18
www. b i o d i ve rs i tyj ou rn a L com
ISSN 2039-0394 (Print Edition)
ISSN 2039-0408 (Online Edition)
Biodiversity
Journal
MARCH 2015, 6 (1): 1-418
bo
w
1 w o r 1 C
Ji v- i . ■ .
O'
|
rt of
FOR NATURALISTIC RESEARCH
AND ENVIRONMENTAL STUDIES
1
rvn 1
n
♦ * 0* *
Carabus (Megodontus) imperiaUs Fischer, 1822 - Russia, SW Altay, Kalbinskiy range
i %
The Genus Carabits Linnaeus, 1758 (Coleoptera Carabidae). Cara bus Linnaeus, 1758 is a genus of beelies of the Family Carabidae. It contains
about ! 00 subgenera and is highly differentiated in about 1 000 species and a very targe number of subspecies. Terrestrial and carnivorous Carabus are
nocturnal predators that feed on snails, earthworms and caterpillars. The genus is widespread in the Oloartic Region but nearly all the species are native
to the Paleartic. Carabus are adapted to live from the sea level up to 5000 m in the mountains, they can be easily found in all kinds of habitats except the
deserts and the areas permanently covered by ice. Unable to fly they usually have reduced to rudimental wings, a very few species occasionally have
individuals capable of flying; C c(a t raws, C. gramriatus and, probably, C mounts, which makes them excellent biogeograph ical indicators, Carabus
are of medium to large size (12 mm to 65 mm) with developed mandibles and long and strong legs; the dorsal surface of the body frequently has a
sculpture formed of three kinds of longitudinal parallel and symmetrical arranged striae and deep points; often the sculpture is very diversified and
shows conspicuously morphological variations: totally erased, irregular, protruding ribs, with large and deep fovea, etc. Very often Carabus arc
colorful and of fascinating beauty. For these reasons they arc so attractive and notable items for collections, in fact they have been collected and studied
since, at least, 200 years. Carabus is one of the most deeply studied groups of Coleoptera, with a huge I iterature on their taxonomy, biology, phylogeny,
ecology and biogeography. Nevertheless, the genus is still far to be completely known, in fact a number of new taxa are described every year especially
from the most remote areas of Asia and many taxonomic problems are far to be solved.
Ivan Kapuzxi. viaCialla 47, 33040 Prepot to (Udine) Italy; email: info@ronchidicialla.it
Cover photo by A. Plutenko
1. Carabus (Megodontus) schoenberri
sajanus Braining, 1927 - Russia, Siberia,
W Sajan Mts., A. Plutenko.
2. Carabus (Megodonfus) leach i panzer!
Dejean, 1 829 - Russia, SW Altay, Kalbin-
skiy range, A. Plutenko.
3. Carabus {Limnocarabus) clatratus au-
ra nie ns is Muller, 1902- Serbia, Voj vo d i -
na, Ruma, Sava river, L Rapuzzi.
Biodiversity Journal, 2015, 6 (1): 3-6
First record of Mesophylax aspersus (Rambur, 1 842) from the
Republic of Kosovo (Trichoptera Limnephilidae)
Halil Ibrahimi, Agim Gashi, Linda G rape i- Koto ri , Astrit Bilalli, Milaim Musliu & Ferdije Zhushi-Etemi
Department of Biology, Faculty of Mathematical and Natural Sciences, University of Prishtina “Hasan Prishtina”, “Mother Theresa”
p.n., 10 000 Prishtina, Republic of Kosovo
’Corresponding author, e-mail: linda_grapci@yahoo.com
ABSTRACT The distribution of Mesophylax aspersus Curtis, 1834 (Trichoptera Limnephilidae) ranges
from Western Europe, Mediterranean region, Madeira, Canary Islands and up to South-
western Asia. According to the present knowledge it is however almost absent from South-
eastern Europe. In this paper we present first record of M. aspersus from the Republic of
Kosovo. This is at the same time first country record of the genus. Unlike many countries
where this species is present abundantly in our case it is extremely rare. A single adult male
specimen of M. aspersus was found in an ultraviolet light trap at the Blinaje Hunting Reserve
on August 23rd 2013. This has been a single specimen of this species caught at this locality
during a one year monthly sampling of caddisflies with UV light traps and entomological
net. Another male specimen has been caught on September 24th 2014 at the same locality.
Streams and rivers in all parts of Kosovo were surveyed during the period 2009-2014 for
Trichoptera species and currently the Blinaje Hunting Reserve is the only locality where this
species has been found. The distributional area of this species has been considerably expan-
ded by this record. The closest country where this species has been recorded is Bosnia and
Herzegovina.
KEY WORDS Mesophylax aspersus ; Kosovo; Trichoptera; Balkan Peninsula.
Received 24.09.2014; accepted 12.12.2014; printed 30.03.2015
INTRODUCTION
The genus Mesophylax McLachlan 1882
(Trichoptera Limnephilidae) is classified according
to Schmid (1955, 1957) in Stenophylacini tribus
close to genera Stenophylax Kolenati, 1848 and
Micropterna Stein, 1874; this genus consists by
only six species in the European fauna (Malicky,
1998; 2004).
Species of genus Mesophylax are mainly dis-
tributed in the Mediterranean area and radiate quite
far to the West, North, East, South-west and South-
east (Malicky, 1998).
Mesophylax aspersus Curtis, 1834 has a distri-
bution mostly limited in countries surrounding the
Mediterranean Sea, occurring from the Canary
Islands to the Near East (e.g. Schmid, 1957;
Botosaneanu, 1974; Dakki, 1987; Bonada, 2004).
From the biological point of view, adults of M.
aspersus emerge in spring and undergo a summer
diapause in caves (Bouvet & Ginet, 1969;
Botosaneanu, 1974; Salavert et al., 2011). They
do not feed during the adult stage, surviving most
probably on the reserves of the adipose tissue
accumulated during the larval phase (Bournaud,
1971).
4
Halil Ibrahimi etalii
MATERIAL AND METHODS
Data sampling and processing
Adult caddisfly specimens were collected with
entomological net and ultraviolet light trap. The
sampling was carried out monthly between March
and December 2013 and only casually during the
autumn of 2014. Ultraviolet light was placed above
the white pan of 60 cm in diameter filled 10 cm
with water with a few drops of detergent. The trap
was placed on stream bank and operated from dusk
until next morning. Collected samples were pre-
served in 80 % ethanol. The specimens were identi-
fied under a stereomicroscope with determination
keys from Malicky (2004) and Kumanski (1985,
1988). Specimens were collected by Halil Ibrahimi
and were determined by Halil Ibrahimi. Specimens
of M. aspersus were verified by Professor Hans
Malicky. The collection is deposited at the Labor-
atory of Zoology of the Faculty of Natural and
Mathematical Sciences, University of Prishtina,
Kosovo.
Study area
The territory of Blinaje Hunting area designated
as special reserve zone is located in central part of
Kosovo, 1 5 km on the western side of Lypjan town.
The total surface of Blinaje special reserve is 5500
ha and stretches in the territory of three municip-
alities: Lypjan, Shtime and Gllogoc. The altitude
within this territory ranges from 670 to 860 m
above sea level. There are 33 artificial lakes present
inside Blinaje special reserve.
The sampling site (Fig. 1) is located at the spring
area of the only stream inside this area which is
adjacent to the biggest lake inside Blinaje special
reserve (42.5185°N, 20.9788°E, and 721 m above
sea level).
RESULTS
Family LIMNEPHILIDAE
Mesophy lax McLachlan, 1882
Mesophylax aspersus Curtis, 1834
A single adult male specimen of Mesophylax
aspersus was found in an ultraviolet light trap at
the Blinaje Hunting Reserve on August 23rd 2013.
This has been a single specimen of this species
caught at this locality during a one year monthly
sampling of caddisflies with UV light traps and
entomological net.
Other species associated with M. aspersus in
this sample are: Potamophylax pallidus (Klapalek,
1899) (10 males, 3 females), Micropterna nyctero-
bia McLachlan, 1875 (4 male, 1 female), Wormal-
dia occipitalis (Pictet, 1834) (1 male), Hydropsyche
saxonica McLachlan, 1884 (2 males) and Hydro-
psyche sp. (5 females); leg. Halil Ibrahimi.
Another male specimen of M. aspersus has been
caught on September 24th 2014 at the same locality
with ultraviolet light trap.
Other species associated with M. aspersus in
this sample are: Potamophylax pallidus (5 males, 2
females), Micropterna nycterobia (1 male, 1
female) and Hydropsyche sp. (2 females); leg. Halil
Ibrahimi.
DISCUSSION AND CONCLUSIONS
In this paper we present first record of Mesophy-
lax aspersus from the Republic of Kosovo. This is
at the same time first country record of the genus.
The distribution of M. aspersus ranges from
Western Europe, Mediterranean region, Madeira,
Canary Islands and up to southwestern Asia (until
Cachemira) (Malicky, 1998; Bonada et al., 2004).
In the Balkan Peninsula the species is however rare.
It has been previously reported from Bulgaria
(Kumanski, 1988) but after a revision of this genus
(Malicky, 1998), the eastern part of the Balkan
Peninsula seems to be inhabited by M. impunct-
antus McLachlan, 1884 and not M. aspersus
(Kumanski, 1997, 2007).
The distributional area of M. aspersus has been
considerably expanded by this record. The closest
country where this species has been recorded is
Bosnia and Herzegovina (Radovanovic, 1935).
This record is almost eight decades old and in
meantime despite detailed investigations in Bosnia
and Hercegovina (eg. Marinkovic-Gospodnetic,
1966, 1970, 1971, 1978; Stanic-Kostroman,
2009), this species hasn’t been found any more. In
Macedonia, a neighboring country to Kosovo, as
in the rest of the eastern part of the Balkan Penin-
First record of Mesophylax aspersus (Rambur, 1842) from the Republic of Kosovo (Trichoptera Limnephilidae)
5
Figure 1. Sampling site in Blinaje Hunting Reserve: A) Adriatic Sea Basin, B) Black Sea Basin, C) Aegean Sea Basin.
sula up to the Western Anatolia is present a sub-
species M. impunctatus aduncus Navas, 1923
(Kumanski, 1997). Thus, in the continental part of
the Balkan Peninsula, Kosovo seems to be the
border line between the distribution of M. as-
persus and M. impunctatus.
The species seems to be very rare in Kosovo.
More than 100 localities (Ibrahimi, 2011; Ibrahimi
et al. 2012 a, 2012 b, 2013) in streams and rivers
in all parts of Kosovo were surveyed during the pe
riod 2009-2014 for Trichoptera species and
currently the Blinaje Hunting Reserve is the only
locality where this species has been found. The
abundance of M. aspersus found in Kosovo also
seems to be low. Out of nearly 1100 caddisfly spe-
cimens caught during 2013 and 2014 in Blinaje
Hunting Reserve, only two specimens belong to M.
aspersus. This is not the case in other areas around
the Mediterranean Sea where this species is
present. For example in the Iberian Peninsula the
species is quiet abundant (Bonada, 2004).
This record is a further contribution to the
inventory of the caddisfly fauna of the Republic of
Kosovo which is one of the poorest investigated
areas in Europe (Pongracz, 1923; Marinkovic-
Gospodnetic, 1975, 1980; Malicky, 1986, 1999;
Ibrahimi, 2007; Ibrahimi & Gashi, 2008; Ibrahimi
et al., 2012 a; Ibrahimi et al., 2012 b; Ibrahimi et
al., 2013; Olah, 2010; Olah et al., 2013a; Olah et
al., 2013b).
ACKNOWLEDGEMENTS
This study was partially financed by the Min-
istry of Education, Science and Technology of the
Republic of Kosovo through the project “Identific-
ation of rare aquatic insects in some spring areas in
Kosovo”, Project holder Halil Ibrahimi.
REFERENCES
Bonada N., Zamora-Munoz C., Rieradevall M. & Prat
N., 2004. Trichoptera (Insecta) collected in Mediter-
ranean River basins of the Iberian Peninsula: Taxo-
nomic remarks and notes on ecology. Graellsia, 60:
41-69.
Bournaud M., 1971. Observations biologiques sur les
Trichopteres cavernicoles. Bulletin mensuel de la
Societe linneenne de Lyon, 7: 196-211.
Botosaneanu L., 1974. Notes descriptive, faunistiques,
ecologiques, sur quelques trichopteres du “trio
subtroglophile” (Insecta: Trichoptera). Travaux de
flnstitut de Speologie “E. Racovitza”, 13: 61-75.
Dakki M., 1987. Ecosystemes deau courante du haut
Sebou (Moyen Atlas): Etudes typologiques et ana-
lyses ecologiques et biogographiques des principaux
peuplements entomologiques. Travaux, Institut des
Sciences, Rabat, Serie de Zoologie 42.
Ibrahimi H., 2011. Faunistical, ecological and biogeo-
graphical characteristics of Kosovo caddisflies
6
Halil Ibrahimi etalii
(Insecta: Trichoptera). PhD Thesis, Faculty of
Mathematics and Natural Sciences, University of
Zagreb, Zagreb, Croatia, 185 pp.
Ibrahimi H., 2007. The biological evaluation of the eco-
logial conditions in the Prishtina River based on ma-
crozoobenthos composition. Master Thesis, Faculty
of Mathematics and Natural Sciences, University of
Sarajevo, Sarajevo, Croatia, 122 pp.
Ibrahimi H., Gashi A., Grapci-Kotori L. & Kucinic M.,
2013. First records of the genus Micropterna Stein,
1873 (Insecta: Trichoptera) in Kosovo with distribu-
tional and ecological notes. Natura Croatica, 22: 147—
155.
Ibrahimi H., Kucinic M., Gashi A. & Grapci-Kotori L.
2012a. The caddisfly fauna (Insecta, Trichoptera) of
the rivers of the Black Sea basin in Kosovo with
distributional data for some rare species. ZooKeys,
182: 71-85. doi: 10.3897/zookeys. 182.2485
Ibrahimi H., Kucinic M., Gashi A., Grapci-Kotori L.,
Vuckovic I. & Cerjanec D. 2012b. The genus Rhy-
acophila Pictet, 1873 (Insecta: Trichoptera) in Kosovo.
Aquatic Insects: International Journal of Freshwater
Entomology, 34 supl: 23-31. doi: 10.1080/01650424.
2012.643021
Ibrahimi H. & Gashi A., 2008. State of knowledge of
investigations on Trichoptera larvae in Kosova.
Ferrantia, 55: 70-73.
Kumanski K., 1985. Trichoptera, Annulipalpia. Fauna
Bulgarica 15, Bulgarska Akademi naNaukite, Sofia,
243 pp.
Kumanski K., 1988. Trichoptera, Integripalpia. Fauna
Bulgarica 19, Bulgarska Akademi naNaukite, Sofia,
354 pp.
Kumanski K., 1997. Contributions to the caddisfly fauna
(Trichoptera) of the central-western part of the
Balkan Peninsula. Lauterbornia, 31: 73-82.
Kumanski K., 2007. Second addition to volume 15
(Trichoptera: Annulipalpia) and volume 19
(Trichoptera: Integripalpia) of Fauna bulgarica.
Historia naturalis bulgarica, 18: 81-94
Malicky H., 2014. Trichoptera, Caddisflies. Fauna Euro-
paea version 2.6, http://www.faunaeur.org
Malicky H., 1986. Beschreibung von vier neuen Kocher-
fliegen-Arten aus der Turkei und aus Jugoslawien
(Trichoptera). Opuscula zoologica fluminensia, 4:
1-7.
Malicky H., 1998. Revision der Gattung Mesophylax
McLachlan. Beitrage zur Entomologie, 48: 115-144.
Malicky H., 1999. Bemerkungen uber die Verwan-
dtschaft von Hydropsyche pellucidula CURTIS (Tri-
choptera, Hydropsychidae). Linzer biologische
Beitrage, 31: 803-821.
Malicky H., 2004. Atlas of European Trichoptera. 2nd
Edition, Springer, Netherlands, 359 pp.
Marinkovic-Gospodnetic M., 1966. New species of
Trichoptera from Yugoslavia. Bulletin Scientifique,
Sec. A- Tome 11, No. 4-6.
Marinkovic-Gospodnetic M., 1970. Descriptions of some
species of Trichoptera from Yugoslavia. Annual of
the Biological Institute of the University in Sarajevo,
23: 77-84.
Marinkovic-Gospodnetic M., 1971. The species of the
genus Drusus in Yugoslavia. Annual of the Biological
Institute of the University in Sarajevo, 24: 105-109.
Marinkovic-Gospodnetic M., 1975. Fauna Trichoptera
SR Srbija. Zbornik radova o entomofauni Srbije, 1:
219-236.
Marinkovic-Gospodnetic M., 1978. The Caddis-Flies
(Trichoptera, Insecta) of Hercegovina (Yugoslavia).
Annual of the Biological Institute of the University
in Sarajevo, 31: 115-131.
Marinkovic-Gospodnetic M., 1980. Fauna Trichoptera
SR Srbija. Zbornik radova o fauni Srbije, 1: 71-84.
Olah J., 2010. New species and new records of Palearctic
Trichoptera in the material of the Hungary Natural
History Museum. Annales Historico-Naturales Musei
Nationalis Hungarici, 102: 65-117.
Olah J., Andersen T., Chvojka P., Coppa G., Graf W.,
Ibrahimi H., Previsic A. & Valle M., 2013a. The
Potamophylax nigricornis group (Trichoptera,
Limnephilidae): resolution of phylogenetic species
by fine structure analysis. Opuscula Zoologica
Budapest, 44: 167-200.
Olah J., Ibrahimi H. & Kovacs T., 2013b. The genus
Chaetopteroides (Trichoptera, Limnephilidae) re-
vised by fine structure analysis of parameres. Folia
Historico Naturalia Musei Matraensis, 37: 93-108.
Pongracz S., 1923. Recesszarnyuak. Neuropteroiden. In:
Csiki Emo Allattani Kutatasai Albaniaban. Explora-
tiones zoologicae ab E. Csiki in Albania peractae. IX.
A. Magyar Tudomanyos Akademia Balkan-Kutata-
sainak Tudomanyos Erdmenyei, 1: 160-166.
Radovanovic M., 1935. Trihoptere Jugoslavije. Glasnik
Zemaljskog Muzeja u Bosni i Hercegovini, 47: 73-84.
Salavert V., Zamora-Munoz C., Ruiz-Rodriguez M. &
Soler J., 2011. Female-biased size dimorphism in a
diapausing caddisfly, Mesophylax aspersus: effect of
fecundity and natural and sexual selection. Ecological
Entomology, 36: 389-395.
Schmid F., 1955. Contribution a l’etude des Limnophil-
idae (Trichoptera). Mitteilungen der Schweizerischen
Entomologischen Gesellschaft Beiheft, 28: 1-245.
Schmid F., 1957. Les genres Stenophylax Kol., Microp-
terna St. et Mesophylax McL. (Tricopt. Limnoph.).
Trabajos del Museo de Zoologia de Barcelona, N.S.
Zool, 2: 1-51.
Stanic-Kostroman S., 2009. Faunisticke, ekoloske i
biogeografske znacajke tulara (Insecta: Trichoptera)
Bosne i Herzegovine. PhD Thesis, University of
Zagreb, Faculty of Mathematics and Natural Sci-
ences, Zagreb, Croatia, 151 pp.
Biodiversity Journal, 2015, 6 (1): 7-10
About the presence of the snow vole, Chionomys nivalis
(Martins, 1 842) (Mammalia Rodentia Cricetidae),in Calabria,
Southern Italy: data review and critical considerations
Armando Nappi 1 & Gaetano Aloise 2
1 M useo civico di Storia naturale, via Cortivacci 2 , 23017 M orbegno, Sondrio, Italy; e-mail: armando.nappi@ alice.it
! M useo di Storia Naturale della Calabria e O rto Botanico. Universita della Calabria, Via P. Bucci s.n., 8 7036 Rende, Cosenza,
Italy; e-mail: g aetan o .alo ise @ unical.it
Corresponding author
ABSTRACT The presence of CllioVlOlTiyS Tlivulis (M artin s , 1 842) (Mammalia Rodentia Cricetidae) in
Calabria, the southern tip of the Italian peninsula, is reported in different literature sources,
but the only Calabrian specimen, from Lago Cecita, Cosenza district, is preserved into M useo
Zoologico “La Specola”, Firenze. A recent examination of this specimen, moreover, has
shown that it is an Arvicold CllfiphibiuS (Linnaeus, 1 75 8 ) juvenile. The distribution of C.
nivalis along the Apennines, requires adequate insights and critical reviews.
KEY WORDS a P ennines; Calabria; ChiOflOmyS nivalis ; distribution.
Received 19.11.2014; accepted 26.0 1.2 0 15; printed 30.03.2015
INTRODUCTION
The snow vole Chionomys TlivCllis (Martins,
1 842) (Rodentia Cricetidae), is a species wide-
spread from south-western Europe through south-
eastern Europe to the W Caucasus, east to Turkey,
Israel, Lebanon, W Syria, W and N Iran and S Turk-
menistan (Musser & Carleton, 2005). In Italy it is
present continuously along the Alps, while the dis-
tribution area of the Apennines is more fragmented
(Amori, 2008). Moreover, on the latter portion,
some bibliographic data, such as few Abruzzo
mountains as well as the Matese Massif, between
Molise and Campania, should be confirmed by
more recent research (Nappi et al., 2007). Further
south, always along the Apennines, the question
about the snow vole presence in the Calabria region,
in the southern tip of the Italian peninsula, subject
of this note, is of particular interest. The presence
of the species in this region, in fact, was considered
uncertain and debated for nearly half a century.
From the literature search, the first value found
is contained in a generic work on mammals, where
snow vole is reported in Italy “ Sullc Alpi 6 SlilV
Appennino, sino alia Calabria' ’ (“on the Alps and
on the Apennines, until Calabria”), without further
details (Scortecci, 1 9 5 3 ). However, this species is
absent in other publications concerning Calabrian
mammals (Costa, 1 8 3 9, 1 845, 1 847, for a correct
dating of the issues of “Fauna del Regno di Napoli”
written by O. G. Costa, see D ’ Eras mo, 1949;
Moschella, 1900; Lucifero, 1 909; Pasa, 1 95 5 ) and
Toschi (1965), expresses some doubts about its
presence in the region.
Afterwards, in a study about some birds of prey
from Sila Grande, Cosenza, is reported the discovery
of the predation rem ains near a nest of huzza rd Buteo
bliteO Linnaeus, 1758, 25. V. 1971, consisting of
“ cranio frammentario, denti e peli di Microtino,
quasi certamente Microtus nivalis” (fragmentary
skull, teeth and hair of Microtine, almost certainly
Microtus nivalis ) , n o w synonymous with Chionomys
8
Armando Nappi & Gaetano Aloise
nivalis (D essi F ulgheri et al., 1 9 72). T his m aterial is
now lost and is no longer verifiable (P. Mirabelli,
p e r s . com.).
Afterwards, the presence of the species in the
reg io n , based on a m u seal specimen, is repo rte d b y
Amori et al. (1986). According to Amori (1993),
“ these sporadic records could be confirmed by fur-
ther and more specific research ” and in an other
review, Amori (1999) indicate the species distribu-
tion, in Italy, from the Alps until the central Apen-
nines. M ore recently, the presence of the snow vole
in Calabria, is reported in a mammals volume of
“Fauna d'ltalia” series (Amori, 2008) and in the
section of this species of the IUCN Red List
(Krystufek & Amori, 2008).
MATERIAL AND METHODS
In this paper, all literature data, that it was pos-
sible to find, were considered. In addition, the only
Calabrian specimen, known by writers, was ana-
lysed. This consist in a liquid preserved body, with
relative skull, into M useo Zoologico “La Specola”,
Firenze, n . MZUF-7448 (Cosenza, Lago Cecita,
18. VIII. 1970, Piero M annucci legit; head-body: 89
m m , t a i 1 : 5 7 m m ; ear: 10 m m ; h i n d f o o t : 2 2 mm).
Of this specimen, skull and teeth morphologies
were analyzed. Third upper molar and first lower
molar, in particular, were compared with the molar
morphotypes, identified by N adachow ski (1991),
just in the genus Chionomys M iller, 1 908.
RESULTS
A recent analysis of the specimen above men-
tioned, has shown that the tooth morphology (Fig.
1) is that typical of water vole Arvicola OITiphibiuS,
(Linnaeus, 1 7 5 8 ) as well as the skull morphology,
r datable to a young specimen of this species (Figs.
2 , 3 ) . A s confirmed, moreover, by comparison with
skulls of juvenile specimens of water vole (Museo
Civico di Storia N aturale di M ilano, nn. 1855, 1863,
1869, 1875, 4134; Museo di Storia Naturale, Uni-
versity della Calabria: nn. SG35, SG 147, AS 164;
Coll. Teriologica G. Aloise: n 615).
DISCUSSION
It seems that the snow vole has colonized the
southern areas of the Italian peninsula already during
very ancient times, perhaps the early middle Pleisto-
cene. This is suggested by a finding at the site of Not-
archirico, within Venosa Basin, Potenza district,
characterized by the presence of archaic elements
such as So rex c fr. runtoneiisis Hinton 1911 , Pliomys
episcopalis Bartolomei, 1 970 and Arvicola Can-
tianUS (Koenigswald 1973) (Sala, 1999). It is also
useful to remember, in this context, the fact that
among the small mammals, in Italy, Calabria is the
southern distribution limit of different species (see
Amori et al., 2008) some of which, of mountain en -
vironm ent, with disjoint areal as Driomys nitedula
Pallas, 1778 (Capizzi & Filippucci, 2008), or frag-
mented areal as Talpa Caeca S avi, 1 8 22 (Aloise &
Cagnin, 2003) and NeOmyS fodienS P enn ant, 177 1
(Aloise et al., 2005). T. Caeca and N . fadiens, pre-
viously considered distributed along the central and
northern Italy, but absent in the southern regions,
were found only recently. Taking into account these
assumptions, the presence of the snow vole, in Ca-
labria, cannot be excluded, even for the lack of re-
search in potentially suitable areas, such as, for
exam pie, P o llin o M as sif an d O rsom arso M o u n tain s .
Based on the results of the present work, and in
the absence of some objective evidence, snow vole
m ust be currently considered to be absent from this
region. On the other hand, until now, all research
related to small mammal fauna, have not yielded
positive results regarding the presence of the species
in Calabria (Lehmann, 1 96 1, 1 964, 1 9 73, 1 97 7;
Aloise et al., 1 98 5; Cagnin et al., 1 9 86; Aloise &
Cagnin, 1 987) and on the basis of more than 3.000
specimens of small mammals, collected over the
entire territory of Calabria during 1983-2013 (Coll.
Teriologica of the M useo di Storia N aturale of the
Universita della Calabria and Coll. Teriologica G.
Aloise), snow vole has never been found. With
regard to Lago Cecita area, alt hough it was also the
subject of several investigations that have provided
more than 300 specimens from traps and 84 speci-
mens fro m rap to rs pellets, C. nivalis has never been
found. It should also be noted that in this area there
are no environments suitable for the snow vole.
Moreover, the presence of water vole, just
around Lago Cecita, is supported by a liquid pre-
served specimen (n. SG 147, Cosenza district,
Spezzano Sila, Valle Capra, 18. VII. 1991) and by a
6 specimen from raptor pellets into Collezione
Teriologica of the Museo di Storia Naturale, Uni-
versita della Calabria, by a stuffed skin (n. 4020,
About the presence of the snow vole, Chionomys nivalis (Mammalia Rodentia Cricetidae), in Calabria, Southern Italy
9
Fig. i. Right first lower molar (left) and right third upper
molar (right) of the specimen MZUF-7448 determined in
this paper as Arvicola amphibius ( see Fig. 2 for details).
Figure 2, 3. Skulls (Fig. 2) and jaws (Fig. 3) in different view
of juvenile of Arvicold amphibius (left) (Calabria, C osenza,
Lago Cecita, 1 8 .V III. 1 9 70, P. Mannucci leg., Museo Zo-
ologico “La Specola”, Firenze, n. MZUF-7448) and
Chionomys nivalis (right) (Emilia Romagna, M odena, M onte
Cirnone, 19. IX. 1990, C. Bertarelli leg., Museo Civico di
Ecologia e Storia Naturale, Marano sul Panaro, n. 296).
14. VII. 1963) into Museo Civico di Storia Naturale
di Milano and by some observations (F. Pellegrino,
2012, pers. com.). The distribution of the snow
vole, in central-southern Italy, certainly requires
adequate deepenings but data about water vole, a
very decreasing species in Italy, recently no longer
found in different historical localities, are equally
interesting (Cagnin, 2008).
ACKNOWLEDGEMENTS
We wish like to thank Paolo Agnelli (Florence,
Italy), G iorgio B ardelli (M ilan, Italy), M ara C agnin
(Rende, Italy), Antonio Gelati (M ara no sul Panaro,
Italy), Michela Podesta (Milan, Italy) and Renzo
Rabacchi (Marano sul Panaro, Italy) who have
provided us some museum specimens, Andrea Maria
Paci ( C itta di Castello, Italy) for the help in the lit-
erature search, Gabrie 11a Bianchi and Livio Ciapponi
(M orb eg no , Italy ) for m a king po ssib le th e realization
of the iconography to one of the us (A.N.).
REFERENCES
Aloise G . & Cagnin M ., 1987. N uovi dati sulla corologia
di ale une entita rilevanti della m ic ro m am m alo fa un a
della Calabria. Hystrix, 2: 1-5.
Aloise G . & Cagnin M., 2003. New southern distribution
limit of Talpa caeca Savi, 1 822 (In s e c ti v o ra ,
Talpidae) in Italy. M ammalian Biology, 68: 235-238.
Aloise G . , Cagnin M. & Contoli L., 1985. Presence de
SoreX minutUS (L., 1 7 6 6 ) (Insectivora, Soricinae) sur
le massif de La Sila Grande (Calabre, Italie).
Mammalia, 49: 297-299.
Aloise G ., Amori G., Cagnin M. & Castiglia R., 2005.
New European southern distribution limit of NeomyS
fadienS (Pennant, 1771) (Insectivora, Soricidae).
Mammalian Biology, 70: 381-383.
Amori G., 1993. Italian Insectivores and Rodents: ex-
tinctions and current status. Supplemento alle Ricerche
di Biologia della Selvaggina, 21: 115-134.
Amori G ., 1999. ChiOHOmyS nivalis (M artins, 1 842). In:
Mitchell-Jones A.J., Amori G., Bogdanowicz W.,
Krystufek B., Reijnders P.J.H., S p itz e n b e rg e r F.,
Stubbe M., Thissen J.B.M., Vohralik V. & Zima J.
The Atlas of European Mammals. Academic Press,
London, 256-257.
Amori G ., 2008. Chionomys nivalis (M artins, 1 842). In:
Amori G ., Contoli L. & Nappi A. Mammalia II.
Erinaceomorpha, Soricom orpha, Lagomorpha,
10
Armando Nappi & Gaetano Aloise
Rodentia. Collana “Fauna d’ltalia”. Vol. XLIV.
Edizioni Calderini, M ilano, 465 - 474.
Amori G., Contoli L. & Nappi A., 2008. Mammalia II.
Erinaceom orpha, Soricomorpha, Lagomorpha,
Rodentia. Collana “Fauna d’ltalia’’. Vol. XLIV.
Edizioni Calderini, M ilano, 736 pp.
Amori G., Cristaldi M . & Contoli L., 1986. Sui Roditori
(Gliridae, Arvicolidae, M uridae) dell’Italia peninsu-
lare ed insulare in rapporto all’ambiente bioclimatico
mediterraneo.Animalia, 11 (1984): 217- 269.
Cagnin M ..Aloise G. & B isazza A 1986. Contributo alia
conoscenza ed all'in q u ad ra m e n to b io g eo g rafic o dei
micromammiferi della Sila Grande (Calabria, Italia).
B io g e o g rap h ia - Lavori della Societa Italiana di
B io g e o g rafia , 1 0 (1 984): 793- 803.
Cagnin M ., 2 0 0 8 . Arvicola amphibiuS (L in n a e u s , 1 75 8).
In: Amori G., Contoli L. & Nappi A. Mammalia II.
Erinaceom orpha, Soricomorpha, Lagomorpha,
Rodentia. Collana “Fauna d’ltalia”. Vol. XLIV.
Edizioni Calderini, M ilano, 445-458.
Capizzi D. & Filipp ucci M.G., 2008. DryomyS Ylitedlllci
(Pallas, 1778). In: Amori G . , Contoli L. & Nappi A.
Mammalia II. Erinaceom orpha, Soricomorpha,
Lagomorpha, Rodentia. Collana “Fauna d’ltalia”.
Vol. XLIV. Edizioni Calderini, Milano, 423 - 431.
Costa O .G ., 1839. Fauna del Regno di Napoli ossiaenu-
merazione di tutti gli animali che abitano le diverse
regioni di questo Regno e le acque che le bagnano
contenente la descrizione de’ nuovi o poco esatta-
mente conosciuti. Quadro delle specie indigene, ed
acclimatizzate della classe de’ M ammiferi. Stamperia
Azzolino e Compagno, Napoli.
CostaO.G., 1845. Fauna del Regno di Napoli ossiaenu-
merazione di tutti gli animali che abitano le diverse
regioni di questo Regno e le acque che le bagnano
contenente la descrizione de’ nuovi o poco esatta-
m ente conosciuti. Genere Arvicola ; Arvicola , Lacep.
Stamperia Azzolino e Compagno, Napoli.
Costa O .G ., 1847. Fauna del Regno di N apoli ossia enu-
merazione di tutti gli animali che abitano le diverse
regioni di questo Regno e le acque che le bagnano
contenente la descrizione de’ nuovi o poco esat-
tam ente conosciuti. Supple mento al catalogo de’
Mammiferi. Stamperia Azzolino e Compagno,
N apoli.
D’Erasmo G ., 1949. Le date di p u b b lie a z io n e della
“Fauna del Regno di Napoli” di Oronzio Gabriele
Costa e diAchille Costa. Rendiconti dell’Accademia
delle Scienze Fisiche e Matematiche (Societa reale
di Napoli), (serie 4), 6: 14-36.
Dessi Fulgheri F., M irabelli P. & Simonetta A.M ., 1972.
Osservazioni prelim inari sui falconidi della Sila
Grande. In: una vita per la natura. Scritti sulla con-
servazione della natura in on ore di Renzo Videsott
nel C in q u a n te n ario del Parco Nazionale del Gran
Paradiso. Edizione sotto l’elgida del W W F,
Came ri no, 141-153.
Krystufek B. & Amori G ., 2008. ChionomyS nivalis. In:
IUCN 2011. IUCN Red List of Threatened Species.
Version 2011.2. www.iucnredlist.org. Downloaded
15/07/2014.
Lehmann E . , 1961. U her die Klein sauger der La Sila
(Kalabrien).ZoologischerAnzeiger, 167: 213- 229.
Lehmann von E., 1964. Eine K le in sau g era u sb e u te vom
Aspromonte (Kalabrien). Beschaftigungen der
B erlinischen G esellschaft N aturforschender Freunde,
4: 37-48.
Lehmann von E., 1973. Die Saugetiere der H oclagen des
Monte Caramolo (Lucanischer Apennin, Nordka-
labrien). Supplemento alle ricerche diBiologia della
Selvaggina, 5: 48-70.
Lehmann von E., 1977. Erganzende Mitteilungen zur
Klein saugerfauna Kalabriens. Supplemento alle
ricerche di Biologia della Selvaggina, 5: 195-218.
Lucifero A., 1909. M ammalia Calabra. Elenco dei M am -
miferi calabresi. Rivista italiana di scienze Naturali,
Siena. Ristampa: 1 98 3. Frama Sud ed., Chiaravalle
Centrale.
Moschella G., 1900. I Mammiferi di Reggio Calabria.
Tip. F. M orello, Reggio Calabria.
Musser G .G . & Carleton M .D ., 2005. Superfamily
Muroidea: ChionomyS nivalis (Martins, 1 842). In:
W ilson D.E. & Reeder D.M . M ammal Species of the
World, Third Edition. The Johns Hopkins University
Press, 969-970.
Nadachowski A., 1991. Systematics, geographic vari-
ation, and evolution of snow voles ( ChionomyS )
based on dental characters. Acta T h e rio lo g ic a , 36:
1- 45.
NappiA.,BertarelliC.,De SanctisA.,Norante N.,PaciA.
M ., Ricci F. & Romano C ., 2007. D ati sulla dis-
tribuzione d ell’ arv ic o la delle nevi ChionomyS flivalis
(Martins, 1 842) (Mammalia, Rodentia, Cricetidae)
lungo l’Appennino c e n tro -s ette n trio n ale . A tti del
Convegno: Biogeografia dell’ A ppennino centrale e
se tten trio n ale trent’anni dopo. Parte II. Biogeographia,
28: 611-618.
Pasa A., 1 955. Ricerche zoologiche sula massiccio del
Pollino (L uc an ia-C alab ria). X M am m iferi. A nnuario
d e 11 ’ Is titu to e Museo di Zoologia dell’U niversita di
Napoli, 7: 1-7.
Sala B ., 1999. Nuovi dati sulla m ic ro te rio fa u n a di Not-
archirico. In: Piperno M . Notarchirico. Un sito del
Pleistocene medio iniziale nel bacino di Venosa,
Edizioni Osanna, 439-441.
Scortecci G., 1953. Animali. M am m iferi. Vol. 2. Edizioni
Labor, Milano.
Toschi A., 1 965. Mammalia. Lagomorpha, Rodentia,
Carnivora, Ungulata. Cetacea. Collana “Fauna d’
Italia”. VII. Edizioni Calderini, Bologna.
Biodiversity Journal, 2015, 6 (1): 11-16
Species composition of carabid communities (Coleoptera
Carabidae) in apple orchards and vineyards in Val d’Agri
(Basilicata, Italy)
Agostino Letardi*, Silvia Arnone, Massimo Cristofaro & Paola Nobili
ENEA, Unita Tecnica sviluppo sostenibile ed innovazione del sistema agroindustriale - Laboratorio gestione sostenibile degli
agro-ecosistemi - Via Anguillarese 3 0 1, 00123 Rome, Italy
Corresponding author, e-mail: agostino.letardi@ enea.it
ABSTRACT An entomological investigation was carried out in an agricultural area, mainly apple
orchards, of the Agri river plain, located in some municipalities ofBasilicata, Italy. Between
2012 and 2014, species richness and composition of carabid assemblages were investigated
on the ground surface of differently managed (abandoned, organic, commercial and IPM)
apple orchards and vineyards. Ground beetles (Coleoptera Carabidae) were sampled by
means of pitfall traps. 1 28 8 individuals belonging to 40 species were collected, representing
two-thirds of the carabid fauna of this area found in our and earlier studies. The species
richness varied between 4 and 20 in the different orchards. The common species, occurring
with high relative abundance in the individual orchards in decreasing order were: PterO-
stichus ( Feronidius) melas (Creutzer, 1 7 99), Pseudoophonus ( Pseudoophonus ) rufipes (De
Geer, 1 774), Brachinus crepitans (Linnaeus, 1 7 5 8), Harpalus ( Harpalus ) dimidiatus
(P. Rossi, 1790) and PoeciluS ( PoecUllS ) ClipreUS (Linnaeus, 1 75 8). Most of the collected
ground beetles were species with a wide distribution in the Paleartic region, eurytopic and
common in European agroecosystem s. The assemblages were dominated by small-medium,
macropterous species, with summer larvae. No endemic species were found.
KEYWORDS ground beetle; pitfall trapping; bioindicators; conservation; agro-ecosystem management.
Received 18.12.2014; accepted 26.01.2015; printed 30.03.2015
INTRODUCTION
In the frame of the ENEA project AGRIVAL
(aree AGRIcole ad alto VALore naturalistico
dell’alta val d’Agri = high nature value farmland in
upper Val d’Agri) (M enegoni et al., 2012, 2014), an
entomological investigation was carried out in an
agricultural area, apple orchards and vineyards, of
the Agri river plain, located in the municipalities of
M arsico V., Tramutola, Grumento N. and Viggiano
(Potenza, Basilicata, Italy).
The aim of this research was to investigate the
Carabid assemblages. In Europe several studies
gave faunal data on similar agro-ecosystems inhab-
iting carabids (Kutasi et al., 2004); these studies
indicate that variations in cultivation management
leads to variations in carabid beetle assemblages.
Although the spatial distribution of carabid beetles
may be primarily determined by microhabitat con-
ditions and biotic interactions at the local scale,
identifying general patterns of carabid responses to
different agro-ecosystem managements may help to
understand how species, functional groups and as-
semblages effectively distribute, and to predict how
they will cope with current and future land-use and
climatic changes (Brandmayr et al., 2011; Kotze et
12
Agostino Letardi et alii
al., 2011). In previous studies in Europe the fol-
lowing species were mentioned as common (Kutasi
et al., 2004): Pseudoophonus rufipes (De Geer,
17 7 4), Harpalus distinguendus (D u fts c h m id , 1 8 1 2 ) ,
Harpalus tardus (Panzer, 1796), Nebria brevicollis
(Fabricius 1 792), PterOStichuS melanarius (Illiger,
17 9 8), Poecdus cupreus (Linnaeus, 1758), Harpalus
affinis (Schrank, 1781).
MATERIAL AND METHODS
Five traps were activated for a week every
month in 12 sampling locations, from May until
October. Sampling started on July, 2012 and ended
on October, 2014. Locations have been selected
according to the type of crop (10 apple orchards
and 2 vineyards) and the managing practices
(traditional, integrated and organic) (Fig. 1, Table
1). Due to logistic unpredictable problems, some
of the locations selected at the beginning of the
experiment (samples Aa, Ab and Ac) were not
suitable anymore, and since 2013 were displaced
with analogous locations, respectively samples G,
H and L.
Ground beetles were sampled using plastic pit-
fall traps (500 ml and 100 mm the diameter of the
top) buried in the soil and filled with 50 ml salt
water. Pitfalls were covered with a 10 x 10 cm
plastic roof to prevent flooding.
The qualitative and quantitative data of the
carabid assemblages, recorded in the orchards of
the selected areas of Val d’Agri, were analyzed in
three different ways: 1) the weighted average of
different species in the total catch of the 12 samples;
2) the sum of the scores was calculated (where the
most abundant species collected in an orchard were
placed in decreasing order, and the dominant
species, with highest relative abundance scored 8,
the second one 7 etc.; the scores from different
orchards were summarised by species: the highest
possible score, if a species was dominant in all
orchards, was [12 x 8J 96); 3) the presence or ab-
sence of the species in the orchards was also in-
vestigated. The most widely distributed species
(which were found in 12 of the 12 investigated
orchards) got 100% ; the species, which was collec-
ted in 9 orchards, got 75% etc.
Carabids were identified to the species level, if
possible, following the nomenclature of Fauna
Figure 1. Locations of sampled farms: Val d’Agri (Basilica ta, Italy). Modified from AGEA 2011.
Carabid communities (Coleoptera Carabidae) in apple orchards and vineyards inVal d’Agri (Basilicata, Italy)
13
FARM
code
Lat. N
Long. E
destination
management
environment
New Ager
A a
40° 16’
15° 53’
apple orchard
conventional
agro-ecosystem
New Ager
Ab
40° 15’
15° 53’
apple orchard
IPM
agro-ecosystem
New Ager
A c
40° 16’
15° 53’
apple orchard
IPM
ecosystem
C ap uti
Ba
40° 17’
15° 49’
apple orchard
conventional
agro-ecosystem
C ap uti
Bb
40° 17’
15° 49’
apple orchard
conventional
near a ditch
F iorenti
C
40° 18’
15° 54’
apple orchard
abandoned
industrial zone
F iorenti
D
40° 18’
15° 54’
vineyard
conventional
agro-ecosystem
D o n z a
E
40° 19’
15° 49’
apple and pear orchard
abandoned
agro-ecosystem
P i s a n i
F
40° 20’
15° 50’
vineyard
organic
agro-ecosystem
T ropiano
G
40° 2 1 ’
15° 49’
apple orchard
conventional
agro-ecosystem
Padula
H
40° 17’
15° 52’
apple orchard
abandoned
agro-ecosystem
B osco G aldo
L
40° 20’
15° 50’
apple orchard
IPM
agro-ecosystem
Table 1. Localization of farms samples and some their characteristics (Val d’Agri, Basilicata, Italy).
Europaea (Vigna-Taglianti, 2013). Specimens, pre-
served in alcohol, are stored in the collection of the
ENEA Casaccia research centre.
RESULTS
Overall, 1,288 individuals have been collected
belonging to 40 carabid species which represent,
according to our elaboration of the available data
(Casale et al., 2006; Letardi et al., 2014a, b), two
thirds of the total carabid fauna reported for this
geographic area. The species richness of the invest-
igated carabid assemblages ranged between 4 and
20 in the different orchards: the weighted averages
of different species in each samples were not stat-
istically significantly different, nevertheless they
show an evident tendency to increase in terms of
biodiversity moving from conventional manage-
ment farms towards organic and semi-abandoned,
re-naturalized ones (Table 2). The relatively high
biodiversity value in the conventional managed
farm New Ager (Aa) could be an exception due to
the very few number of samples (just 4, all during
2012) collected: in 2013 and 2014 it was not pos-
sible to sample inside the New Ager farm, due to
technical logistic impediments.
Qualitative and quantitative data analyses have
been performed among the collected carabid
species following 3 methods: their proportion in the
total catch of the investigated orchards; the scoring
of the seven commonest species in the different
orchards (total scores) and their presence in the
orchards (distribution).
The most abundant species was PteWStichllS
melas (3 3%) followed by Pseudoophonus rufipes
(2 0 % ), Brachinus crepitans ( 14 %), Harpalus di-
midiatus (8%) and Poecilus cup reus (6%). The
species which dominated the carabid assemblages
(with the total scores) were PterOStichuS melas (80),
Pseudoophonus rufipes (7 9), Harpalus dimidiatus
(4 5), Poecilus cupreus (18), Brachinus crepitans
(14), Carabus rossii Dejean, 1 826 (1 1 ) and
Calathus fuscipes (Goeze, 1 777) (10) (Table 3).
Pterostichus melas and Pseudoophonus rufipes
were found in all investigated samples (100%),
Harpalus dimidiatus was found in the 75% of the
different habitats, Poecilus CUpreUS was found in
the 67%, Anchomenus dorsalis (Pontoppidan,
1 7 63) and CalatllUS sp. pr. montivagUS Dejean,
1831 were found in the 58%, while Amara sp . p r.
aenea (De Geer, 1774) and Nebria brevicollis
(Fabricius, 1 792) were also quite common (50%).
14
Agostino Letardi et alii
FARM
code
sam-
ples
average
stan-
dard
dev.
species
num-
ber
New A g er
A a
4
3.00
±1.83
6
New A g er
Ab
4
1 .50
±1.29
4
New A g er
A c
4
2.25
±1.26
5
C ap uti
B a
1 7
1 .65
±1.27
13
C ap uti
Bb
16
1 .75
±1.39
13
F iorenti
C
14
2.93
±1.69
16
F iorenti
D
1 7
2.00
±1.84
18
D onza
E
15
4.60
±2.32
18
Pisani
F
1 7
3.18
±1.70
20
T ropiano
G
12
1 .00
±1.28
10
Padula
H
1 3
2.92
±1.98
17
B osco G aldo
L
1 3
1 .38
±1.12
10
Table 2. Weighted average of species biodiversity.
It can be concluded that four species Pt6W-
stichus melas, Pseudoophonus rufipes, Harpalus
dimidiatus and Poecdus cupreus were among the
commonest species in the investigated samples in
respect of all three approaches.
DISCUSSION
Altogether, as a result of our investigations, 40
carabid species, representing about two-thirds of the
whole carabid fauna reported in this area in our and
previous studies (Casale et al., 2006), were found
in apple orchards and vineyards of the medium area
of the Agri river plain.
Most of the collected carabids, both in the whole
area and in each sample, were species with a wide
distribution in the Paleartic region, eurytopic and
common in European agroecosystems.
The assemblages were dominated by small-
medium, macropterous species, with summer 1
arvae; we didn’t find any endemism (Table 4).
species
Aa
Ab
Ac
Ba
Bb
c
D
E
F
G
H
L
Total
score
P. melas
+
14.3
7.3
41
6.2
21.2
62.6
41 .2
18.8
52.3
15.9
57.8
80
P. rufipes
46.2
28.6
87.8
6.4
87.7
50
10.1
7.4
12.5
8.1
5.7
15.6
79
H. dimidiatus
37.1
5.1
+
14.4
13.1
23.5
+
8.1
6.3
45
P. cupreus
43.1
+
+
+
10.3
+
7.2
+
1 8
B. crepitans
+
6.4
+
+
50.5
14
C. rossii
9
+
+
28.1
+
1 1
C. fuscipes
+
+
5.2
10.5
+
10
A. aenea
+
+
+
5.8
+
+
5
C. convexus
10.4
5
D. clypeatus
5.1
+
4
O. cribricollis
+
5.1
+
4
C. preslii
+
+
5.2
+
3
specimen n°
65
35
41
78
65
52
99
68
96
86
333
270
species n°
6
10
4
17
5
10
13
13
16
18
18
20
Table 3. Relative abundance (%) and the total scores of the most abundant carabid species.
Relative abundance lower than 5% were marked with +.
Carabid communities (Coleoptera Carabidae) in apple orchards and vineyards inVal d’Agri (Basilicata, Italy)
15
Aa
Bb
D
G
Brachinus crepitans
Nebria brevicollis
Trechus quadristriatus
Amara sp. pr. aenea
Calathus circumseptus
Poecilus cupreus
Calathus fuscipes
Nebria brevicollis
Pseudoophonus rufipes
E
Egadroma c fr. marginatum
Poecilus cupreus
Pterostichus melas
Amara sp.pr. aenea
Harpalus dimidiatus
Pseudoophonus rufipes
Pterostichus c fr. nigrita
Anchomenus dorsalis
Harpalus distinguendus
Pterostichus melas
Trechus quadristriatus
Brachinus crepitans
Calathus circumseptus
Ophonus cribricollis
Poecilus cupreus
Ab
C
Calathus sp. pr. montivagus
Pseudoophonus rufipes
Calathus fuscipes
Acinopus megacephalus
Carabus preslii
Pterostichus melas
Harpalus serripes
Anchomenus dorsalis
Carabus rossii
Pterostichus cfr. nigrita
Pseudoophonus rufipes
Brachinus crepitans
Carterus c fr. fulvipes
Pterostichus melas
Calathus cinctus
Chlaenius chrysocephalus
H
Calathus circumseptus
Ditomus clypeatus
Amara sp.pr. aenea
Ac
Calathus fuscipes
Drypta clentata
Brachinus crepitans
Acinopus megacephalus
Calathus sp. pr. montivagus
Harpalus dimidiatus
Calathus cinctus
Harpalus dimidiatus
Carabus convexus
Harpalus distinguendus
Calathus circumseptus
Poecilus cupreus
Carabus preslii
Nebria brevicollis
Calathus sp. pr. montivagus
Pseudoophonus rufipes
Carabus rossii
Ophonus sp.
Carabus preslii
Pterostichus melas
Cychrus italicus
Poecilus cupreus
Carabus rossii
Harpalus dimidiatus
Pseudoophonus rufipes
Cryptophonus tenebrosus
Ba
Pseudoophonus rufipes
Pterostichus melas
Ditomus clypeatus
Agonum sordidum
Pterostichus melas
Harpalus dimidiatus
Amara sp.pr. aenea
Pterostichus c f r. nigrita
F
Harpalus distinguendus
Anchomenus dorsalis
C arabidae sp . 1
Agonum sordidum
Harpalus serripes
Calathus sp.pr. montivagus
Amara sp.pr. aenea
Harpalus sp.
Carabus rossii
D
Anchomenus dorsalis
Ophonus cribricollis
Cryptophonus tenebrosus
Amara sp. pr. aenea
Brachinus sclopeta
Ophonus ( Metophonus ) sp.
Cymindis miliaris
Anchomenus dorsalis
Calathus cinctus
Pseudoophonus rufipes
Harpalus dimidiatus
Brachinus crepitans
Calathus fuscipes
Pterostichus melas
Harpalus sp.
Brady cellus cfr. verbasci
Calathus sp. pr. montivagus
Nebria brevicollis
Calathus cinctus
Carterus c fr. fulvipes
L
Pseudoophonus rufipes
Calathus fuscipes
Harpalus dimidiatus
Anchomenus dorsalis
Pterostichus melas
Calathus sp. pr. montivagus
Harpalus distinguendus
Calathus circumseptus
Pterostichus cfr. nigrita
Ccirterus c fr. fulvipes
Harpalus sp.
Calathus sp. pr. montivagus
Cryptophonus tenebrosus
Harpalus sp .pr. affinis
Carabus preslii
Bb
Cymindis miliaris
Lebia sp.
Carabus rossii
Agonum sordidum
Harpalus serripes
Nebria brevicollis
Harpalus dimidiatus
Anchomenus dorsalis
Nebria brevicollis
Olisthopus cfr. fuscatus
Poecilus cupreus
Brachinus sclopeta
Olisthopus cfr. f uscatus
Poecilus cupreus
Pseudoophonus rufipes
Cychrus italicus
Ophonus cribricollis
Pseudoophonus rufipes
Pterostichus melas
Harpalus dimidiatus
Poecilus cupreus
Pterostichus melas
C arabidae sp . 2
Harpalus serripes
Pseudoophonus rufipes
Trechus quadristriatus
Harpalus sp.
Pterostichus melas
C arabidae sp . 2
Table 3. Species collected in each locality (Val d’Agri, Basilicata, Italy).
The common species in agro-environments
investigated were the same as those usually found
in field crops and which can be considered as
“disturbance- tolerant” species.
The number of captures, qualitative and quantit-
ative data here reported have shown a clear
tendency to be more abundant moving from con-
ventional towards to organic managements, not sup-
ported by a solid statistical analysis; therefore
sampling more distributed in terms oftime and rep-
licates would be necessary to provide more suitable
data.
16
Agostino Letardi et alii
ACKNOWLEDGEMENTS
We wish to thank a large number of entomolo-
gical colleagues of a web forum (www.
entomologiitaliani.net; M. Agosti, L. Badoei,
S. Biondi, S. Cosimi, S. A. Degiovanni, F. Di
Giovanni, L. Forbicioni, G. Franzini, M. Gigli,
G. Giovagnoli, M. Grottolo, P. Leo, C. M anci, J.
Matejiecek, V. Monzini, M. Pavesi,A. Petrioli, N.
Pilon, R. Rattu, R. Sciaky, M . Selis and F. Turchetti)
for their help and suggestions; the owners of the
farms where the study took place; Camilla Nigro,
Piera Damiani and Giuseppe Sassano, of the ALSIA
station of Villa d’AGRI (Italy), for their great sup-
port in field work The study was carried out in the
framework of AGRIVAL Project funded by Italian
Government Grant according to Legge Finanziaria
20 1 0, A groalim entare art. 2 comma 44.
REFERENCES
Brandmayr P., Zetto T. & Pizzolotto R., 2005. I Coleotteri
Carabidi per la valutazione ambientale e la conser-
vazione della biodiversita. APAT, Manuali e linee
guida, 34. I.G.E.R. srl, Roma, 240 pp.
Casale A., Vigna Taglianti A., Brandmayr P. &
Colombetta G ., 2006. Insecta Coleoptera Carabidae
(Carabini, Cychrini, Trechini, Abacetini, Stomini,
Pterostichini). In: Ruffo S. & Stoch F. (Eds.), 2006.
Ckmap (Checklist and distribution of the italian
fauna). Memorie del M useo storia naturale Verona,
2. serie, sezione scienze della vita, 17: 159-164, with
data on CDrom.
Kotze DJ., Brandmayr P., Casale A., D auffy -R ich aid E.,
Dekoninck W., Koivula MJ., Lovei GL., Mossakow-
ski D ., Noordijk J., Paarmann W., Pizzolotto R.,
Saska P., Schwerk A., Serrano J., Szyszko J., Taboada
A., Turin H., Venn S ., Vermeulen R. & Zetto T., 201 1.
Forty years of carabid beetle research in Europe -
from taxonomy, biology, ecology and population
studies to b io ind ic atio n , habitat assessment and
conservation. In: Kotze DJ., Assmann T., Noordijk J.,
Turin H. & Vermeulen R. (Eds.), 201 1. Carabid Bee-
tles as Bioindicators: B iogeographical, Ecological
and Environmental Studies. ZooKeys 100: 55-148.
doi: 10. 3897/zookeys. 1 00.1 523
Kutasi C.S., M arko V. & Balog A., 2004. Species
composition of carabid (Coleoptera: Carabidae)
communities in apple and pear orchards in Hungary.
Acta Phy topathologica et Entomologica Hungarica
39: 71-89.
LetardiA., A r none S., Cristofaro M ., Nobili P., Damiani
P., Nigro C., Sassano G. & Menegoni P., 2014a.
C arabidofaune in meleti e vigneti in Val d’Agri. In:
Mannu R. (Ed.), 2014. Poster del XXIV Congresso
Nazionale Italiano di Entomologia, Orosei (Sarde-
gna), 9-14 Giugno 2014: 90.
LetardiA., A r none S., Cristofaro M ., Nobili P., Damiani
P., Sassano G., Nigro C. & Menegoni P., 2014b.
C arabidofauna per la valutazione di agroecosistem i
ad alto v a lore naturalistico: un caso studio in Val
d’Agri. In: Alba E., BenedettiA., Bucci G., Ciaccia
C., Pacucci C., Pinzari F. & Scarascia Mugnozza G.
(Eds.), 2014. Atti del X Congresso Nazionale sulla
Biodiversita. CNR (Roma, Italy) 3-5 Set 2014.
Abstract-book, Paper # c 7 . 1 5 . [online] URL:
http ://w ww. sisef.it/xbio/
Menegoni P., Iannetta M ., Giordano L., Iannilli V.,
Sighicelli M ., Colucci F., Tronci C ., Trotta C .,
Cristofaro M ., Letardi A., Rapagnani M .R., Arnone
S ., Musmeci S., Nobili P., Ponti L., Imperatrice A.,
Sassano G. & Damiani P., 2014. II progetto AGRI-
VAL: aree AGRIcole ad alto VALore naturalistico
dell’alta val d’Agri. In: Colucci F., Menegoni P.,
Trotta C., 2014. N atura 2000 in Basilicata: percorsi
di “ c o n tarn in azio n e” tra natura, scienza, arte e
cultura dei luoghi. Atti del Convegno di Aliano
(M a ter a), 4-6 aprile 20 13. ENE A : 13 1.
Menegoni P., Iannetta M ., Giordano L., Iannilli V.,
Sighicelli M ., Colucci F., Tronci C., Trotta C. &
Letardi A., 2012. II progetto AGRIVAL “Aree
agricole ad alto valore naturalistico: dalla in d i-
viduazione alia conservazione”, una visione del
rapporto tra agricoltura e biodiversita nella conver-
genza delle politiche agricole ed ambientali. IX
Convegno Nazionale sulla Biodiversita, Valenzano
(BA), 6-7 Settembre 20 12. 34.
Vigna-Taglianti A., 2013. Fauna Europea: Carabidae.
Fauna Europaea version 2.6. URL: http://www.
faunaeur.org
Biodiversity Journal, 2015, 6 (1): 17-26
Paleontologic and stratigraphic data from Quaternary de-
posits of Leghorn subsoil (Italy)
Andrea Guerrini 1 *, Alessandro Ciampalini 1 , Simone Da Prato 2 , Franco Sammartino 1 & Maurizio Forli 3
'Gruppo Archeologico e P ale o n to lo g ic o Livornese, Museo di Storia N aturale del M e d ite rra n e o . Via Roma 234, 57 1 27 Leghorn,
Italy; entail: agl8268@gmail.com
2 CNR-IGG Consiglio Nazionale delle Ricerche. Istituto di Geoscienze e Georisorse. Via Moruzzi 1. 56124 Pisa, Italy
’Societa Italiana di M alacologia, Via Galcianese 20H, 59100 Prato, Italy; e-mail: info@ dodoline. eu
* Corresponding author
ABSTRACT The Authors describe two malacofauna fossils attributable, on biostratigraphic and strati-
graphic base, to Pleistocene and Late Pleistocene, observed by a drilling carried out in the
east of the city of Leghorn, Italy. The malacological fossil association ofPleistocene was low
in number of individuals but well characterized in the number of species; the one attributable
to the Upper Pleistocene is related to contemporary associations already known in literature
for Leghorn subsoil, and shows two species not previously reported. The malacofauna of
the Lower Pleistocene is characteristic of the current coastal muddy debris; Tyrrhenian
malacofauna mainly consists of allochthonous elements, from a “Posidoilio, meadows” and
the deposition al environment is attributable to the Mediterranean current seabeds. The strati-
graphy of the subsoil of the area differs from that known in literature, as it shows a single
level of "Panchina" that rests directly above clay sediments of the Lower Pleistocene.
KEY WORDS M alacofauna; Stratigraphy; Pleistocene; Leghorn.
Received 24.0 1.20 1 5; accepted 25.02.20 1 5; printed 30.03.20 1 5
INTRODUCTION
The present study is part of a project examining
the malacofauna found in sediments forming the
subs tr ate of the city of Leghorn (Italy) which is not
always investigable directly. This is mainly because
of the closure, at the end of the nineteenth century,
of all the quarries, and the rapid development,
during the last century, of the city itself, which
resulted in the obliteration of the last outcrops. In
the second half of 1 900 were published a few pa-
pers of malacology, including one related to the
excavation of the dry dock of the “Torre del Fanale”
(Barsotti et al., 1 974). Therefore, the study of the
malacofauna of Leghorn subsoil can be carried out
only by analysing new successions, even within the
town limits, to refine the knowledge on biotic
fossils and compare them with those already known.
Recently, have been published data on two new
sections, one at the immediate northeast outskirts
of the town in locality “Vallin Buio” (Ciampalini
et al., 2014a), where the Tyrrhenian sediments rest
directly on those of the Pliocene, and the other one
from an excavation inside the town, near the section
studied herein, and called section “via Gram sci”
from its location (Ciampalini et al., 2014b), in
which is highlighted a malacofauna contained in the
top level of the "Panchina" formation.
The present paper describes the discovery of two
unpublished fossil malacofauna in deposits attrib-
Andrea Guerrini etalii
1 8
uted to Low er Pleistocene and the Upper Pleistocene.
These deposits were brought to light during the
execution of a geological drilling for the geotech-nical
characterization of the subsoil and herein named, sec-
tion “via M anasse” (Figs. 1,2). W ithin the sed irn en ts,
in addition to molluscs, were recovered ostracods and
foram inifera, whose study was carried out to assess
the c hron o s tratig rap h ic framework of the sediments
and their paleoenvironmental characterization.
Geological framework
In the subsoil of Leghorn, in a modest layer of
reddish sands (“Sabbie di D o n o r a tic o ” ) , there are
up to two calcarenitic sands (Panchina) which, on
the basis of stratigraphic and faunal evidence, are
generally attributable to the Tyrrhenian (M alatesta,
1942; Barsotti et al., 1974; Ciampalini et al., 2006).
The levels of "Panchina" belong to a morpholo-
gical element known in the literature as "Terrazzo
di Livorno" (Barsotti et al. , 1974; Lazzarotto et al.,
1 990), interpreted as a polycyclic marine terrace
(Federici& Mazzanti, 1995) that developed during
the stage 5 of the marine isotope stratigraphy
("marine isotope stage 5") (Chen et al., 1991;
Antonioli et al., 1999). The subs tr ate of this ter race
consists of sediments related to the Lower Pleisto-
cene that, based on fossil remains found on several
occasions, were attributed to the “Formazione di
Morrona” (Bossio et al., 1981; Dall'Antonia et al.,
2004; Boschian et al., 2006).
MATERIAL AND METHODS
The sediments analyzed originate from a
drilling, the location of which is shown in figure 1,
carried out for the geological and geotechnical
assessment of the subsoil (Fig. 2). The fossil shells
were collected directly from sediments or after
washing them. Considering the small amount of
sedimentary m ate rial available, it has not been pos-
sible to recover a large number of complete speci-
mens of large dimensions.
For measures of shells we used the following
abbreviations: d = m ax irn urn diarn eter; 1 = m ax irn urn
Simplified Description of Map Units
H olocene units
Upper Pleistocene continental units
Upper Pleistocene marine units
Middle Pleistocene continental units
Pliocene units
Miocene units
Ligurian units
LIGURIAN
SEA
LEGHORN
Via
Bacinodi carenaggio
i
Figure 1. Geological sketch map and location of the investigated borehole from “Via Manasse” (43°33'01” N-10°19 , 16”E).
Paleontologic and stratigraphic data from Quaternary deposits of Leghorn subsoil (Italy)
19
width; h = maximum height. The measurements are
in millimeters and the specimens figured are
numbered in Table 1.
Abbreviations used to indicate the marine biotic
communities are according to Peres & Picard
(1964): HP, biocenosis of photophilic algae; SGCF,
biocenosis of coarse sands with influence ofbottom
currents; VTC, biocenosis of coastal terrigenous
muds. The listofTyrrhenian molluscs found in this
study, reported in Table 1, is compared with those
of other locations recently studied, i.e. “Vallin
Buio” and “via Gramsci” (Ciampalini et al., 2014a,
2014b), and with that of the careening basin of the
“Torre del Fanale” which is, to date, the largest
excavation (Barsotti et al., 1 974).
The studied material is deposited, with the cata-
lognumberMSNM 827, at the Museum ofNatural
History of the Mediterranean in Leghorn.
Were weighed 150 g of anhydrous material,
employed for the m icro p alaeo n to lo g ic al analysis.
The samples were disgregated in water at 100 °C,
filtered in sieves with meshes netof 74 pm and then
dried in oven at 70 °C . The m ic ro p ala e o n to lo g ic al
analysis was conducted primarily on foraminifera
and ostracods. As for the b io s tra tig r a p h ic and pa-
laeoecological appearance of identified taxa, ref-
erence is made to the main available papers
(Ruggieri, 1973; Dali 'Antonia et al., 2004; Guernet,
2005; Faranda & Gliozzi, 2008).
RESULTS
In the sequence under consider ation, starting
from ground level, have been recognized seven
lithological intervals listed below, from which three
samples, indicated with the abbreviations MAN1,
M AN2 and M AN 3, respectively, were collected, at
different elevations (Fig. 3), for m icrop alae on to -
logical analysis.
Interval 1 (0-0.80 m): dark brown sands, with
nodules of Mn. In the first 10-15 cm, dark in color
due to the presence of coal, are visible brick frag-
ments that indicate a very recent age;
Interval 2 (0.80-1.10 m): sand with fragments of
wood and sand with gravel;
Interval 3 (1.10-3.80 m): ocher sandy silt with
Fe-M n nodules and rare levels of gravel. This level
becomes darker in the upper part;
Fig. 2. Core photograph, showing the first 5 meters from
ground level; the arrow indicates the portion of the borehole
where it was found the Tyrrhenian malacofauna described
Current Soil
Donoratieo Sands
Rio Maggione
Conglomerates
Castiglionccllo
Calcarenites
(Panchina I)
Monona Formation
(5) Marine molluscs
i-J Samples
Man!
Tt 1 T M 1 T ’
a S G
Figure 3. Stratigraphic column of the borehole from "Via
Manasse”; sample position is shown; a - Shales, S - Sands,
G - G ravels
Interval 4 (3.80-4.50 m): beige calcareous sand,
fossiliferous. From this level the sample MAN1
was taken ;
Interval 5 (4. 5-4. 7 m): gray-blue sand. From it
the sample MAN2 was taken;
Interval6 (4.7-10 m): gray blue clayish siltwith
sandy fossiliferous levels; at 6 m of depth is present
a decimeters level of gravel ocher in color. From
this range the sample MAN3 was taken;
Interval 7 (10-20 m): gray bluish clay silt with
rare fossils.
20
Andrea Guerrini etalii
By m icrop alae o n to lo g ic al analysis conducted on
the sample MAN1, taken within the interval 4, it
was observed a m alacofauna consisting of few
individuals, in good conditions, representing fifteen
species of gastropods and four bivalves; the full list
is shown in Table 1. By comparison of the species
observed with those reported for coeval sections
recently described, and those listed for the larger
“Bacino di Carenaggio” (see Barsotti et al., 1974;
Ciampalini et al. , 2014a, 2014b), it appears the
presence of two species not previously reported, i.e.
Gibbula turbinoides (Deshayes, 1835) and Fusinus
pulchelluS (Philippi, 1 844).
Within Foram inifera, the conservation status
varies widely, from good to bad; frequently, indi-
viduals of Elphidium crispum (Linnaeus, 1 75 8 )
were found associated with common represent-
atives of Elphidium spp., Ammonia parkinsoniana
(d'Orbigny, 1 8 39 ), Ammonia beccarii (L innaeus,
1 75 8), Ammonia spp. The residue of the washing
consists of a medium coarse sand, white in color.
Granules are formed by lithics, quartz and fossil
fragments (molluscs, echinoids, foram inifera) very
elaborate, subspherical and with traces of erosion.
Noteworthy, part of the components of the sand are
cemented forming agglomerates.
The analysis conducted on the sample MAN2,
from interval 5, allowed to recognize a rich and
diversified association of frequent specimens of
Loxoconcha SubmgOSa{R\iggieri, 1 977), with spe-
cimens o f Aurilalanceaeformis {UUczny, 19 69), A.
convexa (Baird, 1850), Pterigo cythereis, Cytherop-
teTOn Sulcatum (Bonaduce, Ciampo et Masoli,
1 9 7 6), C. latum (Muller, 1 894), Paracytheridea c f .
hexalpha (D oruk, 1 9 8 0 ) , and represen tativ es o f the
rare species Cimbaurila cimbaefarmis { Seguenza,
1 8 83 ), among the ostracods. Moreover, specimens
belonging to the Foraminifera genera Elphidium ,
Ammonia , Dorothia , Cassidulina have been collec-
ted. The residue of washing is made up of a sand
with a grain size ranging from fine to coarse and
gray in color. The granules are complex, mainly
made of lithic s .
The sample MAN3, from the interval 6, com-
prises a fossil association rich either in number of
specimens or in number of species. Were found nu-
merous specimens of LoXOCOUCha SubvUgOSa, and
valves belonging to the genera Auvila , PtCVigO-
cythereis, Cytheropteron , Bosquetina , Buntonia
(Ostracoda); specimens belonging to the genera
Elphidium, Ammonia, Dorothia, Cassidulina and to
the species Hyalinea baltic (Schroeter, 1 78 3) (Fo-
raminifera) were collected. In addition, frequent
remains of echinoids, bryozoans and molluscs were
observed. Among these have been identified the
species Turritella tricarinata (b roc chi, 1 8 1 4 ) (Fig.
