergites and stemites. Head prognathous, oval, sli-

ghtly tilted downward; vertex with 2 long setae be-

fore the posterior margin, 3 long setae on each

lateral margin and 4 long setae behind the anterior

margin. Clypeus little convex in lateral view, tran-

sverse, rounded at the anterior margin and particu-

larly at the sides, with two long setae before the

posterior margin, 3 long setae on each side and 3-4

little setae on the anterior margin. Labrum tran-

Forest-ecological aspects of the genus Allardius (Ragusa, 1898) (ColeopteraTenebrionidae) in Sicily and Sardinia

489

Figure 1 . Allardius oculatus from Sicily, Palermo, Addaura.

Figure 2. A. sardininensis from Sardinia, Monte Sette Fra-

telli, Cagliari. Figures 3-6. Monte Inici, Sicily: habitat of A.

oculatus with holes of A. oculatus in branches of oak (4-5).

Figure 7. Monte Sette Fratelli, Sardinia: habitat of A. sardi-

niensis. Figure 8. Larva of A. oculatus (Monte Inici). Figure

9. Larva of A. sardiniensis (Monte Sette Fratelli). Figures 1,

2, photos by M. Romano.

490

Michele Bellavista & Ignazio Sparacio

s verse, slightly eonvex in lateral view, with 2 long

diseal setae and 2 long setae on eaeh lateral margin.

Epipharynx with long, marginal and diseal setae.

Mandible asymmetrieal, strongly selerotized with

apiees distinetly bidentate and with robust teeth at

the base, more developed in the left part. Maxilla

with primary eardo, stipes, maxillary palpus and la-

einia; maxillary palpus three-segmented, with the

last segment pointed and with one long seta on the

outer edge of the seeond segment. Labium with di-

stinet prementum, mentum and submentum. Anten-

nae trimerous; antennomere I very short, wider than

long; antennomere II 4-4.5 times longer than the

first one; antennomere III similar to the seeond but

more slender and rounded at apex with 1-3 very

short setae. Prothorax wider than long with 4-5

setae situated to lateral margin and 2-3 on the dorsal

surfaee and with little and sparse punetures. Fore-

legs somewhat longer and stouter than mid- and

hindlegs; troehanters short, eovered by numerous

long setae; femurs elongated, eovered by numerous

long setae; tibiotarsi eovered by numerous short and

strong spines. Claws brown, pointed at apex, shorter

than tibiotarsi. Abdominal tergites 1-7 with setae

dorso-laterally; dorsal eutiele very bright, wrinkled

transversally and with the little punetures eoneen-

trated mainly in the anterior half of eaeh segment;

spiraeles small and eireular. Abdominal tergites 7

and 8 with large, deep, rounded holes. Abdominal

tergite 8 with two little elevated protuberanees, two

small dimples and two long setae in the middle of

the dorsal surfaee. Abdominal tergite 9 transverse

in dorsal view, irregularly rounded at apex and with

10 little setae, with two prominent, projeeting uro-

gomphi strongly and almost eompletely selerotized,

with 3 long setae situated laterally of eaeh urogom-

phus, 2 in the eenter posteriorly, and 3 on eaeh si-

tuated, respeetively, one to the base, one just above

and another at about mid-length, laterally. Lateral

parts of abdominal tergite 9 also with one small and

rounded prominenee and a larger one, protruding

and hunehed.

Pupa white with darker urogomphi and brown

elaws, mandible apiees and eyes. It is eharaeterized

by well developed lateral proeess and by abdominal

tergite 9 with a pair of urogomphi. These struetures

have an antipredator deviees (Steiner, 1995; Bou-

ehard & Steiner, 2004).

Allardius sardininensis. The observations ear-

ned out on the Sardinian population of A. sardini-

nensis are similar to those performed on A. oculatus

from Sieily and even the main biologieal eharaete-

risties of both larvae and adults are similar. The

branehes of oak attaeked, at least the ones we ob-

served, had a smaller diameter (max 6 em) and the

larvae show some morphologieal differenees (Figs.

17-18). In partieular, the body appears more disten-

ded posteriorly, the dorsal eutiele is less wrinkled

and more dotted, mainly on the abdominal tergite

7, the antennomere III is more elongated and more

rounded at the apex, the elypeus is less rounded at

the sides; the abdominal tergite 9 has holes larger,

deep and rounded; the dimples in the middle of the

dorsal surfaee are larger and deeper.

CONCLUSIONS

Although the present work on the populations

of A. oculatus and A. sardiniensis is still in progress,

nevertheless, it has already allowed us to make im-

portant observations, most of whieh are original and

unpublished, ineluding the deseriptions of the lar-

vae of the two speeies, and of the main biologieal

and eeologieal eharaeteristies of both larvae and

adults. In addition, it was better and more aeeurately

defined the distribution range of A. oculatus in Si-

eily in the light of new loeations surveyed. The lar-

vae deseription showed the absenee at the base of

abdominal segment 9 of small eylinder-shaped or

eone-shaped protuberanees eonfirming that Allar-

dius belong to the subtribe Helopina (see Purehart

& Nabozhenko, 2005).

Notably, we showed a very elear distinet sapro-

xylie aetivities of Allardius and their important

role in the forest eeology. In faet, in reeent years,

the Deadwood is one of the indieators ehosen to

assess the state of forests and the sustainability of

their management (MCPFE, 2003; LEA, 2007);

moreover, its role in the forest eeosystem and bio-

diversity eonservation emerged in an inereasingly

evident manner supported by numerous seientifie

studies (Wermelinger & Duelli, 2002; Mason et

al., 2003; Hahn & Christensen, 2004; Humphrey

et al., 2004; Travaglini et al., 2007; Pignatti et al.,

2009). Henee, the peeuliar biology of Allardius,

espeeially when it is supported by numerous and

stable populations, allows us to eonsider these spe-

eies as the top detritivores of dead wood of the fo-

rests they inhabit, thus stressing the importanee of

Forest-ecological aspects of the genus Allardius (Ragusa, 1898) (ColeopteraTenebrionidae) in Sicily and Sardinia

491

Figure 10. Larvae of Allardius oculatus (Palermo, Addaura): several generations in the wood eompletely degraded. Figure

11. Pupa of A. oculatus (Monte Iniei). Figures 12, 13. A. oculatus (Monte Iniei) during mating. Figures 14-16. Larva of A.

oculatus (Monte Iniei), inpartieular (Figs. 15, 16), abdominal tergite 8-9 in dorsal view (15) and in lateral view (16). Figures

17, 18. Larva of A. sardiniensis (Monte Sette Fratelli), in partieular (Fig. 18), abdominal tergite 8-9 in lateral view.

492

Michele Bellavista & Ignazio Sparacio

their role in the forest eeology in relation to the

presenee of "dead wood".

ACKNOWLEDGEMENTS

Authors are very mueh obliged to F. Fiordilino

(Castellammare del Golfo, Italy) and to M. Romano

(Capaei, Italy).

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Humphrey J.W., Sippola A.L., Lemperiere G., Dodelin

B., Alexander K.N.A. & Butler J.E., 2004. Deadwood

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(Ed.) “Monitoring and indicators of forest biodiver-

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Iwan D., Lillig M., Masumoto K., Nabozhenko M.,

Novak V., Petterson R., Schawaller W. & Soldati F.

2008: Family Tenebrionidae Latreille, 1802. Pp.

105-352. In: Lobl I. & Smetana A. (Eds.), 2008.

Catalogue of Palaearctic Coleoptera. Vol. 5. Tene-

brionoidea. Apollo Books, Stenstrup, 669 pp.

Luigioni P, 1929. 1 Coleotteri dTtalia. Catalogo topogra-

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morie della Pontificia Accademia Seientifica I Nuovi

Lincei, 13: 1-1160.

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Biodiversity Journal, 2012, 3 (4): 493-500

Presence of black rat Rattus rattus (Linnaeus, 1 758) (Mammalia

Rodentia) and possible extinction risk for micro insular popu-

lations of Podarcis sicula (Rafinesque, 1810) (Reptilia Lacerti-

dae): the example of Lachea islet (Sicily, Italy)

Agatino M. Siracusa'*, Veronica Larosa^ & Ettore Petralia^

Department of Biologieal, Geologieal and Environmental Seienees - Seetion of Animal Biology, University of Catania, via Androne

81 - 95124 Catania, Italy; e-mails: amsira@uniet.it, verolarosa@libero.it, e.petralia@libero.it

* Corresponding author

ABSTRACT The black rat Rattus rattus (Linnaeus, 1758) in insular environments represents a threat for

many species of vertebrates, invertebrates and plants, especially in equatorial islands. In the

Mediterranean Basin as regards the herpetofauna, and lizards of the genus Podarcis in parti-

cular, the information available are still few. Since 2006, a study was initiated to verify the

possible impact of the black rat on the micro-insular population of Podarcis sicula (Rafme-

sque, 1810) living in Lachea island, a natural reserve. During 2011 were collected and analy-

zed 2873 excrements of rats and in no case were found remains attributable to Podarcis sicula.

The density of lizards, observed with the technique of the transept, was 0.3 ind./lO m^. During

the period 2006-2011 were analyzed (also by molecular type investigation) a total of 4696

excrements of rat, with no confirmation of predation against Podarcis sicula. However the

rat predation on insects may cause an indirect effect like competition and have negative effects

on populations of invertebrates and therefore also on Podarcis sicula. Moreover the Lachea

island is affected by a moderate tourism. The lizards, in those very frequented areas, show

lower values of the body condition index and a decrease of cells responsible for immune re-

sponse. Although is not documented any form of predation by the black rat, that species is a

potential source of threat and a combined action of the factors mentioned with accidental

events, makes this small micro-insular population of Podarcis sicula as vulnerable to sudden

decreases in numbers.

KEY WORDS Rattus rattus] Podarcis sicula] Lachea island; predation; competition.

Received 12.05.2012; accepted 12.12.2012; printed 30.12.2012

Proceedings of the International Congress “Insularity and Biodiversity”, May 11*-13*, 2012 - Palermo (Italy)

INTRODUCTION

The black rat Rattus rattus (Linnaeus, 1758)

(Mamiualia, Rodentia), a native of the East Asian

region, is now cosmopolitan, widespread both in

continental and insular areas. The first remains at-

tributed to this species in the western Mediterranean

date back about 2400 years ago (Amori et al., 2008;

Masseti, 2008). Currently it is very common in the

Mediterranean islands, and is present in all the me-

dium and large ones and in many of the smaller

ones (Perfetti et al., 2001; Angelici et al., 2009). In

most cases the colonization has anthropochore ori-

gin, although in the islands situated at short distance

from the mainland or other islands, can instead

occur spontaneously (Palmer & Pons, 2001).

494

A.M. Siracusa, V. Larosa & E. Petralia

The black rat has directly caused or contributed

to the extinction of numerous species of birds, small

mammals, reptiles, invertebrates and plants, espe-

cially in equatorial islands (Palmer & Pons, 2001;

Nogales et ah, 2006; St Clair, 2011). It is, among

allochthonous species, the most destructive for Me-

diterranean seabirds, being considered responsible

for the disappearance of colonies of Hydrobates pe-

lagicus (Linnaeus, 1758) and for number decrea-

sing of Calonectris diomedea (Scopoli, 1769)

(Igual et ah, 2006). It can also cause a reduction in

reproductive success of other species of seabirds

such as Puffinus yelkouan (Acerbi, 1827), Puffinus

mauretanicus P. R. Lowe, 1921, or rupicolous birds

as Eleonora’s Falcon Falco eleonorae Gene, 1839

and Pallid ssNifiApus pallidus (Shelley, 1870).

Information are scarce regarding reptiles in ge-

neral and lizards of the genus Podarcis in particu-

lar. In the Balearic Islands are present 43 different

insular populations of Podarcis lilfordi (Gunther,

1874): in the islands without the presence of black

rats this species has statistically significant higher

density (Perez-Mellado et al., 2008). Predation by

black rat also appears to be the cause of the decline

of Podarcis filfolensis kieselbachi (Fejervary,

1924), endemic subspecies of Selmunett island, off

the northeast coast of Malta (Sciberras & Schem-

bri, 2008). Podarcis sicula (Rafinesque, 1810) is

present in Italy in numerous small islands, with po-

pulations showing eco-ethological peculiarities and

having a high degree of vulnerability because of

their isolation (Sindaco et al., 2006). On the Lachea

island there is a population described as Podarcis

sicula ciclopica (Taddei, 1949): it presents distin-

ctive genetic features but not sufficient to justify

the subspecies rank, although further studies are

desirable to clarify the taxonomic status (G. Rap-

pazzo, in verbis).

Since 2006 an investigation was initiated aimed

at studying the possible impact of the black rat on

this micro-insular population of Podarcis sicula,

with the aim of collecting useful information for its

preservation.

MATERIALS AND METHODS

The Lachea island belongs to a small archipe-

lago called “Islands of the Cyclops” at about 200 m

from the coast of Acitrezza (Aci Gastello, Catania,

Sicily), and it has an area of 1 .3 ha. The origin dates

back to the Middle Pleistocene (500,000 to 700,000

years ago) where there was the ancient “pre-Etna

gulf’. The climate is mediterranean type and the ve-

getation has a thermo-xerophilous character; with

around 1 80 plant species its flora is impoverished

due to the insertion of alien species (Ailanthus al-

tissima L., Thuja orientalis L.) and the agricultural

activities practiced until a few decades ago. Since

1998 the island is an Integral Natural Reserve,

whose management is assigned to C.U.T.G. A.N.A.

of Catania University.

From January to December 2011 was carried

out a monitoring of the population of black rat and

Podarcis sicula with the main purpose of verifying

any interference of this Rodent against this species

of lizard. We proceeded to monthly collection of

black rat excrements in four different stations (Fig.

1): station n. 1 (540 m^), situated in the west of the

island is characterized by a predominantly herba-

ceous cover; station n. 2 (555 m^), dominated by

herbaceous and shrub vegetation; station n. 3 (123

m^), concerns a paved area surrounded by a predo-

minantly arboreal vegetation; station n. 4 (75 m^),

consisting predominantly in rocky ground with

sparse and exposed vegetation. This last is frequen-

ted by the Larus michahellis Naumann, 1840,

which nests here. Excrements collected were ana-

lyzed in the laboratory, separating vegetal compo-

nents and animal components (e.g. remains of

insects and birds), giving particular attention to the

possible presence of remains (scales, bone and egg

fragments, skin) of Podarcis sicula. Data were then

reported as percent frequency (F%), equal to the

number of times that a single food category is

found, divided by the total number of excrements

and multiplied by 100 (n/N x 100).

Regarding Podarcis sicula, the survey was co-

vered monthly by a fixed transect (Fig. 2), pre-

viously standardized, at three different time slots

(8:00-9:00, 11:00-12:00 and 15:00-16:00). Counts

were also made by direct sightings, during a period

of 10 min., in 3 different stations, always in the

same time slots (Fig. 3). The first station (42 m^) is

situated in the west side of the island; it is an eleva-

ted area surrounded by small walls built in lava

stone and marly rocks; it presents herbaceous vege-

tation, especially during autumn and winter. The se-

cond station (70 m^) is located in the upper part of

the island (25.35 m a.s.L); it appears flat with trees

Presence of Rattus rattus and possible extinction risk for Podarcis sicula:the example ofLachea islet (Sicily, Italy) 495

that do not exceed 1.5 meters in height, and annual

plants. The third station (120 m^) is located on the

east side of the island, in the area known as Punta

Monaco; this is the zone of maximum solar expo-

sure, and is almost entirely made up of marly rocks;

the vegetation is predominantly herbaceous (Ory-

zopsis miliacea (L.) Asch. et Schweinf, Crithmum

maritimum L., Car Una hispanica globosa Meusel

& Kastner), while at the edge of the station there

are shrubs such as Rubus fruticosus L. and Opuntia

ficus-barbarica A. Berger.

The transept and the stations have been identi-

fied taking into account the ecological diversity,

in order to have an overview on the distribution

and abundance of lizard population of the island.

The transept involves observation and counting of

all individuals contacted along the predetermined

path (at a perpendicular distance of 1 m from it,

both right and left) of length equal to 500 m. The

reliability of the method is linked to the proper

functioning of certain procedures (equiprobability

of detection of all the subjects, certain identifica-

tion of the animals counted, independence of each

observation and precision in measuring the per-

pendicular distance).

The subsequent density estimates were calcula-

ted using the following equation: D = n/2Lw, where

“n” is the number of individuals observed, “L” is

the length of the transept and “w” is the mean of the

perpendicular distance to the line of progression

(Burnham et al., 1980). Throughout the year of

sampling were collected meteorological and cli-

matic parameters, in particular: monthly mean

temperature, monthly minimum and maximum

temperature, and monthly total precipitation, using

data from the weather station of Aci Trezza (7 m

a.s.l.) (www.meteosicilia.it). To assess possible dif-

ferences in the number abundance of black rat ex-

crements, found in the stations, the t-test has been

applied. For the correlation between climate varia-

bles and number of individuals of lizards observed

the nonparametric test of Spearman rank coefficient

(rs) was applied. For comparisons with results from

previous studies were used just monthly samples si-

milar to this research. Data processing was carried

ut by the statistical package Statistics 5.0.

Figure 1. Location of the 4 sampling stations for the black

rat excrements. Figure 2. Transept standard used for the

counting of Podarcis sicula individuals. Figure 3. Location

of the 3 stations used for the sampling of Podarcis sicula.

496

A.M. Siracusa, V. Larosa & E. Petralia

RESULTS

The results of the exerements (n = 2873) analy-

sis are reported in Table 1 . The high number of ex-

erements found in the month of January was

attributed to the non elimination of the previous

ones before the start of sampling.

Excrements contain predominantly plant re-

mains, always present and showing the maximum

frequency; only in a small percentage have been

found birds and insects remains. In no case were

identified remains attributable to Podarcis sicula.

Using collected data was also calculated the

density of the number of excrements found in each

station (Table 2). The average values found in each

station are reported in Table 3.

There were no statistically significant differen-

ces between the abundance of excrement in stations

1 and 4 and stations 2 and 3 (Table 4). The indivi-

duals density of Podarcis sicula observed along the

transect is shown in Table 5 and Tables 6-8 show

the values observed in each station.

Along the transect the highest number of indi-

viduals observed occurs in the month of May. In

winter there is a drastic decrease of individuals re-

corded. The maximum number of individuals ob-

served per month is also significantly positively

correlated with the mean temperature (rs = 0.756,

t(6) = 2.832, P = 0.03) and the maximum tempera-

ture (rs = 0.756, t(6) = 2.832, P = 0.03), while in-

stead it is statistically significantly negatively

correlated with the total precipitation (rs = - 0.903,

t(6) = 5.139, P = 0.00).

The number of individuals observed is also si-

gnificantly positively correlated with the average of

the individuals sum observed at stations (rs = 0.957,

t(6) = 8.103, P = 0.00).

The maximum density observed at the stations

is recorded for the month of May with the first sta-

tion showing 3.6 ind./lO m^ and station 3 showing

Month

n°. excr.

Vegetals

Birds

Insects

Podarcis sicula

January

1604

100.00

2.49

0.00

February

266

100.00

0.00

March

181

100.00

0.00

4.42

0.00

April

107

100.00

0.00

May

100.00

0.00

1.51

0.00

June

100.00

0.00

July

100.00

0.00

2.82

0.00

August

September

309

100.00

0.00

1.94

0.00

October

November

188

December

Nb. Total excre-

ments

2873

Table 1 . Food categories and relative frequency percentage (F%) reported monthly in the black rat excrements (nr = no detection

due to adverse weather conditions; / = not analyzed because excrements rotten). The values for January are shown in italics.

Presence of Rattus rattus and possible extinction risk for Podarcis sicula:the example ofLachea islet (Sicily, Italy) 497

Month

Station 2

Station 3

Station 4

January

1.55

1.14

1.89

5.79

February

0.43

0.20

0.33

0.43

March

0.22

0.14

0.16

0.59

April

0.14

0.12

0.10

May

0.14

0.06

0.05

0.00

June

0.12

0.04

0.08

0.37

July

0.09

0.06

0.10

0.12

August

September

0.39

0.22

0.47

0.75

October

November

0.24

0.12

0.45

0.27

December

Table 2. Monthly

density of blaek rat

exerements found in

eaeh station (ne = no

deteetion due to pre-

senee of nesting

gulls; nr = no detee-

tion due to adverse

weather eonditions).

The values for Ja-

nuary are shown in

italies.

mean

Station 1

10.23

0.13

Station 2

1.18

0.72

Station 3

2.42

1.77

Station 4

10.36

0.26

Table 3. Yearly mean values and standard de-

viation of the number of blaek rat exerements

found at eaeh station.

d.f.

Stat. 1 - Stat. 2

32.726

0.000

Stat. 1 - Stat. 3

11.643

0.000

Stat. 1 - Stat. 4

1.183

0.260

Stat. 2 - Stat. 3

1.717

0.112

Stat. 2 - Stat. 4

31.728

0.000

Stat. 3 - Stat. 1

28.242

0.000

Stat. 3 - Stat. 4

11.743

0.000

Month

n° ind./10 m^

January

0.0 ind./lO m^

February

0.0 ind./lO m^

March

0.1 ind./lO m^

April

May

0.3 ind./lO m^

June

0.2 ind./lO m^

July

0.2 ind./lO m^

August

September

0.2 ind./lO m^

October

November

0.0 ind./lO m^

December

Table 4. Statistieal eomparison (t-test) between

the averages of blaek rat exerements found in

eaeh of 4 sampling stations.

Table 5. Density values oi Podarcis sicuh observed in eaeh month

along the transept (nr = no deteetion due to adverse weather eondi-

tions)

498

A.M. Siracusa, V. Larosa & E. Petralia

1 .0 ind./lO m^. At station 2 the maximum density is

2.1 m^ ind./lO m^ registered in June and July. The

mean value for the three stations together was equal

to 2.2±1.30 ind./lO m^ (n = 3).

DISCUSSION

Based on the results obtained have not been do-

eumented any eases of predation by blaek rat versus

Podarcis sicula. During the whole period of inve-

stigation (2006 - 2011) were analyzed 4696 exere-

ments of rats but never obtaining any evidenee that

would eonfirm the lizard as a trophie souree of this

Rodent in the island (Siraeusa et al., 2010; pres,

stud.; unpublished data). Even moleeular analysis

were negative (Siraeusa et al., 2010).

Rats direet the predation espeeially versus

young lizards; in studies earried out in reeent years

have been observed only sporadieally attempts by

rats to eateh lizards. Moreover, the average density

of blaek rats observed was equal to 39.1 ind./ha, not

partieularly great eompared to other similar-size

Mediterranean islands; all individuals eaptured also

showed very high values of body eondition index

(Petralia et al., 2010).

These eonsiderations and the faet that there

seems to be no eorrelation between the frequeney

of Podarcis sicula (number of individuals eounted

with transeet method) and blaek rat (Table 9),

would indieate that predation is insignifieant.

Should nevertheless be taken into aeeount indireet

effeets that may be partieularly important in a lon-

ger time. Predation of the rat versus inseets and its

eonsumption of plants ean determine an indireet ef-

feet of eompetitive type against Podarcis sicula

espeeially with regard to invertebrates (Perez-Mel-

lado et al., 2008).

Month

n° ind./10 m^

January

0.0 ind./lO m^

February

0.2 ind./lO m^

March

0.2 ind./lO m^

April

May

3.6 ind./lO m^

June

2.4 ind./lO m^

July

2.6 ind./lO m^

August

September

2.1 ind./lO m^

October

November

December

Month

n° ind./10 m^

January

0.0 ind./lO m^

February

0.0 ind./lO m^

March

0.1 ind./lO m^

April

May

2.0 ind./lO m^

June

2.1 ind./lO m^

July

2.1 ind./lO m^

August

September

1.9 ind./lO m^

October

November

December

Month

n° ind./10 m^

January

0.0 ind./lO m^

February

0.5 ind./lO m^

March

0.7 ind./lO m^

April

May

1.0 ind./lO m^

June

0.9 ind./lO m^

July

0.8 ind./lO m^

August

September

0.8 ind./lO m^

October

November

0.2 ind./lO m^

December

Table 6. Number of individuals /lO

of Podarcis sicula observed at

station 1 (nr = no deteetion due to ad-

verse weather eonditions).

Table 7. Number of individuals /lO

m^ of Podarcis sicula observed at

station 2 (nr = no deteetion due to ad-

verse weather eonditions).

Table 8. Number of individuals /lO

m^ of Podarcis sicula observed at

station 3 (nr = no deteetion due to ad-

verse weather eonditions).

Presence of Rattus rattus and possible extinction risk for Podarcis sicula:the example ofLachea islet (Sicily, Italy) 499

2006-20071

2007-2008^

2008-2009^

2011^

Rattus rattus (excr./lO m^)

4.30

8.70

11.40

Podarcis sicula (trans.)

(ind./lO mh

0.2

0.3

Podarcis sicula (stat.)

(ind./10 mh

2.2

Table 9. Abundance

values of Podarcis si-

cula and Rattus rattus

registered between

2006 and 2011 (nr = no

detection) (1 = Fiorini,

2006; 2 = Siracusa et

al., 2010; 3 = Zappala,

2010; 4 = pres. stud.).

Rats also can have negative impacts on popu-

lations of invertebrates and thus indireetly on li-

zards (Pyke et al., 1977). Densities of this rodent

vary aeeording to availability of food of vegetal

origin and size of islands eolonized (VV. AA. in

Martin et al., 2000; Guyot, 1989 in Martin et al.,

2000). The flora diversity and produetivity are also

responsible for both the incoming and the numeri-

eal variations of the speeies, even in small islands

(Palmer & Pons, 2001).

Moreover the population of nesting Larus mi-

chahellis (present with a small eolony on the is-

land), favoring nitrophilous plant speeies and

making available additional food sourees (eareas-

ses, remains of eggs, food remains), allows the

number inerease of rats. Over the past 30 years,

the Larus michahellis population has exploded in

the Mediterranean and serious effeets on eeosy-

stems, espeeially in relation to rare plant and ani-

mal speeies have already been doeumented (Vidal

et al., 1998).

CONCLUSIONS

The Laehea island is affeeted by the tourism,

even if eontrolled. Tourism is eonsidered a potential

souree of threat to different populations of lizards

(Amo et al., 2006), espeeially in small islands. In

areas frequented by human lizards show lower va-

lues of the body eondition index and unfavourable

effeets in the host-parasite interaetion, due to a de-

erease of eells responsible for immune response

(Amo et al., 2006).

Antipredator strategies are expensive in terms

of fitness (Martin & Lopez, 1999) and beeause of

that lizards may suffer an inerease in the risk of pre-

dation (L6pez et al., 2005). Moreover in order to

proteet the reserve from fire risk, trimming is also

exeeuted during the spring.

These interventions eliminate or modify mi-

erohabitats eolonized by different speeies of ar-

thropods, potential prey of Podarcis sicula. For

the eonservation of the herpetofauna, espeeially

in regard to small and isolated populations, is then

reeommended a control of the blaek rat (Salvador

& Veiga, 2005; 2008): after eradieation or control

of this species in 10 Tyrrhenian islands was ob-

served in some eases a sudden inerease in the den-

sity of lizards (Capizzi et al., 2006). Combined

aetions of the faetors listed before, together with

possible stoehastie events, make this small popu-

lation of Podarcis sicula vulnerable to sudden de-

ereases in numbers.

Rattus rattus beeause of its ethologieal and eeo-

logieal features is a potential souree of threat (pre-

datory activity on invertebrates, potentiality to

expand the population, indireet eompetition) even

though has not been doeumented any form of pre-

dation on that Podarcis speeies.

ACKNOWLEDGMENTS

We wish here to thank: Prof. Pietro Pavone, Di-

reetor of the Department of Biologieal, Geologieal,

and Enviromnental Seienees of the Catania Univer-

sity, for the availability of loeals and faeilities pro-

vided at the laboratories of the Department; Prof.

Angelo Messina (President of CUTG AN A Founda-

tion), Prof. M. Carmela Failla (Direetor of CUT-

GANA) and Dr. Emanuele Molliea (Direetor of

RNI “Isola Laehea e Faraglioni dei Cielopi”) to

have authorized the eonduetion of the study within

the Reserve, Mauro Contarino, operator of the re-

serve and eollaborator of Dr. Molliea, for his assi-

500

A.M. Siracusa, V. Larosa & E. Petralia

stance in the field; Prof. Alfredo Petralia and Prof.

Giorgio Sabella for their seientifie support and the

eonstant and eontinuous supervision in all the dif-

ferent phases of this work.

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Biodiversity Journal, 2012, 3 (4): 501-510

Biodiversity loss in Sicilian transitional waters: the molluscs

of Faro Lake

Salvatore Giacobbe

Department of Biologieal and Environmental Seienees, Viale Ferdinando Stagno d'Aleontres, 3 1 - 98166 S.Agata-Messina, Italy;

e-mail: sgiaeobbe@unime.it

ABSTRACT Sediment samplings were carried out in six stations of Faro Lake (Sicily, Italy) during spring

and autumn 1991, and spring 1993, 2006, 2010, to investigate the soft bottom mollusc assem-

blages. The study have provided the first quantitative data on the mollusc fauna of Faro Lake,

to date known only for some dated inventories. Some differences in species composition in re-

spect to the ancient literature were highlighted. The 1991-2010 data sets showed changes in

the mollusc assemblages, which impoverished in composition and structure. Basic indices of

community structure (S, d, H’, J’) indicate a marked decline in the 2006, followed by a recovery

period during 2010. The multivariate analysis, based on a the Bray-Curtis similarity index,

proved that mollusc assemblages sampled prior to the 2006 crisis were more structured and

spatially differentiated than those settled later. The observed decline of endemic taxa and the

concurrent settlement of species introduced by mussel farming, make the mollusc assemblages

in the Faro Lake a case-study for the effect of species introduction in confined environments.

KEY WORDS Coastal ponds; Macrobenthic fauna; changing communities; endemisms; Alien species.

Received 12.05.2012; accepted 27.11.2012; printed 30.12.2012

Proceedings of the L' International Congress “Insularity and Biodiversity”, May 11*-13*, 2012 - Palermo (Italy)

INTRODUCTION

Transitional waters (TWs), since ecotones bet-

ween terrestrial, freshwater and marine ecosystems,

are characterized by highly dynamic processes,

strong physical and chemical dines, and irregular

temporal trends (De Wit, 2007; Evagelopoulos et

al., 2009). These environments, that provide key

ecosystem services, are characterized by high pro-

ductivity levels, whose exploitation attracted human

populations since the Palaeolithic Age (Knottnerus,

2005). Due to the long history of high anthropoge-

nic pressure, European estuaries and coastal areas

are among the most severely degraded systems wor-

ldwide (Lotze et al., 2006). The habitat loss, that

between 1960 and 1995 reached a 50% rate (Airoldi

& Beck, 2007), was also responsible of a large-scale

habitat fragmentation that reflected on the local/re-

gional biodiversity levels. The rarefaction of key-

stone or engineer species, the extinction of endemic

and rare taxa, the burst of pollution indicators, the

spreading of introduced alien species, typically te-

stify a biodiversity loss due to habitat depletion

(Wolff, 1998; Levin et al., 2001).