4 8), Nassarius gigantulus (B ellardi, 1 8 8 2 ) (Figs. 49,
50) and Corbula gibba (O livi, 1 792) (Figs. 5 1, 52).
The residue of washing is a gray sand with a
particle size ranging from fine to coarse. Sand
grains are mainly composed of lithics and fossil
r e m a in s .
DISCUSSION
The result of the micropalaeon to logical ana ly sis
of the samples MAN2 and MAN3, from the
intervals 5 and 6, indicate associations typical of a
marine environment of the shallower part of the
internal platform. Furthermore, the discovery of
significant species as Aurila lance aefarmis,
Cimbaurila cimbaefarmis, Loxoconcha subrugosa,
Hyalinea baltica allow to attribute the lower part
of the succession to the lower Pleistocene, in par-
ticular to the “Calabriano p . p . (Emiliano)”, in
agreement with the presence of the gastropod
NaSSariuS gigantulus that disappears at the top of
the “Emiliano” (Ragaini et al., 2007).
In levels 5 and 6 have been identified, as the
most significant species, Turritella tricarinata,
Nassarius gigantulus and Corbula gibba which fo r m
a common association in sediments of the Lower
Pleistocene attributable to the VTC biocenosis.
Lithological characteristics and micro - macropa-
leontologic associations, allow to report the litholo-
gical interval 4 to the formation known as
"Panchina", widespread in the area, and attributed to
the Late Pleistocene (Tyrrhenian). Fossil Mollusca
referable to the Tyrrhenian, as P ersististrombus latUS
(Gmelin, 1791) and CoUUS emineUS (B o r n , 1 778),
have been reported by Malatesta (1 942) in a sand
fro m th e bo tto m of a we 11 dug in the Hospital of Leg-
horn, in the vicinity of the succession under study.
The small number of bivalves (only four
species) found in this interval is not significant
itself, but, taking into account the fifteen species of
gastropods in common with the other sections
already known in literature, and considering their
sediments, the Tyrrhenian fossil association is
Paleontologic and stratigraphic data from Quaternary deposits of Leghorn subsoil (Italy)
2 1
GASTROPODA Cuvier, 17 95
via Manasse
via Gramsci
(C iam palini et
al., 20 14a)
Vallin Buio
(C iam palini et
al., 2014a)
Bacino care-
naggio (B arsot-
ti et al., 1 9 7 4)
Figures
Tectura virginea to . f. m tiller, 1 7 76)
*
*
4-6
Jujubinus exasperatus (Pennant, 1777 )
*
*
*
*
7, 8
Gibbllla turbinoides (Deshayes, 1832)
*
9-11
Bolma rugosa ( L in n e , 1767)
*
*
*
*
1 2
Tricolia tenuis (Michaud, 1829)
*
*
*
13, 14
Bittium reticulatum (Da Costa, 1 7 7 8 )
*
*
*
*
15-18
Bittium latreillii (Payraudeau, 1 8 2 6)
*
*
19,20
Cerithium vulgatum b rug uiere , 1792
*
*
*
*
Rissoa variabilis (Won Muhifeidt, 1 8 2 4 )
*
*
2 1,22
Rissoa sp.
*
23
Alvania mamillata r is s 0 , 1 8 2 6
*
*
*
24,25
Alvania discors (Allan, 1 8 1 8 )
*
*
*
*
26,27
Alvania cimex (Linnaeus, 1 75 8)
*
*
28,29
Rissoina bruguieri (Payraudeau, 1826)
*
*
3 0, 31
Columbella rustica (Linnaeus, 1758)
*
*
*
32,33
FusinUS pulchellus (Philippi, 1 8 44)
*
34-36
Vexillum ebenus (Lamarck, 1811)
*
*
37,38
BIVALVIA Linnaeus, 1758
Striarca lactea (Linnaeus, 1758)
*
*
*
*
39-42
Glycymeris sp.
*
*
*
*
43,44
Chama gryphoides (Linnaeus, 1758)
*
*
46,47
Parvicardium exiguum (G m eiin, 1 79 1 )
*
*
*
*
45
Table 1. List of fossil molluscs from Tyrrhenian found in the survey of “via Manasse”, compared with
those found in sections of “via Gram sci”, “Vallin Buio and “Bacino di Carenaggio”.
compatible with a biocenosis of the SGCF type,
with specimens from the HP biocenosis. These data
confirm those already found in previous studies on
similar samples (Barsotti et al., 1974; Ciampalini et
a 1 . , 2014a; Ciampalini et al. 2014b) and what re-
ported by Corselli (1981) for the current seabed of
the Gulf of Baratti (LI ).
By comparing the lists of molluscs of the Upper
Pleistocene, relative to the locations of the territory
of Leghorn and reported in the above mentioned
papers (Table 1), it can be seen that the species in
common are nearly all, with the exception of RisSOCl
variabilis (Von Muhifeidt , 1 824) and Vexillum
ebeniiS (Lamarck, 1811), which are absent in the
deposits of “ via Gramsci” and “Vallin Buio”, but
present at the “Bacino di carenaggio”; RisSOitlCl
bruguieri (Payraudeau, 1 826) present in “ v ia G ram -
sci”, but absent in “Vallin Buio” and the “Bacino di
carenaggio” and, finally, Glbbulci tUvbinoidcS and
FusinUS pulchellus absent in all other locations.
As for the failure of a previous report of Bittiutfl
latreillii { Payraudeau, 1826), we assumed that the
22
Andrea Guerrini etalii
Figs. 4-6. Tectura virginea (O . F. M tiller, 1776)d = 3.3, h = 1.7. Figs. 7, 8. Jujubinus exasperatus (Pennant, 1777)d = 4.8, h =
6.5. Figs. 9-11. Gibbulu turbinoides (Deshayes, 1835) d = 3.9, h = 3.6. Fig. 12. BolfflCi TUgOSQ, (Linnaeus, 1767) operculum d
= 6. Figs. 13, 14. Tricolia tenuis (Michaud, 1 8 2 9) d = 4,h = 7.1. Figs. 15-18. BittUim reticuldtlim (Da Costa, 1778), Figs. 15, 16 :
d = 3.2, h =10.8; Figs. 17, 18: d = 2 .6 , h = 7.3. Figs. 19,20. Bittiuni lotveillH (Payrau deau, 1826) d = 2.4, h = 7.9.
Paleontologic and stratigraphic data from Quaternary deposits of Leghorn subsoil (Italy)
23
Figs. 21 , 22 . Rissoa variabilis (Von Miihlfeldt, 1 824) d = 2, h =3.9. Fig. 23. RisSOCl sp. d = 1.6, h = 4.2. Figs. 24, 25. Alvania
mamillata Risso, 1826 d = 3 . 2 , h = 5 . Figs. 26, 27 . Alvania discors (Allan, 18 18 ) d = 2 . 5 , h = 4; Figs. 28 , 29 . Alvania cimex
(Linnaeus, 1758) d = 2.3, h = 4.4; Figs. 30, 31. RisSOinCl bvugllieri (Payraudeau, 1826) d = 2.6, h = 6.8. Figs. 32, 33. Co-
lumbella rustica (Linnaeus, 1 75 8) d = 3.3, h = 4.9. Figs. 34-36. FusiriUS pulchelluS ( Philippi, 1 844) d = 2.0 m m , h =5.5 mm.
24
Andrea Guerrini etalii
Figs. 37, 38. Vexillum ebenus (Lamarck, 1811) d = 4, h = 8.2. Figs. 39-42 Striarca IdCtea (Linnaeus, 1 758), Figs. 39, 40: 1 = 4.9, h =
3.5; Figs. 41, 42: 1 = 6.2, h = 3.9. Figs. 43, 44. Gfycymeris sp. 1 = 4.1, h = 3.9; Fig. 45. Parvicardilim exigUUm ( Gmelin, 1791) 1 =
7.3, h = 6.3; Figs. 46, 47. Chama gryphoideS (Linnaeus, 1758)l=5.3,h = 4.7;Fig.48. Tlirritella tricarimta (B rocchi, 1 8 1 4) d = 4.8,
h = 1 1 ; Figs. 49, 5 0. NaSSCiriuS gigantulllS (B ellardi, 1882)d = 1 .3, h =2.5. Figs. 51,52. Corbuld gibba (Olivi, 1792)1= 9.4,h = 8.3.
Paleontologic and stratigraphic data from Quaternary deposits of Leghorn subsoil (Italy)
25
species may have been confused with B. F6ticU-
latum (Da Costa, 1778). Gibbula turbinoides and
Fusinus pulchellus , exclusive of the succession of
“via Manasse”, are compatible with the habitat of
“posidonieto” as one lives at low depth under the
rocks and on seagrass, and the other between the
Posidonia oceanica ( L.) Delile rhizomes.
The overall available data and m icropalaeonto-
logical analysis confirm, for the level of "Panchina",
a shallow marine habitat at high energy, with rocky
substrates alternating to sandy ones, in proximity of
or mixed to seagrass meadows. Even for the micro-
palaeontological association of the sample collected
in the limes to n e (interval 4), the conservation status
of the fossils and the structure of the granules of
sand confirm the hypothesis of the occurrence of a
marine environment at high energy.
Contrary to other Tyrrhenian fossilmalacofauna
found in the Leghorn area and atthe same s tratigra-
phic level, which are generally poorly preserved
(Ciampalini et al., 2014a, 2014b), the association
of “via Manasse” is composed of specimens little
eroded and often with traces of original coloring.
This, combined with the finer grain size of the
sediments, although still corroborating the hypothe-
sis of a marine environment at high energy, may
indicate a lower transport and consequently a
marine environment more stable and little deeper
than that assumed for the neighboring area of “via
Gramsci”, characterized, instead, by shoals of
"Panchina" quite compact.
The lithological study of the drilling revealed
peculiar stratigraphic characteristics partly different
from those previously reported (Barsottietal. 1974;
Ciampalini et al., 2006; Ciampalini et al., 2014b).
The drilling of “via M anasse” shows, in fact, only
one level of “Panchina” (lithological interval 4),
whereas former studies have often described two
levels, separated by a layer of clay and silt of
continental environment (Barsotti et al. 1 974;
Zanchetta et al., 2004; Ciampalini et al., 2006). On
the other hand, only one level has been observed in
the successions close to the ancient cliffs and at
higher altitudes. The presence of the Lower Pleisto-
cene, at the bottom of the drilling, enriches our
knowledge on the stratigraphy of the area that,
to day, is still poo rly known. As already repo r ted, in
stratigraphic levels below the "Panchina” it is pos-
sible to found sediments of both Pliocene and
Lower or M id die Pleistocene.
CONCLUSIONS
The lithological study of the drilling revealed
seven major lithological intervals. M icropalaeonto-
logical and lith o s tr atig r ap h ic analyses allowed to
attribute the intervals 7-5 to the Form ation of
Morrona (Lower Pleistocene, Calabrian) and the
lithological interval 4 to the Formation of the
“Calcareniti of C a s tig lio n c e llo ” (Upper Pleisto-
cene), also known as "Panchina".
The intervals 3 and 2 are attributed, on the basis
of observations on site, to the formation of con-
glomerates of Rio Maggiore, while interval 1 refers
to the “Sabbie di Donoratico”, present throughout
the Terrace of Leghorn. On the top of the drilling
was identified a soil rich in coal.
In the drilling analysed in this work, was ob-
served only one level of "Panchina" just above the
clay sediments of the Lower Pleistocene.
The analysis of malacofauna present in the
"Panchina" confirms data already known for an
advanced part of the Tyrrhenian cycle. Conversely,
no tropical molluscs, typical of the coasts of N W -
Centre Africa, and characteristic of the Tyrrhenian
baseline (MIS 5e), have been found. Since their
discovery was reported by Malatesta (1 942) at a
depth of about 6 meters (about 12 meters above sea
level), in a well near the present hospital and less
than one km away from “via Manasse”, this result
raises some questions. If for the section of “via
Gramsci”, adjacent to the hospital we could hypo-
thesize the failure in finding the African species
due to the shallow depth of the drilling, little more
than three meters, the same cannot be said for the
drilling of ’’via Manasse”, up to 20 meters above
the ground. At the depth of 6 meters from ground
level “via M anasse” sediments and malacofauna are
attributable to the Lower Pleistocene, sample
MAN3, while in the hospital area both sediments
and malacofauna are Tyrrhenian (Malatesta, 1 942).
The scantiness of recovered materials due to the
nature itself of the sampling carried out, i.e. only a
single drilling, could be the cause of the failure in
finding or recognizing the Tyrrhenian level M IS 5e.
Nevertheless, it is also possible that this level is not
always present in the subsoil of Leghorn.
On the other hand, the lack of the second cal-
carenitic level, probably eroded and replaced by
sediments of the river-type (Conglomerates of Rio
M aggiore), currently occurring above the fossil level,
26
Andrea Guerrini etalii
attests the possibility that important variations in the
stratigraphy of the terrace Leghorn have occurred.
In conclusion, it is not easy to find in the subsoil
of Leghorn the initial part of the transgressive
Tyrrhenian cycle and, consequently, to establish its
relationship with the underlying lithological units.
ACKNOWLEDGEMENTS
We thank Mr. Enrico Ulivi (Lastra a Signa, Flo-
rence, Italy) for the making photographs of
molluscs and Dr. Paolo Russo (Venice, Italy) for
confirmation of the determination of FliSiliUS
pulchellus.
REFERENCES
Antonioli F., Silenzi S., Vittori E. & Villani C ., 1 999.
Sea Level changes and tectonic mobility. Precise
measurements in three coastlines of Italy considered
stable during the last 125 ky. Physics and Chemistry
of the Earth, Part A: Solid Earth And Geodesy, 24:
337-342.
Barsotti G., Federici P.R., Giannelli L., Mazzanti R. &
S alvatorini G ., 1974. Studio del Qua tern ario Livornese,
con particolare riferimento alia stratigrafia ed alle
faune delle formazioni del bacino di carenaggio della
Torre del Fanale. Memorie della Societa Geologica
Italiana, 1 3 : 425-495 .
Boschian G., Bossio A., D a IF Antonia B. & Mazzanti R.,
2006. II Quaternario della Toscana costiera. Studi
c o s tieri, 12: 208.
Bossio A., Giannelli L., Mazzanti R., Mazzei R. &
Salvatorini G., 1981. Gli strati alti del Messiniano, il
passaggio M iocene-Pliocene e la sezione plio-
pleistocenica di Nugola nelle colli ne a NE dei Monti
Livornesi. IX Convegno. Societa Paleontologica
Italiana, (3-8 O ttobre , 1 9 8 1 ) : 55-90.
Chen J.FI ., Curran H .A ., White B. & Wasserburg G.J.,
1991. Precise chronology of the last interglacial
period: 234U-230Th data from fossil coral reefs in
the Bahamas. Bullettin of the Geological Society of
America, 103: 82-97.
CiampaliniA., Ciulli L., Sarti G. & Zanchetta G., 2006.
Nuovi dati geologici del sottosuolo del “Terrazzo di
Livorno”. Atti della Societa Toscana Scienze
Naturali, Memorie, Serie A, 111: 75-82.
Ciampalini A., Da Prato S., Catanzariti R., Colonese
A.C., Michelucci L. & Zanchetta G., 2013. Sub-
surface statigraphy ofmiddle pleistocene continental
deposits from Livorno area and b io s tra t ig ra p h ic
characterization of the Pleistocene substrate
(Tuscany, Italy). Atti della Societa Toscana di Scienze
Naturali, Memorie, Serie A, 120: 39-53,
CiampaliniA., F o rli M . , Guerrini A. & Sam m artino F.,
2014a. The marine fossil malacofauna in a Plio-
Pleistocenic section from Vallin Buio (Livorno,
Tuscany, Italy). Biodiversity Journal, 5: 9-18.
CiampaliniA., F o r 1 i M . , Guerrini A. & Sammartino F . ,
2014b. Una malacofauna tirreniana dal sottosuolo di
Livorno. Bollettino Malacologico, 50: 142-149.
Corse Hi C., 1981. La tanatocenosi di un fondo S. G. C.
F. Bollettino Malacologico, 17 : 1-26.
D all' A ntonia B ., C iam palin i A ., M ichelucci L ., Zanchetta
G. , Bossio A. & Bonadonna F.P., 2 0 04. New insights
on the Quaternary stratigraphy of the Livorno area as
deduced by borehole investigations. Bollettino della
Societa Paleontologica Italiana, 43: 14 1-157.
Faranda C. & Gliozzi E., 2008. The ostracod fauna of
the Plio-Pleistocene Monte Mario succession (Roma
Italy). Bollettino della Societa Paleontologica
Italiana, 47: 2 1 5-267.
Federici P. R ., & Mazzanti R., 1995. Note sulle pianure
costiere della Toscana. Memorie della Societa Geo-
grafica Italiana, 53: 1 65-270.
Guernet C., 2005. Ostracodes et stratigraphie du Neogene
et du Quaternaire mediterra-neens. Revue de Micro-
paleontologie, Paris, 48: 83-121.
Lazzarotto A., M azzanti R. & Nencini C., 1990. Geologia
e morfologia dei Comuni di Livorno e Collesalvetti.
Quaderni del Museo di Storia Naturale di Livorno,
11: 1-85.
Malatesta A., 1942. Le formazioni pleistocenice del
Livornese. Atti Societa Toscana Scienze Naturali.
Memorie, 51: 145-206.
Peres J. M. & Picard J., 1964. Nouveau manuel de
bionomie benthique de la Mer M ed iterranee . Recueil
des Travaux de la S tation m arine d'Endoume, 31: 1-13 7.
Ragaini L., Can tala messa G., Di C elm a C., Didaskalou
O., Impiccini P., Lori P., Marino M., Potetti M. &
Ragazzini S., 2007. First Emilian record of the
b o re a 1 -a f f in i ty bivalve PoTtlciVldicl ilTipreSSCl Perri.
1 9 75 from Montefiore dell’Aso (Marche, Italy).
Bollettino della Societa Paleontologica Italiana, 45:
227-234.
Ruggieri, G ., 1 973. Gli Ostracodi e la stratigrafia del
Pleistocene marino mediterraneo: Bollettino della
Societa Geologica Italiana, 92: 2 1 3-232.
Zanchetta G., Bonadonna F.P., CiampaliniA., Fallick A.
E., Leone G., Marcolini F. & Michelucci L., 2004.
In traty rrhen ian cooling event deduced by non - marine
mollusc assemblage at Villa S. Giorgio (Livorno,
Italy). Bollettino della Societa Paleontologica Italiana,
43: 347-359.
Biodiversity Journal, 2015, 6 (1): 27-40
Stream’s water quality and description of some aquatic
species of Coleoptera and Hemiptera (Insecta) in Littoral
Region of Cameroon
SimeonTchakonte 1 *, Gideon A. Ajeagah 1 , Nectaire Lie NyamsiTchatcho 1,3 , Adama Idrissa Camara 2 , Dramane
Diomande 2 & Pierre Ngassam 1
'Laboratory of Hydrobiology and Environment, Faculty of Science, University of Yaounde I, P.O. BOX 812 Yaounde, Cameroon.
2 Laboratory of Environment and Aquatic Biology, Nangui Abrogoua University, 02 P.O. BOX 801 Abidjan 02, Ivory Coast.
'Department of Aquatic Ecosystems Management, Institute of Fisheries and Aquatic Sciences at Yabassi, University of Douala,
P.O. Box 7236 Douala, Cameroon.
* Corresponding author, e-mail: tchakontesimeoon@yahoo.fr
ABSTRACT Aquatic insects are the dominant taxon group in most freshwater ecosystems and are particu-
larly suitable for large scale and comparative studies of freshwater community responses to
human-induced perturbations. Understanding these responses is crucial for establishing con-
servation goals. In this study, we used three families of aquatic insects (Coleoptera Gyrinidae,
Hemiptera Gerridae and Veliidae) as surrogates to measure the aquatic health of urban streams
in the city of Douala, and we described eight characteristic species. Aquatic insects were
sampled monthly over a 13-month period in two forested sites and ten urbanized sites.
Meanwhile, measurements of the environmental variables were taken. Overall, 20 species
were identified; the family Gerridae was the most diversified with 1 1 species, followed by
Veliidae (5 species), and Gyrinidae (4 species). All these species were present only at the two
forested sites; no species was found in the urbanized area all over the study period. Morpho-
logical description of the eight best indicator species ( Orectogyrus specularis Aube, 1838,
Orectogyrus sp.l, Orectogyrus sp. 2, Eurymetra manengolensis Hoberlandt, 1952, Eurymetra
sp. 1, Eurymetra sp. 2, Rhagovelia reitteri Reuter, 1884 and Rhagovelia sp.) revealed not
described characteristic features and potentially new species. This testified that in Cameroon,
biodiversity of aquatic insects is yet entirely to be investigated, and that there is an urgent need
in their taxonomic revision. Physicochemical analyses revealed the very poor health status of
urban streams with highly polluted water, while suburban streams have unpolluted water. The
results of redundancy analysis revealed that the presence of Gyrinidae, Gerridae and Veliidae
species is undoubtedly favored by the high rate of dissolved oxygen, important canopy cov-
erage and very low organic matter input. It is thus clear that polluted status of urban streams
due to human activities is the primary cause of the extinction of aquatic insect species.
KEY WORDS Aquatic Coleoptera and Hemiptera; morphological features; sensitive species; water pollution.
Received 31.01.2015; accepted 28.02.2015; printed 30.03.2015
INTRODUCTION
Climate change, loss of biodiversity and the
growth of an increasingly urban world population
are main challenges of this century (Muller et al.,
2010). In developing countries, urban population
and anarchic urban land use have dramatically in-
creased over the past few decades. Such population
28
Simeon Tchakonte etalii
growth and urban expansion are placing greater
stresses on the natural environment (Cohen, 2003),
leading to a strong variability on the physical and
chemical features of lotic ecosystems by clearing
riparian vegetation and opening canopy, increasing
inputs of sediments, nutrients, organic matter and
pollutants (i.e., heavy metals), altering flows and
reducing habitat heterogeneity (Xu et al., 2013;
Zhang et al., 2013). Such modifications result into
drastic changes in the biological component and the
ecological functioning of urban streams, with a
deterioration of water quality and loss of sensitive
aquatic biota (Tchakonte et al., 2014). There is
therefore a growing need to better understand and
predict how biotic communities respond to these
disturbances.
As an important functional group in stream eco-
systems that sustains the stability and complexity
of aquatic communities, insects have frequently
been used to indicate changes in the composition of
stream communities that respond to anthropogenic
disturbances since they are sensitive indicators of
long-term environmental changes in water and
habitat quality (Rosenberg & Resh, 1993; Song et
al., 2009; Zhang et al., 2013). Within the insects,
Ephemeroptera, Plecoptera and Trichoptera are well
known as good bioindicators in stream ecosystems
(Rosenberg & Resh, 1993; Foto Menbohan et al.,
2013; Nyamsi Tchatcho et al., 2014), whereas the
use of aquatic Coleoptera and Hemiptera in bio-
monitoring studies is rare. Despite their limited use
in stream biomonitoring, some aquatic Coleoptera
and Hemiptera taxa have been shown as being
sensitive to increase in sediment and organic pollu-
tion (e.g., Hauer & Resh, 1996; Zettel & Tran,
2004). Furthermore, most Hemipteran’s species are
endemic to particular islands or continental regions
and often have extremely limited distributions,
issuing them a bioindicator identity.
In the city of Douala which is the most densely
populated and industrialized area of Cameroon,
urbanization is anarchical with precarious sanitation
systems in shanty quarters; household disposals,
municipal and industrial wastewater and solid wastes
are discharged directly in the environment without
preliminary or adequate treatment (Tening et al.,
2013; Tchakonte et al., 2014).To our knowledge, no
study has so far dealt with diversity, morphological
description and ecological requirement conditions
of aquatic insect of the families Gyrinidae, Gerridae
and Veliidae in Douala rivers. Indeed, these Cole-
optera and Hemiptera accomplish their entire live
cycle in aquatic milieu (except pupal stage of Gyrin-
idae); they are therefore in permanent contact with
the aquatic environment and might reflect even the
most subtle changes occurring in the medium.
This study aimed thus to inventory and to describe
characteristic species of Gyrinidae, Gerridae and
Veliidae in urban and forest streams of Douala city,
in order to provide further information on the
systematic of these families and to offer hypotheses
as to how the species are distributed.
MATERIAL AND METHODS
Study area and sampling stations
Douala city is located at the bottom of the Gulf
of Guinea, along the estuary of the Wouri River.
This city extends between 3°58’- 4°07’ of latitude
North and between 9°34’ - 9°49’ of longitude East,
and presents a flat topography with altitudes
varying between 1.6 and 39 m (Olivry, 1986). The
climate of this region was classified by Suchel
(1972) as a wet tropical type, characterized by a
short dry season (December to February) and a long
rainy season (March to November). Rainfalls are
abundant and regular with the annual average
values varying between 2596 mm and 5328 mm.
The air temperature is relatively high with a monthly
average of approximately 28 °C (Suchel, 1972).
Samplings were carried out monthly, from Septem-
ber 2012 to September 2013 in 12 stations located
in the three larger contiguous watersheds (Nsape,
Tongo’a-Bassa and Mgoua) situated at the left bank
of the Wouri River (Fig. 1).
The watershed of Nsape is located in a peri-
urban area situated at about 30 km away from the
urban centre. This watershed is particularly covered
by vegetation of a secondary dense forest type,
composed of high trees, shrubs and tall grasses (un-
dergrowth) which alternate with some cleared spaces
used for traditional farming purposes. This forested
area is uninhabited and sheltered of any urban/
industrial activity. Two stations identified as N ] and
N 2 were selected in this forested area. Inversely,
Tongo'a-Bassa and Mgoua basins are located in
industrialized areas and are highly polluted by
human activities. Five sampling stations (Tj, T 2 ,
Stream’s water quality and description of aquatic species of Coleoptera and Hemiptera in Littoral Region of Cameroon 29
T 3 , ATj, and AT 2 ) were selected in the Tongo'
a-Bassa catchment. The stations Tj and T 2 are
localized respectively at 350 m upstream and 200
m downstream from the outlet of the effluent of an
industry of chocolate factory and confectionery.
Stations AT ^ and AT 2 are located respectively at
100 m and 3.5 km downstream from the outlet of
effluents coming from a brewery industry, a textile
industry and an industry of manufacture of glasses.
While station T 3 is situated at 500 m downstream
from the junction of the two preceding arms. The
five other stations (M 1? M 2 , AM 1? AM 2 and AM 3 )
were chosen in Mgoua river basin. The stations M 3
and M 2 are respectively localized at 350 m upstream
and 250 m downstream from the outlet of effluents
coming from the great Industrial Centre of Bassa.
Stations AM| and AM 2 are located respectively at
300 m upstream and 150 m downstream from the
outlet of the effluent of a soap and cosmetic factory,
while station AM 3 is situated at approximately 2.4
km downstream of this effluent.
Measurement of environmental variables
At each sampling station, 15 environmental
variables were taken into account. Three physical
parameters were determined to characterize the
habitat. The mean water depth (WD) was measured
30
Simeon Tchakonte etalii
on transects with equal distance interval across the
river sections (Song et al., 2009). Current velocity
(CV) was measured by timing the front of a neutral
non-pollutant dye (blue of methylene) over a
calibrated distance. At each sampling station,
canopy coverage (%) was estimated visually (Rios
& Bailey, 2006).
The measurements of physicochemical paramet-
ers of water at each sampling station were done fol-
lowing APHA (2009) and Rodier et al. (2009)
standard methods. Water temperature, pH, and dis-
solved oxygen (DO) were measured in situ using an
alcohol thermometer, a HACH HQ lid pH-meter,
and a HACH HQ14d oxymeter, respectively. Like-
wise, electrical conductivity (EC) was measured in
situ using a HACH HQ 14d conductimeter. Suspen-
ded solids (SS), turbidity, ammonium (NH 4 + ), ni-
trites (N0 2 ‘), nitrates (NO3'), and phosphates
(PO4 3 ) were measured in the laboratory using
HACH DR/2800 spectrophotometer. The Bioche-
mical Oxygen Demand (BOD5) was measured
using a Liebherr BOD analyzer. In order to assess
the organic pollution level at each sampling station,
the Organic Pollution Index (OPI) was calculated
according to the protocol described by Leclercq
(2001). OPI is based on three ions concentrations
resulting from organic pollution (NH 4 + , N0 2 ", and
P0 4 3 ") and one synthetic parameter (BOD5).
Sampling, identification and representatio-
nof aquatic insect species
Insect samples were collected at each station
using a long-handled kick net (30 cm x 30 cm side,
400 pm mesh-size, 50 cm depth). For each station,
samplings were done in a 100 m stretch following
protocol described by Stark et al. (2001). At each
station, 20 drags of the kick net were done in dif-
ferent micro-habitat, each corresponding to a
surface of 0.15 m 2 (30 cm x 50 cm). The materials
that were collected in the sampling net were rinsed
through a 400 pm sieve bucket and all macroinver-
tebrate individuals were sorted and preserved in
plastics sampling bottles with 70% ethanol. In the
laboratory, all aquatic insects belonging to the
families Gyrinidae (Coleoptera), Gerridae and
Veliidae (Hemiptera) were identified under a
stereomicroscope using appropriate taxonomic keys
(Dejoux et al., 1981; Durand & Leveque, 1981; De
Moor et al., 2003; Stals & De Moor, 2007; Tachet
et al., 2010), and counted.
The specimens intended for representation were
prior immersed in 10% sodium hydroxide
overnight, so as to soften their chitin and lighten
their body. The drawings of general morphology of
characteristic species were carried out under a
stereomicroscope equipped with a drawing tube.
Details of key appendages were drawn using an
optical microscope 100x magnification, equipped
with a drawing tube.
Data analyses
Insect richness, abundances and occurrence
frequencies were used to classify species according
to Dajoz (2000). In order to study the relationships
between environmental variables and the distribu-
tion and dynamic of the eight characteristic insect
species, Canonical Redundancy Analysis (RDA)
was performed based on the data matrix of species
abundances and physicochemical parameters. RDA
is a constrained ordination method, efficient in
directly revealing relationships between the spatial
structure of communities and environmental factors
that might be responsible for that structure (Le-
gendre et al., 2011). Monte Carlo permutations (499
permutations) were done so as to identify a subset
of measured environmental variables, which exer-
ted significant and independent influences on insect
species distribution at p < 0.05. CANOCO for
Windows 4.5 software (Ter Braak & Smilauer,
2002) was used for this analysis.
RESULTS
Environmental variables
The mean values and standard deviation (SD) of
environmental variables measured at each sampling
station are shown in Table 1 . The lower mean val-
ues of water temperature were observed at forested
sites (25.9° C), whereas at urbanized sites, higher
values were recorded, especially downstream from
the outlet of industrial effluents. The mean values
of pH varied between 6.10 (NQ and 8.16 (AM 2 ).
The percentage of dissolved oxygen was overall
higher at suburban sites (>75%) compared to urban
sites, where waters were closed to the hypoxic con-
dition, with mean values oscillating between 2.95%
(T3) and 21.3% (AMQ. Mean values of electrical
conductivity ranged between 13.1 pS/cm (N 2 ) and
Stream’s water quality and description of aquatic species of Coleoptera and Hemiptera in Littoral Region of Cameroon 3 1
1559 pS/cm (ATj). Turbidity and suspended solids
were globally veiy low at forested sites, with mean
values ranging froml4 to 26 NTU and from 4.2 to
7.7 mg/L, respectively. Where as in urban zone,
mean values of these parameters varied between
94.2 NTU ( AM { ) and 259.7 NTU (ATj), and
between 47.9 mg/L (AMj) and 163.5 mg/L (AT|),
respectively for turbidity and suspended solids. The
lowest mean values of nitrates (0.11 mg/L), nitrites
(0.006 mg/L), ammonium (0.09 mg/L) and phos-
phates (0.08 mg/L) were recorded at suburban sta-
tion N j , whereas the highest were registered at the
urban stations ATj (6.98 mg/L), AM 2 (0.26 mg/L),
M | (5.19 mg/L) and AT | (2.2 mg/L), respectively.
Concerning BOD, the lowest value (13.08 mg/L)
was observed in station N | , while the highest values
(218.1 mg/L) were obtained at station AT ] . Mean
values of water’s depth and current velocity fluc-
tuated between 0.22 m (M 3 ) and 0.75 m (AM 3 ), and
between 0.22 m/s (AM 3 ) and 0.89 m/s (ATj),
respectively. At the level of all the sampling stations
situated in urban area, canopy was absent; mean-
while it was estimated to 69% and 73% respectively
at the level of stations N 1 and N2 located in forested
sites. The organic pollution index (OPI) revealed
that organic pollution ranged from low to null at the
forested sites; whereas in urban streams, organic
pollution level was veiy high.
Composition and distribution of species
Overall, 20 species were identified for the three
studied aquatic insect families (Table 2). The family
Gerridae (Hemiptera) was the most diversified with
1 1 species, followed by the family Veliidae (Hemi-
ptera) with 5 species, and the family Gyrinidae (Co-
leoptera) which accounted 4 species. All these
species were caught only at the two forested sites
(N 1 and N2); no species were found at any of the
ten sampling stations located in urban streams, all
over the study period. Among the taxa identified 3
species of Gyrinidae ( Orectogyrus specularis Aube,
1838, Orectogyrus sp. 1 and Orectogyrus sp.2), 5
Variables
Forested sites
Urbanized sites
N,
N,
T,
7;
T
AT,
AT,
M,
M :
AM,
AM,
AMj
Temperature (°C)
Mean
25.9
25,9
29.4
29.2
30.1
32.7
30.54
29.1
29.54
28.8
29.23
29.4
SD
0.79
0.98
1.7
1.61
2.36
2.24
2.12
2.63
2.23
1.78
1.73
2.28
pH(UC)
Mean
6.10
6.19
7.01
6.98
6.72
7.93
6.71
6.92
6.79
6.75
8.16
7.01
SD
0.71
0,93
0,52
0.53
0.73
1.11
0.84
0.63
0.72
0.66
0.91
0.86
DO (%)
Mean
75.4
80.3
14.6
17.7
2,95
4.65
5.5
6.32
5.7
21.3
10,51
5.64
SD
83
10.9
10.!
11.8
1.66
2.47
3
8.73
5.89
12
7.86
6.62
EC (pS/cm)
Mean
132
13,1
397.9
401.1
4753
1559
624.2
403.1
677.5
291.3
478.9
443.7
SD
6.82
3.09
115
117
152
1159
277.5
148
636.6
79
417.1
170
Turbidity (NTU)
Mean
14
26
102.2
116.3
131.6
259,7
173
125.1
119.9
94.2
122.6
106.9
SD
10,5
17.2
108
71
66
187.4
100.6
40
69.6
18.7
86.59
60
SS (mg/L)
Mean
4.2
7,7
62.9
54.9
100.5
163.5
99.2
77
71.62
47,9
72.69
86,9
SD
3.8
4.09
101
85.5
84
112.3
66.1
39.7
27.3
25.9
41.72
72.7
NO/ (mg/L)
Mean
0.11
0,2
2.79
1.53
3.7
6.98
4.43
2.24
3.22
1.91
3.24
3.1
SD
0.16
0,17
5.26
0.58
2.92
4,9
332
2.14
1.49
1.18
3.99
1.3
NO/ (mg/L)
Mean
0,006
0.008
0.23
0.21
0.13
0.21
0.041
0.04
0.08
0.12
0.26
0.2
SD
0.0
0,0
0.46
0.39
0.26
0.31
0.025
0.06
0.1
0,15
0.65
0.29
NHf (mg/L)
Mean
0.09
0.1
4.5
4.56
4.4
4.04
3.12
5.19
4.15
3.1
Z49
3.29
PO/‘(mgfl.)
SD
0.07
0.08
2.53
2.73
2.3!
2.93
1.04
3.16
0.32
2.39
1.86
1.52
Mean
0.08
0.14
1.21
1.06
1.3
2.2
1.4
1.53
1.17
0.9
0.88
1.39
SD
0.13
0.2
0.41
0.38
0.58
1.73
0.69
0.91
0.73
0.43
0.37
0.45
BOD 5 (mg/L)
Mean
13.08
16.2
96.9
156.2
158.5
218.1
176.2
89.6
105
94.6
121.15
139.6
SD
6,9
10,2
50
53
67
63,23
46.24
31.3
27.39
29
22
33
WD (m)
Mean
0.31
0,66
0.37
0.48
0.65
0.26
0,62
0.22
0.43
0.35
0.26
0.75
SD
0.07
005
0.07
0.06
0.04
0.04
0.05
0.03
0.05
0.06
0,05
0.03
CV (m/s)
Mean
0,64
0,48
0.75
0,78
0.57
0.89
0.71
0.38
0.37
0.46
0.45
0.22
SD
0.04
0.05
0.04
0,05
0.04
0.03
0.03
0.03
0.05
0.05
0,04
0,05
OPI Values
Mean
4.63
4.14
1.67
1.73
1.69
1.48
1.81
1.81
1.87
1.98
1.94
1.77
SD
0.53
0.55
0.30
0.26
0.31
0.37
0.25
0.37
0.32
0.31
0.34
0.40
Pollution level
Null
Low
Very high
Table 1. Mean values and standard deviation (SD) of environmental variables measured
at each sampling station during the study period.
32
Simeon Tchakonte etalii
species of Gerridae ( Eurymetra manengolensis
Hoberlandt, 1952, Eurymetra sp. 1, Eurymetra sp.
2, Limnogonus chopardi Poisson, 1941 and Limno-
gonus sp.) and 3 species of Veliidae (Microvelia sp.,
Rhagovelia reitteri Reuter, 1884 and Rhagovelia
sp.) were present simultaneously at the two sub-
urban stations and are considered as characteristic
species. Each of the other species was caught either
at the station N| or at the station N 2 , exclusively.
Morphological description of some charac-
teristic species
Description of species of the genus Orecto-
GYRUS REGIMBART, 1884 (COLEOPTERA GYRINIDAE).
The aquatic insects of the family Gyrinidae are all
holometabolous. Adult Gyrinidae (whirligig beetles)
are highly adapted to the aquatic environment,
being the only beetles that normally use the water
surface film for support. They are, however, equally
at home under the water. Both adults and larvae of
all Gyrinidae are strictly aquatic. The adult gyrinids
are true water beetles with medium-sized to moder-
ately large, ranging from 4-17 mm in length. The
body shape of the adults is ovate or elongate-ovate,
convex, with a sharp lateral edge around the whole
body. This edge separates the hydrofuge dorsal
surface of the insect from its wettable ventral
surface. The lateral edge divides the compound eyes
into dorsal and ventral halves (Fig. 7), with the
dorsal part looking up out of the water, whereas the
ventral part looks down into the water. The antennae
of adult gyrinids are short, stout and highly sens-
itive (Fig. 5). The front legs of gyrinids are long and
adapted for seizing prey. The middle and hind legs
are adapted for swimming: they are shot and dorsov-
entrally compressed, with fringes of swimming
hairs (Fig. 6).
The adult specimens of the genus Orectogyrus
are recognized with their elongate-ovate-convex
body and their last abdominal segment elongate
extending beyond the elytra edge.The upper side of
ORDERS/FAMILIES
SPECIES
Nl
n 2
All urban
stations
COLEOPTERA GYRINIDAE
Aulonogyrus sp.
3*
-
-
Orectogyrus specular is Aube, 1838
9 *
6 *
-
Orectogyrus sp. 1
8 *
15*
-
Orectogyrus sp.2
27 **
19*
-
HEMIPTERA GERRIDAE
Aquarius distanti Horvath 1 899
-
2 *
-
Eurymetra manengolensis Hoberlandt, 1952
-
Eurymetra sp. 1
18*
13*
-
Eurymetra sp.2
4 *
6 *
-
Gerris swakopensis Stal, 1858
-
3*
-
Gerris sp.
-
2 *
-
Hynesionella aethiopica Poisson, 1949
-
2 *
-
Limnogonus chopardi Poisson, 1941
9**
7*
-
Limnogonus sp.
2 *
g**
-
Neo gerris sp.
3*
-
-
Tenagogonus sp.
-
5*
-
HEMIPTERA VELIIDAE
Carayonella hutchinsoni Poisson, 1948
1 *
-
-
Microvelia gracillima Reuter, 1882
2 *
-
-
Microvelia sp.
12 **
6 *
-
Rhagovelia reitteri Reuter, 1884
162***
90**
-
Rhagovelia sp.
31**
42**
-
Table 2. Distribution, abundances and occurrence frequencies of insect species of the families Gyrinidae, Gerridae and Ve-
liidae in different sampling stations; * = rare, ** = accessory, *** = frequent, (-) = absent. For undefined species, the de-
scriptor author’s name of the genus is given.
Stream’s water quality and description of aquatic species of Coleoptera and Hemiptera in Littoral Region of Cameroon 33
elytra is glabrous or (partly) pubescent, black in
color, with distinct metallic shiny portions used as
systematic character. The specimens that we recor-
ded have a pale yellow lateral border on the pro-
notum and elytra. The hindmost two abdominal
sternites are more-or-less laterally compressed and
movable, with a ventral median row of long hairs
used as a rudder for swimming. Three species,
Orecto gyrus specularis Aube, 1838, Orectogyrus
sp. 1 and Orectogyrus sp. 2 were identified. In O.
sp. 1 (Fig. 4), only the last abdominal segment is
extended beyond the elytra, whereas in O. specu-
laris and O. sp. 2 (Figs. 2, 3), it is the two last ones.
Moreover, the ornateness of elytra permitted to
clearly separate these three species. The specimens
of O. specularis measure 9.4 ±0.1 mm in length
and 4. 1 6 ± 0.02 mm in width; the inter-ocular space
measures 1.6 ± 0.001 mm and the pronotum is 1.2
± 0.02 mm in length. For O. sp.l, the body is 7.08
±0.11 mm in length and 3.6 ± 0.01 mm in width;
the inter-ocular space measures 1.12 ± 0.002 mm
and the pronotum is 0.92 ± 0.08 mm in length.
Concerning O. sp. 2, the specimens caught measure
9.54 ± 0.26 mm in length and 4.2 ± 0.02 mm in
width; the mean length of the inter-ocular space is
1.4 ± 0.01 mm and the pronotum is 1.2 ± 0.04 mm
in length. The median silky swimming hairs of the
last abdominal stemite are longer in O. specularis
and O. sp. 2 (800 - 850 pm) as compared to O. sp. 1
(520-680 pm).
Description of species of the genus Rhagove-
lia Mayr, 1865 (Hemiptera Veliidae). The Veliidae
are hemimetabolous insects gliding or treading on
the surface of the water. They are typically charac-
terized by their jointed mouthparts, modified to form
a rostrum or “beak”, which is adapted for piercing and
sucking. The head is short, less than two times longer
than wide, bent downward, and triangular. It has a
distinct longitudinal sulcus mid-dorsally, a jointed
rostrum with 3 segments, a pair of four-segmented
antennae longer than the head, and no ocellus. The
legs are nearly equidistant and the hind-coxae are
distinctly moved apart from each other, with mid-
femora not exceeding or very slightly the end of the
abdomen. The tarsal claws are subapical.
The adult specimens of the genus Rhagovelia
Mayr, 1865 are distinguishable with their body
surface matt and blackish or brownish; all tarsi
three-segmented with the basal segment very short;
mid-tarsi deeply cleft with leaf like claws and hairy
swimming fans arising from the base of the cleft
(Fig. 10). The mesoscutellum is not exposed, covered
by posteriorly-extended pronotal lobe. Two species,
Rhagovelia reitteri Reuter, 1884 and Rhagovelia sp.
were identified for this study.
Adult specimens of R. reitteri collected are mac-
ropterous and their fore-wings (hemelytra) are not
divided into corium and membrane (Fig. 8). How-
ever, these hemelytra can detach during sampling
or identification processes. The specimens of R.
reitteri measure 4.15 ± 0.15 mm in length and 1 .05
± 0.002 mm in width; the inter-ocular space
measures 0.24 ± 0.001 mm and the pronotum is
0.82 ± 0.02 mm in length. Inversely, the individuals
of Rhagovelia sp. are apterous, with stout hind
femora bearing small distinct spines on the inner
margins (Fig. 9). Their body is 4.12 ± 0.2 mm in
length and 1 .2 ± 0.01 mm in width; the inter-ocular
space measures 0.23 ± 0.01 mm. The lengths of the
pro-, meso- and metanotum are 0.81 ± 0.03 mm,
0.24 ± 0.001 mm and 0.24 ± 0.004 mm, respect-
ively. The body is mainly black; pronotum anteri-
orly completely yellow, posteriorly variably
colored, usually black, but in some specimens with
yellowish hind margin, and in smallest specimens
uniformly light reddish brown; connexiva (most
lateral areas of sternites and laterotergites) usually
brown, in smaller specimens yellow; anteclypeus,
rostrum, and proepisterna mainly yellowish;
antenna and legs mainly black, basal half of first
antemiomere, all coxae and trochanters, basal half
of all femora, inner margins of hind femora yellow.
Description of species of the genus Euryme-
tra Esaki, 1926 (Hemiptera Gerridae). The
Gerridae are hemimetabolous insects gliding or
treading on the surface of the water. They are typ-
ically characterized by their jointed mouth parts,
modified to form a rostrum or 'beak', which is
adapted for piercing and sucking. The head is short,
less than two times longer than wide, bent down-
ward, and triangular; it has no longitudinal sulcus,
a jointed rostrum with 4 segments, a pair of four-
segmented antennae longer than the head, and no
ocellus. The mid and hind-legs are distant from
fore-legs and longer than these formers, and their
femora are clearly extended beyond the abdomen.
The hind-coxae are distinctly moved apart from
each other.
34
Simeon Tchakonte etalii
1 mm
0.2mm
Femur
Elongte, grasping
from legs
Compound eyes
Clypeus
Pronotum
Scute Hum
Glabrous & shining
areas
Primrose yellow
Lateral borders
Elytral sutures
Short, flattened
swimming hind legs
Sharp lateral edges
Last abdominal segments elongate
l mm
Trochanter
Enlarged
pedicel
Scape
Tarsus
Tibia
Long, silky
swimming
hairs
0.5 mm
Divided compound eye
0.6mm
Lateral edge of the body
Figures 2-7. Orectogyrus adult. Morphology in dorsal view of O. specularis (Fig. 2), O. sp. 2 (Fig. 3) and
O. sp. 1 (Fig. 4); Fig. 5, antenna; Fig. 6, hind leg; Fig. 7, lateral view of the head showing divided compound eye.
Stream’s water quality and description of aquatic species of Coleoptera and Hemiptera in Littoral Region of Cameroon 35
The adult specimens of the genus Eurymetra
Esaki, 1926 are apterous and distinguishable with
their short, stout and rounded abdomen. Their body
is shiny and rounded, and generally does not exceed
4.5 mm in length. The meso and metanotum are
well distinct and separated by a lateral suture,
whereas the metasternum is reduced to a small
triangular plaque. All tarsi are two-segmented and
the tarsal claws are modified (straight or 'S'-shaped)
in some specimens. At the level of fore-tarsi,
segment 1 is shorter than segment 2 (Fig. 15), whe-
reas in the mid- and hind-tarsi, segment 1 is 3 to 4
times longer than segment 2 (Fig. 14). Three species,
Eurymetra manengolensis Hoberlandt, 1952, Eury-
metra sp. 1 and Emymetra sp. 2 were identified for
this study (Figs. 11-13). The specimens of E.
manengolensis and E. sp.l measure 4.1 ±0.1 mm
in length over 2.48 ± 0.04 mm in width, and 4.04 ±
0.12 mm in length over 2.76 ± 0.07 mm in width,
respectively. For E. manengolensis , the lengths of
the pro, meso- and metanotum are 0.23 ± 0.01 mm,
0.94 ± 0.006 mm and 0.24 ± 0.02 mm, respectively.
Where as in E. sp. 1, the pro-, meso- and metanotum
are 0.25 ± 0.02 mm, 0.94 ± 0.01 mm and 0.22 ±
0.04 mm in length, respectively. For these two
species, 8 abdominal segments are visible in dorsal
view; the edges of thoracic and abdominal tergites
are black in color; a mid-dorsal longitudinal band
is observed on the thoracic and the first two abdom-
inal tergites. Abdominal pleura are well developed
in E. manengolens as compared to E. sp. 1. Con-
cerning E. sp. 2, the individuals caught measure
3.11 ± 0.41 mm in length and 2.17 ± 0.02 mm in
width; 9 abdominal segments are visible in dorsal
view. Their pro-, meso- and metanotum measure
0.22 ± 0.02 mm, 0.97 ± 0.04 mm and 0.23 ± 0.02
mm in length, respectively. Their body is pale
yellow with median sulcus on thoracic tergites.
fore-wing
0.6 mm
Mid-tarsal
segment 3
Mid-tarsal
segment 2
Hairy swimming-
fans
Cleft 0.1 mm
Antennae
Hind-femora
Fore-legs
Longitudinal
sulcus
Compound eyes
Pronotum
Mesonotum
Metanotum
Mid-legs
Abdomen
Figures 8-10. Rhagovelia adult. Morphology in dorsal view of R. reitteri (Fig. 8),
R. sp. (Fig. 9) and detail of mid-tarsal fan (Fig. 10).
36
Simeon Tchakonte etalii
lateral
suture
1.3 mm
Fbia
0.53 mm
Femora
Trochanters
Coxae
Segment 1
Segment 2
Tibia
0.7 mm
Antennae
lore-legs
Compound eyes
Pronotum
Mesonotum
MctanoLum
Abdomen
Abdominal pleura
Mid-legs
Hind-legs
Lateral suture
.3mm
Tarsus
Claw
Segment 2
Segment 1
Taisus
Figures 11-15. Eurymetra adult. Morphology in dorsal view of E. manengolensis (Fig. 11), E. sp.l (Fig. 12)
and£. sp.2 (Fig. 13); Fig. 14, mid-leg; Fig. 15, fore-leg.
Stream’s water quality and description of aquatic species of Coleoptera and Hemiptera in Littoral Region of Cameroon 37
Relationships between environmental vari-
ables and insect species
The results of redundancy analysis (RDA)
revealed that the relationships between the 8
characteristic aquatic insect species and their habitat
conditions follow mainly the first two axes
(F 1=94.9 %; F2=3.6 %) which accounted for 98.5
% of the total variance expressed (Fig. 16). Follow-
ing the first axis (FI) in positive coordinates the
presence and abundances of the 8 characteristic
insect species {Orecto gyrus specularis, Orecto gyrus
sp. 1, Orectogyrus sp. 2, Rhagovelia reitteri,
Rhagovelia sp., Eurymetra manengolensis, Eury-
metra sp. 1 and Eurymetra sp. 2) are positively and
significantly influenced by water depth, high
dissolved oxygen content, important canopy cover-
age and higher values of OPI (i.e., very low organic
matter input). Rhagovelia sp. seems to quite appre-
ciate moderate water flow. Inversely, in negative
coordinates, the presence of these sensitive aquatic
insects is impeded by the polluted status of water
with high values of temperature, pH, turbidity,
electrical conductivity, suspended solids, am-
monium, nitrites, nitrates, phosphates and BOD.
DISCUSSION
This study achieved in Douala watershed
permitted to identify 20 species, all present only at
the two forested sites (N 1 and N2); no species being
found in urban streams. The absence of these
Figure 16. Redundancy analysis biplot showing gathering
of characteristic aquatic insect species in response to envir-
onmental variables; NH4 = ammonium, N02 = nitrites,
N03 = nitrates, and P04 = phosphates. See “Materials and
methods” section for other abbreviations.
Hemiptera (Gerridae Veliidae) and Coleoptera
(Gyrinidae) families in Douala’s urban waterways
is undoubtedly due to their polluted status caused
by the uncontrolled discharge of domestic, muni-
cipal and industrial wastes and sewages in the rivers.
Indeed, the hypoxic condition of water, the very
high values of water temperature, conductivity, tur-
bidity, suspended solids, diverse ions (N 03 ',NH 4 + ,
P0 4 3 '), organic matter input and BOD were re-
gistered at urban stations, and could have been re-
sponsible for the extinction of these aquatic insects.
These observations allowed us to assume that
species of these families might be sensitive to water
pollution and in-stream habitat degradation, since
we hypothesized that these taxa would have histor-
ically been present at these streams before urban-
ization, as they do in suburban streams. Similarly,
Foto Menbohan (2012) reported that species of
these aquatic insect families were absent (or very
rare) in the most of urban streams of the Mfoundi
river basin in Yaounde (Cameroon). Our results are
consistent with those of Compin & Cereghino
(2003) and Song et al. (2009) who showed that a
decrease in Coleoptera species richness in human-
impacted streams is clearly related to changes in
water quality and habitat suitability. Moreover,
aquatic Coleoptera species, especially those belong-
ing to Elmidae, Gyrinidae, and Haliplidae have also
been recognized as good water quality indicators
(Hilsenhoff, 1988; Bote et al., 2002; Sanchez-
Fernandez et al., 2006). Hauer & Resh (1996)
added that many species of these Coleoptera fam-
ilies have been shown to be sensitive to increase in
sediment and organic pollution. Concerning the
aquatic Hemipteran’s families (Gerridae Veliidae),
their use in stream biomonitoring programs is still
worldwide limited.
In this study, these aquatic bugs occurred only
at forested sites and presented high diversity; we
believe that these Hemiptera could be sensitive to
water pollution and their use as bioindicators might
enhance the accuracy of water quality assessments
in urban impaired streams. Our results are in line
with those of Zettel & Tran (2004) who found that
in Vietnam, Rhagovelia polymorpha a congener
species of R. reitteri and Rhagovelia sp. identified
in Douala forested stream, also inhabit small stream
in a forested area, with a moderate to slow water
flow, in partial shade, bottom with rock or sand.
Moreover, the canonical redundancy analysis
38
Simeon Tchakonte etalii
(RDA) revealed that the presence and abundance of
the most characteristic species of these aquatic bugs
are positively and significantly influenced by high
dissolved oxygen content, important canopy cover-
age, low mineralization, very low organic matter
input and current velocity.
Concerning morphological features of the Gyrin-
idae, this study revealed that Orectogyrus sp. 1 and
O recto gyrus sp. 2 differ from the Afrotropical Orec-
togyrus specularis Aube, 1838 and O. camerunensis
Ochs, 1924 known to occur in Cameroon, particu-
larly by the distinct metallic shiny omateness of the
elytra. However, these species are to be compared
to other Afrotropical allotype or paratype occurring
elsewhere, to know whether we are face to new
records or new species. As for the Veliidae, Rhagov-
elia sp. described here differs drastically from R.
reitteri, as it lacks wings. Additionally, Rhagovelia
sp. has stout hind femora bearing short distinct
spines on the inner margins. This former character
makes Rhagovelia sp. to be closer to R. polymorpha
describe by Zettel & Tran (2004), but in R. poly-
morpha the body including legs is silky, with nume-
rous black, semi-erect setae and with short,
appressed yellow pubescence; legs with very long
black setae. Moreover, R. polymorpha is smaller in
size (body length 3. 2-3. 6 mm) as compared to our
specimen (body length 4.12 ± 0.2 mm). The speci-
mens of Eurymetra sp. 1 and Eurymetra sp.2 that we
recorded in this study differ significantly from the
typical E. manengolensis described by Hoberlandt
(1952) in Cameroon Manengouba mount. Abdom-
inal connexiva (pleura) are well developed in E.
manengolensis as compared to E. sp. 1 .
In addition, mid- and hind-coxae are larger in E.
sp. 1 than in E. manengolensis. As for E. sp. 2, the
specimen described here shows some similarities
with the genus Eurymetropsis examined by Poisson
(1965) in terms of morphology (especially size and
color). However, in Eurymetropsis body is more
flattened and often lustrous above, the lateral suture
between the meso- and metanotum is not so keeled,
all tarsal segments are also nearly equal in length,
what distinguish it from our specimen.
CONCLUSIONS
This biological assessment permitted to identify
20 species, all present only at the forested sites; no
species being found in urban streams. This study
highlights that species richness and distribution of
aquatic insect of the families Gyrinidae, Gerridae
and Veliidae in Douala watershed are highly and
negatively influence by polluted status of its urban
streams due to anthropogenic activities which cause
the extinction of the sensitive taxa.We thus believe
that these aquatic Coleoptera and Hemiptera species
are sensitive to water pollution and we suggest that
their use as bioindicators might enhance the accur-
acy of water quality assessments in Cameroon.
Morphological description of our specimens
revealed many undescribed taxa which are probably
new records or new species. This testified that in
Cameroon, biodiversity of aquatic insects is yet
entirely to be investigated, and that there is an
urgent need for a modern taxonomic revision and
establishment of a complete key to the Cameroon-
ian species.
ACKNOWLEDGEMENTS
This study was made possible through the pro-
vision of funding by the International Foundation
for Science (IFS) as well as facilities by the Intra-
African Mobility Program Scholarship (PIMASO).
REFERENCES
APHA, 2009. Standard Methods for the Examination of
Water and Wastewater.America Public Health Asso-
ciation, APHA-AWWAWPCF (Eds.), Pennsylvania,
Washington, 1150 pp.
Bode R.W., Novak M.A., Abele L.E., Heitzman D.L. &
Smith A.J., 2002. Quality Assurance Work Plan for
Biological Stream Monitoring in New York State,
Albany (New York). Stream Biomonitoring Unit,
Bureau of Water Assessment and Management,
Division of Water, Department of Environmental
Conservation, 89 pp.
Cohen J.E., 2003. Human population: the next century.
Science, 302: 1172-1175.
Compin A. & Cereghino R., 2003. Sensitivity of aquatic
insect species richness to disturbance in the Adour-
Garonne stream system (France). Ecological Indic-
ators, 3: 135-142.
Dajoz R., 2000. Precis d’Ecologie, 7e edition, Dunod,
Paris, 615 pp.
Dejoux C., Elouard J.M., Forge P. & Maslin J.L., 1981.
Catalogue iconographique des insectes aquatiques
Stream’s water quality and description of aquatic species of Coleoptera and Hemiptera in Littoral Region of Cameroon 39
de Cote d'Ivoire. Edition de l’ORSTOM, Paris, 178
pp.
De Moor I.J., Day J.A. & De Moor F.C., 2003. Guides
to the Freshwater Invertebrates of Southern Africa,
Volume 8: Insecta II. Hemiptera, Megaloptera,
Neuroptera, Trichoptera & Fepidoptera. Water
Research Commission Report, No. TT 214/03,
Pretoria- South Africa, 219 pp.
Durand J.R. & Feveque C., 1981. Flore et faune
aquatiques de l’Afrique Sahelo-soudanienne. Tome
II. Edition de TORSTOM, Paris, 517 pp.
Foto Menbohan S., 2012. Recherche ecologique sur le
reseau hydrographique du Mfoundi (Yaounde): Essai
de biotypologie. These de Doctorat d’Etat, Faculte
des Sciences, Universite de Yaounde I, 179 pp.
Foto Menbohan S., Tchakonte S., Ajeagah G.A., Zebaze
Togouet S.H., Bilong Bilong C.F. & Njine T., 2013.
Water quality assessment using benthic macro- in-
vertebrates in a periurban stream (Cameroon). The
International Journal of Biotechnology, 2: 91-104.
Hauer F.R. & Resh V.H., 1996. Benthic macroinverteb-
rates. In: Hauer, F.R. &FambertiG.A. (Eds.),
Methods in stream ecology. Academic Press, San
Diego, pp. 339-365.
Hilsenhoff W.F., 1988. Rapid field assessment of or-
ganic pollution with a family-level biotic index. Jour-
nal of North America Benthological Society, 7: 65-68.
Hoberlandt F., 1952. A new species of Eurymetra
(Heteroptera, Gerridae) from the Cameroon. Acta
Entomologica Musei Nationalis Pragae, 26: 1-3.
Feclercq F., 2001. Interet et limites des methodes d’esti-
mation de la qualite de l’eau. Document de travail,
station scientifique des Hautes-Fagnes, Belgique,
44 pp.
Fegendre P, Oksanen J. & TerBraak C.J.F., 2011. Testing
the significance of canonical axes in Redundancy
Analysis. Methods in Ecology and Evolution, 2:
269-277.
Muller N., Werner P. & Kelcey J.G., 2010. Urban biodi-
versity and design. Wiley-Blackwell Publishing Ftd,
West Sussex, UK, 626 pp.
Nyamsi Tchatcho N.F., Foto Menbohan S., Zebaze To-
gouet S.H., Onana Fils M., Adandedjan D., Tcha-
konte S., Yemele Tsago C., Koji E. & Njine T.,
2014. Indice Multimetrique des Macroin vertebres
Benthiques Yaoundeens (IMMY) pour revaluation
biologique de la qualite des eaux de cours d’eau de
la Region du Centre Sud Forestier du Cameroun.
European Journal of ScientificResearch, 123: 412—
430.
Olivry J.C., 1986. “Fleuves et Rivieres du Cameroun”.
Ed. Mesres-Orstom, Paris, 733 pp.
Poisson R., 1965. Catalogue des Heteropteres Gerridae
africano-malgaches. Bulletin I.F.A.N., 27: 1 466—
1503.
Rios S.F. & Bailey R.C., 2006. Relationships between
riparian vegetation and stream benthic communities
at three spatial scales. Hydrobiologia, 553: 153-160.
Rodier J., Fegube B., Marlet N. & Brunet R., 2009. F’
analyse de l’eau. 9e edition, DUNOD, Paris, 1579 pp.
Rosenberg D.M. & Resh V.H., 1993. Freshwater biomon-
itoring and benthic macroinvertebrates. Chapman and
Hall, Fondon, 279 pp.
Sanchez-Femandez D., Abelian P, Mellado A., Velasco
J. & Millan A., 2006. Are water beetles good indic-
ators of biodiversity in Mediterranean aquatic
systems? The case of the Segura river basin (SE
Spain). Biodiversity and Conservation, 15: 4507-
4520.
Song M.Y., Feprieur F., Thomas A., Fek-Ang S., Chon
T.S. & Fek S., 2009. Impact of agricultural land use
on aquatic insect assemblages in the Garonne river
catchment (SW France). Aquatic Ecology, 43: 999-
1009.
Stals R. & De Moor I.J., 2007. Guides to the Freshwater
Invertebrates of Southern Africa, Volume 10: Co-
leoptera. Water Research Commission Report, No.
TT 320/07, Pretoria- South Africa, 275 pp.
Stark J.D., Boothroyd K.G., Harding J.S., Maxted J.R. &
Scarsbrook M.R., 2001. Protocols for Sampling
Macroinvertebrates in Wadeable Streams. New
Zealand Macroinvertebrates working group, report
no. 1, Ministry for the Environment, Sustainable
Management, fund project no. 5103, 57 pp.
Suchel J.B., 1972. Fe climat du Cameroun. These de
Doctorat 3eme cycle, Universite de Bordeaux III,
Paris, 186 pp.
Tachet H., Richoux P, Bournaud M. & Usseglio-Polatera
P., 2010. Invertebres d'eau douce. Systematique,
biologie, ecologie. CNRS editions, Paris, 588 pp.
Tchakonte S., Ajeagah G.A., Diomande D., Camara I. A.,
Konan K.M., & Ngassam P, 2014. Impact of anthro-
pogenic activities on water quality and Freshwater
Shrimps diversity and distribution in five rivers in
Douala, Cameroon. Journal of Biodiversity and
Environmental Sciences, 4: 183-194.
Tening A.S., Chuyong G.B., Asongwe G.A., Fonge B.A.,
Fifongo F.F. & Tandia B.K., 2013. Nitrate and
ammonium levels of some water bodies and their
interaction with some selected properties of soils in
Douala metropolis, Cameroon. African Journal of
Environmental Science and Technology, 7: 648-656.
Ter Braak C.J.F. & Smilauer P, 2002. CANOCO refe-
rence manual and Canodraw for Windows user’s
guide: software for canonical community ordination
(version 4.5). Microcomputer Power, Tthaca, New
York, USA, 500pp.
Xu M., Wang Z., Duan X. & Pan B., 2013. Effects of
pollution on macroinvertebrates and water quality
bio-assessment. Hydrobiologia, 703: 176-189.
40
Simeon Tchakonte etalii
Zettel H. & Tran A.D., 2004. Two new species of
Rhagovelia (Heteroptera: Veliidae) from Vietnam: first
records of the R. papuensis group from south-eastern
Asia. Tijdschrift voor Entomologie, 147: 229-235.
Zhang Y., Zhao R., Kong W., Geng S., Bentsen C.N. &
Qu X., 2013. Relationships between macroinverteb-
rate communities and land use types within different
riparian widths in three headwater streams of Taizi
River, China. Journal of Freshwater Ecology, 28:
307-328.
Biodiversity Journal, 2015, 6 (1): 41-52
High frequency components of the songs of two Cicadas
(Hemiptera Cicadidae) from Sardinia (Italy) investigated by
a low-cost USB microphone
Cesare Brizio
CIBRA - Centro 1 n te rd is c ip lin are di Bioacustica e Ricerche Ambientali d e 11’ U n i v e rs ita di Pavia, ViaTaramelli 24, Pavia, Italy:
e.-mail: cebrizi@ tin.it
ABSTRACT During August 2013, a low-cost ultrasonic USB microphone (Ultra mic 250 by Dodotronic),
was field-tested for its first application ever in C icadom orphan bioacoustics studies. Two
different species were recorded in the ultrasonic domain, with 250 kHz sampling frequency,
one of them also with 96kHz audio recordings for comparison purposes. Ultra mic 250 proved
suitable for field use, while the recording campaign provided the opportunity to con firm the
presence in South-Western Sardinia of two species (Hemiptera Cicadidae), TibicJlCl COTSiCQ,
Corsica B o u 1 a rd , 1 98 3, endemic to Sardinia and Corse, and the widespread Cicada Omi
Linnaeus, 1758. To the best knowledge of the author, those reported are the first field record-
ings of Cicadidae songs encompassing the ultrasonic domain up to 125 kHz and, in particular
for C. Omi, display sound emissions at frequencies above those previously reported in
literature. Even though conceived for the study of C hiropterans, self-contained, low-cost
USB ultrasonic microphones proved usefulin insect bioacoustics investigations.
KEY WORDS Cicadomorpha; ultrasound; bioacoustics.
Received 31.01.2015; accepted 8.03.2015; printed 30.03.2015
INTRODUCTION
It has been known for several years that many
insects species do hear ultrasounds, as for example
in the papers by Conner (1 999), Barber & Conner
(2007), Pollack (2007), Nakano et al. (2008), Sueur
et al. (2008), Corcoran et al. (2009), Nakano et al.
(2009), Takanashi et al. (2010) and Yager (2012),
that successfully demonstrate that ultrasounds play
a sig n ific an t ro le in many contexts, including prey-
predator interaction and male-female communica-
tion. Despite this widely acknowledged fact,
spectral components well above human hearing are
seldom included in field studies about insect songs,
although investigations and description of animal
sounds restrained to a specific frequency window
(such as the human hearing range), may result in an
incomplete or improper representation of their
actual harmonic structure, leading to disputable
conclusions.
Generally speaking, with the notable exception
of the study of Chiropterans, bioacoustics of the
sub-aerial fauna, including insect sounds, has been
fie Id - s tu d ie d mainly within the human hearing
range (conventionally ranging from 20 Hz to 20
kHz - herein under, “audio range”), both for
comparability with published materials, that we
may deem as “historical anthropocentrism”, and for
technical reasons including high cost and complex
handling of the equipment for ultrasound recording,
Cesare Brizio
42
that may require a specific technological stack
including dedicated microphones, preamplifiers,
power sources and recorders, that may prove unsuit-
able for field use.
A recent field expedition of the author to SW
Sardinia, F lu m in im ag g io re ( C a rb o n ia - Ig le s ia s
Province), provided the opportunity to field-test an
in n o v ativ e , lo w cost USB microphone, Dodotronic
Ultramic 250, and resulted in the ultrasound record-
ings here presented to improve bioacoustic know-
ledge on local C icadom orpha, as well as to
document what appears, to the best knowledge of
the author, as the first application of a new class of
cheap, self-contained USB microphones with ultra-
sonic threshold, epitomized by Ultramic 250, in the
field of Hemiptera scientific bioacoustics. As a
further note of interest, the cicada fauna of Sardinia
is still not particularly well studied (J. Sueur, pers.
comm.), and the recordings themselves may con-
tribute to filling this gap.
MATERIAL AND METHODS
Brizio & Buzzetti (2014) reported about the suc-
cessful usage of Ultramic 250 (Fig. 1) in the field
of Orthopteran bioacoustic studies. To test whether
Ultramic application to Cicadomorphan bioacous-
tics would prove equally valid, in August 2013 two
species of cicada from Sardinia were recorded.
All the species reported were recorded within a
15 km range from F lu m in im a g g io re (Carbonia-
Iglesias Province, Sardinia, Italy) (Fig. 2), although
additional recordings of Ciccidci OVVli Linnaeus,
1 758 were subsequently taken in mainland Italy. All
the audio material was obtained by field recording.
Specimens were not captured nor recorded in
constrained conditions.
The capability ofUltramic 250 to deliver accur-
ate recordings of Orthopteran songs was demon-
strated in a separate study (Brizio & Buzzetti,
2014), also by collecting 96 kHz, 16 bit stereo
recordings for comparison purposes. Available
equipment for audio re co rd in g sin clu d ed a Zoom H 1
handheld digital Micro-SD recorder, coupled with
a self-built stick stereo microphone using Panasonic
W M - 64 capsules from an Edirol R-09 digital re-
cord er. A c o u Stic recordings were taken in stereo, 16
bit, with 96 kHz sampling frequency, and thus
capable of covering frequencies up to 48 kHz.
Fig ure 1 . U ltra sound USB re cording set: A s u s Eee PC 1225B
no te book personal computer, USB cable and Dodotronic U 1-
trarnic 250. On the display, SeaWave software by the Uni-
versity of Pavia’s Interdisciplinary Center for Bioacoustics.
Ultrasound monophonic recording at 250 kHz
sampling frequency was performed via a Dodotronic
Ultramic 250 microphone connected via USB cable
to an Asus Eee PC 1 225B notebook personal com-
puter, using SeaWave software by CIBRA-Univer-
sity of Pavia’s “Centro Interdisciplinare di
Bioacustica e R icerche A m bientali” (http ://w w w -3 .
unipv.it/cibra/). Originally received as amplitude
data (mV) by the recording apparatus, software-
normalized spectral energy is expressed in decibels.
Sound pressure is expressed in dB Full Scale, even
though the dB symbol will be used.
Oscillograms, spectrograms and frequency ana-
lysis diagrams were generated by Adobe Audition
1.0 software. All the illustrations refer to Ultramic
250 monophonic recordings unless otherwise noted.
In the recent paper by Brizio & Buzzetti (2014),
some technical requirements of Ultramic 250 (such
as the need to keep the USB cable length under 1 m)
are addressed in more detail. The same paper
proposes a specific operating protocol to ensure
comparability between Ultramic recordings and
audio range recordings available in literature, and
supports the consistency of recordings obtained by
Ultramic and by conventional microphones, while
some cautions are needed due to the poorer fre-
quency response of the ultrasonic-threshold micro-
phone capsule if compared to ordinary microphones.
In day time condition unaffected by Chiropteran
or Orthopteran sounds, background noise floor level
in the ultrasonic domain can be empirically determ-
High frequency components of the songs of two Cicadas from Sardinia (Italy) investigated by a low-cost USB microphone 4 3
ined from frequency analyses as the average level
of the spectral components not attributable to the
sounds emitted by the recorded specimen, and can
easily be measured by recording environmental
sounds in quiet, no wind conditions, pointing the
microphone towards the specimen during silence
pauses. For the recordings here analysed, and for a
“medium gain” setting of Ultramic 250 (see Brizio
& Buzzetti, 2014), noise floor level in spectral
frequency analyses can be placed at around -80 dB
for the entire unaudible range.
In the recording station of Capo Pecora, a 71
kHz, very narrow band continuous emission up to -
65 dB was recorded even in silent conditions and,
being unrelated with the animal sounds here de-
scribed, shall be reported but excluded from any
kind of analysis and will be considered as part of the
background noise, its plausible origin being tele-
communication antennas in the vicinities that may
directly originate the noise, or may induce a spurious
harmonic component in the Ultramic circuitry.
When using Ultramic in the field, it is particu-
larly uneasy to find the ideal recording distance
from a singing specimen (even more so when, as in
the case of the recordings presented here, the
specimen was out of sight) for reasons that include
the incapacity ofthe human ear to take into account
the volume of the inaudible components (as a
consequence, volumes perceived as relatively low
by the unaided ear may saturate the recording) and
the variable intensity of inaudible components
during song emission. As a consequence, even the
smallest variation in the direction of the handheld
microphone pointing towards an unseen specimen
may result in more or less sharp volume changes,
that compose with the natural pattern of volume
variations. Consistent with the scope of this study,
the author strived to attain the closest possible range
and the most precise and constant microphone
heading that could provide a high volume input, as
near as possible to 0 dBfs, from which even the
faintest high frequency harmonics, the most direc-
tional and prone to attenuation even at relatively
short distance - could be extracted and analyzed.
For those reasons, although the oscillogram,
generated in real time by SeaWave, was constantly
monitored during the recording, small oscillations
in volume can be observed.
Adobe Audition software settings, such as
resolution in bands, windowing function and
O Capo P-acora, Horn a$l (Tibicina c, torstca, Cicada omO
Soqai. road for Grugua, approx. 550m as I { Tibicin* c. Corsica)
HKKffl
¥ MW*
Figure 2. Recording stations in southwestern Sardinia, in the territory of the Communes ofArbus
(Capo Pecora) and Flum inim aggiore (Grugua).
44
Cesare Brizio
logarithmic energy plot range (in our case, respect-
ively 16384, Welch Gaussian and 100 dB) used to
generate time-frequency spectrograms were selec-
ted as the best compromise for an accurate graph-
ical rendition unaffected by over-representation of
background noise. As a consequence of the settings
chosen, the lowest significant energy level visual-
ized in the time-frequency spectrograms generated
by Adobe Audition is around -70 dB. In all the
frequency analyses, a heavy line was superimposed
to the illustration at the -70 dB level (Figs. 7, 11
14), marking the level above which spectral
components emerge in the tim e - fre q u e n c y spectro-
grams, and constituting a very conservative threshold
for the safe attribution of those components, well
above the background ultrasonic noise, to the singing
anim al.
To give more evidence even to the faintest
significant spectral components, screenshots from
time-frequency spectrograms (Figs. 8, 12, 13) were
c o n tra s t- e n h a n c e d with Adobe Photoshop by a
procedure involving in sequence: color removal,
image inversion, brightness and contrast adjust-
ment, shadows/highlights adjustment. Those inter-
ventions did not affect the accuracy of time-
frequency rendering, and allowed to highlight the
95 kHz “tail” (see below) to C. OYYli sound units.
RESULTS AND DISCUSSION
Tibicina Corsica Corsica Bouiard, 1 9 8 3
Evidence collected. B io aco u stic al and photo-
graphic.
Figure 3. One of the recorded specimens of Tibicina Cor-
sica Corsica, Genna Bogai, 1 6 . V 111 .2 0 1 3 .
Examined Material. Italy, Sardinia, Genna
Bogai (C arbonia-Iglesias Province), Latitude
39.37373, Longitude 8.49732, 549 m asl and Capo
Pecora (Medio Campidano Province), Latitude
39.450908, Longitude 8.396298, 20 m asl.
Distribution. This subspecies (Lig. 3) is distrib-
uted in Sardinia and Corse (its type locality), while
in mainland Europe (Southern Lrance) it's substi-
tuted by T. Corsica farmairei Bouiard, 19 84.
Remarks. Identification of this species, based
also on visual recognition supported by photo-
graphic evidence, posed no doubt.
Ultrasound recordings took place near Capo
Pecora, in the low shrubs (garrigue) with air
temperatures in the range of 27 °C at around
1 6.00. 96 kHz recordings collected around 11
a.m., in comparable air temperature, along the
road from the Genna Bogai pass to the locality
called Grugua, allowed to verify the consistency
between U ltram ic and ordinary recordings also in
the case of C icadom orphan songs (Pigs. 4, 5). It's
noteworthy that the latter samples include an acous-
tic aggression behaviour as reported by Sueur &
Aubin (2003) for the same species in Corse: the
loud, competitive interaction between two male
specimens, one of them "clicking" and the other
"buzzing" in answer to the "clicks".
The male calling song (oscillogram, Pigs. 4-6)
is typical of T. COYSica. It’s currently believed that
the two subspecies, T. COYSica COYSica and T.
Corsica farmairei can not be separated based on
their songs, that show no appreciable differences
between the insular and the continental subspecies
(J. Sueur pers. comm.).
The whole frequency scope spectrum analysis
(Pig. 7) allows to observe that, besides the cluster
of audible frequency peaks centred around 10 kHz,
the sound pattern can be quite clearly made out up
to around half of the spectrogram , before hitting the
background noise floor observed at around -78 dB.
As explained above, the intensity peak at around 71
kHz is a peculiar noise component to be ignored.
To give evidence of the song’s spectral structure,
figure 7 includes five brackets, C 1/C5, correspon-
ding each to a specific frequency band limited by a
sharp decrease in sound pressure. Within each
cluster (with the exception of C4) two main sub-
clusters can be made out, with the lower frequency
sub-cluster containing the highest sound pressure
High frequency components of the songs of two Cicadas from Sardinia (Italy) investigated by a low-cost USB microphone 4 5
peaks. Components above the background noise
and attributable to the singing specimen can be
made out with sufficient clarity up to 56 kHz.
Although the complexity and peculiarities of
the Cicadomorpha sound apparatus do not allow
for a song with clearly outstanding fundamental
frequencies, as those observed in Orthopteran
songs in Brizio & Buzzetti (2014), it can be easily
recognized how the song acoustic signature, in
frequency bands if not in clearly observable high-
order harmonic frequencies, is observable well
above the audible range.
The time-frequency spectrogram (Fig. 8) gives
further evidence of the presence of ultrasonic, struc-
tured higher- order components replicating the main
audible band centered around 11 kHz.
Cicada orni Linnaeus, 1758
Evidence collected. B io aco u stic al evidence.
Examined Material. Italy, Sardinia, Capo
Pecora (Medio Campidano Province), Latitude
39.450908, Longitude 8.396298, 30 m asl, approx-
imate nearest re cor ding distance 15 m . Italy, Em ilia
Romagna, Poggio Renatico (Ferrara Province),
Latitude 44. 761475, Longitude 11. 473074, 10 m asl,
approximate nearest recording distance 20-25 m.
D is T R ib u T 10 N .T h is species is distributed in all
the Italian territory.
Remarks. The ultrasound recordings took place
near Capo Pecora, collecting the sound of speci-
mens singing from the pine trees and from the
highest shrubs. The following year, further recor-
dings for comparison purposes were obtained in
P o g g io R e n a tic o , in the Padan Plain of mainland
Italy, from specimens singing from English Oaks,
Laurel Oaks, Tree of Heaven AilanthuS ClltissilTlCl
(Mill.) Swingle in an urban private park.
The u nm istakeable calling song (oscillogram,
Figs. 9, 1 0) of C. orni , based on repetitive echemes
and well described in literature (for example by
Sueur et al. (2008)) substantially differs from the
mo re or less continuous, hissing and higher pitched
emission by T. Corsica Corsica.
The frequency analysis of the whole spectral
range of a single echeme (Fig. 11), shows a song
whose conventional subdivision in “bands”, here
proposed as an aid in the observation of the song
structure, isn't as evident as in the song of T.
Corsica Corsica. Apart showing a less defined
pattern, components above the background noise
and attributable to the singing specimen can be
clearly made out up to approximately 80 kHz. By
rescaling the illustration above, one can find a sub-
stantial agreement with Figure 2C in Sueur et al.
(2008), with vibration spectra displaying an higher
relative amplitude around 50 kHz and an increase
towards 80 kHz. As reported by Sueur et al. (2008),
in the high frequency domain the tympanal mem-
brane (TM) of the female C. OWli is driven at its best
re son an ce freq u ency at 50 kHz, a frequency domain
represented by bands C4 and C 5 in Fig. 11 and
Table 2.
Two excerpts from C. orni songs U ltram ic
recordings were compared with frequency analyses
from Sueur et al. (2008), as illustrated in Fig. 12.
Although obtained in different ambient conditions,
the two examined excerpts show some consistent
features w ith the s am pie of C. OfTli tympanal mem-
brane vibration spectra obtained by Sueur et al.
(2008)
- two or three "humps" between 40 kHz and 50
kHz, consistent with some of the male specimens
(faint blue lines)
- sharp energy increase at around 50 kHz
- three "humps" between 50 and 60 kHz, consistent
with several of the female specimens (faint red
line s)
- gradual energy increase towards 80 kHz
Frequencies above 80 kHz were not reported in
Sueur et al. (2008)
Time-Frequency spectrogram (Fig. 13), allows
to observe the sy nchronicity in the emission of the
high-frequency and the audible frequency compon-
ents of the song. Components attributable to the
singing specimen can be made out with sufficient
clarity up to 80 kHz.
As a novelty from an higher frequency range
than that explored by Sueur et al. (2008), a faint
“tail” (a frequency cluster roughly centred at 95 kHz)
lasting around 300 msec was observed immediately
following some of the better defined emissions
(lasting around 100 msec) in the 80 kHz band. By
further contrast enhancement of the time-frequency
spectrogram, and by extending the spectral analysis
to a lapse of time of around 300 msec, encompassing
the “tail”, the presence of this further emission band
can be observed in Fig. 14 and in Fig. 15, emerging
46
Cesare Brizio
Figure 4. Song of Tibitina Corsica Corsica. Oscillogram, calling song -500 msec. Figure 5. Song of T. Corsica Corsica. Calling
song -500 msec. This oscillogram from a 96 kFIz recording of another specimen, obtained by a non-ultrasonic microphone
based on Panasonic WM-64 capsules, is very similar to Fig. 4, and shows an overall good oscillogram consistency between
Ultramic and ordinary recordings. Figure 6. Song of T. Corsica Corsica. Oscillogram, calling song: Sound unit (echerne) -19
msec. Figure 7. Song of T. COfSicCl Corsica. Frequency spectrum analysis of the calling song. Black mann-Harris window type,
FFT size 4096 bytes, 0-125kFlz. Volume window -12dB / - lOOdB. C1/C5: main “bands” or “freq ue nc y clu sters” ob serv ed .
High frequency components of the songs of two Cicadas from Sardinia (Italy) investigated by a low-cost USB microphone 47
Figure 8. Song of Tibidna COVSica COVSiCd. Time-frequency spectrogram, 0-125kHz. The faint peak at 7 1 kHz is a
spurious artifact from an unidentified external source. Figure 9. Song of Cicada OCni. Oscillogram, calling song -870
msec. Figure 10. Song of C. Omi. Oscillogram, calling song: Sound unit (echeme) -120 msec. Figure 11. Song of C.
orm. Frequency spectrum analysis of a single soung unit, B lackm ann-H arris window type, FFT size 4096 bytes,
0-125kHz. Volume range below -20dB. Volume window -12dB / - 1 0 0 d B . C1/C9: main “bands” or “frequency clusters”
observed.
48
Cesare Brizio
Band
Frequency
Hz
Volume
dB
Band
Frequency
Hz
Volume
dB
Band
Frequency
Hz
Volume
dB
C 1
8483
-23 .24
C 1
1 9470
-38.24
C 3
40460
-57.98
C 1
95 82
-20.58
C 1
2 0 3 2 0
-43.16
C 3
44250
-65.95
C 1
103 10
-1 9.98
C 2
22270
-44.30
C 4
463 80
-58.94
C 1
1 0980
-19.18
C 2
25690
-39.33
C 4
5 1080
-60.76
C 1
11960
-29.91
C 2
27280
-41 .60
C 5
54320
-64.69
C 1
1 2690
-33.94
C 2
3 3690
-50.17
C 5
565 1 0
-63.45
C 1
175 10
-37.67
C 3
39420
-61 .36
C 5
56430
-68.53
Table 1 . Song of Tibidna COVSiCd Corsica. Frequency spectrum analysis of the calling song, a selection of the nr ain
observed frequency peaks above -70 dB and their sound pressures from the Ultranric 250 recording.
Band
Frequency
Hz
Volume
dB
Band
Frequency
Hz
Volume
dB
Band
Frequency
Hz
Volume
dB
C 1
2563
-36.98
C 1
1 9770
-50.64
C 5
54500
-63.30
C 1
4882
-1 9.53
C 2
25930
-55.13
C 5
61090
-66.48
C 1
73 85
-24.84
C 3
37900
-66.48
C 7
7 1280
-68.45
C 1
952 1
-33.42
C 4
48090
-63.05
C 8
787 30
-65.22
C 1
10490
-35.22
C 4
49010
-63.78
C 8
795 80
-64.11
Table 2. Song of Cicada Omi. Frequency spectrum analysis of the calling song, a selection of the nr ain observed
frequency peaks above -70 dB and their sound pressures from the Ultranric 250 recording.
Figure 12. Song of Cicada Orni. Two frequency analyses of 300 msec excerpts, centered at -72 dB, from C. Orni U ltranr ic
recordings (solid black lines) are superimposed to average C. Omi male tympanal membrane vibration spectra (blue lines:
male songs, red lines: female songs) measured in laboratory conditions. Illustration modified from Fig. 2 C from Sueur et
al. (2008) - frequency range 20 kFIz- 110 kHz ca.
above the limit of -70 dB and thus becoming observ-
able in the time-frequency spectrogram.
The 80 kHz components do not appear re gularly.
Similarly, the 95 kHz band does not follow every
echeme containing the 80 kHz band. Having not
observed any of the following:
- a total decoupling of the 80 kHz and 95 kHz com-
ponents fro m th e c. Omi ec h e m e s ,
- 80 kHz and 95 kHz components in other Ultramic
rec ord in g s ,
- 80 kHz and 95 kHz components in Ultramic
recordings from the same stations during the pauses
High frequency components of the songs of two Cicadas from Sardinia (Italy) investigated by a low-cost USB microphone 4 9
between C. OTYli song bouts, the author finds much
more probable that those components are an integ-
ral, although occasional, part of C. orni song units
rather than software artefacts emerging at spectro-
gram rendering level, or artefacts from Ultramic.
The s y n c h ro n ic ity of the 80 kHz emission with
some of the echemes, and the appearance of the
highest frequency emissions here reported both
in the frequency analysis mode and in time-
frequency spectrograms corroborate this prelim-
inary conclusion.
In August 20 14, recordings of C. OTYli song,
including the highest frequency components, were
obtained in Poggio Renatico (mainland Italy, Padan
Plain) for comparison purposes, in particular to
investigate the high frequency “tail”. Specimens
were recorded from a slightly higher distance
(around 20 m) than the previous year in Sardinia.
The recordings showed frequent occu rre nces of the
same pattern of irregular high frequency “tails”
observed in August 2013 in Sardinia, affecting a
band of about 20 kHz from around 75 kHz to
around 95 kHz.
For comparison purposes, in the subsequent
night and morning the acoustic/ultrasound back-
ground was recorded in the same location (Poggio
Renatico) of the recordings described above, with
the same settings used during the day, avoiding the
lapse of time from around 9 a.m. to around 9
p . m . when C. orni sings. Screenshots from Adobe
Audition were c o n tra s t- e n h a n c e d with the same
procedure as in figures 16 and 17 for comparison
purposes.
Background recordings taken at around midnight
(Fig. 19) and morning recordings taken at around 8
a.m. (Fig. 20) were examined for any occurrence of
the discontinuous yet well recognizable pattern
observed in the 75 kHz-95 kHz band of C. Omi
recordings. The author observed that:
- none of the background ultrasound components
above 70 kHz exceeded the -70 dB threshold
- the 70 kHz-100 kHz band from the background
recordings doesn't bear any resemblance to the
same band in C. orni recordings. At the same time,
C. orni recordings seem unaffected by the features
appearing in the background recordings.
Background night recordings displayed Chirop-
teran echolocation calls and Orthopteran songs,
with singing species including EumodicOgryllllS
burdigalensis burdigalensis (Latreiiie, 1 804),
Eupholidoptera schmidti (F ieb er, 1 8 6 1 ) , Oecanthus
pellucens (Scopoli,1763). Ultrasounds from passing
bats can be made out at around 30 kHz, while the
regular pattern of Orthopteran songs can be made
out under 30kHz. Frequencies around 68 kHz and
98 kHz display feeble regular pulses of different
duration whose probably anthropogenic origin was
not investigated .
The ultrasound background above 70 kHz recor-
ded in the morning resembled quite closely the
night recording from the same location, with a
persistence of the feeble pulses at around 68 kHz
and 98 kHz. Their regularity in the 8 p.m.-8 a.m.
period may hint at a n o n -b io lo g ic al source.
Surely, a more detailed investigation beyond the
scope of this paper may corroborate or disprove the
author's findings.
CONCLUSIONS
The songs by two cicadas from Sardinia have
been recorded in the field, by a low-cost USB
microphone capable of generating very wide band
(0 to 125 kHz) monophonic recordings, including
both audible and inaudible frequencies. This device,
Ultramic 250, by generating results consistent with
other recording methods and by providing useful
information about the high-frequency components
above 20 kHz and up to 125 kHz, proved as useful
for the investigation of Cicadom orphan songs, as it
proved to be in the study of Orthopteran songs.
The song by T. COTSiCQ. COTSiCQ. showed har-
monic components (bands) up to 56 kHz, while the
song by C. OTlli seems to exceed the lim it o f 8 0 kHz
previously explored in literature, and may include
frequencies in the 100 kHz range.
Getting a full grasp of the intraspecific and
interspecific significance of the ultrasound compon-
ents is beyond the scope of this contribution: it is
reasonable to suppose that the species whose song
is here described may have a sound generating and
receiving capability even in ranges above those
previously reported in literature. Q uestions that can
be addressed include a possible role of ultrasound
components in the evaluation of song direction and
distance by con specifics: a sound source may
be considered omnidirectional when it emits
wavelengths longer than its biggest linear dimen-
sion, while directivity is inversely proportional to
50
Cesare Brizio
Figure 13. Song of Cicada Omi. Enhanced contrast picture of a time-frequency spectrogram , 0-125kHz. White lines give
evidence to the sy nchronicity of audible and inaudible spectral components, up to the frequency cluster centered at around
79 kHz and including a very faint 300 msec “tail” in the 95 kHz range. Figures 14-15. Enhanced contrast spectrogram and
frequency spectrum analysis of 300 msec including the 95 kHz “tail” band. Black mann-Harr is window type, FFT size 4096
bytes, 0-125kHz. Volume range below -20 dB. Volume window -19 dB / - 1 0 2 d B . Figure 16. Song of C. OVVli , comparison
specimen from Poggio Renatico, Padan Plain. Enhanced con trast picture of a time-frequency spectrogram, 0-125 kHz. Black
lines border the frequency cluster centered at around 79 kHz and including a very faint 300 msec “tail” in the 95 kHz range.
High frequency components of the songs of two Cicadas from Sardinia (Italy) investigated by a low-cost USB microphone 5 l
Figures 17-18. Song of Cicada Omi from Padan Plain. Enhanced contrast spectrogram and frequency spec tr urn analysis of
1 800 msec including the 95 kHz “tail” band, B lackm ann-H arris window type, FFT size 4096 bytes, 0-125 kHz. Volume
range below -20 dB. Volume window -15dB/-102 dB. Figure 19. Ultrasound background recording taken at 11:55 p.m. in
the night following the recordings in Poggio Renatico, Padan Plain. Enhanced contrast picture of a tim e -fre q u en c y
spectrogram, 0-125 kHz. See text for comments. Figure 20. Ultrasound background recording taken at 7:55 a.m. in the
morning following the recordings in Poggio Renatico, Padan Plain. Enhanced contrast picture of a time-frequency
spectrogram, 0-125 kHz. See text for comments.
Cesare Brizio
52
wavelength. As reported for example by Miller
(2000, 2002) in the case of K iller W hales OvtiYlUS
OYCa Linnaeus, 1 75 8 (Mammalia Cetacea), as well
as by Jakobsen et al. (2013) for echolocating bats,
for a constant energy and emitter size, an increase
in frequency, that is decrease in wavelengths, fo-
cuses the energy in a beam that is narrower (thus,
more directional) but longer, which at short dis-
tances counteracts the decrease in range due to
increased atmospheric attenuation at higher
frequencies. Unfortunately, field recording condi-
tion and uncontrollable specimen position in the
wild did notallow to draw any conclusion about the
orientation of the singing specimen relative to the
microphone axis, neither to measure the different
relevance of ultrasound components at different
angles between the singing specimen and the
m ic ro p h o n e .
ACKNOWLEDGMENTS
Dr. Jerome Sueur and Prof. Gianni Pavan kin-
dly reviewed the original draft and provided some
important suggestions. This does not imply that
the contributors fully endorse the author's conclu-
sions.
REFERENCES
Barber J.R. & Conner W.E., 2007. Acoustic mimicry in
a predator-prey interaction. Proceedings of the
National Academy of Science, 104: 9331-9334.
Brizio C. & Buzzetti F.M ., 2014. Ultrasound recordings
of some Orthoptera from Sardinia (Italy). Biodiver-
sity Journal, 5: 2 5-3 8.
Conner W., 1999. 'Un chantd'appel amoureux'- acou Stic
communication in moths Journal of Experimental
Biology, 202: 1 7 1 1-1723.
Corcoran A.J., Conner W.E. & Barber J.R., 2009. Tiger
Moth Jams Bat Sonar. Science, 325 (5938): 325-327.
Jakobsen L., Ratcliffe J.M. & Surlykke A., 2013.
Convergent acoustic field of view in echolocating
bats. Nature, 493: 93-96.
Miller P.J.O., 2000. Maintaining contact: design and use
of acoustic signals in killer whales, OfCinilS OVCCl.
PhD dissertation. M assachusetts Institute of Techno-
logy, Woods Hole Oceanographic Institution.
Miller P.J.O., 2002. Mixed-directionality of killer
whale stereotyped calls: a direction of movement
cue? Behavioral Ecology and Sociobiology, 52:
262-270.
Nakano R., Skals N ., Takanashi T., Surlykke A., Koike
T., Yoshida K., M aruyama H., Tatsuki S. & Ishikawa
Y., 2008. Moths produce extremely quiet ultrasonic
courtship songs by rubbing specialized scales.
Proceedings of the N atio nal Academy of Sciences of
the United States of America, 105: 11812-11817.
Nakano R., Ishikawa Y., Tatsuki S ., Skals N ., Surlykke
A. & Takanashi T., 2009. Private ultrasonic whis-
pering in moths. Communicative & Integrative
Biology, 2: 1 23-1 26.
Pollack G.S. & Martins R., 2007. Flight and hearing-
ultrasound sensitivity differs between flight-capable
and flight-incapable morphs of a w in g -d im o rp h ic
cricket species. Journal of Experimental Biology,
210: 3160-3164.
Sueur J. & Aubin T., 2003. Specificity of cicada calling
songs in the genus TibidnU (H em iptera: Cicadidae).
Systematic Entomology, 28: 48 1-492.
Sueur J., Windmill J.F.C. & Robert D., 2008. Sexual
dimorphism in auditory mechanics: tympanal
vibrations of CicCld(l OTlli. Journal of Experimental
Biology, 2 1 1 : 2379-2387.
Takanashi T., Ryo Nakano R ., Surlykke A., Tatsuta H .,
Tabata J., Ishikawa Y. & Niels Skals N ., 2010.
Variation in Courtship Ultrasounds of three Ostvilflid
moths with different sex pheromones. PLoS One,
5(10) :e 13 144
Yager D.D., 2012. Predator detection and evasion by
flying insects. Current Opinion in Neurobiology, 22:
201-207.
Biodiversity Journal, 2015, 6 (1): 53-72
A multi-year survey of the butterflies (Lepidoptera Rhopalo-
cera) of a defined area of theTriestine karst, Italy
Peter F. McGrath
IAP/TWAS, ICTP campus, Strada Costiera 11, 34151 Trieste, Italy; e-mail: mcgrath@twas.org
ABSTRACT A photographic survey of butterflies (Lepidoptera Rhopalocera) was carried out over a period
of three years (2011, 2012 and 2013) in an area around the villages of Malchina, Ceroglie
and Slivia, the municipality of Duino-Aurisina near Trieste, in the Friuli Venezia-Giulia
region, northeast Italy. Historically, this area of the Triestine karst has been influenced by
human activities. Grazing intensity, however, has declined over the past 50-100 years, leading
to encroachment of the forested areas over previously more open grasslands. During the three-
year survey period, sampling intensity, measured as the number of days during which butter-
flies were observed and/or photographed, increased from year to year. In 2012 and 2013,
especially surveys began in February and continued into December. During the three years,
a total of 79 species (Papilionidae, 3; Pieridae, 11; Lycaenidae, 17, Riodinidae, 1; Nymphal-
idae, 37, including 15 Satyrinae; and Hesperiidae, 10), including seven listed as either
endangered or near-threatened in Europe, were identified. Among the species of European
conservation value recorded were: Scolitantides orion, Melitaea aurelia, Melitaea trivia,
Argynnis niobe, Hipparchia statilinus , Coenonympha oedippus and Carcharodus floccifera.
Strong local populations of the following regionally threatened, declining and/or protected
species were also recorded: Euphydryas aurinia , Brintesia circe , Arethusana arethusa,
Hipparchia fagi, Pyronia tithonus and Coenonympha arcania. Such intensive surveys cover-
ing several months of each year provide in-depth knowledge of butterfly fauna in an area of
changing land use, and can provide a benchmark for future surveys against a background of
continued land-use change, as well as other pressures such as climate change.
KEY WORDS Butterflies; Rhopalocera; Triestine karst; environmental change; biodiversity.
Received 31.01.2015; accepted 28.02.2015; printed 30.03.2015
INTRODUCTION
The character of the Triestine karst is determi-
ned by its climate and geology. Climatically, it
represents a transitional area between the Mediter-
ranean and Continental/pre-alpine zones. Geologic-
ally, the underlying limestone rocks contribute to
features such as exposed rocky outcrops, dolinas
(depressions caused by the collapse of underground
caves), thin soils and little surface water (although
some artificial ponds have been created) (Poldini,
1989).
These physical conditions have combined with
historic land-use changes to create the patchwork of
habitats for which the Triestine karst is known today.
The original oak forest was felled in historic times
and for many years the area was heavily grazed.
With a general cessation in grazing, regrowth has
occurred and currently mixed woodlands dominated
by Ostrya carpinifolia Scop., while Carpinus
54
Peter F. McGrath
betulus L., Fraxinus ornus L., Quercus petraea
(Matt.) Liebl. and Q. pubescens Willd. are also wi-
despread. Many areas of open grassland exist, includ-
ing some considered as Mediterranean maquis and
some cut for hay. Other areas are decreasing in size,
however, as bushes and trees, including Cotinus
coggygricL Scop, and Primus mahaleb L., encroach
on formerly grazed or cultivated areas. The grassy
areas that remain contain a mixture of xerothermic
herbaceous species with a peak flowering period
between mid May and mid June (Poldini, 1 989). Nat-
uralised areas of Pinus nigra J.F. Arnold, introduced
for timber in the 1850s, also survive in pockets.
In the dolinas, where temperature inversions
mean that a depth of 60 m is equivalent to an
elevation of 1,500 to 1,600 m above sea level in
winter and 500 m in summer (Touring Club
Italiano, 1999), tree species other than O. carpini-
folia dominate and the microclimate ensures the
survival of glacial relict plant communities.
Meanwhile, close to the villages, small-scale
vineyards and vegetable plots provide mainly for
local consumption. The combination of these phys-
ical and biological conditions has created a unique,
biodiverse environment. Paolucci (2010), for ex-
ample, includes 214 species in his guide to the
butterflies of northeast Italy, including the regions of
Trentino Alto Adige, Veneto and Friuli Venezia
Giulia (the Triveneto) - or some 44% of the 482
European species, the karst playing host to well over
a hundred species.
The encroachment of woodland into open grassy
areas due to the abandonment of formerly grazed
areas, however, continues to change the character
of the Triestine karst, impacting on the fauna and
flora. Van Swaay & Warren (2001), for example,
have noted that the abandonment of agricultural
land and/or changing habitat management affects
some 65% of threatened butterfly species in Europe,
while widespread loss and reduction in size of breed-
ing habitats resulting in habitat isolation and
fragmentation affects 83% of Europe’s threatened
species. Many species listed by Paolucci (2010)
thus exist in fragmented habitats or at the edge of
their ranges.
Overlayed across such biological and anthropo-
genic influences, climate change is also having
noticeable effects on the distribution of many
European butterfly species (Roy & Sparks, 2000;
Roy et al., 2001; Warren et al., 2001; Stefanescu et
al., 2003), and will continue to do so for the fore-
seeable future (Settele et al., 2008).
Given the importance of several Italian locations
for butterfly diversity and conservation (van Swaay
& Warren, 2006), alongside the lack of any sys-
tematic recording scheme in the country (van
Swaay et al., 2012a), this study set out, through
surveying the butterfly fauna of a restricted area of
the Triestine karst, to establish a benchmark against
which future surveys to determine the ongoing
impacts of local land-use and/or climate-induced
changes can be compared.
MATERIAL AND METHODS
Study area
A photographic survey of butterflies (Lepidoptera
Rhopalocera) was earned out over a period of three
years in an area around the villages of Malchina,
Ceroglie and Slivia, the municipality of Duino-
Aurisina near Trieste, in the Friuli Venezia-Giulia
region, northeast Italy, close to the border with
Slovenia (Fig. 1). The highest elevation in the region
is Monte Ermada (323 m) to the west of the surveyed
zone, which is crossed by several rough tracks and
paths. The main paths included in the surveys de-
scribed herein mostly either start from or pass
through Malchina, and include parts of the Gemina
path, the Vertikala, CAI 3 1 and other marked paths
(Fig. 1; Anonymous, 2005), and pass through
various habitats, including vegetable plots, vine-
yards, woodlands, dolinas, and grassland that may
or may not be cut for hay. There are also several
ponds in the study area, in particular one at
Malchina and two close to Slivia.
In Malchina itself, many gardens have nectari-
ferous plants such as Lavandula L., Mentha L. and
Origanum L. that flower especially in July and
attract butterflies from the surrounding areas. The
author’s south-facing garden is one such example.
Equipment
During sampling sessions, pictures were taken
of as many butterflies encountered as possible - if
possible including both upper- and under-wing
views to assist with accurate identification. For the
most part, a Pentax K-k digital camera (typically set
A multi-year survey of the butterflies (Lepidoptera Rhopalocera) of a defined area of the Triestine karst, Italy
55
Figure 1 . Study area. Left: location of the area surveyed in this study in relation to the rest of Italy, the Friuli Venezia Giulia
region and the city of Trieste. The area highlighted in green is shown in more detail to the right. (Outline maps courtesy of
d-maps.com). Right: details of roads plus key tracks and paths and other features of the survey area between Ceroglie, Mal-
china and Slivia north to the border between Italy and Slovenia.
to 200 ASA) was used in tandem with a Sigma 105
mm macro lens. On other occasions, other digital
devices such as a compact camera or smartphone
were used to record specimens. In addition, espe-
cially in 2013 and for those species that are easier
to identify definitively (e.g. Iphiclides podalirius ,
male Anthocharis cardamines , male Colias croceus
or Vanessa atalanta), butterflies identified without
being photographed were recorded as ‘observed’.
Sampling technique/intensity
Surveys were undertaken over three consecutive
years by following the rough tracks, footpaths and
field margins in the survey area. No attempt was made
to quantify the numbers of a given species observed.
Sampling intensity increased during the course of
the three years, as outlined in Table 1 . In most cases,
surveys were carried out for at least 30 minutes and
usually for between 60 to 120 minutes. Surveys were
also typically carried out on hot (for the time of year),
sunny days with minimal cloud cover.
In 2011, photographs were taken ad hoc, with
no attempt to systematically record all sightings,
rather just a few notable occurrences. In addition,
in most cases, the actual sampling actual dates were
not precisely recorded, just the month. In nine
sampling instances, the month is recorded only as
either June or July (Table 1).
In 2012, more intense efforts were made to
photograph or identify all butterflies observed.
Survey dates (59 in total) were accurately recorded
(Table 1).
In 2013, attempts were made to photograph or
identify all butterflies observed. As in 2012,
sampling occasions noted in Table 1 as being under-
taken in the author’s garden often lasted just a few
minutes and tended to be limited to the period of flo-
wering of the Lavandula, Mentha and Origanum
plants. In other cases, butterflies observed during
days when no specific (photographic) survey was
undertaken were also recorded (12 such occasions).
In 2013, including sampling occasions when either
only observations were recorded or when no butter-
flies were seen (despite favourable conditions), a
total of 61 sampling sessions were undertaken
(Table 1).
Identification and analysis
To identify the species recorded, various guide
books were consulted, especially Paolucci (2010)
56
Peter F. McGrath
and Tolman & Lewington (1997). In cases of
uncertainty, experts belonging to the Forum
Entomologi Italiani (http://www.entomologiitaliani.
net) were consulted by posting suitable photographs
online. The author also gratefully acknowledges the
assistance of Lucio Morin, a local butterfly expert,
for help with either the identification or confirm-
ation of the identification of a number of specimens.
Among those species that can be difficult to
distinguish from photographs, L. Morin (pers.
comm.) also confirms that the species found in the
sampling area are Leptidea sinapis, not L. reali,
Colias alfacariensis Ribbe, 1905, not C. hyale
(Linnaeus, 1758), and Plebejus argus, not PI. idas
(Linnaeus, 1761). In the case of white Pieridae,
especially when no suitable photograph was ob-
tained, individuals could often only be identified to
the genus level ( Pieris ). In 2011 or 2012, Pieris
spp. were not regularly recorded, either as photo-
graphs or as ‘observed’. Species names are valid as
per the listing on Fauna Europea (www.faunaeur.
org). It should be noted, however, that Fauna
Europea considers Hamearis lucina (Linnaeus,
1758) as a member of the family Riodinidae,
whereas it is now included among the Lycaenidae
by many authors. The conservation status of the
species observed is based on the European Red List
of Butterflies (van Swaay et al., 2010), the list
provided by van Swaay et al. (2012b) for the
European Habitats Directive, and the list for the
Triveneto region provided by Paolucci (2010).
RESULTS
Environmental variables
A total of 482, 1,208 and 1,657 photographs
were retained from sampling surveys carried out in
2011, 2012 and 2013, respectively. These photo-
graphs accounted for 156, 479 and 738 individual
butterflies in each of the three years, respectively.
In addition, in 2013, some 128 individuals were
recorded as ‘observed’ but not photographed.
During these three years, 79 butterfly species
were recorded. Of these, 45 were recorded in 201 1
when sampling was less intensive, 63 in 2012, and
70 in 2013 (Tables 2, 3 and 4).
Of the 79 species recorded, 3 belonged to the
family Papilionidae; 11 to the Pieridae; 17 to the
Lycaenidae; one to the Riodinidae; 37 to the
Nymphalidae, of which 15 were Satyrinae; and 10
to the Hesperiidae.
In early 2012, no butterflies were observed or
photographed during the single sampling date in
February (12th), although they were on two of three
dates in March (on 1 1th and 24th, but not on 26th).
Likewise, in 2013, no butterflies were observed or
photographed on the February sampling date (16th),
while they were recorded on one of the two
sampling dates in March (on 3rd, but not on 22nd),
and on nine of 10 dates in April (not on 15th).
Among the early-season (up to mid April) species
recorded were Pieris rapae , P. napi, Gonepteryx
rhamni, Libythea celtis , Nymphalis polychloros,
Pararge aegeria and Erynnis tages.
In the second half of April, 15 species were
recorded in 2012 (including one specimen of Zeryn-
thia polyxena on 30 April) and 16 in 2013 (Tables 3
and 4). Among these in 2013 was V. atalanta, which
was also regularly recorded in early March 2014.
With regard to late-season records, in 2012,
butterflies were recorded on 2 and 3 November, but
not 22nd. No sampling was undertaken in Decem-
ber 2012. In 2013, butterflies were recorded on
three of four dates in November (1,10 and 17th, but
not on 24th), and on one of two dates in December
(on 14th but not on 7th). These late-season species
(observed in November and into early Decem-
ber), included C. crocea, L. celtis , V atalanta and
Cacyreus marshalli.
The highest number of species recorded in a
single day was 24 (on 24 August 2013), with more
than 20 species also being recorded on six other
occasions in 2013 (22 June, 13 and 20 July, 16 and
18 August and 9 September). In 2012, the max-
imum number of species recorded in a single day
was 17 (on 17 July).
Comparing the number of species observed
during half-month periods (Tables 2, 3 and 4), 37
species were recorded in the second half of July
2013, with 50 species recorded for the month as a
whole (Table 4). Similarly, in 2012, more species
were recorded in July than any other month (37),
although the diversity was greater in the first half
of the month (29 species compared to 19 in the
second half of the month) (Table 3).
Among the species most commonly recorded
(depending on their respective flight periods) were
I. podalirius , P. rapae and P. mannii, PI. argus,
Polyommatus icarus, Po. bellargus, V. atalanta.
A multi-year survey of the butterflies (Lepidoptera Rhopalocera) of a defined area of the Triestine karst, Italy
57
Melanargia galathea, f. procida, Maniola jurtina
and Coenonympha pamphilus. Among the most
commonly recorded Hesperiidae were E. tages,
Hesperia comma and Ochlodes sylvanus.
Other species were relatively common in some
years, but not recorded in other years. Aporia
crataegi, for example, was recorded in 2011 and
2013 but not in 2012. Likewise, Hipparchia stat-
ilinus and Coenonympha oedippus were recorded
only in 2012, and Aricia agestis and Pontia edusa
only in 2013 (Tables 2, 3 and 4).
Also of note were variant forms of some species.
M. galathea was always present as M. galathea
f. procida, along with a small percentage of f.
leucomelas. Likewise, a small percentage of Argyn-
nis paphia, were f. valesina.
Species recorded rarely (i.e. no more than two
individuals recorded in any one year) in the area
surveyed include Z. polyxena, Callophtys rubi,
Leptotes pirithous, Cupido argiades, Cyaniris
semiargus, Po. daphnis, Scolitantides orion,
Nymphalis antiopa, Aglais io, Polygonia c-album,
Melitaea aurelia, Brenthis hecate, Argynnis adippe,
A. niobe, C. oedippus, Carcharodus alceae,
Carcharodus floccifera and Spialia serorius.
Among these, Z polyxena, S. orion, N. antiopa, M.
aurelia, B. hecate and C. oedippus are notable
owing to their conservation status (see below).
Of particular interest are seven species recorded
in the survey area that are included in the European
Red List of Butterflies (van Swaay et al., 2010). The
conservation status of these species is outlined in
Table 5. In addition, van Swaay et al. (2010) also
note that Euphydryas aurinia, C. oedippus and Z.
polyxena are listed in 1 6, 2 and 1 European LIFE
projects (see http://ec.europa.eu/ environment/life/),
respectively, with special efforts being made
towards their conservation.
A number of other species recorded in the three-
year survey are also of regional conservation in-
terest (Table 6). Other than species such as Callophrys
rubi, N. antiopa and Melitaea trivia that were
recorded infrequently, healthy populations of
vulnerable and locally protected species (including
L. celtis, E. aurinia, Brintesia circe, Arethusana
arethusa, Hipparchia fagi and Coenonympha
arcania ) were recorded in the survey area.
The case of E. aurinia is interesting in that no
individuals were recorded south of the road that
bisects the village of Malchina (SS4); although
never abundant, it was observed in reasonable
numbers in localized areas north of SS4, but never
far (no more than 500 m) from Malchina itself.
Likewise, all individuals of C. oedippus were recor-
ded within an area of radius no more than 150 m,
also to the north of Malchina.
In addition to those species highlighted in Table
6, a further five species found in the survey area are
recorded by Paolucci (2010) as being lower
risk/near threatened (LR/NT) in the Triveneto
region: Cupido alcetas, S. orion, Hamearis lucina,
Melitaea athalia and Minois dryas. Of these, H.
lucina and M. dryas are also relatively common and
well distributed throughout most of the survey area
(Tables 2, 3 and 4).
Likewise, Paolucci (2010) records the following
species as data deficient (DD) in the Triveneto
region: P. mannii, Favonius quercus, C. argiades,
N. polychloros, M. aurelia, A. niobe and C. flocci-
fera. Of these, P mannii and, early in the season,
N. polychloros both maintain reasonable popula-
tions in the survey area (Tables 3 and 4). Thus, the
three-year survey undertaken by the author helps to
fill some of these data gaps.
DISCUSSION
The total of 79 species recorded during the three-
year survey period compares favorably with other
areas of Europe. In the whole of the United King-
dom, for example, there are just 57 resident plus two
regular migrant species (Asher et al., 2001). Wagner
et al. (2013) recorded 49 butterfly species from 27
sites along an altitude gradient in Bavaria, Germany;
while Veronivnik et al. (2011 a) recorded between 42
and 61 species each year during a five-year survey
(2007-2011) of a disused army base at Mlake in Slov-
enia, recording a total of 95 species overall. In north-
ern Italy, Marini et al. (2009) recorded 60 butterfly
species through sampling 44 hay meadow parcels
during a single year (2007) in the Trento region,
while Boriani et al. (2005) sampled nine sites of
three different rural habitat types in Emilia-Ro-
magna in 2002 and 2003, identifying 39 species. The
total also compares well with the 91 butterfly
species recorded by Carrara (1926) following many
years of collection and study in the area around
Trieste (immediately to the east of the area that is
the focus of this study and covering a much larger
area).
58
Peter F. McGrath
Month-
Dates/
Year
Feb
Mar
Mar
Apr
Apr
May
May
Jun
Jun
Jul
Jul
16-28
1-15
16-31
1-15
16-30
1-15
16-31
1-15
16-30
1-15
16-31
2011
0
0
0
1
1
5 1
5 1
3 2
3 2
9 3
9 3
2012
l 4
1
2
1
6 5
4
3
1
7 1
9 1
8 6
2013
l 4
1
l 4
2
8 io
5 11
3 11
6 11
4 11
5 11
77,11
Month-
Dates/
Year
Aug
Aug
Sept
Sept
Oct
Oct
Nov
Nov
Dec
Total
1-15
16-31
1-15
16-30
1-15
16-31
1-15
16-30
1-15
2011
5
5
3
3
2
2
0
0
0
28
2012
4 7
4
3
2
0 8
O 8
2
1
0
59
2013
l7, 11
3 4
2
2 11
3
1
2 11
2 9
2 12
61
Table 1. Sampling intensity broken down into half- month intervals. Sampling occasions marked as ‘in author’s garden’ or
‘observations only’ (see footnotes) were less intense than other occasions that involved excursions along the various paths
highlighted in Fig. 1. 1 Of which 3 occasions in author’s garden, Malchina. 2 Of which 2 occasions in author’s garden,
Malchina. 3 Refers to June and July together, of which 7 occasions in author’s garden, Malchina. 4 No butterflies observed. 5
One sampling date included two periods (30 April, a.m. and p.m.). 6 Of which 6 in author’s garden, Malchina. 7 Of which 1
occasion in author’s garden, Malchina. 8 Owing to other commitments, no surveys were undertaken during October 2012. 9
No butterflies observed. Sampling earned out in evening (17:50-18:20) after warm sunny day. 10 Of which 2 occasions:
‘observations’ only (no photographs) - within Malchina itself. 11 Of which 1 occasion: ‘observations’ only (no photographs)
- within Malchina itself. 12 Of which 1 occasion: ‘observations’ only (no photographs) (14 December) within Malchina itself.
No butterflies recorded on other sampling date (7 December).
TOTAL SPECIES 45
April (l) 1
May (5)
June (3)
June-
July (9)
Aug (5)
Sept (3)
Oct (2)
PAPILIONIDAE
Iphiclides podalirius (Linnaeus, 1758)
X
_X
X X
Papilio machaon Linnaeus, 1758
_x
X X
PIERIDAE
Anthocharis cardamines (Linnaeus, 175 8)
0
Aporia crataegi (Linnaeus, 1758)
_X_
Pieris mannii (Mayer, 1851)
_x_
Pieris rapae (Linnaeus, 1758)
_x_
Leptidea sinapis (Linnaeus, 1758)
X
X
Colias croceus (Fourcroy, 1785)
X
X
x_
Gonepteryx rhamni (Linnaeus, 1758)
_x_
LYCAENIDAE
Favonius quercus (Linnaeus, 1758)
X
Satyrium ilicis (Esper, 1779)
_x_
Lycaena phlaeas (Linnaeus, 1761)
_x_
Cacyreus marshalli Butler, 1898
x_
Cupido argiades (Pallas, 1771)
X
Table 2 (1/2). Summary of butterfly species recorded and observed in the study area in 2011. 1 Figures in brackets indicate no.
of sampling sessions per month (or June/July period). 2 Actual sampling session not recorded, x = Either one or two individuals
photographed during a sampling session; X = 3 or more individuals photographed during a sampling session; o = Observed (but
not photographed) during a sampling session; _ (or blank) = Neither photographed nor observed during a sampling session.
A multi-year survey of the butterflies (Lepidoptera Rhopalocera) of a defined area of the Triestine karst, Italy
59
Species
April (l) 1
May (5)
June (3)
June-
July (9)
Aug (5)
Sept (3)
Oct (2)
Plebejus argus (Linnaeus, 1758)
X
x
X_Xx_
x_
Plebejus argyrognomon (Bergstrasser, 1779)
X
Polyommatus bellargus (Rottemburg, 1775)
_X_
X
xx_
Xxx
Polyommatus icarus (Rottemburg, 1775)
_X_
_xx
X
_x
RIODINIDAE
Hamearis lucina (Linnaeus, 1758)
X
X
X
NYMPHALIDAE
Vanessa atalanta (Linnaeus, 1758)
X
Vanessa cardui (Linnaeus, 1758)
_x_
Limenitis reducta Staudinger, 1901
X
X
Melitaea aurelia Nickerl, 1 850
X
Melitaea didyma (Esper, 1778)
XX_
Euphydtyas aurinia (Rottemburg, 1775)
X
X
Issoria lathonia (Linnaeus, 1758)
X_X
Argynnis paphia (Linnaeus, 1758)
X X
Argynnis adippe (Denis et
Schiffermuller, 1775)
X
Boloria dia (Linnaeus, 1767)
X 2
Brenthis hecate (Denis et
Schiffermuller, 1775)
X
Melanargia galathea procida (Linnaeus, 1758)
X
NYMPHALIDAE, Satyrinae
Minois dryas (Scopoli, 1763)
xxx_x
X
Brintesia circe (Linnaeus, 1775)
_x_
Arethusana arethusa (Denis et
Schiffermuller, 1775)
xxx_x
_x_
Hipparchia fagi (Scopoli, 1763)
XX
xxx_x
X
Hipparchia semele (Linnaeus, 1758)
_x
_x
Lasiommata maera (Linnaeus, 1758)
X
X
_x_
Pararge aegeria (Linnaeus, 1758)
X
Pyronia tithonus (Linnaeus, 1767)
_x_
Maniola jurtina (Linnaeus, 1758)
X
X
_x_
Coenonympha arcania (Linnaeus, 1761)
_X_
Coenonympha pamphilus (Linnaeus, 1758)
X
xx_
xx_
_xXx_
_xx
_x
HESPERIIDAE
Erynnis tages (Linnaeus, 1758)
X
Hesperia comma (Linnaeus, 1758)
_xx
Ochlodes sylvanus (Esper, 1777)
X
X
X
No . of species/month
3
14
11
10
14
21
6
Table 2 (2/2). Summary of butterfly species recorded and observed in the study area in 2011. 1 Figures in brackets indicate no.
of sampling sessions per month (or June/July period). 2 Actual sampling session not recorded, x = Either one or two individuals
photographed during a sampling session; X = 3 or more individuals photographed during a sampling session; o = Observed (but
not photographed) during a sampling session; _ (or blank) = Neither photographed nor observed during a sampling session.
60
Peter F. McGrath
TOTAL SPECIES 63
Feb
(l) 1
March
1-15(1)
March
16-31(2)
April
1-15(1)
April
16-30(6)
May
1-15(4)
May
16-31(3)
June
1-15(1)
June
16-30(6)
PAPILION ID AE
Iphiclides podalirius
X
X
XX
Zerynthia polyxena (Denis et
Schiffermuller, 1 775 ) 2
X
PIERIDAE
Anthocharis card amines
X
Pieris mannii
Pieris napi Linnaeus, 1758
X
X
Pieris rapae
x
X
X
x_x_xx
Pieris sp.
Leptidea sinapis
_xx
Colias alfacariensis Ribbe, 1905
XX
Colias croceus
X
Colias sp.
Gonepteryx rhamni
X
LYCAENIDAE
Favonius quercus
X
Satyrium ilicis
xx_
Callophrys rubi (Linnaeus, 1758)
X
Lycaena phlaeas
Leptotes pirithous (Linnaeus, 1767)
Cacyreus marshalli
Celastrina argiolus (Linnaeus, 1758)
Cupido alcetas (Hoffmannsegg, 1 804)
X X
XX
Scolitantides orion (Pallas, 1771)
X
Plebejus argus
X
X
X
XX
Plebejus argyrognomon
Polyommatus bellargus
X
_x_
XX
Polyommatus icarus
Xx
X
X
Polyommatus sp.
RIODINIDAE
Hamearis lucina
xx_
xxx_
NYMPHALIDAE
Libythea celtis (Laicharting, 1782)
X
X
X
Vanessa atalanta
Vanessa cardui
Nymphalis polychloros (Linnaeus, 1758)
X
X
Table 3 (1/4). Summary of butterfly species recorded and observed in the study area in II-VI.2012. Legend: 1 Figures in
brackets indicate no. of sampling sessions per month. 2 Author provided only for those species not recorded in 201 1 (Table
2). x = Either one or two individuals photographed during a sampling session; X = 3 or more individuals photographed
during a sampling session; o = Observed (but not photographed) during a sampling session; _ (or blank) = Neither photo-
graphed nor observed during a sampling session.
A multi-year survey of the butterflies (Lepidoptera Rhopalocera) of a defined area of the Triestine karst, Italy
61
Species
Feb
(l) 1
March
1-15(1)
March
16-31(2)
April
1-15(1)
April
16-30(6)
May
1-15(4)
May
16-31(3)
June
1-15(1)
June
16-30(6)
Polygonia c-album (Linnaeus, 1758)
X
Limenitis reducta
_x_
Melitaea athalia (Rottemburg, 1775)
X
X
Melitaea aurelia
X
Melitaea cinxia (Linnaeus, 1758)
X X
X
Melitaea didyma
X x_
Melitaea trivia (Denis et Schiffermuller, 1775)
X
X X
Euphydryas aurinia
xxx_
_xx_
Issoria lathonia
X
Argynnis paphia
_X X_
Boloria dia
xxxx
Brenthis daphne (Bergstrasser, 1780)
X
Brenthis hecate
X
Melanargia galathea procida
X
xx_xx_
Minois dryas
Brintesia circe
X
Arethusana arethusa
Hipparchia fagi
Hipparchia statilinus (Hufnagel, 1766)
Hipparchia semele
Lasiommata maera
X
X
X X
Lasiommata megera (Linnaeus, 1767)
Pararge aegeria
X
X
XX
Pyronia tithonus
Maniola jurtina
X
xx_xx_
Coenonympha arcania
_X_
X
X
Coenonympha oedippus (Fabricius 1787)
Coenonympha pamphilus
xxx_x_
XX
X
xX_x
HESPERIIDAE
Carcharodus alceae (Esper, 1780)
X
Erynnis tages
X
XX
_x
Hesperia comma
Ochlodes sylvanus
X
Spialia sertorius (Hoffmannsegg, 1 804)
X
Thymelicus lineola (Ochsenheimer, 1808)
X
X
Thymelicus sylvestris (Poda, 1761)
X
Total spp. for period
0
2
1
5
18
11
7
13
25
Total spp. for month
0
3
20
16
28
Table 3 (2/4). Summary of butterfly species recorded and observed in the study area in II-VI.2012. Legend: 1 Figures in
brackets indicate no. of sampling sessions per month. 2 Author provided only for those species not recorded in 201 1 (Table
2). x = Either one or two individuals photographed during a sampling session; X = 3 or more individuals photographed
during a sampling session; o = Observed (but not photographed) during a sampling session; _ (or blank) = Neither photo-
graphed nor observed during a sampling session.
62
Peter F. McGrath
Species
July
1-15(9)!
July
16-31(8)
Aug
1-15(4)
Aug
16-31(4)
Sept
1-15(3)
Sept
16-30(2)
Oct
(0)
Nov
1-15(2)
Nov
16-30(1)
PAPILIONIDAE
Iphiclides podalirius
_x_x
Zerynthia polyxena (Denis et
Schiffermuller, 1 7 7 5 ) 2
FIERI DAE
Anthocharis cardamines
Pieris mannii
x X
X
_x
Pieris napi
Pieris rapae
X XX
_x
_x
Pieris sp.
_x_x_
Leptidea sinapis
X X
Colias alfacariensis Ribbe, 1905
_X
X
Colias croceus
X
_x
Xx
Colias sp.
X
Gonepteryx rhamni
LYCAENIDAE
Favonius quercus
X
X
_x
Satyrium ilicis
X
Callophrys rubi (Linnaeus, 1758)
Lycaena phlaeas
xx_
Leptotes pirithous (Linnaeus, 1767)
_x
Cacyreus marshalli
X
X
_x_
Celastrina argiolus (Linnaeus, 1758)
X
_xxxx_
x_x_
Cupido alcetas (Hoffmannsegg, 1 804)
X
X X
Scolitantides orion (Pallas, 1771)
Plebejus argus
_X_Xx
_Xx_xXxx
_X_
_X
Plebejus argyrognomon
X
Polyommatus bellargus
X
_XX_
_xXx
X_
Polyommatus icarus
X
_X_X_X
Xx
xx_x
x_X
_X
Polyommatus sp.
x_
RIODINIDAE
Hamearis lucina
X
X X
NYMPHALIDAE
Libythea celtis (Laicharting, 1782)
x_
Vanessa atalanta
X
_x
x_
Vanessa cardui
X
Nymphalis polych loros (Linnaeus, 1758)
Table 3 (3/4). Summary of butterfly species recorded and observed in the study area in VII-XI.2012. Legend: 1 Figures in
brackets indicate no. of sampling sessions per month. 2 Author provided only for those species not recorded in 201 1 (Table
2). x = Either one or two individuals photographed during a sampling session; X = 3 or more individuals photographed
during a sampling session; o = Observed (but not photographed) during a sampling session; _ (or blank) = Neither photo-
graphed nor observed during a sampling session.
A multi-year survey of the butterflies (Lepidoptera Rhopalocera) of a defined area of the Triestine karst, Italy
63
Species
July
1-15(9)!
July
16-31(8)
Aug
1-15(4)
Aug
16-31(4)
Sept
1-15(3)
Sept
16-30(2)
Oct
(0)
Nov
1-15(2)
Nov
16-30(1)
Polygonia c-album (Linnaeus, 1758)
Limenitis reducta
x
x_x_
X_X
Melitaea athalia (Rottemburg, 1775)
_x
Melitaea aurelia
X
Melitaea cinxia (Linnaeus, 1758)
Melitaea didyma
x_
X
Melitaea Mvia (Denis et Schiffermuller, 1775)
Euphydryas aurinia
Issoria lathonia
X
Argynnis paphia
xx_x
_x
Boloria dia
X
_x
Brenthis daphne (Bergstrasser, 1780)
Brenthis hecate
X
Melanargia galathea procida
_xxXX_xx
Minois dryas
X
xxx_
X
X
Brintesia circe
_x_x
_x
_x
X_X
Are thus ana are thus a
_Xxx
xx_
Hipparchia fagi
X
_X_
XX
Hipparchia statilinus (Hufnagel, 1766)
X
X
X
Hipparchia semele
_X
Lasiommata maera
_x_x
XX
X_
Lasiommata megera (Linnaeus, 1767)
X
Pararge aegeria
X
_x
Pyronia tithonus
X xxxx
xxx_
Maniola jurtina
x_xxx_
_x xx_
XX
XXx_
x_x
_X
Coenonympha arcania
X X
Coenonympha oedippus (Fabricius 1787)
XX
Coenonympha pamphilus
X
X
_XX_
X
X_x
XX
HESPERIIDAE
Carcharodus alceae (Esper, 1780)
Erynnis tages
X
Hesperia comma
Xx
X_
Ochlodes sylvanus
XX_XX_X_
X
Spialia sertorius (Floffmannsegg, 1 804)
Thymelicus lineola (Ochsenheimer, 1808)
Thymelicus sylvestris (Poda, 1761)
X
Total spp. for period
29
19
19
14
16
11
3
0
Total spp. for month
37
23
21
3
Table 3 (4/4). Summary of butterfly species recorded and observed in the study area in VII-XI.2012. Legend: 1 Figures in
brackets indicate no. of sampling sessions per month. 2 Author provided only for those species not recorded in 201 1 (Table
2). x = Either one or two individuals photographed during a sampling session; X = 3 or more individuals photographed
during a sampling session; o = Observed (but not photographed) during a sampling session; _ (or blank) = Neither photo-
graphed nor observed during a sampling session.
64
Peter F. McGrath
TOTAL SPECIES 70
Feb
16-28(1)‘
March
1-15(1)
March
16-31(1)
April
1-15(2)
April
16-30(8)
May
1-15(5)
May
16-31(3)
June
1-15(6)
June
16-30(4)
July
1-15(5)
PAPILION ID AE
Iphiclides podalirius
_x_oo_
xO_xo
o_x
o
oXxXx
Papilio machaon
0
PIERIDAE
Anthocharis cardamines
_xx
X
Aporia crataegi
X
xoxxoo
_x
Pieris brassicae (Linnaeus, 1758)
0
Pieris mannii
X
X
_XXo_
Pieris napi
X
X
Pieris rapae
x_xxo_
xo
0
X
Pieris sp.
0 _
ox_o
Pontia edusa (Fabricius, 1777)
Leptidea sinapis
_oo_ox
x_xox
_xx
XX
x_x
Colias alfacariensis
o_x
Colias croceus
o_x
000
oxx_o
Colias sp.
Gonepteryx rhamni
0
X
00
0
LYCAENIDAE
Favonius quercus
Satyrium ilicis
_xxXx
o_xx
_x
Callophrys rubi
X
Lycaena phlaeas
X
_0
Cacyreus marshalli
_ 0 _
Celastrina argiolus
X
XX
_xxxx
Cupido alcetas
X
Scolitantides orion
X
x_
Aricia agestis (Denis et
Schiffermuller, 1775)
XX
Plebejus argus
_ox_
oXx
X_XxX
xX
Plebejus argyronomon
X
Plebejus sp.
Cyaniris semiargus
(Rottemburg, 1775)
X
Polyommatus bellargus
_Xx
_oxxx_
o_xx
Polyommatus daphnis (Denis
et Schiffermuller, 1775)
Polyommatus icarus
_xx
x_xX_x
_xxx
Polyommatus sp.
X
X
X
Table 4 (1/6). Summary of butterfly species recorded and observed in the study area in II-15.VII.20 13. Legend: 1 Figures
in brackets indicate no. of sampling sessions per month. 2 Author provided only for those species not recorded in 201 1 or
2012 (Tables 2 and 3). x = Either one or two individuals photographed during a sampling session; X = 3 or more individuals
photographed during a sampling session; o = Observed (but not photographed) during a sampling session; _ (or blank) =
Neither photographed nor observed during a sampling session.
A multi-year survey of the butterflies (Lepidoptera Rhopalocera) of a defined area of the Triestine karst, Italy
65
Species
Feb
16-28(iy
March
1-15(1)
March
16-31(1)
April
1-15(2)
April
16-30(8)
May
1-15(5)
May
16-31(3)
June
1-15(6)
June
16-30(4)
July
1-15(5)
RIODINIDAE
Ham ear is lucina
X
oo X
NYMPHALIDAE
Libythea celtis
X
0
X
0
Vanessa atalanta
0
0_0_
_X_00
Vanessa cardui
X
_xx
Agalais io (Linnaeus, 1758)
X
Aglais urticae (Linnaeus, 1758)
0
X
_x_
Nymphalis antiopa (Linnaeus, 1758)
Nymphalis polychloros
X
x_
X
Polygonia c-album
0
_x_
Limenitis reducta
X
X
XXX
0_
0
Melitaea athalia
XX
X
Melitaea cinxia
_xX
Melitaea didyma
X X
XX
_x_x_
Melitaea trivia
X
Euphydtyas aurinia
X X
_xx
Issoria lathonia
X
Argynnis paphia
x_
Argynnis niobe ( Linnaeus, 1758)
X
Argynnis sp.
Boloria dia
X
Brenthis daphne
X
XX
X
Melanarga galathea procida
Xxx
XX
oxXxX
Minois dryas
Brintesia circe
oX
_xxxX
Arethusana arethusa
Hipparchia fagi
_x
Hipparchia semele
_x
Lasiommata maera
_xx
xoXXxx
XX
Lasiommata megera
X
X X
o_xxX
Pararge aegeria
x_xx_oo_
X
0
XX
X
X
Pyronia tithonus
Maniola jurtina
_XX
x_Xxxx
XX
_xXxo
Coenonympha arcania
_XX
x_XxXx
o_xx
0 xo
Coenonympha pamphilus
x_X
_Xx
x_xxxx
XX
_xx_x
Table 4 (2/6). Summary of butterfly species recorded and observed in the study area in II-15.VII.20 13. Legend: 1 Figures
in brackets indicate no. of sampling sessions per month (or June/July period). 2 Author provided only for those species not
recorded in 201 1 or 2012 (Tables 2 and 3). x = Either one or two individuals photographed during a sampling session; X =
3 or more individuals photographed during a sampling session; o = Observed (but not photographed) during a sampling
session; _ (or blank) = Neither photographed nor observed during a sampling session.
66
Peter F. McGrath
Species
Feb
16-28(iy
March
1-15(1)
March
16-31(1)
April
1-15(2)
April
16-30(8)
May
1-15(5)
May
16-31(3)
June
1-15(6)
June
16-30(4)
July
1-15(5)
HESPERIIDAE
Carcharodus floccifera
(Zeller, 1847)
Erynnis tages
xox_x
o
0
Hesperia comma
Ochlodes sylvanus
XX
xX
oXxxx
Pyrgus amoricanus
(Oberthiir, 1910)
Pyrgus malvoides (Elwes
et Edwards, 1 897)
Spialia sertorius
X
Thymelicus lineola
Xxx
x_
Thymelicus sylvestris
XX
Total spp. for period
0
2
0
1
16
22
18
30
26
33
Total spp. for month
0
2
16
30
37
Table 4 (3/6). Summaiy of butterfly species recorded and observed in the study area in II-15.VII.2013. Legend: 1 Figures
in brackets indicate no. of sampling sessions per month. 2 Author provided only for those species not recorded in 2011 or
2012 (Tables 2 and 3). x = Either one or two individuals photographed during a sampling session; X = 3 or more individuals
photographed during a sampling session; o = Observed (but not photographed) during a sampling session; _ (or blank) =
Neither photographed nor observed during a sampling session.
Species
July
16-31(7)i
Aug
1-15(1)
Aug
16-31(3)
Sept
1-15(2)
Sept
16-30(2)
Oct
1-15(3)
Oct
16-31(1)
Nov
1-15(2)
Nov
16-30(2)
Dec
1-15(2)
PAPILIONIDAE
Iphiclides podalirius
oX_Xx_x
Papilio machaon
_X_
_x_
0
PIERIDAE
Anthocharis cardamines
Aporia crataegi
Pieris brassicae (Linnaeus, 1758)
Pieris mannii
_xxxx_x
0
_x
Pieris napi
X
_x
Pieris rapae
0
x_
Pieris sp.
0
oox
_0
0_
0
0
Pontia edusa (Fabricius, 1777)
X
_0
0_
Leptidea sinapis
_xx
x_x
x_
_0
Colias alfacariensis
_x_x_
Table 4 (4/6). Summary of butterfly species recorded and observed in the study area in 16.VII-XII.20 13. Legend: 1 Figures
in brackets indicate no. of sampling sessions per month (or June/July period). 2 Author provided only for those species not
recorded in 201 1 or 2012 (Tables 2 and 3). x = Either one or two individuals photographed during a sampling session; X =
3 or more individuals photographed during a sampling session; o = Observed (but not photographed) during a sampling
session; _ (or blank) = Neither photographed nor observed during a sampling session.
A multi-year survey of the butterflies (Lepidoptera Rhopalocera) of a defined area of the Triestine karst, Italy
67
Species
July
16-31(7)‘
Aug
1-15(1)
Aug
16-31(3)
Sept
1-15(2)
Sept
16-30(2)
Oct
1-15(3)
Oct
16-31(1)
Nov
1-15(2)
Nov
16-30(2)
Dec
1-15(2)
PIERIDAE
Colias croceus
XX
X
xo
X
0
Colias sp.
x
Gonepteryx rhamni
x
LYCAEN1DAE
Favonius quercus
0
Satyrium ilicis
Callophrys rubi
Lycaena phlaeas
x_
x_
0
x_
Cacyreus marshalli
_x_
X
Celastrina argiolus
0
xx_
x_
Cupido alcetas
Scolitcintides orion
Arid a agestis (Denis et
Schiffermuller, 1775)
_X_
0
x_x
x_
Plebejus argus
_x_XxXx
0
xxX
_0
Plebejus argyronomon
X
X
Plebejus sp.
X
Cyaniris semiargus
(Rottemburg, 1775)
Polyommatus bellargus
0
XXX
Xx
_x
Polyommatus daphnis (Denis
et Schiffermuller, 1775)
X
Polyommatus icarus
XXx
XXX
Xx
_x
X
X
Polyommatus sp.
RIODINIDAE
Ham ear is lucina
O X
NYMPHALIDAE
Libythea celtis
X
Vanessa atalanta
X
X_0
0_
oxx
X
xo
X
0
Vanessa cardui
Agalais io (Linnaeus, 1758)
Aglais urticae (Linnaeus, 1758)
XX_
Nymphalis antiopa (Linnaeus, 1758)
0
Nymphalis polychloros
Polygonia c-album
Limenitis reducta
oo_xx_x
0
XXX
_x
ox
X
Melitaea athalia
Table 4 (5/6). Summary of butterfly species recorded and observed in the study area in 16.VII-XII.20 13. Legend: 1 Figures
in brackets indicate no. of sampling sessions per month. 2 Author provided only for those species not recorded in 201 1 or
2012 (Tables 2 and 3). x = Either one or two individuals photographed during a sampling session; X = 3 or more individuals
photographed during a sampling session; o = Observed (but not photographed) during a sampling session; _ (or blank) =
Neither photographed nor observed during a sampling session.
68
Peter F. McGrath
Species
July
16-31(7) 1
Aug
1-15(1)
Aug
16-31(3)
Sept
1-15(2)
Sept
16-30(2)
Oct
1-15(3)
Oct
16-31(1)
Nov
1-15(2)
Nov
16-30(2)
Dec
1-15(2)
Melitaea cinxia
Melitaea didyma
X_X
x_
Melitaea trivia
Euphydryas aurinia
Issoria lathonia
_o_x_
X
Argynnis paphia
ox
xo
Argynnis niobe (Linnaeus, 1758)
Argynnis sp.
_ 0 _
Boloria dia
X
Brenthis daphne
Melanarga galathea procida
ox_xx_
Minois dry as
XXX
x_
_x
Brintesia circe
OX XX X
XX
x_
_x
Arethusana arethusa
XxX
Hipparchia fagi
X
Xox
Xx
ox
Hippar chia semele
XX
_X
x_x
Lcisiommata maera
X
xX
_X
Lasiommata megera
xx_
xxX
x_
_x
Pararge aegeria
X
XXX
XX
oX
0_0
X
Pyronia tithonus
Xx_
0
x_x
x_
Maniola jurtina
0
xxX
xX
Coenonympha arcania
0
Coenonympha pamphilus
xx_x
XXX
Xx
_x
_xx
X
HESPERIIDAE
Carcharodus floccifera
(Zeller, 1847)
X
Erynnis tages
_xXXx_
_x_
Hesperia comma
XXX
xX
_x
Ochlodes sylvanus
ooxXx
Pyrgus amoricanus
(Oberthiir, 1910)
X
X
ox
Pyrgus malvoides (Elwes
et Edwards, 1 897)
0
X
x_
Spialia sertorius
Thymelicus lineola
Thymelicus sylvestris
X
Total spp. for period
37
9
31
27
18
8
7
4
4
2
Total spp. for month
50
33
30
10
6
2
Table 4 (6/6). Summary of butterfly species recorded and observed in the study area in 16.VII-XII.20 13. Legend: 1 Figures
in brackets indicate no. of sampling sessions per month. 2 Author provided only for those species not recorded in 201 1 or
2012 (Tables 2 and 3). x = Either one or two individuals photographed during a sampling session; X = 3 or more individuals
photographed during a sampling session; o = Observed (but not photographed) during a sampling session; _ (or blank) =
Neither photographed nor observed during a sampling session.