In the recent years, increased conservation poli-

cies required more and more detailed information

on the environmental quality of coastal and estuarine

environments, with a particular emphasis towards

the biological indicators (Borja et al., 2000). Never-

theless, the necessary background of knowledge on

both the present and past biodiversity levels is often

lacking, and reference conditions along the Euro-

pean coasts are not available (Ponti et al., 2008).

Such evident information gap is particularly marked

502

Salvatore Giacobbe

for some Southern Mediterranean regions, whose

TWs are partially or totally disregarded in the natio-

nal reviews (Basset et ah, 2006) as well as in the

large-seale assessments (Munari & Mistri, 2010).

The Sieilian TWs inelude wetlands, semi-enelo-

sed bays, braekish basins, eoastal ponds, xero-Me-

diterranean lagoons, and marine lakes, submitted to

different types and grade of anthropogenie pressure

(Mazzola et ah, 2010), whose impaet on the loeal

biodiversity is not still quantified. Furthermore, the

present knowledge on the respeetive benthie assem-

blages are inappropriate for a spatio-temporal eom-

parison, due to the different investigation methods

and efforts, whilst historieal data series (Leonardi

& Giaeobbe, 2001) are rarely available.

The Faro Lake, belonging to the “Capo Peloro

e Laghi di Ganzirri” CSI and SPZ area (North-Ea-

stern Sieily), despite of proteeted status, represents

an area in whieh data on benthie assemblages are

laeking. In faet, although in the past times some in-

ventories of speeies were published for algae (Ca-

valiere, 1963), sponges (Labate & Arena, 1964),

eehinoderms (Cavaliere, 1971), and molluses

Figure 1. Upper pane: The Capo Peloro area, between the

Tyrrhenian Sea and the Strait of Messina. The Faro lake is

indieated. Lower pane: Sampling station loeation.

(Spada, 1969; Parenzan, 1979), data about benthie

eommunities are still poorly doeumented. The mol-

luses, whieh inelude at least one exelusive endemie

speeies and some peeuliar eeo-morphotypes

(Priolo, 1965; Parenzan, 1979), were also emplo-

yed as models of environmental eomplexity of Ita-

lian braekish and marine habitats (lannotta et ah,

2009). More reeently, the anthropogenie introdue-

tion of molluses in the Faro Lake was highlighted

by Cosentino et al. (2009), but the eonsequent im-

paet on the indigenous benthie assemblages was

not still investigated.

Aim of the present paper is to provide a first eon-

tribution to the knowledge of the aetual soft-bottom

molluse eommunity in the Faro Lake; to deseribe

their variations on a deeadal seale, based on unpu-

blished data sets; to individuate a possible biodiver-

sity loss due to anthropogenie impaets.

MATERIALS AND METHODS

Study area

The Faro Lake (Fig. 1), although eharaeterized

by redueed dimension (0.263 Km^), represents the

deepest eoastal basin in Italy, reaehing a maximum

depth of 29 m in the funnel-shaped eastern part. By

eontrast, the shallower western side does not ex-

eeed 3 m depth. It reeeives marine waters from the

nearby Strait of Messina, through a narrow and

shallow ehannel, whieh extends for almost 1 km in

length. During the summer period, a transient eon-

neetion with the Tyrrhenian Sea is artifieially rea-

lized by a further ehannel that normally is silted up.

The lake is eharaeterized by a meromietie regime,

with anoxie and sulphidie waters that generally are

eonfined below 15 m depth. The surfaee waters are

mesotrophie, with a predominant heterotrophie

biomass in the partieulate matter (Leonardi et al.,

2009) and regular blooms of photosynthetie baete-

ria just below the oxyeline (Saeea et al., 2008). The

thermo-haline stratifieation is not stable, and the

poorly oxygenated waters ean spread towards the

surfaee in autumn (Sorokin & Donato, 1975). Ex-

eeptional seawater inflows origine strong altera-

tions of the anoxie layer and temporary diseases of

all aerobie organisms (Giuffre & Pezzani, 2005).

The lake was historieally submitted to various

human aetivities, that repeatedly modified the basin

Biodiversity loss in Sicilian transitional waters: the molluscs of Faro Lake

503

morphology along the last ten eenturies (Manga-

naro et ah, 201 1). At the present time, the shallower

beds are affeeted by traditional praetiees of elam

eulture, whilst the deeper zone is affeeted by orga-

nie and earbonate inputs by mussel farming. The

anthropogenie pressure, due to the extensive lan-

dseape urbanization, is also responsible of a mode-

rate ehemieal and biologieal eontamination (Lieata

et al., 2004; Sorgi et al., 2006).

Data sets

Soft bottom samples were eolleeted in the Faro

Lake during spring and autumn 1991, and spring

1993, 2006, 2010, in the framework of loeal “PRA”

and national “PRIN” programs. Sinee the respeetive

sampling plans partially overlapped, six reeurrent

stations were seleeted in the shallower area (1-2.5

m depth) to obtain temporal replieates; the station

3 was exeluded sinee affeeted by elam-eulture aeti-

vities (Fig. 1). Samples were earried out by means

of a 5 dm^ Van Veen grab (three random replieates

inside 10x10 m stations), washed on field by means

of 1 mm mesh sieves, and stored in 70% ethanol.

Molluse fauna was determined at the speeies level,

and the abundanees were submitted to univariate

(not transfonned data) and multivariate (square root

transformed data) statistieal analysis, by means of

PRIMER 6.1 paekage.

In addition, the present distribution of both de-

teeted endemie and introdueed taxa, was eheeked

by direet SCUBA observations during spring-sum-

mer 2010, from 0.5 tolO m depth.

RESULTS

The seleeted data set, reviewed in aeeordanee

with Worms (http://marinespeeies.org/), aeeoun-

ted a total of 46 molluse speeies, most of whieh

were gastropods (30 speeies). Bivalves aeeounted

1 5 speeies, whilst polyplaeophorans were represen-

ted by only one speeies. Fourteen gastropods and

eight bivalves, previously eited by Parenzan (1979),

were not reeorded, whilst nine gastropods, eight bi-

valves and one polyplaeophoran were added (Table

1). Sueh diserepaney did not prove a priori that sub-

stantial ehanges oeeurred in the molluse assemblage

eomposition, sinee the two inventories refleeted dif-

ferent sampling method and effort. Furthermore,

speeies as Haminoea orteai Talavera, Murillo et

Templado, 1987, and Mancikellia parrussetensis

(Giribet et Penas, 1999), were deseribed afterwards

the Parenzan note. Differently, reeent settlement of

the non-native Cerithium scabridum Philippi, 1848,

Crassostrea gigas (Thunberg, 1793), and Anadara

transversa (Say, 1 822), might be in aeeordanee to

the introduetion and spreading of these speeies in

the Mediterranean Sea (e.g. Streftaris & Zenetos,

2006; Croeetta et al., 2009). The present data set

also eonfirmed the oeeurrenee of the exelusive en-

demie Nassarius tinei (Maravigna in Guerin, 1 840),

Jujubinus striatus delpreteanus Sulliotti, 1889, and

Gibbula adansoni sulliottiyiontQvosato, 1888, both

reported in the most reeent eheek-list of Italian pro-

sobranehs (Oliverio, 2008). The peeuliar eeo-mor-

photype Polititapes aureus var. laeta (Poli, 1791),

reported in old literature data (Aradas & Benoit,

1870), was frequently reeorded.

The 46 speeies identified in the 1991-2010 data

set were differently distributed in time and spaee,

and only a minor number of them was found in all

the examined samples. The total number of speeies

per sample, S (Fig. 2), that ranged from one in 2006

to fifteen in 2010, showed a elear deereasing trend

from 1991 to 2006, followed by a remarkable in-

erease in 2010. In this latter year, the highest mini-

mum, maximum and average S values were

reeorded. In general, the spread between minimum

and maximum values was weak for eaeh year, ex-

eept for the 1991, that aeeounted for the seeond hi-

ghest and the seeond lowest S in the whole data set.

Sueh trend was in aeeordanee with Margalef ri-

ehness, d (Fig. 2). Similarly, the Shannon diversity,

H’(Fig. 2), showed a marked deerease from 1991

(highest maximum value) to 2006, and a following

inerease. In 1991 the widest spread between the ma-

ximum and the minimum value was observed,

while the narrowest range was found in 2010. The

Pielou’s index, J’(Fig- 2), further put in evidenee

the 2006 peeuliarity, whose very poor assemblages

ranged from a totally uneven eondition to the hi-

ghest evenness.

In general, the different levels of organization

found inside eaeh annual group of samples were

overeome by the inter-annual ehanges that affeeted

the soft bottom assemblages. Sueh preponderanee

of the inter-annual variability with respeet to the

spatial patehiness was also eonfirmed by the Bray-

Curtis similarity index and related eluster analysis

504

Salvatore Giacobbe

CHECK-LIST

1979

1991-

2010

POLIPLACOPHORA

Acanthochitona crinita (Pennant, 1777)

GASTROPODA

Alvania cimex (Linnaeus, 1758)

Alvania discors (Allan, 1818)

Alvania geryonia (Nardo, 1847)

Alvania lanciae (Caleara, 1845)

Bittium latreillii (Payraudeau, 1 826)

Bittium reticulatum (da Costa, 1778)

Caecum auriculatum de Folin, 1868

Calliostoma sp.

Cerithium renovatum Monterosato, 1884

Cerithium scabridum Philippi, 1 848

Cerithium vulgatum Bruguiere, 1792

Clanculus jussieui (Payraudeau, 1826)

Conus ventricosus Gmelin, 1791

Cy elope neritea (Linnaeus, 1758)

Dikoleps nitens (Philippi, 1844)

Eatonina ochroleuca (Brusina, 1 869)

Ecrobia ventrosa (Montagu, 1803)

Euspira intricata (Donovan, 1804)

Euthria cornea (Linnaeus, 1758)

Fissurella nubecula (Linnaeus, 1758)

Gibberula miliaria (Linnaeus, 1758)

Gibbula adansoni sulliotti Monter., 1888

Gibbula turbinoides (Deshayes, 1835)

Haminoea hydatis (Linnaeus, 1758)

Nassarius corniculum (Olivi, 1792)

Nassarius mutabilis (Linnaeus, 1758)

Nassarius tinei (Maravigna, 1840)

Odostomia scalaris MaeGillivray, 1843

Peringia ulvae (Pennant, 1777)

Phorcus articulatus (Lamarek, 1822)

Phorcus mutabilis (Philippi, 1846)

CHECK-LIST

1979

1991-

2010

Pisania striata (Gmelin, 1791)

Pisinna glabrata (Von Miihlfeldt, 1824)

Potamides conicus (Blainville, 1 829)

Pusillina lineolata (Miehaud, 1832)

Skeneopsis planorbis (Fabrieius 0., 1780)

Thylacodes arenarius (Linnaeus, 1758)

Tricolia pullus (Linnaeus, 1758)

BIVALVIA

Abra alba (W. Wood, 1802)

Abra prismatica (Montagu, 1808)

Abra segmentum (Reeluz, 1 843)

Abra tenuis (Montagu, 1 803)

Anadara transversa (Say, 1 822)

Cerastoderma glaucum (Bmguiere, 1789)

Chamelea gallina (Linnaeus, 1758)

Corbula gibba (Olivi, 1792)

Crassostrea gigas (Thunberg, 1793)

Gastrana fragilis (Linnaeus, 1758)

Gibbomodiola adriatica (Lamarek, 1819)

Gouldia minima (Montagu, 1803)

Loripes lucinalis (Lamarek, 1818)

Mancikellia parrussetensis (G&Pen.,1999)

Modiolus barbatus (Linnaeus, 1758)

Mytilaster minimus (Poll, 1795)

Mytilus galloprovincialis Lamarek, 1819

Ostrea stentina Payraudeau, 1 826

Papillicardium papillosum (Poll, 1791)

Parvicardium exiguum (Gmelin, 1791)

Polititapes aureus var. laeta (Poll, 1791)

Teredo sp.

Venerupis decussata (Linnaeus, 1758)

Table 1. Comparison from the Parenzan (1979) eheek-list

and the present data.

Biodiversity loss in Sicilian transitional waters: the molluscs of Faro Lake

505

Total $

1393 2006 2010

SpecEei richness (Marcalef); d

1991 1995 2006 2010

■ min

■ M

■ Max

Pi*lou*s*venn*u:J*

1.2 1

1991 1995 2006 2010

Figure 2. Minimum (min), maximum (Max) and average (M) values found in the four sampling years, for total number of

speeies for sample, S; Margalef riehness, d; Shannon diversity, H’; and Pielou’s evenness, J’.

Figures 3, 4. 2D MDS ordination for the benthie assemblages (Fig. 3), and bubble plot for the speeies Nassarius tinei

(Fig. 4).

and 2D MDS ordination, which showed the samples

grouped mainly in aeeordanee with the sampling

year (Fig. 3). Similarity grouped the whole 1991,

1993 and some 2006 samples, at 50%, whilst the

2010 grouped apart. At 30% similarity level, the

1993 almost entirely elustered, differently from the

1991 seasonal data set, that showed two main sub-

elusters and two single samples, not eorrelated with

the sampling seasons.

The hypothesis of a temporal shift that affeeted

a lowly diversified benthie assemblage was verified

by means of one-way and two-way ANOSIM tests

(faetors: year, depth, and station), whieh eonfirmed

the diseriminating role played by the faetor “year”

alone (Global R: 0.638; p: 0.1%; number of permu-

ted statisties greater than or equal to Global R: 0).

The Simper test applied to the groups “year” showed

intra-group similarities ranging from a minimum of

2 1 % , in 2006, to a maximum of 6 1 % , in 20 1 0. The

1991 group (Average similarity: 50.74%) was

mainly eharacterized by the endemie gastropods N.

tinei (33.98% eontribution) and G. adansoni sul-

liotti (21.5%), with further 21.06% was due to the

loeal eeo-phenotype P. aurea var. laeta. The

93.46% similarity was reaehed with the eontribu-

tion of five speeies in total. In the 1993 group (Ave-

rage similarity: 50.92%), the 73.29% of eumulative

intra-group similarity was due to the bivalves P.

aurea var. laeta, L. lucinalis and Venerupis decus-

sata (Linnaeus, 1758), whilst a minor eontribution

(8.59%) was done by G. a. sulliotti. Three speeies

of bivalves: Loripes lucinalis (Lamarek, 1818),

Corbula gibba (Olivi, 1792) and Polititapes aureus

var. laeta (Poll, 1781), aecounted 100% of the 2006

internal similarity, whilst in 2010 three gastropod

speeies: Gibbula adansoni sulliotti Monterosato,

506

Salvatore Giacobbe

1888, Haminoea navicula (da Costa, 1778) and He-

xaplex trunculus (Linnaeus, 1758), were responsi-

ble of the 69% intra-group similarity, whieh reaehed

91% with the eontribution of further four species.

The Simper test also indicated a high inter-group

dissimilarity, reaching a maximum of 95% in the

2006 vs. 2010 comparison. Such extreme difference

between the 2006 and 2010 assemblages was essen-

tially due to 13 species (90% total dissimilarity),

with a major contribution of G. adansoni sulliotti

(18. 5 3%). This endemic taxa was the second major

responsible of intergroup dissimilarity (19.47%) in

the 1991 vs. 2006 (total average dissimilarity:

89.35), whilst N. tinei (20.35%) and P. aureus var.

laeta (18.57%), provided the first and third contri-

butions, respectively. Similarly, in 1993 vs. 2006

(total average dissimilarity: 81.54%), G. a. sulliotti

contributed for 15.15%, following to P. a. var. laeta

(22.53%) and preceding V. decussata (13.11%).

Such marked dissimilarity of both 1991 and 1993

groups in respect to 2006, weakly declined in re-

spect to 2010 (69.93% and 74.24%, respectively).

The greater contribution to the 1991 vs. 2010 dissi-

milarity was done by P. a. var. laeta (15.54%), N.

tinei (14.48%) and//, navicula (10.79%), whilst/!

a. var. laeta (15.75%), G. a. sulliotti (11.56%) and

H. navicula (11.51%) were the most responsible of

the 1993 vs. 2010 dissimilarity. The lowest dissimi-

larity, found between the 1991 and 1993 assembla-

ges (62.90%), was especially due to the endemic

taxa P. a. var. laeta (17.42%), G. a. sulliotti

(16.43%) and A^. tinei (16%).The differentiate time-

distribution of the endemic N. tinei in comparison

with the sampling clustering is shown in Figure 4.

Both intra-group similarity and inter-group dis-

similarity testified of remarkable changes that oc-

curred in the soft-bottom mollusc assemblages

since 1991, as also testified by the main univariate

indexes of community structure. Changes funda-

mentally affected the whole soft-bottom assem-

blage, whose spatial variability in 1991, 1993 and

2010 was low in respect to the temporal changes.

The mollusc assemblages found in 2006, characte-

rized by poor faunal composition and structure fur-

ther than by high spatial heterogeneity, were

notably different from those recorded in the prece-

dent and successive times.

In general, the investigated assemblages accoun-

ted a low number of species, as characteristic of

estuarine soft-bottom communities, three to six of

which were responsible of the 90% intra-group si-

milarity. Such characterizing species included some

endemic taxa, whose contribution to the assemblage

composition notably declined in time, since 1991

(three endemic taxa responsible of 76% internal si-

milarity), up to 1993 (two species, 45% similarity)

and 2006 (one species, 25% similarity). The benthic

recovery following the 2006 decline was marked by

the prevalent role of the endemic G. a. sulliotti

(30.90%) together with a very low contribution by

P. aureus var. laeta (2.70%). Notably, the sole en-

demism that was recognized at the species level, N.

tinei, had a primary role in 1991 alone, whilst did

not significantly contribute to the intra-group simi-

larity afterwards. Furthenuore, the sole endemism

that was recurrent in the whole time-series, P. au-

reus var. laeta, did not represent a true taxonomic

entity, but only a local eco-morphotype that is su-

spected to occur in other Mediterranean environ-

ments. A further endemic subspecies, J. striatus

delpreteanus, occurred only once, in 2010. The en-

demisms which loosed their prominent role in the

intra-group similarity were recognized as minor

components in the inter-group dissimilarity, toge-

ther with the non-native species, Cerithium scabri-

dum, Crassostrea gigas and Anadara transversa.

Such low occurrence of non-native species that are

known for their invasiveness (Streftaris & Zenetos,

2006) apparently contrasted with their regular in-

troduction in the Faro Lake by means of mollusc

trade. The extensive survey carried out after the

2010 sampling showed artificial bed of the non-na-

tive oyster, Crassostrea gigas, (Figs. 5, 6) together

with the Manila clam, Venerupis philippinarum

(A. Adams et Reeve, 1850) ( Fig. 7). The progres-

sive naturalization of C. gigas since 2010 was pro-

ved by records of juveniles and small sized adults

in the sampling stations and surrounding lake-floor.

V. philippinarum, despite a more than decadal far-

ming practice, did not naturally recruit in the lake,

differently from A. transversa, which was acciden-

tally introduced with the Manila clam commercial

stocks. For a contrast, the C. scabridum settlement

might be due to secondary natural dispersion from

South-Eastern Sicily (Barash & Danin, 1977).

The survey which allowed to individuate sour-

ces of anthropogenic bio-pollution (Olenin et al.,

2007) also proved that declining endemic taxa were

locally concentrated in some “refuge sites”, that not

necessarily represented undisturbed areas. In parti-

cular, J. s. delpreteanus colonized residual sea-grass

Biodiversity loss in Sicilian transitional waters: the molluscs of Faro Lake

507

meadows of Cymodocea nodosa (Ucria) Ascherson,

1870; whilst N. tinei and G. a. sulliotti were fre-

quently reeorded as opportunistie neerophagous in

the eommercial elam-beds (Figs. 8, 9), due to the

high prey eoneentration and mortality rate. By eon-

trast, other endemisms and pseudo-endemisms

known from old literature data, as the eeo-morpho-

types Conus ventricosus var. ater Philippi, 1836,

whose decline was documented since Parenzan

(1979), and Potamides conicus var. peloritana

Figure 5. Aerial view of the Faro Lake (by Google earth); the arrow indieates a eommereial oyster bed. Figure 6. The same

oyster bed photographed in situ. Figure 7. Underwater view of a eommereial Manila elam bed and (Figure 8) view of Nas-

sarius tinei in the same elam bed. Figure 9. Underwater view of Gibbula adansoni sulliotti feeding upon a dead Callista

chione (Linnaeus, 1758) eommereial elam. Figure 10. Samples of dead Gari depressa from the Faro Lake beds.

508

Salvatore Giacobbe

(Cantraine, 1835), might be totally extinct. Simi-

larly, some autochthonous species as Gari depressa

(Pennant, Mil), which were commercially exploi-

ted in past times (Philippi, 1836), actually can be

recorded as death assemblage only (Fig. 10).

DISCUSSIONS AND CONCLUSIONS

The comparison of four data-sets collected in the

Faro Lake during the latter two decades showed a

notable transformation of the soft-bottom mollusc

assemblages. Such transformation affected both

composition and structure of the mollusc taxocoe-

nosis, involving biodiversity changes. Basic indices

of community structure (S, d, H’, J’) showed a de-

cline that was particularly marked in the 2006, be-

fore a recover in 2010. On a wider temporal scale,

some evidences exists that native mollusc popula-

tions, known as local eco-morphotypes, were extinct

or strongly reduced in the last century. Lacking the

proves that human activities caused such an abrupt

and generalized decline, the 2006 benthic depletion

might be explained as a transient effect of diseases

due to anoxic layer alterations (Giuffre & Pezzani,

2005), also facilitated by intensive mussel farming

and related organic sedimentation increase.

The benthic re-colonization, that involved side

supply processes from the nearby coastal areas, pro-

vided stenohaline and moderately euryhaline spe-

cies of marine origin, whilst eur 3 walent taxa spread

from the connected Ganzirri Lake. Nevertheless,

such latter reservoir of local biodiversity is lacking

of the major endemic taxa, as N. tinei and J. striatus

delpreteanus, whose surviving is tied to limited “re-

fuge sites” inside the same Faro basin. On the re-

gional scale, habitat fragmentation prevents genic

fluxes from other Sicilian TWs, but allochthonous

taxa are daily introduced by mussel trade from re-

mote source. Since local sources of biodiversity and

non-native taxa introduction support re-coloniza-

tion by means of unpredictable recruitments, the

2010 biodiversity recovery produced an increased

number of species, but a less structured living as-

semblage with respect to the 1991 mollusc taxocoe-

nosis. The introduced species did not show their

potential invasiveness so far, representing a smaller

part of the 2010 assemblage, and a direct link bet-

ween their settlement in the Faro Lake and loss of

the endemic taxa was not proved. Nevertheless, the

mollusc farming, which is widely responsible of in-

troduction in Western Mediterranean (Galil, 2008),

severely impacts both on the habitat structure and

dynamics at the water-sediment interface, thus fa-

vouring the opportunistic taxa. Other anthropogenic

impacts, as chemical/organic pollution, that also af-

fects the Faro Lake ecosystem, are less recognizable

in their effects. The anthropogenic -induced decline

of the well adapted endemisms, altering the com-

munity structure, negatively affects the multiple

species relationships, thus reducing the effective-

ness of the “priority effect” to contrast biological

invasions (Case, 1990).

Concomitant bio-pollution and native species

decline are linked aspects of a “biotic homogeniza-

tion” that notably affects worldwide biogeography

(Olden, 2006), and involves under-evaluated effects

on the present ecosystem functioning. Although the

measures of mollusc diversity in the Faro Lake did

not show dramatic changes in the last decades,

strong signals of biodiversity reduction were detec-

ted. The high number of menaced endemic taxa and

ecological adaptations, make the mollusc assembla-

ges in the Faro Lake a case-study for the effect of

species introduction in confined environments.

ACKNOWLEDGEMENTS

This research has been supported by the Mes-

sina University, and sponsored by the “Provincia-

Regionale di Messina” the management official

body of the reserve. Facilities have been provided

by the mussel farms FARAU s.r.l. and SACOM

s.r.l. A relevant contribution was also provided con-

cerning field and in laboratory by the researchers

A. Cosentino and M.C. Mangano, and by the stu-

dents A. Rindone, S. Giccone, S. De Matteo, A. Spi-

nelli, M. Cintorino and M. Cavallaro.

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Biodiversity Journal, 2012, 3 (4): 511-520

Rediscovery and re-evaluation of a ‘‘ghost” taxon: the case of

Rissoa galvagni Aradas et Maggiore, 1 844 (Caenogastropoda

Rissoidae)

Danilo Scuderi' & Bruno Amati^

'Via Mauro de Mauro, 15b - 95032 Belpasso, Catania, Italy; e-mail: danscu@tin.it

^Largo Giuseppe Veratti, 37/D - 00146 Roma, Italy; e-mail: bmno.amatil955@libero.it

ABSTRACT The taxonomy of species of the family Rissoidae has always been source of debate and only

a few of rissoid genera have been recently comprehensively revised. The need of revisional

work is particularly obvious in the case of the genus Crisilla Monterosato, 1917, taxon sho-

wing open nomenclatural issues along with uncertainty and difficulty in delimitation of its

species. In this study we revise the status of Crisilla pseudocingulata (Nordsiek, 1972) in

the light of the recent rediscovery of type material of Rissoa galvagni (Aradas et Maggiore,

1844), to which the former is here compared. Based on observations on dimensions, colour

and sculpture of the teleoconch and on the distinctive protoconch characters, C. pseudocin-

gulata is here regarded as junior synonym of C. galvagni. Additional data on the morpho-

logy and colouration of the head-foot as well as on and the variability of shell features are

provided, contributing to an updated description of the taxon.

KEY WORDS a-taxonomy; Aradas collection; Crisilla; marine microgastropods; Mediterranean Sea.

Received 12.05.2012; accepted 13.10.2012; printed 30.12.2012

Proceedings of the P' International Congress “Insularity and Biodiversity”, May 11*-13*, 2012 - Palermo (Italy)

INTRODUCTION

The caenogastropod family Rissoidae Gray,

1 847 (Rissooidea) is one of the most diversified and

widespread marine mierogastropod families, inelu-

ding thirty-one available genus-group taxa, found

from polar waters to the tropies and from the inter-

tidal to the deep sea (Ponder, 1985). The generie-

level elassifieation of the family remained long

based almost entirely on shell eharaeters (Thiele,

1929-31; Wenz, 1938-44; Coan, 1964) until head-

foot, radular and anatomieal information was inelu-

ded (Ponder, 1968; 1985).

Moleeular data, when employed at the generie

level, have even suggested the possibility that Ris-

soidae are not a monophyletie group, but rather an

assemblage of at least two family-level elades (Cri-

seione & Ponder, 2013). Most of their a-taxonomy.

however, still relies on shell morphology, as shown

by the generie revisions available (e.g. Verduin,

1976; 1982a; 1983; 1985; 1986 for Fremin-

ville inDesmarest, 1814; Verduin, 1984; 1988; van

Aartsen & Verduin, 1982; van der Linden & Wa-

gner, 1985 for Setia H. Adams et A. Adams, 1852).

The few attempts to question the presently ae-

eepted elassifieation by eombining anatomieal and

moleeular data have revealed the eonsiderable li-

mits of a purely eonehologieal approaeh (Criseione

et al., 2009; Criseione & Patti, 2010). One major

issue is the high degree of eonvergenee shown by

rissoids in several shell eharaeters (sueh as proto-

eoneh morphology and seulpture, presenee and

number of axial ribs, ete.) often not refleeting the

aetual phylogenetie relationships. When studying

taxonomy of Rissoidae, present-day malaeologists

also need to faee some more “historieal” problems.

512

Danilo Scuderi & Bruno Amati

Most Mediterranean rissoid species, for example,

were described several decades ago (often by ama-

teurs) and the interpretation of their original de-

scriptions is quite often a challenging task. In some

cases the alleged loss of type material and the erro-

neous interpretation of the original description have

led to a considerable proliferation of nomina dubia

in current check-lists and revisions of Mediterra-

nean and Macaronesian rissoids (Verduin, 1984;

1988; Moolenbeck et ah, 1991). Cases of valid taxa

being neglected are also not uncommon (see for in-

stance '"Rissod’' scillae Aradas et Benoit, 1876 ex

Seguenza G. in Verduin, 1984 and Gaglini, 1994).

We here examine the case of Rissoa galvagni

Aradas et Maggiore, 1 844, not 1 843 as reported by

Verduin (1984) and clarify ed by van Aartsen &

Giannuzzi Savelli (1987), described for the shores

of Ognina (Central Mediterranean, Ionian Sea, Ca-

tania), whose type material has long been considered

lost (Verduin, 1984). The taxon was discussed by

Verduin (1984) based on the examination of two lots

of topotypic material, allegedly attributed to Aradas

collection. One lot included only specimens of Cri-

silla pseudocingulata (Nordsieck, 1972) and the

other mainly specimens of this latter species and a

few ones of other congeneric species (Table 1).

None of those specimens was however considered

to match the original description and R. galvagni

was listed as nomen dubium. After the publication

of Verduin’s revision (1984) the taxon was no longer

mentioned in the literature, slipping into obscurity.

Although it was first described for the waters of

Ibiza (Balearic Islands), Crisilla pseudocingulata is

also abundant along the Eastern coast of Sicily (Io-

nian Sea) and its distinctive shell features make it

easy to differentiate it from other congeners. Many

of the most active malacologists in the XIX century

(such as Benoit, Philippi, Monterosato and Aradas)

extensively collected microgastropods along E Si-

cily and it appears surprising that C. pseudocingu-

lata was only found and described over a century

later. The composition of the material studied by

Verduin and the overlapping distribution of the two

taxa have always represented good evidence for a

case of synonymy of C. pseudocingulata and R. gal-

vagni (DS pers. obs.), but nothing could be proven

until now, when the type material of the latter taxon

was found amongst the lots of a recently re-disco-

vered section of the Aradas collection (MBAC)

(Scuderi, 2007). Based on the examination of this

material and additional material of R. galvagni and

C. pseudocingulata we here provide a revised and

updated description of R. galvagni and discuss the

evidence for the synonymy of the two taxa.