A multi-year survey of the butterflies (Lepidoptera Rhopalocera) of a defined area of the Triestine karst, Italy
69
Among the species not recorded by Carrara
(1926) is C. marshalli, a South African species
introduced into Italy in 1997 via horticultural trade
in its host plant, Pelargonium (Balletto et al., 2005).
C. marshalli has been recorded from nearby Udine
and Tarcento as well as Slovenia since 2008
(Bemardinelli, 2008; Verovnik et al., 2011b) and
thus is likely to have arrived in the province of
Trieste around the same time.
Some 13 European countries, including France,
Germany, the Netherlands, Sweden and the UK, are
implementing butterfly recording schemes in at-
tempts to build long-term data sets on species abun-
dance. To date, however, Italy is not among these
countries (van Swaay et al., 2012a, Butterfly Con-
servation Europe: http://www.bc-europe.eu/index.
php?id=339, accessed 14 March 2014). Such
schemes, which also record abundance, are valuable
for detecting population changes over the long-
term, including those influenced by climate change
(Roy & Sparks, 2000; Roy et al., 2001; Warren et
al., 2001; Stefanescu et al., 2003). However, a case
has also been made for recording schemes that meas-
ure presence rather than abundance (Casner et al.,
2014), as is the case in the current study (although
some inferences on abundance can perhaps be made
based on repeated sightings over a short time
period). This study has also identified several species-
rich 1 km transects that could be used as standard
transects in a regular recording scheme for the area
as per current guidelines (van Swaay et al., 2012a).
Among the 79 species recorded in this survey,
some 14 are of conservation concern either in the
region or more widely in Europe (Tables 5 and 6).
Of particular note are E. aurinia and C. oedippus.
In the case of E. aurinia , a number of individuals
were recorded in each of the three years of the sur-
vey, indicating a stable, healthy population, even if
it did not cover the whole of the survey area. While
C. oedippus was recorded only in 2012, several in-
dividuals were found, indicating a relatively small
but potentially healthy population that appears,
however, to be isolated from any other local popu-
lations. Both species were found in patches of rough
vegetation and field margins of the cultivated area
close to Malchina. It can also be noted that neither
species was recorded from the Trieste area in the
early 20th century (Carrara, 1926). Targeted sur-
veys timed to coincide with peak flight periods of
these two species and across a wider area than the
areas identified by the author in this survey would
provide useful additional information on the im-
portance of the location for these two species.
These two species are also among the 34 species
considered by van Swaay & Warren (2006) when
developing a list of Prime Butterfly Areas (PBAs)
for conservation priority in Europe. When selecting
their 43 1 PBAs, van Swaay & Warren (2006) took
into account two types of area: discrete sites that
support one or more target species; and wider areas
(such as mountain ranges or valley systems) where
a target species occurs as scattered populations that
may well be connected as a single metapopulation.
Indeed, a possible C. oedippus metapopulation has
been recorded at sites around Komen, some 8 km
from Malchina across the border in Slovenia (Celik
& Verovnik, 2010). In Italy, C. oedippus is known
from around 100 sites, although many are con-
sidered under threat, mostly by natural reforestation
(Bonelli et al., 2010). Further studies in and around
the survey area would also help to confirm if other
species recorded only rarely in the area were part
of other significant metapopulations.
Given the presence of both E. aurinia and C.
oedippus in the survey area, the area of the Triestine
karst around Malchina could be considered for pos-
sible inclusion as a PBA. This would add to the
cluster of PBAs already identified in the Friuli
Venezia Giulia/Slovenia/Istria region. The fact that
the area also habours a number of other species at
risk regionally, including strong populations of L.
celtis , B. circe , An arethusa, //. fagi and C. arcania,
as well as populations of other species such as H.
statilinus and Pyronia tithonus (Tables 2, 3, 4, and
6) adds to the value of the area.
Species
European
(EU25) status
EU27 status 1
Scolitantides onion
LC
NT
Melitaea aurelia
NT
LC
Melitaea trivia
LC
NT
Argymnis niobe
LC
NT
Hipparchia statilinus
NT
NT
Coenonympha oedippus
EN
LC
Carcharodus floccifera
NT
LC
Table 5. European-level conservation status of endangered
and threatened butterfly species recorded in the survey area
(from van Swaay et al., 2010 and 2012b). LC = Least
concern; NT = Near threatened; EN = Endangered. 1 EU27
includes also Bulgaria and Romania
70
Peter F. McGrath
As a designated Natura 2000 site (see Natura Net-
work Viewer: http://natura2000.eea.europa.eu/#),
much of the survey area is theoretically protected
from development. In practice, however, the on-
going abandonment of agricultural fields and succes-
sion to more overgrown/wooded areas (Poldini,
1989) or other threats such as construction of new
housing continue to erode suitable butterfly habitats.
As mentioned earlier, the abandonment of agricul-
tural land and/or changing habitat management
affects many of Europe’s threatened butterfly
species, while other important threats include climate
change, increased frequency and intensity of fires
and tourism development (van Swaay et al., 2010).
Indeed, in 2012, several areas close to the survey area
were affected by fire (Tosques, 2012a; 2012b).
Habitat loss is, however, regarded as the greatest
threat to butterflies. Van Swaay & WaiTen (2006), for
example, highlight that even species targeted for con-
servation are declining not only within PBAs, but also
within protected areas. Likewise, in the UK, Warren
et al. (2001) demonstrated that, despite the positive ef-
fects of climate change on range expansion, for three-
quarters of 46 species considered, these gains were
outweighed by the negative effects of habitat loss.
Van Swaay & Warren (2006) conclude that le-
gislation alone is not enough to maintain threatened
populations, but that practical conservation meas-
ures are also urgently needed. Such measures
should include sound habitat management of key
sites allied with sympathetic management of sur-
rounding areas, such as the continuation of tradi-
tional agriculture and forestry practices. They also
recommend that populations of target species are
monitored and that research is conducted to identify
appropriate habitat management techniques - with
appropriate financial support. In contrast, Navarro
& Pereira (2012) argue that ‘rewilding’ (defined as
“the passive management of ecological succession
with the goal of restoring natural ecosystem pro-
cesses and reducing human control of landscapes”)
of abandoned farmland should be considered as a
possible land management option in Europe, partic-
ularly on marginal areas. However, they also recog-
nize that such passive forest regeneration will cause
some species to decline in abundance while others
would increase, i.e. there would be both ‘winner’
and ‘loser’ species.
In the survey area considered here, the greatest
Species
Status in Triveneto region
Comments re: area surveyed 1
Zerynthia polyxena
Very local, EN, protected at EU level
One individual photographed in 2012
Callophrys rubi
LR but in decline
Rare. Recorded once in 2012 and once in 2013
Libythea celtis
Scarce, VU
Good local populations
Nymphalis antiopa
DD/EN - population at lower altitudes EN
One individual observed in 2013
Melitaea trivia
VU, protected in FVG 2
Never common. Recorded twice in 2012 and
once in 2013
Euphydryas aurinia
NT, protected in FVG at EU level
Reasonable population localized to parts of
survey area
Bren this hecate
VU
Very rare. Recorded twice in 2012 only
Brintesia circe
EN, threatened, very local
Good local population
Arethusana arethusa
NT, protected in FVG
Good local population
Hipparchia fagi
VU, locally common , EN in Alto Adige
Good local population
Hipparchia statilinus
DD/LR, can be locally common
A few individuals recorded in 2012 only
Pyronia tithonus
Very local distribution, VU/EN
Found regularly, but never more than one or
two individuals
Coenonympha arcania
ER/NT, common - populations in hill/
mountain areas of FVG less threatened
Good local population
Coenonympha oedippus
VU, protected at EU level
A few individuals recorded in 2012 only
Pyrgus amoricanus
NT, only local populations
Recorded intermittently in 2013 only
Table 6. Triveneto-level conservation status of protected, endangered, threatened and vulnerable butterfly species recorded
in the survey area (from Paolucci, 2010). LR = Lower risk; NT = Near threatened; VU = Vulnerable; EN = Endangered;
DD = Data deficient. 1 For additional details, refer to Tables 2, 3 and 4. 2 FVG = Friuli Venezia Giulia
A multi-year survey of the butterflies (Lepidoptera Rhopalocera) of a defined area of the Triestine karst, Italy
71
threat to local butterfly populations and diversity of
species remains the natural reforestation that is on-
going since the decline of grazing in the area. Similar
effects are occurring to local bird communities, with
specialist grassland species such as the rock partridge
Alectoris graeca (Meisner, 1804), grey partridge
Perdix perdix (Linnaeus, 1758) and ortolan bunting
( Emberiza hortulana Linnaeus, 1758) having gone
locally extinct, populations of skylark (Alcmda
arvensis Linnaeus, 1758) and tawny pipit Anthus
campestris (Linnaeus, 1758) under threat, and num-
bers of com bunting ( Emberiza calandra Linnaeus,
1758), red-backed shrike ( Lanius collurio Linnaeus,
1758) and nightjar ( Caprimulgus europaeus Lin-
naeus, 1758) much reduced. Concomitantly there
have been increases in species frequenting scmb and
woodland, such as the nightingale, blackcap, black-
bird, chaffinch and melodious warbler (Parodi,
1999). However, exactly which type of management
practices are most suited for maintaining both faunal
and floral diversity in the area, is unknown.
Based on research in Germany on a comparable
grassland site with shallow soil in a warm, dry tem-
perate climate, Romemiann et al. (2009) concluded
that neither mowing nor various mulching regimes
properly conserved the structure of wildflower pop-
ulations developed over many years of grazing in
species-rich semi-natural grasslands. However, they
did recommend mulching twice per year, as this gen-
erated the most similar floristic and functional plant
community compared to the original grazing regime.
In contrast, regarding the conservation of another
endangered grassland-specialist insect species, Saga
pedo (Pallas, 1771) (Orthoptera, Tettigoniidae) that
is also present in the survey area (Fontana &
Cussigh, 1996; author’s observations), from their stu-
dies in the Czech Republic, Holusa et al. (2013) re-
commended either extensive rotational grazing or
using scythes to cut grass in a traditional way to main-
tain open areas of natural grassland. Alternatively,
partial machine mowing (one-third to one-half of
specific areas) each September could be considered.
Unfortunately it is more than likely that the cur-
rent situation of abandonment and neglect of once
grazed and cultivated areas is likely to continue in
the survey area for the foreseeable future. Similarly,
Bonelli et al. (2010), discussing the conservation of
C. oedippus populations across Italy, note that nat-
ural reforestation is best prevented by developing suit-
able, but costly, management plans, “which for the
moment remain only on paper, in the best of cases.”
The same is likely true for large parts of the
Triestine karst, despite the undoubted conservation
value for butterfly species, as reported here.
ACKNOWLEDGEMENTS
The author gratefully acknowledges the as-
sistance of Lucio Morin, a local butterfly expert, for
help with the identification of a number of speci-
mens and for providing valuable comments on the
text. In addition, a number of other members of the
Forum Entomologi Italiani assisted with identifica-
tion of various specimens via photographs posted
to the online forum (www.entomologiitaliani.net).
Paul Tout and Rudi Verovnik also provided valuable
comments on the text. The contents of this article
represent the views of the author and do not reflect
the official opinion of TWAS or IAP.
REFERENCES
Anonymous, 2005. Carta topographica per escursionisti
no. 47: Carso Triestino e Isontino. Tabacco, Udine.
Asher J., Warren M., Fox R. Harding P., Jeffcoate G. &
Jeffcoate S. (Eds.), 2001. Millennium Atlas of
Butterflies in Britain and Ireland. Oxford University
Press, Oxford, UK, 456 pp.
Balletto E., Bonelli S. & Cassulo L., 2005. Mapping the
Italian butterfly diversity for conservation. In: Kuhn
E., Feldmann R., Thomas J.A. & Settele J.(Eds.),
December 2005. Vol. 1: General Concepts and Case
Studies on the Ecology and Conservation of Butter-
flies in Europe - Conference Proceedings. UFZ
Leipzig-Halle, 71-76.
Bonelli S., Canterino S. & Balletto E., 2010. Ecology of
Coenonympha oedippus (Fabricius, 1787) (Lepidop-
tera: Nymphalidae) in Italy. Oedippus, 26: 25-30.
Boriani L., Burgio G., Marini M. & Genghini M., 2005.
Faunistic study on butterflies collected in Northern Italy
rural landscape. Bulletin of Insectology, 58: 49-56.
Bernardinelli I., 2008. Prima segnalazione di Cacyreus
marshalli per il Friuli Venezia Giulia. Notiziario
ERSA 3/2008: Available at: http://www.ersa.fvg.it/
informativa/notiziario-ersa/anno/2008/numero-3/
prima-segnalazione-di-cacyreus-marshalli-per-il-
friuli-venezia-giulia/ (accessed 10 March 2014).
Carrara G., 1926. Macrolepidotteri del territorio di
Trieste. Atti del Museo Civico di Storia Naturale -
Trieste, 11: 63-116.
Casner K.L., Forister M.L., Ram K. & Shapiro A.M.,
2014. The utility of repeated presence data as a
surrogate for counts: a case study using butterflies.
Journal of Insect Conservation, 18: 13-27.
72
Peter F. McGrath
Celik T. & Verovnik R., 2010. Distribution, habitat pref-
erences and population ecology of the False Ringlet
Coenonympha oedippus (Fabricius, 1787) (Lepidop-
tera: Nymphalidae) in Slovenia. Oedippus, 26: 7-15.
Fontana R & Cussigh F., 1996. Saga pedo (Pallas) ed
Empusa fasciata Brulle in Italia, specie rare da pro-
teggere (Insecta Orthoptera e Mantodea). Atti dell’
Accademia Roveretana degli Agiati, 246B: 47-64.
Holusa J., Kocarek P. & Vlk R., 2013. Monitoring and
conservation of Saga pedo (Orthoptera: Tetti-
goniidae) in an isolated northwestern population.
Journal of Insect Conservation, 17: 663-669.
Marini L., Fontana P., Battisti A. & Gaston K., 2009.
Agricultural management, vegetation traits and
landscape drive orthopteran and butterfly diversity in
a grassland-forest mosaic: a multi-scale approach.
Insect Conservation and Diversity, 2: 213-220. doi:
10. 1111/j. 1752-4598.2009. 00053.x
Navarro L.M. & Periera, FI.M., 2012. Rewilding aban-
doned landscapes in Europe. Ecosystems 15: 900-
912. doi: 10. 1007/sl002 1-012-9558-7
Paolucci P., 2010. Le farfalle delFItalia nordorientale.
Museo di Storia Naturale e Archeologia di Montebel-
luna, Treviso / Cierre Edizione, Verona, 240 pp.
Parodi R., 1999. Gli Uccelli della Provincia di Gorizia.
Museo Friulano di Storia Naturale, Udine, 356 pp.
Poldini L., 1989. La vegetazione del carso Isontino e
Triestino - Studio paessaggio vegetale fra Trieste,
Gorizia e i territori adiacenti. Edizioni Lint, Trieste,
313 pp.
Romermann C., Bernhardt-Romermann M., Kleyer M.
& Poschlod P., 2009. Substitutes for grazing in semi-
natural grasslands - do mowing or mulching represent
valuable alternatives to maintain vegetation struc-
ture? Journal of Vegetation Science, 20: 1086-1098.
Roy D.B. & Sparks T.H., 2000. Phenology of British
butterflies and climate change. Global Change
Biology, 6: 407-416.
Roy D.B., Rothery P., Moss D., Pollard E. & Thomas
J.A., 2001. Butterfly numbers and weather: predicting
historical trends in abundance and the future effects
of climate change. Journal of Animal Ecology, 70:
201-217.
Settele J., Kudrna O., Harpke A., Kuehn I., van Swaay
C., Verovnik R., Warren M., Wiemers M., Hanspach
J., Hickler T., Kuehn E., van Haider I., Veling K.,
Vliegenthart A., Wynhoff I. & Schweiger O., 2008.
Climatic Risk Atlas of European Butterflies. Biorisk
1 (Special issue), 710 pp.
Stefanescu C., Penuelas P. & Filella I., 2003. Effects of
climatic change on the phenology of butterflies in the
northwest Mediterranean Basin. Global Climate
Biology, 9: 1494-1506.
Tolman T. & Lewington R., 1997. Collins Field Guide -
Butterflies of Britain and Europe. HarperCollins,
London, 320 pp.
Tosques R., 2012a. Brucia il Carso sopra Sistiana. II Pic-
colo, 22 August 2012. http://ilpiccolo.gelocal.it/
cronaca/20 1 2/08/22/news/brucia-il-carso-sopra-
sistiana- 1.5579458 (accessed 13 March 2014).
Tosques R., 2012b. Bruciati 10 ettari; Ieri altre fiamme.
II Piccolo, 22 August 2012. http://ilpiccolo.gelocal.
it/cronaca/20 1 2/08/23/news/bruciati- 1 0-ettari-ieri-
altre-fiamme- 1.5583593 (accessed 13 March 2014).
van Swaay C. & Warren M., 2001. Implementing the Red
Data Book of European Butterflies: the identification
of Prime Butterfly Areas. Proceedings of the Section
Experimental and Applied Entomology of the
Netherlands Entomological Society (NEV), Amster-
dam, 12: 129-134.
van Swaay C. & Warren M.S., 2006. Prime Butterfly
Areas of Europe: an initial selection of priority sites
for conservation. Journal of Insect Conservation, 10:
5-11. doi: 10. 1007/s 1084 1-005-7548-1
van Swaay C., Cuttelod A., Collins S., Maes D., Lopez
Munguira M., Sasic M., Settele J., Verovnik R., Ver-
strael T., Warren M., Wiemers M. & Wynhof I.,
2010. European Red List of Butterflies. Publications
Office of the European Union, Luxembourg, 60 pp.
van Swaay C.A.M., Brereton T., Kirkland P. & Warren
M.S., 2012a. Manual for Butterfly Monitoring.
Report VS2012.010, De Vlinderstichting/Dutch But-
terfly Conservation, Butterfly Conservation UK &
Butterfly Conservation Europe, Wageningen, 14
pp. http://www.bc-europe.eu/upload/Manual
Butterfly_Monitoring.pdf (accessed 13 March 2014).
van Swaay C., Collins S., Dusej G., Maes D., Lopez
Munguira M., Rakosy L., Ryrholm N., Sasic M.,
Settele J., Tomas J.A., Verovnik R., Verstrael T.,
Warren M., Wiemers M. & Wynhoff I., 2012b. Dos
and don'ts for butterflies of the Habitats Directive of
the European Union. Nature Conservation 1: 73-153.
doi: 1 0:3 897.natureconservation. 1.2786
Verovnik R., Kosmac M. & Valid P., 2011a. Mlake - a
hotspot of butterfly diversity in Slovenia. Natura
Sloveniae, 13: 17-30.
Verovnik R., Polak S. & Seljak G., 2011b. On the pres-
ence and expansion of an allochthonous butterfly
species in Slovenia - the geranium bronze ( Cacyreus
marshalli (Butler 1898)). Acta entomologica slov-
enica, 19: 5-16.
Wagner K.D., Krauss J. & Steffan-Dewenter I., 2013.
Butterfly diversity and historical land cover change
along an altitudinal gradient. Journal of Insect
Conservation, 17: 1039-1046.
Warren M.S., Hill J.K., Thomas J.A., Asher J., Fox R.,
Huntley B., Roy D.B., Telfer M.G., Jeffcoate S.,
Harding P., Jeffcoate G., Willis S.G., Greatorex-
Davis J.N., Moss D. & Thomas C.D., 2001. Rapid
response of British butterflies to opposing forces of
climate and habitat change. Nature, 414: 65-69.
Biodiversity Journal, 2015, 6 (1): 73-78
First observations on the herpetological and theriological
fauna of Alimia Island (Rhodes Archipelago, Aegean Sea)
Mauro Grano 1 *, Cristina Cattaneo 2 & Augusto Cattaneo 3
'Via Valcenischia 24, 00141 Roma, Italy; e-mail: elaphe58@yahoo.it
2 Via Eleonora d’Arborea 12, 00162 Roma, Italy; e-mail: cristina.cattaneo76@libero.it
3 Via Cola di Rienzo 162, 00192 Roma, Italy; e-mail: augustocattaneo@hotmail.com
'Corresponding author
ABSTRACT This note is a preliminary study on the herpetological and theriological fauna of Alimia Is-
land (Rhodes Archipelago, Aegean Sea). Are described seven species of reptiles and three
of micromammals. Is also provided a short botanical characterization of the island.
KEY WORDS Alimia; Dodecanese; Aegean island; Rhodes.
Received 02.02.2015; accepted 05.03.2015; printed 30.03.2015
INTRODUCTION
Currently there is no scientific literature regard-
ing herpetological and theriological fauna of Al-
imia Island (Rhodes Archipelago, Aegean Sea), and
the following reported data are completely new.
MATERIAL AND METHODS
Alimia Island has been the subject of a partial
survey in August 2014. Since this island is uninhab-
ited, the authors stayed in the nearby island of
Chalki.
Due to the great difficulties in reaching Alimia,
surveys were carried out only in two days, and, for
the discontinuity and precariousness of the con-
nections, the authors of this paper, in the study
of theriological fauna, could not use live-traps
(Sherman) and photo-traps. Therefore were carried
out exclusively field research, by examining osteo-
logical remains and inspecting glass bottles found
in situ.
Study area
Alimia is a small island essentially calcareous,
located in the Aegean Sea, north of Chalki and west
of Rhodes and is one of the 1 63 islands that com-
pose the Dodecanese Archipelago (Sokratis, 2006).
It is part of Peripheral Unit of Rhodes and adminis-
tratively belongs to the Municipality of Chalki.
Its geographical coordinates are: longitude
27°42’24.11” E; latitude 36°16’26.26” N. The
island has an area of 7.42 km 2 , a coastline of 2 1 km
and a maximum height of 274 m above sea level
(Fig. 1). Alimia is provided of two large creeks:
Imborios and Agios Georgios. Just for the presence
of these two natural harbors, in ancient times Alimia
was called Eulimna (from Greek limen = harbor)
(Blackman et al., 2014).
The island is split in two unequal parts by a fail
(Stefanini & Desio, 1928). In southern part a nar-
row strip of land connects the peninsula of Tigani
to the rest of the island (Rackham & Vemicos,
1991). Alimia lacks superficial hydrography; there’s
only a small retrodunal pond of brackish water in
74
Mauro Grano etalii
the bay of Agios Georgios (Fig. 2). Together with
the surrounding small islands and Chalki, it’s in-
cluded in the European Network “Natura 2000” as
SPA, Special Protection Area, with GR42 10026
code. Moreover, with the Official Gazette 991
GG/B of 27 May 1999, Alimia was officially de-
clared archaeological site of national interest for the
presence of remains belonging to the Neolithic,
such as the Paleochristian Basilica in the bay of
Imborios and the Post-Byzantine church of Agios
Georgios. The uncontaminated nature of the coasts
is testified by the presence in this island of the now
rare monk seal, Monachus monachus (Di Turo,
1984; Marchessaux & Duguy, 1977). Currently the
island is uninhabited and is used by the inhabitants
of Chalki, as in the past, for sheep and goats grazing
(Iliadis, 1950). Regular connections either with
Chalki or with Rhodes are not provided, indeed,
Alimia knows only a sporadic tourism made by
private boats.
Botanical aspects. Alimia is characterized by
wide low shrubs in which Juniperus phoenicea L.
and Pistacia lentiscus L. are the most distinctive
elements. These species have pulvinated aspect,
especially near the coast. In places where shrubs
becomes more thin and open, thrives a phiygana
almost exclusively characterized by Thymbra cap-
itata (L.) Cav., to which sometimes is associated
Teucrium capitatum L. and more sporadically
Salvia fruticosa Mill.; Origanum onites L. and
Sarcopoterium spinosum (L.) Spach were infre-
quently observed. The scarcity of the arboreal
element is highlighted by the presence of scarce and
localized clusters of Pinus brutia Ten. The steepest
zones of Alimia have terraces once used for olive
growing, now hardly visible as covered by the
current vegetation. Iliadis (1950) informs that once
this island was used not only as grazing land, but
also for the production of fodder plants, grain, oil
and figs (Cattaneo & Grano, in press).
RESULTS
REPTILIA
Hemidactylus turcicus turcicus (Linnaeus, 1758)
This gecko has a Mediterranean chorotype
(Sindaco & Jeremcenko, 2008). Populations intro-
duced by humans are also known for some states
in United States and South America. Essentially
nocturnal species, is often visible in trophic activity
during evening hours. The few individuals ob-
served at Alimia were found among the ruins of
Ag. Georgios village, under wood planks in shaded
sites with relative humidity. Due to the lack of elec-
tricity in the island, the species could not be ob-
served near light sources.
Mediodactylus kotschyi (Steindachner, 1870)
This species, until a short time ago known as
Cyrtopodion kotschyi , has been recently subject of
taxonomic review (Rosier, 2000). It has an E-
Mediterranean chorotype (Sindaco & Jeremcenko,
2008). Are currently recognized 27 subspecies and
in Chalki and Alimia should be present Mediodac-
tylus kotschyi beutleri (Baran et Gruber, 1981). This
subspecies is typical of southwest Turkey and
eastern Aegean islands. However, the validity of
such subspecies could be under discussion by
modern molecular genetic studies (Kasapidis et al.,
2005). This gecko carries out semidiurnal activities,
choosing stony and arid habitats. It is usually found
on soil and on dry-stones walls (Beutler, 1981). At
Alimia Mediodactylus kotschyi was observed into
Ag. Georgios village, on dry-stone walls used as
enclosures for sheep and goats grazing.
Stellagama stellio daani (Beutler et Fror, 1980)
The chorotype of this reptile is Mediter-
ranean/Arabian (Sindaco & Jeremcenko, 2008).
The recent genus Stellagama is considered mono-
specific and includes the only species S. stellio
(Baig et al., 2012). Seven subspecies are currently
acknowledged, two of which are present in Greece:
S. stellio stellio (Linnaeus, 1758) and S. stellio
daani. The first one has been found in five Cyclades
islands (Delos, Mikro Rhematiaris, Mykonos, Rinia
and Tinos) and in the Ionian islands of Corfu and
Paxi (Spaneli & Lymberakis, 2014); the second one
was found in other Cyclades islands (Paros, Naxos,
Despotico e Antiparos), in most of the eastern
Aegean islands and in Thessaloniki, in the north of
mainland Greece (Spaneli & Lymberakis, 2014). In
Alimia S. stellio daani (Fig. 3) was found relatively
frequent. Adult and young specimens have been
observed during activity and thermoregulation
First observations on the herpetological and theriological fauna ofAlimia Island (Rhodes Archipelago, Aegean Sea) 75
Figure 1. Alimia Island, Rhodes Archipelago, Aegean Sea.
between rocks and among the mins of Ag. Georgios
village.
Ablepharus kitaibelii kitaibelii (Bibron et
Bory, 1833)
A single specimen of this lizard, with a Balcanic
and W- Anatolic chorotype (Sindaco & Jeremcenko,
2008), was observed near a dry-stone wall, close
Ag. Georgios beach. Ablepharus kitaibelii appears
to be a mainly hygrophilous species (Cattaneo,
1998), as generally lives on wet soil and in under-
wood bedding of conifers forest (Broggi, 2002;
Wilson & Grillitsch, 2009). However, both to Al-
imia and to nearby Chalki island, A. kitaibelii has
been found in extreme aridity. In Chalki indeed it
has been very often observed on rocky and arid soil
and also on stone walls inside Imborios village. In
this latter case, however, the research by the above
mentioned species of a degree of humidity inside
the village is plausible.
Anatololacerta oertzeni pelasgiana (Mertens,
1959)
Anatololacerta oertzeni (Werner, 1904) is a
lizard with a Mediterranean chorotype (Sindaco &
Jeremcenko, 2008). It consists of six subspecies. At
Rhodes and in the adjacent islands there’s the sub-
species A. oertzeni pelasgiana (Fig. 4). Like most
Mediterranean reptiles, this lizard is particularly
active in springtime, while in summer exposes one-
self less frequently. However the young, very typ-
ical for the blue color of the tail (Fig. 5), are also
Figure 2. Ag. Georgios village, Alimia Island.
active in the summertime and in the hottest hours
(Wilson & Grillitsch, 2009). Indeed, in Alimia the
young specimens have been seen more frequently,
especially observed up on the walls of the houses
of Ag. Georgios village and on dry stone walls. On
the contrary adults not exposed oneself to direct
sunlight, but stayed inside abandoned houses in
conditions of light and shadow.
Ophisops elegans (Menetries, 1832)
Small lizard with Mediterranean/Iranian choro-
type (Sindaco & Jeremcenko, 2008). Eight sub-
species are currently recognized including one for
Greece: O. elegans macrodactylus (Berthold,
1842). Ophisops elegans (Fig. 6) resulted the most
common reptile in Alimia, generally observed on
soil and especially at the basis of juniper bushes.
Dolichophis sp.
Regarding snakes only a partial exuvia was
found at the basis of a dry stone wall bordering the
abandoned Ag. Georgios village. Stmctural and
chromatic features of the specimen (dark scales with
a light middle strip) assign it unequivocally to the
genus Dolichophis Gistel, 1868. Even if was im-
possible to make a more detailed meristic examina-
tion of the specimen (being the exuvia incomplete),
however, for exclusive biogeographic considera-
tions, we can assume that the exuvia is attributable
to D. jugularis and more precisely to D. jugularis
zinneri Cattaneo, 2012. This subspecies is indeed
present in Rhodes and in the islands of its ar-
76
Mauro Grano etalii
chipelago, such as Chalki, Simi and Tilos (Cattaneo,
2012). Therefore the presence of D.jugularis zinneri
also at Alimia could be argued with good probability.
MAMMALIA
Rattus sp.
hi the immediate surroundings of the big boulders
which form the basis of Kastro (180 m s.l.m.) have
been found two long bones of Rattus sp.: a femur
and a tibia. The failure to find other osteological
remains didn’t allow the distinction between Rattus
rattus Linnaeus, 1758 and R. norvegicus Berkenhout,
1769. Both species live in Rhodes and R. rattus is
present also in the nearby islands of Chalki
(Massed, 2012). In the Dodecanese area is reported
the presence of R. norvegicus for Kos and R. rattus
for Tilos, Karpathos, Kos and Astypalaia (Angelici
et al., 1992; Masseti & Sara, 2002).
Mus musculus Schwarz et Schwarz, 1943
Mus musculus commonly called house mouse, is
an anthropocore and highly invasive regarded
species. This species native of Asia, is present in all
continents, except Antarctica (Masseti, 2012).
Figures 3-6. Reptiles from Alimia Island. Figure 3. Stellagama stellio daani. Figure 4. Anatololacerta oertzeni
pelasgiana (adult). Figure 5. A. oertzeni pelasgiana (young). Figure 6. Ophisops elegans.
First observations on the herpetological and theriological fauna ofAlimia Island (Rhodes Archipelago, Aegean Sea) 77
Mus musculus is included in the list, compiled by
IUCN, of 100 world’s worst invasive alien species
(Lowe et al., 2000). This small rodent can live in
very diversified habitat due to the presence of
human’s commensals populations so-called “indoor”
and wild populations called “outdoor” (Amori et al.,
2008). The osteological remains related to this
species, which consist of the skull, a hemimandible,
a scapula, some vertebrae and some ribs, were found
inside a dark glass bottle of beer among the ruins of
the Ag. Georgios village. Some peculiar features,
such as the presence of the notch on the external side
of the upper incisors, the presence of only two rows
of tubercles on the molars of the hemimandible and
of a single root in the upper molar and, moreover,
the dimension of various finds, the appearance and
the reduced sizes of the braincase, have allowed the
attribution of these finds to the species M. musculus
(Toschi, 1965; Amori et al., 2008).
Suncus etruscus Savi, 1822
Suncus etruscus is the smallest living terrestrial
mammal, characterized by a weight of about 2 g
and by a length which rarely reaches to 5 cm (ex-
cluding tail). It is a typical species of Mediterranean
bio-climatic zones, where it lives in environments
characterized by dry stone walls and rocks (Amori
et al., 2008), situation which moreover has also
been found at Alimia. The distribution range of this
species includes countries of the Mediterranean
basin and extends to Pakistan and India. In the
African continent reaches Natal and Tanzania. Has
been found a single find of this small mammal, a
hemimandible, in the same bottle where the remains
relating to Mus musculus were found. The shape
and size of hemimandible and the height mandibu-
lar coronoid, less than 3.2 mm, were the distinctive
features for the attribution to the species S. etruscus
(Amori et al., 2008).
CONCLUSIONS
As already noted, due to difficulties in reaching
Alimia, the survey on the island may not have been
capillary. It seems that because the arid nature and
lack of active watercourses of this island, Amphi-
bians are totally absent. A comparable situation is
also in the nearby island of Chalki, which has very
similar environmental features (Buttle, 1995; Cat-
taneo, 2009). All specimens of various species of
reptiles were found in Ag. Georgios bay, among the
ruins of the abandoned village. Some specimens of
S. stellio and A. oertzeni have also been observed
in close proximity of a group of military buildings
abandoned in the peninsula of Tigani. The herpeto-
fauna of Alimia has proved to be interesting anyway
and, considering the small size of the island, sub-
stantially consistent. The species that hosts are
clearly of Rhodian matrix; includes two taxa more
than the nearby and much bigger (about six times)
island of Chalki (A. oertzeni and Ophisops elegans )
(Buttle, 1995; Cattaneo, 2007, 2009), but the latter
is more distant from Rhodes. The coexistence of
seven species of reptiles in this small island, dry and
without human presence, represents a perfect model
of sympatry and of optimal utilization of resources.
It is also worth noting that the dense interactive
network of the island (to whom contribute four
species of lizard and at least three of micromammals)
could also allow the survival to a second ophidic
species. In this regard is worth remembering that
Boettger (1888) reported the news, provided by von
Oertzen, about the possibility of the existence of
Montivipera xanthina in Chalki. Researches carried
by us and by others (Joger & Nilson, 2005) have
ruled out this possibility, but fact remains that the
indication of von Oertzen could refer to another
nearby island (in this instance Alimia), assimilated
to Chalki or confused with this, following a lapsus
linguae. Besides the authority and reliability of the
German author don’t put doubts about the authen-
ticity of the news. So that’s why is desirable in the
future that researches are carried out in this direc-
tion, in order to clarify the enigma.
ACKNOWLEDGMENTS
The authors wish to thank Flavio Rocchi and
Giuliano Milana for helping in the determination of
the finds related to micromammals.
REFERENCES
Angelici F.M., Pinchera F. & Riga F., 1992. First record
of Crocidura sp. and Mus domesticus and notes on
the Mammals of Astipalaia Island (Dodecanese,
Greece). Mammalia, 56: 159-161.
78
Mauro Grano etalii
Amori G., Contoli L. & Nappi A., 2008. Fauna d’ltalia.
Mammalia II: Erinaceomorpha, Soricomorpha,
Lagomorpha, Rodentia. Calderini, Bologna, 736 pp.
Baig K.J., Wagner R, Annajeva N.B. & Boheme
W., 2012. A morphology-based taxonomic revision
of Laudakia Gray, 1845 (Squamata: Agamidae).
Vertebrate Zoology, 62: 213-260.
Beutler A., 1981. Cyrtodactylus kotschyi. Agaischer
Bogenfingergecko. In: Bohme W. (Ed.), Handbuch
der Reptilien und Amphibien Europas. Band 1,
Echsen 1 . Akademische Verlagsgesellschaft, Wies-
baden, 53-74.
Blackman D., Rankov B., Baika K., Gerding H. &
Pakkanen J., 2014. Shipsheds of the Ancient Mediter-
ranean. Cambridge University Press, 624 pp.
Boettger O., 1888. Verzeichnis der von Hern E. von
Oertzen aus Griechenland und aus Kleinasien mitge-
brachten Batrachier und Reptilien. Sitzungsberichte
der koniglich preussischen Akademie der Wis-
senschaften zu Berlin, 5: 139-186.
Broggi M.F., 2002. Herpetological notes on the Do-
decanese islands of Symi and Sesklia (Greece).
Herpetozoa, 15: 186-187.
Buttle D., 1995. Herpetological notes on the Dodecanese
islands of Chalki and Symi, Greece. British Herpet-
ological Society Bulletin, 52: 33-37.
Cattaneo A., 1998. Gli Anfibi e i Rettili delle isole greche
di Skyros, Skopelos e Alonissos (Sporadi settentri-
onali). Atti societa italiana di Scienze Naturali del
Museo civico di Storia naturale di Milano, 139: 127—
149.
Cattaneo A., 2007., Appunti di erpetologia rodiota. Atti
del Museo civico di Storia Naturale di Trieste, 53:
11-24.
Cattaneo A., 2009. L’ofidiofauna delle isole egee di
Halki e Tilos (Dodecaneso) con segnalazione di un
nuovo fenotipo di Dolichophis jugularis (Linnaeus)
(Reptilia Serpentes). II Naturalista siciliano, 36:
131-147.
Cattaneo A., 2012. II colubro gola rossa dell’arcipelago di
Rodi: Dolichophis jugularis zinneri subsp. nova (Rep-
tilia Serpentes). II Naturalista siciliano, 36: 77-103.
Cattaneo C. & Grano M., in press. Indagine preliminare
sulla flora di Chalki e Alimia. Annali del Museo
Civico di Rovereto.
Di Turo P, 1984. Presenza della foca monaca ( Monachus
monachus) nell’area mediterranea con particolare rifer-
imento alia Puglia. Thalassia Salentina, 14: 66-84.
Official Gazette, 991 GG / B / 27/05/1999.
Iliadis K., 1950. H XdkKr|rr|<; AcoSeKavpaou (Iaxopia -
Aaoypacpia - pOpKaieOipa). AOpva, xopo<; A. siKoveg,
Xapxqg, 560 pp.
Joger U. & Nilson G., 2005. Montivipera xanthina (Gray,
1849) - Bergotter. In: Joger U. & Stiimpel N. (Eds.),
Handbuch der Reptilien und Amphibien Europas,
Bd. 3/IIB, Schlangen (Serpentes) III. Aula-Verlag,
Wiebelsheim, 63-76.
Kasapidis P, Magoulasa A., Mylonas M. & Zouros E.,
2005. The phylogeography of the gecko Cyrtopodion
kotschyi (Reptilia: Gekkonidae) in the Aegean ar-
chipelago. Molecular, Phylogenetics and Evolution,
3: 612-623.
Lowe S., Browne M., Boudjelas S. & De Poorter M.,
2000. 100 of the World’s Worst Invasive Alien
Species a selection from the Global Invasive Species
Database. Published by The Invasive Species Special-
ist Group (ISSG) a specialist group of the Species
Survival Commission (SSC) of the World Conserva-
tion Union (IUCN), 12 pp.
Marchessaux D. & Duguy R., 1977. Le phoque moine,
Monachus monachus (Hermann, 1799), en Grece.
Mammalia, 41: 419-440.
Massed M., 2012. Atlas of terrestrial mammals of the
ionian and aegean islands. De Gruyter, Berlin/
Boston, 302 pp.
Massed M., Sara M., 2002. Non-volant Terrestrial Mam-
mals on Mediterranean Islands: Tilos (Dodecanese,
Greece), a Case Study. Bonner zoologische Beitrage,
51: 261-268.
Mertens R., 1959. Zur Kenntnis der Lacerten auf der
InselRhodos. Senckenbergiana biologica, 40: 15-24.
Rackham O. & Vernicos N., 1991. On the ecological
history and future prospects of the island of Khalki.
In: Grove A.T., Moody J. & Rackham O., Creta and
the South Aegean Islands (effects of changing climate
on the environment). Geography Department,
Downing Place, University of Cambridge, E.C.
Contract EV4C-0073-UK, 347-361.
Rosier H., 2000. Zur Taxonomie und Verbreitung von
Cyrtopodion kotschyi (Steindachner, 1870) in Bul-
garien (Sauria: Gekkonidae). Gekkota, 2: 3-19.
Sindaco R. & Jeremcenko V.K., 2008. The Reptiles of the
Western Paleartic. 1. Annotated Checklist and Distri-
butional atlas of the turtles, crocodiles, amphisbaenians
and lizard of Europe, North Africa, Middle East and
Central Asia. Monografie della Societas Herpetologica
Italica - 1. Edizioni Belvedere, Latina (Italy), 580 pp.
Sokratis G., 2006. Geography guide of Greece. Forumers
of skyscraperscity.com. Athens, 89 pp.
Spaneli V. & Lymberakis P, 2014. First record of Stel-
lagama stellio (Linnaeus, 1758) from Crete, Greece.
Herpetology Notes, 7: 367-369.
Stefanini G. & Desio D., 1928. Le Colonie. Rodi e le
isole Italiane dell’Egeo. Utet, Torino, 463 pp.
Toschi A., 1965. Fauna d’ltalia - Mammalia VII: Lago-
morpha, Rodentia, Carnivora, Artiodactyla, Cetacea.
Calderini, Bologna, 647 pp.
Wilson M.J. & Grillitsch H., 2009. The herpetofauna of
Simi (Dodecanese, Greece) (Amphibia, Reptilia).
Herpetozoa, 22: 99-113.
Biodiversity Journal, 2015, 6 (1): 79-82
First record of Rugulina fragilis (Sars G.O., 1878) from the
Mediterranean Sea (Mollusca Gastropoda Pendromidae)
Francesco Giusti 1 , Carlo Sbrana 2 & Luigi Romani 3 *
'Via 25 Aprile 19 /E , 57017 Colle Salvetti, Leghorn, Italy; e-mail: jojcub@yahoo.it
2 Via Sette Santi 1, 57100 Leghorn, Italy; e-mail: car letto.nicchi@tisca li.it
3 Via delle ville 79, 55013 Lam mari, Lucca, Italy; e-mail: luigir omani78@gmail.com
Corresponding author
ABSTRACT Several shells of RllguUnCl fragilis (Sars G.O., 1 878) (Mollusca Gastropoda Pendromidae)
are reported from the Tuscan Archipelago. This is the first record of the species from the
Mediterranean Sea.
KEY WORDS Mediterranean Sea; new records; Pendromidae; Rligulina fragilis ; Tuscan Archipelago.
Received 02.02.2015; accepted 05.03.2015; printed 30.03.2015
INTRODUCTION
Pendromidae Waren, 1991 is a small family of
vetigastropods whose systematic position is not yet
well understood (Bouchet P. & Rocroi J.P., 2005).
It includes two genera, Pendwma Dali, 1927 and
Rugulina Palazzi, 1 988. The latter comprises few
species (Gofas, 2014), only two of them belong to
the European fauna: Rugulina fragilis (Sars G.O.,
1 87 8) and R. UlOUteWSatoi (van Aartsen & Bogi,
1987). Their troubled nom enclatural and taxonomic
histories are well documented by Waren (1991).
Only R. monterosatoi has previously been found in
the Mediterranean Sea.
MATERIAL AND METHODS
All material was picked up from bottom samples
trawled by local fishermen. Shells were studies with
a stereomicroscope. Photos were taken with a di-
gital photocamera. The protoconch whorls are
counted according to the method of Verduin (1977).
ABBREVIATIONS AND ACRONYMS. Dp:
total diameter of the pro to conch (in pm); H: max-
imum height (in mm); Nwp: number of whorls of
the protoconch; Nwt: number of whorls of the
teleoconch; W: maximum width (in mm); APC:
Attilio Pagli collection (Lari, Italy); CBC: Cesare
Bogi collection (Leghorn, Italy); CSC: Carlo
Sbrana collection (Leghorn, Italy); LCC: Lrancesco
Chiriaco collection (Leghorn, Italy); LGC:
Lrancesco Giusti collection (Leghorn, Italy);
RRC: Romualdo Rocchini collection (Pistoia,
Italy).
RESULTS AND DISCUSSION
Taxonomy
Class Gastropoda Cuvier, 1795
Subclass Vetigastropoda S alvini-Plaw en , 1980
Lamily Pendromidae Waren, 1991
G enus Rugulina P alazzi, 198 8
80
Francesco Giusti et alii
Rugulina fragilis (Sars G .0 1878) (Figs. 1-3,5)
Adeorbis fragilis G.O. Sars, 1878: 2 1 3, tab. 22, figs.
1 9a-c (Fig . 4)
Rugulina fragilis-. Waren, 1991: 71—73, figs 11A-E,
1 3 A , B
Rugulina fragilis-. Beck et al., 2006: 47
Rugulina fragilis-. Hoffman et al., 2010: 49, figs. 1-3
Original description. " Testa tenuis et fragilis,
albida, leviter rufescens, exacte trochiformis, spira
elevata, anfractibus 4 convexis, ultimo permagno
et amplo basi leviter applanata, sutura profunda,
apertura patula, oblique expansa, forma ovato-
elliptica, labro externo tenuissimo, obliquo,
columella cequaliter incurvata, umbilico magno et
projundo crista nulla a basi difinito. Superficies vix
nitida, lineis spiralibus, elevatis, regularibus
obducta. Diam. basis 2,0 mm; altit. 1,7 mm."
[The shell is thin and fragile, whitish, slightly
reddish, perfectly trochiform, with elevated spire, 4
convex whorls, the last is large and wide with the
base slightly flattened, suture deep, aperture wide,
obliquely expanded, ovate-elliptic, the external lip
is very thin, oblique, the columella is regularly
curved, the umbilicus is large and deep, there is no
keel on the base. Surface barely shining, regularly
covered with raised spiral lines. Diameter at the
base 2.0 mm; height 1.7 mm.]
Examined material. Rugulina fragilis : off
Capo Corso (Corsica, France) 600 m, 2 shells in
CSC, 6 shells in FG C , 1 shell in A PC . Rugulina cf .
fragilis-. off Capo Corso (Corsica, France) 600 m,
l shell in csc. Rugulina monterosatoi-. off
Gorgona Island (Feghorn, Italy) 400 m, 4 shells in
RRC, 1 shell in CSC, 1 shell in FCC; 3 shells off
Gorgona Island (Feghorn, Italy) 300 m, in APC; 13
shells, Alboran Sea (Spain) 160 m, in CBC; 4
shells off Giglio Island (Grosseto, Italy) 400 m, in
CBC; 4 shells off Capraia Island (Feghorn, Italy)
500 m , in FG C .
Description of the examined shells. Small,
thin and fragile, broadly conical (height: 1.00-1.65
mm;width: 1.15-2.05 m m ), w hitish and semitrans-
parent. Protoconch (0.5-0. 6 whorls; diameter about
185 pm) protruding, paucispiral, smooth, tilted,
border with the teleoconch clear. Teleoconch whorls
(2. 2-2. 6) convex, fairly expanding, suture deep.
Aperture broad, ovate, prosocline (seen laterally).
Outer lip sharp. Columella curved, simple. Base
quite flattened. Umbilicus wide. Sculpture of subtle
spiral threads (9-14 on the last whorl), more close-
set in the periumbilical zone. Shell surface further
ornamented with a somewhat net-shaped micros-
culpture of irregular discontinuous lines.
Remarks. The mediterranean shells match R.
fragilis in all respects. The only similar species is
the cogeneric R. monterosatoi, which is constantly
smaller (maximum W : 0.85 mm; maximum H: 0.80
mm). Its spiral sculpture has only 2 strong perium-
bilical cords forming a sort of keel and 1 adapical
thread. The protoconch are comparable in size (Dp:
about 185 pm) (Aartsen van & Bogi, 1987; Waren,
1991; pers. obs.) (Figs. 7-9). Comparing similar-
sized shells of R. fragilis and R. monterosatoi, the
latter has a more globular outline, a more depressed
spire, less expanded whorls, the outline of the last
whorl appears more squarish due to the spiral
sculpture, the umbilicus is smaller, and the proto-
conch less tilted (Figs. 5-6). Note that the shell in
fig. 6 is gerontic, being larger for the species and
having the last whorl slightly loose. It is neverthe-
less clearly different from R. fragilis.
A shell sim ilar to R. fragilis (Figs. 1 0- 1 3) w as
found in the same bottom sample (H: 1.20 mm;
W: 1.40 mm; Nwt: 1.7). It differs from the latter
in having a less conical outline, a stronger micros-
cultpure, a protoconch not tilted, larger in dia-
meter (Nwp: 0.6; Dp: about 270 pm). Being an
unique specimen, we prefer to leave its status
open .
Rugulina fragilis is previously known from the
Northern Atlantic Ocean, ranging from E Greenland
to Norway (Waren, 1991; Hoffman et al., 2010),
and the Seine seamount, off the morroccan coasts
(Beck et al., 2006). This is the first record from the
Mediterranean Sea. Although no living specimens
were found, the shells are in good conditions.
Rugulina fragilis should be added to the recent
mediterranean malacofauna.
ACKNOWLEDGMENTS
The authors are grateful to Cesare Bogi
(Feghorn, Italy), Romina Rocchini (Pistoia, Italy),
Attilio Pagli (Fari, Italy), Alessandro Raveggi
(Florence, Italy), Francesco Chiriaco (Feghorn,
Italy), Stefano Bartolini (Florence, Italy) for the
First record of Rugulina fragilis from the Mediterranean Sea (Mollusca Gastropoda Pendromidae)
81
Figures 1-3: Rugulina fragilis , off Capo Corso, 600 m, 1.90 mm x 1.50 mm; Fig. 1: frontal view; Fig. 2: apical view; Fig.
3: basal view. Fig. 4: AdeOfbis fragilis (from Sars, 1878, pi. 22, figs. 19a, modified). Fig. 5: R. fragilis, off Capo Corso, 600
m, 1.15 mm x 1.00 mm. Fig. 6: R. lllOllteWSatoi, off Giglio Island, 400 m, 1.00 mm x 1.00 mm. Figs. 7-9: R. monte WSatoi,
off Gorgona Island, 400 m, 0.77 mm x 0.77 mm; Fig. 7: frontal view; Fig. 8: apical view; Fig. 9: pro to conch. Figs. 10 - 13 :
R. c f. fragilis , same locality as Figs. 1 - 3 , 1.40 mm x 1.20 mm; Fig. 10: fro n tal view; Fig. 11: apical view; Fig. 12: pro to conch;
Fig. 13: details of the sculpture.
82
Francesco Giusti et alii
loan of material. Sincere thanks are due to Cesare
Bogi for useful advices, to Stefano Bartolini for di-
gital photographs and to Enzo Campani (Leghorn,
Italy) for reading the manuscript. Thanks to Patrick
LaFollette (Cathedral City, U.S.A.) for editing the
English manuscript.
REFERENCES
Aartsen van J. J. & BogiC., 19 87 . Dcironici mOTlteroSCltoi
a new Mediterranean gastropoda. Bollettino Malaco-
logico, 22: 27 3-27 6 .
Beck T., Metzger T. & Freiwald A., 2006. Biodiversity
inventorial atlas of macrobenthic seamount animals.
Eu-ESF project OASIS, 126 pp. Oceanic seamounts:
an integrated study; E V K 2 -C T-2 00 2 -00 0 7 3 , http://
wwwl.uni-hamburg.de/OASIS /Pages /publications/
BIAS.pdf.
Bouchet P. & Rocroi J.P., 2005. Classification and nomen-
clator of gastropod fam dies. M alacologia, 47: 1-397.
Gofas S., 2014. RllgulinCl Palazzi, 1988. Accessed
through: World Register of Marine Species at
http ://w w w.m arinespecies.org/aphia. php?p =
taxdetails& id= 1 3 8 326 on 2014-12-10
Hoffman L.,Van Heugten B. & Lavaleye M.S.S., 2010.
Skeneimorph species (Gastropoda) from the Rockall
and Hatton Banks, northeastern Atlantic Ocean.
Miscellanea M alacologica, 4: 47-61.
Sars G. O ., 1 878. Bidrag til kundskaben om Norges
arktiske fauna: 1. Mollusca regionis Arcticae Norve-
giae. Oversigt over de i Norges arktiske region
forekommende bloddyr. Christiania, A .W . Brpgger
XV + 466 pp., 34 pis.
Verduin A., 1977. On a remarkable dimorphism of the
apices in many groups of sympatric, closely related
m arin e g as tropod species. Baste ria, 41: 91-95.
Waren A., 1991. New and little known Mollusca from
Iceland and Scandinavia. Sarsia, 76: 53-124.
Biodiversity Journal, 2015, 6 (1): 83-86
Colinatys Ortea, Moro et Espinosa, 20 1 3 from Eastern Medi-
terranean Sea (Opisthobranchia Haminoeidae)
Luigi Romani 1 *, Stefano Bartolini 2 & Alessandro Raveggi 3
'Via delle ville 7 9, 55013 Lammari, Lucca, Italy; e-mail: luigir omani78@gmail.com
2 V ia Ermete Zacconi 16, 50137 Flo re nee, Italy; e-mail: stefmaria.bartolini@alice.it
3 V ia Benedetto Varchi 67, 50132 Florence, Italy, e-mail: sandro. firenze@ libero.it
Corresponding author
ABSTRACT Two shells of the genus ColillCltyS Ortea, Moro et Espinosa, 2013 (Opistobranchia Hami-
noeidae), similar to ColillCltyS Cllciyoi (Espinosa et Ortea, 2004), type species of the genus,
are reported from Larnaca, Cyprus. The presence of the species in the Mediterranean Sea
is discussed.
KEY WORDS Colinatys ■ H am inoeidae; new records; Mediterranean Sea.
Received 02.02.201 5; accepted 05.03.20 1 5; printed 30.03.20 1 5
Colinatys sp.
Examined material. 2 shells from Larnaca, Cy-
prus, depth 43 m, May, 2011, picked from bioclast ic
bottom samples collected by SCUBA near wreck of
ferry MS Zenobia, 34°53'52"N 33°39'25"E. Spe-
cimen 1 , H = 1.35 mm,W = 0.85 mm. (Figs. 1-4),
in Alessandro Raveggi collection; specimen 2, H =
1.60 mm,W = 1.20 (Figs. 1-6), in Stefano B artolini
co llec tio n .
Description. Shell small, translucent, colorless,
involutely coiled, subey lindrical-py riform , trun-
cated, periphery below center of the smoothly roun-
ded body whorl.Aperture longer than spire, narrow
posteriorly, widening anteriorly. Umbilicus narrow,
partially obscured by the slightly flared columellar
lip. Spire concave, nearly covered by final whorl,
leaving a narrow opening through which the proto-
conch can be seen. Outer lip sharp, straight to
slightly concave above periphery, convex below.
Sculpture of weakly encised, whitish spiral bands
of irregular widths, interrupted by stronger closely
packed orthocline axial growth lines, dividing the
bands into of rows of squared to elongated pits.
Within the apical depression only axial sculpture is
evident. The shell surface has a wrinkled, weakly
reticulated appearance. The whitish appearing
spiral bands are visible within the aperture through
the translucent shell (Figs. 1-6).
DISCUSSION
No European O pisthobranchs nor alien indo-
pacific species recorded from Cyprus (Ozturk et al.,
2004; Tsiakkiros & Zenetos, 2011) have similar
shells.
Considering that many alien marine organisms
have settled in the Eastern Mediterranean during
recent years (Zenetos et al., 2010), an extensive
bibliographic survey was carried out of shelled
opisthobranchs of the Indo-Pacific and neighboring
areas but was unsuccessful in finding similar
species (Issel, 1 869; Hedley, 1 899a-c; Habe, 1964;
84
Luigi Romani et alii
Maes, 1967; Keen, 1971; Kay, 1979; Powell, 1979;
Kilburn & Rippey, 1 982; Sharabati, 1984; Lin &
Qi, 1 985; Springsteen & Leobrera, 1 986; Kay &
S choenberg-D ole, 1991; Higo et al., 1 999, 200 1;
Jansen, 2000; Okutani, 2000; Hasegawa, 2001,
2005; Hasegawa et al., 2001a-b; Qi, 2004; Dharma,
20 0 5; Thach, 2005; Poppe, 2008; Sasaki, 2008;
Valdes, 2 0 08; Yonow, 20 0 8, 2012; Zenetos et al.,
2010 ).
Surprisingly, we found that our shell most closely
resem ble ColiflCltyS Cllciyoi (Espinosa et O rtea, 2004)
(Figs. 7-11), known from Cuba, Florida and
Bahamas (Espinosa & Ortea, 2004; Ortea et al.,
2013; Red fern, 2013). The genus ColinatyS Ortea,
Moro et Espinosa, 2013 was erected for this species
on anatomical grounds, which was originally as-
signed to AtyS Montfort, 18 10, then transferred to
RetUSCl T. Brown, 1827 (Valdes et al., 2006; Rosen-
berg et al., 2009). No other species have been assigned
to the genus.
Figures 1-6. Colincitys sp.,Larnaca (Cyprus).Figs. 1-4: 1.35 m m , Fig s. 5 , 6 : 1.60 mm; Figs. 7-11. Colincitys aluyoi (E spino sa
et Or tea, 2004), Bahamas, Figs. 7, 8: 1.50 mm, Figs. 9, 10: 1.00 mm. Figs. 11: 2.00 mm (from Redfern, 2013, modified).
Colinatys Ortea,Moro et Espinosa, 2013 from Eastern Mediterranean Sea (Opisthobranchia Haminoeidae)
85
Our shells match agree with the conchological
characters of Colinatys, but we prefer not to assign
them to alayoi as doubts on conspecificity remain
due to differencens of the shell colour pattern (C.
Cllayoi has a more marked “checkerboard” pattern),
absence of anatomical information and very long
distance from typical range. Additional material,
particularly live collected specimens for anatomical
comparison, are needed to establish the presence of
an established population and to clarify its status
and relationships. Whether the present species is
Mediterranean, Lessepsian, or of other origin is
unknown, so we prefer to consider Colinatys sp. a
cryptogenic species (Carlton, 1996).
ACKNOWLEDGEMENTS
We thanks Maria Scaperrotta (Florence, Italy)
who sorted the sediment samples and A ttilio Pagli
(Lari, Italy) for his help in searching for biblio-
graphical sources. We are grateful to Leopoldo
Moro (Santa Cruz de Tenerife, Spain), Jesus Ortea
(Oviedo, Spain) and Enzo Campani (Leghorn,
Italy) for useful suggestions, Thanasis Manousis
(Epanomi, Greece), John Varnava (Dhekelia,
Cyprus), Costantinos Kontadakis (Athens, Greece)
for providing information. We are endebted to
Patrick I LaFollette (Cathedral City, U.S.A.) for
the critical reading and the english revision of the
manuscript. Colin Redfern (Boca Raton, U.S.A.)
kindly has given permission to use the photos of
Colinatys alayoi.
REFERENCES
Carlton J.T, 1996. Biological invasions and cryptogenic
species. Ecology, 77: 1 653- 1 655.
Dharma B., 2005. Recent & fossil Indonesian shells.
H ackenheim , Ge rm any: ConchB ooks, 424 pp.
Espinosa J. & Or tea J., 2004. Nuevas e species de molus-
cos gasteropodos m a r in os (M oil u sea: Gastropoda) de
las Bahamas, Cuba y el Mar Caribe de Costa Rica.
Revista de la Academia Canaria de Ciencias, 15:
207-2 16.
Habe T., 1964. Shells of the W ester n Pacific in color. Vol.
II. Osaka, Hoikusha. 23 3 pp.
Hasegawa K., 2001. Deep-Sea Gastropods ofTosa Bay,
Japan, Collected by the R/V Kotaka-M aru and
Tansei-M aru during the Years 1 997-2000. National
Science Museum monographs, 20: 121-165.
Hasegawa K., 2005. A Preliminary List of Deep-Sea
Gastropods Collected from the Nansei Islands,
Southwestern Japan. National Science Museum
monographs, 29: 137-190.
Hasegawa K ., Hori S. & Ueshima R., 2001 a. A Prelim-
inary List of Sublittoral Shell-bearing Gastropods in
the Vicinity of Shimoda, Izu Peninsula, Central
Honshu, Japan. Memoirs of the National Science
Museum, 37: 203-228.
Hasegawa K ., Saito H ., Kubodera T. & Xu F., 200 1b.
Marine Molluscs Collected from the Shallow Waters
of Hainan Island, South China Sea, by China-Japan
Joint Research in 1 997. National Science Museum
monographs, 21: 1-44.
Hedley C., 1899a. The Mollusca of Funafuti. Part I.
Australian Museum Memoir, 3: 395-488.
Hedley C., 1899b. The Mollusca of Funafuti, Part II.
Australian Museum Memoir, 3: 489-510.
Hedley C., 1899c. The Mollusca of Funafuti (supple-
ment). Australian Museum Memoir, 3: 547-570.
Higo S., Callomon P. & Goto Y., 1 999. Catalogue and
bibliography of the marine shell-bearing Mollusca of
Japan. Elle Scientific Publications Osaka, 749 pp.
Higo S ., Callomon P. & Goto Y., 2001. Catalogue and
Bibliography of the Marine Shell-Bearing Mollusca
of Japan. Gastropoda Bivalvia Polyplacophora
Scaphopoda Type Figures. Elle Scientific Publica-
tions, Yao, Japan, 208 pp.
IsselA., 1869. Malacologia del Mar Rosso. Biblioteca
Malacologica, Pisa, 387 pp., 5 pis.
Jansen P., 2000. Seashells of south-east Australia.
Capricornia Publications, Lindfield, NSW, 118 pp.
Kay A.E., 1979. Hawaiian M a line Shells. Reef and Shore
Fauna of Hawaii, Section 4: Mollusca. Bernice Pauahi
Bishop Museum Special P u blic atio n.64: 653 pp.
Kay A.E. & Schoenberg-Dole O., 1991. Shells of Hawaii.
University of Hawaii Press, Honolulu, 87 pp.
Keen A.M., 1971. Sea Shells ofTropical West America:
Marine Mollusks From Baja California to Peru.
Second edition. Stanford University Press: Stanford,
C A . xiv + 1 064 pp .
Kilburn R.N. & Rippey E., 1982. Seashells of Southern
Africa. MacMillan, Johannesburg, i-xi. 1-249 pp.
Lin G. & Qi Z . , 1985. A preliminary survey of the
Cephalaspidea (Opisthobranchia) of Hong Kong and
adjacent waters. In: M orton B . & Dudgeon D . (Eds.).
The M alacofauna of Hong Kong and Southern China.
II. Hong Kong: Hong Kong University Press, 113 —
124.
M aes V.O., 1967. The littoral marine mollusks of Cocos-
Keeling Islands (Indian Ocean). Proceedings of the
Academy of Natural Sciences of Philadelphia, 119:
93-2 1 7, 26 pis.
Okutani T., 2000. Marine Mollusks of Japan. University
Press Tokai. Tokyo, xlviii + 1173 pp.
86
Luigi Romani et alii
Ortea J., Moro L. & Espinosa J., 2013. Nueva especie de
Notodiaphana Thiele, 1931 del Oceano Atlantico y
nueva ubicacion generica para AtyS Cllciyoi Espinosa
& Ortea, 2004 (Gastropoda: O p is th o b ra n c h ia : Ceph-
alaspidea). Re vista de la Academia Canaria de
Ciencias,25: 15-24.
Oztiirk B., Buzzurro G. & Avni Benli H., 2004. Marine
molluscs from Cyprus: new data and checklist.
Bollettino Malacologico, 39: 49-78.
Poppe G. T., 2008. Philippine M arine M ollusks. Volume
3 (Gastropoda - Part III). Conchbooks. Hackenheim,
Germany, 702 pp., 708-1 01 4 pis.
Powell A.W.B., 1979. The New Zealand Mollusca:
Marine Land and Freshwater Shells. Collins,
Auckland, 468 pp., 80 pis.
Qi Z., 2004. Seashells of China. China Ocean Press.
Beijing, viii + 418 pp.
Red fern C., 2013. Bahamian Seashells: 1161 Species
from Abaco, Bahamas. Bahamianseashells.com,
Boca Raton, Florida, 501 pp.
Rosenberg G., Garcia E.F. & Moretzsohn F., 2009.
Gastropods (Mollusca) of the Gulf of Mexico. In:
Felder D.L. & Camp D .K . (Eds.), The Gulf of
Mexico: Origins, Waters and Marine Life. Texas A.
&M.University Press, 579-699.
Sasaki T., 2008. Micromolluscs in Japan: taxonomic
composition, habitats, and future topics. Zoosympo-
sia, 1 : 147-232.
Sharabati D., 1984. Red Sea Shells. KPI Limited London,
12 8 pp .
Springsteen F.J. & Leobrera F.M ., 1 986. Shells of the
Philippines. Carfel Seashell Museum, Manila, 377 pp.
Thach N .N ., 2005. Shells of Vietnam. Conch Books,
Hackenheim, Germany, 3 3 8 pp + 91 pis.
Tsiakkiros L. & Zenetos A., 2011. Further additions to
the alien mollusc fauna along the Cypriot coast: new
opisthobranchia species. Acta A driatica, 52: 1 15-124.
Valdes A., 2008. Deep sea “ c e p h a la s p id e a n ” hetero-
branchs (Gastropoda) from the tropical southwest
Pacific. In: Heros V., Cowie R .H . & Bouchet P.
(Eds.), Tropical Deep-Sea Benthos 25. Memoires du
Museum national d'H is to ire naturelle, 196: 587-792.
Valdes A., Ham an n J., Behrens D. & DuPont A. 2006.
Caribbean Sea Slugs. A Field Guide to the Opistho-
branch M ollusks from the Tropical Northwestern
Atlantic. Sea Challengers, Gig Harbor, WA, 289 pp.
Yonow N ., 2008. Sea Slugs of the Red Sea. Pensoft
Series Faunistica, 304 pp.
Yonow N . , 2012. Opisthobranchs from the western
Indian Ocean, with descriptions of two new species
and ten new records (Mollusca, Gastropoda). Zoo-
Keys, 1 97: 1-129.
Zenetos A., Gofas S., Verlaque M., Cinar M.E., Raso
J.E.G., Bianchi C .N ., Morri C ., Azzurro E., Bile-
cenoglu M ., Froglia C., Siokou I., Viola n ti D., Sfriso
A., San Mar tin G.,GiangrandeA., Katagan T., B alles-
tero s E ., Ramos-EsplaA.,Mastrototaro F., Ocana O.,
Zingone A. & Gambi N ., 2010. Alien species in the
Mediterranean Sea by 2010. A contribution to the
application of European Union's Marine Strategy
Framework Directive (MSFD). Part I. Spatial distri-
bution. Mediterranean Marine Science, 11: 381-493.
Biodiversity Journal, 2015, 6 (1): 87-94
Reports of Haliotis Linnaeus, 1 758 (Mollusca Vetigastropoda)
from the Middle Miocene of Ukraine
Maurizio Forli 1 *, Alexander Stalennuy 2 & Bruno Dell’Angelo 3
'Via Grocco 16, 59100 Prato, Italy; e-mail: info@dodoline.eu
2 Bandem 90/4, 46011 Ternopil, Ukraine; e-mail: stalenmiy@rambler.ra
3 Via Santelia 55, 16153 Genoa, Italy; e-mail: brano.dellangelo@chitons.it
’Corresponding author
ABSTRACT Two species of Haliotidae are described and illustrated from the Maksymivka quarry near
Ternopil (Ukraine), a site characterized by its peculiar Middle Miocene (Badenian) coralgal
facies. The first species, Haliotis volhynica Eichwald, 1829, has a wide geographical
distribution that extends from the Paratethys of Central Europe to the Ukraine, and is quite
common in the Maksymivka site. Another different species of Haliotis Linnaeus, 1758 was
recently found at Maksymivka, only two specimens in several years of research. This
species was already reported by Krach (1981) from Poland as Haliotis tuberculata tauro-
planata Sacco, 1897, a species from the Burdigalian of Piedmont that differs from the
Maksymivka species by several characters. We leave this rare species indeterminate at
specific level because of the scarcity of material known to date.
KEY WORDS Gastropoda; Haliotidae; Haliotis', Miocene; Ukraine.
Received 02.02.2015; accepted 05.03.2015; printed 30.03.2015
INTRODUCTION
Haliotidae Rafinesque, 1815 is a family of
marine gastropods consisting of 56 living species
and at least 35 fossil ones (Geiger & Groves, 1999;
Geiger, 2000). The genus Haliotis Linnaeus, 1758
is the only one for the family and is known from
Upper Cretaceous (Maastrichtian) (Sohl, 1992) to
Recent. Strausz (1966) and Geiger & Groves
(1999) proposed to refer all the 11 fossil European
taxa known at that time (Haliotis anomiaeformis
Sacco, 1896; H. benoisti Cossmann, 1895; H.
lamellosa Lamarck, 1822; H. lamellosoides Sacco,
1896; H. monilifera Bonelli, 1827; H. neuvillei
Bial de Bell, 1909; H ovata Bonelli, 1827; H
tauroplanata Sacco, 1897; H. torrei Ruggieri,
1989; H. tuberculata Linnaeus, 1758; H. volhynica
Eichwald, 1829) to H. tuberculata volhynica ,
because the Recent species (H. tuberculata tuber-
culata) with its Atlantic and Mediterranean popu-
lations is known to be extremely plastic in its shell
morphology, and most material of European fossil
specimens fall within the range of variation within
the Recent species.
Aim of the present work is to illustrate some
specimens of Haliotis recently found at the Mak-
symivka quarry (Ukraine) from the Middle
Miocene (Badenian). A revision of the fossil cited
species of Haliotis is needed to define the status of
the described taxa. The reports of Haliotis from the
Miocene of Ukraine are scarce, relating to H.
volhynica or H. tuberculata volhynica (Zelinskaya
et al., 1968; Krach, 1981) and H. tuberculata tauro-
planata (Krach, 1981).
88
Maurizio Forli et alii
MATERIAL AND METHODS
The Maksymivka quarry near Ternopil
(Ukraine) (Fig. 1) is well known in literature for its
peculiar Middle Miocene (Badenian) coralgal facies
and its fauna (Radwanski et al., 2006; Studencka &
Jasionowski, 2011; Gorka et al., 2012). It embraces
an area of several square kilometers over a distance
of about one kilometer (Radwanski et al., 2006: fig.
3). The reef exposed in this quarry is a member of
the unique reef structure (almost 300 km long)
formed within the Paratethyan realm, and distribu-
ted widely in the north-eastern and eastern borders
of the Carpathian Foredeep Basin in Western
Ukraine, Moldova and north-east Romania (Gorka
et al., 2012: fig. 1). The coralgal facies at Mak-
symivka is characterized by a complex structure:
particular coralgal buildups of variable size (from
centimetres of rodolith forms, to several metres
thick), composed of red-algal (lithothamnian)
colonies associated locally with sparse hermatypic
corals. A survey of all these peculiar features/com-
ponents is well reported by Radwanski et al., 2006.
Other organisms associated with reefs are represen-
ted by mollusks (bivalves and gastropods), crabs,
foraminifera, annelids, bryozoans and echinoderms.