ACRONYMS AND ABBREVIATIONS. Bruno

Amati collection, Rome, Italy (BAG); Italo Noffoni

collection, Roma, Italy (INC); Museo del Diparti-

mento di Biologia Animale delfUniversita, Catania,

Italy (MBAC); Museo Civico di Zoologia, Roma,

Italy (MCZR); U.S. National Museum, Washington,

USA(USNM); Danilo Scuderi collection, Catania,

Italy (DSC); d = diameter of protoconch nucleus;

D = maximum diameter of protoconch; DO, = dia-

meter of protoconch first half whorl; DN-1 = diame-

ter of teleoconch; dry = dry shell; L = teleoconch

height; live = live collected specimen; M = aperture

height; N = number of whorls; SEM = scanning

electronic microscope.

MATERIALS AND METHODS

This study was based on dry shells, including

types of R. galvagni (MBAC), further museum ma-

terial and shells collected by the authors mainly in

the Ionian Sea and Southern Spain from 1982 to

2012 and housed in their private collections.

Shells were observed by stereo microscope,

photographed and measured by means of microme-

tre readings. Morphological characters of adult

shells (dimension, colouration, sculpture) were as-

sessed from a representative number of specimens

from each lot. Adults were recognised by a com-

plete apertural lip. Standard teleoconch parameters

(L, M, DN-1) were measured on the type material

and on further 20 specimens from three different lo-

calities. Protoconch parameters (d and DO) (Ver-

duin, 1982b) were measured and the number of

whorls (N), including protoconch, was counted

(precise to 0.1 as shown in Verduin, 1982b). Details

of protoconch and teleoconch microsculpture were

recorded using the SEM. Live collected specimens

of C. pseudocingulata were also observed under the

stereomicroscope and features of their head-foot re-

corded by means of colour drawings.

RESULTS

Observations and measurements of the studied

dry material (Tables 2, 3) revealed a total identity

Rediscovery and re-evaluation of a “ghost” taxon: the case of Rissoa galvagni (Caenogastropoda Rissoidae)

513

between samples R. galvagni and C. pseudocingu-

lata in shell size, morphology, maero-, mieroseul-

pture and eolouration. The evidenee eolleeted

supported the synonymy of the two taxa and the

reasons for this operation are diseussed below. An

updated deseription of the taxon is provided in the

systematie seetion based on the above mentioned

data on the shell and on new observations on the

head- foot. Twelve shells ineorreetly assigned to R.

galvagni were found in one MBAC lot

(AAC.005.02), isolated and plaeed in separate

boxes. These speeimens were not further exami-

ned in this study.

SYSTEMATICS

Family RISSOIDAE Gray, 1847

Genus Crisilla Monterosato, 1917

Crisilla galvagni (Aradas et Maggiore, 1844)

Rissoaria galvagni Aradas et Maggiore, 1844

Rissoa galvagni Aradas et Maggiore, 1844 in Calcara,

1845: 28

Cingula maculata Aradas et Benoit, 1870 not Montero-

sato, 1869

Cingula concinna Aradas et Benoit, 1870 not Montero-

sato, 1869

Rissoa picta Aradas et Benoit, 1870 not Jeffreys, 1867

Rissoa depicta Aradas et Benoit, 1870 not Manzoni, 1868

Rissoa galvagni Aradas et Maggiore, 1 844 in Aradas &

Benoit, 1870: 210

Rissoa galvagni var. maculata Monterosato, 1872 not

Monterosato, 1869

Rissoa galvagni var. concinna Monterosato, 1872 not

Monterosato, 1869

Rissoa galvagni var. picta Monterosato, 1872 not Jef-

freys, 1867

Rissoa galvagni var. depicta Monterosato, 1872 not Man-

zoni, 1868

Rissoa galvagni var. maculata Seguenza, 1873 not Mon-

terosato, 1869

Rissoa galvagni var. concinna Seguenza, 1873 not Mon-

terosato, 1869

Rissoa galvagni var. maculata Monterosato, 1875 not

Monterosato, 1869

Rissoa galvagni var. concinna Monterosato, 1875 not

Monterosato, 1869

1 Rissoa granulum Monterosato, 1875 not Philippi, 1844

Rissoa galvagni var. depicta Monterosato, 1875 not Man-

zoni, 1868

Rissoa galvagni var. callosa Monterosato, 1875 not Man-

zoni, 1868

Rissoa galvagni Aradas in Seguenza, 1876:180

Rissoa {Cingula) pulcherrima BDD, 1884 not Jeffreys,

1848

Cingula galvagnii Aradas et Maggiore, 1839 in Locard,

1886:266

Rissoa {Cingula) semistriata (Cams, 1893) not Montagu,

1808

Setia {Crisillosetia) pseudocingulata Nordsieck, 1972

Cingula (s.l.) pseudocingulata (Nordsieck, 1972) in Ver-

duin, 1984:50

Alvania {Crisilla) pseudocingulata (Nordsieck, 1972) in

Ponder, 1985:43-44.

Crisilla pseudocingulata (Nordsieck, 1972: Setia) in

CLEMAM (Oct 2012)

Rissoa galvagni Aradas et Maggiore, 1 844 incertae sedis

in CLEMAM (Oct 2012)

Material examined. Type material. Leetotype

here designated. Rissoa galvagni Aradas et Mag-

giore, 1844, Central Mediterranean, Ionian Sea,

Catania, Ognina, MBAC n° AAC.005.a. Paraleeto-

types: same data as leetotype (MBAC n°

AAC.005.00, 20 dry).

Other museum material: Rissoa galvagni Aradas

et Maggiore, 1844, (MBAC n° AAC.005.01, 10

dry); (AAC.005.02, 84 dry); Rissoa maculata Mon-

terosato, 1869 sub nomine calvagni (sie) Aradas,

Central Mediterranean, Ionian Sea, Catania, Ognina

(MCZR sine numera eabinet of typieal material, 3

dry). Newly eolleeted material: Central Mediterra-

nean, Ionian Sea, Punta Tonnara, Brueoli, -2m, roek

seraping (DSC, I dry); Catania porto, -2m, roek sera-

ping (DSC, 7 live); Catania porto, -10m, roek sera-

ping (DSC, 1 live); Riposto, -2m, roek seraping

(DSC, 12 live); Aeitrezza, Laehea Island, -2/-40m,

roek seraping (DSC, 344 dry, 21 live); S. Giovanni

Li Cuti, -l/-20m, roek seraping and shell grit (DSC,

77 dry, 12 live); Capo Molini, beaeh, shell grit (DSC,

206 dry); Aeitrezza, -2/-50m, roek seraping and fi-

shing nets byeateh (DSC, 408 dry, 46 live); Pozzillo,

-5/1 Om, shell grit (DSC, 2 dry); Ognina, roek sera-

ping and shell grit (DSC, 7 dry, 10 live); Catania “Ta-

vemetta”, shell grit (DSC, 9 dry); Aei Castello (DSC,

1 live); SW Mediterranean, Cabo de Palos, Mureia,

Spain, -5m (BAC, 2 dry; INC, 20 dry).

Description. Shell (Figs. 1, 2, 4, 5) small,

height (L) 1.4 - 2.1 mm, diameter (DN-1) 0.8 - 1.2

mm, height of aperture (M) 0.7-1 mm (Table 2),

solid, semitransparent, ovate-eonieal. 4. 5-4. 8

whorls moderately flat to eonvex, with a shallow

suture slightly eanalieulated. Protoeoneh (Fig. 3)

514

Danilo Scuderi & Bruno Amati

paucispiral (1.4 - 1.5 whorls), height 0.27 - 0.30

mm, nucleus slightly tilted, diameter of nucleus (d)

0.10 - 0.15 mm, diameter of first half whorl (DO)

0.20 - 0.24 mm, diameter maximum of protoconch

(D) 0.31 - 0.38 mm. Protoconch microsculpture of

one subapical spiral chord formed by dense granu-

les and separating protoconch in two sections:

upper one more shouldered, lower one more roun-

ded. Minute granules, from irregularly distributed

to regularly arranged into fine chords, also present.

Protoconch/teleoconch transition sharp, prosocline.

First teleoconch whorl sculpture of two, rarely

three, well-spaced spiral chords with interspaces de-

void of sculpture. Chords increasing in number from

second whorl, reaching up to 14-17 on body whorl,

always irregularly spaced and of different width; 8-

10 reaching the aperture only on its subsutural and

basal edges. Teleoconch microsculpture of dense

growth lines. Aperture large, ovate-rounded, shorter

than half of shell height, internally smooth; no labial

rib present on outer lip. Umbilical chink often pre-

sent. Shell colour yellowish-whitish, paler towards

base. Colour pattern of reddish rhomboidal spots ar-

ranged in two (upper whorls) to four (body whorl)

spiral rows. First row opistocline, below upper su-

ture; second row prosocline, above lower suture

(upper whorls) and above aperture (body whorl);

third row opisthocline, above aperture; fourth row

prosocline, irregular weak, at shell base.

Head-foot (Fig. 7) background colour white,

yellow stripes extending from eyes to ciliated ce-

phalic tentacles; head with two yellow spots bet-

ween tentacles, marked light brown stripe running

from snout to a half of head, large yellow spot on

opercular suspensor muscle, visible through thin

operculum. Three small metapodial tentacles on po-

sterior end of foot.

Remarks. According to the distributional data

collected here, C. galvagni occupies the West Me-

diterranean to Ionian Sea, and it is locally abundant

on the Eastern shores of Sicily. Some shells collec-

ted in Malta (6 dry, DSC) and Lampedusa Island (28

dry, DSC) revealed a considerable degree of distin-

ction from the typical C galvagni, but further stu-

dies are required to identify the sources of this

differentiation. Among the material of C. galvagni

here studied, a morph can be isolated from the typi-

cal C galvagni based on the following features:

smaller on average (L = 1 .6 mm, DN-1 =0.95 mm),

less slender, with more compressed base, whorls less

shouldered and more rounded, weak umbilical chink

always present. However, the morph shares proto-

conch features, chromatic pattern and habitat with

the typical form. Among Mediterranean rissoids,

shells of C. galvagni morphologically resemble

some smooth morphs of C. simulans (Locard 1886).

This latter species, however, is slightly smaller, it

has a more rounded protoconch, different teleoconch

macro- and microsculpture and head-foot colour.

C. picta (Jeffreys, 1867) and C. callosa (Man-

zoni, 1 868) from the Macaronesian region are very

similar to each other and to C. galvagni. Aradas &

Benoit (1872-76) synonymised C. galvagni with the

former. Compared to C. galvagni both Macarone-

sian species have thicker shells, more flattened

whorls and base, and lack the umbilical chink.

These two, almost similar, Macaronesian species

are however well distinguished by Verduin (1984),

and treated as separated species by Gofas et al.

(2001). Shells of C. galvagni with weaker sculpture

may resemble those of Rissoa maculata Montero-

sato 1869 (now Setia) due to their similar size and

morphology. Monterosato (1872) synonymised the

two taxa but later (1878) treated them as distinct

species. In the same paper (1878) another Mediter-

ranean “smooth rissoid”, Rissoa amabilis (now

Setia) was introduced for Rissoa pulcherrima A. A.

(being the taxon preoccupied by Rissoa pulcher-

rima Jeffreys 1848 from the Atlantic region).

Priolo (1952) misinterpreted Monterosato’s con-

clusions and erroneously subsumed all three taxa

under Cingula pulcherrima, adding further confu-

sion to the matter. Our data agree with Montero-

sato’s (1878) views and support the separation of

C. galvagni, S. maculata and S. amabilis.

DISCUSSION

Crisilla was erected by Monterosato (1917) to

separate species of Cingula, in which he listed the

only C. trifasciata (J. Adams, 1 800), from all the

numerous Mediterranean and Macaronesian species

morphologically similar to C. semistriata. It was

treated as sub-genus of Alvania Risso, 1 826 by Pon-

der (1985) but it was later given genus rank by Bou-

chet & Waren (1993) based on the morphological

homogeneity of shells of its species. All currently

accepted species of Crisilla have always been

source of taxonomic debate and their identification

Rediscovery and re-evaluation of a “ghost” taxon: the case of Rissoa galvagni (Caenogastropoda Rissoidae)

515

Label

Lot number

Verduin’s identification

n. es

R. galvagni Ar./. . .

182032

C. pseudocingulata

R. galvagni Ar./. . .

183211

C. pseudocingulata

“numerous”

C. maculata

C. turriculata

Table 1. Samples of Rissoa galvagni of USNM observed and identified by Verduin (1984).

TELEOCONCH

PROTOCONCH

Specimen

identifier

Dn-1

Leetotype

1.7

0.8

1.1

0.11

0.21

0.35

4.5

Paraleetotypes

1.6

0.8

1.0

0.14

0.24

0.35

4.5

1.6

0.8

1.2

0.11

0.22

0.35

4.5

2.0

0.9

1.2

0.12

0.22

0.34

4.5

1.8

0.8

1.1

0.14

0.24

0.35

4.8

1.7

0.8

1.0

0.11

0.20

0.35

4.8

1.8

0.9

1.1

0.14

0.24

0.32

4.6

1.5

0.8

1.0

0.14

0.24

0.32

4.5

1.9

0.9

1.1

0.11

0.21

0.35

4.6

1.8

0.9

1.1

0.12

0.21

0.32

4.5

1.6

0.8

1.0

0.12

0.2

0.32

4.6

2.0

1.0

1.1

0.15

0.22

0.32

4.6

1.8

0.9

1.2

0.14

0.21

0.35

4.2

1.8

0.9

1.2

0.12

0.24

0.35

4.5

1.7

0.8

1.0

0.14

0.22

0.38

4.6

1.7

0.8

1.1

0.14

0.21

0.35

4.8

1.6

0.7

1.0

0.12

0.21

0.37

4.6

1.8

0.8

1.1

0.12

0.21

0.35

4.6

1.5

0.7

1.0

0.12

0.22

0.34

4.6

1.6

0.8

0.9

0.14

0.21

0.34

4.2

1.9

0.8

1.1

0.11

0.20

0.32

4.4

Average

1.7

0.8

1.1

0.13

0.22

0.34

4.5

Standard

deviation

0.1

0.01

0.02

0.16

Table 2. Shell dimensions and whorl eount (see text for abbreviations) of the type material of Crisilla galvagni

(MB AC n°AAC.005.00)

516

Danilo Scuderi & Bruno Amati

has always been a diffieult task. In the XIX eentury

only few speeies now ineluded in this genus were

known and were eommonly regarded (along with

speeies of Setia) as minute and weakly seulptured

members of Rissoa (Aradas & Maggiore, 1840-

1844; Philippi, 1844; Caleara, 1845; Jeffreys,

1867). Later Monterosato (1872, 1875, 1878,

1884a, 1884b) reported order into the matter by re-

vising the taxonomy of many European Rissoidae,

deseribing new taxa and updating the distribution

(Mediteranean vs. non-Mediterranean) of new and

extant speeies, ineluding those of Crisilla.

The taxonomieal validity of ''Rissoa'' galvagni

was debated for almost one and a half eenturies

(Aradas & Benoit, 1872-76; Granata-Grillo, 1877;

Monterosato 1872 and 1878; Priolo, 1952; Verduin,

1984), but due to the loss of its type material and its

similarity with other rissoids the issue had remained

TELEOCONCH

PROTOCONCH

Specimen

identifier

Dn-1

Acitrezza

1.9

0.9

1.1

0.11

0.21

0.32

4.4

Cabo de Palos

1.9

0.9

1.1

0.12

0.21

0.34

4.4

S. G. Li Cuti

1.7

0.8

1.1

0.10

0.2

0.31

4.4

S. G. Li Cuti

1.7

0.8

1.2

0.11

0.21

0.32

4.6

1.5

0.8

0.12

0.22

0.34

4.5

1.6

0.8

1.0

0.12

0.23

0.34

4.6

1.5

0.7

1.0

0.12

0.22

0.32

4.5

1.8

0.8

1.2

0.13

0.24

0.32

4.6

1.8

0.9

1.1

0.10

0.20

0.32

4.6

1.9

0.9

1.1

0.11

0.21

0.33

4.4

1.8

0.8

1.0

0.14

0.24

0.32

4.5

Aeitrezza

2.0

1.0

1.1

0.12

0.24

0.32

4.8

1.8

0.9

1.2

0.14

0.24

0.32

4.5

1.7

0.8

1.0

0.12

0.24

0.32

4.6

2.0

0.9

1.2

0.15

0.24

0.32

4.8

1.6

0.8

0.9

0.12

0.22

0.34

4.5

1.9

0.9

1.2

0.15

0.24

0.34

4.8

1.6

0.8

1.1

0.11

0.21

0.32

4.4

1.4

0.7

0.8

0.12

0.21

0.32

4.0

1.7

0.8

1.1

0.11

0.20

0.32

4.4

Average

1.7

0.8

1.1

0.12

0.22

0.32

4.51

Standard

deviation

0.6

0.1

0.01

0.18

Table 3. Shell dimensions and whorl eount (see text for abbreviations) for newly eolleeted material of Crisilla galvagni',

values of speeimens illustrated in figs. 2-5 are highlighted.

Rediscovery and re-evaluation of a “ghost” taxon: the case of Rissoa galvagni (Caenogastropoda Rissoidae)

517

Figures 1-12. Crisilla galvagni. Fig. 1. Lectotype of “Rissoa galvagni” (h: 1.7 mm, DN-1: 1.06 mm). Fig. 2. SEM photograph

of shell from Acitrezza, h: 1.9 mm, DN-1: 1.1 mm; Fig. 3. Idem, detail of the protoconch. Fig. 4. Cabo de Palos, h: 1.9 mm,

DN-1: 1.08 mm. Fig. 5. S. Giovanni Li Cuti, h: 1.7 mm, DN-1: 1.05 mm. Fig. 6. Specimen of S. maculata labelled “Rissoa

galvagni” in the Monterosato’s Collection (MZR), h: 2 mm, DN-1 : 1.2 mm. Fig. 7. Drawing of C. galvagni from S. Giovanni

Li Cuti. Fig. 8. Label in Aradas’ handwriting of the lot n° AAC.005.00 (MBAC) from which the lectotype was selected.

Fig. 9. Label of the lot n° AAC.005.01 (MBAC). Figs. 10-12. R. galvagni, original labels, Monterosato’s collection (MCZR):

Monterosato’s handwriting (Fig. 10); Benoit’s handwriting (with notation “R. granulum Phil.” in Monterosato’s handwriting)

(Fig. 11); idem, back: notation in Monterosato’s handwriting (“Tipo dato dall’autore al Cav.er Benoit e da questo al Dr. Ti-

beri”) (Fig. 12). Scale bar: 1, 2, 4, 5, 6 = 0.5 mm; 3 = 0.1 mm.

518

Danilo Scuderi & Bruno Amati

so far unresolved. The homogeneity of the type ma-

terial (eontaining exelusively shells of R. galvagni),

was not observed for all lots of Aradas eolleetion

studied. One of them (MBAC AAC.005.02) eontai-

ned a mixture of other rissoid speeies similar to R.

galvagni. Among the material of Monterosato’s

eolleetion we found lots labelled ''Rissoa galva-

gnr (Figs. 6, 10, 11, 12) instead eontaining shells

of S. maculata.

These lots were most probably donated to Mon-

terosato by shells eolleetors, sueh as Tiber! and Be-

noit, who were also Aradas’ elose friends and

eollaborators. This is a elue of the extreme diffieulty

to aehieve a eoiTeet speeies identifieation of R. gal-

vagni, even for its deseriptor, whieh explains the

eonfiision emerged in the following studies exami-

ning this taxon. Another potential issue affeeting the

eorreet interpretation of R. galvagni was the poor

preservation state of its types, as not uneommon for

types of many other European rissoids (Verduin,

1984). The type material of R. galvagni eontained

a large amount of worn shells and it was therefore

neeessary to seleet a leetotype whose features

agreed at best with the original deseription. Our

operation provides the taxon with the required no-

menelatural stability. Two previous misinterpreta-

tion of the R. galvagni biased Verduin’s (1984)

eonelusions about the validity of this taxon. Priolo

(1952) and later Nordsieek (1972) erroneously pla-

eed it in synonymy respeetively with Setia pulcher-

rima (Jeffreys, 1848) and Rissoa soluta Jeffreys,

1 867 not Philippi, 1 844. Both authors did not study

the types of R. galvagni.

The material of R. galvagni studied by Verduin

(1984) was judged by him as not eorresponding to

the original deseription beeause it differed from it

by its shell not being smooth and by its different

shell height. Our observations on the type material

and on further lots of the Aradas eolleetion revealed

that their smooth shell surfaee is a result of these

shells being extremely worn. More aeeurate exami-

nation reveals that a seulptured shell is indeed a fea-

ture of C. galvagni. In addition, the shell seulpture

of Crisilla speeies is mueh weaker than that of other

rissoid (sueh as Alvania) and also quite variable,

ranging from eonsiderably seulptured to rather de-

void of seulpture (Ponder, 1985, Oliver et al., 2012).

Verduin’s (1984) eonelusion was henee based on a

very polymorphie eharaeter and therefore not sati-

sfaetorily supported.

Also his deeision (Verduin, 1984) was heavily

affeeted by a misinterpretation of the dimensions of

the type material. The shell height of R. galvagni

was provided by Aradas & Maggiore ( 1 844) in “Si-

eilian lines”, a measurement unit of length adopted

in the island, differing from the “line” used in the

rest of Italy and in most European eountries and

whieh was the one that Verduin (1984) had in mind.

The length in mm of the Sieilian line ean be in-

ferred from the deseription of Testacella haliotidea

(Benoit, 1857-62: 45-46): “... the higher speei-

mens are not less wide than 1 8 lines, whieh eorre-

spond to almost 41 mm”. This means that a

Sieilian line eorresponds to almost 2.5 mm. It is

also likely that the shell height in the original de-

seription of R. galvagni (less than half a line) was

published with a typographieal error. Aradas &

Maggiore (1844) reported R. mandralisci [= Pi-

sinna glabrata (Von Muehlfeldt, 1 824), eurrently

member of the Anabathride] as the smallest known

rissoid, with a maximum height of half a line. As-

suming that no other rissoid was eonsidered to

have a smaller size by those authors, the height of

R. galvagni should have been reported not as “less

than” but as “more than half line”. This latter value

eorresponds to about 1.5/1. 7 mm, whieh is the

average shell height of the shells of type material

as measured here (Table 2).

Being the issues related to shell seulpture and

size in R. galvagni solved and being all remaining

eharaeters ineluded in this taxon’s deseription eor-

responding to those of C. pseudocingulata, we here

eonsider this latter taxon to be a junior synonym of

the former one [ICZN, 1985 Artiele 24 (a)]. This

eonelusion is strongly supported by the observation

of the types and by their similarity with all other

material of C. pseudocingulata here studied.

ACKNOWLEDGEMENTS

We are grateful to Alberto Zilli and Massimo

Appolloni (MCZR) for their assistanee during the

study of the material of Monterosato’s eolleetion.

We are also indebted to Prof. Giorgio Sabella (Di-

partimento di Biologia Animate, University of Ca-

tania) and to Italo Nofroni (Rome) for providing

aeeess to the type material of Aradas’ eolleetion

(MBAC) and to his private eolleetion respeetively.

Stefano Palazzi (Modena, Italy) is thanked for ha-

Rediscovery and re-evaluation of a “ghost” taxon: the case of Rissoa galvagni (Caenogastropoda Rissoidae)

519

ving provided material and suggestions. Thanks are

also due to Franeeseo Criseione (Australian Mu-

seum, Sydney) and another anonymous referee who

provided useful eomments and suggestions to the

improvement of the manuseript.

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Biodiversity Journal, 2012, 3 (4): 521-526

New observations on the taxonomy, biology and distribution

of Tricolia landinii Bogi et Campani, 2007 (Gastropoda Vetiga-

stropoda)

Danilo Scuderi' &Agatino Reitano^

'Via Mauro de Mauri, 15b Piano Tavola - 95032 Belpasso, Catania, Italy; e-mail: danscu@tin.it

Via Gravina, 77 - 95030 Tremestieri Etneo, Catania, Italy; e-mail: tinohawk@yahoo.it

ABSTRACT Tricolia landinii Bogi et Campani, 2007, is here reviewed on the basis of both shell morphology

and observations of the living animals. This taxon, is here aseertained, it was deseribed on the

basis of only shell eharacters of young specimens, without the study of external soft parts. New

data about adult shell morphology, living animal and distribution of this minute species are

here furnished, together with a detailed iconography. On the basis of these characters T. landinii

appears more similar to the T. tingitana group, rather than to T nordsiecki, as underlined in the

original description. In the light of the adult shell morphology here reported and of the living

animal’s features, a complete analysis of the entire group of this “small Tricolia’', with com-

parisons to the close resembling species, is here furnished.

KEY WORDS Tricolia landinii', Phasianellidae; juveniles; Mediterranean; re-description; adult shell.

Received 12.05.2012; accepted 03.11.2012; printed 30.12.2012

Proceedings of the P' International Congress “Insularity and Biodiversity”, May 11*-13*, 2012 - Palermo (Italy)

INTRODUCTION

The dichotomic key of classification of species

of Tricolia Risso, 1826 made by Gofas (1986) was

based on shell and radular characters while a second

contribution (Gofas, 1993) was based on both shell

and external soft parts characters of species.

According to these papers European species of

Tricolia are generally arranged in two different

groups: one comprises larger species,

T. miniata (Monterosato, 1884),

T. petiti (Craven, 1882),

T. pullus (Linnaeus, 1758),

T. speciosa (Muhlfeldt, 1824)

T. tenuis (Michaud, 1 829)

This first group is taxonomically rather stable,

being constituted by well-known species almost ea-

sily to recognise, even if some more accurate stu-

dies should better define the taxonomical status of

some taxa as T. pullus azorica (Dautzenberg, 1889),

T. pullus canarica Nordsieck, 1973, T. pullus picta

(da Costa, 1778), or some morphs of T. tenuis.

A second group instead is constituted by seven

species all of small dimensions:

T. algoidea (Pallary, 1 920)

T. deschampsi Gofas, 1993

T. entomoeheila Gofas, 1993

T. landinii Bogi et Campani 2007

T. nordsiecki (Talavera, 1978)

T. punctura Gofas, 1993

T. tingitana Gofas, 1982.

A part the small dimensions, all these species

share a similar globose shape of the shell, a similar

creamy colour with darker strips and stains and a

rocky shallow water habitat. T. tingitana, endemic

to S- Spain, was recently cited from eastern Medi-

terranean, on the basis of abundant both living and

fossil materials collected in several localities of the

522

Danilo Scuderi &Agatino Reitano

eastern eoast of Sieily and Calabria (Seuderi &

Russo, 2003). After few years T. landinii Bogi et

Campani 2007 was deseribed as a new speeies ende-

mie from E-Sieily and eompared with T. nordsiecki

as the most similar speeies, beeause of its sharp ou-

tline, flat protoeoneh and first teleoeoneh whorl and

umbilieus without keel.

More reeently, further studies on the growth of

the shell and the living of speeies of Tricolia here

eondueted aseertained the identy of the Sieilian po-

pulation of T. tingitana (sensu Seuderi & Russo,

2003) with T. landinii. The deseription of this latter

speeies, in faet, was based on young speeimens, as

eould be argued by the eharaeters showed in the ori-

ginal deseription eompared to those of the materials

here studied. The deseription given by Bogi &

Campani (2007) based on young speeimens renders

an image of T. landini as more similar to flattened

shell Tricolia, i.e. T. nordsiecki, rather than to the

T. tingitana group, whieh share the presenee of a

more high spired shell. This eaused misidentifiea-

tions and taxonomieal problems linked to the eor-

reet identifieation of this speeies.

In the present paper a more aeeurate study on

abundant material of T. landinii, reeent to fossil

shells and living eolleeted speeimens, allowed a

more preeise definition of this speeies and a re-de-

seription, whieh eomprises the growing stages of

the speeies, detailed informations on shell as well

as external soft body parts, a more appropriate

eomparison to the elosest similar speeies and a

eomplete distribution of the speeies, whieh seem

not to be peeuliar of Sieily.

ACRONYMS AND ABBREVIATIONS. The

materials used for this study are deposited in the

following private eolleetions: Agatino Reitano

eolleetion, Italy, Catania (ARC); Danilo Seuderi

eolleetion, Italy, Catania (DSC); live = live eollee-

ted speeimen; dry = dry shell.

MATERIALS AND METHODS

This study was based on both dry shells and li-

ving speeimens eolleeted by the authors mainly in

the Ionian Sea (E-Sieily) 1984 to 2012 and housed

in their private eolleetions. Further materials from

Messina Strait and Tyrrhenian Sea are here eonsi-

dered eonspeeifie with those of Sieily on the basis

of shell eharaeters only.

Living eolleeted materials were stored in aqua-

rium, observed while erawling and drown. Photo-

graphs were obtained at the stereoseope with a

Nikon Coolpix 4500. Images were adjusted with a

eommon Image editing software. Systematies in the

present paper follow Clemam (2012).

RESULTS

Observations at the steroseope allowed to study

the shell eharaeters of T. landinii and its growth sta-

ges. Living animals furnished further important

eharaeters whieh give us the eertainty that a speeies

different from others S- Spain elose similar speeies

is involved.

Here follows a re-deseription of the adult shell

eharaeters based on newly eolleeted materials and

on living animal, whieh adjust as for some morpho-

logieal eharaeters and eomplete the previous de-

seription of this taxon, laeking the soft body

features as diseriminating eharaeter.

SYSTEMATICS

elass GASTROPODA Cuvier, 1795

ordo VETIGASTROPODA Salvini-Plawen &

Haszprunar, 1987

Family PHASIANELLIDAE Swainson, 1840

Genus Tricolia Risso, 1826

Tricolia landinii Bogi et Campani, 2007

Examined Material. Italy, Sieily: Cajto, Cata-

nia, -2/3m, samples obtained after Gobius efr. pa-

ganellus Linnaeus, 1758 stomaeh eontents (DSC, 4

live); Ognina, Catania, tide pools, -0.2m, samples

obtained after washing algae and shell grit (ARC

and DSC, 220 live; 450 dry); Cannizzaro, Catania,

-0. l/45m, samples obtained after washing algae and

shell grit (ARC 25 live; 150 dry), Aeieastello, Ca-

tania, tide pools, -0.1m, samples obtained after wa-

shing algae (ARC 25 live); Aeitrezza, Catania,

-0.2/2m, samples obtained after seraping of the

roeky substrate on surfaees of 20x20 em (DSC, 156

live, 138 dry); Capo Mulini, Catania, tide pools,

-0.1m, samples obtained after washing algae (ARC

25 live) and shell grit (DSC 200 dry); S. Giovanni

Li Cuti, Catania, -l/2m, samples obtained after wa-

New observations on the taxonomy, biology and distribution of Tricolia landinii (Gastropoda Vetigastropoda)

523

Figures 1-9. T. landinii. Figs. 1, 2. Adult shell from Aeitrezza (2.2 x 2.0 mm). Fig. 3. Adult speeimen seen from aside

showing external lip (height 1.8 mm). Fig. 4. Not fully grown entirely blaekish speeimen seen from aside showing external

lip (height 1.3 mm). Figs. 5-6. Not fully grown speeimens eorresponding to the original deseription (Fig. 5 height 1.1 mm;

Fig. 6 height 1.2 mm). Fig. 7. Juvenile stage (height 0.7 mm). Fig. 8. Speeimen from Calafuria, Tuseany (height 1.1 mm).

Fig. 9. Drawing of living animal with detail of right and left neek lohes (height 1.8 mm).

524

Danilo Scuderi &Agatino Reitano

shing algae and shell grit (DSC 25 live, 120 dry);

Ganzirri, Messina, -50m, samples obtained after re-

siduals of fishing nets and shell grit (DSC, 2 live; 4

dry). Italy, Calabria: Seilla, Reggio Calabria, tide

pools, -0.2m, samples obtained after washing algae

and shell grit (ARC and DSC, 15 live; 46 dry); Laz-

zaro, Reggio Calabria, shell grit (DSC, 13 dry). Ra-

vagnese, Reggio Calabria, fossil (DSC, 1). Italy,

Tuseany: Calafuria, Pisa, -2m, samples obtained

after seraping of the roeky substrate on surfaees of

20x20 em (DSC, 2 live).

Description. Shell. Teleoeoneh of adult speei-

mens (Figs. 1-3) reaehing 2.4 x 2.0 mm eonstituted

by 3.5 whorls, the last very ample. Seulpture of

dense growth lines, of whieh some more relevant

are evident on the first tele-whorl as thin axial ribs

riblets (Figs. 4, 6, 8); surfaee of shell eovered by

faint spiral mieroseulpture, deteetable only at high

magnifieation of the stereoseope, but almost smooth

on the body whorl, with exeeption of the umbilieal

area whieh bears a narrow but deep umbilieus sur-

rounded by a marked umbilieal keel. Protoeoneh

flat, eonstituted approximately by one whorl 200

pm in diameter, eovered by subtle spiral treads.

Colour. Protoeoneh almost whitish; first tele-

whorl blaekish, beeoming ereamy in subsequent

whorls, with dark blaek or purplish axial flames ir-

regularly arranged and white dots arranged in two

spiral rows, the first abapieally and the latter ada-

pieally. Uniformly red-brown and blaekish speei-

mens (Fig. 4) are known, some others are laeking

the white dots.

Animal (Fig. 9). Entirely green, with three pairs

of epipodial tentaeles almost of the same length;

right neek lobe broad, with edge not notehed and

left neek lobe double fingered. Opereulum round,

white, relatively thiek and paueispiral, with nueleus

externally visible (Figs. 1, 5, 7).

Biology and Distribution. Type loeality: Por-

tieello. Villa San Giovanni (Reggio Calabria), Strait

of Messina, Italy.

The speeies is present along the Ionian eoast of

Sieily, exeluding the ealeareous roeky shore of the

Southern part of the Island. Further material from

Messina Strait and Tuseany seems to demonstrate

the not endemie status of the speeies, though the

identifieation of this material needs eonfirmation

with the observation of the living animal features.

The speeies eommonly lives in shallow roeky sho-

res, 0,10/4-6 m depth, on the red algae of the spe-

eies Pterocladiella capillacea (S.G. Gmelin) San-

teliees & Hommersand, 1997.

Paleontology. The fossil material aseribable

to this speeies, found at Ravagnese, Reggio Cala-

bria, whieh belongs to Tyrrhenian Stage suggests

the presenee of this speeies in the Messina Strait

area sinee middle/late Pleistoeene.

CONCLUSIONS

Aeeording to the original deseription (Bogi &

Campani, 2007) T. landinii is eharaeterised by: ge-

neral outline remarkable low, protoeoneh and first

tele-whorl flattened, prosoeline and sinuous outline

of external lip, faint umbilieus without any keels.

The examination of a large quantity of topoty-

pie, both eomplete and not full grown speeimens of

T. landinii allowed us to aseertain the juvenile spe-

eimens on whieh the original deseription of this

speeies was made and thus to re-deseribe it. Juve-

nile speeimens, in faet, are eharaeterised by the al-

ways whitish protoeoneh whorls whieh beeome

almost blaek with the eonstruetion of the first teleo-

eoneh whorl and show a faint umbilieus, without

any keel and in an almost blaek basal area (Fig. 7).

The seeond tele -whorl assumes the eolour of adult

speeimens, beeoming yellowish with red-brownish

longitudinal strips and white dots. The feature and

eolour of this stage, slightly more than 1 mm high,

perfeetly eorresponds to the original deseription of

holotype of T. landinii.

The adult shell is bigger, exhibits a general not

flattened form, forni, with a narrow umbilieus bor-

dered by a sharp keel in a white area and the eolo-

ration above deseribed for the seeond tele-whorl,

whieh eould be rather variable eonsidering hun-

dreds speeimens of an entire population.

The whole shell assumes therefore a form mar-

kedly different from that deseribed in the original

deseription, being more elosely to the T. tingitana

group of speeies, whieh eomprises T. tingitana, T

deschampsi, T entomocheila and T. punctura, rather

than to T. nordsiecki (Fig. 10) or juvenile stages of

T. miniata, to whieh was originally eompared. In

partieular the shell of T. landinii differs from that

of T. entomocheila for the absenee of the deep noteh

in the subsutural part of the outer lip. The shell of

T. landinii also differs from that of T. punctura.

New observations on the taxonomy, biology and distribution of Tricolia landinii (Gastropoda Vetigastropoda)

525

Figure 10. Schematic drawing of the types of shell in Tricolia tingitana (A) and T. nordsiecki (B) species and of the living

animals of the T. tingitana group: T. deshampsi (1), T. tingitana (2) and T. landinii (3). Black arrows indicate the second

epipodial tentacles (absent in the former species); red arrows indicate the left neck lobe.

which lives in the Strait of Messina too, beeause

of its smaller dimensions, in having well rounded

whorls, less laterally eompressed; T. punctura has

a different eolour pattern, being almost unique

among speeies of this group of small Tricolia

(Gofas, 1993).

The adult shell of T. landinii resembles more

that of T. tingitana and T. deschampsi. The former

of this two speeies is reported as endemie to the

S-Spain (Gofas, 1982). More reeently T. tingitana

was erroneously eited for E-Sieily (Seuderi &

Russo, 2003) on the basis of the elose resem-

blanee of the shell, whieh is almost indistingui-

shable from that of T. landinii. Studies here

eondueted allowed us to aseertain the identity of

material of T. tingitana sensu Seuderi & Russo

(2003) with T. landinii, mainly on the basis of the

features of the living animal (Fig. 9).

Moreover, the presenee of 6-7 spiral eords on

the first teleoeoneh whorl and of three shallow si-

nuosities on the outer lip allow the distinetion of

shells of T. deschampsi from those of T. tingitana

and T. landinii.

Here, the examination and eomparison of the

living animals allowed a better separation among

these speeies aeeording to the following differen-

ees (Fig. 10): T deschampsi has only two pairs of

epipodial tentaeles, while T. landinii and T. tingi-

tana have three pairs of epipodial tentaeles. The

eolour of the main parts of the body soft parts of

T. tingitana is almost purplish while in T. landinii

is entirely green. Moreover in the former speeies

the left neek lobe brings 5-7 long digitations, while

in T. landinii is only double fingered.

Further not Sieilian material, here attributed to

T. landinii on the basis of shell eharaeters only, ex-

526

Danilo Scuderi &Agatino Reitano

tends the distribution of this speeies to the entire

western eoasts of Italy.

ACKNOWLEDGEMENTS

We would like to thank the anonymous referees

who improved the manuseript whith useful eom-

ments and suggestions.

REFERENCES

Bogi C. & Campani E., 2007. Tricolia landinii, una

nuova specie per le coste orientali della Sicilia. Ibe-

rus, 25: 27-31.

Clemam, 2012. Taxonomic Database on European Ma-

rine Molluscs, (http://www.somali.asso.fr/clemam/

index.php). (last access: 08.12.2012).

Giannuzzi Savelli R., Pusateri R, Palmeri A. & Ebreo C.,

1997. Atlante delle conchiglie marine del Mediterra-

neo, Vol.l: Archaeogastropoda. Edizioni La Conchi-

glia, Roma, 125 pp.

Gofas S., 1982. The genus Tricolia in the Eastern Atlantic

and the Mediterranean. Journal of Molluscan Studies,

48: 182-213.

Gofas S., 1986. Taxonomic des Tricolia Mediterraneen-

nes. Lavori della Societa Italiana di Malacologia, 22:

179-194.

Gofas S., 1993. Notes on some Ibero-Moroccan and Me-

diterranean Tricolia (Gastropoda, Tricoliidae), with

description of new species. Journal of Molluscan Stu-

dies, 59:351-361.

Monterosato T. di Maria di, 1884. Nomenclatura generica

e specifica di alcune conchiglie mediterranee. Pa-

lermo, Virzi, 152 pp.

Sabelli B., Giannuzzi Savelli R. & Bedulli D., 1990.

Catalogo annotate dei molluschi marini del Medi-

terraneo. Libreria naturalistica bolognese, Bologna,

1-3.

Scuderi D. & Russo G.F., 2003. Due nuovi Gasteropodi

per le acque italiane: Melibe fimbriata Alder & Han-

cock, 1864 e Tricolia tingitana Gofas, 1982 (Mollu-

sca: Gastropoda). Biologia Marina Mediterranea, 10:

618-621.

Biodiversity Journal, 2012, 3 (4): 527-542

A new record for the Italian fauna: Plagyrona placida (Shuttle-

worth, 1 852) from Sardinia and Southern Italy (Gastropoda

PulmonataValloniidae)

Simone Cianfanelli'*, Gianbattista Nardi^ & Marco Bodon^

'Museo di Storia Naturale, Sezione Zoologica de "La Specola", Universita di Firenze, Via Romana 17, 50125 Firenze, Italy;

e-mail: simone.eianfanelli@unifi.it

^Via Bosehette 8/ A, 25064 Gussago, Breseia, Italy; e-mail: gbnardi@libero.it

^Dipartimento di Seienze Ambientali dell'Universita di Siena, Via RA. Mattioli 4, 53100 Siena, Italy; e-mail: mabodon@tin.it

*Corresponding author

ABSTRACT Plagyrona placida (Shuttleworth, 1852) is a terrestrial minute species with a wide but frag-

mented distribution, known from some of the Macaronesian Islands of the Canaries and the

Madeira archipelago, from Corsica Island, from some European countries (Portugal, Albania

and Greece) and from Northern Africa (Algeria). This species has been recently discovered

in Italy (Sardinia, Campania and Calabria) for the first time; data of sampling and the charac-

teristics of the Italian populations are discussed in this note. P. placida lives in the Mediter-

ranean forest or bush environments, but its specific habitat is not known because it has been

found, at least in Italy, in alluvial debris collected along streams and in litter. Even if this spe-

cies has not been recorded until now, the undisturbed habitat and its rarity suggest that it may

be native to Italy, and not accidentally introduced by man through trees used for reforestation

or through imported vegetables, as already happened for others small species.

KEY WORDS Valloniidae; Plagyrona placida; Italy; Sardinia; Campania; Calabria.

Received 12.05.2012; accepted 12.12.2012; printed 30.12.2012

Proceedings of the L' International Congress “Insularity and Biodiversity”, May 11*-13*, 2012 - Palermo (Italy)

INTRODUCTION

The family Valloniidae Morse, 1864, ineludes

helieoid minute molluses, usually living in the soil

or in the litter, with eosmopolitan distribution (Sehi-

leyko, 1998).

In Europe it is represented by the gonora Acan-

thinula Beek, 1 847, Gittenbergia Giusti et Manga-

nelli, 1986, Plagyrona Gittenberger, 1977,

Spermodea^QSiQvlmid, 1902, ValloniaPdsso, 1826

and Zoogenetes Morse, 1864, with about 15 known

speeies (Sehileyko, 1998; Bank, 2011a; Holyoak

& Holyoak, 2012). While the genus Vallonia inelu-

des several speeies, the genera Acanthinula, Sper-

modea and Plagyrona inelude few speeies and the

remaining genera inelude only monospeeifie taxa

(Gerber, 1996; Falkner et al., 2001; Bank et al.,

2002; Bank, 2011a; Holyoak & Holyoak, 2012).

In Italy, until now, six speeies belonging to the

family Valloniidae are known: Acanthinula acu-

leata (Muller, 1774), Gittenbergia sororcula (Be-

noit, 1859), Vallonia costata (Muller, 1774), V.

enniensis (Gredler, 1856), V. pulehella (Muller,

1774) and V. sueviea Geyer, 1908, the last one only

reeently reported (Manganelli et al., 1995; Bank,

2011b; another taxon, V. exeentrica Sterki, 1893,

has been eonsidered as eonspeeifie with V pulehella

by some authors). A seventh speeies, Plagyrona

528

S. ClANFANELLI, G. NaRDI & M. BODON

placida (Shuttleworth, 1 852), is now reported in

this paper from Sardinia, Campania and Calabria.

P. placida has a fairly wide but fragmented di-

stribution, having been reported for the Canary Is-

lands (El Hierro, La Gomera, La Palma, Tenerife)

(Wollaston, 1878; Ibanez et al., 2001; Bank et al.,

2002; Brito & Fraga, 2010), the islands of Madeira

and Porto Santo (Wollaston, 1878; Walden, 1983;

Bank et al., 2002; Cameron et al., 2007; Seddon,

2008), Portugal (Servain, 1880; Silva e Castro,

1887; Loeard, 1899; Gittenberger, 1977; Oliveira,

2008, 2009, 2010; Torres & Oliveira, 2010; Bank,

2011a; Holyoak & Holyoak, 2012), Corsiea (Rip-

ken & Bouehet, 1998; Falkner et al., 2002), Albania

(Reisehhiitz et al., 2008; Ferher & Eross, 2009;

Bank, 201 le) and Greeee (Gittenberger, 1989;

Bank, 201 Id); moreover, it was also surveyed in

north-eastern Afriea, in Algeria (Bourguignat, 1863,

1864; Gittenberger, 1977).

Being of small size, rather rare and loealized,

therefore diffieult to find, P. placida might have

eseaped non speeifie searehes performed in other

eountries, so that its known distribution may have

been underestimated. In Madeira and Porto Santo

the speeies has been found also as a fossil (Cook et

al., 1993; Cameron et al., 2006) so, in these plaees,

it is undoubtedly to be eonsidered a native speeies;

on the other hand, in other eountries, its auto-

ehthony is not proven with eertainty, even if there

are no elements in favour of a passive diffusion.

Historical notes

"Helix placida" was deseribed by Shuttleworth

(1852) on the basis of some speeimens found on the

island of Tenerife, in the Canary arehipelago. A se-

eond deseription of this entity was made by Bour-

guignat (1863) who, eolleeting this minute snail in

Algeria, named it "Helix debeauxiana" (taxon now

aeknowledged as a junior synonym of Plagyrona

plaeida; Walden, 1983). Lowe (1855) established a

"variety" of" Helix pusilla", taxon deseribed by him

some years before from Madeira (Lowe, 1831),

whieh he ealled "Helix pusilla var. p. serieina". But,

whereas "Helix pusilla" Lowe, 1831 is a younger

synonym of Paralaoma servilis (Shuttleworth,

1852) (Falkner et al., 2002), the new taxon "Helix

pusilla var. p. serieina" ean be attributed to Plagy-

rona placida (Bank et al., 2002).

Then reports of "Helix debeauxiana" followed

from Portugal (Servain, 1880) and deseriptions of

"Helix luseana" (Paiva, 1866) from the island of

Madeira and "Helix bussaeona" (Silva e Castro,

1887) from Portugal, too, all nowadays eonsidered

as synonyms of the speeies (Gittenberger, 1977;

Bank et al., 2002).

Gittenberger (1977) deseribed the new genus Pla-

gyrona, on the basis of the peeuliar spiral miero-seul-

pture on protoeoneh and teleoeoneh of the shell, to

separate "Helix debeauxiana" by the others Vallonii-

dae. In that work he briefly deseribed also the radula.

In the last deeades P placida was surveyed for

the fauna of Franee, having been found in a site of

Corsiea, near Coti-Chiavari, in the SW of the island

(Ripken & Bouehet, 1998), in an Albanian site, pla-

eed E of Rrogozhine (Reisehiitz et al., 2008) and in

Greeee, in the Ionian islands of Kerkyra, Ithaea and

Cephalonia (Gittenberger, 1989).

Finally, Holyoak & Holyoak (2012) have reeen-

tly separated the populations of Portugal into two

different taxa, with the deseription of a new speeies

Plagyrona angusta Holyoak & Holyoak, 2012. This

speeies has been established on a few morphometrie

eharaeters of the shell, as a narrower diameter and

a smaller umbilieus, and, even though it is not al-

ways well distinguishable from P placida, the sym-

patry of the two taxa may support its distinet

speeifie status. Unfortunately, even if the authors

found living speeimens of both taxa, they have not

reported any anatomieal data and therefore the ana-

tomy of Plagyrona is still unknown, so the alloea-

tion of this genus to the family Valloniidae still has

some margin of uneertainty.

While for the European Mediterranean areas no

detailed information about the habitat of eolleetion

of P placida is available, it is known for Portugal,

Algeria and for the Maearonesian Islands. In Por-

tugal the speeimens have been sieved from leaf-lit-

ter on roeky slopes of limestone eovered by bushes

and herbs (Seddon & Tattersfield, 1992) or in

humid roeky limestone habitats underneath deei-

duous woodlands, on trunks and branehes eovered

with epiphytie mosses or on low limestone roeks

eovered of mosses (Holyoak & Holyoak, 2012). In

Algeria the speeies has been found under leaves,

among hypnoid mosses eovering oak trees, prefer-

ring upper sides of large horizontal branehes (Bour-

guignat, 1863; Letorneux, 1870), while in the

islands of Madeira and in the Canary Islands the

speeies has been found in laurel forests on mosses

and liehens on damp trunks (Shuttelworth, 1852;

A new record for the Italian fauna: Plagyrona placida from Sardinia and S. Italy (Gastropoda Pulmonata Valloniidae) 529

Paiva, 1866; Wollaston, 1878), in rocky sites above

the ground on damp tree trunks or in leaf-litter or

in litters, mosses and branehes of laurels (Seddon

& Holyoak, 1993; Seddon, 2008) or in humid laurel

forests or Erica arborea L. serubs, on mosses or

trunks (Holyoak & Holyoak, 2012).

MATERIALS AND METHODS

Shells of P. placida were eolleeted in alluvial

debris or in litter: the sediments were sieved using

deereasing mesh sieves and the speeimens were se-

parated visually or using a binoeular mieroseope.

The photographs of the shells have been taken

with the aid of a binoeular mieroseope and related

software. Details of the protoeoneh and teleoeoneh

were obtained from samples mounted on alumi-

nium supports eovered by eonduetive glue, sputter-

eoated with graphite and gold, and examined using

a seanning eleetron mieroseope (SEM). All dimen-

sions (shells height, shells diameter, aperture height

and aperture diameter) were measured using a mi-

erometer in the light mieroseope.

The eolleetion data are listed as follows: loeality,

altitude, munieipality and abbreviation of the pro-

vinee in parentheses, UTM eoordinates (ED 50),

eolleetors and dates, number of speeimens in paren-

theses. Names of the loealities were taken from the

offieial map of Italy by I.G.M.I.; UTM eoordinates

were taken from the same maps or deteeted by GPS.

The examined material is presently preserved in

the following eolleetions: Museo di Storia Naturale

deirUniversita di Firenze, sezione di Zoologia de

"La Speeola", Via Romana 17, Florenee, Italy

(MZUF); M. Bodon, Via delle Eriehe 100/8, Genoa,

Italy, (MBC); S. Cianfanelli, Via Monferrato 3, Flo-

renee, Italy (SCC); E. Talenti, Piazza Parri 4, Ineisa,

Florenee, Italy (ETC); G. Nardi, Via Bosehette 8/A,

Gussago, Breseia, Italy (GNC).

RESULTS AND DISCUSSION

Plagyrona placida (Shuttleworth, 1852)

Helix placida 1852: 140

Helix placida, Pfeiffer, 1853: 82-83

Helix pusilla var. p. serieina Lowe, 1855: 176

Helix Debeauxiana Bourguignat, 1863: 183-184,

PL 19, figs. 13-16

Helix Debeauxiana, Bourguignat, 1864: 308, 329

Helix luseana Paiva, 1866: 342-343, PL 11, fig. 9

Helix luseana, Paiva, 1867: 80, PL 2, fig. 3

Helix Debeauxiana, Letoumeux, 1870: 277, 279

Patula plaeida, Pfeiffer, 1870: 40-41, 62, PL 120,

figs. 9-12

Patula placida, Mousson, 1872: 25, PL 2, figs. 9-12

Helix placida, Pfeiffer, 1876: 139

Patula placida, Wollaston, 1878: 63, 87-88, 282,

331,481,570

Helix Debeauxiana, Servain, 1880: 61-62

Helix Debeauxiana, Silva e Castro, 1887: 246 (re-

eords to eonfirm aeeording to Holyoak & Ho-

lyoak, 2012)

Helix bussaeona Silva e Castro, 1887: 246

Helix Debeauxiana, Tryon, 1887: 28-29, 275. PL 6,

figs. 31-33

Helix luseana, Tryon, 1887: 31, 275. PL 6, figs. 59-60

Helix plaeida, Tryon, 1887: 51, 275. PL 9, fig. 94

Helix (Punetum) debeauxiana, Westerlund, 1889: 8

Helix (Punetum) bussaeona, Westerlund, 1889: 9

Pyramidula (Pyramidula) bussaeona, Pilsbry,

1894: 44

Pyramidula (Pyramidula) debeauxiana, Pilsbry,

1894: 44

Patula (Punetum) debeauxi, Kobelt, 1898: 47, PL

225, fig. 1434

Helix bussaeona, Kobelt, 1898: 47

Helix Debeauxiana, Loeard, 1899: 72

Helix Bussaeona, Loeard, 1899: 73

Plagyrona debeauxiana, Gittenberger, 1977: 297-

303, Figs. 3, 4; PL 1, figs. 1, 2; PL 2, figs. 1-6

Plagyrona plaeida, Walden, 1983: 266, 268

IPlanogyra sororcula, Palazzi, 1988: 17 (not Helix

sororcula 1859)

Plagyrona placida, Gittenberger, 1989: 14, Figs. 4, 5

Plagyrona placida. Cook et al., 1990: 50, 72, 74, 76

Plagyrona placida, Fidalgo & Callopez, 1990: 80

Plagyrona placida, Ripken & Bouehet, 1998: 15

Plagyrona placida, Seddon & Tatterslield, 1992: 259

(reeord from Algarve to co nfir m according to Ho-

lyoak & Holyoak, 2012)

Plagyrona placida. Cook et al., 1993: 83, 93, 96,

99-103

Plagyrona placida, Seddon & Holyoak, 1993: 326

Plagyrona placida, Goodfriend et al., 1994: 319

Plagyrona placida, Sehileyko, 1998: 98-99, Fig. Ill

Plagyrona placida, Cameron & Cook, 2001: 262

Plagyrona placida, Falkner et al., 2001: 34

Plagyrona placida, Ibanez et al., 2001: 147

530

S. ClANFANELLI, G. NaRDI & M. BODON

Plagyrona placida. Bank et al., 2002: 102, 140,

151, 157, 174, 187, 195

Plagyrona placida, Falkner et al., 2002: 37, 107

Plagyrona placida, Albuquerque de Matos, 2004: 38

Plagyrona placida, Cameron et al., 2006: 31-33,35,41

Plagyrona placida, Cameron et al., 2007: 15, 19

Plagyrona placida, Oliveira, 2008: 41 (reeords to

eonfinn aeeording to Holyoak & Holyoak,

2012)

Plagyrona placida, Reisehiitz et al., 2008: 38

Plagyrona placida, Seddon, 2008: 37; PL 5D; Map

37, 125

Plagyrona placida, Feher & Eross, 2009: 27

Plagyrona placida, Kappes et. al., 2009: 585

Plagyrona placida, Oliveira, 2009: 55-56 (reeords

to eonfirm aeeording to Holyoak & Holyoak,

2012)

Plagyrona placida, Brito & Fraga, 2010: 188

Plagyrona placida, Fontaine et al., 2010: 24

Plagyrona placida, Oliveira, 2010: 42-43 (reeords

to eonfirm aeeording to Holyoak & Holyoak,

2012)

Plagyrona placida, Torres & Oliveira, 2010: 32 (re-

eords to eonfirm aeeording to Holyoak & Ho-

lyoak, 2012)

Plagyrona placida. Bank, 2011a

Plagyrona placida. Bank, 201 le: 25

Plagyrona placida. Bank, 20 lid: 14

Plagyrona placida, Gargominy et al., 2011: 326

Plagyrona placida, Welter-Sehultes, 2011

Plagyrona placida, Holyoak & Holyoak, 2012:

153-165, Figs. 1 A-C, 3E,F

Plagyrona placida, Welter-Sehultes, 2012: 205

Description. Shell (Figs. 1-11) very small (1.2- 1.6

mm in height; 1 .6-2.3 mm in diameter; Table 1), de-

pressed, with 3% - 3 V 2 eonvex and slowly expanded

whorls, separated by a deep suture; spire not eleva-

ted; last whorl little wide and slightly deseending

near the aperture. Protoeoneh not protruding, with

the surfaee eovered by many thin spiral striae and

spiral groves, erossed with more spaeed and less

marked growth lines; teleoeoneh eovered by dense

periostraeal ribs, elearly visible and equal to eaeh

other, and by thin spiral lines, sometimes seareely

visible. Aperture roundish, with oblique outer peri-

stome, not thiekened and not refieeted, interrupted

in the parietal portion. Umbilieus large, eorrespon-

ding to 3/10 of the maximum shell diameter. Perio-

straeum light brown in eolour, with weakly whitish

bands, more evident in not reeent shells.

Examined Material (Fig. 12). Sardinia: Rio

Abba Frida (left tributary of the Rio Melis, tributary

of the Rio San Giorgio), 3.5 km E-SE from Perda-

sdefogu along the eonneeting road to the SP 125,

470 m a.s.l. (Perdasdefogu, OG); UTM: 32S

NJ4191, S. Cianfanelli & E. Talenti leg. 23.05.2011

(44 shells from alluvial debris, MZUF GC/41424;

2 shells from alluvial debris, SCC; 2 shells from al-

luvial debris, ETC; 14 shells from litter, MZUF

GC/41786); G. Nardi & A. Braeeia leg. 12.04.2012

(2 shells from alluvial debris, GNC).

Campania: near Nerano, 200 m a.s.l. (Massa

Lubrense, NA); UTM: 33T VE4493, M. Bodon,

E. Bodon & S. Cianfanelli leg., 30.12.2012 (3

shells, SCC).

Calabria: Lao river, 100 m upstream of the

bridge in loeality Campieello, 290 m a.s.l. (Laino

Castello and Papasidero, CS); UTM: 33S WE7918,

M. Bodon & E. Bodon leg. 23.07.2005 (1 shell,

MZUF GC/41804; 3 shells, MBC).

The others speeies of molluses, eolleeted in the

sites deseribed above, are listed in Table 2.

Comparative Notes. All Italian speeimens are

here attributed to P. placida, even if the Sardinian

population shows a more eonieal shell (mean shell

height/shell diameter = 0.71), resembling to that of

P. angusta (Figs. 1-3; Table 1). Anyway, for the

maximum shell diameter (2.30) and for the large

umbilieus, also this population is here identified as

P placida.

Among other Valloniidae (Figs. 13-18), the

genus Vallonia differs from Plagyrona placida for

the more depressed shell and the enlarged and stron-

gly thiekened peristome. The most similar speeies

for the seulpture of the teleoeoneh, Vallonia costata,

is distinguished by the presenee of thieker ribs (pe-

riostraeal ribs) alternated with thinner ribs (radial

striae) (Figs. 13, 14; Giusti & Manganelli, 1986,

Fig. 7, Table 3; Gerber, 1996, Fig. 64a), while the

shell of Plagyrona is eovered by periostraeal ribs

all with the same thiekness (Figs. 6-11; Gittenber-

ger, 1989, Figs. 4, 5). Moreover, in V. costata, ra-

mified radial striae are present on the teleoeoneh,

while they are absent in Plagyrona.

Gittenbergia sororcula is very similar to V. co-

stata, but it shows a thin peristome, it does not have

the ramified radial striae on the teleoeoneh, al-

though a spiral and malleated miero- seulpture is

present on the protoeoneh (Figs. 15, 16; Giusti &

Manganelli, 1986, Fig. 6, Table 3); finally, it also

A new record for the Italian fauna: Plagyrona placida from Sardinia and S. Italy (Gastropoda Pulmonata Valloniidae) 53 1

Figures 1-5. Shells of Plagyrona placida. Figs. 1-3: speeimens eolleeted in the valley of the Rio Abba Frida (Perdasdefogu,

OG), Sardinia, S. Cianfanelli & E. Talenti leg. 23.05.2011, MZUF GC/41424, GC/41786. Figs. 4, 5: alluvial debris of the

Lao river, eolleeted in loeality Campieello (Laino Gastello and Papasidero, CS), Calabria, M. Bodon & E. Bodon leg.,

23.07.2005, MBC.

532

S. ClANFANELLI, G. NaRDI & M. BODON

Figures 6-11. Shells of Plagyrona placida photographed by seanning electron microscope (SEM). Fig. 6: apical view. Fig.

7: frontal view. Fig. 8: magnification of protoconch in lateral view. Figs. 9, 10: magnification of protoconch in oblique view.

Fig. 11: magnification of teleoconch in apical view. Figs. 6-8: specimen collected in the valley of the Rio Abba Frida), Sar-

dinia, S. Cianfanelli & E. Talenti leg. 23.05.2011, MZUF GC/41424. Figs. 9-11: specimen collected in locality Campicello,

alluvial debris of the Lao river, Calabria, M. Bodon & E. Bodon leg. 23.07.2005, MZUF GC/41804.