Almost all of the organisms of originally aragonitic
shells were dissolved as a result of post sedimentary
diagenesis and are now preserved in the form of
moulds and/or imprints (see Gorka et al., 2012: fig.
7A for a massive coralline-algae boundstone with
embedded Haliotis shells).
The shells were collected manually inside the
reef structure, paying particular attention to
prevent breakage, and after were cleaned and
measured (Table 1). The measurements are in
millimeters (mm) and in degrees for the angle.
About the references we considered only those
works where species have been not only recorded,
but also figured.
ABBREVIATIONS AND ACRONYMS. AS:
Alexander Stalennuy collection, Temopil, Ukraine;
BD: Bruno Dell’Angelo collection, Genoa, Italy;
BS: Bellardi and Sacco collection, Museo di Geo-
logia e Paleontologia, University of Turin (now
stored at the Museo Regionale di Scienze Naturali
BELARUS
UKRAINE
Matj|ymivka/Temopirs'ka oblSs^craina^ Maksymivka
RUSSIA
49'36’1 3.49"N 25'54'32.70"E elev 364 m Alt 3.48 km
Figure 1. Maksymivka quarry near Ternopil (Ukraine), from Google earth.
Reports of Haliotis Linnaeus, 1 758 (Mollusca Vetigastropoda) from the Middle Miocene of Ukraine
89
of Turin), Italy; MF: Maurizio Forli collection,
Prato, Italy; MZB: Museo di Zoologia, University
of Bologna, Italy; L: length; W: width; H: max-
imum height of the shell, measured from the base to
the top of the spire; a: angle of inclination of the ini-
tial part of the spire; LAV: ratio between L and W.
SYSTEMATICS
Classis GASTROPODA Cuvier, 1795
Ordo VETIGASTROPODA Salvini-Plawen et
Haszprunar, 1987
Familia HALIOTIDAE Rafmesque, 1815
Genus Haliotis Linnaeus, 1758
Type species: Haliotis asinina Linnaeus, 1758
Haliotis volhynica Eichwald, 1829
Figs. 2-14
1829. Haliotis volhynica Eichwald: 294, pi. 5, fig. 18.
1856. Haliotis volhynica Eichw. - Hornes: 510, pi.
46, fig. 26
1928. Haliotis volhynica Eichw. - Friedberg: 530,
pi. 34, figs. 8, 9
1937. Haliotis volhynica Eichwald - Davidaschvili:
540, pi. 1, fig. 5
1954. Haliotis tuberculata lamellosoides Sacco -
Csepreghy-Meznerics: 10, pi. 1, fig. 24
1955. Haliotis (Haliotis) volhynica Eichw. - Korob-
kov: pi. 2, fig. 3
1960. Haliotis (Haliotis) tuberculata var. lamel-
losoides Sacco - Kojumdgieva & Strachimirov:
84, pi. 28, fig. 9
1966. Haliotis tuberculata volhynica Eichwald-
Strausz: 26, fig. 16c
1967. Haliotis volhynica Eichw. - Bielecka: 132, pi.
8, figs. 3, 4 (fide Bahik, 1975).
1968. Haliotis volhynica Eichwald - Zelinskaya et
al.: 95, pi. 27, fig. 1
1979. Haliotis (Sulculus) volhynica Eichwald -
Jakubowski & Musial: 61, pi. 5, fig. 5
1981. Haliotis tuberculata Eichwald - Krach: 39,
pi. 11, figs. 1-3
2012. Haliotis tuberculata Linnaeus - Gorka et al.:
163, figs. 7a, 15a, b
Table 1 . Measurements of the examined specimens and their repository. See in Abbreviations and Acronyms.
90
Maurizio Forli et alii
Figures 2-14. Haliotis volhynica Eichwald, 1829 from the Middle Miocene (Badenian) of Maksymivka quarry (Ukraine).
Figures 2-4: specimen 4 from Table 1. Figure 5: specimen 10 from Table 1. Figures 6, 7: specimen 1 1 from Table 1. Figures
8-10: specimen 6 from Table 1. Figures 11-14: specimen 1 from Table 1.
Reports of Haliotis Linnaeus, 1 758 (Mollusca Vetigastropoda) from the Middle Miocene of Ukraine
91
Figures 15-18. Haliotis sp. from the Middle Miocene (Badenian) ofMaksymivka quarry (Ukraine). Fig. 15: specimen 1 from
Table 1. Figures 16-18: specimen 2 from Table 1. Figures 19-22. Haliotis tuberculata tauroplanata Sacco, 1897, Early
Miocene (Burdigalian) of Torino hills (Piedmont, Italy), BS. 082.01.004.
92
Maurizio Forli et alii
Examined Material. Maksymivka: 15 specimens
(MF, BD, MZB) (Table 1).
Description. Shell of medium size (L max about
70 mm), widened-oval, spire slightly raised, tilted
about thirty degrees relative to the plane formed by
the edge of the shell (a). Apex positioned on the
left, moved to the center, about a third of the spiral
width corresponding to that point. Regularly con-
vex outer surface, with ornamentation spiral consti-
tuted by main cords detected, which develop at the
beginning almost intermittently and then, becoming
evident and irregularly wavy both in the sense of
the spire and in height, form knobs scattered or
aligned in radial folds more or less signed. The
spiral cords are separated by evident furrows,
sometimes side by side, giving rise to spiral cords
of smaller width of the main, intersecting with
growth striae more or less marked, form small
imbricated lamellae, arranged irregularly. Openings
with marginal conical-tubular protrusions pro-
nounced, regularly spaced from each other by
spaces almost equal to that of the size of the base
of the next opening, which maintain the propor-
tions, as the size of the shell, of which the last four
open. Outer lip, from the edge of the keel formed
by the openings up to the basal cords more
pronounced, tilted and slightly concave. Columellar
callus flattened with weak concavity.
Distribution. Haliotis volhynica Eichwald, 1829
is a species with a wide geographical distribution
that extends from the Paratethys of Central Europe
(Austria, Romania, Bulgaria, Poland) to the
Ukraine, with chronostratigraphic distribution lim-
ited to the Middle Miocene. It is particularly abund-
ant in organogenic limestones of Ukraine related to
sediments deposited at depths corresponding to the
infralitoral.
Remarks. The morphological characteristics of H.
volhynica , at least of the specimens we examined
and compared to those from the literature, are fairly
constant, in particular the position of the apex and
the spiral evolution, which is more rounded and
closed with respect to that of Haliotis sp., as well
as the ornamentation consisting of marked and
tubercolate spiral cords (Csepreghy-Meznerics,
1954; Zelinskaya et al., 1968; Krach, 1981; Gorka
et al., 2012). Even by the drawings can be found
the same morphological characters that seem
peculiar to H. volhynica , such as in Strausz (1966)
or in Friedberg (1928) by the shape of the internal
moulds of the shell.
Specimens found at Maksymivka are well char-
acterized and show a small degree of variability,
and are different from the Recent species H. tuber-
culata tuberculata L., 1758 with its Atlantic and
Mediterranean populations, contrary to what
previously expressed by many authors about H.
volhynica, considered as a subspecies H. tubercu-
lata volhynica (Strausz, 1966; Geiger & Groves,
1999) or directly as H. tuberculata (Gorka et al.,
2012). Agreeing to Landau et al. (2003) we consider
H. volhynica a separable species from the Recent
H. tuberculata.
Haliotis sp.
Figs. 15-18
1981. Haliotis tuberculata tauroplanata [non Sacco,
1897] - Krach: 40, pi. 11, figs. 4-7.
Examined Material. Maksymivka: 2 specimens
(AS, MF) (Table 1).
Description. Shell of small size, elongate-oval, with
width about half the length. Spire little high with
apex positioned a little more to the left than H.
volhynica, about a quarter of the spiral width cor-
responding to that point. Regularly convex surface
of the shell, with spiral ornamentation of the first
two whorls consisting of 4-5 slender main cords
spaced between them, which then become about ten
or so, just signed, sometimes forming very small
knobs, which disappear as increasing the spiral size,
leaving the remaining surface of the shell, smooth
or with sparse spiral cords just signed. Openings, of
which the last four open, with marginal conical-
tubular protrusions less raised. Outer lip, from
under the keel of the openings to the edge columel-
lar, with slightly convex profile. It was not possible
to examine the internal part because it is filled with
cemented limestone.
Distribution. Paratethys, Middle Miocene (Bad-
enian): Poland, Weglinek (Krach, 1981); Ukraine,
Maksymivka (this work).
Remarks. Only two specimens of a second species
of Haliotis were recently found at Maksymivka,
and this despite the numerous samples taken in
recent years by one of the authors (AS). This second
Reports of Haliotis Linnaeus, 1 758 (Mollusca Vetigastropoda) from the Middle Miocene of Ukraine
93
species may therefore be considered quite rare,
unlike H. volhynica which instead is found quite
commonly, although it is hard to find complete and
in fair condition specimens.
Haliotis sp. from Maksymivka differs from H.
vohlynica mainly by a different ratio L/W (1.34-
1.58 for H. volhynica vs. 1.79 for Haliotis sp.) that
gives rise to a more open spire and by the different
ornamentation, without radial folds and evident
spirals cords, if not in the early part of the spire. The
similarity with H. tuherculata tauroplanata Sacco,
1897 from the Early Miocene (Burdigalian) of
Turin hills (Piedmont, Italy) (Figs. 19-22) is
evident when considering the general shape of the
shell, but it is a bit less when comparing the two
different spiral ornamentations. In Haliotis sp. the
first part of the spire shows some slender spiral
cords and small tubercles which disappear as
increasing the spiral size, while there are not present
in H. tuberculata tauroplanata which has also a
spiral ornamentation made up of flattened cords
which extend over the entire surface of the shell.
Moreover Haliotis sp. has the anterior margin
almost straight, while that of H. tuberculata tauro-
planata is more convex and gives a more ellipsoid
profile to the shell.
We consider the specimen reported by Krach
from Weglinek (Poland) as H. tuberculata tauro-
planata conspecific to the present species.
CONCLUSIONS
The findings of H. volhynica from Maksymivka
confirm that abalone species, as taxa typical for
high-energy, rocky environments, are one of
the most characteristic and abundant group of
gastropods among Late Badenian free-living
reef-dwellers. Particularly important is to be con-
sidered the recent discovery of a second species of
Haliotis from the same site, only two specimens in
several years of research that led to the discovery
of many specimens of H. volhynica. This latter
seems the more frequent species of gastropods
found at Maksymivka, though most of the speci-
mens are not complete or are so matted in the rock
that they can not be extracted.
This second species, already reported in the past
by Krach (1981) from Poland as H. tuberculata
tauroplanata , show differences with the species of
Sacco from the Burdigalian of Piedmont, and must
be considered as a different species. We leave it
indeterminate at specific level because of the
scarcity of material known to date, waiting for more
material to give a specific determination and to
confirm or not the differences with the species from
the Miocene of Italy.
ACKNOWLEDGEMENTS
The authors wish to thank Daniele Ormezzano
(Museo Regionale di Scienze Naturali of Turin,
Italy) for helping during the visit to the Bellardi
Sacco collection.
REFERENCES
Baluk W., 1975. Lower Tortonian gastropods from
Korytnica, Poland. Palaeontologia Polonica, 32: 1-
186.
Bielecka M., 1967. Trzeciorzqd poludniowo-zachodniej
czqsci Wyzyny Lubelskiej [The Tertiary of the south-
western part of the Lublin Upland], Biuletyn
Panstwowego Instytutu Geologicznego, 206: 115-
188.
Csepreghy-Meznerics I., 1954. Helvetische und torton-
ische Fauna aus dem ostlichen Cserhatgebirge. An-
nales de flnstitut Geologique de Hongrie, 41: 1-185.
Davidaschvili L.S., 1937. On the ecology of animals of
the middle Miocene reefs of Ukrainian SSR.
Problems of Paleontology, 2-3: 537-563.
Eichwald E., 1829. Zoologia Specialis quam expositis
animalibus turn vivis, turn fossilibus potissimum
Rossiae in Universum, et Poloniae in specie... Typis
Josephi Zawadzki, Vilnae, 3 14 pp., 5 pis.
Friedberg W., 1911-1928. Miqczaki miocenskie ziem
Polskich. Czesc I. Slimaki i Lodkonogi. (Mollusca
Miocaenica Poloniae. Pars I. Gastropoda et Scapho-
poda). Museum Imienia Dzieduszyckich, Lwow-
Poznan, 631 pp.
Geiger D.L., 2000. Distribution and Biogeography of the
recent Haliotidae (Gastropoda: Vetigastropoda)
World-wide. Bollettino Malacologico, 35: 57-120.
Geiger D.L. & Groves L.T., 1999. Review of fossil
Abalone (Gastropoda: Vetigastropoda: Haliotidae)
with comparison to Recent species. Journal of
Paleontology, 73: 872-885.
Gorka M., Studencka B., Jasionovski M., Hara U.,
Wysocka A. & Poberezhskyy A., 2012. The Medobory
Hills (Ukraine): Middle Miocene reef systems in the
Paratethys, their biological diversity and lithofacies.
94
Maurizio Forli et alii
Biuletyn Panstwowego Instytutu Geologicznego,
449; 147-174.
Hornes M., 1851-1870. Die fossilen Mollusken des
Tertiar-Beckens von Wien. Abhandlungen der K. K.
Geologischen Reichsanstalt, Wien, 3: 1-42, pis. 1-5
(1851), 43-208, pis. 6-20 (1852), 209-296, pis. 21-
32 (1853), 297-384, pis. 33^10 (1854), 383-460, pis.
41-45 (1855), 461-736, pis. 46-52 (1856); 4: 1-479,
pis. 1-85 (1870).
Jakubowski G. & Musial T., 1979. Lithology and fauna
of the Middle Miocene deposits of Trzqsiny
(Roztocze Tomaszowskie Region, South-eastern
Poland). Prace Muzeum Ziemi, 32: 37-70, pis. 1-6.
Kojumdgieva E. & Strachimirov B., 1960. Tortonien; Le
Tortonien du type viennois. Les fossiles de Bulgarie,
7: 13-246.
Korobkov I. A., 1955. Spravochnik i metodicheskoe
rukovodstvo po tretichnym molljuskam. Brjuchonogie.
Gostoptechizdat, Leningrad, 795 pp.
Krach W., 1981. The Badenskie utwory rafowe na Roz-
toczu Lubelskim [The Baden reef formations in Roz-
tocze Lubelskie], Wydawnictwa Geologiczne, 121:
5-115.
Landau B., Marquet R. & Grigis M., 2003. The early
Pliocene Gastropoda (Mollusca) of Estepona,
southern Spain. Part 1 : Vetigastropoda. Palaeontos,
3: 1-87.
Radwanski A., Gorka M. & Wysocka A., 2006. Middle
Miocene coralgal facies at Maksymivka near Ter-
nopil (Ukraine): A preliminary account. Acta Geolo-
gica Polonica, 56: 89-103.
Sohl N. F., 1992. Upper Cretaceous gastropods (Fissu-
rellidae, Haliotidae, Scissurellidae) from Puerto
Rico and Jamaica. Journal of Paleontology, 66: 414-
434.
Strausz L., 1966. Die Miozan-Mediterranen Gastropoden
Ungams. Akademiai Kiado, Budapest, 693 pp.
Studencka B. & Jasionovski M., 2011. Bivalves from
the Middle Miocene reefs of Poland and Ukraine: A
new approach to Badenian/Sarmatian boundary in
the Paratethys. Acta Geologica Polonica, 61: 79-
114.
Zelinskaya V.A., Kulichenko V.G., Makarenki D.E. &
Sorochan E.A., 1968. Gastropod and scaphopod
mollusks of the Paleogene and Miocene of the
Ukraine. Paleontologiceskij Spravocnik, 2: 1-282.
Biodiversity Journal, 2015, 6 (1): 95-104
Planktonic and Fisheries biodiversity of Alkaline Saline crater
lakes of Western Uganda
Mujibu Nkambo 1 *, Fredrick W. Bugenyi 2 , Janet Naluwayiro 1 , Sauda Nayiga 3 , Vicent Kiggundu 1 , Godfrey
Magezi 1 &Waswa Leonard 4
'Aquaculture Research and Development Center-Kajjansi, National Fisheries Resources Research Institute (NaFIRRI), Uganda
department of Biological Sciences, College of Natural Sciences (CONAS), Makerere University, Kampala, Uganda
3 Islamic University In Uganda (IUIU)
4 Katwe Kabatoro community, Uganda
^Corresponding author, email: mnkambo@yahoo.co.uk
ABSTRACT Eight (8) selected saline crater lakes in Western Uganda were sampled for fish biodiversity.
Water samples collected from each of these lakes were analysed for zoo- and phyto-
planktonic composition and abundance. In situ, physico-chemical parameters including
average depth, salinity, temperature, conductivity, Dissolved Oxygen and pH were taken at
each sample collection point. The Mean ± SD of the different parameters ranged between
0.2±0.0 m and 2.3±0.3 m for average depth, 0.0±0.0 mgl' 1 and 205.0±15.3 mgl" 1 for salinity,
27.9±0.3°C and 34.4±2.4°C for temperature, 18.6±0.1 mscm-1 and 106.3±3.5 mscm-1 for
conductivity, 1.7±0.4 mgl 1 and 6.0±1.0 mgf 1 for Dissolved Oxygen and 9.6±0.1 and 11.5±1.0
for pH. With the exception of the Lakes Bagusa, where Anabaena circinalis Rabenhorst ex
Bornet et Flahaulwas found to dominate the algal biomass, and Bunyampaka and Nyamunuka
where no Spirulina platensis (Nordstedt) Gomont was found, the rest of the studied lakes had
S. platensis dominating their algal biomass. All lakes showed very low zooplankton abund-
ances and biodiversity, with Lake Kikorongo (the one with the highest zooplankton biod-
iversity) having Brachionus calyciflorus Pallas, 1766 as the most abundant, only ranging
between 50 to 100 individuals/litre. None of the lakes had fish at the time of sampling.
KEY WORDS Zooplankton; Phytoplankton; Fish; saline; alkaline; lakes.
Received 11.02.2015; accepted 18.03.2015; printed 30.03.2015
INTRODUCTION
Minute free-floating organisms found in various
water bodies can be referred to as planktons and
have been reported to be the main food for fish
(Lind, 1965). Planktons have been reported to play
pivotal roles in the biosphere in terms of both primary
and secondary production (Boero et al., 2008).
Plant-like minute organisms continuously drifting
in the water are referred to as phytoplankton while
the minute animal-like organisms, unable to syn-
thesize food are referred to as zooplankton. Plank-
tons are not only food organisms for fish fry, fin-
gerlings and adult fish but also influence key abiotic
features in aquatic systems (Joshi, 2009).
Saline systems have been reported to have a
generally low biodiversity (Hammer, 1986), with
diatoms being more dominant among algal biomass
in alkaline saline systems (Stenger-Kovacs et al.,
2014). Rotifera, Cladocera, Copepoda and Anostraca
species generally are the dominant zooplankton in
saline systems with their biodiversity decreasing
96
Mujibu Nkambo etalii
with increasing salinity (Hammer, 1993). In partic-
ular, East African saline lakes have been reported
to show more rotifers in their zooplankton as-
semblages than either Copepods or Cladocerans,
with the dominant species of Rotifera, Copepoda
and Cladocera reported to change with the salinity
gradient (Green, 1993). Alkaline saline crater lakes
are considered very productive environments
(Harper et al., 2003; Grant, 2006), with prokaryotic
photosynthetic primary production suggested to be
the driving force behind nutrient recycling in these
systems (Jones & Grant, 1999). Community even-
ness decreases with increasing nutrient concentra-
tion, with the few favored species being dominant
(Harper et al., 2003). Abundance of certain species
like Dunaliella sp. dominate the saline waters of
Utah lake in the USA (Larson & Belovsky, 2013),
while ‘ SpirulincC Arthrospira fusiformis (Voronikhin)
Komarek et J.W.G.Lund has been reported to be
dominant in lake Bogoria which is a hypersaline
lake in Kenya, east Africa (Harper et al., 2003;
Matagi, 2004) with no macro-zooplankton and
lesser flamingo, Phoeniconaias minor (E. Geoffroy
Saint-Hilaire, 1798), as the only grazers (Harper et
al., 2003).
Several fish species more especially amphi-
haline species have been described to have physiolo-
gical mechanisms which enable them to migrate
between freshwater and sea water, with many other
species with ability to tolerate, adapt or even accli-
mate to salinity, alkalinity and ionic compositions
levels outside the conventional freshwater and sea-
water conditions (Brauner et al., 2013). Whereas
both native and exotic species were found in waters
with salinities less than 30 mgU in a study of fish
distribution in inland saline waters in Victoria,
Australia, no inland fish species were found at salin-
ities above 30 mgl' 1 (Chessman & Williams, 1974).
Flamingo lakes with salinity levels below 20 mgl 1
are reported to have fish species of commercial
value (Hadgembes, 2006). Several species were
said to tolerate salinities as high as 15,000 mgl 1
with only the nine-spined stickle back resisting at
salinities of 20,000 mgl' 1 (Rawson & Moore, 1944).
Oreochromis alcalicns alcalicus (Hilgendorf,
1905), O. alcalicus grahami (Trewavas, 1983), and
O. amphimelas (Hilgendorf, 1905), have been
reported to be endemic in lakes Magadi and Natron
which are among the East African saline lakes
(Matagi, 2004).
A number of environmental factors including
salinity and nutrients in hypersaline systems may
be potential factors which do affect biodiversity in
saline environments (Larson & Belovsky, 2013).
Saline lakes show limited species complement in
micro-organisms contrary to the considerable biod-
iversity in micro-organisms (Harper et al., 2003).
Larson & Belovsky (2013) reported salinity and
nutrient concentration in hypersaline lakes as
among the strong determinants of phytoplankton
diversity, with species richness decreasing with
increasing salinity and increasing with increasing
nutrient concentration. Despite the inverse propor-
tionality between saline and aquatic biodiversity,
the relationship between salts is still not well
understood (Derry et al., 2003; Rios-Escalante,
2013). Contrary to the numerous fish and plank-
tonic biodiversity studies in fresh water systems,
very little of such studies has been conducted in
these unique saline systems (Jones & Grant, 1999;
Larson & Belovsky, 2013). The aim of this study
is, therefore, to investigate fish and planktonic bio-
diversity in selected saline crater lakes of western
Uganda as a way of providing more information on
fish and planktonic biodiversity in saline systems.
MATERIAL AND METHODS
Study area
Lakes considered in this study are small unique
water bodies found in Katwe-Kikorongo volcanic
field in western Uganda. Lake Katwe
(029.87033°E, 00.13217°S), is the largest among
these lakes with an average area of 2.5 km 2 (Nixon
et al., 1971). Other lakes considered in this study were
Katwe Munyanyange (029.8859 1°E, 00. 1351 3 °S),
Nyamunuka (029.98743°E, 00.09344°S), Bagusa
(030.17958°E, 00.09793°S), Murumuri (029.99186°E,
00.07323°S), Maseche (030.19019°E, 00.09355°S),
Bunyampaka (030.12819°E, 00.03765°S) and Kiko-
rongo (030.01228°E, 00.01 190°S). Among the
studied lakes, Bagusa and Kilcorongo were at the
lowest and highest altitude, 884 m and 939 m,
respectively, above sea level (a. s. 1). The majority
of these lakes are alkaline and saline in nature with
dominant anions being carbonates and sulphates
(Nkambo et al., 2015). These lakes exhibit consid-
Planktonic and Fisheries biodiversity of Alkaline Saline crater lakes of Western Uganda
97
erable temporal variations in volume and surface
area, with their total depth ranging between <1-6
m (Kirabira et al., 2013).
Zooplankton and phytoplankton diversity
Data collection in this study was done between
the 26th of February and 3rd of March, 2014, a
period towards the end of the dry season in this
region. A Global Positioning System (GPS) unit
(GARMIN 12XL) was used to take GPS coordin-
ates and the Altitude / elevation above sea level of
the different sampling points.
Zooplankton samples were obtained by filtering
four liters of water collected from every set geo-
referenced sampling point through a 50 pm mesh
zooplankton net. The samples obtained after filtra-
tion were preserved in 95% ethanol and earned to
the National Fisheries Resources Research Institute
(NaFIRRI) laboratory, Jinja, for identification to
the lowest possible taxonomic level and counted
under an inverted microscope. Using a Van Dorn
water sampler, water samples for phytoplankton
analysis were collected at a depth of 0.5 m in lakes
whose average depth was more than 1 m. For the
very shallow lakes (depth < 0.5 m), surface water
samples were collected for zoo and phytoplankton
analysis. 500 ml of the collected water samples
were preserved using Lugol’s solution in pre-rinsed
Nalgene bottles which were kept in a cooler box
containing dry ice and later transferred to the
National Fisheries Resource Research Institute
(NaFIRRI) laboratory in Jinja. In the laboratory,
phytoplanktons in the collected water samples
were identified to the lowest possible taxonomic
level and the wet biomass of each of the identified
group determined.
Selected physical and chemical parameters
(depth, temperature (T°C), dissolved oxygen
concentration (DO), pH, Conductivity (Cond) and
salinity) were also measured in-situ at the lake
surface and bottom. Where lakes were too shallow,
physico-chemical measurements were taken at the
lakes surface. Water temperature, dissolved oxygen
concentration and conductivity were measured
using a YSI oxygen/temperature/conductivity meter
(Model YSI 5 50 A), pH was determined using an
OAKTON pH Tester 30, while salinity was meas-
ured with a refractometer. The depth was determ-
ined using a portable depth finder (Hondex PS-7).
Fish diversity
All the study lakes, deeper than 0.5 m were
sampled for fish by setting gill nets and seine nets
in the evening at 5 pm and removing them in the
following morning at 7 am. In addition we also
asked to the people belonging to the communities
around each of the studied lakes whether they have
ever seen or got any fish from these lakes.
RESULTS
Physico-chemical parameters
Lakes Munyanyange, Nyamunuka, Murumuri,
and Bunyampaka were found to be very shallow
(depth<0.5 m) at the time of sampling, whereas,
lake Kikorongo was the deepest. The highest
measured dissolved oxygen (DO) was 6.0±1.0 mgl' 1
in Lake Kikorongo while Munyanyange and Muru-
muri had the lowest and second lowest DO (1 .7±0.4
and 1.7±0.5 mgl" 1 , respectively). All the sampled
lakes were found to be alkaline with pH ranging
between 9.58±0.1 (lake Bunyampaka) and 11.5±1.0
(Nyamunuka). The highest temperatures ranged
between 28.9±0.4°C and 34.4±2.4°C. Salinity was
between 0 mgl' 1 ( lake Kikorongo) and 205.0±15.3
mgl' 1 (Nyamunuka) Conductivity ranged between
10.5±0.6 mscm' 1 (Nyamunuka) and 106.3±3.5
mscm' 1 (Murumuri) (Table 1).
Phytoplankton diversity
A total of twenty nine (29) phytoplankton
species were found in the eight study lakes. Out of
them, nineteen (19) belonged to Cyanophyceae,
commonly known as Cynobacteria (blue-green
algae (BG)) which are predominantly photosynthe-
tic prokaryotes containing a blue pigment in addi-
tion to the chlorophyll (WHO, 1999). Six (6)
belonged to Chlorophyceae commonly referred to
as Chlorophyta (green algae (G)). Four (4) be-
longed to Bacillariophyceae, commonly referred to
as diatoms (D). Lakes Maseche (912,347 pgL" 1 ) and
Bagusa (210,290 pgL' 1 ) were found to have the
highest and second highest algal biomass. These
were followed by lakes Katwe and Murumuri
which had algal biomass of 90,653 pgL 1 and
86,240 pgL 1 , respectively. Lakes Katwe and
98
Mujibu Nkambo etalii
Lake
Depth (m)
D.O (mgl 1 )
pH
Temp. (°C)
Salinity (mgl 1 )
Cond (mscm 1 )
Katwe
2.1±0.7
2.6±0.2
9.9±0.1
27.9±0.3
180±67.8
104.5±6.4
Munyanyange
0.2±0.1
1.7±0.4
10.8±0.4
34.4±2.4
101.0±7.1
59.7±8.2
Nyamunuka
0.2±0.2
2.6±0.3
11.5±1.0
30.5±3.1
205.0±15.3
10.5±0.6
Murumuri
0.2±0.0
1.7±0.5
ll.lil.3
32.0±0.8
162.8±34.2
106.3±3.5
Bunyampaka
0.2±0.1
2.20±0.6
9.6±0.1
30.33±1.5
199.50±16.4
103.90±4.3
Bagusa
1.9±0.5
3.2±0.8
10.5±0.4
32.1±2.0
199.5±16.4
103.9±4.3
Maseche
1.3±0.2
2.9±0.4
10.9±0.4
30.0±0.7
92.3±7.6
71.2±1.3
Kikorongo
2.3±0.3
6.0±1.0
10.4±0.0
28.9±0.4
0 . 0 ± 0.0
18.6±0.1
Table 1. Mean±SD of the selected measured physico-chemical parameters in the selected studied saline crater lakes.
Bunyampaka showed the highest algal biodiversity
while lakes Maseche and Nyamunuka had the
lowest biodiversity. Cyanophyceae (BG) dominated
the algal composition of all the studied lakes. With
the exception of Lake Munanyange, were no Chloro-
phyceae species (G) were found, the other lakes
showed Chlorophyceae and Bacillariophyceae (D)
species in relatively small abundances in compar-
ison to Cyanophyceae. With the exception of Lake
Bagusa where Anabaena circinalis was found to be
dominant, and Bunyampaka and Nyamunuka where
no Spirulina platensis was found, in the rest of the
lakes S. platensis was dominant (Table 2).
Zooplankton diversity
Lakes Kikorongo, Maseche, and Katwe were
found to have zooplanktons belonging to Rotifera.
Lakes Maseche and Kikorongo showed also
zooplanktons belonging to Copepoda and none of
the studied lakes was found to have cladocerans at
the time of sampling. Lakes Munyanyange, Maseche,
Murumuli, Katwe and Nyamunukahad small cysts
which could not be identified. Water samples
from lakes Bunyampaka and Bagusa had neither
zooplankton nor un-identified cysts (Table 3).
Fish diversity
N one of the selected saline crater lakes con- si-
dered in this study had fish at the time of sampling.
With the exception of Lake Kikorongo, in which the
African catfish, Clarias gariepinus (Burchell,
1822), was observed (sometimes) during the rainy
season, none of the other studied lakes was reported
to have fish, ever.
DISCUSSION
Phytoplankton diversity
In the present study, blue-green (Cyanobacteria)
algae are dominant in all the lakes, with Spirulina
R J.F. Turpin ex M. Gomont being the most domin-
ant phytoplankton in the majority of them (Table 2).
This is in agreement with earlier studies done in
alkaline, saline crater lakes which reported Spirulina
platensis to be the most dominant (Hecky &
Kilham, 1973) in constrast with the reported domin-
ance of algal biomass by diatoms in anthroposaline
lakes in Romania and Bolivia (Stenger-Kovacs et
al., 2014). The dominance of Cyanobacteria in
these harsh environments can be attributed to their
ability to withstand extreme water conditions like
very high temperatures, pH and salinity. Some
Cyanobacteria species have got special adaptations
like ultraviolet absorbing sheath pigments which
increase their fitness in relatively exposed environ-
ments; indeed they have been reported to occur in
waters that are salty, brackish or fresh, in cold or
hot springs and in environments where no other
Planktonic and Fisheries biodiversity of Alkaline Saline crater lakes of Western Uganda
99
Taxa
Group
Taxa
Bagusa
Buny-
ampaka
Munya-
nyange
Maseche
Katwe
Kiko-
rongo
\va mu-
ii uka
Mil ru-
in uri
BG
Planktolyngbya limnetica
(Lemmermann) Komarkova-
Legnerova et Cronberg
15845
2,773
-
8,216
7,343
-
-
7,805
BG
Aphanocapsci sp.
522
274
548
-
342
-
365
730
BG
Spirulina platensis
(Nordstedt) Gomont
12,324
-
25,880
879,909
66,856
43,133
-
69,012
BG
Anabaena sp.
15,649
219
-
-
3,286
-
-
-
G
Stichococcus sp.
3.912
96
-
-
787
-
-
-
D
Centric diatoms
104
292
37
146
730
-
292
37
BG
Planktolyngbya circumcreta (G.S.
West) Anagnostidis et Komarek
440
411
1,438
-
-
-
205
411
BG
Chroococcus sp.
391
137
-
365
-
23
-
-
BG
Anabaena circinalis
Rabenhorst ex Bomet et Flahault
157,273
11.867
9,859
5,112
6,207
548
38,085
913
G
Stichococcus sp.
3,668
342
274
479
2,465
1,575
1,404
G
Nephrochlamys rostrata
Nygaard, Komarek, J.Kristiansen
et O.M. Skulberg
162
-
-
-
-
1,506
-
-
BG
Tiny blue green
-
101
151
-
-
228
-
-
BG
Anabaenopsis tanganyikae (G.S.
West) Woloszynska et V. V.Miller
-
55
-
-
-
-
-
-
BG
Pseudoanabaena sp.
-
411
411
-
103
-
308
-
BG
Anacystis limnetica
(Lemmermann) Drouet et Daily
-
183
365
-
274
137
342
297
BG
Romeria sp.
-
55
-
-
-
-
-
-
D
Nitzschia acicularis
(Kiitzing) W. Smith
-
274
-
-
23
-
137
23
G
Closterium acerosum
Ehrenberg ex Ralfs
-
-
411
411
205
205
411
-
BG
Oscillatoria tenuis
C. Agardh ex Gomont
-
-
-
17,253
-
-
-
-
BG
Aphanocapsa nubila
Komarek et H.J. Kling
-
-
-
456
-
91
-
183
BG
Coelosphaerium kuetzingian um
Nageli
-
-
-
-
1,826
-
-
4,747
BG
Planktolyngbya undulata
Komarek et R. Kling
-
-
-
-
103
-
-
-
G
Ankistrodesmus falcatus
(Corda) Ralfs
-
-
-
-
103
-
-
-
BG
Chroococcus dispersus
(Keissler) Lemmermann
-
-
-
-
-
68
-
-
D
Cyclostephanodiscus sp.
-
-
-
-
-
37
-
-
D
Navicula gastrum
(Ehrenberg) Kiitzing
-
-
-
-
-
856
-
-
BG
Merismopedia tenussima
Lemmermann
-
-
-
-
-
-
-
292
G
Monoraphidium contortum
(Thuret) Komarkova-Legnerova
-
-
-
-
-
-
22
BG
Aphanizomenon flosaquae Ralfs
ex Bomet et Flahault
-
-
-
-
-
-
-
365
Total
210,290
17,488
39,372
912,347
90,653
48,065
44,698
86,240
Table 2. Mean Wet biomass (pg/1) concentrations of the different phytoplanktons in the selected Alkaline,
saline lakes considered in this study. BG = Blue green algae, G = green algae, D = Diatoms.
100
Mujibu Nkambo etalii
microalgae occur (WHO, 1999). Allelopathy can
be another factor to explain the dominance of Cy-
anobacteria in these lakes. Freshwater Cyanobac-
teria like Oscillatoria sp. have been reported to
have exudates which can inhibit green alga
Chlorella vulgaris Beyerinck [Beijerinck] (Leao et
al., 2010). In the same way Cyanobacteria in these
saline crater lakes might be influencing the algal
biodiversity through allelopathy. Matagi (2004)
reported Spirulina ( Arthrospira fusiformis ) to be the
most successful algae in colonizing alkaline, saline
lakes found in the Eastern Rift valley. Lake Lonar,
an inland alkaline saline crater lake in India was
reported to have its phytoplankton biomass domin-
ated by Spirulina platensis (Satyanarayan et al.,
2007; Siddiqi, 2007; Yannawar & Bhosle, 2013).
Lakes Nakuru, Bogoria and Elmenteita, which are
alkaline-saline lakes in Kenya, were characterized
by mass growth of Cyanobacteria including Arth-
rospira fusiformis (Harper et al., 2003; Ballot et
al., 2004). The Presence of Cyanobacteria in these
alkaline-saline lakes is in conformity with the
findings of the present study where Spirulina domi-
nates the phytoplankton biomass. Jones & Grant
(1999) reported Spirulina spp. to be among the
main contributors to primary production in moder-
ately saline lakes while studies by Hadgembes
(2006) documented Spirulina to be one of the unique
Cyanobacteria occurring in East African saline lakes.
The extreme inhospitable conditions in alkaline,
saline crater lakes mean that the biodiversity in
these systems is limited to organisms with special
adaptations to survive such extreme conditions
(Matagi, 2004). Primary production in Flamingo
lakes of East Africa was reported to be dominated
by A. fusiformis with Ecloth iorhodospira sp. some-
times playing a key role (Jones & Grant, 1 999; Ma-
tagi, 2004). Nitzchia sp. and Navicula sp. are some
of the other algal species found in these lakes
(present study) or in other highly alkaline and saline
environments (Matagi, 2004). Chroococcus sp. is
another species of Cyanobacteria recorded in this
Munya-
yange
Kiko-
rongo
Maseche
Murumuli
Bunyam-
paka
Katwe
Bagusa
\va mu-
ii u ka
ROTIFERA
Brachionus calyciflorus
Pallas, 1766
-
+++
-
-
-
+
-
-
Brachionus angularis
Gosse, 1851
-
-
+
-
-
-
-
-
Lecane lima
(Muller, 1776)
-
+
-
-
-
-
-
-
Trichocerca cylindrica
(Imhof, 1891)
-
-
-
-
-
-
-
-
Synch eat a sp.
-
+
-
+
-
-
-
-
COPEPODA
nauplii
-
+
-
-
-
-
-
-
cyclopoid copepodite
-
-
+
-
-
-
-
-
CLADOCERA
Moina micrura
Kurz, 1874
-
-
-
-
-
-
-
-
unidentified cysts
+
-
+
+
-
+
-
+
Table 3. Mean zooplankton abundance (Individuals per litre) in the different studied saline crater lakes.
+ = 1 to 10 individuals /l, ++ = 10 to 50 individuals/1, +++ = 50 to 100 individuals/1, - = 0 individuals/1.
Planktonic and Fisheries biodiversity of Alkaline Saline crater lakes of Western Uganda
101
study which was reported by Jones & Grant (1999)
to play a key role in primary production in soda
lakes of East Africa.In studies aimed at reconstruct-
ing relationships between diatoms assemblages
with salinity, it was observed that salinity might not
be the primary cause of the shift in the diatom
assemblage but a factor highly related to drivers of
species shift (Saros & Fritz, 2000).
In the same way salinity might be playing a key
role in determining the Cyanobacteria species
occurring in the alkaline saline environments, but
other possible drivers should not be overlooked. In
a study to determine the extent to which salinity
influences community structures in saline systems,
salinity was reported to play a less significant role
in the determination of composition, species
richness and biodiversity (William, 1998). For
example, the highest algal biodiversity observed in
lakes Katwe and Bunyampaka and the least algal
biodiversity (in lakes Maseche and Nyamunuka)
might be attributed to the daily anthropogenic
disturbances experienced by the first two lakes
during the process of salt extraction, while the other
two are lakes with no daily anthropogenic disturb-
ance. It should be noted that communities around
lakes Katwe and Bunyampaka continuously extract
salt from these lakes on a daily basis and such
disturbances might be impacting various algal
species differently.
Zooplankton diversity
All the studied lakes showed a very low zoo-
plankton biodiversity, with only Brachionus calyci-
florus in lakes Kikorongo and Katwe; Brachionus
angularis in Fake Maseche, and Syncheata sp. in
lakes Kikorongo and Murumuli (the only species of
the phylum Rotifera present, see Table 3). Brachio-
nus plicatilis (Muller, 1786) and Paradiaptomas
africanus (Daday, 1910) (= Lovenula africana)
have already been reported to be characteristic zo-
oplankton in alkaline saline crater lakes (Hecky &
Kilham, 1973). Matagi (2004) listed Lovenula afri-
cana (Daday, 1910), Brachionus dimidiatus (Bryce,
1931), B. plicatilis (Muller, 1786) and chironomids
as the dominant macro-invertebrates in the highly
alkaline, hypersaline flamingo lakes of east Africa.
All of these were absent in our study. Nauplii in Fake
Kikorongo and cyclopoid copepodites in Fake Ma-
seche were the only Copepoda members observed.
Other macro-zooplanktons like cladocerans, neck-
ton fauna, and crustacean decapods were conspicu-
ously absent, probably due to the highly alkaline
pH. Fake Bogoria in Kenya which is an alkaline,
saline lake was reported to have no macro-zooplank-
ton with lesser Flamingo, Phoeniconaias minor as
the only grazer occasionally visiting these lakes in
high numbers (Harper et al., 2003). The absence of
crustacean decapods in a limnological study of lake
Fonar an alkaline, saline crater lakes in India was
attributed to a pH shock (pH > 10-11, alkaline
death point) (Siddiqi, 2007). An inverse proportion-
ality between biodiversity and salinity was reported
by Deny et al., (2003); Farson & Belovsky (2013),
and Rios-Escalante (2013); this might be the explan-
ation for the slightly high zooplankton biodiversity
in Fake Kikorongo since its salinity was found to
be very low at the time of sampling (Table 1). In
field and laboratory experiments designed to exam-
ine the consequences of climate-induced salinity
increases on zooplankton abundance and diversity
in coastal lakes, severe disturbances in zooplankton
community structure and abundance were caused
by even veiy small salinity changes, with even very
small increments in salinity capable of leading to
biodiversity depletion (Schallenberg et al., 2003).
Similarly, the low zooplankton biodiversity in these
lakes might be attributed to the high salinity levels
(see Table 1).
The unidentified cysts found in some of the
lakes might be dormant stages of zooplanktons
released as a mean of surviving extreme environ-
mental conditions. These dormant stages return to
life on the set of suitable conditions within the
lakes. Some zooplanktons, particularly members of
the order Anostroca lik eArtemia Feach, 1819, have
been reported to release dormant embryos in form
of cysts. Indeed, all Artemia species and strains
reproduce ovoviviparously (by generating live
nauplii) under favorable conditions, and oviparously
when dormant embryos are released in form of
cysts to withstand the harsh unfavorable envir-
onmental conditions (Ghomari et al., 2011; Ben
Naceur et al., 2012).
Fish diversity
Although some fish species like Oreochromis
alcalicus alcalicus, O. alcalicus grahami, and O.
amphimelas were reported to inhabit lakes Magadi
102
Mujibu Nkambo etalii
and Natron which are both alkaline and saline in
nature (Matagi, 2004), no fish were found in these
lakes during the study. Hecky & Kilham (1973)
also reported cichlid fish like Alcolapia grahami
(Boulenger, 1912) to occur in some of the alkaline,
saline lakes. The absence of fish in the different
studied lakes at the time of sampling can be attrib-
uted to the extreme environmental conditions like
the very high temperatures, salinity and alkalinity.
Previously, the complete absence of fish in Lake
Lonar was reported to be correlated with extreme
environmental physico-chemical parameters (Siddiqi,
2007). Many lakes, being shallow, experience very
high variations in volume and surface area with
some of these lakes reported to evaporate to dryness
during the extreme dry seasons (Nkambo et al.,
2015). This makes difficult, if not impossible, for
fish to survive during the dry seasons. Moreover,
the absence of appropriate food organisms in form
of zooplankton (i.e. Brachionus plicatilis ) could be
another probable reason for the absence of fish (see
Table 3).
Some of the Cyanobacteria found in these lakes
lik q Anabaena sp, Anabaenopsis V.V. Miller, 1923
and Oscillataria Vaucher ex Gomont, 1892, have
been reported to release cyanotoxins with lethal
effects on mammals (WHO, 1999; Lyra et al.,
2001). Although several research works on cyano-
toxins have focused on humans and other livestocks
(Leao et al., 2010), it is possible that these cyano-
toxins have similar toxic effects on aquatic organ-
isms including fish. Anabaena sp. have been
reported to release anatoxin-a which is a neurotoxin
reported to have lethal effects when tested on
Cyprinus carpio Linnaeus, 1758 larvae (Oswawald
et al., 2007). The presences of toxic Cyanobacteria
might be hindering fish occurrence in these lakes
considered under this study. With the exception of
Lake Kikorongo, which sometimes receives flood
waters from the neighboring lake Gorge (Hecky &
Kilham, 1973; Mungoma, 1990; Nkambo et al.,
2015), the rest of the lakes considered under this
study are located in closed basins with no connec-
tions to other lakes or rivers. This implies that these
lakes have no possibilities of being seeded with fish
by inflowing waters from other natural systems.
The reported occurrence of the African Catfish,
Clarias gariepinus, in Lake Kikorongo in the rainy
season might be due to flood waters from Lake
Gorge. In fact, Nkambo et al., (2015) reported
incoming flood waters from Lake Gorge to cause a
reduction in the salinity of Lake Kikorongo. A
reduction in salinity might make this lake condu-
cive for fish survival during the wet season. It is
possible that catfish brought along with floods
remain in this lake during the rainy season and die
off in the dry season as the water conditions become
extremely unbearable.
CONCLUSIONS
Our findings are in agreement with earlier
studies by Hecky & Kilham (1973), Matagi (2004),
Satyanarayan et al., (2007), Siddiqi (2007) and Yan-
nawar & Bhosle (2013) which reported Cyanobac-
teria to dominate algal biomass in saline systems,
in contrast with earlier studies by Stenger-Kovacs
et al., (2014), which reported diatoms to dominate
the algal biomass in saline lakes in Romania and
Bolivia. Lakes considered under this study had
very low zooplankton abundance and diversity with
Lake Kikorongo, which had the highest zooplank-
ton biodiversity, showing the rotifer Brachionus
calyciflorus, the most abundant, ranging only
between 50 to 100 individuals /litre. None of the
study lakes had fish at the time of sampling.
In our opinion, this information on fish and
planktonic biodiversity in these alkaline, saline
systems is very useful in providing the ecological
basis for the management of the lakes. Data on
dominant zoo and phytoplanktons in the lakes can
be used as bio-indicators in assessing the ecological
status, as well as the impact of climate change on
these unique systems.
Recommendations
Further comprehensive studies are needed to
assess the effect of season variability on fish and
planktonic biodiversity. Detailed studies of the daily
anthropogenic disturbances on the algal and
zooplankton biodiversity in these lakes during the
salt extraction process are required to give a better
understanding of the changes in planktonic compos-
ition, species abundance and biodiversity due to
anthropogenic disturbances.
ACKNOWLEDGEMENTS
Special thanks are reserved for the following
Planktonic and Fisheries biodiversity of Alkaline Saline crater lakes of Western Uganda
103
institutions of Uganda: National Agricultural Re-
search Organisation (NARO), National Fisheries
Resources Research Institute (NaFIRRI), Kajjansi
Aquaculture Research and Development Center
(KARDC), Department of Biological Science,
Makerere University and Katwe-Kabatoro Com-
munity, for the various supports offered will
carrying out this research.
REFERENCES
Ballot A., Kotut K., Wiegand C., Metcalf S. J., Codd
G. A. & Pflugmacher S., 2004. Cyanobacteria and
cyanobacterial toxins in three alkaline Rift Valley
lakes of Kenya-Lakes Bogoria, Nakura and Elmen-
teita. Journal of Plankton Research, 26: 925-935.
Ben Naceur H., Ben Rejeb Jenhani A. & Romdhane
M.S., 2012. Impacts of salinity, temperature and pH
on the morphology of Artemia (Branchiopoda:
Anostraca) from Tunisia. Zoological studies, 51:
453-462.
Boero F., Bouillon J., Gravili C., Miglietta M.P., Parsons
T. & Piraino S., 2008. Gelatinous plankton: irregu-
larities rule the world (sometimes). Marine Ecology
Progress series, 356: 299-310.
Brauner C.J., Gonzalez R.J. & Wilson J.M., 2013.
Extreme environments: hypersaline, alkaline and
ion-poor waters. In: McCormick S.D., Farrell A.,
Brauner CJ. (Eds.). Fish Physiology. Academic Press,
Boca Raton, Volume 32, pp. 435-476. ISBN:
9780123969514
Chessman B.C. & Williams W.D., 1974. Distribution of
fish in Inland saline waters in Victoria, Australia.
Australian Journal of Marine and Freshwater
Research, 25: 167-172.
Derry A.M., Prepas E.E. & Herbet P.D.N., 2003. A
comparision of zooplankton communities in saline
lakewater with variable anion composition. Hydro-
biologia, 505: 199-215.
Ghomari, S.M., Selselet G.S., Hontaria F. &Amat F.,
2011. Artemia biodiversity in Algerian Sebkhas.
Faboratoire de Technologie et Nutrition. Mostaganem,
Algeria, Universite de Mostaganem.
Grant W.D., 2006. Alkaline Environments and Biod-
iversity, in extremophiles. Encyclopedia of Fife
Support Systems (EOFSS). Charles G. & Nicolas G.
(Eds.), Oxford, UK, Eolss Publishers.
Green J., 1993. Zooplankton associations in East African
lakes spanning a wide salinity range. Developments
in Hydrobiology, 87: 249-256.
Hadgembes T., 2006. Environmental Threats to East
African Saline Lakes and their likely conservation
strategies. Addis Ababa, Ethiopia, academia.ed.
Hammer U.T. (Ed.), 1986. Saline Ecosystems of the
World, Dordrecht Publishers.
Hammer U.T., 1993. Zooplankton distribution and abund-
ance in saline lakes of Alberta and Saskatchewan,
Canada. International Journal of Salt Lake Research,
2: 111-132.
Harper M.D., Childress B.R., Harper M.M., Boar R.R.,
Hickley P., Mills S.C., Otieno N., Drane T., Vareschi
E., Nasirwa O., Mwatha W.E., Darlington J.P.E.C &
Escute-Gasulla X., 2003. Aquatic biodiversity and
saline lakes: Lake Bogoria National Reserve, Kenya.
Hydrobiologia, 500: 259-276.
Hecky R.E. & Kilham P., 1973. Diatoms in Alkaline,
Saline lakes: Eology and Geochemical Implications.
Limnology and Oceanography, 18: 53-71.
Jones E.B. & Grant W.D., 1999. Microbial diversity and
ecology of soda Lakes of East Africa. Microbial
Biosystems: New Frontiers. Proceedings of the 8th
international Symposium on Microbial Ecology. Bell
C.R., Brylinsky M. & Johnson-Green P. (Eds.),
Halifax, Canada, Atlantic Canada Society for Micro-
bial Ecology.
Joshi B.D., 2009. Biodiversity and Environmental
Management, APH Publishing.
Kirabira J.B., Kasedde H. & Semukuuttu D., 2013.
Towards The Improvement of Salt Extraction At
Lake Katwe. International Journal of Scientific &
Technology Research, 2: 76-81.
Larson A.C. & Belovsky E.G., 2013. Salinity and
nutrients influence species richness and evenness of
phytoplankton communities in microcosm experi-
ments from Great Salt Lake, Utah, USA. Journal
of Plankton Research, 35: 1154-1166.
Leao P.N., Pereira A.R., Liu W.-T., Ng J., Pevzner P.A.,
Dorrestein P.C., Konig G.M., Vasconcelos V.M. &
Gerwick W.H., 2010. Synergistic allelochemicals from
a freshwater cyanobacterium. Proceedings of the
National Academy of Sciences, 107: 11183-11188.
Lind E.M., 1965. The phytoplankton of some Kenya
Waters. Journal of the East African Natural History
Society, 25: 76-91.
Lubzens E., Tandler A. & Minkoff G., 1989. Rotifers as
food in aquaculture. Hydrobiologia, 186-187: 387-400.
Lyra C., Suomalainen S., Gugger M., Vezie C., Sundman
P., Paulin L. & Sivonen K., 2001 . Molecular charac-
terization of planktic cyanobacteria of Anabaena,
Aphanizomenon, Microcystis and Planktothrix gen-
era. International Journal of Systemic Evolutionary
Microbiology, 51: 513-526.
Matagi S.V., 2004. A biodiversity assessment of the
Flamingo Lakes of eastern Africa. Biodiversity, 5:
13-26.
Mungoma S., 1990. The alkaline, saline lakes of
Uganda: a review. Hydrobiologia, 208: 75-80.
104
Mujibu Nkambo etalii
Nixon H.P., Morton H.W. & von Knorring O., 1971.
Tychite and Northupite From Lake Katwe, Uganda.
Bulletin of the Geological Society of Finland, 43:
125-130.
Nkambo M., Bugenyi F.W., Naluwairo J., Nayiga S. &
Waswa L., 2015. Limnological Survey of the Al-
kaline, Saline crater lakes of Western Uganda.
Journal of Natural Science Research, 5: 53-61.
Oswawald J., Rellan S., Carvalho A.R, Gago A. &
Vasconcelos M., 2007. Acute effects of an anatoxin-
a producing cynobacterium on juvenile fish -Cyprinus
carpio L. Toxicon, 49: 693-698.
Rawson D.S. &Moore J.E., 1944. The Saline Lakes of
Saskatchewan. Canadian Journal of Research, 22:
141-201.
Rios-Escalante P.D.L., 2013. Crustacean zooplankton
species richness in Chilean lakes and ponds (23°-5 1°
S). Latin American Journal of Aquatic Research, 41:
600-605.
Saros J.E. & Fritz S.C., 2000. Nutrients as a link between
ionic concentration/composition and diatom distribu-
tions in saline lakes. Journal of Paleolimnology, 23:
449-453.
Satyanarayan S., Chaudhari PR. & Dhadse S., 2007.
Limnological study on Lonar lake: A Unique Black-
ish Crater Lakes in India. Proceedings of Taal 2007:
The 12th World Lake Conference: 2061-2066.
Schallenberg M., Hall C.J. & Burns C.W., 2003.
Consequences of climate-induced salinity increases
on zooplankton abundance and diversity in coastal
lakes. Marine ecology. Progress series, 251: 181—
189.
Siddiqi S.Z., 2007. Limnological Profile of High-Impact
Meteor Crater Lake Lonar, Buldana, Maharashtra,
India, an Extreme Hyperalkaline, Saline Habitat.
Proceedings of Taal 2007: The 12th World Lake
Conference: 1597-1613.
Stenger-Kovacs, C., Lengyel E., Buczko K., Toth F.M.,
Crossetti L.O., Pellinger A., Zsuzsa Doma Z. &
Padisak J., 2014. Vanishing world: alkaline, saline
lakes in Central Europe and their diatom assemblages.
Inland Waters, 4: 383-396.
WHO, 1999. Toxic Cyanobacteria in the Environment.
Toxic Cyanobacteria in Water: a guide to their public
health consequences, monitoring and management.
Chorus I. & Bartram J. (Eds.). E. & F.N. Spon,
London, ISBN 0-419-23930-8.
William W.D., 1998. Salinity as a determinant of the
structure of biological communities in salt lakes.
Hydrobiologia, 381: 191-201.
Yannawar B.V. & Bhosle B.A., 2013. Cultural Eutrophic-
ation of Lonar Lake, Maharashtra, India. Internatio-
nal Journal of Innovation and Applied Studies, 3:
504-510.
Biodiversity Journal, 2015, 6 (1): 105-106
New species of the genus Cyclostremiscus Pilsbry et Olsson,
1 945 from Central Philippines (GastropodaTornidae)
Ivan Perugia
via Roncalceci n.152, 48125 Ravenna (Filetto), Italy; e-mail; ivanperugia@virgilio.it
ABSTRACT Cyclostremiscus Pilsbry et Olsson, 1945 is a genus of the family Tornidae (Gastropoda
Rissooidea) established for very small shells of prosobranch molluscs generally living in
tropical seas. The new species here described was found in Cebu, Philippine, locatity Tongo
Point near Moalboal, in a modest quantity of seagrass beached after a windy day.
KEYWORDS Gastropod a; Tornidae; Cyclostremiscus-, Moalboal; Philippines.
Received 19.02.2015; accepted 21.03.2015; printed 30.03.2015
INTRODUCTION
Cyclostremiscus Pilsbry et Olsson, 1945 is a
genus of the family Tornidae established for very
small sells of prosobranch molluscs generally living
in tropical seas. The family Tornidae from Carib-
bean area has been extensively studied and illus-
trated by Rubio et al. (2011).
A new species of this genus from Philippine is
described in the present paper.
ACRONYMS. MHNUK: Natural History Mu-
seum of London, United Kingdom. MNHN: Mu-
seum National d'Histoire Naturelle Paris, France.
PC: I. Perugia collection, Ravenna, Italy.
SISTEMATICS
Superfamily TR U N C ATE LL O ID E A Gray, 1840
Family TORNIDAE Sacco, 1896
Subfamily V IT R IN ELL IN A E Bush, 1897
Genus Cyclostremiscus Pilsbry et Olsson, 1945
Type species: Vitrinella panamensis C.B. Adams,
1852
Cyclostremiscus albachiarae n. sp.
Examined material. Holotype, Cebu (Philip-
pine), locatity Tongo Point near Moalboal,
29. XI. 2007, I. Perugia legit, in a modest quantity
of seagrass beached after a windy day, MNHN -IM -
30079. Paratypes 1-5, same data of holotype,
M N HN -IM -3 00 80 . Paratypes 6-26, same data of
holotype (PC ).
Description of holotypus. Shell of small size
(Figs. l-4),diam eter 1.5 mm, height 1.0 mm, much
wider than high, discoid, rounded, relatively strong,
vitreous, colourless. Protoconch of 1.5 whorls,
rough, not elevated, max about 350 microns in dia-
meter. Teleoconch: spire with 2 whorls, umbilicus
open and deep, spiral and axial sculpture present on
the entire surface. Spiral sculpture formed by strong
prominence keels placed one on dorsum, one on
periphery, one on base and another delimiting the
umbilicus. Axial sculpture of numerous thin thick
arcuate riblets, surmounting the keels, itself inter-
sected by almost obsolete spiral lines. Aperture
rounded, outer lip with 3 prominences caused by
the end of spiral keels; anal sulcus well defined.
106
Ivan Perugia
Figures 1-4. CyclostremiscUS albachiarae n. sp., holotype from Moalboal, Cebu, Philippines.
Variability. The paratypes do not show sub-
stantial morphological differences compared to the
holotype. 36 specimens found have all the same
size, diameter 1.5 mm, height 1.0 mm.
Etymology. Dedicated to my granddaughter
Albachiara Perugia (Ravenna, Italy).
Remarks. For Cyclostremiscus albachiarae n. sp.
was possible a single comparison with Cyclostvema
gyalum Melvill, 1904 (MHMUK 1904.7.29.13)
which presents three similar keels on body-whorl
but is larger (about 5 mm in diameter), has a fine
spiral sculpture, not axial and the peripheral keel is
slightly waved at the margin (giving a slightly
stellate outline when viewed from above).
Besides the work of Rubio et al. (2011), other
contributions to the knowledge of the family
Tornidae were made by Melvill (1904), Adam &
Knudsen (1969), Bosch et al. (1995), and Bouchet
& Rocroi (2005).
REFERENCES
Adam W. & Knudsen J., 1969. Quelches genres de mol-
lusques prosobranches rnarins inconnu ou peu connus
de l’Afrique occidentale. Bulletin Institut Royal des
Sciences Naturelles de Belgique, 44: 1-69.
Bosch D.T., Dance S.P., Moolenbeek R.G. & Oliver
P.G., 1 995. Seashells of eastern Arabia. Motivate
Publishing Dubai, 296 pp.
Bouchet P. & Rocroi J.P., 2005. Classification and
nomenclator of gastropod fam dies. Malacologia, 47:
1-397.
Melvill J.C., 1904. Descriptions of Twenty-eight
Species of Gastropoda from the Persian Gulf, Gulf
of Oman, and Arabian Sea, Dredged by Mr. F.W.
Townsend, of the Indo-European Telegraph Sevice,
1900-1904. Proceedings of the M alacologic al
Society, 6: 158-1 69.
Rubio F., Fernandez-G arces R. & Rolan E., 2011. The
Family Tornidae (Gastropoda, Rissooidea) in the
Caribbean and Neighboring Aras. Iberus, 29: 1-230.
Biodiversity Journal, 2015, 6 (1): 107-114
New taxonomic data on some populations of Carabus
Macrothorax} morbillosus Fabricius, 1 792 (Coleoptera Cara-
bidae)
Ivan Rapuzzi 1 & Ignazio Sparacio 2
'via Cialla 47, 33040 Prepotto, Udine, Italy; email: info@ronchidicialla.it
2 via E. Notarbartolo 54, 90143 Palermo, Italy; email: isparacio@inwind.it
ABSTRACT In this work we give new taxonomic data on some, little known, populations of Carabus
( Macrothorax ) morbillosus Fabricius, 1792 (Coleoptera Carabidae). In particular, C. morbil-
losus lampedusae Bom, 1925 described from Lampedusa Island (Sicilian Channel, Italy) is
reconsidered a valid subspecies and are designated the lectotype and paralectotypes. Similarly,
Carabus morbillosus bruttianus Bom, 1906 described from Southern Calabria is considered
a distinct subspecies, including the populations of C. morbillosus from North-Eastern Sicily.
KEY WORDS Coleoptera; Carabus ; taxonomy; W-Mediterranean.
Received 11.02.2015; accepted 18.03.2015; printed 30.03.2015
INTRODUCTION
Carabus ( Macrothorax ) morbillosus Fabricius,
1792 (Coleoptera Carabidae) locus typicus
"Mauretania" is a widely W-Mediterranean distrib-
uted species (La Greca, 1964; 1984; Vigna Taglianti
et al., 1993; Parenzan, 1994) including several
populations, even insular, more or less fragmented
and differentiated, widespread in Southern France,
southern Spain, Morocco, Algeria, Tunisia, Corse,
Sardinia, Tyrrhenian central Italy, Southern Ca-
labria, Sicily, Sicilian islands, and Malta.
From the biogeographic point of view, as the
whole subgenus Macrothorax Desmarest, 1850, C.
morbillosus is considered a "Tyrrhenian" element
(Jannel, 1941 Antoine, 1955; La Greca, 1964; 1984;
Casale et al., 1982) with all connected hypotheses
and opinions on the origin and spread of the group.
Some populations also seem to be originated from
passive transport and, later, acclimatized (see Ca-
sale et al., 1989).
Currently, in Italy, are reported: C. morbillosus
morbillosus in Sardinia, Lampedusa and some
stations in Central Italy; C. morbillosus alternans
Palliardi, 1825 (locus typicus: Sicily) in Sicily,
Sicilian islands, and Calabria (Casale et al., 1982;
Vigna Taglianti, 1995; Vigna Taglianti et al., 2002).
In the "European Fauna" (Vigna Taglianti, 2015) the
populations of North Africa, Sardinia and Lampedusa
are attributed to C. morbillosus constantinus
Kraatz, 1899 (locus typicus: Constantine, Algeria).
Carabus morbillosus is an euriecious species
living in open areas or with sparse vegetation, often
in ruderal areas, urban gardens and crops, under
stones and debris, from the sea level up to about
1000 m a.s.l. Carabus morbillosus alternans is
found in forests and wooded fields, as oak and eu-
calyptus groves (Bosco Ficuzza, Piazza Armerina,
Bosco di Santo Pietro, etc.). It is present almost all year
long, mainly active from September to April-May.
In this work we provide new taxonomic data on
some little known populations of C. morbillosus.
108
Ivan Rapuzzi & Ignazio Sparacio
In particular, C. morbillosus lampedusae Bom,
1925 described from Lampedusa Island, is recon-
sidered as "bona subspecies" and its Lectotype and
Paralectotyes are designated; the same is also for C.
morbillosus bruttianus Born, 1906, described from
Southern Calabria to which we attribute also the
populations of C. morbillosus alternans from
North-Eastern Sicily.
ABBREVIATIONS AND ACRONYMS. ETZH:
Entomological Collection of ETH, Zurich, Switzer-
land; ex/s: exemplair/s; RC: Ivan Rapuzzi collec-
tion, Prepotto, Italy; SC: Ignazio Sparacio collec-
tion, Palermo, Italy.
SYSTEMATICS
Carabus ( Macrothorax ) morbillosus lampedusae
Bom, 1925
Examined material. Carabus morbillosus lam-
pedusae. ETZH: male (Fig. 1). Length: 30.10 mm;
width elytra: 10.85 mm; length elytra: 18.80 mm;
width pronotum: 7.80 mm; length pronotum: 6.20
mm. Three labels: "Insel Lampedusa" (handwritten
by Bom on circular label); "53.565" (print label),
"Carabus morbillosus lampedusae?" (handwritten
in blu color, more recent and different calligraphy
than original labels from Born); red label with
present designation of the “Lectotype”. Paralecto-
types, 2 males (ETZH), two labels each specimens:
"53.566" (print label); length: 30.50 mm; width
elytra: 11.70 mm; length elytra: 19.00 mm; width
pronotum: 7.60 mm; length pronotum: 6.20 mm.
"Carabus morbillosus lampedusae?" (handwritten in
blu color, more recent and different calligraphy
than original labels from Born). "53.567" (print
label); length: 30.30 mm; width elytra: 11.35 mm;
length elytra: 18.80 mm; width pronotum: 7.95
mm; length pronotum: 6.00 mm. "Carabus mor-
billosus lampedusae?" (hand- written in blu color,
more recent and different calligraphy than original
labels from Bom); red label with present designa-
tion of the “Paralectotype”.
Other examined material. Carabus morbillosus
lampedusae. ITALY, SICILY. Lampedusa Island
(Agrigento), 15.V.1983, I. Sparacio legit, 3 males
and 2 females (CS); idem, 10.11.2013, G. Mara-
ventano legit, 2 males and 2 females (CS); idem,
III.2014, T. La Mantia legit, 2 males and 4 females
(CS); Lampedusa (RC), 1 female; Lampedusa
Island, 4.II.1994, M. Romano legit, 2 males and 1
female (RC); Lampedusa Island, XI.2012, A. Corso
legit, 3 males and 1 female (RC).
Carabus morbillosus costantinus. TUNISIA.
Saouaf-El Fahs, 10. V. 1992, 1 female (CS);
Hammamet, 25.IV. 1998, 2 males (CS); El Fahs-
Zaghouan, 28. IV. 1998, 2 females (CS); Tabarka,
3/9.VI.1996, 10.V.1992, 5 males and 6 females
(CS); Tunisi, Cap Gammarth, 4.IV.2014, 5 males
and 4 females (CS), Tunisia, Bezeste, III. 1982, 1
male (RC); Tunisia, Ain Draham, 11/20. VI. 2008,
G. Sama & R Rapuzzi legit, 1 female (RC).
Remarks. Carabus lampedusae was originally
described by Bom (1925) as a subspecies of C.
morbillosus from Lampedusa Island (Sicilian
Channel, Italy) without designation of the holotype.
We had the opportunity to examine three male spe-
cimens preserved in the Entomological Collection
of ETH Zurich, ex Bom collection from Lampedusa
Island, and we have designated the lectotype and
paralecto types. All the three specimens are well
preserved.
From the systematic point of view, C. morbil-
losus lampedusae was considered mostly a variety
or a synonym of the nominate subspecies of North
Africa (Luigioni, 1929 sub morbillosus v. lampedu-
sae ; Breuning, 1932-36: sub natio costantinus ;
Porta, 1949 sub var. constantinus; Magistretti, 1965
sub C. morbillosus morbillosus natio costantinus ;
Casale et al., 1982 sub C. morbillosus morbillosus ;
Deuve, 2004 sub costantinus ).
Based on the examined material, we believe
"lampedusae" a valid subspecies of C. morbillosus ,
separate either from North African populations,
which is related to, or from the Sicilian ones, that
are more differentiated in morphology. In particular,
C. morbillosus lampedusae differs from the popu-
lations of North Africa by its squat and convex
body-shape, a darker and less bright color, with a
dominant chromatic variety characterized by dark
pronotum and dark green elytra in the middle, and
red on the sides; pronotum has wider and deeper
basal dimples with hind angles more sinuate
on the sides; the 1st elytral interstria shows
shallow points, well distinct and little confluent.
Hence, C. morbillosus lampedusae would be
comprised within the group of autochthonous
species of Lampedusa, of apparent North African
origin, morphologically differentiated in insularity
New taxonomic data on Carabus (Macrothorax) morbillosus Fabricius, 1792 (Coleoptera Carabidae)
109
/a Ml
Tk&JU F
■^rz iJutC-OAXllA Q&U)
Figure 1. Carabus ( Macrothorax ) morbillosus lampedusae Bom, 1925, lectotype with original labels.
Figure 2. Carabus (M) morbillosus bruttianus Bom, 1906 lectotype with original labels.
conditions. In the Conclusion section (see below),
we report morphological characters that distinguish
these populations to each other and those charac-
terizing the populations under study in this work.
Carabus {Macrothorax) morbillosus bruttianus
Bom, 1906
Examined material. ITALY, CALABRIA.
Carabus morbillosus bruttianus. Lectotype male,
Sta Eufemia d’Aspromonte, Calabrien (ETHZ)
(Fig. 2); idem, Paralectotype female (ETHZ).
Other examined material. Calabria. Reggio
Calabria dintomi, I. Sparacio legit, 8.XI.1999, 5
males (RC); Reggio Calabria dintomi, 5.III.2004,
I. Sparacio legit, 2 males and 1 female (RC);
Reggio Calabria dintomi, 8.XI.1999, 6 males and 5
females (SC); Reggio Calabria: Campo Calabro,
8.XI.1999, 6 males and 2 females (SC); Torrente
Zagarella, 8.XI.1999, 8 males (Fig. 5) and 11 fe-
males (SC); Gioia Tauro, 9.XI.1999, 13 males and
10 females (SC). ITALY, SICILY. Messina dintomi,
4.XI.2001, 13 males (Fig. 4) and 9 females (SC);
Messina, Monte Ciccia, 4. III. 2004, 7 males and
7 females (SC); Messina: Colle San Rizzo,
4.XI.2001, 3 males and 1 female (SC); Messina:
Faro, 4. III. 2004, 2 males and 2 females (SC); 3
males (SC); Messina, Torrenova, VIII.2013, A.
Tetamo legit, 1 male (SC); Messina dintomi,
4.XI.2001, I. Sparacio legit, 1 male and 1 fe-
male (RC); Messina, Monte Ciccia, 4.III.2004, I.
Sparacio legit, 2 males (RC); Lipari, Isole Eolie,
V.2014, R Lo Cascio legit, 3 males and 1 female (SC).
Carabus morbillosus alternans. ITALY, SICILY.