SITES

H/D

Abba Frida (Per-

dasdefogu, OG),

Sardinia

1.42 ±0.13

(1.20- 1.60)

1.99 ±0.21

(1.65 -2.30)

0.71 ±0.04

(0.67 - 0.79)

0.78 ±0.07

(0.65 - 0.85)

0.81 ±0.06

(0.70 - 0.90)

Campicello (Laino

Castello/Papasidero,

CS), Calabria

1.25

(1.20- 1.30)

1.90

(1.85 - 1.95)

0.66

(0.65 - 0.67)

0.73

(0.70 - 0.75)

0.73

(0.70 - 0.75)

Table 1. Shell size (mm) in Italian specimens of Plagyrona placida'. mean ± standard deviation and range (in parenthe-

sis). H = shell height; D = shell diameter; h = aperture height; d = aperture diameter; N = number of measured shells.

A new record for the Italian fauna: Plagyrona placida from Sardinia and S. Italy (Gastropoda PulmonataValloniidae) 533

differs from Plagyrona for the more depressed shell

and unequal ribs.

Acanthinula aculeata presents a higher shell

than Plagyrona, an expanded and slightly thickened

peristome and periostracal ribs with visible flexible

spines, placed in the middle part of each whorl, al-

ways alternated with thin radial striae and spiral

lines (Figs. 17, 18; Gittenberger, 1977, Figs. 3, 4;

Giusti & Manganelli, 1986, Table 3).

There are other species belonging to other fami-

lies, among those present in the Italian fauna, with

similar small size and depressed shells, resembling

to Plagyrona. The species of the genus Pyramidula

Fitzinger, 1833, like P. pusilla (Vallot, 1801) and/!

rupestris (Drapamaud, 1801), family Pyramiduli-

dae, are darker in colour (reddish-brown or purple),

with sculpture of shell surface less pronounced and

no spiral striae (Gittenberger & Bank, 1996, Figs.

5-15). Punctum pygmaeum (Drapamaud, 1801), fa-

mily Punctidae, differs in the smaller size and more

depressed spire (Cianfanelli, 2009, Fig. 90 B). Pa-

ralaoma servilis (Shuttleworth, 1 852), family Pun-

ctidae, perhaps the taxon showing the most similar

shell in colour, sculpture and size, differs from P.

placida because of the more depressed shape and

the less deep sutures, but especially because of the

unequal radial ribs, as the main periostracal ribs are

more spaced from each other (Figs. 19, 20; Giusti,

1973, Table 5).

Four of these entities have been found in Sardi-

nia and three in Calabria, in the same sites where P.

placida was collected (Table 2).

CONCLUSIONS

The three Italian sites, where Plagyrona placida

has been recently discovered, are the result of rese-

arch on the distribution of the malacological fauna,

not targeted, but extended to many Italian areas.

The records were made in sedimentary sub-

strate (leaf-litter and alluvial debris), where shells

only (without living animals) have been collected.

Although fresh material, therefore certainly coming

from habitats next to the collection sites, it is not

possible to establish with certainty the microhabitat

Figure 12. Distribution of Plagyrona placida in Europe and Northern Afriea: reeent populations known in the seientifie li-

terature (blaek dots); fossil populations known in the seientifie literature (white stars); reeent populations found in Italy (red

dots). A few data from seientifie literature need to be eonfirmed (see Holyoak & Holyoak, 2012).

534

S. ClANFANELLI, G. NaRDI & M. BODON

Figures 13-20. Other species similar to Plagyrona placida, in frontal view (left) and magnification of protoconch and first

whorls (right). Figs. 13, 14: Vallonia costata, Betania, M. Stabut (Tolmezzo, UD), Friuli-Venezia Giulia, 400 m a.s.l., UTM:

33T UM4741, S. Cianfanelli leg. 5.4.1988, MZUF GC/10553, SEM support MZ/1 76/1; Figs. 15, 16: Gittenbergia sororcula.

Fig. 15: at the crossroads between Rif Fasanelli, Rif Colleruggio and Rif De Gasperi, Pollino Mountains (Rotonda, PZ),

Basilicata, 1650 m a.s.l., UTM: 33S WE9518, S. Cianfanelli, E. Talenti & R. Martignoni leg. 28.10.1993, MZUF GC/6131,

SEM support MZ/176/3. Fig. 16: Val di Luce (Abetone, PT), Tuscany, 1590 m a.s.l., UTM: 32T PP3087, S. Cianfanelli &

E. Lori leg. 24.9.2008, MZUF GC/26585, SEM support MZ/245/4. Figs. 17, \S: Acanthimila aculeata, Passo Porrai, Monte

Cucco (Costacciaro, PG), Umbria, 910 m a.s.l., UTM: 33T UJ1805, S. Cianfanelli & M. Calcagno leg. 2.2.1992, MZUF

GC/2635, SEM support MZ/176/4. Figs. 19, 20: Paralaoma servilis, Ombrone river, S. Pantaleo (Pistoia), Tuscany, 65 m

a.s.l., UTM: 32T PP5264, S. Cianfanelli & E. Lori leg. 22.2.2007, MZUF GC/24152, SEM support MZ/245/2-3.

A new record for the Italian fauna: Plagyrona placida from Sardinia and S. Italy (Gastropoda PulmonataValloniidae) 535

Family

Species

Abba Frida,

Sardinia.

Alluvional

debris

Abba Frida,

Sardinia.

Litter

Nerano,

Campania.

Litter

Campicello,

Calabria.

Alluvional

debris

Cochlostomatidae

Cochlostoma spp.

Cochlostomatidae

Cochlostoma sardoum (Westerlund, 1890)

Aciculidae

Acicula lineolata banki Boeters, Gittenberger

et Subai, 1989

Aciculidae

Platyla similis (Reinhardt, 1880)

Pomatiidae

Pomatias elegans (Muller, 1774)

Carychiidae

Carychium biondii Paulucci, 1882

Carychiidae

Carychium tridentatum (Risso, 1 826)

Pyramidulidae

Pyramidula rupestris (Drapamaud, 1801)

Vertiginidae

Vertigo antivertigo (Drapamaud, 1801)

Vertiginidae

Vertigo pygmaea (Drapamaud, 1801)

Vertiginidae

Vertigo angustior Jeffreys, 1830

Vertiginidae

Columella aspera Walden, 1966

Vertiginidae

Columella edentula (Drapamaud, 1805)

Vertiginidae

Truneatellina callieratis (Scacchi, 1883)

Vertiginidae

Truneatellina eylindriea (Femssac, 1807)

Orculidae

Sphyradium doliolum (Bmguiere, 1792)

Oreulidae

Pagodulina pagodula (Des Moulins, 1830)

Chondrinidae

Granopupa granum (Drapamaud, 1801)

Chondrinidae

Rupestrella philippii (Cantraine, 1840)

Lauriidae

Lauria eylindraeea (Da Costa, 1778)

Lauriidae

Lauria sempronii (Charpentier, 1837)

Argnidae

Argna biplieata biplicata (Miehaud, 1831)

Argnidae

Agardhiella truncatella (Pfeiffer, 1841)

Valloniidae

Vallonia pulehella (Muller, 1774)

Valloniidae

Aeanthinula aculeata (Muller, 1774)

Valloniidae

Plagyrona plaeida (Schuttleworth, 1852)

Enidae

Merdigera obseura (Muller, 1774)

Punctidae

Punetum pygmaeum (Drapamaud, 1801)

Punctidae

Paralaoma servilis (Shuttleworth, 1852)

Helicodiscidae

Lucilla scintilla (Lowe, 1852)

Helicodiseidae

Lucilla singleyana (Pilsbry, 1 890)

Discidae

Discus rotundatus rotundatus (Muller, 1774)

Zonitidae

Vitrea contracta (Westerlund, 1871)

Zonitidae

Vitrea subrimata (Reinhardt, 1871)

Zonitidae

Retinella olivetorum icterica (Tiberi, 1872)

Zonitidae

Oxychilus draparnaudi (Beck, 1837)

Zonitidae

Oxychilus oppressus (Shuttleworth, 1878)

Zonitidae

Mediterranea hydatina (Rossmassler, 1838)

536

S. ClANFANELLI, G. NaRDI & M. BODON

Family

Species

Abba Frida,

Sardinia.

Alluvional

debris

Abba Frida,

Sardinia.

Litter

Nerano,

Campania.

Litter

Campicello,

Calabria.

Alluvional

debris

Zonitidae

Daudebardia brevipes (Drapamaud, 1805)

Zonitidae

Daudebardia rufa (Drapamaud, 1805)

Euconulidae

Euconulus fulvus (Muller, 1774)

Femssaciidae

Cecilioides acicula (Muller, 1774)

Femssaciidae

Cecilioides petitiana (Benoit, 1862)

Femssaciidae

Cecilioides sp.

Suhulinidae

Rumina decollata (Linnaeus, 1758)

Oleacinidae

Poiretia dilatata dilatata (Philippi, 1836)

Testacellidae

Testacella scutulum Sowerhy, 1820

Clausiliidae

Medora dalmatina poUinensis Nordsieck, 2012

Clausiliidae

Cochlodina kuesteri (Rossmassler, 1836)

Clausiliidae

Siciliaria paestana paestana (Philippi, 1836)

Clausiliidae

Papillifera papilaris papillaris (Muller, 1774)

Cochlicellidae

Cochlicella acuta (Muller, 1774)

Cochlicellidae

Cochlicella barbara (Linnaeus, 1758)

Hygromiidae

Xerotricha conspurcata (Drapamaud, 1801)

Hygromiidae

Candidula cavannae (Paulucci, 1881)

Hygromiidae

Hygromia cinctella (Drapamaud, 1801)

Hygromiidae

Ichnusotricha berninii Giusti et Manganelli, 1987

Hygromiidae

Cernuella cisalpina (Rossmassler, 1837)

Hygromiidae

Cernuella virgata (Da Costa, 1778)

Hygromiidae

Xerosecta dohrni (Paulucci, 1882)

Hygromiidae

Trochoidea pyramidata (Drapamaud, 1805)

Hygromiidae

Monacha parumcincta (Menke, 1828)

Helicodontidae

Helicodonta obvoluta (Muller, 1774)

Helicidae

Chilostoma planospira setulosum (Briganti, 1 825)

Helicidae

Marmorana serpentina (Femssac, 1821)

Helicidae

Marmorana fuscolabiata (Rossmassler, 1842)

Helicidae

Cantareus apertus (Bom, 1778)

Helicidae

Cornu aspersum aspersum (Muller, 1774)

Table 2. Species of molluscs collected with Plagyrona placida, in the three Italian sites.

where P. placida lives in the Italian sites; however,

it might be represented by roeky habitats or tree

trunks and branehes eovered by mosses, in Medi-

terranean maquis, in oak forests (Quercus ilex L.)

or in mesophilie deeiduous forests.

The environments seem to be intaet, espeeially

in Campania (a Mediterranean maquis of ilexes

with ealeareous eliff) and in Calabria, the Lao

River Valley (a proteeted area established as a Na-

ture Reserve, Ministerial Deeree 423 dated

21 .07. 1987), whieh ineludes one of the most intaet

waterways whieh have the greater eeologieal im-

portanee for all Southern Italy. Moreover, the as-

soeiated malaeofauna (Table 2) is eharaeterized by

A new record for the Italian fauna: Plagyrona placida from Sardinia and S. Italy (Gastropoda PulmonataValloniidae) 537

the presence of typical litter entities, not very com-

mon or requiring a well preserved habitat, such as

Columella aspera Walden, 1966 (reported in Italy,

until now, only from Elba Island; Manganelli et al.,

1995), Pagodulina pagodula (Des Moulins, 1830),

Lauria sempronii (Charpentier, 1837), Agardhiella

truncatella (Pfeiffer, 1841) (it has never been repor-

ted for Southern Italy), Daudebardia brevipes (Dra-

pamaud, 1805) andZ). rufa (Drapamaud, 1805), or

endemic species such as Cochlostoma sardoum

(Westerlund, 1890), Carychium biondii Paulucci,

1882, Oxychilus oppressus (Shuttleworth, 1878),

Cochlodina kuesteri (Rossmassler, 1836), Siciliaria

paestana (Philippi, 1836), Candidula cavannae

(Paulucci, 1881), Ichnusotricha berninii Giusti et

Manganelli, 1987, Xerosecta dohrni (Paulucci,

1882), Chilostoma planospira (Lamarck, 1822) and

Marmorana fuscolabiata (Rossmassler, 1842).

All this suggests that P. placida could be native;

but the presence of non-native species, such as Pa-

ralaoma servilis, Lucilla scintilla (Lowe, 1852) and

L. singleyana (Pilsbry, 1890), even if only few spe-

cimens were collected and only in alluvial debris

(never in litter), still leaves some doubt about the

fact that P placida may actually be native in the Ita-

lian sites where it was found. Furthermore, it should

be noted that, while P. placida is a rare species, so

that the animal is still unknown and living speci-

mens have never been collected in Italy, small non-

native species of litter and soil such as P servilis,

L. scintilla and L. singleyana, once introduced, be-

come abundant species, often dominant compared

to the native species, they can reach very high den-

sities and they can often spread quickly and widely

in their environments, so they are now present in

almost all European countries and, in Italy, in most

regions (Bodon et al., 2004; Cianfanelli, 2009;

Bank, 2011a; Cianfanelli & Bodon, in press).

P placida is a new taxon that must be added to

the checklist of terrestrial Mollusca of the Italian

fauna (Bodon et al., 1995; Manganelli et al., 1995);

since 1995 such list has undergone several incre-

ments (Eikenboom, 1996; Giovannelli, 1996; Man-

ganelli & Favilli, 1996; Manganelli at al., 1997;

Giusti & Manganelli, 1998; Riedel, 1998; Cianfa-

nelli et al., 2000; Colla et al., 2000; Manganelli et

al., 2000; Beckmann, 2002; Carr, 2002; Colla &

Stoch, 2002; Falkner et al., 2002; Gerber, 2002;

Beckmann & Falkner, 2003; De Mattia, 2003; Zal-

lot, 2003; Beckmann, 2004; Bodon et al., 2004;

Cianfanelli et al., 2004; De Mattia, 2005; Ferreri et

al., 2005; De Mattia & Prodan, 2006; Garominy &

Ripken, 2006; Gittenberger & Eikenboom, 2006;

Lo Brano & Sparacio, 2006; Nordsieck, 2006; De

Mattia, 2007; Nordsieck 2007a, 2007b; Reitano et

al., 2007; Beckmann & Falkner, 2008; Bodon &

Cianfanelli, 2008; Falkner, 2008; Falkner & Nie-

derhofer, 2008; Gavetti et al., 2008; Falkner & von

Proschwitz, 2009; Nardi, 2009; Nitz et al., 2009;

Reitano et al., 2009; Bodon et al., 2010; Feher et.

al., 2010; Halgass & Vannozzi, 2010; Kokshoom &

Gittenberger, 2010; Liberto et al., 2010; Manganelli

et al., 2010; Nitz et al., 2010; Pfenninger et al.,

2010; Bank, 2011b; Colomba et al., 2011; De Mat-

tia et al., 2011; Nardi, 2011; Nardi & Bodon, 2011;

Nordsieck, 2011a, 2011b, 2011c; Nordsieck, 2012;

Reise et al., 2011; Colomba et al., 2012; Liberto et

al., 2012; Niero et al., 2012; Evangelista et al., in

press). Moreover, the current list needs to be revised

and confirmed following the more modem methods

of investigation.

ACKNOWLEDGEMENTS

We thank Antonio Braccia, Emanuele Bodon,

Micaela Calcagno, Elisabetta Lori and Enrico Ta-

lenti for their help during the research; Maurizio

Ulivi for technical assistance; Giuseppe Manganelli

for literature.

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Biodiversity Journal, 2012, 3 (4): 543-554

Terrestrial gastropods of the minor islets of the Maltese Ar

chipelago (Mollusca Gastropoda)

David R Cilia'*, Arnold Sciberras^ Jeffre)^ Sciberras^ & Luca Pisani'^

'29, Triq il-Palazz l-Ahmar, Santa Venera, Malta

^133, Amest, Areade Street, Paola, Malta

^24, Camilleri Court 5, Triq il-Marlozz, Mellieha (Ghadira), Malta

"^9, Milner Street, Sliema, Malta

*Corresponding author: dpeilia@gmail.eom

ABSTRACT For this study, the terrestrial malaeofauna of minor islets of the Maltese arehipelago was inve-

stigated. A number of new reeords were found and synthesized with previous reeords to produee

a comprehensive list of species. A brief commentary on the population, environment, habitat,

and morphology for most species is given.

KEY WORDS Maltese islands; insularity; Gastropoda; systematics; taxonomy; new records.

Received 12.05.2012; accepted 20.10.2012; printed 30.12.2012

Proceedings of the P' International Congress “Insularity and Biodiversity”, May 11*-13*, 2012 - Palermo (Italy)

INTRODUCTION

The Maltese arehipelago eonsists of three

large islands, Malta, Ghawdex (Gozo) and Kem-

muna (Comino, and a number of smaller islets

and roeks surrounding them. For this study, Kem-

muna and most of the islets supporting maero-

phytes were investigated for terrestrial molluses.

From east to west, these islands are: Skoll tal-

Barbaganni, Haifa Roek, Tac-(2awl Roek, Kem-

munett (Cominotto), Kemmuna (Comino), Small

Blue Lagoon Roek, Large Blue Lagoon Roek, Pi-

geon Roek, Selmunett (St. Paul’s Islands), and

Ta’ Fra Ben (Figs. 1-12).

The terrestrial gastropod speeies riehness on

these islands varies depending on their dimen-

sions and relative exposure. The molluses on na-

tural island reserves of Filfla and General’s Roek

have been subjeets of intensive studies in the past

(e.g. Soos, 1933; Holyoak, 1986; Beekmann,

1987, 1992; Thake & Sehembri, 1989) and are not

investigated here.

MATERIALS AND METHODS

In the eourse of about fourteen years (1998-

2012), the authors eolleeted terrestrial molluses

from the investigated islands mainly as part of a

survey dealing with vegetation, the results of whieh

ean be seen in Seiberras & Seiberras (2010). The

majority of sites are inaeeessible, and visits to these

loeations were mainly aehieved by swimming and

elimbing. Snail speeimens and soil sampled from

the sites were sealed in waterproof plastie bags and

labelled; these were later eleaned, identified and

studied.

Speeimens were also reeovered from stored

soil samples and from previous surveys in order to

assemble a eoneise-as-possible list of reeords. Li-

terature on the terrestrial molluses of the Maltese

islands, with partieular attention to Sehembri

(1983) and Giusti et al. (1995), was eonsulted in

the proeess. Supraspeeifie elassifieation follows

Bouehet & Roeroi (2005) and Kokshoom & Git-

tenberger (2010).

544

D.P. Cilia, A. Sciberras J. Sciberras & L. Pisani

Study area: the islands

To allow for a more readable text, authorities for

molluse speeies were omitted in this seetion. The

eomplete nomenelature is available in the seetion

dealing with systematies.

Shall tal-Barbaganni (36°1'39"N, 14°19'36"E)

(Fig. 1). Sparsely populated by Inula chrithmoides

L., this is the smallest member of the Gozitan ar-

ehipelago, and is frequently eovered by sea spray

in rough weather. Only one terrestrial molluse, Fe-

russacia folliculum, was reeovered fi*om this island.

Haifa Rock 14°19'52"E) (Fig. 2).

The seeond largest island of the Gozitan group,

Haifa Roek supports about 20 speeies of maerophy-

tes on a sparse terra rossa soil eover.

Tac-Cawl Rock (36°01'33"N, 14°18'58"E)

(Fig. 3). Tac-Cawl Rock is not an island in the ac-

curate sense of the word, since a narrow Upper Co-

ralline Fimestone isthmus connects it to the Gozitan

mainland. Due to this, it is hypothesized that indi-

viduals in the terrestrial mollusc populations present

on it are subject to occasional migration and genetic

mixing with the mainland population. Nevertheless,

the isthmus is frequently inundated by waves or

tidal fluctuations, especially during the winter

months. The rock supports about 20 species of ma-

crophytes.

Kemmunett (Cominotto) (36°00'49"N,

14°19T3"E) (Fig. 5). The largest island of the Kem-

muna group (bar Kemmuna itself), Kemmunett

hosts close to 50 macrophyte species in the large

variety of microhabitats, including cliffs, disturbed

ground, garigue and steppe, concentrated within its

small surface area.

Small Blue Lagoon Rock (36°00'40"N,

14°19'25"E) (Fig. 6). This sloping landmass featu-

res a low species richness, both in macrophytic and

in terrestrial molluscan species. A few patches of

terra rossa support the sparse clumps of vegetation

present.

Large Blue Lagoon Rock (36°00'39"N,

14°19'31"E) (Fig. 7). Fike the island preceding it,

this rock is a steep slope with vegetation predomi-

nant on its upper half, anchored mainly in karstic

terra rossa pockets. The geology of the two islands

and the vegetation present are similar, though the

molluscan species richness here is much more pro-

nounced.

Pigeon Rock 14°19'45"E) (Fig. 8).

This is a very steep, high and relatively inaccessible

islet with an interesting macrophytic community,

hosting three endemic taxa. It is also interesting as

regards its molluscan representatives - it is the only

island in the Kemmuna archipelago, and indeed out

of all islands investigated, to host a population of

the endemic calciphile Murella melitensis.

Selmunett (35°57'55"N, 14°24'03"E) (Fig. 10).

Also known as St. Paul’s Islands, Selmunett is a

group of two connected landmasses that are sepa-

rated by the sea during rough weather. Boasting a

high macrophytic diversity (about 100 species), it

is expected that the molluscan species richness re-

flects this; indeed, the 12 species of gastropods re-

corded here reveal that this landmass has the

highest diversity out of all the islands investigated

(with the exception of Kemmuna and Kemmunett,

the former of which is much larger). In an edition

of the journal Potamon devoted to the natural hi-

story of the same islands, Schembri (1983) men-

tions 7 species of landsnails: Theba pisana,

Eobania vermiculata. Helix aspersa, Pomatias sul-

catus melitense, Rumina decollata, Lampedusa sy-

racusana, Trochoidea ealcarata and Troehoidea

schembri. These names are herein reproduced as

they appear in the original paper; current taxonomy

clumps the latter two names into one species. The

finds of Rumina decollata and Theba pisana were

not replicated for the present study, and they are

also listed as absent from the islands in Giusti et al.

(1995). On the other hand, the records of two spe-

cies of Cernuella, Oxychilus draparnaudi and Pa-

pillifera bidens constitute new records. The terrain

of the islands is varied and alkaline terra rossa soil

and xerorendzina soils characterize most of the hi-

gher parts (Savona Ventura, 1983, Farrugia Randon,

2006).

Ta^ Fra Ben (35°57'35"N, 14°25'43"E) (Fig.

11). This landmass is situated off Qawra in north-

western Malta and is connected by a very thin

isthmus. Molluscan species diversity is low, corre-

Terrestrial gastropods of the minor islets of the Maltese Archipelago (Mollusca Gastropoda)

545

Figure 1. Skoll tal-Barbaganni as viewed from the south. Figure 2. Haifa Roek. Figure 3. Tac-Cawl Roek. Figure 4. Map

of the minor islets of Gozo: q = Haifa Rock; r = Skoll tal-Barhaganni; s = Tac-Cawl Rock. Figure 5. Kemmunett, as vie-

wed from Kemmuna. Figure 6. Small Blue Lagoon Rock as viewed from the Kemmuna mainland.

546

D.P. Cilia, A. Sciberras J. Sciberras & L. Pisani

Comino

Figure 7. Large Blue Lagoon Rock. Figure 8. Pigeon Rock. Figure 9. Map of the minor islets of Comino: 1 = Pigeon Rock;

m = Large Blue Lagoon Rock; n = Small Blue Lagoon Rock; o = Kemmunett. Figure 10. Selmunett. Fig. 11. Ta’ Fra Ben. Fi-

gure 12. Map of the minor islets of Malta: i = Selmunett; h = Ta’ Fra Ben.

Terrestrial gastropods of the minor islets of the Maltese Archipelago (Mollusca Gastropoda)

547

spending to the poor vegetation eover eonsisting

mainly of halophytes, predominantly Arthrocne-

mum macros tachyum (Morie.) Moris.

Kemmuna (Comino). The third largest island

of the Maltese arehipelago, Kemmuna has the lar-

gest speeies riehness of all. Past studies and eollee-

tions have been helpful in determining the overall

faunal diversity of this island; still, a new reeord of

the very eommon Cantareus aspersus was noted

during the present researeh.

ABBPIEVIATIONS. Skoll tal-Barbaganni (BR);

Haifa Roek (HR); Tac-Cawl Roek (CR); Kemmu-

nett (Cominotto) (KT); Small Blue Lagoon Roek

(SBL); Large Blue Lagoon Roek (LBL); Pigeon

Roek (PR); Selmunett (St. Paul’s Islands) (ST); Ta’

Fra Ben (FB); Kemmuna (Comino) (KM); David

Cilia (DC); new reeord (nr).

RESULTS

The results of the present researeh, ineluding se-

veral new reeords, were synthesized with those in

Giusti et al. (1995) to formulate a grid with all the

observations (Table 1). The number of speeies on

eaeh island was then eompared to the number of

maerophytes on the same island aeeording to Borg

(1927), Camilleri (1990), Lanfraneo (2002), and

Seiberras & Seiberras (2009, 2010) (Fig. 13). These

results are elaborated upon in the ‘Systematies’ see-

tion below.

SYSTEMATICS

Clade Littorinimorpha Golikov & Starobogatov, 1975

Family Pomatiidae Newton, 1891 (1828)

Subfamily Pomatiinae Newton, 1891 (1828)

Tudorella VischQV, 1885

Tudorella melitense (Sowerby, 1843)

Present reeords: HR, CR (nr), KT (nr), SBL (nr),

LBL (nr), PR (nr), ST, KM

Remarks: This opereulate speeies is ubiquitous.

The speeifie status of whieh is based on Pfenninger

et al. (2007), was found on most of the islands in-

vestigated. It demonstrates a toleranee to hypersa-

line eonditions, as its presenee elose to the first

oeeurrenees of Inula chrithmoides L. shows. Its

absenee on ST, where suitable habitats are availa-

ble, remains unexplained, though it is present on

the opposite headland of Mistra on mainland

Malta. Speeimens of Clibanarius sp. hermit erabs

bearing T. melitense shells were seen around KM

eoasts, elose to the Ghemieri peninsula and Santa

Marija Bay.

Clade Stylommatophora A. Sehmidt, 1855

Family Chondrinidae Steenberg, 1925

Subfam. Granariinae Kokshoom & Gittenberger, 2010

Granopupa Bottger, 1889

Granopupa granum (Drapamaud, 1801)

Present reeords: KT (nr), ST, KM

Remarks: The presenee of this eommon gastro-

pod on ST and KM eonfirms earlier reeords by Giu-

sti et al. (1995).

Family Enidae Woodward, 1903 (1880)

Subfamily Eninae Woodward, 1903 (1880)

Tribe Chondrulini Wenz, 1923

Mastus Beek, 1837

Mastus pupa (Linne, 1758)

Present reeords: KT (nr), LBL (nr), ST, KM

Remarks: This gastropod was generally found

at the base of grass tufts and in assoeiated loosely

aggregated soils, mostly as single speeimens. The

KT, LBL and KM speeimens are more slender than

mainland populations in Malta and Gozo.

Family Ferussaeiidae Bourguignat, 1883

Ferussaeia Risso, 1 826

Ferussacia (s. sir) folliculum (Sehroter, 1784)

Present reeords: BR (nr), KM

Remarks: Sehroter’s 1784 publieation, preee-

ding Gmelin’s 1791 work (the souree for whieh this

speeies’ deseription is often mistakenly attributed),

lists this speeies as F folliculum, whieh is the eor-

reet spelling as opposed to F. folliculus. Therefore,

Welter-Sehultes’ opinion regarding nomenelature

548

D.P. Cilia, A. Sciberras J. Sciberras & L. Pisani

(2012) is followed here. Surprisingly, this was the

only terrestrial gastropod species recovered from

BR, where it is found in relatively high population

densities beneath leaf debris surrounding Inula

chrithmoides stems, incidentally also the only plant

which can survive on this rock (Sciberras & Sciber-

ras, 2010). The hardiness of F. folliculum is partly

attributable to the hardy epiphragm formed during

adverse conditions.

Cecilioides VQXussdiC, 1814

Cecilioides acicula (Muller, 1774)

Present records: none.

Remarks: Reported for ST by Giusti et al.

(1995), this subterranean snail was not encountered

during the present research.

Family Subulinidae Fischer & Crosse, 1877

Subfamily Rumininae Wenz, 1923

Rumina Risso, 1 826

Rumina decollata (Linne, 1758)

Present records: HR, CR (nr), KT (nr), LBL (nr),

PR (nr)

Remarks: Individuals of this omnivorous spe-

cies in various stages of growth were found on se-

veral of the islands investigated. Schembri (1983)

mentions this species for ST, but this find was not

repeated during the present study. Dead slender

shells reminding one of R. saharica Pallary, 1901

were occasionally recovered in sympatry with the

present species, but the absence of live specimens

and the intrapopulation variability of R. decollata

shells makes a case for the presence of R. saharica

highly hypothetical. On the other hand, R. saharica

was recently discovered on other circum-Sicilian is-

lands (Liberto et al., 2012).

Family Clausiliidae Gray, 1855

Subfamily Alopiinae Wagner, 1913

Tribe Medorini Nordsieck, 1997

MuticarialAXY^holm, 1925

Muticaria macrostoma (Cantraine, 1835)

Present records: HR, CR*, KT, LBL*, PR*, KM

Remarks: Several individuals of this species

were found in crevices and overhangs across the

investigated islands. Intermediates of some of the

four subspecies mentioned by some authors (e.g.

Nordsieck, 2007; Colomba et al., 2010) exist in se-

veral cases, in fact, some of the investigated speci-

mens correspond to the nominate form (e.g. ST),

others are intermediate (e.g. PR) while others are

closer to the form oscitans Charpentier, 1852 (e.g.

CR, HR). In the light of this research, unpublished

records (DC) and other publications that discuss

this situation (Giusti et al., 1995), M. macrostoma

is herein recognized as a species complex referable

to one taxon until more molecular data becomes

available.

Tribe Delimini Brandt, 1956

Papillifera Hartmann, 1842

Papillifera bidens affinis (Philippi, 1836)

Present records: HR, CR (nr), KT (nr), LBL (nr),

ST (nr), KM

Remarks: This clausiliid was found on lime-

stone substrates and at the bases of tufts of grass.

The absence of this mollusc on ST from previous

surveys is remarkable, as where present, the snail

occurs in sizeable populations; however, Selmunett

remains under considerable anthropogenic in-

fluence, and the presence of Papillifera Hartmann,

1 842 (which, unlike other clausiliids, is highly sub-

ject to passive dispersal by man (Nordsieck, unda-

ted)) could be a recently induced phenomenon.

Family Oxychilidae Hesse, 1927 (1879)

Subfamily Oxychilinae Hesse, 1927 (1879)

Oxychilus Fitzinger, 1833

Oxychilus (s. str.) draparnaudi (Beck, 1837)

Present records: ST (nr), KM

Remarks: Several individuals of O. draparnaudi

were found beneath stones and in soil on ST. Some

of the shells recovered contained unidentified cole-

opteran larvae.

Family Pristilomatidae Cockerell, 1891

DYrea Fitzinger, 1833

Terrestrial gastropods of the minor islets of the Maltese Archipelago (Mollusca Gastropoda)

549

SBL

LBL

Cantareus apertus

Cantareus aspersus

Caracollina lenticula

Cecilioides acicula

Cernuella caruanae

Cernuella cisalpina

Cochlicella acuta

Eobania vermiculata

Ferussacia folliculum

Granopupa granum

Mastus pupa

Murella melitensis

Muticaria macrostoma

Oxychilus draparnaudi

Papillifera bidens

Rumina decollata

Sphincterochila candidissima

Theba pisana

Trochoidea spratti

Tudorella melitense

Vitrea sp.

Xerotricha conspurcata

Number of species

Table 1: Terrestrial molluse speeies on the smaller islands of the Maltese arehipelago. The asterisks (*) denote a presenee,

aeeording to a synthesis of the present study and that by Giusti et al. (1995).

550

D.P. Cilia, A. Sciberras J. Sciberras & L. Pisani

Vitrea sp. sensu Giusti et al., 1995

Present reeords: none

Remarks: This undeseribed speeies, native to

Malta and the Aeolian islands, was reported for KM

and ST by Giusti et al. (1995), although it was not

eneountered during the present researeh on either

of the islands.

Family Sphineteroehilidae Zileh, 1960 (1910)

Subfamily Sphineteroehilinae Zileh, 1960 (1910)

Sphincterochila AncQy, 1887

Albea Vallary, 1910

Sphincterochila (Albea) candidissima

(Drapamaud, 1801)

Present reeords: KT (nr), KM

Remarks: S. candidissima was found on both

islands but at very low densities on the former,

generally inhabiting terra rossa soil on garigue

with eonsiderable exposure to the sun. The shade

of shrubs of Thymbra capitata (L.), Capparis

orientalis Veil!., Euphorbia melitensis Pari, and Pi-

stacia lentiscus (L.) offer temporary respite to a

number of speeimens.

Family Trissexodontidae Nordsieek, 1987

Caracollina Beek, 1837

Caracollina (s. sir.) lenticula (Miehaud, 1831)

Present reeords: HR, CR (nr), KT, KM, ST

Remarks: The presenee of C. lenticula on CR is

eonfirmed by two bleaehed shells found under a roek.

Family Coehlieellidae Sehileyko, 1972

CochlicellaFemsssLC, 1821

Cochlicella (s. str.) acuta (Muller, 1774)

Present reeords: KT (nr), SBL (nr), PR (nr), KM

Remarks: Several individuals of this speeies

were found in the eentral part of KM. Two fresh

speeimens were obtained from KT and SBL eaeh.

Family Hygromiidae Tryon, 1866

Subfamily Geomitrinae Bottger, 1 909

Tribe Troehoideini Nordsieek, 1987

Trochoidea Brown, 1 827

Trochoidea (s. six)spratti (Pfeiffer, 1846)

Present reeords: HR, CR (nr), KT, SBL (nr),

LBL (nr), PR (nr), FB (nr), KM, ST

Remarks: This snail eomes in a variety of eon-

ehologieal forms that have formed the basis of se-

veral deseriptions (e.g. Treehmann, 1938;

Beekmann, 1987). The speeimens from HR are

wide as they are tall, eorresponding to the nomino-

typieal form, those from SBL and LBL are flatter

and more globose, eorresponding to the form

schembrii Pfeiffer, 1 848, while the speeimens from

ST laek the evident ribbing observable in the for-

mer populations, somewhat similar to the form cal-

carata Benoit, 1860. Where found, the speeies

generally oeeurs in eonsiderable quantities.

Subfamily Hygromiinae Tryon, 1866

Tribe Helieellini Ihering, 1 909

Xerotricha Monterosato, 1 892

Xerotricha conspurcata (Drapamaud, 1801)

Present reeords: none

Remarks: Reported for KM by Giusti et al.

(1995), this speeies was not eneountered during the

present researeh.

Tribe Hygromiini Tryon, 1866

Cernuella Sehliiter, 1838

Cernuella (s. str.) cisalpina (Rossmassler, 1837)

Present reeords: ST (nr)

Remarks: The single speeimen found on ST is

eharaeteristieally whitish with a pinkish lip, similar

to individuals in populations on mainland Malta.

Xeroamanda Monterosato, 1 892

Cernuella (Xeroamanda) caruanae (Kobelt,

1882)

Present reeords: ST (nr), KM

Remarks: The single dead shell found on ST

eonstitutes a new reeord of this supposedly Maltese

endemism for the island.

Terrestrial gastropods of the minor islets of the Maltese Archipelago (Mollusca Gastropoda)

551

Figure 13. Tudorella melitense, h: 12.6 mm, 8.3 mm; Large Blue Lagoon Roek, Comino. Figure 14. Mastus pupa, h: 12.7

mm, D: 5,3 mm; Large Blue Lagoon Roek, Comino. Figure 15. Muticaria macrostoma, h: 12 mm, D: 3.6 mm; Haifa Roek,

Gozo. Figure 16. Muticaria macrostoma, h: 12.8 mm, D: 3.8 mm; Selmunett, Malta. Figure 17. Trochoidea spratti, h: 5.3

mm, D: 7,1 mm; Large Blue Lagoon Roek, Comino. Figure 18. Trochoidea spratti, h: 4.9 mm, D: 6,7 mm; Small Blue La-

goon Roek, Comino. Figure 19. Murella melitensis, h: 11.3 mm, D: 17,1 mm; Pigeon Rook, Comino.

552

D.P. Cilia, A. Sciberras J. Sciberras & L. Pisani

Family Helicidae Rafinesque, 1815

Subfamily Helicinae Rafinesque, 1815

Tribe Murellini Hesse, 1918

Pfeiffer, 1877

Murella melitensis (Ferussae, 1821)

Present reeords: PR (nr)

Remarks: This speeies was until reeently elas-

sified undQV Marmorana Hartmann, 1844. Fioren-

tino et al. (2010) separate the two genera on the

basis of moleeular data and love-dart morphology.

The three individuals of this highly variable speeies

found on PR show very faint ribbing whieh is ab-

sent from other populations in Malta and Gozo, to-

gether with the highly eharaeteristie brown

mottling on a white baekground and a brown spot

at the eonjunetion of the peristome and the eolu-

mella. The absenee of the speeies from the KM

mainland and the surrounding isles may indieate

that this represents a reliet population.

Tribe Thebini Wenz, 1923

Theba Risso, 1826

Theba pisana pisana (Muller, 1774)

Present reeords: HR (nr), CR (nr), KT (nr), KM,

FB (nr)

Remarks: FB supports a very dense population

of this speeies, generally gathered on or beneath the

base of Arthrocnemum macrostachyum (Morie.)

Moris, shrubs. Sehembri (1983) mentions this spe-

eies for ST, but this find was not repeated during

the present study.

Tribe Helieini Rafinesque, 1815

EobaniaYiQSSQ, 1913

Eobania vermiculata (Muller, 1774)

Present reeords: HR, KT (nr), LBL (nr), PR (nr),

FB (nr), ST, KM

BR HR CR KT SBLLBL PR ST FB

■ Molluscs

n Plants

Figure 20. A graphical representation of the correlation between the number of terrestrial mollusc and macrophyte

species (except Kemmuna, Comino).

Terrestrial gastropods of the minor islets of the Maltese Archipelago (Mollusca Gastropoda)

553

Remarks: On ST, this is the eommonest snail

speeies. On the other hand, no live speeimens were

found on LBL, with the reeord being based on three

subfossil shells.

Cantareus Risso, 1826

Cantareus apertus (Bom, 1778)

Present reeords: KT (nr), SBL (nr), KM

Remarks: Single individuals of this speeies were

eneountered in moist grass or aestivating elose to

the soil. The speeies seems to prefer more open ha-

bitats than its eongener, mentioned below.

Cantareus aspersus (Muller, 1774)

Present reeords: KT (nr), SBL (nr), KM (nr)

Remarks: The presenee of the speeies on SBL is

inferred from several fragments from adult shells.

The population of KT eonsists of individuals thie-

ker and taller than mainland shells.

The plaeement of Helix aspersa Muller, 1774

into the genus Cantareus here follows the moleeular

analyses of Koene & Sehulenburg (2005), Manga-

nelli et al. (2005), and Wade et al. (2006; 2007),

where phylogenetie trees support a plaeement of the

taxon together with C. apertus, Eobania and Otala

Sehumaeher, 1817 in a monophyletie elade. These

results support the anatomieal data elaborated upon

by Giusti et al. (1995).

CONCLUSIONS

A number of 22 species of terrestrial molluscs,

all of which occur on mainland Malta and most of

which occur also in Gozo, were recorded from the

ten islands investigated. The species richness of

non-marine molluscs on each of the islands is cor-

related to the island’s size, variety of habitats, and

floral diversity (Fig. 20).

It is interesting to note that although the area of

most of the islands is relatively small, biodiversity

and abundance of individuals is generally high; ho-

wever, this does not mean that the islands (even

close ones, such as those in the Kemmuna group)

have a similar ecosystem, as can be seen from the

species recovered on each. Present and future work

on the distribution and abundance of these molluscs

in each particular case may be used to observe

trends and assess threats to conservation, especially

in the cases of endemic or rare species.

ACKNOWLEDGEMENTS

The authors would like to thank Diane Portelli,

Romario Sciberras, and Esther Sciberras for various

forms of assistance on the field and outside of it and

also Salvatore Giglio for the photographs of the

molluscs and anonymous reviewers for their sug-

gestions.

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Biodiversity Journal, 2012, 3 (4): 555-570

Terrestrial molluscs from the R.N.I. ‘‘Grotta Conza” (Pa-

lermo, Sicily) (Gastropoda Architaenioglossa Pulmonata)

Agatino Reitano'*, Fabio Liberto^ Salvatore Giglio^ Rosario Grasso'^ & Maria Teresa Spena'^

'Via Gravina, 77 - 1-95030 Tremestieri Etneo, Catania, Italy; e-mail: tinohawk@yahoo.it

^Strada Provinciate Cefalu - Gibilmanna, 93 - 1-90015 Cefalu, Palermo, Italy; e-mail: fabioliberto@alice.it

^Contrada Settefrati, 1-90015 Cefalu, Palermo, Italy; e-mail: hallucigenia@tiscali.it

''Dipartimento di Scienze Biologiche, Geologiche ed Ambientali, Universita degli Studi di Catania, Via Androne, 8 1 - 1-95 124 Catania,

Italy; e-mail: rosagra@unict.it, marisaspena@hotmail.com

*Corresponding author

ABSTRACT The results of a study on the fauna of terrestrial molluscs from the R.N.I. "Grotta Conza" (Pa-

lermo, North-Western Sicily) are here described. Research has allowed us to compile a chec-

klist of 41 species, 18 of which are endemic to Sicily. For each species ecological and

distributional data are provided.

KEY WORDS Terrestrial molluscs; North-Western Sicily; endemic species; biodiversity.

Received 12.05.2012; accepted 21.10.2012; printed 30.12.2012

Proceedings of the P' International Congress “Insularity and Biodiversity”, May 11*-13*, 2012 - Palermo (Italy)

INTRODUCTION

The Integral Natural Reserve, R.N.I. "Grotta

Conza" is eharaeterized by the presenee of a karst

eave of great speleologieal, paleontologieal and bio-

logieal importanee and an area outside the eave

with rupieolous vegetation, Mediterranean maquis,

grassland and forests of native and non-native spe-

eies planted on lithosols (lithie xerorthents). The

knowledge about terrestrial molluse fauna in this

area is searee, and in the malaeologieal literature

only a few speeies are reported (Beekmann, 2004;

La Mantia & Rizzo, 2009). The researeh earried out

during the years 2007-2012, in the area of the R.N.I.

"Grotta Conza", allowed us to eompile the first

eheeklist of terrestrial molluses in the natural re-

serve and to observe the status of these speeies.

Study area

The Natural Reserve "Grotta Conza" (Palermo,

Sieily, eadastral number Si Pa 60, 38°11'13"N,

13°16'44"E) was established by D.A. Territorio e

Ambiente n°292/44 del 16/05/1995 (Suppl. Ord.

G.U.R.S. n.4, 20/01/1996) and is managed by Club

Alpino Italiano - Sieilia. It is ineluded in seetion

594040 (Tommaso Natale) of the Regional Teehni-

eal Map (seale 1 : 10.000). The reserve is loeated bet-

ween 100 and 220 m of altitude; it has an area of

12.34 heetares and is divided into a small “Zone A”,

where the entranee of the eave is loeated, and a

wider “Zone B”. The area is eharaeterized by a Me-

diterranean elimate, i.e., as “Csa” in the Koppen

(1936) elimate elassifieation, with hot, dry summers

and mild to eool wet winters. The eave opens at an

elevation of 175 m, and extends for a total length

of 100 m; sloping 30° upwards with a +30 m height

differenee; the inner part of the eave is eonneeted

to the external environment by small burrows (few

eentimeters in diameter). The eave is developed wi-

thin Upper Triassie limestone roek (Seiaeea Forma-

tion; see Italian Geologieal Map 1:50.000 leaf 594

Partinieo). This formation is eomposed of lime-

stone, dolomitie limestone, stromatolitie and lofe-

556

A. Reitano, F. Liberto, S. Giglio, R. Grasso & M.T Spena

ritic dolomite, megalodon limestone, algal biolitite,

and eoral biolitite. The eave is known for Paleoli-

thie and Neolithie artifaets and Pleistoeenie mam-

mal faunal remains (De Stefani, 1941; Mannino et

al., 1986), and for invertebrate troglobiotie fauna

(Brian, 1959; Caruso, 1982; 1995; Caruso & Costa,

1978; Spena, 2007; Krapp et al., 2010; Cottarelli

et al., 2012). The eave is strongly affeeted by out-

side thermal fluetuations due to its small size

(Mannino et al., 1986; Spena, 2007; Cottarelli et

al., 2012), and as a result of the karstie eonditions,

the surrounding area is devoid of running water.

The eave is fossil; it is never flooded, and the water

input to the eave is exelusively due to rainfall ex-

pressed as temporary drips (Cottarelli et al., 2012).

In the past, the area elose to the eave was eultivated

with olive, almond and earob trees, but today na-

tive shrub vegetation (oak and broom) is reelai-

ming the area (Figs. 1-6).

The rest of the territory of the reserve is oecu-

pied by non-native speeies {Pinus, Cupressus, Eu-

calyptus, Acacia) planted by the “Azienda

Regionale Foreste Demaniali”. There are also

aspeets of grassland (Gianguzzi et al., 2009). Ove-

rall, herbaeeous plants make up 50% of the taxa li-

sted by Gianguzzi et al. (2009), followed by

perennial grasses, partieularly hemieryptophytes

(24%) and geophytes (13%). There are 14 endemie

taxa, amongst whieh, of partieular importanee, one

ean mention Asperula rupestris Tineo (exelusive to

North-Western Sieily), Cymbolaria pubescens

(Presl) Cufod, Helichrysum panormitanum Guss.,

Echium italicum siculum (Laeaita), and Eryngium

bocconii Lojae. The soils are mainly made of roehy

outerops and lithosols (lithie xerorthents).

MATERIALS AND METHODS

The research in the field allowed us to make

ecological observations on molluscs, and the fin-

ding of live specimens and shells for subsequent

morphological and taxonomic classifications. Inside

the cavity, different sampling methods were used,

namely retrieval by sight, pitfall traps, and baiting.

For visual collection, a careful search through the

entire cave and in all possible microhabitats was

carried out. Animals and shells were therefore col-

lected on calcareous crusts on the walls, on clay,

under large and small stones naturally lying on the

ground, under organic remains (dung, wood, orga-

nic matter in decomposition of natural plant and

animal) (Bucciarelli, 1961) and in the vicinity of

small water-filled containers. This was aimed at fin-

ding species with different ecological requirements.

The other technique used was to put, within the

cave, pitfall traps in environments characterized by

different substrates and different morphology, in

order to sample in all possible habitats (soil, sludge,

and clastic elements with variable granulometry).

This collection technique requires special precau-

tions, especially inside the cave, because, if misu-

sed, it can cause serious damage to wildlife and

degrade communities. Consequently, the traps were

frequently and carefully monitored in such a way

as to avoid any excess catch. The research carried

out in the territory of the reserve spanned both the

area of the Integral Reserve, "Zone A", and “pre-ri-

serve Zone B". In both areas, samples were taken

in different habitat types, using retrieval by sight

and pitfall-traps (Figs. 7-8). Within the second zone,

research was carried out in different microhabitats

including hypolithic environment, litter and rock

outcrops in the area in front of the entrance.

The species have been classified in the labora-

tory through the morphological analysis of the shell

and the genitalia with the aid of a Leica microscope

MZ12.5. Photos were taken with a digital camera.

Taxonomical references are based on the checklist

of the Italian fauna (Bodon et al., 1995; Manganelli

et al., 1995, 1998, 2000) and the checklist of

"Fauna Europaea", version 1.1. (Bank, 2011, avai-

lable at: http://www.faunaeur.org). Chorological

categories are those proposed by La Greca (1962)

and later elaborated upon by Vigna Taglianti et al.

(1993, 1999) and Parenzan (1994). All species here

treated are accompanied by notes on ecology and

biology. Since taxonomic revisions were not the

scope of this work, taxonomic comments were re-

duced or delayed to works still in preparation. All

examined specimens are preserved in the collec-

tions of the authors.

RESULTS

SYSTEMATICS

GASTROPODA

Ordo Architaenioglossa Haller, 1890

Familia Cochlostomatidae Kobelt, 1902

Terrestrial molluscs from the R.N.I.“Grotta Conza” (Palermo, Sicily) (Gastropoda Architaenioglossa Pulmonata) 557

Figures 1-6. R.N.I. “Grotta Conza” environments. Figure 1. Entraee to Conza eave view by the inside. Figure 2. Idem, view

by the outside. Figure 3. Wall of ealeareous roek. Figure 4. Grassland and maquis with Euphorbia dendroides. Figure 5. Bru-

shwood. Figure 6. Slope with rupieolous vegetation.

558

A. Reitano, F. Liberto, S. Giglio, R. Grasso & M.T Spena

Cochlostoma paladilhianum paladilhianum

(Saint Simon, 1869)

Endemic species of North-Western Sicily, with

a subspecies pirajnoi (Paulucci, 1789) present on

Favignana island (Zilch, 1958; Alzona, 1971;

Bodon et ah, 1995). It lives on limestone walls and

rocks. On the area of the reserve it is rather locali-

zed to vertical rocky walls.

Familia Aciculidae J.E. Gray, 1850

Acicula benoiti (Bourguignant, 1864)

Endemic species of Sicily. It lives in the crevices

of limestone rocks and in underground environ-

ments (Boeters et al., 1989; Reitano et al., 2009).

Only empty shells were found by sieving debris col-

lected in the inner part of the cave and in the crevi-

ces of rocks outside the cave (Fig. 9).

Ordo Fittorinimorpha Pchelintsev, 1963

Familia Pomatiidae Newton, 1891

Pomatias elegans (O.F. Muller, 1774)

European species, recently reported for Nor-

thern Tunisia (Ben Romdhane et al., 2008). In Sicily

it lives in many kinds of biotopes: dune environ-

ments, from mid-mountain forests, in public gar-

dens areas, in the litter shrub formations and

riparian forest. It is considered to belong to "kera-

toconchae" (Sacchi, 1952; Giusti & Castagnolo,

1982), being able to resist long periods of drought

by burying itself in the soil or hiding under dead

leaves, stones and vegetal debris. Fiving specimens

were found under stones in the most humid places

and with relatively greater vegetation cover.

Tudorella panormitana (Sacchi, 1954)

Endemic species of the mountains of Palermo,

distributed from a few meters above sea level up to

medium altitudes, from Trabia to Monte Palmeto

(Sacchi, 1954; Alzona, 1971). In Sicily another spe-

cies T. multisulcata is present, which is widespread

in the mountains of Trapani and Southern Sicily.

T. panormitana (Fig. 10) is morphologically

characterized by the white or pale yellow shell (in

T. multisulcata, the colour is variable from orange

to dark red) and a reticulate sculpture of raised spi-

ral threads and radial ribs {T. multisulcata has spiral

striae only a little more pronounced than the radial

ribs). Fossils of Pleistocenic Tudorella Fischer,

1885, found in the mountains of Palermo, have the

same morphological characters of the shell of T. pa-

normitana. Fiving and fossil specimens of the

mountains of Trapani have the typical characters of

T. multisulcata (Sacchi, 1954) (personal data). The

two species are distinct also as regards genital mor-

phology; T. panormitana has a penial apex which

is much thinner and more elongated with respect to

that of T. multisulcata (Fo Brano & Sparacio,

Figure 7. R.N.I. “Grotta Conza” environments: pitfall traps. Figure 8. Idem, pitfall traps hidden under a stone.

Terrestrial molluscs from the R.N.I.“Grotta Conza” (Palermo, Sicily) (Gastropoda Architaenioglossa Pulmonata)

559

2006). On the basis of recent molecular studies of

Pfenninger et al. (2010), it seems that the current

distribution of Tudorella in the Western Mediterra-

nean is due to the fragmentation of the original di-

stribution area that occurred in the late Oligocene

and Miocene, and dispersive phenomena that oc-

curred in the late Miocene and Pleistocene. The mo-

lecular study of Pfenninger et al. (2010) also

highlights the existence of significant genetic di-

stances between populations of T. panormitana and

T. multisulcata. T panormitana is a xeroresistent

species typical of limestone landscapes; in the re-

serves living specimens were found under stones,

especially in grassland.

Infraclassis Pulmonata Cuvier in Blanville, 1824

Ordo Stylommatophora A. Schmidt, 1855

Familia Cochlicopidae Pilsbry, 1900

Hypnophila cylindracea (Calcara, 1840)

The genus Hypnophila Bourguignat, 1858 in Si-

cily is represented by three endemic species: H.

emiliana (Bourguignat, 1858) from the Aegadian

Islands, H. incerta (Bourguignat, 1858) from the

Aeolian Islands and H. cylindracea with a disjunct

distribution limited to North-Western Sicily. The

species belonging to the genus Hypnophila are con-

sidered of great biogeographical interest for their

distribution in the Mediterranean basin (Giusti &

Manganelli, 1984). H. cylindracea lives in crevices

of limestone or under stones in damp niches rich in

plant debris. Many empty shells were found in the

inner part of the cave (Fig. 11).

Familia Pleurodiscidae Wenz, 1923

Pleurodiscus balmei (Potiez & Michaud, 1838)

Species with very fragmentary East Mediterra-

nean distribution. In Italy it is reported for the

Southern regions and Sicily (Alzona, 1971; Man-

ganelli et al., 1995; Hallgass & Vannozzi, 2009).

This xeroresistant species inhabits open environ-

ments, but also tree stumps or wet karstic or vol-

canic caves. This trogloxene species is common

inside the cave.

Familia Chondrinidae Steenberg, 1925

Granopupa granum (Drapamaud, 1801)

European-Mediterranean-Turanian species wi-

despread along the Italian peninsula, G. granum is

a xeroresistent species quite common in Sicily. In

the reserve shells were found under rocks in areas

with sparse vegetation cover.

Rupestrella rupestris margritae (Beckmann,

2002)

R. rupestris (Philippi, 1836) is an endemic spe-

cies of Sicily. Beckma nn (2002) distinguishes five

subspecies: R. rupestris rupestris, widespread in

Western Sicily, R. rupestris coloba (Pilsbry, 1918)

known for Levanzo island, R. rupestris carolae

Beckmann, 2002, from the “Rocca di Cefalu”, R.

rupestris lamellosa Beckmann, 2002, from the area

between Sciacca and Caltabellotta and R. rupestris

margritae Beckmann, 2002 from the territory of

Isola delle Femmine. The R. rupestris specimens

found in the reserve show the typical morphological

characters of the subspecies margritae: shell sleek,

with a small and roundish mouth, with well-formed

columellar tooth not visible in frontal view (Fig.

12). Rupestrella species are rupicolous, calciophilic

and xeroresistant; in the reserve, R. r margritae

lives on limestone walls with small populations.

Rupestrella homala homala (Westerlund, 1 892)

Endemic species of Sicily, with three subspe-

cies (Beckmann, 2002): R. homala homala (We-

sterlund, 1892), widespread in North-Western

Sicily, R. homala massae Beckmann, 2002, known

only for the southern slopes of Monte Cofano (Cu-

stonaci) and R. homala falkneri known for the area

between Termini Imerese and Caccamo. R. homala

homala is distinguished from R. rupestris by more

convex whorls, greater convexity of the whorls,

less marked suture, and a more stocky silhouette

due to a wider last whorl (Fig. 13.) Biology is si-

milar to that of the previous species. R. homala has

been described for Monte Gallo, a place about 1 .5

km far from the reserve.

Familia Vertiginidae Fitzinger, 1833

560

A. Reitano, F. Liberto, S. Giglio, R. Grasso & M.T Spena

Truncatellina callicratis (Scacchi, 1833)

European-Mediterranean-Turanian species, wi-

dely distributed in the Mediterranean region (Giusti

et al., 1995). It lives in very dry limestone landsca-

pes with herbaceous vegetation. Some shells have

been found through the debris picked up in the cre-

vices of rocks, under rocks and at the base of cliffs.

Familia Enidae B.B. Woodward, 1903

Mastus pupa (Linnaeus, 1758)

Species with Holomediterranean distribution,

present in Southern Italy, Sardinia and Sicily (Al-

zona, 1971; Manganelli et al., 1995). It is a ther-

mophilic species, present in a wide variety of

habitats. In the reserve is common, especially in

areas with open grassland, or between stones in

rocky environments.

Familia Ferussaciidae Bourguignat, 1883

Cecilioides (Cecilioides) cff. acicula (O.F. Miil-

ler, 1774)

Species with Turanian-European-Mediterranean

distribution, extended to Macaronesia. The genus

Cecilioides Ferussac, 1814 comprises endogenous

species that usually live buried under stones, bet-

ween plant roots and on the soil of caves. Many

shells have been found both in the inner part of the

cave and in the debris picked up out from the cave,

under rocks or in crevices of the limestone walls

(Fig. 14).

Hohenwarthiana aradasiana (Benoit, 1862)

Endemic species of Sicily (Giusti, 1973; Man-

ganelli et al., 1995). It occurs in endogenous habi-

tats, buried under stones or among the roots of the

plants. Shells of H. aradasiana were found in the

same habitats of the previous species (Fig. 15).

Ferussacia folliculus (Gmelin, 1791)

Species with Mediterranean distribution, thermo-

philic, inhabiting arid environments with little ve-

getation cover. In the reserve both living specimens

and empty shells were found under rocks or debris.

Familia Subulinidae P. Fischer & Crosse, 1877

Rumina decollata (Linnaeus, 1758)

The genus Rumina Risso, 1 826 is represented in

Sicily with two species: R. saharica Pallary, 1901,

recently reported for Marettimo island (Liberto et

al., 2012) andR. decollata, widespread throughout

the island. It has a Mediterranean distribution, and

it is present throughout the Italian peninsula and the

islands. It is a typical Mediterranean species belon-

ging to keratoconchae, which populates natural en-

vironments but also degraded and anthropized ones.

It is omnivorous and even necrophilic. It was intro-

duced in many areas of Central-Northern America

to be used in the biological control of the invasive

helicid Cornu aspersum (Cowie, 2001). It lives fre-

quently under stones, among organic debris, in dry

walls, and under dead trunks, bushes and shrubs. It

is a thermophilic and xeroresistant species thanks

to the thick white epiphragm and to its habit of bu-

rying itself into the soil during summer.

Familia Clausiliidae J.E. Gray, 1855

Siciliaria leucophryna (L. Pfeiffer, 1862)

Endemic species of Sicily, limited to the sur-

roundings of Sferracavallo (Palermo). This species

has been confused with S. eminens (Schmidt, 1 868),

particularly by Boettger (1879) and Benoit (1882),

however Nordsieck (2002) specified its distribution

area in the neighbourhoods of Sferracavallo and se-

lected a lectotype (Senckenberg Museum, Frankfurt

67478, coll. O. Boettger ex-Dohm ex-L. Pfeiffer).

S. leucophryna is a calciophilic specie, that occurs

in the crevices of the rocks and under stones in sha-

ded places (Fig. 18).

Papillifera bidens affinis (Philippi, 1836)

P. bidens is a very common species in Sicily and

inhabits many natural and anthropized biotopes. It

is a calciophilic species, but can form large popu-

lations in non-calcareous environments (like the

Terrestrial molluscs from the R.N.I.“Grotta Conza” (Palermo, Sicily) (Gastropoda Architaenioglossa Pulmonata)

561

Figure 9.Acicula benoitih: 4.35 mm, D: 1.33 mm ; Figure 10. Tudorella panormitana h: 19 mm, D: 12.7 mm . Figure 11. Hyp-

nophila cylindracea h: 6.2 mm , D: 2.1 mm . Figure 12. Rupestrella r margritae h: 4.14 mm , D: 2 mm . Figure 13. R. h. homala

h: 3.45 mm, D: 2 mm. Figure 14. Cecilioides efr. acicula h: 6.16 mm, D: 1.45 mm. Figure 15. Hohenwarthiana aradasiana h:

4.3 mm, D: 1.45 mm. Figure 16. Daudebardia b. sicula D: 3.65 mm. Figure 17. Oxychilus fuscosus h: 7.7 mm, D: 14 mm.

562

A. Reitano, F. Liberto, S. Giglio, R. Grasso & M.T Spena

crystalline rocks of Peloritani Mountains, the vol-

canic rocks of the Iblean plateau and Mount Etna).