Palermo . Carini, 9.X.1978, 3 males and 4 females;
idem, 1.V.1979, 1 male (SC); Palermo, 18.X.1978,
1 male and 1 female; idem, 7.II.1979, 1 male; idem,
30.III.1980, 1 female (SC); Palermo: Sferracavallo,
Grotta Conza, 3.XI.1978, 2 females (SC); Godrano,
25.XI.1978, 1 male and 2 females (SC); Palermo:
Sferracavallo, 14.1.1979, 2 males; idem, 13.V.1980,
1 female (SC); Piana degli Albanesi, 8.II.1979, 3
males and 3 females; idem, 6.II.1992, 1 male and 3
females; idem, 15.IV. 1995, 1 female (SC); Palermo:
Favorita, Vallone del Porco, 3.III.1979, 1 male and
2 females (SC); Bosco Ficuzza, 21. XI. 1979, 1 male
(Fig. 3) and 1 female; idem, 9.II.1987, 1 female;
110
Ivan Rapuzzi & Ignazio Sparacio
idem, 28.1.1989, 1 female (SC); Altofonte,
1. III. 1981, 1 male and 1 female (SC); Altofonte:
Poggio San Francesco, 13. III. 1981, 1 male (SC);
Cefalu, 7.XI.1987, 1 female (SC); Palermo:
Mondello, 18.IX.1988, 1 male (SC); Capaci,
25.11.1989, 1 female (SC); Monreale: Giacalone,
12.1.1992, 3 males and 1 female (SC); Ficuzza:
Bivio Lupo, 25.11.1992, 1 male; idem, 13.XI.2001,
2 males (SC); Santuario di Gibilmanna, 23. X. 1994,
1 male (SC); Lercara Friddi: S. Caterina A.D.,
24.X.2004, 2 males (SC); Bagheria: Monte Catal-
fano, 14.X.2006, 1 female (SC); Roccamena: Maran-
fusa, 25.IV.2008, 1 female (SC); Prizzi, 12.VI.2009,
1 male and 1 female (SC); Palermo: Micciulla,
4.IV.2010, T. La Mantia legit, 1 male (SC); Rocca
Entella, 18.XI.2011, 3 males (SC); Ficuzza: Gorgo
del Drago, 25.XI.2012, 2 females (SC); Diga Poma,
10.XI.20 13, 1 male and 1 female (SC); Trabia: Pizzo
Cane, XI.2014, 2 males and 1 female (SC); Cefalu,
Settefrati, VI. 1984, 1 male (RC); Ficuzza, Godrano,
XI/XII.2010, I. Rapuzzi & L. Caldon legit, 70 exs
males and females (RC); Palermo, 13.III.1992, 1 fe-
male (RC); Gibilmanna, 500 m, VII. 1984, 1 exs
(resti) (RC); Piana d. Albanesi, 700/800 m, III. 1988,
1 male and 1 female (RC); Isnello, 700 m, III. 1988,
2 males (RC). Trapani . Erice, 12.XI.1972, M. Romano
legit, 1 male and 1 female (RC); Campobello di Maz-
ara, Cave di Cusa, 28.XI.2009, 3 males and 3 females
(RC); Mazara costiera, 13.1.1985, 1 male and 2 fe-
males (RC); Capo Granitola, 30.1.1986, V. Castelli
legit, 2 males and 3 females (RC); Mazara,
3.II.1985, V. Aliquo’ legit, 1 male and 1 female
(RC); Selinunte, 24.XI.2002, I. Rapuzzi & L. Cal-
don legit, 2 males and 6 females (RC); Segesta,
15.11.201 1, 1. Rapuzzi & L. Caldon legit, 4 males and
2 females (RC); Santa Ninfa, XI.2009, 1. Rapuzzi &
L. Caldon legit, 1 male and 1 female (RC); Mazara
del Vallo, 28.XI.1979, 2 males and 3 females (SC);
San Vito Lo Capo (Trapani), 14.X.1984, 3 males
(SC); Monte Cofano, 14.X.1984, 1 female (SC); Cas-
tellammare del Golfo, 12.XII.1984, 2 males and 1
female (SC); Cave di Cusa, 14.XII.2003, 2 males;
idem, 3 1 .XII. 1 988, 3 females (SC); Foci Fiume Belice,
20.IV. 1989, 1 female (SC); Foci Fiume Birgi,
6.XI.1993, 1 female (SC); Selinunte, 3.XII.1995, 1
male and 1 female (SC); Valderice, 1.2014, 4 males
Figure 3. Carabus ( Macrothorax ) morbillosus alternans male, Bosco Ficuzza, Palermo, Sicily. Figure 4. Carabus morbillosus
bruttianus male, Messina surroundings, Sicily. Figure 5. Carabus morbillosus bruttianus male, Torrente Zagarella, Reggio
Calabria, Calabria (Photos by M. Romano).
New taxonomic data on Carabus (Macrothorax) morbillosus Fabricius, 1792 (Coleoptera Carabidae)
111
Figure 6. Scatterplot of the relationship between length and
width of elytra (upper) and pronotum (bottom) for three po-
pulations examined. Are shown the regression lines with the
associated confidence intervals (95%). Values of Correlation
coefficents for width/length of elytra in the three populations
are: Calabria 0.68*, Messina 0.25, Sicilia 0.78*. Those for
width/length of pronotum are: Calabria 0.15, Messina 0.42*,
Sicilia 0.46*. (* P< 0.05).
Pronotum Standardized
coefficients
Pooled- within-groups
correlations
Root 1
Root 2
Root 1
Root 2
Elytra Lenght -0.782
1.007
-0.971
0.239
Pronotum Width -0.305
-1.238
-0.790
-0.613
Eigenval 1.346
0.038
Cum. Prop 0.973
1
Table 1. Standardized coefficients (left) e Pooled-within-
groups correlations (right) for the two variables selected by
correspondence analysis.
Means of Canonical
Variables
Pop. Root 1 Root 2
Calabria 1.480 0.256
Messina 0.570 -0.235
Sicily -1.283 0.067
Table 2. Means of Canonical Variables for the three
examined populations.
Cases
Percent correct
Calabria
Messina
Sicily
Calabria
56.3
9
7
0
Messina
73.1
4
19
3
Sicily
76.7
0
7
23
Total
70.8
13
33
26
Table 3. Classification Matrix. The first column shows the per-
centages of observations properly attribuited to each popula-
tion using discr im inant analysis. The remaining columns show
the number of cases falling into each population (diagonally,
cases correctly classified).
and 3 females (SC). Agrigento . Agrigento,
21.1.1973, 1 male (RC); Agrigento: Valle dei Templi,
1.1972, 1 male and 1 female (RC); Agrigento: Valle
dei Templi, 9.II.1987, 1 female; idem, 2.1.1989, 1
male and 1 female (SC). Caltanissetta . Monte Ca-
podarso: F. Imera meridionale, 5.VI.2006, 1 male
(SC); Ponte Cinque Archi, 14.11.2015, 2 males (SC).
Enna . Valguamera (Enna), 25.IX.1979, 1 male (SC);
Piazza Armerina: Monte Rossomanno, 10. III. 2008,
1 female (SC); Piazza Armerina, XI.2009, 1. Rapuzzi
& L. Caldon legit, 1 female (RC). Syracuse . Vendi-
cari, 18.VIII.1993, 2 males and 1 female (SC);
Priolo, 28.XI.2010, 3 males; idem, 5 .111.2011, 1 fe-
112
Ivan Rapuzzi & Ignazio Sparacio
10 . 5 -
10.2
9 . 9 -
9.6
I =:■
GO
w
7.75
7.50
7 25 -
7.00
6.75
Elytra Width (EW)
i
»
i
i
>
Calabria Messina Sicily
18 . 0 -
17 . 5 -
17 . 0 -
16 . 5 -
16 . 0 -
Elytra Length (EL)
*
c
>
i
.b
<
a J
>
Calabria Messina Sicily
Pronotum Width (PW)
r
\_r
>
i
b
a
*
>
6.0
5.8 -
6 6 -
Pronotum Length (PL)
*
*
*
*
>
Calabria Messina Sicily
Calabria Messina Sicily
Figure 7. Comparison of means, and confidence intervals (95%) among the three popultions for the four variables examined.
For the variables selected by discriminant analysis (PW and EL) are shown the values of significance in multiple compar-
isons. PW: Messina/Calabria t = 3.211**; Sicilia/Calabria t = 7.426***; Sicilia/Messina t = 4.772***. EL: Messina/Calabria
t = 2.412*; Sicilia/Calabria t = 8.520***; Sicilia/Messina t = 6.983***.
male (SC); Magnisi, 28.XI.2010, 2 males and 1 fe-
male (SC); Palazzolo Acreide, 5 .111.20 1 1 (SC);
Vizzini, 14.11.2015, 1 male (SC). Messina . Nebrodi
Mts., North from Capizzi, 1250 m, 23 .VII. 1991, 1
male (RC).
Discriminant analysis. We also made a biome-
tric study on 30 male specimens of C. morbillosus
alternans from Sicily, with the exception of the north-
easternmost regions, 16 male specimens of C. mor-
billosus bruttianus from Southern Calabria and 26
males attributed to C. morbillosus bruttianus from
Messina surroundings. The following measures
were examined: pronotum width (PW), pronotum
length (PL), elytra width (EW), and elytra length
(EL). In figure 6 are shown the graphs of the rela-
tionships between length and width of elytra and
pronotum for the three populations. Our findings
showed significant results when comparing C. mor-
billosus alternans and C. morbillosus bruttianus
(Calabria and Messina); whereas slight differences
were observed between the two populations of C.
morbillosus bruttianus from Calabria and Messina.
In order to identify which one of the four
morphometric characters used allows to distinguish
the three populations of C. morbillosus it was used
the discriminant functions analysis. Variable selec-
tion was done by the "Forward stepwise". The res-
New taxonomic data on Carabus (Macrothorax) morbillosus Fabricius, 1792 (Coleoptera Carabidae)
113
ulting model shows a discriminating value not high
but still significant (Wilks' Lambda: 0.4107060
approx. F (4.136) = 19.05342; p <, 0000). Variables
selected from the analysis are: EL and PW. Partial
lambda values (0.761599 and 0.932681, respect-
ively) indicate that EL followed by PW have the
most discriminating power among the three popu-
lations examined. The analysis produced two linear
functions, Rootl and Root2, the first appears negat-
ively correlated mainly with “elytra length” and, to
a lesser extent, with “pronotum width” (Table 1)
and discriminates the population of Calabria from
that of Sicily (Table 2). The second is negatively
correlated with pronotum width (Table 1) and,
although possess a low discriminatory power,
partially contributes to distinguish the population
of Calabria from that of Messina (Table 2).
In addition, the Mahalanobis distance between
the centromeres of the three populations, although
it is significant for all comparisons, shows high
values only between Calabria and Sicily (Mess-
ina/Calabria 1,116 *; Sicily/Calabria 8,004 ***; Si-
cily/Messina 3,673 *** . (P: *** < 0.001; ** < 0.01;
* < 0.05). The classification matrix (Table 3) shows
that more than 70% of the specimens from Messina
and the rest of Sicily are properly classified, while
this percentage drops to around 56% for specimens
from Calabria.
To assess whether the averages of each of the
two variables identified with the discriminant ana-
lysis significantly differ among the three popula-
tions, it has been carried out the analysis of variance
(Fig. 7). Multiple comparisons were performed with
the correction of Turkey. For both characters
ANOVA was significant (pronotum width: Df =
2/62, F = 23.61, P <le-04 ***. Elytra Length Df =
2/62, F = 37.14, P <le-04 * **). Multiple compar-
isons between populations are highly significant
except for the comparison Messina/Calabria for EL
that is barely significant (p = 0.0477).
Remarks. Born (1906) describes Carabus mor-
billosus bruttianus from Calabria, locus typicus St.
Eufemia d'Aspromonte, distinguishing it from the
Sicilian populations of Palermo known as C. mor-
billosus servillei (= C. morbillosus alternans). Sub-
sequently both Porta (1923) and Luigioni (1929)
report it as a distinct "variety" of C. morbillosus
from Calabria. In particular, Porta (1923) reiterates
the morphological differences already reported by
Born (1906) in its original description. However,
latest Authors consider C. morbillosus bruttianus as
a synonym of C. morbillosus alternans of Sicily
(Magistretti, 1965; Casale et al., 1982; Vigna
Taglianti, 1995; Vigna Taglianti, et al., 2002).
The examination of numerous specimens from
different places near Reggio Calabria (Southern Ca-
labria), allowed us to confirm the morphological
characteristics of this taxon, which results morpho-
logically distinct from the neighboring populations
of C. morbillosus alternans of Sicily. Populations
attributable to C. morbillosus bruttianus are also
present near Messina (north-eastern Sicily), de-
scribed as C. borni Krausse, 1908 (= sicanus Csiki,
1927; nom. pro borni Krausse). Porta (1923) reports
this taxon as a distinct "variety" of North-Eastern
Sicily, thus distinguishing it, geographically, from
the remaining populations of South-Western Sicily.
The populations of Messina are, in fact, mor-
phologically distinct from the remaining Sicilian
ones attributed to C. morbillosus alternans and,
rather, similar to the Calabrian populations of C.
morbillosus bruttianus from which differ only in a
few minor characters, especially by color and shape
of pronotum.
CONCLUSIONS
Actually, the C. morbillosus population of
Sicily, Sicilian islands, and the nearby Southern Ca-
labria, turns out to be more diversified than con-
sidered up to now. In most of the islands, is con-
firmed the presence of C. morbillosus alternans
which is very well-differentiated and distinct from
all the other races of the species; C. morbillosus
bruttianus is present in Southern Calabria, in the
territories of North-Eastern Sicily (Messina and
surrounding area) and, as to our knowledge, even
in Lipari in the Aeolian Islands. In Lampedusa
Island there is an island subspecies, C. morbillosus
lampedusae , similar to North African populations
of C. morbillosus .
At the moment, the populations covered by this
work can be distinguished as outlined below:
1 . Pronotum wider and arched forward with ma-
ximum width in the fore third. Primary intervals sa-
lient and very short, secondary ribs raised and wide,
tertiary intervals broken down into lines of evident
granules, 1st elytral interstria with small tubercles
114
Ivan Rapuzzi & Ignazio Sparacio
and confluent points in the form of irregular furrow.
Aedeagus apex distinctly more elongated, narrow
and slightly curved morbillosus costantinus
-. Squat and convex body-shape, less bright in
color and dark. Pronotum with basal dimples large
and deep, sinuate at sides before hind angles.
Primary intervals wider, 1st elytral interstria with
points on the surface, well separate from each
other morbillosus lampedusae
2. Pronotum distinctly narrower forward with
maximum width at the center. Primary intervals
elongated and slightly salient, secondary ribs de-
pressed, tertiary intervals less raised than secondary
ones; 1st elytral interstria with wide points, deep,
very distinct, sometimes juxtaposed with each
other. Apex of aedeagus shorter, wider and curved.
Shape great and flattened on the back, brilliant;
elytra elongate, rounded and dilated in the rear
third; elytra apex short and sightly sinuate at sides...
morbillosus alternans
-. Smaller and convex on the back of the elytra,
less shine; pronotum narrower and slightly rounded
forward with maximum width in the fore half; disc
with evident points and transverse wrinkles thin and
sparse; elytra short and ovalish, primary intervals
in granules shorter and less raised; elytral apex
stretched and clearly sinuate at sides
morbillosus bruttianus
ACKNOWLEDGEMENTS
We thank Dr. Andreas Muller (ETHZ) and Dr.
Franziska Schmid (ETHZ) for the loan of the type
specimens from Born Collection of Carabus
{Macrothorax) morbillosus lampedusae and C. (M.)
morbillosus bruttianus', Dr. Antonio Adorno
(University of Catania, Italy), Dr. Andrea Corso
(Syracuse, Italy), and Dr. Marcello Romano
(Capaci, Italy).
REFERENCES
Antoine M., 1955. Coleopteres Carabiques du Maroc, I.
Memoire de la Societe des Sciences Naturelles du
Maroc, 1: 1-177.
Born P., 1906. Ueber einige Carabus. Formen aus
Calabrien. Insekten-Borse, 23: 1-6.
Born P., 1925. Carabus morbillosus lampedusae nov.
subspec. Societas entomologica, 7: 25-26.
Breuning S., 1932-1936. Monographic der Gattung
Carabus L. Best. - Tab. Europ. Coleopt., 104-110,
Reitter, Troppau, 1610 pp., 41 tav.
Casale A., Sturani M. & Vigna Taglianti A., 1982. Cole-
optera Carabidae. I. Introduzione, Paussinae, Cara-
binae. Fauna d ’Italia, XVIII. Edizioni Calderini,
Bologna, 500 pp.
Casale A., Bastianini M. & Minnniti M., 1989. Sulla
presenza in Toscana di Carabus ( Macrothorax ) mor-
billosus Fabricius (Coleoptera, Carabidae, Carabini)
e sul suo significato zoogeografico). Frustula
entomologica, 10 (1987): 61-12.
Deuve T., 2004. Illustrated Catalogue of the Genus Ca-
rabus of the World (Coleoptera, Carabidae). Pensoft
Publishers, Sofia-Moscow, 461 pp.
Jannel, 1941. Coleopteres Carabiques (Premiere partie).
Faune de France, 39. Lechevalier, Paris, 571 pp.
La GrecaM., 1964. Le categorie corologiche degli ele-
menti faunistici italiani. Atti Accademia Nazionale
Italiana di Entomologia, Rendiconti, 11: 231-253.
La Greca M., 1984. L’origine della fauna italiana. Le
Scienze, 187: 66-79.
Luigioni P., 1929. I Coleotteri d’ltalia. Catalogo sinon-
imico-topografico-bibliografico. Memorie della
Pontificia. Accademia di scienze ” I Nuovi Lincei”,
Roma, 2, 13: 1-1160.
Magistretti M., 1965. Coleoptera Cicindelidae, Carabi-
dae. Fauna d’ltalia, VIII. Edizioni Calderini,
Bologna, 512 pp.
Parenzan P., 1994. Proposta di codificazione per una
gestione inormatica dei corotipi W-paleartici con
particolare riferimento alia fauna italiana. Entomolo-
gica, 28: 93-98.
Porta A., 1923. Fauna Coleopterorum Italica. lAdephaga.
StabilimentoTipografico Piacentino, Piacenza, 286 pp.
Porta A., 1949. Fauna Coleopterorum Italica. Supple-
mentum 2. Stabilimento Tipografico Soc. an.
Giacomo Gandolfi, Sanremo, 388 pp.
Vigna Taglianti A., 1995. Coleoptera Archostemata,
Adephaga 1 (Carabidae). In: Minelli A., Ruffo S. &
La Posta S. (Eds.), Checklist delle specie della fauna
italiana. 44. Calderini. Bologna.
Vigna Taglianti A., 2015. Fauna Europaea: Carabus
( Macrothorax ) morbillosus Fabricius 1792. In:
Audisio, R, 2015 Fauna Europaea: Carabidae Fauna
Europaea version 2.6.2, Fauna Europaea: Name Search
Vigna Taglianti A., Casale A. & Fattorini S., 2002. I
Carabidi di Sicilia ed il loro significato biogeografico
(Coleoptera, Carabidae). Bollettino dell' Accademia
Gioenia di scienze naturali, 35 (361): 435M64.
Vigna Taglianti A., Audisio P.A., Belfiore C., Biondi M.,
Bologna M.A., Carpaneto G.M., De Biase A., De
Felici S., Piattella E., Racheli T., Zapparoli M. & Zoia
S., 1993. Riflessioni sui corotipi fondamentali della
fauna W-paleartica ed in particolare italiana. Biogeo-
graphia, 16: 159-179.
Biodiversity Journal, 2015, 6 (1): 115-117
About the presence of the Haifa Grouper Hyporthodus haifen-
sis (Ben-Tuvia, 1 953) (Perciformes Serranidae) in the Strait of
Messina, Italy, Mediterranean Sea
Rosi Barbagallo 1 , Francesco Turano 2 & Riccardo Delle Fratte 3 *
'Via A lcantara 11, 95045 Misterbianco, Catania, Italy; e-mail: lailar@live.it
2 ViaAliquo Taverriti 28, 89131 Reggio Calabria, Italy; e-mail; cicciotura no@gmail.com
3 Via dell’Imbrecciato 95, 00149 Rome, Italy; e-mail: riccardodellefratte@gmail.com
Corresponding author
ABSTRACT In this paper is reported for the first time the presence of the Haifa Grouper, HyporthodllS
haifensis (B en-Tuvia, 1953) (Perciformes Serranidae) in the w aters of the Strait of M essina,
Italy which confirms the expansion process of the species toward the northern part of the
Mediterranean Sea.
KEYWORDS Epinephelinae; HyporthodllS haifensis', Lampedusa; Pellaro; Serranidae.
Received 26.12.2014; accepted 09.02.2015; printed 30.03.2015
INTRODUCTION
The subfamily Epinephelinae belonging to the
family of marine bony fish Serranidae includes
many genera, among which Hyporthodus Gill,
1861. The species belonging to this genus are com-
monly called "groupers" as some of their close re-
latives of the genus EphinephelliS Bloch, 1793.
Haifa Grouper, Hyporthodus haifensis (Ben-
Tuvia, 1953) is a marine fish, demersal and gen-
erally present in water depths between 90 and 220
meters (Froese & Pauly, 2014; Heemstra & Randall,
1993). The body has large pelvic and pectoral fins,
often bordered with white, tail shape rather roun-
ded, body color dark brown. Its distribution area
appears to be the Eastern Atlantic from the coast of
Angola in the south, to those in the southern part of
Portugal.
Hyporthodus haifensis is also present in the
southern part of the Mediterranean Sea (where the
species arrived entering through Gilbraltar) from
the coast to the south of Spain, along almost all the
Mediterranean coast of North Africa to Lebanon,
Israel, Turkey and southern Greece (Heemstra &
Randall, 1993). The only indication of its presence
in Italian waters was published in 2000 for the
Lampedusa Island, Pelagie Archipelago, Sicily
Channel (Azzurro et al., 2000). Considering the
characteristics of Lampedusa and its geographical
location, we can say that the island represents
a bridge between the Italian and North African
territory. This led to assume, even at the tim e of the
report of H. haifensis in Lampedusa, a future
movement of the species in the direction of the
waters of Sicily and the Italian mainland (Heemstra
& Randall, 1993).
MATERIAL AND METHODS
The sighting occurred during a dives with scuba
equipment. The camera equipment consisted of
Canon G-15, Fantasea housing and Sea & Sea
flashes.
116
Rosi Barbagallo et alii
Figures 1-3. Young specimen of Epinephelus haifensis (Pellaro, Reggio Calabria, Italy), 35 m deep, XI. 2014,
photographed by day (Figs. 1, 2) and during the night (Fig. 3).
RESULTS AND CONCLUSIONS
The discovery of this species has occurred in
Pellaro (Reggio Calabria, Italy) in the month of
November 2014 on a backdrop of mixed sand and
mud, 35 meters deep and with a water temperature
of about 23 °C. The animal was a young specimen
(Fig. 1) of about 25 cm in overall length, stationing
around a small wreck of about 3 meters in length.
The identification was made possible by count-
ing the number of soft rays in the anal fin, which is
not less than nine; another distinctive character of
the species (but only valid for young specimens) is
that the pelvic fins are not long enough to reach the
anus. No wonder for the low depth at which the
specimen was found infact, despite belonging to a
species which prefers stationing at depths beyond
90 meters, it is well known the habit of young grou-
pers to colonize shallow waters, within 30 meters.
Also in the Strait of Messina, thanks to its weather
and sea conditions or currents, peculiar at all, many
species, such as the Longspine snipefish MaCWratfl-
phosUS scolopax (Linnaeus, 1758) (Syngnathi-
formes C entriscidae), generally station at depths
About the presence of the Haifa Grouper Hyporthodus haifensis in the Strait of Messina, Italy, Mediterranean Sea 1 1 7
much lower than those where they are usually found
in other areas of the Mediterranean Sea.
The specimen was found in the same spot in
three separate dives performed during the same
month of November and then photographed by day
(Fig. 2) and during the night (Fig. 3). The latter
photo was taken in a subsequent night dive, during
which it was found the same specimen in the same
place. The image depicts the specimen during sleep
and is interesting because, as is well known, fish
change their colors at night using special cells
called chrom atophores. This picture shows exactly
the colors taken at night by Haifa’s grouper, which
is rarely documented.
This new record of H. haifensis is in continuity
with the previous one recorded in Lampedusa,
confirming the expansion process of the species
toward the northern part of the Mediterranean Sea.
This expansion was predicted by some authors
(Heemstra & Randall, 1993) and will therefore be in-
teresting to continue to monitor future developments.
It is also very likely that often the presence of
specimens of this species may not be noticed, as
they may be easily confused with specimens of
other M editerranean species.
REFERENCES
Azzurro E., Andaloro F. & Marino G., 2000. Presenza
della cernia di Haifa, Epiliephelus haifensis (Serran-
idae: Epinephelinae), nel Mediterraneo centrale.
Biologia marina m editerranea, 7: 786-789.
Froese R. & Pauly D., 2014. Fish Base. W orld W ide Web
electronic publication, www.fishbase.org, (08/2014).
http://www.fishbase.org/summary/Epinephelus-
haifensis.html (last accessed: 20.12.2014).
Heemstra P.C . & Randall J.E., 1993. FAO Species
Catalogue. Vol. 16. Groupers of the world (family
Serranidae, subfamily Epinephelinae). A n annotated
and illustrated catalogue of the grouper, rockcod,
hind, coral grouper and lyretail species known to date.
FAO Fisheries Synopsis, Rome, 125 (16), 382 pp.
Biodiversity Journal, 2015, 6 (1): 119-120
Monograph
Preface
Speciation and Taxonomy:
Neotropical Primate diversity
Marc Van Roosmalen 1 & Mason Fischer 2
Tstr. do CETUR, 2445 Tarama Manaus- AM CEP 69.022-155, Brazil; email: marc.mvrs@gmail.com
2 Mason Fisher Photograhy; email: mason@masonfischerphotography.com
Received 14.01.2015; accepted 25.01.2015; printed 30.03.2015
Proceedings of the 2nd International Congress “Speciation and Taxonomy”, May 1 6th- 1 8th 2014, Cefalu-Castelbuono (Italy)
During the last three decades Dr. Van Roos-
malen has surveyed by boat, canoe, and on foot
entire basins of some major tributaries of the
mighty Amazon River in order to study primate
diversity and distributions across the entire Amazon
Basin (including large parts of the Precambrian
Guiana and Brazilian Shields).
This way he tested and empirically came to
fully validate Alfred Russel Wallace's river-barrier
hypothesis first laid down in his 1852 account On
the Monkeys of the Amazon. Wallace points at the
larger rivers he sailed as the principal evolutionary
cause of the Amazon's uniquely rich extant primate
diversity and complex biogeography, for many
rivers together with their floodplains effectively
block off gene flow between populations along
opposite riverbanks.
As the Amazon represents a largely pristine and
vast natural realm not (yet) modified by human
interference, no better place to retrace evolutionary
processes that may have acted upon primates
(including our own ancestors) and other mammals
since the Pliocene. Moreover, Van Roosmalen's
biodiversity surveys revealed a number of new
monkeys from all over the Amazon (described
elsewhere) and other megafauna (somefrom the
Rio Aripuana Basin described here), among which
even a new genus - the dwarf marmoset Callibella
humilis M. van Roosmalen, T. van Roosmalen,
Mittermeier et de Fonseca, 1998.
This peaceable, non-territorial, enigmatic,
second smallest monkey in the world (here depicted
in the upper left corner) occupying the smallest
distribution of any monkey on the planet stands at
the base of the phylogenetic tree of all extant
marmosets. Interestingly, the whole family of
advanced Callitrichidae (i.e., the genera Cebuella
Gray, 1866 Callithrix Erxleben, 1758, Mico
Thomas, 1920, Saguinus Hoffmannsegg, 1807,
Leontopithecus Lesson, 1840) exhibits social group-
ings that fiercely defend a common living space or
territory (Fig. 1).
Speciation, radiation and rate of metachromic
bleaching among primatesseem to be related to ter-
ritoriality and social rather than sexual selection (as
is the case in other mammals and birds). Con-
sequently, the strictly territorial Amazonian mar-
mosets (Mico), tamarins (Saguinus), lion tamarins
(Leontopithecus), ouistitis (Callithrix), titis (Cal-
licebus Thomas, 1903) and howling monkeys
(Alouatta Lacepede, 1799) are the most diversified,
species-rich and colorful genera among a total of
19 New- World monkeys, in striking contrast to
Goeldi's Monkey Callimico and dwarf marmoset
120
Marc Van Roosmalen& Mason Fischer
Figure 1. Neotropical Primate diversity - Amazon Basin, Brazil.
Callibella - the only two Neotropical primate coat coloration (eumelanin black, brown and/or
genera being monotypic and archetypic in skin and agouti).
Biodiversity Journal, 2015, 6 (1): 121-122
Monograph
Introduction
Speciation and Taxonomy: digressions at the
edge of a meeting
Pietro AN cata
Dipartimento di Scienze Biologiche, Geologiche e Ambientali, sez. Biologia Animale, Universita di Catania, via Androne 81,
95124 Catania, Italy
Received 18.02.2015; accepted 02.03.2015; printed 30.03.2015
Proceedings of the 2nd International Congress “Speciation and Taxonomy”, May 1 6th- 1 8th 2014, Cefalu-Castelbuono (Italy)
The title of the meeting organized by the Biod-
iversity Journal leads itself to reflections upon the
cognitive relation between man and nature. Two
different ways of looking at biodiversity are
approached: explanation of its origin and classific-
ation of its diversity. The first looks into the pro-
cesses that have led to the formation of that
extraordinary, wonderful, tragic and cruel world
that we call life. The result has been the impressive
system of knowledge of the biological evolution.
We are confident, on the other hand, that the pro-
cesses of life are independent of any our interpreta-
tion. The second meets our need to name and
describe the living beings appearing as separate
discrete entities. The outcome of this activity is
taxonomy, an object of our mind, whose first sys-
tematic form dates back to the Systema Naturae,
elaborated by Linnaeus far before the emergence
of the evolutionary theory, nearly actualizing the
first job of Adam (Genesis 2, 19).
Speciation is a crucial event of evolution: ge-
netic variations and adaptations to different envir-
onmental contexts have produced a multiplicity of
species and a great diversity of living organisms.
Taxonomy is able to represent only a time confined
image of the result of this process; however its sys-
tem becomes a new subject of our knowledge with
implications on our perception of the diversity of
life. May rules and methods of taxonomy affect the
comprehension of the life evolution? Indeed, we
can suppose that, in the interaction between the
static description of biodiversity and the analysis of
its development, the mechanism of our mind plays
a relevant role in guiding the thoughts towards the
established knowledge.
Phylogenetic analys is try to connect taxonomy
to speciation and contributes to its redefinition.
However both phylogeny and taxonomy respect a
tacit postulate whose rational foundation is not
considered problematic: similarity among beings
indicates a common origin and the chain of repro-
ductive events brings us to the common life origin.
And, if life, in the famous primordial soup, had
originated uncountable times, as an unavoidable
consequence of the properties of inorganic matter,
as the inorganic molecules arise from chemical
reactions rigorously determined by their context?
Should we hypothesize that at least a part of biod-
iversity was determined by distinct origins in the
primordial soup?
Removing this heretic thought, arisen from the
hesitations of my mind, let us consider a less worry-
ing problem of our taxonomic system: the indefin-
iteness of taxonomic categories, detectable from a
comparison between different phyla, classes or
orders. Are taxonomic categories pure classifica-
tion tools or they attempt to measure some aspect
of diversity? Molecular analysis has allowed the
122
Pietro Alicata
measurement of the genetic distances among taxa
and to calculate the time of their separation, even
in absence of paleontological data. But, the results,
despite the sophisticated mathematical methods
utilized, are grossly inadequate: solution of
problems at the species or subspecies level is
possible, but the phylogenetic trees remain highly
hypothetical.
Current taxonomy tries to represent the surpris-
ing phenotypic diversity of beings that has a
magnitude many times larger than the diversity of
genetic material. It would be most likely possible
to redefine the taxonomic categories according to
the level of phenotypic diversity. This would
require a free access to an exhaustive species’ docu-
mentation (description, figures, ecological notes,
and so on). Some farseeing scientists are pursuing
this aim for a few taxa.
However there are good reasons to preserve the
stability of taxonomy. For example, the great role
of taxonomy in nature conservation strategies: one
cannot preserve any living organism that does not
have a name. The prerequisite for the creation of
the IUCN Red List of Threatened Species is precise
taxonomic knowledge and changes in taxonomy
(for example variation in synonymy) can determine
changes in the status of a species. Also the level of
nature protection in a territory may be increased by
a new taxonomic evaluation of a biological species.
Of course there is a great need for taxonomy experts
to monitor the populations of protected species and
to evaluate the status of habitats relevant for nature
conservation. But, may these considerations of
mine reveal a conflict of interest?
At the end, we will have a good solution if,
while many mathematical minds endeavor to elab-
orate models to resolve evolutionary puzzles, tradi-
tional taxonomy continues to fulfil the biblical job of
giving a name to animals and plants, whose shapes,
colours and adaptations always attract the interest
of numerous enthusiastic scientists, as proven by
the success of this journal and of the meeting.
Biodiversity Journal, 2015, 6 (1): 123-138
Monograph
Taxonomy faces speciation: the origin of species or the fading
out of the species?
Alessandro Minelli
Department of Biology, University of Padova, Via Ugo Bassi 58 B, I 35131 Padova, Italy; email: alessandro.minelli@unipd.it
ABSTRACT Efficient field sampling and new investigation tools, including barcoding and other molecular
techniques, are bringing to light an unexpected wealth of new species, including sets of
morphologically quite uniform, but genetically distinct cryptic species. On the other hand,
increasing appreciation of the dynamic nature of the species and a better knowledge of
speciation processes and introgression phenomena challenges the taxonomists’ efforts to
shoehorn all diversity of life into a formal classification of which the species would be the
basic unit. Unfortunately, there is probably not a single best notion of species, either in theory
or in practice.
KEY WORDS barcoding; cryptic species; hybridization; speciation; species concepts.
Received 25.01.2015; accepted 26.02.2015; printed 30.03.2015
Proceedings of the 2nd International Congress “Speciation and Taxonomy”, May 1 6th- 1 8th 2014, Cefalu-Castelbuono (Italy)
THE SPECIES - A SOLID PILLAR OF
OUR REPRESENTATION OF LIVING
NATURE?
A substantial percentage of recent books and art-
icles in zoology, botany, palaeontology, biogeo-
graphy and ecology may suggest that the species has
passed undamaged through the Darwinian revolu-
tion. Although everybody, or so, in these disciplines
is likely ready to accept that species are products of
evolution, in practice a great many professionals
describe and analyze the living world of the past and
present time in terms not that different from those
of Linnaeus and the other pre-Darwinian authors.
In the title-page of his magnum opus, Linnaeus
(1758) announced an arrangement of his Systema
naturae per regna tria naturae, secundum classes,
ordines, genera, species. Firmly placed at the
bottom of the hierarchy, the species category was
thus proposed as the fundamental unit of classifica-
tion. In the author’s creationist and largely fixist
views, species have been created at the beginnings
of time and the naturalist’s job is to piously explore
Nature with the aim of completing their inventory.
What does survive to our time, of this reassuring
pre-Darwinian conception of biological diversity?
Little, if anything, in theoiy, but quite a lot in practice.
This is true both of the approach with which taxonom-
ists continue Linnaeus’ project for a global inventory
of biodiversity and of the perspective from which
most of their colleagues in biology, ecology, biogeo-
graphy and stratigraphy look at the extant or extinct
forms of life that are the object of their studies.
To be sure, there are also the ‘professionals of
the species problem,’ that is, biologists - but also
philosophers of biology - who take very seriously
the Darwinian challenge and specifically focus on
all those contexts where the boundaries between
species are less precise or less complete, and often
largely arbitrary.
The species problem has, in fact, two main
aspects. One is conceptual, the other is practical. The
124
Alessandro Minelli
conceptual aspect of the species problem is how the
species can, or should be defined, provided that this
question can be eventually answered to the general
satisfaction of biologists and philosophers alike. The
practical aspect is, how species are recognized by
taxonomists working on the different groups of
organisms and, most important, whether taxonom-
ists can all agree on a single species concept, to be
adopted as the universal currency in describing the
diversity of life. A comparison of taxonomic practice
as performed by leading specialists in a diversity of
taxa, from mammals to fungi, from bacteria to
flowering plants, has abundantly demonstrated that
the entities called species in a group have little in
common with the entities called species in another
group (Claridge et al., 1997). Unfortunately, this
heterogeneity is concealed under the (nearly)
universal use of Linnaean binomens. It is thus all too
easy to take taxonomic species as a set of broadly
comparable units, of which we can make statistics
for the most different purposes, e.g. biodiversity
assessments and comparisons of extant or extinct
faunas and floras. This practice should be best
avoided (Minelli, 2000) but we do not have a real
substitute for it; the global biodiversity estimates
offered below are not exempt from this ‘original sin.’
In this article I will focus on this practical aspect
to the species problem, mostly taking examples from
papers published in 2014: with this temporal restric-
tion I only wish to stress the lively interest surround-
ing these questions. The relevant literature is
enormous, and rapidly increasing with the increas-
ing availability of morphological and especially
molecular methods, and their massive application to
the most diverse kin ds of organisms. Enormous is
also the literature about the conceptual aspects of the
species problem, but I will only mention here two
articles (Bemardi & Minelli, 2011; Mallett, 2013) to
which I refer the interested reader and summarize in
Table 1 the most important among the more than 20
different species concepts proposed to date.
HOW MANY SPECIES?
Even if we temporarily ignore the problems
caused by the lack of a satisfactory species concept
applicable to every kind of living things and thus
simply frame the question in terms of taxonomic
(named) species, it is difficult to say how many
species we know at present and, still worse, how
many species still await description. Estimates of
‘valid’ described species range between 1.5 million
and 2 millions; a document issued in 2011 by the
International Institute for Species Exploration gave
a figure of 1,922,710 species as described through-
out 2009.
Something, however, must be wrong with many
of these estimates. In the last few decades, the
number of new species described each year has
been in the order of 17500 (International Institute
for Species Exploration, 2012). This means that
since 1985 about half a million new entries have
been added to the list of described species. The net
increase has been sensibly smaller, because of the
number of nominal species that in the same time
interval have been recognized to be just synonyms
of other species. However, the net increase has been
probably in the order of 350 000-400 000, whereas
the most recent estimates of the number of de-
scribed species are not correspondingly larger than
the estimates produced 30 years ago.
In the last two decades of the XX century a
number of papers offered estimates of the number
of living species that still await description, one of
the first and most often cited being May (1988).
Some estimates were based on the percentage of
undescribed species in small but dense samplings
in areas and habitats with high diversity, e.g.
Hodgkinson & Casson (1991) for tropical insects
and Grassle & Maciolek (1992) for deep-sea
animals. Other estimates included ecological
considerations, such as the degree of hostplant spe-
cialization of phytophagous insects, as in Erwin’s
(1982) pioneering paper or Stork’s (1988) revisita-
tion of the same. One of the most recent papers on
the subject adjusts the estimates to ~8.7 million (±1 .3
million SE) eukaryotic species globally, of which
~2.2 million (±0.18 million SE) are marine (Mora et
al., 2011); another, more sensible one (Scheffers et
al., 2012) acknowledges the plurality of unknown or
poorly known factors, as a consequence of which
uncertainty remains between a global total as low as
2 million species, microbes excluded, and estimates
as high as 50 millions and over.
PROLIFERATION OF NEW SPECIES
Strong catalysts favouring the description of
new species are the new megajoumals specifically
Taxonomy faces speciation: the origin of species or the fading out of the species?
125
devoted to taxonomy. In zoology, the leading role
of Zootaxa and ZooKeys is by now unrivalled and
undisputed. According to the journal’s editor-
in-chief, in 2010 Zootaxa contributed about 20% of
all animal taxa described that year as new, that is, a
number in the order of 4000 (Zhang, 2011).
Launched a few years after their zoological
equivalents, Phytotaxa and PhytoKeys have been
also rapidly growing and by now outcompete the
biggest journals long established in the field.
According to Zhang et al. (2014), the total number
of new plant taxa described in 2011 was 6024 (of
which 575 in Taxon, 473 in Phytotaxa, 183 in
Novon, 169 in the Botanical Journal of the Linnean
Society); in 2012, the total was 6647 (of which 632
in Phytotaxa, 465 in Systematic Botany, 340 in
Phytoneuron, 301 in the Kew Bulletin, 267 in
Taxon); in 2013 the number decreased to 5116 (of
which 501 in Phytotaxa, 248 in PhytoKeys, 199 in
Biodiversity Research and Conservation, 196 in the
Botanical Journal of the Linnean Society). In
discussing these numbers, it is necessary to consider
that these include taxa proposed at any taxonomic
rank.
Despite the large and largely unknown
degree of uncertainty surrounding the estimates
mentioned above, these are nevertheless
important. Besides the fact that these figures help
bringing the urgency of biodiversity conservation
to the public attention, estimates of gaps of
knowledge to be filled can stimulate targeted
efforts aiming at filling them.
Some research groups are currently addressing
this specific problem through well-planned field
work in lesser investigated and species-rich areas,
with special regard to hyperdiverse taxa such as
weevils. For example, a German team, supported
by local investigators in tropical areas, has recently
produced a couple of excellent papers on the
wingless weevils of the genus Trigonopterus .
Previous to the most recent researches, this
genus included 9 1 described species ranging from
Sumatra to Samoa and from the Philippines to
New Caledonia. Of these, 50 species of Trigonop-
terus had been described from New Guinea, the
center of the genus’ diversity. But new targeted
samplings in seven localities across New Guinea
have resulted in the recognition of 279 Trigonop-
terus species, most of which new to science;
of these, a first set of 101 species have been
described by Riedel et al. (2013). Another 98 new
species of Trigonopterus have been described in a
paper (Riedel et al., 2014) devoted to materials
recently collected in Indonesia (Sumatra, Java,
Bali, Palawan, Lombok, Sumbawa, Flores), a large
area from where only one species of Trigonopterus
was previously known.
Perhaps less expected, there are also large num-
bers of undescribed species in the Lepidoptera,
especially among the so-called micros. A recent
study of the gelechioid genus Ethmia in Costa Rica
revealed the presence of 22 undescribed species
in addition to 23 described in the past (Phillips-
Rodriguez et al., 2014).
Virtually unfathomed is, in some specialists’
view, the world of Fungi, of which the number of
existing species is estimated between 1.5 and 5
million, i.e. 15 to 50 times the number of currently
described species. The wealth of undescribed fungal
diversity is not limited to the microscopic forms: a
recent study reported the identification of at least
126 species (and potentially up to 400) within a
taxon of macrobasidiolichens currently regarded as
one species ( Dictyonema glabratum (Sprengel) D.
Hawksw. also known as Cora pavonia E. Fries)
(Lucking et al., 2014).
The use of new investigation tools such as
barcoding (discussed below) is precious, indeed, in
revealing the existence of a multiplicity of cryptic
species hitherto shoehorned under one species
name. I give here four examples, three of which
from papers published last year.
In polychaetes, for examples, cryptic species
crop up with virtually every accurate study. The
detailed review published by Nygren (2014)
includes several dozen examples, of which only the
most conspicuous ones (those with >5 cryptic
species inferred to be present within a taxon curren-
tly treated as a single species) are listed in Table 2.
The taxonomic complexity revealed by this study
is probably nothing more than the tip of a huge
iceberg of species diversity in the annelids. Most of
the ciyptic diversity discovered to date in poly-
chaetes is still formally undescribed, one of the few
exceptions being the five species of Archinome
listed in the Table.
Impressive are the results of some studies focus-
ing on individual genera, where a systematic use of
barcoding procedures has revealed an astonishing
diversity of species, morphologically very uniform,
126
Alessandro Minelli
as in some amphipods living in desert spring of the
southern Great Basin of California and Nevada,
USA, where 33 ‘provisional species’ have been
recognized within a clade hitherto referred to the
one species, Hyalella azteca Saussure, 1858 (Witt
et al., 2006).
A cornucopia of cryptic species, to use the
words of the authors (Winterbottom et al., 2014) has
been discovered in a DNA barcode analysis of the
gobiid fish genus Trimma. Here, 473 specimens
initially assigned to 52 morphological species
revealed the presence of 94 genetic lineages sep-
arated by a sequence divergence usually typical of
inter- rather than intraspecies differences.
To a quite smaller extent, but still worth men-
tioning here, new species are still being described
at a sensible rate even in groups such as mammals,
where a long tradition in taxonomy could be expec-
ted to have adequately accounted for extant species
diversity. Taxonomic unrest is obviously larger in
species-rich clades such as rodents or bats. For
example, several new species of the bat genus
Miniopterus have been recently described from
Madagascar and the neighbouring Comoros
archipelago, and at least seven out of the 18
species-level taxa recognized in the most recent
study still require formal taxonomic treatment
(Christidis et al., 2014).
TESTING THE BARCODE
“In 2003, Paul Hebert, researcher at the Univer-
sity of Guelph in Ontario, Canada, proposed “ DNA
barcoding” as a way to identify species. Barcoding
uses a very short genetic sequence from a standard
part of the genome the way a supermarket scanner
distinguishes products using the black stripes of the
Universal Product Code (UPC). Two items may look
very similar to the untrained eye, but in both cases
the barcodes are distinct. [. . .] The gene region that
is being used as the standard barcode for almost all
animal groups is a 648 base-pair region in the mi-
tochondrial cytochrome c oxidase 1 gene (“COl ”).
COI is proving highly effective in identifying birds,
butterflies, fish, flies and many other animal groups.
COI is not an effective barcode region in plants be-
cause it evolves too slowly, but two gene regions in
the chloroplast, matK and rbcL, have been approved
as the barcode regions for plants.”
This is the way this technique is described, in
very simple terms, in the official Barcode of Life
website http://www.barcodeoflife.org/.
During the last few years, DNA barcoding has
become a popular method for the identification of
species. How efficient and reliable is it? The ques-
tion can be reasonably asked in respect to groups
and areas for which an exhaustive taxonomic treat-
ment was already available, based on morphology,
and the recent barcoding effort has covered a large
percentage of the species recognized thus far.
In the case of insects, most published DNA bar-
coding studies focus on species of the Ephemerop-
tera (Ball et al., 2005; Stahls & Savolainen,
2008), Trichoptera (Zhou et al., 2011), Lepidoptera
(deWaard et al., 2009; Hausmann et al., 2011a,
2011b; Strutzenberger et al., 2011), Hymenoptera
(Smith & Fisher, 2009; Zaldivar-Riveron et al.,
2010) and Coleoptera (Raupach et al., 2010, 2011;
Greenstone et al., 2011; Astrin et al., 2012;
Woodcock et al., 2013).
Raupach et al. (2014) have recently tested the
efficiency of DNA barcoding for the Heteroptera
of Central Europe. Based on a conventional quant-
itative threshold currently accepted as a minimum
molecular difference between two species, they
found that species identification based on barcod-
ing sequences is correct in a 91.5% of cases. In 21
cases, the molecular distance between two tradi-
tionally accepted species is lower (in ten cases,
actually zero). To the contrary, intraspecific dif-
ferences larger than the conventional species-level
threshold have been found for 16 species tradi-
tionally regarded as valid. These results suggest
that the barcode cannot be blindly accepted as a
tool that allows quasi-automatic identification of
all species, but at the same it turns to be a useful
tool to discover taxa, or groups of closely related
taxa, that are in need of in-depth revision. In
particular, Raupach et al.’s study has provided
evidence for ongoing hybridization events within
various genera (e.g. Nabis, Lygus, Phytocoris ) as
well as the putative existence of cryptic species, e.g.
within the aradid Aneurus avenius (Dufour, 1833)
and the anthocorid Orius niger (Wolff, 1811).
Much larger success was obtained by Huemer
et al. (2014) in the identification via barcode
of 1004 species of Lepidoptera shared by two
European countries, Austria and Finland, ca. 1600
km apart. Correct identification was possible for
Taxonomy faces speciation: the origin of species or the fading out of the species?
127
98.8% of the taxa. However, deep intraspecific
divergence, larger than the conventional threshold
accepted as separating intra- from interspecific dif-
ference, was detected in as many as 124 taxonomic
species hitherto recognized based on morphology.
Authors concluded that despite the intensity of past
taxonomic work on European Lepidoptera, nearly
20% of the species shared by Austria and Finland
require further work to clarify their status.
The information obtained by systematically
applying the barcoding method to groups for which
traditional taxonomy is inadequate has different
consequences. For example, this technique has
been applied to the biting midges (Ceratopo-
gonidae) of the county of Finnmark in northern
Norway. Results indicated the presence of 54
species, of which 14 likely new to science, 16 new
to Norway, and one new to Europe (Stur &
Borkent, 2014). Another study involved a New
World genus of Curculionidae ( Conotrachelus ).
Two sets of specimens were compared, those emer-
ged from some 17 500 seeds collected in six
Central American rain forests and those collected
in the same forests using interception traps that
capture flying insects. Barcoding data suggested
the presence of 17 species in the trapped samples,
and 48 species among the specimens obtained from
the attacked seeds. Tittle hope to use previous
knowledge to identify them, however, as the
barcoding of representatives of 24 species from
museum collections provided matches for only
three of the 1 7 species from the traps and no match
at all for the putative 48 reared species (Pinzon-
Navarro et al., 2010).
Overall, barcoding methods have proven much
less informative for plants than the results obtained
from animals would have allowed to hope. A near
complete failure has been a study on willows ( Salix )
species, using two to seven plastid genome regions.
Of the 7 1 Holarctic species in that study, only one
has a unique barcode (Percy et al., 2014)!
THEORY-DRIVEN SPECIES INFLATION
This legitimate, welcome progress in the appre-
ciation of species diversity in lesser investigated
groups contrasts, to some extent, with a recent pro-
liferation of ‘new species’ proposed by some au-
thors in a revisitation of the taxonomy of popular
mammal clades such as carnivores and ungulates.
The theoretical background advocated by the
zoologists responsible for this ‘taxonomic inflation’
is the phylogenetic species concept, according to
which any arguably monophyletic and practically
diagnosable lineage deserves to be considered (and
eventually named) as a distinct species. With the
increasing use in taxonomy of molecular techniques
(e.g. barcoding), finding a differential trait between
two populations, e.g. a single nucleotide difference,
has become all too easy.
A first application to mammals of the phylogen-
etic species concept led Cracraft et al. (1998) to
raise the Sumatran tigers to species status {Pan-
ther a sumatrae Pocock, 1929) based on three dia-
gnostic sites in the mitochondrial cytochrome b
gene. Shortly thereafter, Mazak & Groves (2006)
added a third tiger species, the Javan tiger P. sonda-
ica (Temminck, 1844), to the previously estab-
lished P tigris (Finnaeus, 1758) and P. sumatrae.
Similarly, based on mtDNA and their analysis of
morphological diagnosability, Groves & Grubb
(2011) distinguished three species of European red
deer: Cervus elaphus Finnaeus, 1758 (West European
red deer), C. pannoniensis Banwell, 1997 (East
European red deer) and Cervus corsicanus Er-
xleben, 1777 (Corsico-Sardinian and North- African
red deer). Moreover, these are only a fraction of
the total of 12 species recognized by these authors
for the entire red deer/wapiti complex. Further
examples of oversplitting caused by the applic-
ation of the phylogenetic species concept include
the 1 1 species of klipspringer recognized within
one traditional species, Oreotragus oreotragus
(Zimmermann, 1783), based on size differences
and different sexual dimorphism, and the splitting
of the mainland serow Capricornis sumatraensis
(Bechstein, 1799) into six species (Groves &
Grubb, 2011). Zachos et al. (2013), who are very
critical of this trend in mammal taxonomy, acknow-
ledge however that in other groups more than one
species must be in fact recognized, as in the case
of the African elephants (the forest elephant
Loxodonta cy clods Matschie, 1900 and the
savanna elephant Loxodonta africana (Blumen-
bach, 1797); cf. Rohland et al., 2010), and the
giraffe, within which six or more distinct species
should be probably recognized (Groves & Grubb,
2011).
128
Alessandro Minelli
Agamospecies Concept
an operational, morphologically defined
unit in organisms that reproduce
asexually or by uniparental repro-
duction (without fertilization)
Cain (1954)
Biological Species Concept
a group of interbreeding natural
populations, reproductively isolated
from other similar groups
Dobzhansky (1935, 1937, 1970),
Mayr (1940, 1942, 1963, 1970),
Mayr & Ashlock, 1991)
Cladistic Species Concept
a group of organisms bounded by
two events of speciation or by a
speciation and an extinction event
Ridley (1989)
Cohesion Species Concept
the most inclusive group of organ-
isms within which genetic and/or
demographic exchange can occur
Templeton (1989)
Ecological Species Concept
a set of populations isolated through
occupation of a specific ecological
niche
Van Valen (1976).
Evolutionary Species Concept
an evolutionary lineage of popula-
tions in ancestor-descendant relation-
ship, that maintains its identity vs.
other lineages so defined, and with
its own specific evolutionary trends
and historical destiny
Simpson (1951, 1961)
Genetic Species Concept
the largest reproductive community
of sexual interfertile individuals that
share a common gene pool; or a
field for gene recombination
Dobzhansky (1950),
Carson (1957)
Hennigian Species Concept
a reproductively isolated natural pop-
ulation, or group of natural popula-
tions, issued from the dissolution of
a stem species in a speciation event,
that ceases to exist for extinction or
speciation
Meier & Willmann (2000)
Least Inclusive Taxonomic Unit
a taxonomic group defined on the
basis of apomorphies
Pleijel & Rouse (1999),
Pleijel (2000).
Morphological Species Concept
a community or a number of related
communities, whose distinctive mor-
phological characters are, in the opin-
ion of a competent systematist, suf-
ficiently defined to qualify it or them
with a specific name
Regan ( 1 926)
Phylogenetic Species Concept -
diagnosable version
the smallest diagnosable grouping of
organisms, within which there is a
pattern of ancestor-descendant rela-
tionship
Cracraft(1983)
Taxonomy faces speciation: the origin of species or the fading out of the species?
129
Phylogenetic Species Concept -
a monophyletic group of individuals
Rosen (1978), De Queiroz &
monophyly version
characterized by one or more auta-
pomorphies
Donoghue (1988)
Phenetic Species Concept
a set of organisms that are pheno-
typically similar and that look
different from other sets of organisms
Sneath (1976)
Recognition Species Concept
a group of organisms that share a
common fertilization system, or
better, a Specific Mate Recognition
System
Paterson (1979, 1985)
Table 1. A selection of species concepts, with short definitions, mainly in accordance with Bemardi & Minelli (2011) and
Mallett (2013), and some key references. Concepts that specifically apply to extinct organisms (the Successional Species
Concept in the two versions: George’s (1956) Chronospecies Concept, and Simpson’s (1961) Paleospecies Concept) are
not included.
Current taxon name(s)
Inferred number
of species
Archinome jasoni Borda et al., 2013, A. tethyana Borda et al., 2013, A. levinae Borda
et al., 2013, A. rosacea (Blake, 1985), A. storchi Fiege et Bock, 2009
5
Branchiomma spp.
11
Capitella capitata (Fabricius, 1780)
12+
Eumida sanguinea (Orsted, 1843)
11
Harmothoe imbricata (Linnaeus, 1767)
6
Leitoscoloplos pugettensis (Pettibone, 1957)
5
Marenzelleria viridis (Verrill, 1873), M. bastropi Bick, 2005, M. neglecta Sikorski et
Bick, 2004, M wireni Augener, 1913, M. arctia (Chamberlin, 1920)
5
Marphysa sanguinea (Montagu, 1815)
5
Ophryotrocha labronica Bacci et La Greca, 1961
14
Owenia fusiformis Delle Chiaje, 1844
5
Palola spp.
16
Sabellastarte spp.
7
Scoloplos armiger (Muller, 1776)
5-6
Syllis alternata Moore, 1908
5
Table 2. Cryptic diversity revealed in some polychaete ‘species’ taxa by recent molecular investigations
(data compiled from Nygren, 2014, Table SI).
130
Alessandro Minelli
TRICKY SPECIES COMPLEXES
Better investigated groups reveal a complexity
of interrelationship within which any formal taxo-
nomic arrangement is likely to remain provisional,
or at least arbitrary. Species complexes are particu-
larly intractable when the reproductive behavior of
some of the forms involved deviates from the typ-
ical biparental scheme. Exemplary in this respect is
the complex of the European green frogs, which
includes a number of hybridogenetic entities whose
survival strictly depends on an uninterrupted
availability of sperm from a closely related bipar-
ental species, as in the case of the Edible Frog, i.e.
the hybridogenetic Pelophylax lclepton esculentus
(Linnaeus, 1758). This hybrid between the Pool
Frog Pelophylax lessonae (Camerano, 1882) and
the Marsh Frog Pelophylax ridibundus (Pallas,
1771) is fertile, but usually unable to produce
balanced gametes of the two sorts, whereas it
usually survives by female hybrids mating with
males of one of the parental species, usually P.
lessonae (e.g., Spolsky & Uzzell, 1986; Christiansen,
2009). Local conditions are indeed extremely
diverse and are hardly amenable at a conventional
taxonomic treatment. In Central and Western
Europe the hybrid P. esculentus lives in sympatry
with the parental species P. lessonae (LE-system),
but there are also gamete-exchanging systems of P
ridibundus/P. esculentus (RE) and P. ridibundus/
P. lessonae/P. esculentus mixed populations (RLE)
(reviewed by Gunther, 1991; Plotner, 2005), and
also rare all-hybrid populations (EE-system) repro-
ductively independent of the parental forms (Graf &
Polls Pelaz, 1989) but dependent for sperm on the
presence of triploid individuals; the latter are
obtained when diploid eggs produced by diploid
hybrid females (LR) are fertilized by haploid sperm
of diploid or triploid males (LR, LLR, LRR) (Arioli
et al., 2010).
The taxonomic treatment of uniparental organ-
isms is generally difficult and controversial. Lin-
naean species are quite pacifically recognized in
some groups, e.g. in bdelloid rotifers, but in this
group thelytokous parthenogenesis is a very old
phenomenon and a number of largely fixed differ-
ences among strains have been fixed, that allow
recognizing species- and genus-level taxa around
which there is not much dispute. Things are differ-
ent in groups where parthenogenesis, or apomyxis,
is a recent phenomenon and phenotypic differences
between clonal strains are much more subtle and
their taxonomic evaluation much more subjective.
In the case of brambles ( Rubus spp.) and dandelions
( Taraxacum spp.) thousands of names have been
introduced to accommodate slightly divergent phen-
otypes at what some specialists consider the taxo-
nomic rank of species. In many instances, however,
uniparental reproduction is accompanied by vari-
ation in ploidy level and/or by morphological and
molecular distances comparable to those ordinarily
existing between related bisexual species, or even
larger. An interesting example has been recently
illustrated by Marotta et al. (2014) in the freshwater
oligochaetes of the genus Tubifex. Despite the
occurrence of different reproductive mechanisms
(biparental reproduction vs. thelytoky), many
populations referable to this genus have been tradi-
tionally classified as a single species Tubifex tubifex
(Muller, 1774). Under this name, however, is con-
cealed an unexpected diversity, as suggested by a
careful karyological and molecular analysis of
samples collected in just one limited area, the
Lambro River near Milano. Alongside a diploid
form, for which a distinct name ( T. blanchardi
Vejdovsky, 1891) is available in the literature, the au-
thors found several polyploid lines (3n, 4n, 6n), with
karyological differences matching with large molecu-
lar divergence in the 16S rRNA and COI sequences.
It will be no surprise if this diversity will eventually
emerge as just the tip of a still unfathomed iceberg.
The identification of gene flow between related
species is very important when taxa of economic and
especially medical or veterinary importance are
involved. Fontaine et al. (2015; see also Clark &
Messer, 2015) have recently demonstrated introgres-
sion in a medically important group of sibling species
of Afrotropical mosquitos ( Anopheles gambiae
Giles, 1902, A. coluzzii Coetzee et al., 2013 and A.
arabiensis Patton, 1905) that differ in behaviour and
thus in medical importance. Allele exchanges
between these malaria vectors have been found to
involve most of their autosomal genes, it is therefore
possible that traits enhancing vectorial capacity may
be gained through interspecific gene flow.
SPECIATION
Sooner or later, the taxonomist must confront
the issue of speciation, traditionally a focal issue
Taxonomy faces speciation: the origin of species or the fading out of the species?
131
in evolutionary biology, thus basically approached
through the tools of population genetics. Even-
tually, even the good practicing taxonomist who is
happy applying Regan’s (1926) morphological
species concept (cf. Table 1) is brought by the
intricacies of his/her study material to admit how
right was Darwin when he acknowledged that ‘Wo
line of demarcation can be drawn between species
... and varieties' ’ (Darwin, 1859, p. 469). It is
beyond the scope of this article to present here
even a short summary of current awareness, and
current debates, on the issue of speciation. The
interested reader is referred to Coyne & Orr’s
(2004) monograph, which is both a synthesis of
modern understanding of speciation problems, a
guide to older literature and a solid background
against which to read the literature of the last
decade. I will thus skip the traditional main issues,
beginning with the geographic scenarios of spe-
ciation (allopatric, parapatric, sympatric). I will
only glean from the very recent literature some
exemplary cases that show how cautious should
be the taxonomist in front of the temporal and
spatial change to which natural populations are
subjected. The more we know about these aspects,
the more critical should be our attitude towards a
taxonomic delimitation of species.
A first warning concerns the tempo of evolu-
tion. An unwarranted generalization of Darwin’s
depiction of evolution as proceeding through the
gradual accumulation of changes happening at a
very slow and essentially uniform pace led in the
past to assume that a speciation event should take
on the average some hundred thousand years or
more. There is no reason, however, for us to expect
that living nature adopts an essentially uniform
pace of change. Indeed, we have now well-docu-
mented proofs of very rapid speciation events, and
also of extremely conservative species pairs whose
remote splitting is concealed under an amazing
degree of morphological stasis. As a consequence,
the taxonomist must be cautious in inferring related-
ness from morphological, ecological or biogeo-
graphic evidence without the further support of
molecular estimates of divergence times.
Consider, for example, that the divergence
between two species of amphioxus, both currently
classified in the same genus, Branchiostoma
floridae Hubbs, 1922 and B. lanceolatum (Pallas,
1774), has been estimated at 186-189 million
years (Canestro et al., 2002), whereas the origin
of the whole radiation of extant Brassicaceae
(3709 species; Warwick et al., 2006) is probably
not older than 40 million years (Couvreur et al.,
2010; Franzke et al., 2011), and perhaps even
younger, around 16 million years (Franzke et al.,
2009). This can be compared to the 22.4 million
years through which the hummingbirds (338 living
species) have been apparently radiating from their
last common ancestor (McGuire et al., 2014). Still
very long times, indeed, if compared to the 100
000 years, or so, within which the cichlids of Fake
Victoria have radiated into a species flocks of five
hundred species at least (Verheyen et al., 2003;
Genner et al., 2007).
GENES INVOLVED IN SPECIATION
Research on the genes more directly involved
in speciation is attracting increasing interest, but
convincing generalizations are still difficult to
obtain.
Problems in fixing the boundary between two
closely related taxa that broadly, but not completely
exhibit the character of distinct species are often
due to the fact that some parts of their genome are
more readily and extensively affected by introgres-
sion, whereas other parts are much more resilient.
A classic case - Carrion Crow ( Corvus corone
Linnaeus, 1758) vs. Hooded Crow ( Corvus cornix
Linnaeus, 1758) - has been carefully investigated
by Poelstra et al. (2014). These authors have found
that only a small number of narrow genomic islands
are not affected by gene flow. As mirrored by these
birds’ livery, gene expression divergence between
them is concentrated in pigmentation genes ex-
pressed in gray versus black feather follicles.
Despite its limited genetic basis, this trait is critic-
ally important, however, as it affects mate choice
and thus color-mediated prezygotic isolation.
In pairs of stick insect populations adapted to
different host plants and undergoing parallel speci-
ation, Soria-Carrasco et al. (2014) found thousands
of small genomic regions, most of which unique to
individual population pairs, to be significantly
diverging between populations. These authors have
also detected parallel genomic divergence across
population pairs involving an excess of coding genes
with specific molecular functions.
132
Alessandro Minelli
STABILITY OF SPECIES IN THE FACE
OF INTROGRESSION
While the existence of introgression between
locally sympatric related species is well docu-
mented in a large number of animals and plant
species pairs, very little is known about the long-
term effects of a gene flow continuing over centur-
ies. A recent study of two widely hybridizing tree
species, the white spruce (Picea glauca (Moench)
Voss) and Engelmann spruce (P. engelmannii Parry
ex Engelm.) in western North America, suggests
that these two species have a long history of hybrid-
ization and introgression, dating to at least 2 1 000
years ago, nevertheless they still maintain their
distinct species identity (De La Torre et al., 2014).
The boundaries between closely related species
are sometimes permeable in one direction only. For
example, brown bear ( Ursus arctos Linnaeus, 1758)
and polar bear ( Ursus maritimus Phipps, 1774) are
genetically distinct, but evidence of polar bear
genes has been found in the brown bear population
of the Admiralty, Baranof and Chicagof Islands off
Alaska, whereas no evidence of brown bear genes
has been found in the local polar bear population
(Cahill et al., 2015). Another example of asym-
metric introgression has been recently described
between a pair of freshwater fish, the North Amer-
ican darters Etheostoma caeruleum Storer, 1845
and Etheostoma spectabile (Agassiz, 1854) (Zhou
& Fuller, 2014).
HYBRIDIZATION
Opportunities for hybridization between closely
related biological species are not restricted to
species pairs that have being diverging only in recent
time, witness a fern from the French Pyrenees
( Cystocarpium x roskamianum Fraser-Jenk), a re-
cently formed hybrid whose parental lineages
diverged from each other ca. 60 million years ago,
and are currently classified in different genera (Cys-
topteris and Gymnocarpium) (Rothfels et al., 2015).
Due to both climatic and biological reasons,
hybrid zones are not fixed in space. Detailed
evidence of moving hybrid zones has summarized
by Buggs (2007) for the following pairs of taxa
(nomenclature updated where necessary):
MAMMALIA
Cervus nippon nippon Temminck, 1838 - Cer-
vus elaphus Linnaeus, 1758
AVES
Poecile carolinensis (Audubon, 1834) - Poecile
atricapillus (Linnaeus, 1766)
Hippolais polyglotta (Vieillot, 1 8 1 7) - Hippolais
icterina (Vieillot, 1817)
Vermivora pimis (Linnaeus, 1766) - Vermivora
chrysoptera (Linnaeus, 1766)
Corvus corone corone Linnaeus, 1758 - Corvus
corone cornix Linnaeus, 1758
Quiscalus quiscula quiscula (Linnaeus, 1758) -
Quiscalus quiscula versicolor Vieillot, 1819
SQUAMATA
Pholidobolus montium (Peters, 1 863) - Pholido-
bolus affinis (Peters, 1863)
Sceloporus tristichus (Cope, 1875) - Sceloporus
cowlesi Lowe et Norris, 1956
AMPHIBIA
Pseudophryne bibroni Gunther, 1859 -
Pseudophryne semimarmorata Lucas, 1892
Triturus cristatus Laurenti, 1768 - Triturus
marmoratus (Latreille, 1800)
Plethodon glutinosus (Green, 1818) - Plethodon
jordani Blatchley, 1901
OSTEICHTHYES
Pseudorasbora parva (Temminck et Schlegel,
1846) - Pseudorasbora pumila Miyadi, 1930
INSECTA
Heliconius hydara Hewitson, 1867 - Heliconius
erato petiverana (E. Doubleday, 1847)
Anartia fatima (Fabricius, 1793) - Anartia
amathea (Linnaeus, 1758)
Solenopsis invicta Buren, 1972 - Solenopsis
richteri Forel, 1909
Orchelimum nigripes Scudder, 1875 - Orche-
limum pulchellum Davis, 1909
Allonemobius socius (Scudder, 1877) - Allonemo-
bius fasciatus (De Geer, 1773)
Limnoporus dissortis (Drake et Harris, 1930) -
Limnoporus notabilis (Drake et Hottes, 1925)
Geomydoecus aurei Price et Hellenthal, 1981 -
Geomydoecus centralis Price et Hellenthal, 1981
Taxonomy faces speciation: the origin of species or the fading out of the species?
133
CRUSTACEA
Orconectes rusticus (Girard, 1 852) - Orconectes
propinquus (Girard, 1852)
ANGIOSPERMAE
Helianthus annuus L. - Helianthus bolanderi A.
Gray
Mercurialis annua L. diploid - Mercurialis
annua L. hexaploid
Occasionally, the peculiar geographical distri-
bution of a set of populations offers the opportunity
to investigate different stages of an ongoing speci-
ation process. This happens with the so-called ring
species, where the two extremes, say A and E, of a
series of progressively differentiated populations
have recently come in contact but fail to interbreed.
This happens generally when the whole complex is
distributed, ring-like, around an inhospitable area,
such as very high mountains, or an exceedingly arid
area. Ring species are extremely rare in plants:
recently, Cacho & Baum (2012) have presented the
Caribbean slipper spurge ( Euphorbia tithymaloides)
as the first example among the flowering plants.
More numerous are the zoological examples, as
summarized by Irwin et al. (2001). These authors
listed seventeen examples where the populations at
the opposite ends of the chain overlap without any
sign of hybridization, or nearly so. In many cases
the two extreme forms have been given distinct
specific names, whereas in other cases taxonomists
still treat all the populations involved in the ring as
belonging to the same Linnaean species: one ex-
ample, among a number of possible ones, of the
danger of inferring evolutionary status from simply
considering the current taxonomic status (i.e., the
nomenclature) of a set of populations.
Irwin et al.’s (2001) list includes a number of
birds: Crested Honey-buzzard Pernis ptilorhyncus
(Temminck, 1821) and Barred Honey-buzzard P.
celebensis Wallace, 1868; Herring Gull Larus
argentatus Pontoppidan, 1763 and Lesser Black-
backed Gull L. fuscus Linnaeus, 1758 (with some
hybridization); Ringed Plover Charadrius hiaticula
Linnaeus, 1758 and Semipalmated Plover C. semi-
palmatus Bonaparte, 1825; Collared Kingfisher
Todiramphus chloris (Boddaert, 1783) andMicrone-
sian Kingfisher T. cinnamominus (Swainson, 1821);
Eurasian Skylark dGw<7<2 arvensis Linnaeus, 1758,
Japanese Skylark A. japonica Temminck et Schle-
gel, 1848 and Oriental Skylark (A. gulgula Franklin,
1831); Greenish Warbler Phylloscopus trochiloides
(Sundevall, 1837); Chi ffchaff Phylloscopus colly-
bita ( Vieillot, 1817) and Mountain Chiffchaff {P.
sindianus W. E. Brooks, 1880); Sulawesi Triller
Lalage leucopygialis Walden, 1872, Pied Triller L.
nigra (J. R. Forster, 1781), and White-shouldered
Triller L. sueurii (Vieillot, 1818); Brown Thornhill
Acanthiza pusilla (Shaw, 1790) and Tasmanian
Thornhill d. ewingii Gould, 1844; Large Tree-finch
Camarhynchus psittacula Gould, 1837 and Medium
Tree-finch C. pauper Ridgway, 1890.