In Sicily it is represented by three subspecies: P. bi-

dens ajjinis, widespread in Sicily, P. bidens tinei

(Westerlund, 1878) in South-Western Sicily, and/!

bidens rudieosta (O. Boettger, 1878) in the Central-

Estern Sicily. The systematics of Sicilian popula-

tions of P. bidens is probably more complex and

requires a modem revision. In the reserve the sub-

species P. bidens ajjinis, described for the surroun-

dings of Palermo, is abundant.

Familia Oleacinidae H. & A. Adams, 1855

Poiretia dilatata dilatata (Philippi, 1836)

Species with S-European distribution, limited to

the central-eastern sector (Central and Southern Italy,

Sicily, the Peloponnese, and Crete); in Sicily it is pre-

sent as nominal subspecies (Subai, 1980). It lives in

forest and riparian environments, less frequently

under stones or at the base of cliffs. Occasionally it

is also present in caves. P. dilatata eats both plants

and other molluscs that they actively hunt (Liberto

et al., 2010). A few shells were found in the most sha-

ded and vegetated areas of the reserve.

Familia Discidae Thiele, 1931

Discus rotundatus rotundatus (O.F. Muller, 1774)

Species with European-Mediterranean distribu-

tion extending to Macaronesia, and present in al-

most all Italian territory (Alzona, 1971; Manganelli

et al., 1995). It inhabits deciduous forests, under lit-

ter or under rotting wood, and mderal environments

under debris. In the reserve, D. rotundatus is un-

common and localized amongst the bushes close to

the entrance of the cave.

Familia Pristilomatidae T. Cockerell, 1891

Vitrea cfr. subrimata (Reinhardt, 1871)

Species with European-Mediterranean distribu-

tion spreading throughout Italy. V. subrimata com-

prises a complex of populations which differ by the

structure of flabelliform appendices of the inner

wall of the proximal penis and therefore needing a

modem revision (Manganelli et al., 1995; Ferreri

et al., 2005). It inhabits mainly natural environ-

ments, usually cavities in soil, or under partly con-

cealed rocks or boulders. In the reserve, only empty

shells were found by sieving debris collected under

rocks or in crevices.

Familia Oxychilidae P. Hesse, 1927

Daudebardia brevipes sicula (Bivona, 1839)

Endemic subspecies of Central-Southern Italy

and Sicily, described for the area around Palermo

(Manganelli et al., 1995), occurs in litter of forests,

also under stones and in wet and shaded debris. Few

specimens and empty shells were found under sto-

nes, in damp environments, especially with auto-

chthonous forest cover (Fig. 16).

Oxychilus (Oxychilus) fuscosus (Rossmassler,

1838)

Endemic species of North-Western Sicily, distri-

buted along the mountains of Palermo in damp en-

vironments, especially with forest cover. The shell

of O. fuscosus is characterized by the well-rounded

last whorls, a rather narrow umbilicus and espe-

cially by the presence in the upper part of the shell

of a reticulate microsculpture consisting of very fine

longitudinal furrows crossed by marked and irregu-

lar furrows and growth lines (Fig. 17). This micro-

sculpture is well visible near the suture and

gradually disappears towards the periphery of the

last whorl of the spire. The dimensions of the shell

are variable from one population to another, and in

favorable environments the maximum diameter can

reach 35 mm. Specimens of O. fuscosus were found

at the entrance of the cave, in front of the entrance

and under the rocks in shaded and vegetated places.

Familia Milacidae Ellis, 1926

Milax nigricans (Philippi, 1836)

Species with Holomediterranean distribution,

widespread and common throughout Sicily, in na-

Terrestrial molluscs from the R.N.I.“Grotta Conza” (Palermo, Sicily) (Gastropoda Architaenioglossa Pulmonata)

563

tural and anthropized habitats. This species was de-

scribed for the surroundings of Palermo, therefore

the population of the reserve is topotypic.

Tandonia sowerbyi (A. Ferussac, 1823)

The genus Tandonia Lessona et Pollonera, 1882

is widespread in Sicily with two species: T sower-

byi with Holomediterranean- Atlantic distribution

and T. marinellii Liberto, Giglio, Colomba & Spa-

racio, 2012, at present only known for the North-

Western Sicily (Trapani mountains). These two

species have a very similar external appearance and

can be classified with certainty only after a morpho-

logical examination of the genitalia (Liberto et al.,

2012). T sowerbyi is found in open hilly and forest

environments, but also in gardens and greenhouses.

Some living specimens were found under rocks in

the grassland of the reserve and in front of the en-

trance of the cave.

Familia Agriolimacidae H. Wagner, 1935

Deroceras (Deroceras) panormitanum (Les-

sona et Pollonera, 1882)

D. panormitanum was considered a species very

variable in colour and shape of the genitalia (Giusti

1973; 1976; Giusti & Manganelli, 1990; Giusti et

al., 1995; Wiktor, 2000; Reise et al, 2011). We be-

lieve that D. panormitanum s.l. is a complex of dif-

ferent species, which require a detailed morphological

and molecular analysis. D. panormitanum s. 1. is very

common in Sicily, where it lives in a wide range of

environments. In the reserve two morphotypes live

in simpatry, they differ in size of the body and in

genital morphology.

Familia Trissexodontidae H. Nordsieck, 1987

Caracollina (Caracollina) lenticula (Mi-

chaud, 1831)

Species with Mediterranean distribution. In Sicily

it is very common from a few meters above sea level

up to medium altitudes. In the reserve living speci-

mens and empty shells were found under stones.

Familia Cochlicellidae Schileyko, 1972

Cochlicella {Cochlicella) acuta (O.F. Muller, 1774)

Species with Mediterranean- Atlantic distribu-

tion, C. acuta is thermophilic, typical of the coast,

but it can live at medium altitudes in natural or an-

thropized environments. In the reserve, living spe-

cimens were found in the xeric areas.

Familia Hygromiidae Tryon, 1866

Monacha consona (Rossmassler, 1839)

Endemic Sicilian species with the locus typicus

restricted to the surroundings of Palermo (Forcart,

1965). It has a globular shell, medium size, with a

generally open umbilicus, and genitalia with a ty-

pical scheme of Monacha Fitzinger, 1833 sensu

strictu, meaning without the retractor muscle, with

vaginal appendix and mucous glands. Rather hy-

grophilous species, uncommon in the reserve.

Szentgalya gregaria (Rossmassler, 1839)

Endemic Sicilian species (Manganelli et al.,

1995; Liberto et al., 2010), characterized by a small

to medium-sized sub-globose depressed shell, with

a generally closed umbilicus (Fig. 19) and lacking

of genital digitiform glands, vaginal appendix

(which are present in M. consona) and retractor mu-

scle. S. gregaria is less hygrophilous than M. con-

sona and may form abundant populations also in

xeric environments. When it is inactive it takes re-

fuge a few centimeters under the soil. Liberto et al.

(2010) elevate Szentgalya Pinter, 1977 to the rank

of a full genus but Bank (2011) and Falkner et al.

(2011) do not accept Szentgalya either as genus or

as subgenus. We prefer to follow the choice of Li-

berto et al. (2010) waiting to publish our additional

data. S. gregaria is included in the checklist of the

International Union for Conservation of Nature

(lUCN) as a “near threatened” species. The species

is quite common in xeric grasslands of the reserve.

Trochoidea (Trochoidea) caroni (Deshayes,

1830)

Species endemic to Southern Italy and Sicily

(Manganelli et al., 1995), also reported as fossil for

the Maltese Islands (Giusti et al., 1995). Currently,

it is considered a species with a certain variability

564

A. Reitano, F. Liberto, S. Giglio, R. Grasso & M.T Spena

in shape and size of the shell, ineluding different

populations whieh probably need a modem syste-

matie review. Relatively widespread in Sieily, from

eoastal areas to medium altitudes, espeeially in

grassy habitats. Uneommon in the reserve (Fig. 20).

Trochoidea {Trochoidea) pyramidata (Drapar-

naud, 1805)

Speeies with W- Mediterranean distribution, re-

ported for peninsular and insular areas of Italy. It is

thermophilie and xeroresistant, eommon in low to

medium altitudes. It inhabits herbaeeous vegetation

and shmbs, even several inehes from the ground, in

multiple environments, often degraded, natural and

anthropized. This speeies shows a remarkable po-

lymorphism of the shell, both in size and eolour

(Giusti, 1973). T. pyramidata is common and wide-

spread in the reserve, especially on the grassland.

Schileykiella reinae (Pfeiffer, 1857)

Endemic species of the N-W Sicily with di-

sjunct distribution. The genus Schileykiella Man-

ganelli, Sparacio et Giusti, 1989 includes two other

species: S. parlatoris (Bivona, 1839), widespread

in Sicily, and S. bodoni Cianfanelli, Manganelli et

Giusti, 2004, known only for Marettimo island

(Cianfanelli et al., 2004). S. reinae is a calciophilic

and hygrophilous species that occurs in forest en-

vironments, Mediterranean maquis, at the base of

limestone cliffs, or under rocks and debris. In the

reserve only empty shells were found (Fig. 21).

Xerotricha conspurcata (Drapamaud, 1801)

Species with Mediterranean distribution, repor-

ted in Italy for peninsular regions, Sicily and Sar-

dinia. It is very common in Sicily under stones and

debris from various coastal areas to medium altitu-

des. It lives in natural or antropized environments.

Cernuella (Cernuella) cfr. cisalpina (Rossmassler,

1837)

Species with an European-Mediterranean distri-

bution (Manganelli & Giusti, 1987; Manganelli et

al., 1995). The systematic position of the group ci-

salpina-virgata of the genus and subgenus Cer-

nuella Schliiter, 1838 is somewhat complex

because of the variability of the shell, a character

which has been fundamental for the description of

many taxa. Cernuella (C.) cisalpina is a complex

of small or medium sized species with many forms

that require a systematic review (Manganelli &

Giusti, 1987; Favilli et al., 1995; Giusti et al., 1995;

Ferreri et al., 2005). In Sicily it is common, from

back dunes up to medium altitudes, in natural and

anthropized environments.

Cernuella (Cernuella) cfr. virgata (Da Costa, 1778)

Species with European-Mediterranean distri-

bution (Manganelli & Giusti, 1987; Manganelli et

al., 1995). Many taxa with uncertain taxonomic

value, generally characterized by medium to large

sized shells, are attributed to C. virgata. Thermo-

philic and xeroresistant, it is widespread in Sicily,

at low and medium altitudes, in different natural

environments, even degraded and anthropized,

often in numerous colonies. In the reserve is quite

common in grassland.

Familia Helicidae Rafinesque, 1815

Chilostoma macrostoma (Rossmassler, 1837)

Species endemic to the Palermo Mountains, the

supraspecific status of which is yet to be determined.

Some aspects of the taxonomy and nomenclatural

position of this entity have recently been subject of

deepenin (Lo Brano & Sparacio, 2006; Colomba et

al., 2008; Liberto et al., 2010). It inhabits natural fo-

rest and rocky environments. A few shells were col-

lected in the crevices of the rocks of the reserve.

Murella platychela platy chela (Menke, 1830)

Endemic species of the North-Western Sicily,

with several subspecies; the nominal subspecies

was described for Monte Cuccio (Pfeiffer, 1931), a

few kilometers South of the area covered by this

study. It is strictly calciophilic and inhabits rocky

sites. Specimens were found on the rocky walls of

the reserve (Fig. 22).

Eobania vermiculata (O.F. Muller, 1774)

Species with Holomediterranean distribution.

Terrestrial molluscs from the R.N.I.“Grotta Conza” (Palermo, Sicily) (Gastropoda Architaenioglossa Pulmonata)

565

Figure 18. Siciliaria leuchophryna h: 20.5 mm, D: 4.8 mm. Figure 19. Szentgalya gregaria h: 6 mm, D: 9.4 mm. Figure 20.

Trochoidea (T.) caroni h: 9.45 mm , D: 9.8 mm. Figure 21. Schileykiella reinae h: 3.5 mm, D: 6.4 mm. Figure 22. Murella

platychela h: 16.2 mm , D: 23 mm. Figure 23. Erctella mazzullii h: 28.5 mm, D: 27 mm.

566

A. Reitano, F. Liberto, S. Giglio, R. Grasso & M.T Spena

Dispersed by man in many non-Mediterranean

eountries. It is thermophilie, eommon in many types

of environments, both natural and anthropized,

from a few meters above sea level up to medium al-

titudes. In the reserve living speeimens have been

found under roeks or in ereviees of limestone roek.

Cantareus apertus (Bom, 1778)

Speeies with Mediterranean distribution, present

in almost all the Italian peninsula, Sieily and Sardi-

nia. It is thermophilie, eommon in open, arid, even

anthropized and degraded environments, at low and

medium altitudes. During the summer it seeks re-

fuge under the ground, elosing the opening of the

shell with a robust white and eonvex epiphragm. In

the reserve it was found in grassland.

Cornu aspersum (O.F. Muller, 1774)

Speeies with European-Mediterranean distribu-

tion, present in almost the whole peninsular and in-

sular Italian territory, but passively introdueed by

man in many other eountries (Giusti et al., 1995).

Reeent moleeular studies (Guiller & Madee, 2010)

have made it possible to reeonstmet the phytogeny

of the populations of C. aspersum in the Mediter-

ranean area, reeognizing the Kabylia area (North-

Western Algeria), as the probable original region of

the speeies; however, no populations from Sieily

were analyzed in this study. The populations of Si-

eily and eireum-Sieilian islands have a eertain mor-

phologieal variability that requires further study and

investigation. C. aspersum sensu latu inhabits many

habitats: the dune system, the Mediterranean ma-

quis, dry meadow hills, the forests of high mountain

(Lo Brano & Sparaeio, 2006), and mderal or an-

thropized areas. C. aspersum is eommon and wide-

spread throughout the reserve.

Erctella mazzullii (De Cristofori & Jan, 1832)

Endemie speeies of the Palermo Mountains, oe-

eurring from Monte Pellegrino to Monte Palmeto

(Terrasini). Reeently, the E. mazzullii eomplex has

been the subjeet of a multidiseiplinary revision

work (morphologieal, moleeular, paleontologieal,

and biogeographieal) (Colomba et al., 2011) that

has allowed the re-evaluation of the genus Eretella

Monterosato, 1894 and the reeognition, as valid

speeies, of E. mazzulli (De Cristofori & Jan, 1832),

E. eephalaeditana (Giannuzzi-Savelli, Oliva et Spa-

raeio, 2012), endemie of the Roeea of Cefalu, and

E. insolida (Monterosato, 1892), widespread in the

mountains of Trapani. The speeies of Eretella genus

are ealeiophilie and saxieavous; in faet they live ex-

elusively on limestone roeks, on whieh they dig pe-

euliar tunnels using aeid to dissolve the hard

Mesozoie limestone. Tunnels are used as a refuge

from the summer temperature and from predators

during periods of inaetivity. The speeies is rather

rare and loealized within the reserve, on the walls

of limestone (Fig. 23).

CONSIDERATIONS AND CONCLUSIONS

The present study has allowed us to eompile,

for the Natural Reserve "Grotta Conza", a eheeklist

of 41 speeies of terrestrial molluses, a signifieant

number in relation to the limited extension of the

reserve (12.34 heetares). The high number of spe-

eies is mainly related to three faetors: 1) the reserve

is North faeing, whieh makes this area subjeet to

the main eool and humid air masses eoming from

the North-West; 2) the lithologieal nature of the en-

vironment eonsisting of limestone; 3) the riehness

of mierohabitats.

The humidity and the availability of ealeium are

fundamental for the biology of terrestrial molluses,

while the diversity of environments (slopes with

seattered boulders, roeks, vertieal eliffs, gorges,

eaves) and the diversity of the vegetation (xerie gra-

ssland, Mediterranean maquis, patehes of serub, ru-

picolous vegetation), provide mieroenvironments

suitable for the demands of the different speeies.

From the eeologieal point of view, most of the

molluses are either ealeiophilie and rupieolous {Co-

ehlostoma paladilhianum, Rupestrella rupestris mar-

gritae, R. homala homala, Murella platyehela

platyehela and Eretella mazzullii), ealeiophilie and

interstitial {Aeicula benoiti, Hypnophila eylindraeea,

Ceeilioides aeieula and Hohenwartiana aradasiana),

ealeiophilie and typieal of roeky masses (Tudorella

panormitana, Siciliaria leueophryna, Papillifera bi-

dens, and Chilostoma maerostoma), xerophilie and

xeroresistant (Truneatellina eallieratis, Pleurodiseus

balmei, Mastus pupa, Rumina deeollata, Xerotrieha

eornspureata, Cernuella ef. eisalpina, Cernuella ef.

virgata, Szentgalya gregaria, Troehoidea earoni.

Terrestrial molluscs from the R.N.I.“Grotta Conza” (Palermo, Sicily) (Gastropoda Architaenioglossa Pulmonata)

567

Trochoidea pyramidata, Cantareus apertus, Grano-

pupa granum, Ferussacia folliculum, Caracollina

lenticula, and Cochlicella acuta), hygrophilous (Po-

matias elegans. Discus rotundatus, Vitrea cf. subri-

mata, Oxychilus fuscosus, Daudebardia brevipes

sicula, Schileykiella reinae, and Monacha consona),

and species with a larger ecology, also anthropophilic

(Milax nigricans, Tandonia sowerbyi, Deroceras pa-

normitanum, Poiretia dilatata, Eobania vermiculata,

and Cornu aspersum).

The chorological data (Table 1) highlight the

predominance of endemic taxa (41%), followed by

species with Mediterranean (34%), European (20%)

and wide distribution (5%). This composition can

be explained by the geographical location of the re-

serve in the northern mountains of Palermo (North-

Western Sicily). This region, as a result of the

succession of marine ingressions and regressions of

the Pleistocene and the complex topography of the

area, has undergone long periods of geographic iso-

lation leading to speciation processes, followed by

reunification with the near areas and then re-colo-

nization by European and Mediterranean species.

However, some endemisms have an even older,

Mio-Pliocenic origin: T. panormitana is phyloge-

netically related with North-West African and Sar-

CHOROTYPES

N. OF

SPECIES

CHOROTYPES WITH WIDE DISTRIBUTION

4.88

EUROPEAN-MEDITERRANEAN-TURANIAN extended to

MACARONESIA

4.88

CHOROTYPES WITH EUROPEAN DISTRIBUTION

19.51

EUROPEAN-MEDITERRANEAN-MACARONESIAN

2.44

EUROPEAN-MEDITERRANEAN

12.19

EUROPEAN

2.44

S-EUROPEAN

2.44

CHOROTYPES WITH MEDITERRANEAN DISTRIBUTION

34.15

MEDITERRANEAN

12.19

WEST MEDITERRANEAN

2.44

EAST MEDITERRANEAN

2.44

HOLOMEDITERRANEN

7.32

HOLOMEDITERRANEAN-ATLANTIC

4.88

SOUTHERN APENNINE-SICILIAN

4.88

ENDEMISMS

41.46

SICILIAN

41.46

TOTAL

100%

Table 1 . The chorological data of the terrestrial molluscs from R.N.I. “Grotta Conza”.

568

A. Reitano, F. Liberto, S. Giglio, R. Grasso & M.T Spena

dinian Tudorella (Pfenninger et al., 2010); Erctella

shows morphological and ecological affinities with

mpieolous and saxieavous Helieidae from Kabylia

(North-Eastern Algeria) referable to Helix suba-

perta (Aneey, 1893) (Colomba et al., 2011); while

Hypnophila is distributed on disjoint ealeareous

platforms, a phenomenon best explained in light of

the series of geologieal events taking plaee in the

Western Mediterranean region towards the end of

the Oligoeene and the early Plioeene (Giusti &

Manganelli, 1984). The endemie taxa inelude 17

speeies and subspeeies. Two of them, S. leuco-

phryna and R. rupestris margritae, have their di-

stribution limited to limestone eoastal environments

between Sferravallo and Isola delle Femmine and

are therefore worthy of speeial attention and pro-

teetion. T. panormitana, C. macrostoma, and E.

mazzullii are endemie speeies with a distributional

area limited to the mountains of Palermo. C. pala-

dhilianum, H. cylindracea, R. homala, O.fuscosus,

S. reinae, and M. platychela are endemies to North-

Western Sieily; finally, H. aradasiana, P bidens af-

Jinis, M. consona, S. gregaria, and D. panormitanum

s.l. are endemie to Sieily with a wider distribution.

Some of these endemie speeies are ineluded in

the Red List of Threatened Speeies of the Interna-

tional Union for Conservation of Nature (lUCN)

(Falkner et al., 20 11). The E. mazzulli eomplex is

eonsidered "endangered"; S. gregaria, S. reinae, R.

rupestris are elassified as “near threatened”, and R.

homala and C. paladilhiamim are eonsidered “vul-

nerable”. The main threats are the destruetion of

the natural environment through quarries for the

extraetion of marble and gravel, the eonstruetion

of roads, urban development, and uneontrolled

fires. The presenee of the R.N.I. "Grotta Conza" is

therefore a very important tool for the eonservation

and proteetion of these speeies and the environ-

ment in whieh they live.

In partieular, we believe it is desirable to take

all appropriate measures to prevent fires, and to thin

or eradieate non-native plant speeies (i.e. Pinus,

Cupressus, diVid Eucalyptus), whieh eause the aeidi-

fieation of the soil, making it inhospitable to some

endemie snail speeies (Z). brevipes sicula, O.fusco-

sus, S. reinae and M. consona). In addition, a thiek

reforestation of Pinus would eompete with native

tree speeies, whieh would reduee the development

of a suitable undergrowth neeessary to meet the

eeologieal needs of almost all speeies of molluses

linked to the ground.

ACKNOWLEDGMENTS

The authors wish to thank M. Vaeearella, Presi-

dent of the Club Alpino Italiano, Sieilian seetion,

and G. Mento, Direetor of the R.N.I. “Grotta

Conza”, for supporting and eneouraging biologieal

researeh. Funding was provided by the “Assesso-

rato Regionale Territorio e Ambiente della Regione

Sieiliana” to R. Grasso. We are greatefiil to Roberto

Viviano and Arturo Viviano for their support on the

field and to the operators Gianluea Chiappa and

Giuseppe Lo Dieo.

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Biodiversity Journal, 2012, 3 (4): 571-582

Molecular studies on the genus Medora H. et A. Adams, 1 855

from Italy (Gastropoda Pulmonata Clausiliidae)

M. Stella Colomba'*, Fabio Liberto^ Agatino Reitano^ Walter Renda^ Giuseppe Pocaterra^ Armando

Gregorini' & Ignazio Sparacio^

'Universita di Urbino “Carlo Bo”, Dipartimento di Scienze Biomolecolari, via Maggetti 22 (loc. Sasso) - 61029 Urbino (PU), Italy

^Strada Provinciale Cefalu-Gibilmanna n° 93 - 90015 Cefalu, Palermo, Italy

^Via Gravina 77 - 95030 Tremestieri Etneo, Catania, Italy

"•via Bologna 18/a - 87032 Amantea, Cosenza, Italy

^Via XXV Aprile 19 - 40018 San Pietro in Casale, Bologna, Italy

^Via E. Notarbartolo 54 int. 13 - 90145 Palermo, Italy

*Corresponding author: mariastella.eolomba@uniurb.it

ABSTRACT In Italy, the genus Medora H. et A. Adams, 1855 includes two species: M. italiana (Kiister,

1847) andM dalmatina (Rossmassler, 1835). In particular, populations ascribed to M. italiana

were, and still are, the focus of several works aiming at better understanding the real taxonomic

value of these entities and defining their presence on the Italian territory. In order to contribute

to the improvement of the current knowledge on the organization of the genus at different ta-

xonomic levels, several populations were investigated by analysing 16S rDNA, COI and ITS2

gene partial sequences. Phylogenetic reconstructions were obtained by the Maximum Likeli-

hood algorithm. Although further studies are needed, preliminary data suggest that the genus

Medora shows a much more complex and articulate differentiation than hypothesized so far.

KEY WORDS Medora; Medora italiana complex; 16S rDNA; COI; ITS2; molecular phylogeny.

Received 12.05.2012; accepted 02.11.2012; printed 30.12.2012

Proceedings of the P' International Congress “Insularity and Biodiversity”, May 11*-13*, 2012 - Palermo (Italy)

INTRODUCTION

The genus Medora H. et A. Adams, 1855 has a

typieal trans-Adriatie distribution (Gridelli, 1950;

La Greea, 1964; La Greea, 1984; Vigna Taglianti et

ah, 1993; Parenzan, 1994) with different speeies

seattered in the North-Western Balkan peninsula,

Italian peninsula and Sieily (Giusti et al., 1986; Rei-

tano et al., 2007; Nordsieek, 2007; 2009; 2012;

Bank, 2011). Regarding Italy, Nordsieek (1970)

eonsidered M. italiana (Kiister, 1 847) of the Cen-

tral-Southern Apennines distinet fromM albescens

(Menke, 1830) of the Balkan peninsula, as the two

taxa differ by a few anatomieal eharaeters, mainly

by the insertion mode of the divertieulum of the

bursa eopulatrix. In addition, he assigned to M. ita-

liana the following subspeeies: M. i. italiana (loeus

typieus: Piedimonte Matese, Caserta, Campania),

M. i. punctulata (Kiister, 1850) (loeus typieus:

Monte Tiriolo, Catanzaro, Calabria), M. i. garga-

nensis (A.J. Wagner, 1918) (loeus typieus: Pulsano,

Monte Sant’Angelo, Foggia, Apulia), M. i. milet-

tiana Giusti, 1967 (loeus typieus: Monte Miletto,

Campobasso, Molise) and M. i. kobelti Nordsieek,

1970 (loeus typieus: Gola di Romagnano, Potenza,

Basilieata).

A subsequent revision of the genus Medora in

Italy and Yugoslavia was published by Giusti et al.

(1986) based on morphologieal features, anatomieal

eharaeters and allozymes (i.e. variant forms of an

572

M.S. COLOMBA, F. Liberto.A. Reitano.W. Renda, G. Pocaterra.A. Gregorini & I. Sparacio

enzyme eoded by different alleles at the same

loeus). Aeeording to these authors, the statistieal

analysis of eonehologieal eharaeters allowed to re-

eognize two speeies, M albescens with two subspe-

eies and M. dalmatina (Rossmassler, 1835) with

three subspeeies; the anatomieal analysis of genita-

lia was of poor diagnostie value distinguishing only

two speeies, M. albescens and M. dalmatina, wi-

thout any elements at the subspeeifle level; and, fi-

nally, allozymes allowed to diseriminate between

two speeies with numerous subspeeies: M. albe-

scens with five subspeeies: albescens, almissana

(Kiister, 1 847) and clissana (Branesrk, 1 897), in ex-

Yugoslavia, italiana and punctulata, in Italy, and M.

dalmatina with three subspeeies, dalmatina, ortho-

pleura (Westerlund, 1878) and drasnicensis H. Nor-

dsieek, 1970, all in Croatia, Bosnia and

Herzegovina. In eonelusion, Giusti et al. (1986)

suggested that: 1) it was not possible to distinguish

M. italiana from M. albescens', 2) Medora albe-

scens was widespread in the Balkan peninsula with

several subspeeies and in Italy with two subspeeies:

M. a. italiana in the eentral part of Italy, and M. a.

punctulata showing a disjunetive distribution with

some populations in Northern and Central Apenni-

nes and others in Calabria; and 3) the examined po-

pulation of Gargano resulted indistinguishable from

that of Ospo (Slovenia) and probably was of anthro-

pie origin (see Giusti et al., 1986: 260). However,

the authors also suggested that several subspeeies

of Medora eould be eonsidered good speeies aeeor-

ding to the eleetrophoretie analysis results, but due

to the absenee of morphologieal eharaeters diseri-

minating among different taxa they preferred to

lump them all togher (Giusti et al., 1986: 260, 316).

After nearly ten years, Manganelli et al. (1995)

reported for the Italian peninsula only two speeies

without subspeeies: M. albescens and, as first re-

eord for Italy, M. dalmatina whose presenee, stres-

sed by Foreart (1965) and Alzona (1971), had been

ruled out by Nordsieek (1970) and Giusti et al.

(1986). Later on, Nordsieek (2007) stressed onee

again that M. albescens is a Balkan taxon (sub M.

macascarensis albescens) and eonfirmed for Italy,

as stated previously (Nordsieek, 1970), M italiana

with the subspeeies mentioned above. Reitano et al.

(2007) deseribed another additional subspeeies, M.

italiana peloritana Reitano, Liberto et Sparaeio,

2007 from Monte Veneretta, Messina (NE Sieily).

Colomba et al. (2008) earned out a preliminary

study on moleeular geneties of Italian populations

of Medora, highlighting a good differentiation of

the populations studied, espeeially the Southern

ones. Bank (2011) reported for Italy M. italiana

with the five subspeeies eited above, while not ei-

ting M. dalmatina. Reeently, Nordsieek (2012) de-

seribed a new subspeeies of M. dalmatina'. M.

dalmatina pollinensis Nordsieek, 2012 from Sou-

thern Italy, namely Basilieata and Calabria (S. Lo-

renzo Bellizzi, Cerehiara di Calabria, Gole di

Raganello (near Civita), Orsomarso and Papasi-

dero) and, aeeording to unpublished preliminary

moleeular data (Hausdorf, unpubl.), antieipated that

not only M. italiana would be a distinet speeies

fromM macascarensis (Sowerby, 1828) but also

that M. i. garganensis should be elevated to the spe-

eies level. Finally, Cianfanelli et al. (2012) reported

M. dalmatina pollinensis from Campieello, Laino

Castello and Papasidero (Cosenza, Calabria) in al-

luvional debris.

The present study was earned out to eharaete-

rize, by moleeular analyses, the genus Medora in

Italy, with partieular attention to the M. italiana

eomplex aiming at understanding the real taxono-

mie value of the populations aseribed to it and better

defining the presenee of this group in the Italian ter-

ritory. To this purpose, we studied nearly all taxa

eurrently eonsidered subspeeies of M. italiana in-

eluding M. i. italiana from Piedimonte Matese, M.

i. punctulata from Monte Consolino and Monte Ti-

riolo, M. i. peloritana from Monte Veneretta, M. i.

garganensis from San Mareo in Lamis (Foggia), M.

i. milettiana from Monte Miletto, plus several M.

italiana populations from Leonessa (Rieti), Posta

(Rieti), Rubbiano (Aseoli Pieeno) and Gualdo Ta-

dino (Perugia). Finally, a few additional populations

of other speeies of the genus Medora were ineluded,

i.e., M. dalmatina pollinensis from Civita, Orso-

marso and Papasidero (Cosenza), and M. macasca-

rensis (= albescens) from Makarska (Croatia).