The other taxa in the list are rodents (Deer Mouse
Peromyscus maniculatus (Wagner, 1845); Pocket
Mice Perognathus amplus Osgood, 1900 and P.
longimembris (Coues, 1875), a bee Hoplitis producta
(Cresson, 1864), a group of butterflies Junonia
coenia Hiibner, [1822] and J. genoveva (Cramer,
1780)//. evarete (Cramer, 1782)) and a fruit fly {Dro-
sophila paulistorum Dobzhansky et Pavan, 1949).
In the case of the salamander Ensatina
eschscholtzii Gray, 1850, some hybridization
between the end forms of the ring has been
reported, and past but still recognizable hybridiza-
tion has been found in the ring of the Japanese pond
frogs Rana nigromaculata Hallowell, 1861 and R.
brevipoda Ito, 1941.
In still other cases, there is no reproductive isol-
ation between the two, now overlapping, terminal
forms of the ring; as a consequence, a hybrid zone
is formed. The cases listed by Irwin et al. (2001)
include birds Crimson Rosella Platycercus elegans
(Gmelin, 1788), Adelaide Rosella P adelaidae
Gould, 1841, Yellow Rosella P jlaveolus Gould,
1837, Great Tit Parus major Linnaeus, 1758, a
mammal House Mouse Mus musculus Linnaeus,
1758 and two millipedes Rhymogona silvatica
(Verhoeff, 1894) and R. cervina (Verhoeff, 1910).
Several ring species (putative ones as well as
confirmed ones) have been extensively studied over
the last few years. No wonder, the actual interrela-
tionships among the involved populations are often
more complex than in the simple model outlined
above. For example, in the case of the Greenish
Warbler Phylloscopus trochiloides (Sundevall,
1837) species complex Alcaide et al. (2014) have
recently revisited the status, and the history, of the
ring of populations distributed around Tibet. The
two extreme, reproductively isolated forms co-
existing in central Siberia are connected through a
134
Alessandro Minelli
southern chain of populations showing a gradient of
genetic and phenotypic traits. The authors demon-
strate that the gene flow has been interrupted in the
past at more than one location around the ring,
whereas the two Siberian forms have occasionally
interbred. Eventually, this little bird displays a con-
tinuum from slightly divergent contiguous popula-
tions to almost fully reproductively isolated species.
RETICULATION
Patterns of hybridization and introgression
among closely related taxa take sometimes a
reticulated structure. A recently investigated ex-
ample involving the biogeographical history of the
Eurasian species of Fraxinus has revealed the
occurrence of an ancient reticulation between
European and Asian species as well as other ancient
reticulation events between F. angustifolia Vahl and
F. excelsior L. and the other species of the section
Fraxinus. Some of these events would have oc-
curred during the Miocene, following climatic
variations that may have led these species to expand
their distribution range, eventually coming into
contact (Hinsinger et al., 2014).
SPECIATION REVERSED
Incomplete speciation and ongoing gene flux
between partially isolated populations may cause
divergence to be stopped and even reversed. Well-
documented cases of reversed speciation are, how-
ever, very limited. An example has been described
by Bhat et al. (2014) for the European whitefish
Coregonus lavaretus (Linnaeus, 1758), of Lake
Skrukkebukta in Northern Norway. This freshwater
fish is highly polymorphic and in several lakes it
has independently differentiated into sympatric
morphs that specialize on different food (plankton
vs. benthos) and are to some extent reproductively
isolated and genetically differentiated. In 1993,
Lake Skrukkebukta was invaded by another Core-
gonus species, the vendace Coregonus albula
(Linnaeus, 1758). A zooplanktivorous specialist,
this fish displaced the planktivorous whitefish from
the pelagic niche pushing it into the benthic habitat
already inhabited by the benthivorous whitefish
morphs. As a consequence, within three generations
(15 years) the genetic differentiation between the
two whitefish morphs has dramatically dropped: the
invasion of a superior trophic competitor has thus
caused incipient speciation to reverse. An overview
of cases of speciation reversal was provided a few
years ago by Seehausen et al. (2008).
REFERENCES
Alcaide M., Scordato E.C., Price T.D. & Irwin D.E.,
2014. Genomic divergence in a ring species complex.
Nature, 511: 83-85.
Arioli M., Jakob C. & Reyer H.U., 2010. Genetic di-
versity in water frog hybrids ( Pelophylax esculentus)
varies with population structure and geographic
location. Molecular Ecology, 19: 1814-1828.
Astrin J.J., Stiiben P.E., Misof B., Wagele J.W., Gimnich
F., Raupach, M.J. & Ahrens, D., 2012. Exploring
diversity in cryptorhynchine weevils (Coleoptera)
using distance-, character- and tree-based species
delineation. Molecular Phylogenetics and Evolution,
63: 1-14.
Ball S.L., Hebert P.D.N., Burian S.K. & Webb J.M.,
2005. Biological identifications of mayflies (Eph-
emeroptera) using DNA barcodes. Journal of the
North American Benthological Society, 24: 508-524.
Bernardi M. & Minelli A., 2011. II concetto di specie e
la paleontologia: una rassegna introduttiva. Ren-
diconti Online della Societa Geologica Italiana,
Volume speciale 13: 1-24.
Bhat S., Amundsen P.-A., Knudsen R., Gjelland K.0.,
Fevolden S.-E., Bernatchez L. & Praebel K., 2014.
Speciation reversal in European whitefish ( Corego-
nus lavaretus (L.)) caused by competitor invasion.
PLoS ONE, 9(3): e91208.
Buggs R.J.A., 2007. Empirical study of hybrid zone
movement. Heredity, 99: 301-312.
Cacho N.I. & Baum D.A., 2012. The Caribbean slipper
spurge Euphorbia tithymaloides : the first example of
a ring species in plants. Proceedings of the Royal
Society B, 279: 3377-3383.
Cahill J.A., Stirling I., Kistler L., Salamzade R., Ersmark
E., Fulton T.L., Mathias Stiller M., Green R.E. &
Shapiro B., 2015. Genomic evidence of geographic-
ally widespread effect of gene flow from polar bears
into brown bears. Molecular Ecology, DOI:
10. 1111/mec. 13038.
Cain A.J., 1954. Animal species and their evolution.
Hutchinson University Library, London, ix+190 pp.
Canestro C., Albalat R., Hjelmqvist L., Godoy L., Jor-
nvall H. & Gonzalez-Duarte R., 2002. Ascidian and
amphioxus Adh genes correlate functional and mo-
lecular features of the ADH family expansion during
Taxonomy faces speciation: the origin of species or the fading out of the species?
135
vertebrate evolution. Journal of Molecular Evolution,
54: 81-89.
Carson L.H., 1957. The species as a field for gene recom-
bination. In: E. Mayr, Editor, The species problem.
American Association for the Advancement of
Science, Washington: 23-38.
Christiansen D.G., 2009. Gamete types, sex determina-
tion and stable equilibria of all-hybrid populations of
diploid and triploid edible frogs ( Pelophylax escu-
lentus ). BMC Evolutionary Biology, 9: 135.
Christidis L., Goodman S.M., Naughton K., Appleton B.,
2014. Insights into the evolution of a cryptic radiation
of bats: dispersal and ecological radiation of
Malagasy Miniopterus (Chiroptera: Miniopteridae).
PLoS ONE, 9(3): e92440.
Claridge M.F., Dawah H.A. & Wilson M.R. (Eds.), 1997.
The species: the units of biodiversity. Chapman &
Hall, London: xvi+439 pp.
Clark A. G. & Messer P.W., 2015. Conundrum of jumbled
mosquito genomes. Science, 347: 27-28.
Couvreur T.L.P., Franzke A., Al-Shehbaz I. A., Bakker
F.T., Koch M.A. & Mummenhoff K., 2010.
Molecular phylogenies, temporal diversification, and
principles of evolution in the mustard family
(Brassicaceae). Molecular Biology and Evolution,
27: 55-71.
Coyne J.A. & Orr H.A., 2004. Speciation. Sinauer Asso-
ciates, Sunderland, MA: xiii+545 pp.
Cracraft J., 1983. Species concepts and speciation
analysis. Current Ornithology, 1: 159-187.
Cracraft J., Feinstein J., Vaughn J. & Helm-Bychowski
K., 1998. Sorting out tigers ( Panthera tigris ): mi-
tochondrial sequences, nuclear inserts, systematics,
and conservation genetics. Animal Conservation, 1 :
139-150.
Darwin C., 1859. On the origin of species by means of
natural selection, or the preservation of favoured
races in the struggle for life. Murray, London: x+502
pp.
De La Torre A.R., Roberts D.R. & Aitken S.N., 2014.
Genome-wide admixture and ecological niche
modelling reveal the maintenance of species bound-
aries despite long history of interspecific gene flow.
Molecular Ecology, 23: 2046-2059.
De Queiroz K. & Donoghue M.J., 1988. Phylogenetic
systematics and the species problem. Cladistics, 4:
317-338.
deWaard J.R., Landry J.-F., Schmidt B.C., Derhoussoff
J., McLean J.A. & Humble L.M., 2009. In the dark
in a large urban park: DNA barcodes illuminate
cryptic and introduced moth species. Biodiversity
and Conservation, 18: 3825-3839.
Dobzhansky T., 1935. A critique of the species concept
in biology. Philosophy of Science, 2: 344-355.
Dobzhansky T., 1937. Genetics and the origin of species.
Columbia University Press, New York, xvi+364 pp.
Dobzhansky T., 1950. Mendelian populations and their
evolution. American Naturalist, 84: 401-418.
Dobzhansky T., 1970. Genetics of the evolutionary
process. Columbia University Press, New York,
ix+505 pp.
Erwin T.L., 1982. Tropical forests: their richness in
Coleoptera and other arthropod species. Coleopterists
Bulletin, 36: 74-75.
Fontaine M.C., Pease J.B., Steele A., Waterhouse R.W.,
Neafsey D.E., Sharakhov I.V., Jiang X., Hall A.B.,
Catteruccia F., Kakani E., Mitchell S.N., Wu Y.-C.,
Smith H.A., Love R.R., Lawniczak M.K., Slotman
M.A., Emrich S.J., Hahn M.W. & Besansky N.J.,
2015. Extensive introgression in a malaria vector
species complex revealed by phylogenomics.
Science, 347: 1258524.
Franzke A., German D., Al-Shehbaz I. A. & Mummen-
hoff K., 2009. Arabidopsis family ties: molecular
phylogeny and age estimates in the Brassicaceae.
Taxon, 58: 425-437.
Franzke A., Lysak M.A., Al-Shehbaz I. A., Koch M.A. &
Mummenhoff K, 2011. Cabbage family affairs: the
evolutionary history of Brassicaceae. Trends in Plant
Science, 16: 108-116.
Genner M.J., Seehausen O., Lunt D.H., Joyce D.A.,
Shaw P.W., Carvalho G.R. & Turner G.F., 2007. Age
of cichlids: new dates for ancient lake fish radiations.
Molecular Biology and Evolution, 24: 1269-1282.
George T.N., 1956. Biospecies, chronospecies and
morphospecies. In: P.C. Sylvester-Bradley, Editor,
The species concept in paleontology. The Systematics
Association, London: 123-137.
Graf J.-D. & Polls Pelaz M., 1989. Evolutionary genetics
of the Rana esculenta complex. In: R.M. Dawley &
J.P. Bogart (Eds.), Evolution and ecology of unisex
vertebrates. New York State Museum, Albany, NY:
289-302.
Grassle J.F. & Maciolek N.J., 1992. Deep-sea species
richness: regional and local diversity estimates from
quantitative bottom samples. American Naturalist,
139:313-341.
Greenstone M.H., Vandenberg N.J. & Hu J.H., 2011.
Barcode haplotype variation in North American
agroecosystem lady beetles (Coleoptera: Coccinelli-
dae). Molecular Ecology Resources, 11: 629-637.
Groves C. & Grubb P., 2011. Ungulate taxonomy. The
Johns Hopkins University Press, Baltimore: ix+317
pp.
Gunther R., 1991. Europaische Wasserfrosche (Anura,
Ranidae) und biologisches Artkonzept. Mitteilun-
gen aus dem Zoologischen Museum in Berlin, 67:
39-53.
136
Alessandro Minelli
Hausmann A., Haszprunar G. & Hebert P.D.N., 2011a.
DNA barcoding the geometrid fauna of Bavaria
(Lepidoptera): successes, surprises, and questions.
PLoS ONE, 6: el7134.
Hausmann A., Haszprunar G., Segerer A.H., Speidel W.,
Behounek G. & Hebert P.D.N., 2011b. Now DNA-
barcoded: the butterflies and larger moths of Ger-
many (Lepidoptera: Rhopalocera, Macroheterocera).
Spixiana, 34: 47-58.
Hinsinger D.D., Gaudeul M., Couloux A., Bousquet J.,
Frascaria-Lacoste N., 2014. The phylogeography of
Eurasian Fraxinus species reveals ancient transcon-
tinental reticulation. Molecular Phylogenetics and
Evolution, 77: 223-237.
Hodkinson I.D. & Casson D., 1991. A lesser predilection
for bugs: Hemiptera (Insecta) diversity in tropical
rain forests. Biological Journal of the Linnean
Society, 43: 101-109.
Huemer P., Mutanen M., Sefc K.M. & Hebert P.D.N.,
2014. Testing DNA barcode performance in 1000
Species of European Lepidoptera: large geographic
distances have small genetic impacts. PLoS ONE,
9(12): ell5774.
International Institute for Species Exploration, 2011.
SOS - State of Observed Species. Arizona State
University, Tucson, AZ; available at http://www.esf.
edu/species/SOS.htm
International Institute for Species Exploration, 2012.
Retro SOS 2000-2009. Arizona State University,
Tucson, AZ; available at http://www.esf.edu/species/
SOS.htm
Irwin D.E., Irwin J.H. & Price T. D., 2001. Ring species
as bridges between microevolution and speciation.
Genetica, 112-113:223-243.
Linnaeus C., 1758. Sy sterna Naturae per regna tria
Naturae secundum classes, ordines, genera, species,
cum characteribus, differentiis, synonymis, locis.
Editio decima, reformata. Laurentius Salvius,
Holmiae, Tomus I: [4]+l-823+[l] pp.; Tomus II:
[4J+825-1384 pp.
Lucking R., Dal-Forno M., Sikaroodi M., Gillevet P.M.,
Bungartz F., Moncada B., Yanez-Ayabaca A., Chaves
J.L., Coca L.F. & Lawrey J.D., 2014. A single macro-
lichen constitutes hundreds of unrecognized species.
Proceedings of the National Academy of Science of
the United States of America, 111: 11091-11096.
Mallet J., 2013. Species, concepts of. In: S.A. Levin,
Editor, Encyclopedia of biodiversity, Volume 6.
Waltham, MA, Academic Press: 679-691.
Marotta R., Crottini A., Raimondi E., Fondello C. &
Ferraguti M., 2014. Alike but different: the evolution
of the T ubifex tubifex species complex (Annelida,
Clitellata) through polyploidization. BMC Evolution-
ary Biology, 14:73.
May R.M., 1988. How many species are there on earth?
Science, 241: 1441-1449.
Mayr E., 1940. Speciation phenomena in birds. American
Naturalist, 74: 249-278.
Mayr E., 1942. Systematics and the origin of species
from the viewpoint of a zoologist. Columbia Univer-
sity Press, New York: xiv+334 pp.
Mayr E., 1963. Animal species and evolution. The
Belknap Press of Harvard University Press,
Cambridge: xiv+797 pp.
Mayr E., 1970. Populations, species, and evolution: an
abridgment of animal species and evolution. The
Belknap Press of Harvard University Press,
Cambridge: xv+453 pp.
Mayr E. & Ashlock P.D., 1991. Principles of systematic
zoology. McGraw-Hill, New York: xx+475 pp.
Mazak J.H. & Groves C.P., 2006. A taxonomic revision
of tigers ( Panthera tigris ) of Southeast Asia.
Mammalian Biology, 71: 268-287.
McGuire J.A., Witt C.C., Remsen J.V. Jr, Corl A.,
Rabosky D.L. Altshuler D.L. & Dudley R., 2014.
Molecular phylogenetics and the diversification of
hummingbirds. Current Biology, 24: 1-7.
Meier R. & Willmami R., 2000. The Hennigian species
concept. In: Q.D. Wheeler, R. Meier, Editors,
Species concepts and phylogenetic theory: a debate.
Columbia University Press, New York: 30-43.
Minelli A., 2000. The ranks and the names of species and
higher taxa, or, a dangerous inertia of the language
of natural history. In: M.T. Ghiselin & A.E. Leviton,
editors, Cultures and institutions of natural history.
Essays in the history and philosophy of science.
California Academy of Sciences, San Francisco:
339-351.
Mora C., Tittensor D.P., Adi S. & Simpson A.G.B., 2011.
How many species are there on earth and in the
ocean? PLoS Biology, 9 e 100 1127.
Nygren A., 2014. Cryptic polychaete diversity: a review.
Zoologica Scripta, 43: 172-183.
Paterson H.E.H., 1979. A comment on ‘mate recognition
systems.’ Evolution, 34: 330-331.
Paterson H.E.H., 1985. The recognition concept of
species. In: E.S. Vrba, Editor, Species and speciation
(Transvaal Museum Monograph 4). Transvaal
Museum, Pretoria: 21-29.
Percy D.M., Argus G.W., Cronk Q.C., Fazekas A.J.,
Kesanakurti P.R., Burgess K.S., Husband B.C.,
Newmaster S.G., Barrett S.C.H. & Graham S.W.,
2014. Understanding the spectacular failure of DNA
barcoding in willows ( Salix ): does this result from a
trans-specific selective sweep? Molecular Ecology,
23: 4737-4756.
Phillips-Rodriguez E., Powell J.A., Hallwachs W. &
Janzen D.H., 2014. A synopsis of the genus Ethmia
Hiibner in Costa Rica: biology, distribution, and
Taxonomy faces speciation: the origin of species or the fading out of the species?
137
description of 22 new species (Lepidoptera, Gelech-
ioidea, Depressariidae, Ethmiinae), with em-
phasis on the 42 species known from Area de
Conservacion Guanacaste. ZooKeys, 461: 1-86.
Pinzon-Navarro S., Barrios H., Murria C., Lyal C.H.C.
& Vogler A.P., 2010. DNA-based taxonomy of larval
stages reveals huge unknown species diversity in
neotropical seed weevils (genus Conotrachelus ):
relevance to evolutionary ecology. Molecular
Phylogenetics and Evolution, 56: 281-293.
Pleijel F., 2000. Phylogenetic taxonomy, a farewell to
species, and a revision of Heteropodarke (He-
sionidae, Polychaeta, Annelida). Systematic Biology,
48: 755-789.
Pleijel F. & Rouse G.W., 1999. Least-inclusive taxo-
nomic unit: a new taxonomic concept for biology.
Proceedings of the Royal Society of London B, 267:
627-630.
Plotner J., 2005. Die westpalaarktischen Wasserfrosche.
Laurenti-Verlag, Bielefeld, 160 pp.
Poelstra J.W., Vijay N., Bossu C.M., Lantz H., Ryll B.,
Mueller I., Baglione V., Unneberg P., Wikelski M.,
Grabherr M.G., Wolf J.B.W. 2014. The genomic
landscape underlying phenotypic integrity in the face
of gene flow in crows. Science, 344: 1410-1414.
Raupach M.J., Astrin J.J., Hannig K., Peters M.K.,
Stoeckle M.Y. & Wagele, J.-W., 2010. Molecular
species identifications of Central European ground
beetles (Coleoptera: Carabidae) using nuclear rDNA
expansion segments and DNA barcodes. Frontiers in
Zoology, 7: 26.
Raupach M.J., Hannig K. & Wagele J.W., 2011. Identi-
fication of Central European ground beetles of the
genus Bembidion (Coleoptera: Carabidae) using
DNA barcodes: a case study of selected species.
Angewandte Carabidologie, 9: 63-72.
Raupach M.J., Hendrich L., Kuchler S.M., Deister F.,
Moriniere J. & Gossner M.M., 2014. Building-up of
a DNA barcode library for true bugs (Insecta: Hemi-
ptera: Heteroptera) of Germany reveals taxonomic
uncertainties and surprises. PLoS ONE, 9(9):
el 06940.
Regan C., 1926. Organic evolution. Report of the British
Association for the Advancement of Science 1925:
75-86.
Ridley M., 1989. The cladistic solution to the species
problem. Biology and Philosophy, 4: 1-16.
Riedel A., Sagata K., Surbakti S., Tanzler R. & Balke M.,
2013. One hundred and one new species of Trigonop-
terus weevils from New Guinea. ZooKeys, 280: 1-
150.
Riedel A., Tanzler R., Balke M., Rahmadi C. &
Suhardjono Y.R., 2014. Ninety-eight new species of
Trigonopterus weevils from Sundaland and the
Lesser Sunda Islands. ZooKeys, 467: 1-162.
Rohland N., Reich D., Mallick S., Meyer M., Green R.E.,
Georgiadis N.J., Roca A.L. & Hofreiter M., 2010.
Genomic DNA sequences from mastodon and woolly
mammoth reveal deep speciation of forest and
savanna elephants. PLoS Biology, 8 (12): el000564.
Rosen D.E., 1978. Vicariant patterns and historical
explanation in biogeography. Systematic Zoology,
27: 159-188.
Rothfels C.J., Johnson A.K., Hovenkamp P.H., Swofford
D.L., Roskam H.C., Fraser- Jenkins C.R., Windham
M.D. & Pryer K.M., 2015. Natural hybridization
between genera that diverged from each other approx-
imately 60 million years ago. American Naturalist,
185: 433-442.
Scheffers B.R., Joppa L.N., Pimm S.L. & Laurance W.F.,
2012. What we know and don’t know about Earth’s
missing biodiversity. Trends in Ecology and Evolu-
tion, 27: 501-510.
Seehausen O., Takimoto G., Roy D. & Jokela J., 2008.
Speciation reversal and biodiversity dynamics with
hybridization in changing environments. Molecular
Ecology, 17: 30-44.
Simpson G.G., 1951. The species concept. Evolution, 5:
285-298.
Simpson G.G., 1961. Principles of animal taxonomy.
Columbia University Press, New York: xii+247 pp.
Smith M.A. & Fisher B.L., 2009. Invasion, DNA
barcodes, and rapid biodiversity assessment using the
ants of Mauritius. Frontiers in Zoology, 6: 31.
Sneath P.H.A., 1976. Phenetic taxonomy at the species
level and above. Taxon, 25: 437-450.
Soria-Carrasco V., Gompert Z., Comeault A.A., Farkas
T.E., Parchman T.L., Johnston J.S., Buerkle C.A.,
Feder J.L., Bast J., Schwander T., Egan S.P., Crespi
B.J. & Nosil P. ,2014. Stick insect genomes reveal
natural selection’s role in parallel speciation. Science,
344: 738-742.
Spolsky C. & Uzzell T., 1986. Evolutionary history of
the hybridogenetic hybrid frog Rana esculenta as
deduced from mtDNA analyses. Molecular Biology
and Evolution, 3: 44-56.
Stahls G. & Savolainen E., 2008. MtDNA COI barcodes
reveal cryptic diversity in the Baetis vernus group
(Ephemeroptera, Baetidae). Molecular Phylogenetics
and Evolution, 46: 82-87.
Stork N.E., 1988. Insect diversity: facts, fiction and
speculation. Biological Journal of the Linnean
Society, 35: 321-337.
Strutzenberger P., Brehm G. & Fiedler K., 2011. DNA
barcoding-based species delimitation increases
species count of Eois (Geometridae) moths in a well-
studied tropical mountain forest up to 50%. Insect
Science, 18: 349-362.
Stur E. & Borkent A., 2014. When DNA barcoding and
morphology mesh: Ceratopogonidae diversity in
138
Alessandro Minelli
Finnmark, Norway. ZooKeys, 463: 95-131.
Templeton A.R., 1989. The meaning of species and
speciation: a genetic perspective. In: D. Otte, J.A.
Endler, Editors, Speciation and its consequences.
Sinauer, Sunderland: 3-27.
Van Valen L., 1976. Ecological species, multispecies, and
oaks. Taxon, 25: 233-239.
Verheyen E., Salzburger W., Snoeks J. & Meyer A., 2003.
Origin of the superflock of cichlid fishes from Lake
Victoria, East Africa. Science, 300: 325-329.
Warwick S.I., Francis A. & Al-Shehbaz I. A., 2006.
Brassicaceae: species checklist. Plant Systematics
and Evolution, 259: 249-258.
Winterbottom R., Hanner R.H., Burridge M. & Zur M.,
2014. A cornucopia of cryptic species - a DNA
barcode analysis of the gobiid fish genus Trimma
(Percomorpha, Gobiiformes). ZooKeys, 381: 79-111.
Witt J., Threloff D.S., Doug L. & Hebert P.D.N., 2006.
DNA barcoding reveals extraordinary cryptic di-
versity in an amphipod genus: implications for desert
spring conservation. Molecular Ecology, 15: 3073-
3082.
Woodcock T.S., Boyle E.E., Roughley R.E., Kevan P.G.,
Labbee R.N., Smith A.B.T., Goulet H., Steinke D. &
Adamowicz S.J., 2013. The diversity and biogeo-
graphy of the Coleoptera of Churchill: insights from
DNA barcoding. BMC Ecology, 13: 40.
Zachos F.E., Apollonio M., Barmann E.V., Festa-
Bianchet M., Gohlich U., Habel J.C., Haring E.,
Kruckenhauser L., Lovari S., McDevitt A.D., Pertoldi
C., Rossner G.E., Sanchez- Villagra M.R., Scandura
M. & Suchentrunk F., 2013. Species inflation and
taxonomic artefacts. A critical comment on recent
trends in mammalian classification. Mammalian
Biology, 78: 1-6.
Zaldivar-Riveron A., Martinez J.J., Ceccarelli F.S., De
Jesus-Bonilla V.S., Rodriguez-Perez A.C., Resendiz-
Flores A. & Smith M.A., 2010. DNA barcoding a
highly diverse group of parasitoid wasps (Braconidae:
Doryctinae) from a Mexican nature reserve. Mito-
chondrial DNA, 21 (SI): 18-23.
Zhang Z.-Q., 2011. Accelerating biodiversity descrip-
tions and transforming taxonomic publishing: the
first decade ofZootaxa. Zootaxa, 2896: 1-7.
Zhang Z.-Q., Christenhusz M.J.M., Esser H.-J., Chase
M.W., Vorontsova M.S., Lindon H., Monro A. &
Lumbsch H.T., 2014. The making of world’s largest
journal in systematic botany. Phytotaxa, 191:
001-009.
Zhou M. & Fuller R.C., 2014. Reproductive isolation
between two darter species is enhanced and asymme-
tric in sympatry. Journal of Fish Biology, 84: 1389—
1400.
Zhou X., Robinson J.L., Geraci C.J., Parker C.R., Flint
Jr. O.S., Etnier D.A., Ruiter D., DeWalt R.E., Jacobus
L.M. & Hebert P.D.N., 2011. Accelerated construc-
tion of a regional DNA-barcode reference library:
caddisflies (Trichoptera) in the Great Smoky Moun-
tains National Park. Journal of the North American
Benthological Society, 30: 131-162.
Biodiversity Journal, 2015, 6 (1): 139-146
Monograph
The town as a concentrated source of redaimable water and
materials. Opportunities for an engineered conservation
strategy
Salvatore Nicosia
Universita degli Studi di Palermo, DICAM - Dipartimento di Ingegneria Civile, Ambientale, Aerospaziale e dei Materiali, Viale
delle Scienze Ed. 8, 90128 Palermo, Italy; e-mail: salvatore.nicosia@ unipa.it
ABSTRACT A fierce theoretical debate is ongoing about the human species’ existence itself being sustain-
able for Earth and for living world. In the meanwhile cities, which are considered to concen-
trate the mankind's ecological footprints, are steadily growing and gathering huge populations
worldwide. This paper assumes that margins do exist to relieve man’s burden on Nature to
some extent, and that, regardless of our general concept of the matter, these margins should
be exploited. The focus of this note is on beneficial use of waste water and waste to spare
new resources and to create filter areas close to towns or belts around them. A brief reference
is made to some official declarations and indices published on biodiversity in anthropic
environments, such as the one from UNEP.
KEY WORDS Anthropic environments; Biodiversity; City planning; Resources; Urban Ecology.
Received 28.12.2014; accepted 30.01.2015; printed 30.03.2015
Proceedings of the 2nd International Congress “Speciation and Taxonomy”, M ay 1 6th- 1 8th 2014, Cefalu-Castelbuono (Italy)
INTRODUCTION
Exactly forty years have elapsed since the issue
of the ever most popular synthesis of criticism to
unrestrained use of resources (Meadows et al.,
1974). Pressure and unbalancing actions on natural
cycles have not relented yet; and much of them are
debited to cities, the exemplar man-made environ-
ment. It is documented that actually cities are
steadily growing worldwide (WHO, 2014). The
urban population in 2014 accounted for 54% of the
total global population, up from 3 4% in 1 9 60; w ith
all the related needs for areas, for every kind of
supplies and for waste management.
At least relieving measures are due and urgent,
and some of them were envisaged as early as 1971
by Odum E.P. (1969) and by Odum H.T. (1983). It
is still a matter of debate whether a urban ecology
can exist, and, if it can, whether it obeys to the
general laws of ecology or to its own special rules.
Anyway, in a pragmatic approach at least some
indicators and indices ought to be agreed, in order
to give a transparent measure of the environmental
benefits achieved trough certain actions.
CITIES, MANKIND AND RESOURCES
Historically, villages, towns and cities have been
made by men for themselves
- to develop broader and more free exchanges of
goods, manpower and skills
- to find customers for technological artefacts
- to benefit of more qualified services; of medical
care; of higher education
- to build up wealth
- to feel safer, etc .
140
Salvatore Nicosia
Cities obviously use land that often formerly
belonged to some other species. The ratio (covered
area/people living in) can actually be lower in
towns made of tall buildings than in sprawled ones;
indeed, this was the concept of Le Corbusier and
other architecture M asters. Denser tow ns, how ever,
have less possibilities of growing orchards, veget-
able gardens, firewood lands, fisheries, etc.; so they
need to fetch resources from far, unless flat roofs
are used to this purpose. The two arrangements can
obviously coexist in different quarters of the same
town (Fig. 1 ).
Cities necessarily draw most of the resources
they need from far; freshwater first. Figure 2 shows
the orders of magnitude of the materials and energy
exchanges of a middle capital town peopled by
about 600 000, per annum.
Is hardly possible that the used resources can be
given back to their sources or original places or
states. Restitution is usually not feasible because,
for instance: water is drawn from higher elevation
sources and discharged into lower water bodies;
foodstuffs are partly simply eaten, partly discarded;
energy is downgraded in its use.
DOES URBAN CIVILIZATION NECES-
SARILY THREATEN BIO DIVERSITY?
COULD IT BE HELPFUL IN SOMEWAY?
It is likely that diversity decreases where and
when
- a portion of a vital primary resource, e.g., high
quality water, is diverted;
- great amounts of secondary resources such as
wastewater and organic matter, although treated,
are discharged into limited seawater volumes;
- nuisances like warming; lighting; noise; men’s
stamping on, or walking through; traffic; navig-
ation etc. advantage opportunistic species like
rats, wasps, seagulls, ravens, magpies and drive
away the most sensitive ones.
Some blames to towns, however, appear still
ill-founded. As for now it looks more an article of
faith than a demonstrated and explained fact, that
at sea the outfall of a constant - discharge sewer
after treatment; or a storm sewer;would generate
more harmful gradients of salinity and turbidity,
than a river with its natural alternation of dry-
weather flows and flood flows.
Some positive effects that a well intentioned
town can develop towards wildlife are:
- Storing fresh water and, possibly, treated wastewa-
ter, smoothening floods/droughts
- Mitigating climate through managed green areas,
leading to less ecological stress
- Providing shelter to timid species
- Pouring organic matter in oligo-trophic biotopes,
resulting in an enhanced food pyramid.
The debate on urban civilization against bio-
diversity urges us to define - in a really scientific
and consistent way - what the civilization could be
like; how much of resources it strictly ought to
require/use; and how bio-diversity is to be quantified
(©suitable Indicators and Indices).
The concept of integrating town and nature
through filter ecosystems was enunciated and
gradually better defined in the Seventies by Odum
Figure 1. A historical walled town (left) with its circular gradient of buildings, green belt and surrounding
countryside; a modern town (right) with regulated quarters planned for different functions.
The town as a source of reclaimable water and materials. Opportunities for an engineered conservation strategy 1 4 1
Waste: 300 000 t
Freshwater: 60
million cubic meters
Foodstuff,
diverse materials
Energy: 720 000
MWhe + 54 10*
meters methane (=
600 000 MWht)
Low-temperature
Energy
Waste water: 50 million
cubic meters w ith 2 600 1
nitrogen and 440 1
compounds
Figure 2. The city of Palermo as a case-study: some major flows of materials and energy (civil uses only).
Figure 3. Left: The reservoir Piana degli Albanesi (610 m above sea level) is a multi-purpose water source for uses
in the plain of Palermo. Right: Example of heat losses in old-fashioned, low-tech town heating systems.
Figure 4. Plume of suspended solids at the mouth of a river (left) and flowing out from an on-shore outfall (right).
142
Salvatore Nicosia
E.P. (1969) and by Odum H.T. (1983). Naveh
(1982) used the term techno-ecosystem to represent
systems where technology and ecology are associ-
ated. These are the realm of ecological engineering.
Figure 5 suggests a possible application of these
concepts.
THE ACTUAL ORDERS OF MAGNITUDE
A medium-large town can provide, to itself and
to its surroundings, treated wastewater enough to
turn a squalid channel into a 2 m 3 /s steady flow
watercourse; or 100 hectares brownfield into a
wetland. This benefit is not entirely free, since 1 m 3
treated wastewater contains the embedded energy
of about 0.4 kWh; but most of this amount should
have been expended anyway, just to meet the
quality requirements at the discharge point.
The same town can also provide to itself and to
its surroundings 40 000 t compost: enough for 2 -f-
4 000 hectares soil being annually amended. What
to do with such engineered ecosystems is to the
environmental biologists’ expertise.
Two vast chapters apart are those of green roofs
and of underwater barriers laid for aquatic fauna
breeding and growing.
Green roofs can control urban climate; reduce
and smoothen water runoff; give shelter and ecolo-
gical corridors to animals and spontaneous plants,
and more; provided that mechanical energy (usually
drawn as electrical, actually) is supplied to lift
stored rainwater from the underground reservoirs.
Underwater barriers have been experienced, invest-
igated and discussed too much for requiring treat-
ment here .
RESHAPING TOWNS AND SETTING
THEM TO WORK FOR NATURE. INDICAT-
ORS AND INDICES OF ACHIEVEMENTS
Since urbanization is fundamentally changing
the nature of our planet, preserving biodiversity on
this new urban world requires going well beyond
the traditional conservation approaches of protect-
ing and restoring what we think of as “natural eco-
systems,” and trying to infuse or mimic such
elements in the design of urban spaces.
After two official sources: CBD - the UN’s Con-
vention on Biological Diversity; and the Working
Group CBO - Cities and Biodiversity Outlook; “...
unprecedented opportunities lie ahead in making
urban expansion greener. Cities have a large
Waste recycle
facilities
Water and
waste-water
treatment
plants
Groundwater
recharge areas
Playgrounds, parks
civic amenity sites
Civic allotments for
vegetables and
fruit-trees growing
Wetlands and vast
parks, wildlife-
oriented
Figure 5. A urban area featuring an inner and an outer ecological filter areas, plus one filter belt in between.
The town as a source of reclaimable water and materials. Opportunities for an engineered conservation strategy 1 4 3
potential to generate innovations and governance
tools and therefore can, and must, take the lead in
sustainable development. Many of the opportunities
can be found in nature based solutions, using ecosys-
tems in novel ways to address some of the most pres-
sing challenges, such as climate change, water and
food security, and poverty relieving. The way forward
involves reimagining cities as places of biodiversity,
and as sources for unique valuable services, rather
than only sinks that mark large ecological footprints .”
After CB O (based at S toe kholm U niversity, SE),
rich biodiversity can exist in cities; but it cannot be
taken for granted that it will be the same as before
urbanization. Habitat conversion often leads to the
loss of “sensitive” species dependent on larger,
more natural clusters of habitat for survival.
Cities already represent in themselves a new
class of ecosystems shaped by the dynamic interac-
tions between ecological and social systems. There
is a suite of “cosmopolitan” species, skilled gener-
alists that are present in most cities around the
world. The net result is sometimes termed “biotic
homogenization.”
It is still a matter of debate whether a urban eco-
logy can exist, and, if it can, whether it obeys to
the general laws of ecology or to its own special
rules. Anyway, in a pragmatic approach at least
some indicators and indices ought to be agreed, in
order to give a transparent measure of the environ-
mental benefits achieved trough certain actions.
Among the indicators of diversity we will cite
here the Singapore Index (SI), 2008.
This is a self-assessment tool for cities to
benchmark and monitor the progress of their biod-
iversity conservation efforts against their own
individual baselines.
It comprises:
a) the “Profile of the City”, which provides com-
prehensive background information on the city;
b) 23 indicators based on the guidelines and
methodology provided.
The scoring of the Index is quantitative in nature;
a maximum score of 4 has been allocated to each
indicator, and with the current count of 23 indicat-
ors, the total possible score of the Index is 92 points.
The year in which a city first undertakes this
scoring program will be taken as the baseline year.
The future applications of the index will be meas-
ured against the baseline to chart its progress in
conserving biodiversity.
For 7 of the indicators, a statistical treatment
will be applied to sample data sets coming from
Figure 6. A possible ecological succession for derelicted land turned into wetland.
144
Salvatore Nicosia
Figure 7. Left to right, clockwise: windrow composting; compost handling; plant germination.
several cities, to ensure the scoring ranges estab-
lished are unbiased and fair to a broad spectrum of
cities of different characteristics, over a wide
geographical range.
URBAN ECOLOGY; GOVERNING BIOD-
IVERSITY IN CITIES: A NOBLE COM-
MITMENT OR A PURE DREAM?
We have now to look at the frame within which
the actions for a sustainable town are developed, in
order to judge about the theoretical substantiation
of them and to forecast how far the pragmatic
approach outlined above can arrive.
Among the optimistic sources we are quoting
here a statement from CBO: “ There is a need far
redefining the role of cities so that they increasingly
provide stewardship of marine, terrestrial and
freshwater ecosystems elsewhere. Developing the
concept of nature based - solutions entails exploring
a deeper dimension of how attributes of ecosystems,
such as diversity, modularity and redundancy may
be interpreted, applied and used ’ .
Another affirmative statement comes from Jari
Niemela (1999): “ The question arises whether a
distinct theory of urban ecology is needed for
understanding ecological patterns and processes in
the urban setting. The answer is no; however, due
to the intense human presence approaches that
include the human aspect are useful in studying
urban systems ” .
Collins et al. (2000) raise serious doubts and
develop a strong criticism of these perspectives. For
these Authors, in studying urban systems the intense
human presence certainly obliges to approaches that
include the human aspect; still, even such tentative
integrated approaches could reveal themselves a
dead way. Quoting the Collins’ words: “ From the
perspective of a fieldecologist examining a natural
ecosystem, people are an exogenous, perturbing
force. Human beings - and especially their cities,
seemingly so "artificial"- fail to fit neatly into eco-
logical theory. People mobilize some nutrients and
deplete others, produce pollutants, drive species
extinct, promote the survival of others, change the
composition of the atmosphere and alter land-
scapes. In cities people create habitats that never
before existed, divert water, increase temperatures
and, by intent or by accident, manipulate the
communities of other species found within city
boundaries and beyond (...)”.
Still after Collins and coworkers, “ We lack a
method of modelling ecosystems that effectively
incorporates human activity and behaviour. And the
processes and dynamics within cities largely elude
The town as a source of reclaimable water and materials. Opportunities for an engineered conservation strategy 145
**
COMPONENTS
INDICATORS
VARIABLES
SCORE
MAXIMUM
■ L\D.: ■ Diversity of ecosystems
'INI'.! t Diversity of
1ND.I » Diversity of ecosystems
ecosystem;
BASIS OF SCORING
irvD.:i
Diversity of
ecosystems
rationale for selec tion of
INDICATOR
HOW TO CALC IX ATT
A) Eased on the estimation that
INDICATOR
realisticallv. nycttycan accommodation
The number of natural ecosystems found in
to about 10 natural ecosystems, within us
a cm 1 saves an ladic anon of the (favors
Number of na tural
boundaries the scoriae would be
MAXTMIM
ran re of niches for native flora and fauna
ecosystems found in the citv
SCORE
Since different ecomtem; are found in
0 point * 0 natural ecosystem
l Native
different geographical regions any
1 point * 1-5 ecosystems
Biodiversity in
sciennikallv acceptable tertetmil and
WHERE TO GE T DATA
2 points - -L6 ecosystems
4
the City
tninne ecosystems including fosests
(tropical subtropical monsoon temperate
FOR C ALC ULATIONS
a points - T -9 ecosystems
4 points * 10 and more ecosystems
lowland montane, primary secondary.
Possible sources of data on
lTND.:i
etc ). mangroves freshwater swamp; peat
sutural areas include
swamp; natural grasslands mers steam:
government aeenr.es m
B) Baseline of 100
lake;, rocky shores, beach, mud-flats, sand
charge of biodiveraty, cits-
dunes sea zrsss beds, corals, etc . can be
municipalities, urban
C] Traffic hoe ssstem of increase, neutral
computed in die calculation of this index
p! arming agencies
biodiversity ceitres. nature
and decrease
groups. uniseisittes.
publications. etc
Table 1. The Singapore Index (SI): example of the working tables: a local ecosystems inventory.
producers; 1st
order consumers
producers; 1st
order consumers
Figure 8. Three types of heterotrophic system (the one upper left in every picture) and their relationships with the
surroundings. Tapping of fossil fuels to feed mechanical farming makes the main difference between the three.
146
Salvatore Nicosia
an understanding based on traditional ecological
theories.
For most [natural] ecosystems the overall
calculation is fairly well balanced between inputs
and outputs. Urban energy budgets [instead],
dominated as they are by deliberate human energy
imports and by losses via fossil-fuel burning, do not
resemble the energy budgets of any other ecosystem
on earth ” .
Figure 8 is an attempt to depict this concept.
CONCLUSIONS
Much of the blame put on town ought actually
to be put on the human way of life.
We believe that man, as a second-level or vertex
consumer, is by no means the only species on Earth
whose life is heterotrophic, or that lives in crowded
com m unities.
It is true that his weaknesses (like the need for
shelter and warmth and the inability to nourish
himself of raw food) and his strengths (such as his
unique ability to handle fire, and to make, build up
and transfer knowledge, etc.) are peculiar. All this
increases the singularities of mankind, but in our
opinion does not entail any definition of supposed
peculiar human ecological niches.
The only fundamental difference that we see
stays in that, that men are not innocent in their be-
haviour, and ecological feedbacks to their actions
are usually overbalanced by their obstinacy. Man-
kind usually neglects or denies the biosphere’s
response to its actions, and, if compelled, is more
willing to force further than to ease. The renounce
to such unjustified self-exemption from taking feed-
backs into account; and a consistent commitment in
respecting and saving the other species’ lives and
spaces, even in towns, at least as a mitigation meas-
ure, is the tribute that mankind still owes to Nature.
The Author’s speech was dedicated to the luminous
memory of Giovanni Falcone and Francesca
Morvillo Falcone, and of the men of their escort.
Fallen at Capaci, Palermo, May the 23rd, 1992.
REFERENCES
CBO - Cities and Biodiversity Outlook - Stockholm
Resilience Centre. Stockholm University (SE).
www.cbobook.org.
Collins J.P., Kinzig A., Grimm N.B., Fagan W.F., Hope
D.,Wu J. & BorerE.T., 2000. A New Urban Ecology
- Modeling human communities as integral parts of
ecosystems poses special problems for the develop-
ment and testing of ecological theory. American
Scientist, September- October, Volume 88, Number 5.
Meadows D.H., Meadows D.L., Randers J. & Behrens
W.W.III, 1974. The Limits to Growth: A Report for
the Club of Rome's Project on the Predicament of
Mankind. Potomac Associates, Virginia (U.S.A.).
Naveh Z., 1982. Landscape ecology as an emerging
branch ofhuman ecosystem science. In: Advances in
Ecological Research 12. Academic Press, London,
pages 189-237. See especially Section D.
Niemela J., 1999. Texts posted in the website ofUniver-
sity of He Is in ki (SF) - D epartm ent of E nvironm ental
Sciences (www. helsinki.fi/urbanecologyresearch/
m ember s/niem ela.htm )
Odum E.P., 1969. The Strategy of Ecosystem Develop-
ment. An understanding of ecological succession
provides a basis for resolving man's conflict with
nature. Science, 164, 3877: 262-270.
Odum H.T., 1 983. Systems ecology; An introduction.
John Wiley and Sons, New York.
WHO-OMS Global Health Observatory www. who.
int/gho/urban_health/situation_trends/urban_
p op u la tion_gro w th_ text/e n.
World Resources Institute, World Conservation Union,
and United Nations Environment Programme -
"Global Biodiversity Strategy", Rio de Janeiro
Earth Summit 1992. http://www.cbd.int/authorities/
getting in volved/cbi.shtm 1.
Biodiversity Journal, 2015, 6 (1): 147-160
Monograph
Pest management of citrus fruits in Sicily (Italy) through in-
terventions of biological control. The example of the biofact-
ory of Ramacca, Catania
Giuseppe Greco
Ente di Sviluppo Agricolo della Regione Siciliana (E.S.A.). via Liberta 203, 90 1 43 Palermo, Italy
ABSTRACT Since 2007, in Sicily, plant health protection against citrus mealybugs is taking place through
the Biofactory of Ramacca, in the Plain of Catania, a property of the Institute for Agricultural
Development of the Sicilian Region (i.e. Ente per lo Sviluppo Agricolo, E.S.A.). The
Biofactory is unique being aimed to produce industrial quantities of auxiliary insects and is
a center of European interest because it is fully organized to provide means ofbiological fight
imposed by the Directive 1 28/2009/EC, which requires, from 1 January 2014, farms to
comply with the application of general principles of integrated pest management. In this paper
we examine structural features of the Biofactory, breeding techniques empoyed and results
obtained in the period 2007-201 3, which allowed many companies, from 200 to 360 ( i.e.
20% -35% of the regional surface operating in organic citrus production) to be able to employ
biological weapons against pest insects. We analyze dynamics and results of production
deriving from the approval and adoption, by the owner (E.S.A.), of a new "discipline" that
governs the assignment of insects to farmers at a very low price to balance E.S.A.'s purposes,
which is both to ensure adequate performance in order to pursue institutional support to
agriculture and, considering the Insitute’s economic nature, to partially cover the production
costs incurred to ensure the service. The continuity of the project is assured by the ongoing
program for the period 20 1 3-2020 with an enlargement of the array of entomological
production aimed at intercepting the needs of new productions (i.e. greenhouse horticulture,
vines, ornamental and fruit trees).
KEY WORDS pest management; biological control; Biofactory; Ramacca; Sicily.
Received 05.07.20 1 4; accepted 12.10.2014; printed 30.03.2015
Proceedings of the 2nd Interna tional Congress “Speciation and Taxonomy”, May 16th-18th 2 0 14, Cefalu-Castelbuono (Italy)
INTRODUCTION
A biofactory (or commercial insectary) is a
structure in which takes place the breeding of
arthropods on an industrial scale, aimed at the
production of living organisms to be released in
large amounts into the environment in the context
of techniques of biological control and integrated
pest management. On the contrary, the insectary is
a breeding realized for scientific purposes.
The multinationals of chemistry have never seen
welcome the birth of b io fa c to r ie s , because the
organic product stands as alternative to the use of
pesticides (Tremblay, 1988; Pollini et al., 1988;
Goidanich et al., 1990; Flint, 1991; Grafton-
C ard w ell & Reagan, 1995; Pollini, 1998;Ferrariet
al., 2000, 2006; M asutti & Zangheri, 200 1; Muc-
cinelli, 2006; Penny & Cranston, 2006).
148
Giuseppe Greco
There are reports of a first biofactory already in
1916 in Santa Paula, California, the “Lim oneira
Company”. In 1931, there were 16 and produced
especially insects antagonist to citrus mealybugs
like the coc cine Hid ClJptolaemUS montrOUZieri that
is bred and successfully launched today.
In Northern Europe biofactories are used for
biological control in greenhouses: here the chemical
control had shown its serious limitations in the
effectiveness of and compatibility with healthy
products. In fact, the glass or plastic covers are an
insurmountable physical barrier for antagonists of
harmful species, warming accelerates the develop-
ment of both plants and pests, the collection of the
products can not be reconciled with respect to the
"waiting period" fixed by law between chemical
treatment and collection and, not least, greenhouses
turn out to be "gas chambers” for the farmers who
w ork therein .
A careless use of chemical products in agricul-
ture with the aim to maximize the production has
led over the years to a number of disorders that have
resulted in considerable damage to the environment
and to humans. Many chemicals have been banned
and the defense of the plants has been oriented to
the use of alternative methods equally effective and
safeguarding the ecosystems (De Bach etal., 1969;
Viggiani, 1 977; Chiri, 1 987; Walde et al., 1 9 89;
Celli et al., 1991; Hoffmann & Frodshan, 1993;
Luck et al., 1996; Murdoch et al., 1996; Ferrari, et
al., 2000; Vacante & Benuzzi, 2004; So r ribas et al.,
2008, 2010; Tena & Garcia-M ari, 2011).
Figure 1. Biofactory of Ramacca, Catania, Italy, Institute for
Agricultural Development of the Sicilian Region (E.S.A.).
The first biofactories in Europe born in England
and Holland around 1960 and, since then, have
always grown both in number and in quantity of
species bred and used. Today in Europe there are 26
biofactories with more than 30 species raised and
excellent qualitative-quantitative standards.
In Italy there are only two biofactories: the
first (in order of construction) is in Cesena
(1 987/90) while the second is in Sicily, in the Plain
of Catania (Figs. 1, 2) in the territory of Ramacca
(200 1 /03) (Greco 2014a, b). The latter is mainly
distinguished by the quality and quantity of its
products supplied aiming more at the diffusion of
breeding techniques rather than for commercial
purposes. Both biofactories serve an agricultural
area which is considerably increasing in size, and
achieve agricultural productions with the least
possible impact on the territory, sustainable for the
planet, whereas in other parts of the world, biolo-
gical control has totally replaced chemical poisons.
Yet here, in the Mediterranean, people are not
deeply aw are of the benefits of this resource and the
many solutions it offers, but the products of a
biofactory are going to become even more relevant
in the light of Directive 128/2 009 / EC establishing
a framework for Community action to achieve a
sustainable use of pesticides.
This Directive was transposed into Italian law
by Legislative Decree 150 of 14 August 2012. Since
1 January 2014, professional users of p h y to s an itary
products (art. 19) should apply the general prin-
ciples of integrated pest management required
Figure 2. Biofactory of Ramacca, Catania, Italy:
biofactory corridor.
Pest management of citrus fruits in Sicily through interventions of biological control: the biofactory of Ramacca, Catania 1 4 9
among which is reported, as technical and funda-
mental element, the use of biological means of
struggle.
EXPERIENCE IN SICILY, AT THE CENTER
OF THE MEDITERRANEAN: THE BIOFACT-
ORY OF RAMACCA
In 1 996 the Sicilian Region has commissioned
the Institute for Agricultural Development (i.e.,
Ente per lo Sviluppo Agricolo, E.S.A.) to study the
possibility of implementing attive interventions of
biological control. From that date until today
E.S.A. carried out:
1) a preliminary plan for measures ofbiological
control of Ceratitis CQ.pitQ.tCl (Mediterranean fruit
fly) atregional scale, prepared in collaboration with
the FA O / IAEA Agriculture and Biotechnology
L aboratory.
2) the planning of a biofactory alternative to the
first one, to be built in Ramacca (Catania), aimed
at the production of 3 species of insects beneficial
to citrus cultivation {Aphytis melinus, Cryptolaemus
montrouzerii , Leptomastix dactylopii) and 1 insect
to be employed in horticulture. DiglipHuS iSQeQ
Walker, 1 8 3 8 (H ym enoptera: Eulophidae).
Actually, it was funded and implemented only
the second project in which the biofactory of
Ramacca is designed to be a flexible pole of pro-
duction of material (insects) to be used in agricul-
ture for most programs of biological or integrated
control. For its start-up phase of production, have
been considered, as reference, those crops that,
more than others, are susceptible to these kinds of
initiatives for their technical and economic charac-
teristics: citrus and protected horticulturals. There-
fore the biofarm has been designed and equipped
for the production of:
a) 3 insect species beneficial to biological
control programs for citrus cultivation (Aphytis
melinus , Criptolaemus montrouzerii , Leptomastix
dactylopii)-,
b) 1 insect used for integrated pest management
of vegetables and flowers grown under cover
( Diglyphus isaea).
The facto ry is located in the territory of
Ramacca (Catania), Margherito district, on a total
area of approximately 3.5 hectares that can be
potentially increased and improved in case of
changed conditions of the market.
The biofactory is composed of:
a) 1 shed of2,500 sqm (72 ml, 10.00 mix 34,30)
which houses cells in a controlled and conditioned
environm ent;
b) 6 greenhouses, each of 100 sqm ca. (10.00
ml x 10.00 ml), five of which are used for the
production of Diglyphus isaea and one for Lyri-
Omiza (guest of Diglyphus), this latter room is
placed at a safe distance to avoid contamination
between competitors since both species are raised
in purity.
The 6 greenhouses are heated, to prolong the
production season even in the coldest months
(January and February), and equipped with an
adequate irrigation system to allow the cultivation
of bean plants in pots placed on anti-algae cloths;
c) 1 office building of 350 sqm (ml 34.30 ml x
10.00 ml).
The shed is composed of 36 rooms including
cells, work rooms, service corridors, warehouse,
workshop, toilets and trasformer room, central
cooling and boiler. Cold storage and processing
rooms are 28, divided as follows:
9 for Aphytis melinus-,
6 fo r Criptolaemus montrouzerii
4 fo r Leptomastix dactylopii
9 in common for Criptolaemus montrouzerii
and Leptomastix dactylopii.
BREEDING TECHNIQUES OF INSECT
PRODUCTS IN BIOFACTORY
Aphytis melinus De Bach, 1959
H ym enoptera Aphelinidae
Aphytis melinUS (Figs. 3, 4) is a parasitoid of
Aonidiella aurantii Maskell, 1 8 79 (Rhynchota
Homoptera D ia sp id id ae ) , or California red scale, a
major pest of citrus, but it can also parasitize other
species such Diaspididae Aonidiella citrina
(Coquillett, 1891) and AspidiotUS nerii Bouche,
1 8 3 3 (Flanders, 1 953; De Bach & Argyriou, 1 967;
Abdelrahm an, 1974; Rosen & Eliraz, 1978; Rosen
et al., 1 979; Fuck et al., 1 982; Orphanides, 1 984;
Yu et al., 1986; Opp & Fuck, 1986; Reeve, 1987;
Yu & Fuck, 1 98 8; Rodrigo & Garcla-Marl, 1990,
150
Giuseppe Greco
Figure 3 . Aphytis melinus
(Photo by “Centrale O rto fru ttic o la of Cesena, Italy).
Figure 4. Climate cabinets with AspidiotUS nerii br e d on
pumpkins for developing of Aphytis tnclinUS.
1992; Hare & Luck, 1994; Heimpel & Rosenheim,
1995;Tumminellietal., 1996; Gottlieb etal., 1998;
Pekas et al., 2003; Pasotti et al., 2004; Rodrigo et
al., 2004; Pina, 2007; Pina T. & Verdu M.J., 2007;
Vacas et al., 2009; Vanaclocha et al., 2009).
Agricultural use of the insect: A. tfieliflUS is
launched at the adult stage and disperses easily in
all the citrus grove, possessing excellent research
skills. In citrus infected is good practice to make a
winter treatment with white oil at 2-2.5%; this
allow s to reduce, albeit only partially, the wintering
population of the cochineal. The parasitoid is
launched following a pattern that includes a series
of consecutive launches after the flightdetection of
cochineal males in late April-early May. When the
plan of biological control is set up, in the first year
are expected about 10-12 launches, 2/3 of which to
be carried out in A p ril-M ay - Ju n e until mid-July,
while the remaining 3 or 4 launches take place from
m id - S e p te m b e r to throughout October. In the
months of April, May and June, launches can be
made every two weeks, moving on to a weekly
frequency when temperatures increase. 8,000 to
1 2,000 parasitoids per hectare, for a total of 100 to
150,000 / ha for production season are launched. In
2-3 years the intensity of the pest is reduced so that
is possible to reduce proportionally the number of
lauches, limiting them exclusively to the spring-
summer period. It is very im portant to pay attention
to chemical treatments performed before and to
those that will take place.
Breeding techniques and production cycle in
biofactory: breeding of Aphytis IfielinUS is made in
climate cabinets, using the p arth e n o g e n e tic strain
of Aspidiotus nerii bred on pumpkins.
Pumpkins are kept in cells furnished with metal
shelves; the environment of the cells is adjusted so
as to have 13 ± 1 °C and 50 ± 5% RH; pumpkins
are previously washed and disinfected.
The production process has a duration of about
60 days, breeding is carried out in two areas: one
for the multiplication of the host and one for the
production of the parasitoid. Even the AspidiotUS
nerii (host) is reared in cells whose furniture is
made of metal shelving with lozenges. The nymphs
of AspidiotUS are then collected and placed in a jar
before inoculating other pumpkins. The environ-
m ental conditions for the breeding of AspidiotUS are
the following: temperature 26 ± 1 °C, RH 50% ± 5.
At the 4 5th day, before the spill of nymphs, 10%
of pumpkins are brought in the cells for develop-
ment of AspidiotUS for harvesting nymphs to be
used for the inoculation of pumpkins, whereas the
remaining 90% is iplaced in plastic bins for
the production of A. TUoUnUS. Pumpkins are put in
contact with A. 1716 Un US for 2 4 h.
The adults are taken after 24 h, blowing carbon
dioxide to saturation. After inoculation, pumpkins
can be placed in the two cells intended for the pro-
duction deirA. melinus, air-conditioned to 26 ± 1
°C and 50 ± 5% RH. After 10-15 days, A. melinus
newborn are collected after release of carbon dio-
xide. Insects fall to the bottom of the cabinets and
are put within cylinders where are measured volu-
metrically. Adults collected are packaged in trays
of 10,000 or 25,000 insects containing honey as
Pest management of citrus fruits in Sicily through interventions of biological control: the biofactory of Ramacca, Catania 1 5 1
Figure 5. Leptomastix dactylopii
Figure 6 . PlaUOCOCCUS dtri
(Photo by “Centrale O rto fru ttic o la of Cesena, Italy).
(Photo by “Centrale O rto fru ttic o la of Cesena, Italy).
feed. Packages can be stored for a few days in the
refrigerator ventilated at 15 °C. The production
ratio is 1 : 3 .
Leptomastix dactylopii Howard, 1 8 8 5
Hymenoptera Encyrtidae
Parasitoid (Fig. 5). Endophagous of PlatlOCOC-
CUS citri Risso, 1 8 1 3 (Rhynchota Homoptera
P s eu d o c o c c id ae) (Fig. 6) (Chandler et al., 1 9 80;
Tingle & Copland, 1 98 8, 1 989). The United States
are its country of origin and its cycle in nature takes
place on mealybugs, P. ficUS Signoret, 1 8 7 5, P. vitis
Ezzat et McConnell, 1963 and, in laboratory
conditions, spread also over other hosts.
Natural cycle and agricultural use o f the insect:
at 25 °C, and 75% humidity, the cycle of L. daC-
tylopH takes about 21 days. Adults, 12 hours after
the flicker, begin to mate. Females move on the
pseudococcid colony seeking - measuring them by
antennas - for the nymphs with appropriate shape
and age where to inject the eggs (one for each
victim). From each egg comes out a larva that, in
13 days, making three mutes and th rough four larval
stages, becomes pupa, at first light in colour, then
darker. After a week from 'pupation, the adult
flickers. Particularly remarkable it is that the larva
produces chitin and hardens the outer wall by an
aeroscopic plate from which it breathes atmospheric
oxygen. At the end of metamorphosis, by the
chewing apparatus severs an operculum placed in
anal position of the host and flickers. L. dactylopii
is an insect yellow honey with three simple eyes.
Its sizes range from 0.5 to 6 mm (11 an tenno meres).
Males have longer and silky antennae with 10
antennomeres, females shorter and hairless (11
antennom eres). L. dactylopii is marketed at the
adult stage and can be used on citrus fruits in com-
bination with Criptolaemus montrouzieri and on
ornamental plants infested by PlaUOCOCCUS dtri.
Breeding techniques and production cycle in
biofactory: the production cycle of L. dactylopii
takes place entirely in climate cabinets. The host is
P. dtri ( mealybugs) which is bred on potato sprouts
etiolated in areas separate from those of the para-
sitoid. For storage of potatoes are used cells condi-
tioned to 5 °C and 50 +/- 5% relative humidity.The
breeding cycle of the parasitoid lasts 9-10 weeks.
In the first stage, are produced etiolated shoots of
potato which, after 2-3 weeks, are infested with the
citrus mealybug. When nymphs are ready, L.
dactylopii is inoculated. After 20 days the adults
are collected with aspirators and packed in jars of
100 individuals. Insects can be stored at 15 ° C, if
well fed with appropriate diets.
Cryptolaemus montrouzieri m u is ant, 185 0
Coleoptera C occinellidae
Polyphagous predator (Hodek & Honek, 1 996.
Milan Vargas, 1999) thatcan live at the expense of
several P seu d oc o c c id s or even other insects (Figs.
7, 8). The adult measures about 5-6 mm has
black elytra, while the head, chest, abdomen and
152
Giuseppe Greco
Figure 7. Cryptolaemus montrouzieri
(Photo by “Centrale O rto fru ttic o la of Cesena, Italy)
extremities of the elytra are orange. At a constant
temperature of 25 °C females live about 60 days
and, during this time, lay 60 to 120 eggs.
Eggs are located close to the cottony ovisacs of
the prey so that and the young larva, just shelled,
can easily reach its preferred food: eggs and young
nymphs of the pest.
The Coccinellidae larva goes th rough four
stages before pupating (by attaching to a support)
after which it becomes an adult. It has a waxy
coating to camouflage itself onto the colonies of P.
CltVl, but cannot be mistaken for its larger size and
its mobility.The cycle from egg to adult lasts, at 25
°C, 35 days. It is an insect native to Australia and
therefore sensible to harsh winters; it has already
acclimatized in many areas of southern Italy and, in
the islands, winters as an adult.
Agricultural use of the insect: Cryptolaemus is
sold at the adult stage. On citrus fruit it is used in
association with Leptomastix daCtylopii e specially
in the hotbeds of infestation, which are out of
control of the parasitoid. In the field, it is employed
from June up to August (3 months). Cryptolaemus
could be used also on ornamental crops in green-
houses or in potted plants; on this item, it is devel-
oping an interesting market in northern Europe.
The production cycle of Cryptolaemus takes
place entirely in climate cabinets. The host is P.
citri ( mealybugs) which is bred in purity on
etiolated shoots of potato in a separate room. As
P. dtri is used as host also by LcptOIflClStix, room s
designated for P. dtvi production are used for both
insects.
Figure 8. Larvae of Cryptolaemus montruozierii on
potato sprouts infested by PlttnOCOCCUS citVl.
In particular, in a section of the biofactory, there
are cells for the storage of potatoes (at 13 °C and
60% RH); and in another section, cells for the
development of the tubers and, still, other cells for
the development of P. dtvi (at 25 °C and RH of 60
± 5%) that will serve to feed both the auxiliaries
( Leptomastix and Cryptolaemus ) .
In another area of the building there are cells
for development of P. dtri, cells for collection of
Cryptolaemus and processing room s. The breeding
cycle of predator lasts 10-13 weeks. In the first
ph ase P. dtri is bred in purity on etiolated sprouts
of potato. In breeding cells, potatoes are made
germinate in the dark for 2-3 weeks; the shoots are
infested with P. dtri and the infestation is let to
develop for 3-4 weeks; finally there is the inoculum
with Cryptolaemus. Adults, collected after 35 days
with vacuum cleaners, are packaged in cans from
100 to 200 units. They are then counted volumet-
ric ally . Insects can be s to red at 15 ° C, even up to a
month if well fed with an appropriate diet.
MANAGEMENT BIOFACTORY
In 2006, the managing of the biofactory of
Ramacca began with the finding of the head-
breeding strains (Aphytis melinus, Cryptolaemus
montrouzieri Leptomastix dactylopii and Diglyphus
ISaea ) and of interm ediate entomological m ate rials
(. Aspidiotus , Planococcus , Liriomyza , etc.) of which
such insects are parasitoids or predators. As
planned, entomological breeding aimed, from the
Pest management of citrus fruits in Sicily through interventions of biological control: the biofactory of Ramacca, Catania 1 5 3
beginning, at the production of Aphytis lUelUlUS,
Leptomastix dactylopii and Cryptolaemus mon-
tWUZien. At first it was even started a production
of DiglyphliS isciea (greenhouse parasitoid on Livi-
omyza trifolii , L. bryoniae and L. huidohernsis ) then
abandoned because of the uneconomic production
cycle.
Until 2011 the entomological material was
distributed free to farmers th rough per ip heral com-
panies belonging to E.S.A. (i.e. SOPAT, Offices for
the A ntiparasitic Fight) and to the Department of
Agriculture and Forestry (SOAT, OMP).
The criteria developed by the Administrative
Department of biofac to ry included a distribution of
the product to farmers cultivating citrus, to organic
or converting to organic farms, and to farms that
apply and implement criteria of integrated pest man-
agement, according to a programming technique
agreed with local Institutes that provide agricultural
technical assistance (ESA, SOAT and the Office of
Agriculture and Forestry).
The reaching of full production was expected by
the third year (29 March 2009), during which it has
been programmed the full activity of the building
with the following annual production levels:
Aphytis melinus 6 7,200,000 individuals;
Cryptolaemus montrouzieri 350,000 individuals;
Leptomastix dactylopii 1,0 0 0,0 0 0 individuals;
Diglyphus isaea 1,9 0 0,0 0 0 individuals.
Data management in the period 2006-2011
During the period 2006-20 1 1 (Fig. 9), insects
have been distributed free to regional farms and
other applicants who had a purpose in the public in-
terest, including regional and national Universities,
Trends observed in citrus treated with insects of the Biofactory of Ramacca in 2006-2011
4.000
3500
t 3.000
| 2 500
M 2.000
1.500
1.000
500
orange groves lemon groves mandarin groves clementines groves
c
EH
□ 2006
R 2007
□ 2008
□ 2009
■ 2010
□ 2011
Number of farms, by size classes, that used
Leptomastix dactylopii in 201 1
y
E
s_
£
o
o
80
60
40
20
0<x<5 5 <x<10 >10
Size classes (in hectares)
Number of launches of Leptomastix dactylopii car lied
out by the companies served by the biofactory,
grouped by size classes. Data processing 2(11 1
No, of
launches
Size classes (in hectares)
Figure 9. Data management in the period 20 0 6- 201 1 (Source E.S.A.).
154
Giuseppe Greco
Regional Departments, Development Services,
Institutes or Development Agencies of other Italian
regions.
M axim um productions were distributed in 20 10-
2011, mostly to citrus farms, for a total of more than
4,300 hectares distributed in 325-355 entities.
Noteworthy, as for the 20 10-20 1 1 data, there is a
significant increase in production ( + 50% compared
to 2010), correlated with a stabilization of the
“protected” area, amounting to 4,361 hectares (-28
hectares compared to 2010); the maintenance of the
substantial number of seasonal launches can be
explained by a kind of loyalty of the users who, in
manifesting an appreciable degree of satisfaction,
show confidence in using alternative means of
organic production.
Data management in the period 2012-2013
In 2011, it was suggested to apply a reduced
price to Sicilian farms. This is to contribute to the
costs of production that, every year, E.S.A. supports
to ensure its performance. So it was approved and
put into effect a new "Discipline" which regulates
the sale of insects to farmers at a "price of contri-
bution", in order to proceed, gradually, to com-
pensate production costs. The "price of contribu-
tion", which ranks, by definition, below the values
of the free market, reconciles the needs of the
Istitute, which has to ensure adequate performance
in providing institutional support to agriculture,
with its financial nature aimed to partially cover the
costs of production. This regulation does not
exclude the transfer of beneficial insects also in
favor of other subjects, in different places (extra-
regional) and, possibly, for different purposes
(agricultural as well as commercial or public). In
this case, the above mentioned constraints do not
apply, so thatE.S.A. can set the products at different
prices (to be considered net of shipping), commen-
surate with market values.
Application of new "Rules" recorded a drop in
distribution in 201 2-201 3, and, during a period of
6 years of free distribution, it obviously resulted in
a big change of the demand of the three species. A
first effect of the new regime can be seen in the
production levels of 20 12-2013. In particular (Fig.
io), the amount of Aphytis melinus, Leptomastix
dactylopii and Cryptolaemus montrouzieri -
although often reached high profiles above those of
feasibility - stood at levels significantly lower than
those of2011,i.e. 139-149 million, 672-1766 thou-
sand and 233-277 thousand individuals, respect-
ively. In 2012, production reached 119% of what
expected in steady-state c o n d itio n s ,(i.e . +19% ).
Briefly, these results can be explained with a
production trend that had to take into ac c o u n t u ser s ’
requests, which resulted in a change of strategies
and productive quality (when possible) that af-
fected, for example, the extent and availability of
traditional raw materials to be acquired (potatoes,
Observed production of Aphytis melinus compared
with expected production
250.000 000
£ 200.000.000
-5 150.000.000
—♦—observed production
t 100000.000
p ex per Ini prodiiclUm
a
Ji 4 1
50.000.000
2006 2007 2000 2009 2010 2011
sears
Observed production of Cryptolaemus montrouzieri
compared with expected production
oWnnl prwbHrllwi
rXprflrd JiiothirUnr
years
Observed production of Leptomastix dactylopii
compared with expected production
6 . 000.000
w 5.000 000
S 4,000.000
B
o 3.000.000
* 2.000 000
1 . 000.000
2006 2007 2008 2009 2010 2Qn
years
■ observed prodnri ion
expected production
Figure 10. Development of production of Aphytis melinUS ,
Leptomastix dactylopii and Cryptolaemus montrouzieri than
expected feasibility (Source E.S.A.).