Choosing moleeular markers is eertainly a eru-

eial point when studying moleeular evolution. Ge-

nerally, mitoehondrial genes are mostly employed

as markers in phylogenetie studies of elosely related

speeies. This is beeause mitoehondrial genes show

eonsiderable advantages ineluding: (i) abundanee

in the tissues, (ii) easiness in being manipulated,

(iii) inheritanee in single eopy; (iv) high mutation

rates, whieh makes them partieularly useful in this

type of analysis. With regard to the eytoehrome oxi-

Molecular studies on the genus Medora from Italy (Gastropoda Pulmonata Clausiliidae)

573

dases, just because of the activities they perfomi,

the functional products of these genes are highly

conserved. This feature associated with high rates

of mutation, especially in third position, makes both

COI and COII extremely well suited to investigate

the phylogenetic relations among taxa separated in

relatively recent times. Mitochondrial genes, howe-

ver, represent only a small part of the entire ge-

nome, hence, to ensure a more thorough and

detailed phylogenetic analysis, they are often stu-

died with nuclear genes (typically 18S, 5.8S and

28S rDNAs or the internal transcriptional spacers,

ITSl and ITS2). On this basis, we decided to inve-

stigate the phylogenetic relationships among the

Italian taxa of the genus Medora by comparing ri-

bosomal partial sequences of mitochondrial (16S

rDNA and cytochrome oxidase c subunit I, COI)

and nuclear (second internal transcriptional spacer,

ITS2) genes, in order to shed some light on this in-

teresting group whose systematics is still unclear

and (in some respects) controversial.

MATERIALS AND METHODS

Specimens

Sixty-six (66) specimens were fixed in 70-90%

ethanol. Particularly, fourty-six M. italiana from

Piedimonte Matese (Caserta, Campania), Monte

Consolino, Stilo (Reggio Calabria, Calabria),

Monte Veneretta (Messina, Sicily), Monte Tiriolo

(Catanzaro, Calabria), Monte Miletto (Campo-

basso, Molise), San Marco in Lamis (Foggia, Apu-

lia), Leonessa (Rieti, Latium), Posta (Rieti,

Latium), Rubbiano (Montefortino, Fermo, Marche)

and Gualdo Tadino (Perugia, Umbria); fifteen M.

dalmatina pollinensis from Papasidero, Orsomarso

and Civita (Cosenza, Calabria); and five M. maca-

scarensis from Makarska (Croatia) (Table 1). Each

population was labelled respectively as PMA,

CON, PEL, TIR, MIL, GAR, REA, RIE, RUB, GT,

PAP, RSO, CIV and MCR, using consecutive num-

bers for specimens (up to five per site). Samples

were stored at -20 °C in test tubes. For each indivi-

dual, the entire animal was used for total DNA ex-

traction (by Wizard Genomic DNA Purification Kit,

Promega). For each population, para-voucher spe-

cimens - sensu Groenenberg et al. (2011) i.e. diffe-

rent specimens than the ones used for DNA

analysis, but from the same sample or population -

were stored by Dr. Colomba (University of Urbino).

16S rDNA fragments (283-292 bp) were amplified

by the primers MED16S_F (5’-ACTGTGCAA-

AGGTAGCATAA-3’) and MED16S_R (5’-CCAA-

CATCGAGGTCACAA-3’) (present paper); COI

amplicons (616-658 bp) were obtained by the uni-

versal internal primers ECO 1490 (5’-GGTCAACA-

AATCATAAAGATATTGG-3’) and HC02198

(5 ’-TAAACTTCAGGGTGACCAAAAAATCA-3 ’)

as in Folmer et al. (1994); ITS2 fragments (445-531

bp) were amplified by the primers ITS2_F (5’-

ACATTGAAC ATCGACATCTTGA-3’) and

ITS2_R (5’-CTCCGCTTAGTAATATGCTTAA-3’)

(present paper). The three molecular loci were am-

plified in 50 pi reactions by the following (slightly

different) PCR protocols: (i) 95 °C for 5 min; 95 °C

for 1 min, 55 °C for 1 min, 72 °C for 1 min (30 cy-

cles), 72 °C for 10 min (16S rDNA and ITS2); (ii)

95 °C for 5 min; 95 °C for 1 min, 50 °C for 1 min,

72 °C for 1 min (35 cycles), 72 °C for 10 min

(COI). To remove primers and unincorporated nu-

cleotides, the amplified products were purified by

the Wizard SV gel and PCR Clean-up kit (Pro-

mega). Sequencing of the purified PCR products

was carried out using automated DNA sequencers

at Eurofins MWG Operon (Germany). Sequence

chromatograms of each amplified fragment were

browsed visually for reading mistakes by the se-

quencer. All sequences were visualized with BioE-

dit Sequence Alignment Editor 7 (Hall, 1999),

aligned with the ClustalW option included in this

software and double checked by eye. Nucleotide di-

versity within groups (n = Pi and 7i:j(2; Pi corrected

according to Jukes and Cantor) and nucleotide di-

vergence between groups corrected according to

Jukes and Cantor (Dxyj^^), using the full set of all

sequences were computed with DnaSP 5 (Librado

& Rozas, 2009). Phylogenetic analyses were con-

ducted in MEGA 5 (Tamura et al., 201 1) using the

Maximum Likelihood algorithm. The best-fit evo-

lution models of nucleotide substitution resulted Ta-

mura 3 -parameter + gamma distribution for 16S

rDNA and COI; and Kimura 2-parameter + gamma

distribution for ITS2. Support for nodes was calcu-

lated by the Bootstrap method (1000 replicates) and

expressed as percentages. Muticaria syracusana

and Muticaria neuteboomi 16S rDNA, COI and

ITS2 partial sequences (GenBank IDs: HQ696866-

HQ696869, AY382II7) were used as outgroup to

574

M.S. COLOMBA, F. Liberto.A. Reitano.W. Renda, G. Pocaterra.A. Gregorini & I. Sparacio

Figure 1. Consensus tree for 16S rDNA gene partial sequenees showing the evolutionary history inferred by using the Maximum

Likelihood method based on the Tamura 3-parameter model + Gamma distribution (G= 0.28, 5 eategories). The tree is drawn to

seale, with braneh lengths measured in the number of substitutions per site. Bootstrap pereentages are shown on the nodes.

Molecular studies on the genus Medora from Italy (Gastropoda Pulmonata Clausiliidae)

575

0.02

100

RS03 IT3

RS04 ITSi

RS02 ITS2 1

RS01 ITS2

PAP1 ITS2

RS05 ITS2

CIV1 ITS2

CIV2 ITS2

CIV3 ITS2

CIV4 ITS2

CIVS ITS2

PAP2 ITS2

PAP3 ITS2

AP4 its:

PS ITS

TIR5 ITS2’

ONI ITS2

CON2 ITS2

CONS ITS2

CON4 ITS2

CONS ITS2

PEL1 ITS2

H PEL2 ITS2

PEL3 ITS2

PEL4 ITS2

M. pollinemis

M. punctiilata

M. milettiana

M. macascarensis

MUTSYR ITS2

M. garganensis

Out Group

Figure 2. Consensus tree for ITS2 gene partial sequenees obtained by using the Maximum Likelihood method based on the Kimura

2-parameter model + Gamma distribution (G= 0.46, 5 eategories) The tree is drawn to seale, with braneh lengths measured in the

number of substitutions per site. Bootstrap pereentages are shown on the nodes.

576

M.S. COLOMBA, F. Liberto.A. Reitano.W. Renda, G. Pocaterra.A. Gregorini & I. Sparacio

0.1

M punctulata

M. macascarensis

M. milettiana

M. ita liana

M. garganemis

CIV1 COI

CIV5 COI

RS01 CCjl

RS02 CC

RS03 Cp\

RS04 £01

RSp5 COI

MUTNEU CO(

M pol linens is

100 L MUTSYR COI

Out Group

Figure 3. Consensus tree for COI gene partial sequenees obtained by using the Maximum Likelihood method based on the Tamura

3-parameter model + Gamma (G= 0.17, 5 eategories) The tree is drawn to seale, with braneh lengths measured in the number of

substitutions per site. Bootstrap pereentages are shown on the nodes.

Molecular studies on the genus Medora from Italy (Gastropoda Pulmonata Clausiliidae)

577

root the phylogenetic trees. Sequences generated in

this study were released in GenBank (IDs:

KC833899-KC834014,KC853238-KC853281).

RESULTS

Mitochondrial genes (16S rDNA and COI) sho-

wed a phylogenetic signal strong enough to distin-

guish unambiguously all taxa and mostly resolve

their phylogenetic relations; in addition, ITS2 was

particularly useful to separate all taxa at the specific

level. In order to be able to better decipher the phy-

logenetic framework, particular attention has been

devoted to genetic distances. Differences between

DNA sequences are frequently reported as p distance

(i.e. proportion, calculated as number of nucleotide

differences divided by the total number of sites). Un-

fortunately, p is a rather rough estimate of the actual

number of evolutionary changes occurred, since the

same site may have undergone more than one mu-

tations over time. Hence to try to correct the risk of

underestimation, divergence between two sequences

is therefore not simply measured as a percentage of

different nucleotides, but, taking account of possible

events of multiple substitutions and back-substitu-

tions, is calculated using suitable mathematical mo-

dels which consider the stochastic nature of gene

mutation. In this study we used the Jukes and Cantor

correction which assumes equal base frequencies

and equal mutation rates.

All populations under study were analysed for

the 16S rDNA marker. As shown in figure 1, the to-

pology of the maximum likelihood consensus tree

revealed several clearly distinct clusters: ''pollinen-

sis'' including the populations of Papasidero, Orso-

marso and Civita; ''garganensis'' with the

population of San Marco in Lamis; ''milettiana''

with the population of Monte Miletto; ''macasca-

rensis’’’’ with the population of Makarska; and, fi-

nally, the ''Medora italiana complex” which

comprises a few sub-clusters. These latters, indica-

ted with alphabet letters, are respectively, cluster A

including the populations of Monte Tiriolo, Monte

Consolino and Monte Veneretta; cluster B with the

scientific name

Localities

Latitude

Longitude

exx.

MCR

M. macascarensis

Croatia, Makarska, Kotisina, 350 m slm, IV.2008,

G. Pocaterra

43017-29" N

17°02'48" E

RUB

Medora sp. 1

I, Marche, Fermo, Montefortino Rubbiano, 580 m slm,

IV.2009, G. Pocaterra

42°55'43.60" N

13°18'31.18"E

RIE

Medora sp. 1

I, Lazio, Rieti, Posta, Fiume Velino, Romualdo, 604 m

slm, X.2010, A. Hallgass & G. Pocaterra

42°28'39"N

13°05'08"E

Medora sp. 2

I, Umbria, Perugia, Gualdo Tadino, La Rocchetta, 600

m slm, IV.2009, G. Pocaterra

43°13'42.58" N

12°48'13.39" E

REA

Medora sp. 2

I, Lazio, Rieti, Leonessa, Monti Reatini, 1107 m slm,

9.IX.2009,A. Hallgass

42°31'52.65"N

12°57'03.08" E

MIL

M. milettiana

I, Molise, Campobasso, Campitello Matese, Monte Mi-

letto, 1620 m slm, X.2008, A. Hallgass

41°27'41.23"N

14°22'43.95" E

PMA

M. italiana

I, Campania, Caserta, Piedimonte Matese, Gole del To-

rano, 274 m slm, 7.VIII.2008,W. Renda & G. Pocaterra

41°21'45.23"N

14°22'46.53" E

GAR

M. garganensis

I, Puglia, Loggia, San Marco in Lamis, 5.VI.2008,

M. Perilli

CIV

M. pollinensis

I, Calabria, Cosenza, Civita, Gole del Raganello, 340 m

slm, 13.VI1.2008, W. Renda

39°49'45.80" N

16°19'00.69" E

PAP

M. pollinensis

I, Calabria, Cosenza, Papasidero, Valle del Lao, 192 m

slm. 13.IX.2008,W. Renda

39°52'00.05" N

15°54'04.19"E

RSO

M. pollinensis

I, Calabria, Cosenza, Orsomarso, Valle dell' Argentine,

131 m slm, 13.IX.2008, W. Renda

39°48'01.36''N

15°54'26.91"E

TIR

M. punetulata

I, Calabria, Catanzaro,Tiriolo, Monte Triolo, 720 m

slm, 21.11.2009, W. Renda

38°57'04.58" N

16°31'04.59"E

CON

M. punetulata

I, Calabria, Reggio Calabria, Stilo, Monte Consolino,

666 m slm, 3.VII.2008, W. Renda

38°28'54.11"N

16°27’53.57"E

PEL

M. punetulata

peloritana

I, Sicily, Messina, Castelmola, Monte Veneretta, 750 m

slm, 26.IX.2008, A. Reitano & W. Renda

37°52'10.72'' N

15°16'02.50" E

Table 1 . Medora populations collected in Croatia and Italy.

578

M.S. COLOMBA, F. Liberto.A. Reitano.W. Renda, G. Pocaterra.A. Gregorini & I. Sparacio

CIV

CON

GAR

MIL

MCR

MNEU

MSYR

PAP

PEL

PMA

REA

RIE

RSO

RUB

TIR

CIV

CON

16%

GAR

16%

13%

14%

13%

MIL

14%

11%

MCR

14%

11%

MNEU

27%

26%

25%

24%

MSYR

26%

25%

27%

25%

26%

24%

PAP

16%

15%

14%

27%

26%

PEL

15%

13%

25%

23%

14%

PMA

17%

15%

10%

26%

18%

REA

14%

12%

26%

14%

RIE

16%

13%

25%

15%

RSO

18%

17%

28%

18%

17%

19%

RUB

15%

10%

12%

27%

26%

15%

11%

18%

TIR

15%

12%

25%

23%

14%

17%

Table 2. Matrix showing the number of base differenees per site (expressed in %) from averaging over all 16S rDNA

partial sequenee pairs between groups.

CIV

CON

GAR

MCR

MIL

MNEU

MSYR

PEL

PMA

RSO

TIR

CIV

CON

17%

GAR

17%

12%

MCR

16%

10%

12%

MIL

17%

10%

11%

10%

MNEU

19%

MSYR

19%

20%

18%

19%

20%

PEL

17%

12%

10%

19%

20%

PMA

16%

10%

13%

11%

10%

21%

11%

RSO

18%

17%

18%

21%

20%

18%

17%

TIR

17%

12%

10%

19%

20%

11%

18%

Table 3. Matrix showing the number of base differenees per site (expressed in %) from averaging over all COI partial

sequenee pairs between groups.

Molecular studies on the genus Medora from Italy (Gastropoda Pulmonata Clausiliidae)

579

population of Piedimonte Matese; cluster C sho-

wing the populations of Posta and Rubbiano; and

eluster D with the populations of Leones sa and

Gualdo Tadino. Genetie distanees range from 2%

to 19% (Table 2). Based on tree topology, we be-

lieve that, within the M. italiana eomplex, elusters

A (CON + TIR + PEL), B (PMA), C (RJE + RUB)

and D (GT+ REA) are, respeetively, separate spe-

eies. When eomparing PMA (eluster B) to the po-

pulations of eluster A, the genetie divergenee is

7-8% (16S rDNA, Table 2) or 10-11% (COI, Table

3). Henee, M. italiana from Piedimonte Matese ean,

with reasonable eertainty, be eonsidered a separate

speeies. Cluster A ineludes the populations of

Monte Tiriolo (loeus typieus of ''punctulata'"),

Monte Consolino, and Monte Veneretta whieh,

based on their genetie divergenees (Table 2) ean all

be aseribed to the same speeies whieh we refer to

as M. punctulata. In addition, both p distanees and

tree topology reveal a eertain degree of internal dif-

ferentiation whieh may suggest to eonsider them

subspeeies, as proposed by Reitano et al. (2007) for

the taxon from Monte Veneretta ('"peloritana’’' he-

rein labelled PEL). Regarding the populations in-

eluded within eluster C, indieated as Medora sp.l,

taking into aeeount the p distanees between them

(6.3%, for 16S rDNA), they eould even belong to

two distinet speeies. At the moment, however, we

eannot give a definitive answer to the question

whieh has to be analysed in-depth by larger samples

and more detailed studies (eurrently in progress).

Finally, eluster D, Medora sp. 2, eneloses two po-

pulations that we eonsider eonspeeifie (p distanee

for 16S rDNA, 5%). In this regard it is interesting

to note that the populations referred to as RIE (elu-

ster C) and REA (eluster D) are both from Latium,

respeetively from Posta (Rieti) and Leonessa

(Rieti), thus suggesting that in this region two Me-

dora speeies might oeeur in areas whieh are only

1 3 Km far. In eonelusion then, the M. italiana eom-

plex would inelude M. italiana, M. punetulata and

at least two other additional speeies at present re-

ported as Medora sp.l and Medora sp. 2. Finally,

the Maximum Likelihood eonsensus tree seems to

strongly support the speeifie rank of milettiana,

garganensis, maeasearensis and pollinensis.

As regards ITS2 and COI genes, although our

data are only partial beeause not all populations have

been studied yet, nevertheless, obtained results de-

serve attention as, although limited, seem to eonfirm

the big pieture outlined by the analysis of 1 6S rDNA

sequenees. Partieularly, ITS2 proved very useful in

diseriminating taxa at the speeifie level. Indeed, the

Maximum Likelihood analysis (Fig. 2) elearly di-

stinguishes pollinensis, milettiana, garganensis and

maeasearensis, eonfirming their speeifie rank and,

interestingly, separates M. italiana from Piedimonte

Matese (PMA) from the populations of M. punetu-

lata of Monte Consolino, Monte Tiriolo and Monte

Veneretta (CON, TIR, PEL). The same pattern is

shown by the Maximum Likelihood eonsensus tree

based on COI partial sequenees analysis (Fig. 3).

DISCUSSIONS AND CONCLUSIONS

The present work represents, as to our kno-

wledge, the first attempt to eonduet a full and tho-

rough moleeular analysis of the genus Medora from

Italy. Our findings, though preliminary and subjeet

to further study, however, allow us to suggest some

interesting eonsiderations both on the systematies

of the genus and on the paleo-biogeographie events

that, over time, originated the eurrent distribution

of Medora populations on the Italian territory (Fig.

4). First of all, it is interesting to note that all three

genes (16S rDNA, COI and ITS2) eonfirm the spe-

eifie rank of M. italiana, M. milettiana, M. garga-

nensis, M. punetulata andM pollinensis, underlining

that the genus Medora, in Italy, ineludes mueh more

speeies than reported so far. Given the distribution

(trans-Adriatie) of Medora and the faet that the Ita-

lian populations are little differentiated morpholo-

gieally (Giusti et al., 1986), it has been suggested

that Medora speeimens might have reaehed the Ita-

lian territories relatively reeently. At the end of the

Plioeene, the Apennines were already eonneeted to

the Alps and extended up to the Matese area with

the more southerly regions fragmented in a vast ar-

ehipelago (Sgrosso, 1998; Boeealetti et al., 2005).

The Italian peninsula was therefore well separated

from the Balkan peninsula, but with the first glaeia-

tion, the Adriatie Sea dried up almost eompletely

allowing most Eastern speeies to passage to Italy.

Moreover, glaeial eyeles oeeurred throughout the

Pleistoeene, joined to the major territorial extension

of the Apennine and to the paleo-elimatie pheno-

mena relevant to this period, allowed a progressive

spread and differentiation of these organisms in the

Italian territory up to the most southern regions (i.e.

580

M.S. COLOMBA, F. Liberto.A. Reitano.W. Renda, G. Pocaterra.A. Gregorini & I. Sparacio

Calabria, with the populations attributable to M.

punctulata). M. punctulata peloritana (from

North-Eastern Sieily) represents, therefore, the

southernmost population of the genus and, proba-

bly, the most reeently differentiated. Our data eon-

firm the affinity between peloritana and the

populations from Calabria (see also Reitano et ah,

2007; Nordsieek, 2012) and it is interesting to note

that in all examined eladograms, M punctulata pe-

loritana is always distinet and sometimes (as in the

ease of 16S rDNA) also well separated from the Ca-

labrian populations as a whole.

More eomplex eould be instead the position of

M. garganensis in relation to the geologieal vieis-

situdes of the “Abruzzi-Apulia paleobioprovinee"

whieh the Gargano is part of (efr. Azzaroli, 1982;

Rieehetti et al., 1992; Angelone, 2007). In faet, M.

garganensis might represent one of the earliest po-

pulations settled in Italy, or at least have had an evo-

lutionary proeess distinet from that of the other

populations sinee the Gargano area underwent long

periods of isolation.

The populations reported for Italy as M. dalma-

tina (Manganelli et al., 1995) were deseribed by

Nordsieek (2012) as a distinet subspeeies: M dal-

matina pollinensis. Notably, as already evideneed

by Giusti et al. (1986), the eharaeter of the initial

portion of the spiral fin forked ean be observed in

Figure 4. M. dalmatina

(star: Vrgorac, Croatia)

and Medora popula-

tions studied in this

paper (see also Table 1).

Dot 1) M macascaren-

sis (Makarska, Kotisina,

Croatia). Dots 2-14

(Italy): 2) M. sp. 1

(Montefiorito, Rub-

biano); 3) M. sp. 1

(Posta, Fiume Velino; 4)

M. sp. 2 (Gualdo Ta-

dino, La Roeehetta); 5)

M. sp. 2 (Leonessa,

Gola di Rio Fuggio);

6) M. milettiana (Cam-

pitello Matese, Monte

Miletto); 7) M. ita-

liana (Piedimonte Ma-

tese); 8)M garganensis

(San Mareo in Lamis,

Gargano); 9) M. polli-

nensis (Civita, Gole del

Raganello); 10) M. pol-

linensis (Cosenza, Pa-

pasidero); 11) M polli-

nensis (Cosenza, Orso-

marso, Italy); 12) M.

punctulata punctulata

(Tiriolo, Monte Triolo);

13) M punctulata ssp.

(Stilo, Monte Conso-

lino); 14) M. punctulata

peloritana (Castelmola,

Monte Veneretta).

Molecular studies on the genus Medora from Italy (Gastropoda Pulmonata Clausiliidae)

581

different populations of italian Medora whieh ean-

not be aseribable to '' dalmatina" and, one of the

anatomical characters of M. dalmatina, the length

of the zone of contact between the wall of the distal

portion of the epiphallus and that of the proximal

portion of the penis (DEP=PP) often differs among

populations of M dalmatina of the Balkan penin-

sula and Medora of Italy only by about 0.2 mm.

Moreover, convergence phenomena between Italian

and Balkan species have already been documented

and confirmed with molecular evidence in inverte-

brate taxa which had differentiated along the Apen-

nine chain (Sparacio, 2000; Audisio et al., 2009);

and, in addition, the Pollino massif, where M. pol-

linensis occurs, is a geographic area of high biodi-

versity. Hence, although its affinity with M.

dalmatina could be real, nevertheless waiting for

further molecular evidence and considering that our

data (Figs. 1-3) suggest a quite articulate differen-

tiation among the three examined populations of M.

pollinensis, at the moment we prefer to consider it

as a distinct taxon rather than a subspecies of M.

dalmatina of the Balkan peninsula.

Given that our results are to be further validated

by (i) increasing the number of populations and of

specimens per population, (ii) implementing the

phylogenetic reconstruction using an additional mi-

tochondrial gene, i.e. 12S rDNA, and (iii) perfor-

ming also the Bayesian analysis, obtained results

(although still preliminary) seem to indicate that:

- M. i. garganensis and M. i. milettiana consi-

dered (so far) subspecies of M. italiana can be ele-

vated to the specific rank as M. garganensis and M.

milettiana (for M. garganensis see also Nordsieck,

2012 ).

- M. italiana from Piedimonte Matese can be

considered a distinct species, also distinct from M.

maeasearensis.

- The populations of Monte Tiriolo are a sepa-

rate species, which we refer to as M. punetulata in-

cluding the populations of Monte Consolino and

Monte Veneretta.

From the above, it is clear that the genus Me-

dora shows a much more complex and articulate

differentiation than hitherto hypothesized by mor-

phological surveys (Nordsieck, 1970; Giusti et al.,

1986; Nordsieck, 2012) and that, in an attempt to

clarify its organization and internal structure, at va-

rious taxonomic levels, a more detailed analysis in-

cluding a higher number of molecular markers and

additional Medora populations both from Italy (for

example "''kobeltr) and Balkans (as M. dalmatina),

is required.

ACKNOWLEDGEMENTS

We are grateful to Marco Bodon (Genoa, Italy),

Alessandro Hallgass (Rome, Italy), and Matteo Pe-

rilli (San Marco in Famis, Foggia, Italy) for contri-

buting to specimens collection in the field.

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GENBANK ACCESSIONS PUBLISHED IN THE BIODIVERSITY JOURNAL 3, 2012

Muticaria brancatoi (Gastropoda Pulmonata Clausiliidae) - Colomba et al., 2012, 3: 251-258

COI - GenBank ID: KC550118, KC550119, KC550120

Tandonia marinellii (Gastropoda Pulmonata Milacidae) - Liberto et al., 2012, 3: 201-228

COI - GenBank ID: KC550121

Tandonia ef. sowerbyi (Gastropoda Pulmonata Milaeidae) - Liberto et al., 2012, 3: 201-228

COI - GenBank ID: C550122-C550123

Medora spp. (Gastropoda Pulmonata Clausiliidae) - Colomba et al., 2012, 3: 571-582

Medora sp. 16S rDNA - GenBank ID: KC833914-KC833917

Medora sp. 16S rDNA - GenBank ID: KC833943-KC833946

Medora sp. 16S rDNA - GenBank ID: KC833956-KC833959

Medora sp. 16S rDNA - GenBank ID: KC833947-KC833950

Medora garganensis 16S rDNA - GenBank ID: KC833909-KC833913

Medora italiana 16S rDNA - Gen B ank ID: KC833938-KC833942

Medora maeasearensis 16S rDNA - GenBank ID: KC833918-KC833922

Medora milettiana 16S rDNA - GenBank ID: KC833923-KC833927

Medora pollinensis 16S rDNA - GenBanlc ID: KC833899-KC833903

Medora pollinensis 16S rDNA - GenBank ID: KC833928-KC833932

Medora pollinensis 16S rDNA - GenBank ID: KC833951-KC833955

Medora piinetulata 16S rDNA - GenBank ID: KC833904-KC833908

Medora punctulata 16S rDNA - GenBank ID: KC833960-KC833964

Medora punetulata peloritana 16S rDNA - GenBank ID: KC833933-KC833937

Medora garganensis ITS2 - GenBank ID: KC833975-KC833979

Medora italiana ITS2 - GenBank ID: KC834000-KC834004

Medora maeasearensis ITS2 - GenBank ID: KC833980-KC833984

Medora milettiana ITS2 - GenBank ID: KC833985-KC833989

Medora pollinensis ITS2 - GenBank ID: KC833990-KC833994

Medora pollinensis ITS2 - GenBank ID: KC833965-KC833969

Medora pollinensis ITS2 - GenBank ID: KC834005-KC834009

Medora punetulata ITS2 - GenBank ID: KC834010-KC834014

Medora punetulata ITS2 - GenBank ID: KC833970-KC833974

Medora punctulata peloritana ITS2 - GenBank ID: KC833995-KC833999

Medora garganensis COI - GenBank ID: KC853248-KC853252

Medora italiana COI - GenBank ID: KC853267- KC853271

Medora maeasearensis COI - GenBank ID: KC853253-KC853256

Medora milettiana COI - GenBank ID: KC853257-KC853260

Medora pollinensis COI - GenBank ID: KC853272-KC853276

Medora pollinensis COI - GenBank ID: KC853238-KC853242

Medora punetulata COI - GenBank ID: KC853277- KC853281

Medora punetulata COI - GenBank ID: KC853243-KC853247

Medora punctulata peloritana COI - GenBank ID: KC853261-KC853266

584

NEW TAXA PUBLISHED IN THE BIODIVERSITY JOURNAL 3, 2012

Carabus (Eurycarabus) faminii romanoi n. sp. (Coleoptera Carabidae)

Rapuzzi et Sparacio, 2012, 3: 85

LISD-um:lsid:zoobank.org:act:5B4C04D5-E617-4CC4-8A5F-63C0CCD8AAC5

Laubuca brahmaputraensis n. sp. (Cypriniformes Cyprinidae)

Kulabtong, Suksri et Nonpayom, 2012, 3: 94

LISD-um:lsid:zoobank.org:act:2EFB6F23-0E99-4CDC-BC7E-3A0D44F54656

Erctella cephalaeditana n. sp. (Gastropoda Helicidae)

Giannuzzi-Savelli, Sparacio et Oliva, 2012,3: 109

LISD-um:lsid:zoobank.org:act:A669BFC6-46B2-4305-A70C-EDC47A5AF3B8

Petaloconchus (Macrophragma) laurae n. sp. (Mollusca Gastropoda Vermetidae)

Scuderi, 2012 , 3: 124

LISD-um:lsid:zoobank.org:act:BB644892-5C91-4F72-955E-F22F4292F3C2

Hemiplecta belerangn. sp. (Gastropoda Pulmonata Ariophantidae)

Cilia et Abbas, 2012 , 3: 138

FISD - um:lsid:zoobank.org:act:432BC97B-3 1 4 1 -4D4F-B2 1 6-72DF 1 FCA2222

Tandonia marinellii n. sp. (Mollusca Gastropoda Milacidae)

Fiberto, Giglio, Colomba et Sparacio, 2012, 3: 203

LISD-um:lsid:zoobank.org:act:DB8C6222-E38E-4068-AE3E-2136063A0B80

Muticaria brancatoi n. sp. (Gastropoda Pulmonata Clausiliidae)

Colomba, Reitano, Fiberto, Giglio, Gregorini et Sparacio, 2012, 3: 252

LISD-um:lsid:zoobank.org:act:5DE3972E-C981-4124-B95D-F3C340687C31

Carabus (Apotomopterus) francottei n. sp. (Coleoptera Carabidae)

Rapuzzi, 2012, 3: 243

LISD-um:lsid:zoobank.org:act:4472CD03-3844-45EC-A09B-6D734DDB950C

Carabus (Apotomopterus) eccoptopteroides n. sp. (Coleoptera Carabidae)

Rapuzzi, 2012, 3: 244

LISD-um:lsid:zoobank.org:act:2A70E98C-DD77-46A2-8C94-5BlFA95C1011

Carabus (Morphocarabus) hummeli putyatini n. ssp. (Coleoptera Carabidae)

Rapuzzi, 2012, 3: 479-486

LISD-um:lsid:zoobank.org:act:559CB2E2-C4A2-47C0-86B8-F56998C2843B