Pest management of citrus fruits in Sicily through interventions of biological control: the biofactory of Ramacca, Catania 1 5 5
var. “Spunta” and “Desiree” and pumpkins var.
“ B u ttern u t”). Another cause is to be found in distri-
bution fees, which were fixed in the absence of
solid experience of huge productions and, therefore,
of necessary and useful market information. F in ally,
the price of each insect certainly influenced the
users’ choise. For example the price / effectiveness
or cost/utility ratio for Aphytis melinUS was con-
sidered, by the regional users, more convenient
than those fixed for CryptolaemUS montWUZieri and
Leptomastix dactylopii.
Profile of user companies in 2012-2013
Quantitative aspects of each entomological
entity distributed to regional farms are of course
also reflected on land statistics. In fact, Users (i.e.
farms), primarily engaged in citrus cultivation, were
more than 200 (213 to 298), for an area of at least
2,300 Ha. Just to quantify, 2,152 Ha of orange
groves, 313 ha of lemon groves, 91 Ha of m andarin
groves and 5 1 of clem entine groves took advantage
from the service provided by E.S.A.
The new payment system had a negative impact
not only on the lemmon groves of Syracuse: also
other citrus groves suffered a regression of land
extensions which reached its peak in the areas
planted with orange trees. It also follows, that the
biological defense against the citrus mealybug, (P
citri) and red scale (A. aurantu ) by Aphytis melinus,
Cryptolaemus montrouzieri and Leptomastix dac-
tylopii , decreased to 2, 4 41, 1,0 27 and 540 Has, re-
spectively, Siracusa and Catania remain the
provinces where biological fight is mainly per-
formed, followed by an increasing number of far ms
in Agrigento province. Hence it is indirectly con-
firmed that the location of the Bio factory (Ramacca,
Catania) is in line with the geographical distribution
of its real users.
The profile of the more than 298 farms that, in
2012, took advantage of the service of the Biofact-
ory of Ramacca is best represented in figure 11.
Companies that follow programs of integrated
biological defense or integrated fight in citrus and
benefit of the insects provied by the biofactory have
predominantly a size less than 5 hectares (161, 48
and 27 farms can be listed for A. melinUS , L. daC-
tylopii an d C. montrouzieri, respectively). M edium-
sized companies were those that, in 2012,
performed more seasonal launches of Aphytis
melinUS w ith an average of about 3.8; but also the
other companies showed average values (3-4
seasonal launches).
For C. montrouzieri the number of launches is
inversely proportional to the company size, ranging
fro m ab o u t 1 for sm a 11 fa rm s to 0.6 fo r larg er ones.
A sim ilar pattern was confirm ed for Leptomastix
dactylopii w ith about. 0.8 launches for companies
under 5 Ha and. 0.4 launches for larger ones.
Average launches < 1 reveal a partial use of insects
(for organic control) that, in these cases, are not
employed on the entire surface of the citrus grove.
The new payment system had an impactalso on the
number of launches that, with reference to 2006-
2011 data, appear in decline. This could be due to
a more parsimonious use of the “organic product”
but also to a kind of “users’ loyalty” (i.e., farmers
despite the new regulation, continue to show a
certain degree of satisfaction).
EVOLUTION OF SERVICE AND PRO-
SPECTS FOR SEVEN YEARS FROM 2013
TO 2020.
The last items briefly discussed in the previous
paragraph, led E.S.A. to review the current huge
production and proceed, after an initial experi-
mental phase, to the diversification of production,
to improve the bouquet offered. In this contest
E.S.A. has already started a project that will be
developed in the period 20 1 3-2020. In particular,
the service aimed at breeding and producting huge
quantities of Aphytis melinus, Criptolaemus
montrouzerii and Leptomastix dactylopii is con-
firmed, re-thinking of new production levels, based
on all the variables mentioned before. Moreover,
seven additional experimental activities have also
been designed one for each year, to be held simul-
taneously with the aforementioned base production,
aimed at increasing the entomological list to be
employed in other contests, as v itic u ltu r a 1, orna-
mental and floricoltural. Each experiment involves
the development of procedures for the breeding of
the following auxiliaries (see below) to be per-
formed, in proper conditions, for the production of
huge quantities of insects.
1 ) Cryptolaemus montruozierii larvae (predators
of P. dtri, citrus mealybug);
156
Giuseppe Greco
Number of farms, by size classes, that used
Aphytis melinus in 2011
Number of farms, by size classes, that used
Aphytis melinus in 2012
SO
70
£ 00
* 50
tM
(m 40
J. 30
= 20
10
0<x<5 5<x<10 >10
Size classes (in hectares)
0<xc5 5*x«lO >10
Size classes (in hectares)
Number of farms, by size classes, that used
Cn ptolaem us nwntrou zieri in 2(111
Number of farms, by size classes, that used
Cryptolaemus montrouzieri in 2012
60
<*>
50
£
40
£
30
O
O
20
£
10
0<X<5 Scx-clQ >10
Size classes (in hectares)
2 50
s
s 10
<2
<5 30
o s
IS
(KM 5**<1S >10
Size classes (in hectares)
Number of farms, by size classes, that used
Leptomastix dactylopii in 2011
Number of farms, by size classes, that used
Leptomastix dactylopii in 2012
80
S 60
o
o
a
40
20
0*x<5 6<x<10 >10
Size classes (in hectares)
GA M
s ■
■s ..
c
o ,
>10
Size classes (in hectares)
Figure 11 . Number of companies, sorted by size classes, th at used Aphytis fflclinUS,
Criptolaemus montrouzerii and Leptomastix dactylopii in 2011 and 2012 (Source e .s .a .).
2) Chrysoperla earned Stephens, 1 836 (Neurop-
tera Chrysopidae) predator of aphids (Benuzzi &
Nicoli, 1988; Osservatorio agroambientale di
Cesena, 1991; Nicoli & Galazzi, 2000);
3) Anagyrus pseudococci (Girault 191 5) (Hy-
menoptera Encyrtidae) parasitoid of PldllOCOCCUS
vitis and ornamental mealybugs, P. ficUS, Pseudo-
coccus longispinus, Ps. affinis , Rhizoecus falcifer
(Avidov etal., 1967; Rosen & Rossler, 1966; Islam
& Jahan, 1993a, b;Blumberg etal., 1995; Islam &
Copland, 1 997, 2000).
4) Encarsia formosa Gahan, 1924 (Hymenop-
tera A phelinidae), parasitoid of w hiteflies as Tridleur-
odes vaporarorium (Westwood, 1 8 5 6 ) (H em ip ter a
A ley rod id ae);
5) Lindorus lophantae (Biaisdeii, 1 8 9 2 ) (Coie-
optera C occinellidae) (generic predator of mealy-
bugs, also active against Aonidiella aurantii) -,
Pest management of citrus fruits in Sicily through interventions of biological control: the biofactory of Ramacca, Catania 1 5 7
6) OrilAS laevigatus (Fieber, 1 8 60) (Flemiptera
A n th o c o rid ae ) predator of thrips (Tawfik & Ata,
1 973; Tavella et al., 1991; Villevieille & Millot,
1991; Chatnbers et al., 1 993; Vacante & Tropea
Garzia, 1993a-b;Meir acker van den, 1994;Alauzet
et al., 1994; Tavella et al., 1994; Frescata & Mexia,
1 995 ; Tom m asini & Nicoli, 1 995);
7) larvae of ChUoCOrUS bipUStlllatUS (Linnaeus,
1758) (Coleoptera Coccinellidae)(predators of
Coccus esperidum (brown soft scale), Ceroplastes
sinenesis (Chinese wax scale), Ceroplastes rusci
(fig wax scale), SflisSCtifl olcUC (Black scale),
Carnuaspis bekii (Purple scale), Aspidiotus blacks
(Oleander scale), Chrisomphalus dictyospermi
(Morgan’s scale), AoYlidlclla aUVantii (California
red scale).
REFERENCES
Abdelrahm an I., 1 974. Growth, development and innate
capacity for increase in Aphytis ell iyS O Hipll Clli M ercet
and A. melinus DeB ach, parasites of California red
scale , Aonidicllci Cllirantii (M ask .) in relation to tem-
perature. Australian Journal of Zoology, 22: 2 1 3-230.
AlauzetC., Dargagnon D. & M alausa J.C., 1994. Bionom-
ics of the polyphagous predator: OvillS IciCVigCltliS
(Het.: A n th o co rid ae) . Entomophaga, 39: 3 3-40.
Avidov Z., RosslerY. & Rosen D., 1967. Studies on an
Israel strain of, AnagyruS pseudoCOCCi (G irault) (Hy-
menoptera: Encyrtidae). II. Some biological aspects,
Entomophaga, 12: 111-118.
Blum be rg D., Klein M . & Mendel Z., 1995. Response by
encapsulation of four mealybug species (Horn opt era:
Pseudococcidae) in parasitization by, AnclgyrilS
pseudoCOCCi. Phytoparasitica, 23: 157-163.
C elli G ., M aini S. & Nicoli G ., 1991. La fabbrica degli
insetti. Franco Muzzio Editore, Padova, 3 3 3 pp.
Chandler L.D., Meyerdirk D.E., Hart W.G. & Garcia
R.G., 1 980. Laboratory studies on the development
of the parasite, AnClgyrilS pseudoCOCCi (G irault)
on in sec tary -reared , PICUIOCOCCUS dtvi (Risso),
Southwest Entomology, 5: 99-103
Chatnbers R.J., Long S. & Helyer N.L., 1993. Effective-
ness of OviuS loCVigUtUS (Hem.: A nthocoridac) for
the control of F rotlkl UliclUl OCCidciltClUs on cucumber
and pepper in the UK. Biocontrol Science and
Technology, 3: 295-307.
Chili A. A., 1987. Enemigos natu rales de los Afidos:
D epredadores. Manejo Integrado de Plagas. En:
Revista del Proyecto MIP/CATIE No. 4 pag 32. 1987.
Collier T.R., 1995. Host feeding, egg maturation, resorp-
tion, and longevity in the parasitoid Aphytis melinilS
(Hymen optera: Aphelinidae). Annals of the Entomo-
logical Society of America, 88: 206-2 14.
De Bach P. & A rg y rio u L.C., 1967. The colon iza tio n and
success in Greece of some imported Aphytis spp.
( H y m .: Aphelinidae) parasitic on citrus scale insects
(Horn. D iaspid id ae). Entomophaga, 12: 325-342.
De Bach P., Rosen D. & Kennet C.E., 1969. Biological
control of coccids by introduced natural enemies.
Huf faker C.B. (Eds.). Biological Control, 165-194.
D ifesa integrata Agrumi - Regione Siciliana - Assessor-
atoAgricoltura e Foreste-Osservatorio perle malattie
delle piante, A cireale.
Flanders S.E., 1 953. Aphelinid biologies with implica-
tions for taxonomy. Annals of the Entomological
Society of America, 46: 84-94.
Flint M .L., 1991. Integrated Pest M anagement for CitVUS.
University of California, Division of Agriculture and
Natural Resources, No. 3303, 144 pp.
Frescata C. & Mexia A., 1 995. Biological control of
Western F lo w er T h rip s w ith OfiuS IdCVigCltUS ( Heter-
optera A nthocoridae) in organic strawberries in
Portugal. In: Parker B.L. et al. (Eds.) - Thrips biology
and management. Plenum Press, New York, 249 p p .
Goidanich G., Bruno Casarini B. & UgoliniA., 1990. La
difesa delle piante da frutto. Bologna, Edizioni
agricole, 1202 pp .
Gottlieb Y., Zchorl-Feln E ., Faktor O. & Rosen D., 1998.
Phylogenetic analysis if parthenogenetic-inducing
Wolbachia in the genus Aphytis (Hymenoptera:
Aphelinidae). Insect Molecular Biology, 7: 393-396.
G ra fto n -C aid w e 11 E.E. & Reagan C .A ., 1 995. Selective
use of insecticides for control of armored scale
(Horn optera: Diaspididae) in San Joaquin Valley
California Citrus. Journal of Economic Entomology,
88 : 1 7 1 7-1 725.
Greco G., 2014a. A biofactory in Sicily for the biological
eradication of the CCTOtitis CCipitCltCl in the Mediter-
ranean countiries. www.entesviluppoagricolo.it
Greco G ., 2014b. La Biofabbrica di Ramacca. www.en-
tesviluppoagricolo.it
H are J.D. & Luck, R.F., 1 994. Environmental variation
in physical and chemical cues used by the parasitic
w asp . Aphytis mcliviUS, for host recognition. Entomo-
logia E xp erim en talis et Applicata, 72: 97-108.
Heimpel G.E. & Rosenheim J.A., 1 995. Dynamic host
feeding by the parasitoid Aphytis Hie l in US: the balance
between current and future reproduction. Journal of
Animal Ecology, 64: 153-1 67.
Hoffmann M . P. & FrodshanA.C., 1993. Natural Enemies
of Vegetable Insect Pest, Cooperative Extension,
Cornell University, Ithaca, N.Y., 63 pp.
Hodek I. & Honek A., 1996. Ecology of C occinellid ae,
Dordrecht, Kluwer Academic Publishers.
Islam K.S. & Copland M .J.W., 1997. Host preference and
progeny sex ratio in a solitary koinobiont mealybug
158
Giuseppe Greco
en d o p aras ito id , AnagyrilS pSeildoCOCCi (Girault), in
response to its host stage. Biocontrol Science and
Technology, 7 : 449-456.
Islam K.S. & Copland M.J.W., 2000. Influence of egg
load and oviposition time interval on the host dis-
crimination and offspring survival of, AfiagyrUS
pS6lldoCOCCi ( Hymenoptera: Encyrtidae), a solitary
endoparasitoid of citrus mealybug , PlcillO COCCUS dtri
(Hemiptera: Pseudococci dae). Bulletin of Entomolo-
gical Research, 90: 69-75.
Islam K.S. & Jahan M., 1993a. Oviposition and develop-
ment of the mealybug parasitoid, AflClgyrUS pseildo-
COCCl (Girault) at different constant temperatures.
Pakistan Journal of Scientific and Industrial Re-
search, 36: 322-324.
Islam K.S. & Jahan M., 1993b. Influence ofhoneydew
of citrus mealybug ( PlciYlO COCCUS dtvi) on searching
behavior of its parasitoid. AnClgyVUS pSeudoCOCCi ,
Crop Protection, 63: 743-746.
Masutti S. & Zangheri S., 2001. Entomologia generate
ed applicata. Padova, Cedam, 978 pp.
Luck R.F., Podoler H. & Kfir R., 1 982. Host selection
and egg allocation behavior by Aphytis melUlUS and
Aphytis lingnanensis: comparison of two facultat-
ively gregarious parasitoids. Ecological Entomology,
7: 397-408.
Luck R.F. & Podoler H 1985. Competitive exclusion of
Aphytis lingnanensis by Aphytis melinus ■. potential
role of host size. Ecology, 66: 904-913.
Luck R.F., Forster L.D. & Morse J.G., 1996. An ecolo-
gically based IPM program for citrus in California’s
San Joaquin Valley using augmentative biological
control. Proceedings International Society of Citri-
culture, pp. 499-503.
Ferrari M ., M arcon E. & M enta A., 2000. Lotta biologica.
Controllo biologico ed integrato nella pratica fitoiat-
rica. Edagricole, Bologna, 366 pp.
Ferrari M ., M arcon E. & M enta A., 2006. Fitopatologia,
entomologia agraria e biologia applicata. Bologna,
Edagricole.
Mario Muccinelli, 2006. Prontuario dei fitofarmaci.
Undicesima edizione. Bologna, Edagricole, 2006.
Massimo Benuzzi & Giorgio Nicoli (1 98 8 ). ChvySO-
perla carnea. In: Lotta biologica e integrata nelle
colture protette (Strategic e tec niche disponibili).
Centrale Ortofrutticola alia Produzione, Cesena,
73-78.
Meiracker R.A.F. van den, 1994. Induction and termina-
tion of diapause in OvillS predatory bugs. Entomolo-
gia experim entalis et applicata, 73: 1 27-1 37.
Milan Vargas o ., 1999 . Cryptolaemus montrouzieri
(Coleoptera: Coccinellidae) como ControlBiologico
de la Chinche Harinosa Rosada del Hibiscus.
Moreno D.S. & Luck R.F., 1992. Augmentative releases
of Aphytis melinus (Hymenoptera: Aphelinidae) to
suppress California red scale (Homoptera: Diaspididae)
in southern California lemon orchards. Journal of
Economic Entomology, 85: 1112-1119.
Murdoch W.W., Briggs C.J. & Nisbet R.M ., 1996. Com-
petitive displacement and biological control in para-
sitoids: a model. American Naturalist, 148: 807-826.
Nicoli g. & Gaiazzi d ., 2000 . Chrysoperla carnea. in:
Nicoli G. & Radeghieri P. (a cur a di). Gli ausiliari
nell'ag ric oltu ra sostenibile. Calderini Edagricole,
Bologna.
Opp S .B . & Luck R.F., 1 986. Effects of host size on
selected fitness components of Aphytis melhlUS and
Aphytis lingnanensis (Hymenoptera, Aphelinidae).
Annals of the Entomological Society of America, 79:
700-704.
Orphan ides G.M., 1984. Competitive displacement
between Aphytis spp. (Hym. Aphelinidae) parasites
of the California red scale in Cyprus. Entomophaga,
29: 275-28 1 .
Osservatorio agroambientale di Cesena (1991). CrisOpe.
In: Guida al rico no scim en to degli organismi utili in
agricoltura. Centro ServiziAvanzati per l'Agricoltura
(Centrale Ortofrutticola di Cesena) e dell'Osser-
vatorio agroambientale di Cesena, Bologna, 40-41.
Pasotti L., Perrotta G ., Raciti E., Saraceno F., Sciacca V.
& Tumminelli R., 2004. Validazione e applicazione
in Sicilia Orientale di un modello di sviluppo della
cocciniglia rossa forte degli agrumi, Aonidiella
aurantll (Maskell), basato sull’accumulo di gradi-
giorno. Atti III Giornate di studio - M etodi numerici.
statistici e informatici nella difesa delle colture
agrarie e delle foreste. Firenze 24-26 nov. 2004:
177-181.
Pekas A., Tena A.,AguilarA. & Garcia-Mari, F., 2010.
Effect of Mediterranean ants (Hymenoptera: Form i-
cidae) on California red scale Aonidiella aiirantH
(Hemiptera: Diaspididae) populations in citrus
orchards. Environmental Entomology, 39: 827-834.
Penny J. & Cranston P.S., 2006. Lineamenti di entomo-
logia. Bologna, Zanichelli, 514 pp.
Pina T., Martinez B. & Verdii, M.J., 2003. Field para-
sitoids of Aonidiella aurantii (Homoptera: Dia-
spididae) in Valencia (Spain). IOBC /W P R S B u lie tin ,
26: 1 09-1 15.
Pina T., 2007. Control biologico del piojo rojo de Ca-
lifornia Aonidiella aurantii (Maskell) (Hemiptera:
Diaspididae) y estrategias reproductivas de su
principal enem igo natural Aphytis chryS Omphali
Mercet (Hymenoptera: Aphelinidae), 384 pp.
Pina T. & Verdu M.J., 2007. El piojo rojo de California
Aonidiella aurantii ( Maskell) y sus parasitoides en
citricos de la Comunidad Valenciana. Boletin de
San id ad Vegetal Plagas, 33: 357-368.
P ollin i A ., Ponti I. & Franco Laffi F., 1988. F ito fag i delle
piante da frutto. Verona, Edizioni L' inform atore
ag rario , 18 8 p p .
Pest management of citrus fruits in Sicily through interventions of biological control: the biofactory of Ramacca, Catania 1 5 9
Pollini. 1998. Manuale di entom ologia applicata.
Bologna, Edagricole, 1462 pp.
Reeve J.D., 1987. Foraging behavior of Aphytis Me l in US:
effects of patch density and host size. Ecology, 68:
530-538.
Rodrigo E. & Garcia-Mari F., 1 992. Ciclo biologico de
los diaspinos de c Itric o s Aonidiella QUYClintii ( M a sk .) ,
Lepidosaphes beckii ( n ew m .) y Parlatoria pergandei
(Com st.) en 1990. Boletin de Sanidad Vegetal Plagas,
18: 31-44.
Rodrigo E. & Garcia-Mari F., 1990. Comparacion del
ciclo biologico de los diaspinos Parlatoria pCTgCUldH,
Aonidiella aurantii y Lepidosaphes beckii (Ho-
moptera, Diaspididae) en citricos. Boletin de Sanidad
Vegetal Plagas, 16: 25-35
Rodrigo M.E., Garcia-Mari F., Rodriguez-Reina J.M.
& OlmedaT., 2004. Colonization of growing fruitby
the armored scales Lepidosaphes beckii, Parlatoria
pergandii and Aonidiella aurantii (Horn., Dia-
spididae). Journal of Applied Entomology, 128:
569-575 .
Rosen D. & RosslerY., 1996. Studies on an Israel strain
of , AnagyrilS pseudococci (Girault) (Hymenoptera,
Encyrtidae). I. Morphology of the adults and
develop mental stages. Entomophaga, 11: 2 69 - 27 7 .
Rosen D., DeBach P. & Junk W. (Eds.), 1 979. Species
of Aphytis of the World (Hymenoptera: Aphelinidae),
801 pp.
Rosen D. & Eliraz A., 1 978. Biological and systematic
studies of developmental stages in Aphytis (Hymen-
optera: Aphelinidae). I. Developmental history of
Aphytis chilensis Howard. Hilgardia, 46: 77-95.
Sorribas J.. Rodriguez R. & Garcia-Mari F., 2010. Para-
sitoid competitive displacement and coexistence: link-
ing spatial and seasonal distribution with climatic
conditions. Ecological Applications, 20: 1101-1113.
Sorribas J., Rodriguez R.. Rodrigo E. & Garcia Mari F.,
2008. N iveles de parasitismo y especies de para-
sitoides del piojo rojo de California Aonidiella
aurailtll (Hemiptera: Diaspididae) en citricos de la
Comunidad Valenciana. Boletin de Sanidad Vegetal
Plagas, 34: 201-210.
Tavella L., Arzone A. & Alma A., 1991. Researches on
Orius laevigatus (Fieb.), a predator of Frankliniella
occidental^ (Perg.) in greenhouses. A prelim inare
note. Bulletin IOBC/W PRS, 14: 65-72.
Tavella L., Arzone A. & Giordano V., 1994. Indagini
biologiche su OrillS laevigatllS ( Fieber) (Rhynchota:
a nth o co rid ae) p re da to re di Frankliniella Occident alis
(Pergande) (Thysanoptera). Atti XVII Congresso
Nazionale Italiano di Entom ologia Udine 13-18
giugno 1994: 519-521.
Tawfik M.ES. & A ta A .M ., 1 9 73. The life-history of
OrillS laevigatUS (Fieber). Bulletin de la Societe
entom ologique d'Egypte, 57: 145-1 5 1.
Ten a A. & Garcia-Mari F., 2011. Current situation of
citrus pests and diseases in the Mediterranean Basin.
IOBC/W PRS B ulletin, 62:365 -3 78.
Tingle C.C.D. & Copland M.J.W., 1 988. Predicting
development of the mealybug parasitoids, AnagyruS
pseudococci , Leptomastix dactylopii and Leptomas-
tidea abnormis ( H y m . Encyrtidae) under glasshouse
conditions. En tom ologia Experimentalis etApplicata,
46, 19-28.
Tingle C.C.D. & Copland M.J.W., 1 989. Progeny pro-
duction and adult longevity of the mealybug para-
sitoids , Anagyrus pseudococci , Leptomastix dacty-
lopii and, Leptomastidea abnormis (Hym. Encyrti-
dae) in relation to temperature. Entomophaga 34:
111 - 120 .
Tommasini M .G. & Nicoli G., 1995. Evaluation of OrillS
spp. as biological control agents of thrips pests: Initial
experiments on the existence of diapause in OrillS
laevigatUS. Medicine Faculty Landbouw, University
Gent (B ), 60/3a: 90 1-908.
Tremblay E ., 1 988. Entomologia applicata, vol. II, pars
I. 2a ed. Napoli, Liguori Editore, 1988.
Turn m inelli R., Conti F., Saracen o F., Raciti E. &
Schiliro, R., 1996. Seasonal development of Califor-
nia red scale (Homoptera: Diaspididae) and Aphytis
melinUS DeBach (Hymenoptera: Aphelinide) on
citrus in Eastern Sicily. Proceedings of the Interna-
tional Society of Citriculture, Sun City, South Africa,
493-498.
Vac an te V. & Tropea Garzia G., 1993a. Prime osser-
vazioni sulla risposta funzionale di OrillS laevigatllS
(Fieber) nei controiio di Frankliniella occidentalis
(Pergande) su peperone in serra fredda. Colture
protette, 22 (suppl. n.l ): 33-36.
Vac an te V. & Tropea Garzia G., 1993b. Ricerche di
laboratorio sulla biologia di OrillS laevigatus
(Fieber). Colture protette, 22 (suppl. n.l): 37-38.
Vacas S A lfaro C., Navarro-Llopis V. & Prim o J., 2009.
The first account of the mating disruption technique
for the control of California red scale, Aonidiella
aurantii Maskell (Homoptera: Diaspididae) using
new biodegradable dispensers. Bulletin of Entomo-
logical Research, 99: 41 5-423.
Vanaclocha P., Urbaneja A. & Verdu M.J., 2009.
Mortalidad natural del piojo rojo de California,
Aonidiella aurantii , en citricos de la Comunidad
Valenciana y sus parasitoides asociados. Boletin de
Sanidad Vegetal Plagas, 35: 59-71.
Viggiani G., 1977. Lotta biologica e integrata nella difesa
fito sanitaria vol. I e vol. II. Liguori Editore, Napoli.
5 17+445 pp.
Villevieille M . & Millot P., 1991. Lotte biologique co litre
Frankliniella occidentalis avec Orius laevigatus sur
fraisier. Bulletin IOBC-WPRS, 14: 57-64.
Walker A. K., 1994. A review of the pest status and
160
Giuseppe Greco
natural enemies of TllvipS pollTli. Biocontrol News
and Information, 15: 7N-10N.
Vacante V. & Benuzzi M., 2004. Produzione massale e
impiego dei principali ausiliari nella lotta biologica
in orticoltura e agrumicoltura. Aracne Edit rice,
Roma, 33 pp.
W aid e S.J., Luck R.F., Yu D.S. & Murdoch W.W., 1989.
A refugee for red scale: the role of size-selectivity by
a parasitoid wasp. Ecology, 70: 1700-1706.
Yu D.S., 1986. The interactions between California red
scale Aonidiella ciurcintii (Maskell), and its para-
sitoid s in citrus groves of inland southern California.
Ph.D. dissertation, University of California, River-
side, 105 pp.
Yu S. & Luck R.F., 1988. Temperature-dependent size
and development of California red scale (Homoptera:
Diaspididae) and its effect on host availability for the
ec to p ara s ito id . Aphytis TYielinUS DeBach (Hymen-
op t e r a : Aphelinidae). Environmental Entomology,
17: 154-161 .
Biodiversity Journal, 2015, 6 (1): 161-164
Monograph
Implantation of Stagg beetles hostels in the city of Geneva
(Switzerland)
Giulio Cuccodoro* & Mickael Blanc
Museum d’histo ire naturelle, Departement d’Entomologie, route de Malagnou 1, 1208 Geneva, Switzerland
Corresponding author
ABSTRACT Brief presentation of our ongoing project of implementation of a network of “Stagg beetles
hostels” in the city of Geneva (Switzerland) aiming at consolidating the last large populations
of big woodboring beetles CerCllTlbyX Cerdo L innaues, 1 758 (Coleoptera C eram by cidae) and
LuCCinilS C6WUS L innaues, 1 75 8 (Coleoptera Lucanidae) in Switzerland.
KEYWORDS stagg beetles hostels; Geneva; woodboring beetles; LuCCMUS CerVUS; CCYCUTlbix C6rdo.
Received 05.12.2014; accepted 1 5.02.20 1 5; printed 30.03.20 1 5
Proceedings of the 2nd International Congress “Speciation and Taxonomy”, May 1 6 th - 1 8 th 2014, Cefalu-Castelbuono (Italy)
INTRODUCTION
The large woodboring beetles Cercimbyx CCvdo
Linnaeus, 1 75 8 (Coleoptera C eram by cidae) and
LllCCinuS cervus Linnaeus, 1 75 8 (Coleoptera Lu-
canidae) are associated to oaks and beechs. Their
larvae feeding on decaying wood take 3 to 5 years
to develop into imagoes, which live only one Sum-
mer and hardly disperse further that half kilometer
from their native tree. In North and Central Europe,
they can complete their life cycle only in trees big
enough to protect larvae from the Winter frosts.
During the 20th century their distribution has
however drastically regressed due to intensive
exploitation of those trees for timber as well as
changes in agriculture techniques.
Because of the large array of other useful and
more elusive organism s benefiting of their favoured
habitat (i.e. senescent oaks and beechs), they are
considered ombrella species and received therefore
protected status in Europe since 1992. In Switzer-
land, where they received federal protected status
only in 2011 (OFEV, 2011), they can be found
today essentially in the southern part of the country,
CerCUnbyX Cerdo with only sc arse populations in the
cantons of Geneva, Valais and Tessin.
Despite a limited and rather urbanized territory,
the canton of Geneva has the privilege to host the
most abundant Swiss populations of both of these
emblematic beetles. Amazingly, they are essen-
tially found in the hearth of the city itself rather
than in the surounding countryside, where most
old trees were cut in the early 20th century for tim-
ber and adaptation of the landscape to mechanized
agriculture.
Renewal of suitable natural habitats better
distributed in space and time in the countryside is a
matter of several decades. Meanwhile, as falling
branches put citizens and other park’s users at risk,
senescenth oaks and beechs in town are gradually
cut and removed. Asa result, despite their apparent
abundance in the city of Geneva, these urban popu-
lations of large wood-boring beetles are each day
less abundant and more fragmented, in such a way
that their disapparition seems programmed if
nothing is attempted today.
162
Giulio Cuccodoro & Mickael Blanc
MATERIAL AND METHODS
We thus decided to try to consolidate the urban
populations of these protected beetles by promoting
in the city of Geneva the set up of a network of
“stagg beetles hostels”. Already implemented
successfully in several other places elsewhere in
Switzerland and Europe, « woodboring beetles
hostels» consist basically in 1 meter long logs half
buried in the ground recreating artificially the
appropriate conditions for female oviposition and
subsequent developpem ent of the larvae.
However installation of such large and lasting
structures in the heart of a densely urbanised city
faces various technical and social problems. In first
instance stagg beetles hostels are more cumbersome
than birdhouse: they require several square meters
of land, should remain over a decade to be really
efficient, and as such mustbe installed only in close
concertation with all relevant city services (urban-
ists, gardeners, maintenance, etc.). Therefore we
dedicated a lot of time and energy explaining urban-
ists, gardeners and maintenance workers that stagg
beetles hostels were as necessary as easy to build,
but moreover that they consist indeed in an inev-
itable new element of the urban furniture of the
Geneva of the 21th century.
Second and certainly mostchallenging problem
is that most citizens perceive insects as a nuisance,
a source of danger (punctures, vectors of diseases)
or as revelators of defective sanitary conditions.
Lucanus Scopoli, 1 763 and CevCDTlbyX Linnaeus,
1 75 8 unfortunately don’t escape this “rule”, which
is even exacerbated by their quite impressive size
in such a way that they are often mistaken as
dreadful exotic beasts fallen off a plane from Africa
or elsew here.
Therefore we accompanied our project from the
beginning with a real campaign for « social rehab-
ilitation » of these large woodboring beetles. In first
instance we made a call to the citizens for observa-
tions in the frame of a participatory inventory
aiming at 1) bringing presence of these beetles to
the knowledge of citizens, 2) teaching to recognize
them, 3) explaining they role, 4) drawing attention
to their patromonial status and the responsibility of
Geneva citizens regarding their respect and protec-
tion, and 5) offering people the possibility to con-
tribute directly to this issue by the transmitting their
own observations.
Besides we took every opportunity to talk to
school classes and publish in daily newspapers
small articles declining these thema. A WEB page
specially dedicated to that project and relaying per-
manently our call for observations was also created
(www .ville-ge.ch/m hng/coleopteres_bois_geneve.php).
RESULTS
It took some two years from the origin of our
project in 2011 to the construction of the first stagg
beetle hostel in Geneva, which was achieved on the
17th of April 2013 by gardeners of the city of
Geneva assisted by childrens. Installed at the foot
of three big senescent oaks colonized by both
Cerambyx and Lucanus in the “Parc La Grange”,
the most scenic and visited pare of the city of
Geneva, this Stagg beetles hostel is agremented
with a graphic pannel summarizing the biology, role
and status of these beetles, with a flash code address-
ing smartphone users directly to ourWEB page for
further informations. The device was completed
with an attractive giant oak LuCdUUS sculpted by a
local artist aiming both at catching attention of the
visitors and favouring perenniality of the hostel,
which symbolizes to us the participatory involve-
ment of the Geneva citizens for a more rational
management of their environment.
CONCLUSIONS
This realisation will take all its meaning only if
drawn in the future. In effect to modify the social
perception of large woodboring beetles from the
status of unwanted frightening pests to majestic
useful animals being integral part of the environ-
mental identity of the citizens will be certainly a
long-term process. In this perspective participatory
involvement of the public in a “continuous asses-
ment” via regular calls for inform ations in the daily
media seems very important to us. However it’s
obvious to our eyes that the best way to accelerate
adhesion rate to the cause of large woodboring
beetles w ill consist in penetreting public education
programs, and that from the earliest school
classes.
Nevertheless it appeared that once explained
the environmental issues and technical feasability
Implantation of Stagg beetles hostels in the city of Geneva, Switzerland
163
Figure 1. First Stagg beetles hostel of Geneva (Switzerland): co n s tru c tio n w ith ch ildre n .
Figure 2. First Stagg beetles hostel of Geneva (Switzerland): sculpture by Sylvio Asseo.
164
Giulio Cuccodoro & Mickael Blanc
Figure 3. First Stagg beetles hostel of Geneva (Switzerland) as completed (logs, sculpture, and didactic pannel).
of the project, most professional actors of the city
services concerned were enthousistic to contribute
at their level to its realization. The best proof we
can present is that additional 7 stagg beetles hostels
have been installed by the gardeners in other parks
of the city in 2014, and more are planned for 2015.
Next steps will be to increase the density the
network of stagg beetles hostels with a target of
one each 300 m in order to enhance gene flows
between each individual population, then to imple-
ment corridors of stagg beetles hostels favoring
natural dispersal of these beetles through the
suburban crown toward the country side they used
to belong. Meanwhile we already work with fore-
stal autorities in order to promote the plantation of
oaks and beeches in the contryside with a better
scaling in space and time of suitable habitats for
these magnificent insects.
The project received the “Geneva cantonal
award forsustainable developmentedition 2014”.
REFERENCES
OFEV, 2011. Liste des especes prioritaires au niveau
national. Especes prioritaires pour la conservation au
niveau national, etat 2010. Office federal de 1 ’ en-
viron n e m e n t. L ’ e n v iron nement pratique n ° 1 1 0 3 , 132
pp.
CEE, 1 992. Directive 92/43/CEE du Conseil, du 21
mai 1992, concern ant la conservation des habitats
naturels ainsi que de la faune et de la flore sauvages.
CEE. Journal officiel n° L 206 : 0007-0050.
Biodiversity Journal, 2015, 6 (1): 165-170
Monograph
Requalification of coastal plant landscape of South-Eastern
Sicily, Italy: the case of Marina di Priolo
Angelo Zimmitti 1 , Rosaria Mangiafico 2 & Pietro Pitruzzello 3 *
'Via Garibaldi 2 1, 96010 Melilli, Syracuse, Italy
! A ssociazione Anteo, via Concerie 52, 96010, Syracuse. Italy
Universita degli Studi di Catania, Cutgana, Polo Biotecnologico via Santa Sofia 98, 95123 Catania, Italy
Corresponding author, e-mail: p itr u z z e llo p iero @ gmail.com.
ABSTRACT In this paper the A uthors examine the psarn m ophilous vegeta ti on and the degrees ofnaturalness
of the coastal plant landscape of a part of the South-Eastern littoral in Sicily. This area is char-
acterized by considerable human pressure due to the presence of a large industrial center and
beach tourism. The recent construction of the garden next to the beach, made mainly with
ornamental plants has contributed to further amend the original physiognomy of the coastal
landscape. Were analyzed, with phy to sociological me thod, psamm ophilous plant com m unities
and zonation of vegetation. The results of the analysis show a impoverishment of flora and a
progressive decline in the psam m ophilous communities mainly due to the constant leveling
the beach in summer. The authors propose a series of actions aimed at the requalification and
conservation of coastal vegetation landscape of the investigated area.
KEY WORDS plant landscape; re q u alific a tio n ; littoral; human pressure.
Received 15.10.2014; accepted 30.01.2015; printed 30.03.2015
Proceedings of the 2nd International Congress “Speciation and Taxonomy”, M ay 1 6 th - 1 8 th 2014, Cefalu-Castelbuono (Italy)
INTRODUCTION
In the present work we analyzed the plant land-
scape of M arina di Priolo, a stretch of sandy coast
between Marina di Melilli and Magnisi peninsula,
about 6 km north of Syracuse (Sicily, Italy) (Fig. 1).
In a not too distant past the area was used for the
production of salt in the salt marshes of Magnisi,
placed in a large basin behind the dunes adjacent to
the study area. A lthough reduced from its original
e x ten t, this im p o rta n t hum id e n v iro n mentis p rote c -
ted through the establishmentofthe R.N.O. "Saline
di Priolo" managed by the L.I.P.U. (D.A. n. 807/44
of 1 2/2 8/2000). Since the 50s of last century, the
area has undergone significant environmental
change mainly due to the progressive establishment
of one of the largest petrochemical industrial cen-
ters of Europe. The massive industrialization of the
area has also led to the growth of urban centers and
neighboring persistent anthropogenic coastal envir-
onment that, in recent years, was also affected by
the profound transformations related to the increase
in to u rism .
The recent creation of a green area called
"Garden of the Sea", adjacent to the beach, consists
mainly of ornamental species, some exotic, helped
to further modify the original structure of the
coastal landscape. The purpose of this research is
the cognitive analysis of the dune environment,
spatial seriation of psamm ophilous plant com-
munities and their state of preservation. Based on
the results obtained, we propose actions for the
rehabilitation and protection of plant landscape of
the site investigated.
166
Angelo Zimmitti etalii
The area of study. From the perspective of geo-
log ic a 1- s tr u c tu r a 1 the area of Marina di Melilli,
Syracuse (Sicily, Italy) is part of the Hyblean Plat-
eau and the local stratigraphic succession is repres-
ented by ceno-neozoic carbonate rocks (Carbone et
al., 1 9 8 6). Examining the th erm o - p lu v io m e trie data
fro m th e n e arb y statio n of Syracuse, the dim a te o f
the study area is Mediterranean, with mild, rainy
winters and hot, dry summers (Zampino et al.,
1997). While, as evidenced by Scelsi& S p am p in a to
(1 998) bioclimate is in the range inferior thermo-
mediterranean dry type.
MATERIAL AND METHODS
The methodological approach used for phytoso-
ciological study of the psam m ophilous vegetation
is that of the Sigmatista School of B raun-B lanquet
(B raun-B lanquet, 1964), while for syntaxonomical
framing were followed proposals of Brullo et al.
( 2002 ).
The collected samples were determined accord-
ing to the Flora of Italy (Pignatti, 1 9 82 ), prepared
and preserved in the herbarium of the Ecomuseo dei
Monti Climiti Melilli (Laboratory of Nature and
E n v iro n m e n tal) .
RESULTS
Despite the heavy distortions of anthropogenic
nature, the investigations carried out m ade it possible
to identify, in the least disturbed stretches of coast-
lin e , d ifferen tcomm un itie s of psammoph ilo u s plan ts
that, despite impoverished of m any typical elements,
hint at some aspects of the original plant landscape
and suggest effective conservation measures for the
protection and rehabilitation of ecosystem s. Through
the observations made could be detected, proceed in g
from the aphytoic zone inland, a first strip of tero-
phitic h alo nitro p h ilo u s vegetation, parallel to the
coast-line, which is closely pioneer, ascribable to the
Salsolo-Cakiletum maritimae, characterized by the
dominance of Ccikile maritima Scop, associated with
Salsola kali l . and Polygonum maritimum l .
The next strip, attributable to the CyperO-
AgVOpy return juncei, is characterized by herbaceous
perennial plants of low embryo dunes. The associ-
ation p h y sio g n o m ic ally is characterized by the do-
minance of Elytrigia juncea ( l . ) n evskiwhich is as-
sociated with Sporobolus virginicus { L.) K until and
Achillea maritima (L .) Ehrend. et Y.P. Guo. The ve-
getation parallel to the latter strip is dominated by
Centaurea sphaerocephala l. and Onosis natrix
subsp. ramosissima (D esf .) Batt.; are also present
Pancratium maritimum l., Euphorbia terracina l.
and LotUS CytisoideS. It is a plant comm unity ascrib-
able to the Centaureo-Ononidetum ramosissimae,
chain aephytic and h em ic rip to p h y tic vegetation nor-
mally c o n fin ed on th e dunes furth er in lan d with little
movement, the expansion of which is favored by
human disturbance (M in is s ale & Sciandrello, 2010).
Proceeding inland, the psam m ophilous series
is interrupted by a road parallel to the coastline.
The analysis also revealed a degradation of the
psam m ophilous vegetation due to the leveling and
trampling of the dunes in the vicinity of the holi-
day season. The persistent action of scraping in
sandy shore led to the demise of mobile dunes with
typical vegetation with Ammophila arenaria (L.)
Link, therefore, observing the current vegetation
con firm s th e absence of th e typical z o n a tio n of dune
environments like those along the Ionian coast of
south-eastern and far less degraded (see Brullo et
al., 1988; M inissale & Sciandrello, 2010). The plant
communities found are ranked according to the
following syntaxonomical scheme:
CAKILETEA MARITIMAE R.Tx & Preising in
Br.-B 1. & R.Tx 1952
CAK1LETALIA IN T E G R IF O L I A E R.Tx ex
Oberd. 1 949 corr. R iv as -M artin ez , Costa &
Loidi 1992
CAKILION MARITIMAE Pignatti 1953
Salsolo-Cakiletum maritimae Costa & m ansanet
1981 corr. Rivas-M artinez et al. 1992
AMMOPHILETEA Br.-B 1. & R.Tx ex Westhoff et
al. 1946
AM M OPHILETALIA Br.-Bl. 1933
AM M OPHILION AUSTRALIS Br.-Bl. 1921 em.
Gehu, Rivas-M artinez & R.Tx in Rivas-M artin ez
et al. 1980
Cypero capitati-Agropyretum juncei Kuhnhoitz-
Lordat (1 923 ) Br.-Bl. 1933
CRUCIANELLETALIA MARITIMAE Sissing
1974
ON ON ID IO N RAMOSISSIMAE Pignatti 1952
Centaureo-Ononidetum ramosissimae Br.-B 1 . &
F re i in F re i 19 3 7
Requalification of coastal plant landscape of South-Eastern Sicily, Italy: the case of Marina di Priolo
167
Figure 1. The area of study: Marina di Priolo, Syracuse
( S ic ily , Italy ) .
Figure 3. Marina di Priolo, Syracuse (Sicily, Italy):
p s am m o p h ilo u s vegetation.
CONCLUSIONS
The research suggests a number of measures
aimed at the improvement and protection of plant
landscape of the study area:
- allocation of a minimum unit of surface pro-
tection to the progressive development of natural
v e g e ta tio n . T h e "minimun dynamic area " is d e fin e d
as the balance between the effects of disturbance
and the area required for the development of the
psam mophilous community. In our case, the situ-
ation found suggests to preserve space as widely as
possible to enable us to reconstruct the seriation of
vegetation and restore the dune system.
Figure 2. Marina di Priolo, Syracuse (Sicily, Italy):
waste left along the beach.
Figure 4. Marina di Priolo, Syracuse (Sicily, Italy):
the green area caled “Garden of the Sea”.
This could be achieved by:
- elimination of non - native flora, both spontan-
eous and ornamental, present in the area concerned
with habitat restoration through the use of native
species from propagation material (seed), local
germ plasm collected in a special center or in a
neighboring area less anthropized and comparable
with the examined site. It is therefore proposed a
renaturalization especially in the "Garden of the
Sea", by converting the area into a natural garden
characterized by the recovery of plant comm unities
typical of dune environments having a dual role:
eco-functional and didactic educational.
- development of a seaside tourism compatible
with the environmental restoration of the site.
168
Angelo Zimmitti etalii
Salsolo-Cakiletum maritimae
Releve Number
1
2
3
S urfac e (m q )
50
50
20
Slope (% )
50
40
40
Char. Ass.
Salsola kali l .
1
+
+
3
Char. Euphorbion peplis & Cakiletea maritimae
Cakile maritima scop.
4
3
3
3
Polygonum maritimum l .
1
+
1
3
Xanthium strumarium italicum (m oretti) d . Love
+
+
2
Chamaesyce peplis (L . ) p ro k h .
+
1
Companions
Sporobolus virginicus (L .) k unth
+
+
+
3
Achillea maritima (l .) Ehrend. & y.-p. Guo
1
+
2
Table 1. The area of study, Marina di Priolo, Syracuse (Sicily, Italy):
Salsolo-Cakiletum maritimae (Date 2 0 .x .2 0 1 2 ) .
Cypero capitati-Agropyretum juncei
Releve Number
1
2
3
S urfac e (m q )
30
30
30
Slope (% )
70
60
60
Char. Ass.
Elytrigia juncea (l .) Nevski
4
3
3
3
Sporobolus virginicus (L .) k unth
2
1
3
3
Achillea maritima (L.) Ehrend. & y.-p. Guo
+
1
Char. Ammophilion & Ammophiletea
Eryngium maritimum l .
1
+
+
3
Pancratium maritimum l .
+
+
+
3
Echinophora spinosa l .
+
+
2
Silene nicaeensis ah.
+
+
2
Companions
Cakile maritima Scop.
+
+
+
3
Polygonum maritimum l .
+
+
2
Table 2. The area of study, M arina di Priolo, Syracuse (Sicily, Italy):
Cypero capitati-Agropyretum juncei (Date 20.x. 2012).
Requalification of coastal plant landscape of South-Eastern Sicily, Italy: the case of Marina di Priolo
169
Centaureo- Ononidetum ramosissimae
Releve Number
l
2
3
S urfac e (m q )
1 5
20
20
Slope (% )
100
100
100
Char. Ass.
Ononis hispanica ramosissima (Desf.)FortheretPodiech
4
4
3
3
Centaurea sphaerocephala l .
2
1
2
3
Char. Crucianelletalia & Ammophiletea
Euphorbia terracina l .
2
2
1
3
Pancratium maritimum l .
1
+
+
3
Elytrigia juncea (L .) Nevski
1
1
+
3
Silene nicaeensis a 11 .
1
+
+
3
Sporobolus virginicus (L .) k unth
1
+
2
Ononis variegata l .
+
+
2
Cyperus capitatus v a n d e 1 .
1
1
Companions
Anisantha rigida (Roth) h y 1 .
2
2
1
3
Silene colorata Poir.
2
1
1
3
Lagurus ovatus l .
1
1
+
3
Vulpia fasdculata (Forssk.) Fritsch
1
1
+
3
Scolymus liispanicus l .
1
+
1
3
Glebionis coronaria (l.) Spach
1
+
+
3
Dittrichia viscosa (L .) g reuter
1
+
2
Cutandia maritima (L .) b arbey
+
+
2
Table 3. The area of study, Marina di Priolo, Syracuse (Sicily, Italy):
Centaureo-Ononidetum ramosissimae (d ate 1 5 .iv.20 1 3 ).
The recommended actions will help restoring
psammophilous communities also improving the
ecological continuity between the dune environ-
ment and the wetland of RNO "Saline di Priolo"
permitting, at the same time, visitors to perceive a
higher degree of naturalness of the environment
compared to the current situation of degradation.
The proposed objectives are part of a broader scope
of environmental restoration of the examined area,
connecting with an ongoing project concerning the
work of restoring of the f o rm e r tenement E S P E S I ,
located on the peninsula M agnisi, to be allocated to
the visitor center and guest house (PO FESR 2007-
2013 axis 3 ob. specific 2.1; program agreement
31/08/2011 between Department of Environment
and Regional Authorities of the “Enti gestori delle
Riserve Siciliane”).
REFERENCES
Braun-BlanquetJ., 1964. Pflanzensoziologie. 3 ed, Springer
Verlag, Wien, 865 pp.
170
Angelo Zimmitti etalii
Brullo S.,De SantisC.,FurnariF.,Longhitano N.& R onsis-
valle G 1 988. La vegetazione dell’Oasi della Foce del
S im e to (S ie ilia orien tale). B rau n - B lan qu etia, 2 : 165-1 88.
Brullo S., Giusso Del Galdo G., Minissale P., Siracusa
G. & Spam pinato G., 2002. Considerazioni sintasso-
nomiche e fito g e o g rafic h e sulla vegetazione della Si-
cilia. Bollettino Accademia Gioenia di Scienze
Naturali, 3 5 (3 6 1 ): 3 25-3 5 9.
Carbone S., Grasso M . & Lentini F., 1 9 86. Carta geolo-
gica del settore n o rd -o rie n ta le ibleo. Scala 1:50.000.
S.E.L.C.A., Firenze.
Minissale P. & Sciandrelo S., 2010. Flora e vegetazione
terrestre della Riserva Naturale di Vendicari (Sicilia
Sud-Orientale). L'Area Protetta di Vendicari. Atti del
Convegno Celebrativo per il 35° anno di fondazione
dell'Ente Fauna Sicilia 8°, pp.128.
Pignatti S., 1982. Flora d 'Italia . I — 1 1 1 , Edagricole,
Bologna, 2324 pp.
Scelsi F. & Spam pinato G ., 1 998. C a ra tte ris tic h e biocli-
matiche dei Monti Iblei. Bollettino Accademia
Gioenia di Scienze Naturali, 29 (35 3 ): 27-43.
Zampino S . , Duro A., Piccione V. & Scalia C., 1997.
Fitoclima della Sicilia. Termoudogrammi secondo
W alter & Lieth. Atti 5° W orkshop Prog. Strat. C .N .R.
"Clima, Ambiente e Territorio nel M ezzogiorno"
A m alfi. 2: 7-54.
Biodiversity Journal, 2015, 6 (1): 171-174
Monograph
The use of flora, vegetation and habitats in the studies of
Environmental Impact Assessment
Pietro Minissale
Dipartimento di Scienze Biologiche, Geologiche e Ambientali, Sez. Biologia Vegetale, Universita di Catania, via A. Longo 19,
95125 Catania (Italy); e-mail: p.minissale@unict.it
ABSTRACT The paper examines local flora, vegetation and habitats in order to highlight the plant
component’s role as not only an indicator of the quality and state of the environment, but also
as an extremely useful element in restoration activities required by environmental impact
studies. Some methodological proposals have been done as objective criteria in the assessment
procedures.
KEY WORDS local flora; biodiversity; indicators; restoration.
Received 18.06.2014; accepted 08.09.2014; printed 30.03.2015
Proceedings of the 2nd International Congress “Speciation and Taxonomy”, May 16th- 18th 2014, Cefalu-Castelbuono (Italy)
INTRODUCTION
In the studies of environmental impact, the bi-
otic component as a whole (and which includes
man) is normally the cornerstone on which the im-
pact generated from plans and projects is assessed.
This paper examines the plant component (limited
to the vascular flora and plant communities) in
order to highlight the way it plays a key role as in-
dicator of the quality and state of the environment,
as an accurate sensor of the impacts but also as an
extremely useful element in restoration activities
required or proposed by environmental impact stud-
ies. It also intends to make a few methodological
proposals for the use of objective criteria in the
assessment procedures.
MATERIAL AND METHODS
The paper is a brief methodological review,
resulting from experience gained on several impact
assessments developed in recent years on the island
of Sicily, which is representative of the Mediter-
ranean region. These assessments have allowed sa-
lient features of plant biodiversity to be recognized
and taken into account in an impact assessment
study in order to minimize the effects of exploitation
of plant biodiversity in favour of conservation policies.
INDICATORS OF THE QUALITY AND
STATE OF THE ENVIRONMENT
The assessment of the quality or the degree of
naturalness of a study area is crucial in making a
considered judgment on the quality and intensity of
the impact that the implementation of a plan or pro-
ject leads to. Plants species and plant communities
fully comply with these requirements.
Plant species are, in fact, indicators of the qual-
ity and state of the environment since each taxon
of the flora of a study area is placed into specific
habitats. The narrow endemic species, or otherwise
rare or included in the national or regional red lists,
are usually associated with the most natural and
172
Pietro Minissale
sensitive habitats effected by human actions. In this
way, they indicate the presence of important hab-
itats to be protected. Furthermore, these plants are
the preferred subjects for appropriate impact as-
sessments (Rossi et al., 2014). In contrast, the most
trivial synanthropic species are present in habitats
with predominantly anthropogenic determinism
such as farmland, edges of the road, landfills, etc.
and therefore they are not veiy useful in the context
of environmental assessments or, at the very least,
they indicate the lack of floristic elements to be
protected. In this case traditional agricultural
landscape as a whole, rather than the natural one,
will be focused on for the impact assessment
(Barbera & Cullotta 2012).
Plant communities, better than single species,
are very fine and accurate indicators of the type,
quality and state of the environment since they are
an expression of ecological factors such as climate,
soil, and anthropogenic influence which allows
them to exist within the framework of the vegeta-
tion series (1), where discrete units (with statisti-
cally uniform floristic composition) can usually be
recognized (Pott, 2011). Plant communities are also
the most characteristic and diagnostic element of
habitat, which is considered as a uniform parts of
the ecosystem and is almost always detectable in
large to small scale on cartography.
In most cases, their identification is easier than
a single species whose presence may be due to
chance. For this reason, plant communities are the
driving elements for impact assessments as they
may have different value in the conservation policies
and different sensitivity to human actions. The spa-
tial mosaic of habitats is also useful in assessing the
potential fauna of an area and therefore, it allows
an opinion on the ecosystem as a whole to be
expressed (Sabella, 2015).
A topic generally not highlighted for environ-
mental impact studies is the great local diversity of
the indicators mentioned above. In Italy, for example,
it is possible to recognize at least three biomes,
(1) A series of vegetation is made up of all the plant com-
munities related by dynamic relationships that could occur
in a ecologically homogeneous space with the same poten-
tial vegetation, having the same physical conditions (i.e.
meso-climate, soil type, geomorphology). It is dependent on
processes of vegetational succession, management and
extreme events (e.g. fire, storm damage, volcanic eruption).
Mediterranean, Alpine and Temperate. There are
also significant differences in the flora and vegeta-
tion which exist in smaller territories as highlighted
by several authors (Greuter, 2010; Blasi et al., 2010;
Blasi & Frondoni, 2011). Due to this fact and al-
though the method of investigation and assessment
may be the same, the contribution of regional
specialists who can better understand or highlight
floristic and vegetational peculiarities is required.
With regard to the assessment methodologies and
to be able to converse with other specialists, it is
necessary to use objective and quantifiable criteria
as much as possible. One of these is the “floristic
vegetational value” for plant species with endemism,
rarity, and/or endangered taxa, while for plant com-
munities and habitat the evaluation depends on
vegetation series position and biogeographical
significance. A proper scale of values, to be assigned
to these biotic elements, must be compared with the
induced changes by a plan or project, allowing more
calibrated matrices to be created.
As explained regarding indicators, the botanist
works on two main levels: plant species and plant
communities, recognizable as habitats. A third level,
the landscape, and in particular plant landscape,
should be considered but this competence is to be
shared with other specialists such as agronomists,
geologists and architects. However, if we consider
the natural plant landscape for which the recogni-
tion of vegetation series is key, the environmental
analyst with a botanical background remains the
only acceptable specialist.
The sources for plant species are the national
and regional floras, the red lists, and the lists of pro-
tected species by laws and directives. However,
these lists are often deficient because they ignore
many important species. The most striking case is
that of Annex 2 of the European Directive 92/43 in
which many rare endemic species are not men-
tioned. A likely reason is that in the 1990s the spe-
cialists involved in making up this list did not fully
understand its importance. At the level of plant
communities, the list of habitats of Community
interest is very useful (listed in Annex 1 of the
above mentioned directive) where the deficiencies
seem fairly small. As highlighted above, the need
to know the vegetation series of each area is of great
importance in order to safeguard and properly
assess any mature stages if present. In Italy, a refer-
ence element is Blasi (2010a, 2010b) who indicates,
The use of flora, vegetation and habitats in the studies of Environmental Impact Assessment
173
based on the collaboration of many regional spe-
cialists, all the vegetation series of the national ter-
ritory. Also in this case, the general pattern must be
checked and adjusted for each individual case study.
ENVIRONMENTAL COMPENSATIONS
The main purpose of providing environmental
compensation for the damage caused to nature
through building and construction projects is to
maintain the quality of the environment (Persson,
2013). This approach has been used to a large extent
in Germany and the USA since the 1970s, and the
EU has adopted several directives dealing with
environmental compensation. Therefore, if environ-
mental impact assessments show even a modest loss
to the habitats or ecosystems, this gives the analyst
the opportunity to propose compensatory actions
that should result in the recovery of damaged hab-
itats or improvement of neighbouring ones not
directly affected by the plan or project. These com-
pensatory activities, when related to environmental
restoration, cannot be wasted or nullified by the
planting of species that are not relevant to the site
but they require a highly skilled design as will be
explained in the following paragraph.
RESTORATION ACTIVITIES
The use of plant species for environmental
restoration is an opportunity, not only to mitigate
or compensate the impact of construction or infra-
structure work but also to trigger or facilitate the
recovery of habitats often in decline. This assertion
is valid only if the restoration activities are set in a
rigorous way, i.e. taking into account the vegeta-
tion series and local potential vegetation. Once
again, the local plant diversity is the driving
element of the interventions. These assumptions
are widely accepted in northern Europe and North
America (Persson, 2013), but it is hard for them to
be established in the Mediterranean region, and in
Italy in particular.
At present, most of the infrastructure work
(especially linear ones such as roads and railways)
are marked by sometimes alien invasive plants.
The presences of these alien plants are, in many
cases, a legacy of the past (i.e. work created
decades ago) but in recent cases, such as the
Catania-Siracusa highway completed in 2009, on
the slopes, potentially invasive species such as
Cortaderia selloana have been planted with
some native plants (Basnou, 2009; Domenech et
al., 2005).
Nevertheless, there are some pioneering activit-
ies developed in Sicily, which are following the new
direction of environmental restoration (La Mantia
et al., 2012; Barbera et al., 2013) and bodes well for
the future.
CONCLUSIONS
With the present paper, an attempt to highlight
what the salient points in an environmental impact
assessment regarding the flora and vegetation has
been carried out. The outlined framework emphas-
izes the importance of taking into account the
above-mentioned elements (species, communities,
habitats) in all evaluations and proposal steps of
a work as they are the perfect sensors of any
positive/negative impact, the indicators of environ-
mental quality, and the main protagonists in envir-
onmental restoration and mitigation activities.
On these basis, the environmental analyst must
be able to better guide or mitigate the project’s
actions, considering that, in spite of any attempt to
contain the rate of destruction or alteration of nat-
ural resources, the transforming activity of man and
its resulting impacts on the biosphere, both large
and small-scale, will never end.
REFERENCES
Barbera G. & Cullotta S., 2012. An Inventory Approach
to the Assessment of Main Traditional Landscapes in
Sicily (Central Mediterranean Basin). Landscape
Research, 37: 539-569.
Barbera G., Di Leo C. & Scuderi L., 2013. Problems and
perspectives on the use of native species: landscape
restoration project within the “international Verdura
Golf & SPA Resort of Sciacca” (Agrigento, Sicily)
Workshop: Ensuring the survival of endangered
plants in the mediterranean island 18°-20° April
2013, Orto Botanico di Catania, Sicily.
Basnou C., 2009. Cortaderia selloana (Schult. & Schult.)
Asch & Graebn., pampas grass (Poaceae, Magnolio-
phyta). In: DAISIE: Handbook of alien species in
Europe Dordrecht, Netherlands: Springer, 346 pp.
174
Pietro Minissale
Blasi C. (Ed.), 2010a. La vegetazione d’ltalia. Palombi
e Partner s.r.l. Roma.
Blasi C. (Ed.), 2010b. La vegetazione d’ltalia. Carta delle
Serie di vegetazione , scala 1:500.000. Palombi e
Partner s.r.l., Roma.
Blasi C. & Frondoni R., 2011. Modem perspectives for
plant sociology: The case of ecological land classi-
fication and the ecoregions of Italy. Plant Biosystems,
145 (suppl.): 30-37.
Blasi C., Marignani M., Copiz R., Fipaldini M., Bonac-
quisti S., Del Vico E., Rosati L. & Zavattero L., 2010.
Important Plant Areas in Italy: from data to mapping.
Biology Conservation, 144: 220-226.
Domenech R., Vila M., Pino J. & Gesti J., 2005. Histor-
ical land-use legacy and Cortaderia selloana invasion
in the Mediterranean region. Global Change Biology,
11: 1054-1064.
Greuter W., 2010. Lectio magistralis: “Flora mediterranea:
una in multo, multum in una”. Pp. 19-30 in: Confer-
imento della Laurea specialistica ad honorem in Bio-
logia ed Ecologia vegetale. Universita degli Studi di
Palermo, Palermo.
La Mantia T., Messana G., Billeci V., Dimarca A., Del
Signore M., Leanza M., Livreri Console S., Mara-
ventano G., Nicolini G., Prazzi E., Quatrini P, Sangue-
dolce F., Sorrentino G. & Pasta S., 2012. Combining
bioengineering and plant conservation on a Mediter-
ranean islet. Iforest, 5: 296-305.
Persson J., 2013. Perceptions of environmental compens-
ation in different scientific fields, International
Journal of Environmental Studies, 70: 4, 611-628.
Pott R., 2011. Phytosociology: a modern geobotanical
method. Plant Biosystems, 145 (Suppl. 1): 9-18.
Rossi G., Montagnani C., Abeli T., Gargano D., Pemzzi
L., Fenu G., Magrini S., Gennai M., Foggi B.,
Wagensommer R .P, Ravera S., Cogoni A., Aleffi M.,
Alessandrini A., Bacchetta G., Bagella S., Bartolucci
F., Bedini G., Bernardo L., Bovio M., Gastello M.,
Conti F., Domina G., Farris E., Gentili R., Gigante
D., Peccenini S., Persiani A.M., Poggio L., Prosser
F., Santangelo A., Selvaggi A., Villani M.C., Wilhalm
T., Zappa E., Zotti M., Tartaglini N., Ardenghi
N.M.G., Blasi C., Raimondo F.M., Venturella G.,
Cogoni D., Puglisi M., Campisi P, Miserere L.,
Perrino E.V., Stmmia S., Iberite M., Lucchese F.,
Fabrini G. & Orsenigo S., 2014. Are Red Lists really
useful for plant conservation? The New Red List of
the Italian Flora in the perspective of national con-
servation policies. Plant Biosystems, 148: 187-190.
Sabella G., 2015. The use of the entomo fauna in the stud-
ies of Environmental Impact Assessment and Eval-
uation of Impact. Biodiversity Journal, 6: 175-184.
Biodiversity Journal, 2015, 6 (1): 175-184
Monograph
The use of the entomofauna in the studies of the Environmen-
tal Impact Assessment (E.I.A.) and Assessment of Impact
(A.I.)
Giorgio Sabella , Oscar Lisi & Fabio Massimo Viglianisi
Dipartimento di Scienze Biologiche, Geologiche e Ambientali, sez. Biologia Animale, Universita di Catania, via Androne 81,
95124 Catania, Italy; e-mail: sabellag@unict.it; olisi@unict.it; fabiovgl@unict.it
’Corresponding author
ABSTRACT The paper highlights the entofauna’s role as not only as an indicator of the environmental
quality, but also as an useful component in the studies of the Environmental Impact Assess-
ment (E.I.A.) and Assessment of Impact (A.I.). Some approaches and tools, with particular
emphasis on Sicily, are proposed in regards to the use of the entomofauna in the assessment
procedures.
KEY WORDS Environmental Impact Assessment; Impact Assessment; Entomofauna; tools; Sicily.
Received 09.01.2015; accepted 19.03.2015; printed 30.03.2015
Proceedings of the 2nd International Congress “Speciation and Taxonomy”, May 1 6th- 1 8th 20 1 4, Cefalu-Castelbuono (Italy)
INTRODUCTION
This paper is a brief methodological review, res-
ulting from the experience gained on several impact
assessments elaborated in the recent years in Sicily.
In Europe there are three different types of en-
vironmental impact assessment: 1) S.E.A. (Stra-
tegic Environmental Assessment), based on the
Directive 2001 /42/EC on the assessment of the ef-
fects of certain plans and programs on the environ-
ment; 2) E.I.A. (Environmental Impact Assessment)
based on the Directive 2014/52/UE, concerning the
impacts assessment of public and private projects
on the environment. The Directive determines the
authorization of certain projects affecting the envir-
onment to an assessment by the competent national
or regional authority. This assessment must identify
the direct and indirect effects of these projects on
the following: human, fauna, flora, soil, water, air,
climate, landscape, material resources and cultural
heritage, and the interaction between these compon-
ents; 3) A.I. (Assessment of Impact), regarding the
assessment of plans and projects significantly
affecting the sites of the Natura 2000 Network.
This evaluation is based on the Directive
92/43/EEC (Habitat Directive), on the conservation
of natural habitats and of wild fauna and flora, and
on the Directive 2009/1 47/EC on the conservation
of wild birds (ex Directive 79/409 EEC). Particu-
larly, the Assessment of Impact is defined and regu-
lated by the article 6, of the Habitat Directive, that
in the third paragraph reads: “Any plan or project not
directly connected with or necessary to manage the
site but likely to have a significant effect, individu-
ally or in combination with other plans or projects,
is subjected to impact assessment on the site respect
to the conservation objectives of this site”. Through
these guidelines the EU seeks to ensure biodiversity
by conserving natural habitats, wild fauna, vegeta-
tion and flora in the territory of the Member States.
176
Giorgio Sabella etalii
In the studies of evaluation of environmental
impact, the analysis of the biotic component is
fundamental to asses the impact generated from
plans and projects; therefore it is obvious that the
fauna is one of the minimum contents required for
the preparation of environmental impact studies,
together with vegetation, flora (see Minissale,
2015) and ecosystems.
THE FAUNA IN THE ENVIRONMENTAL
IMPACT STUDIES
Before delving into the issues related to the
wildlife analysis in the environmental impact
studies, the definition of the fauna's concept is
necessary. According to La Greca (1995), the fauna
is: “A set of species and subspecies of vertebrates
and invertebrates, each divided into one or more
populations, living in a certain territory not captive
or farmed (indigenous species) and included in
natural ecosystems, the presence of which in that
area is due to historical events (paleogeographic
and paleoclimatic), or to evolutionary processes in
situ (autochthonous species or subspecies), or to
indigenation of exotic species”.
The study of wildlife shows numerous and com-
plex problem: a) veiy large number of animals,
especially invertebrates (over 80% of the animals
belong to the phylum Arthropoda, inside of which
more than 75% belong to the class Hexapod); b)
basic knowledge in general unsatisfactory, even for
protected areas; c) difficulty of making a quick
faunistical list of a region, even with small
extension; d) necessity to adopt different and very
specialized sampling methods related to the animals
mobility and the different habitats they occupy; e)
difficulty in developing maps for wildlife.
In relation to these issues, two methodologies
are used for the study of fauna in the environmental
impact studies: 1) Ecosystem approach (review of
certain natural habitats of particular interest in
relation to the associated fauna component); 2) List
of species (for a more detailed discussion of the
topic see Sabella & Petralia, 2012). Really, the
zoologists must interpret the territory as a mosaic
of areas that provide real or potential opportunities
(trophic, reproduction, shelter, etc.) for the various
wildlife species.
The attention of the zoologists involved in the en-
vironmental impact studies, generally, focuses on the
terrestrial vertebrates (especially birds), because it is
the best known component of the wildlife and
responds to the needs to assess the environmental
quality in relation to the targets of the impact studies.
THE USE OF ENTOMOFAUNA: AP-
PROACHES AND TOOLS
The insects are generally poorly used in the en-
vironmental impact studies. This component,
instead, on account of its species richness (more
than 1 million of taxa known heretofore), of its
ubiquitous occurrence and very different diet
(predators, phytophagous, saprophagous, parasites,
pollinators, etc.) and of its diverse and articulated
ecological requirements is suited for the environ-
mental impact studies (Rosenberg et al., 1986). The
study of entomofauna, in many cases, provides
more detailed information on the fine structure and
functioning of the ecosystems and/or allows to
study in more detail the habitats or the microhabit-
ats of particular naturalistic value (dunes and back-
shore, springs, ripicolous environments, soil, caves,
rotting stumps, hollow of old trees, etc.), which are
sometimes very important in the environmental im-
pact studies, and also in the territorial planning and
in the nature conservation policy (Gobbi, 2000).
In relation to the high species number and to the
great diversity of the environments in which they
occur, the study of insects present all the problems
outlined above in a more accentuated, so much so
that, at first glance, it would seem almost impossible
to use the insects in the assessment procedures. For
this reasons guidelines for the use of insects in im-
pact evaluation are, generally, lacking, with excep-
tion regarding the freshwater ecosystems (Adham et
al., 2009; Walters, 2011; Barman, 2014) and the agro
ecosystems (see for example Caoduro et al., 2014).
To do this you have, first, to give up the idea of
establishing a more or less full list of insect species
of a territory, even if small. In any case, this idea is
unworkable for all faunistical studies. But ignoring
entomofauna cannot solve this problem. In the last
years, however, many tools have become available
for use the insects in the environmental management
and also in the impact assessment studies. For a
review of the various sampling methodologies of
entomofauna used in environmental monitoring and
a case study see Burgio et al. (2013).
The use of the entomofauna in the studies of the Environmental Impact Assessment (E.I.A.) and Assessment of Impact (A.I.) 177
It is clear that for assessment studies can be used
only a small fraction of all insect species that occur
in the study area and should be considered those
with conservation problems (IUCN status, inclusion
in international conventions or European directive
annexes) and/or scientific value (endemic, sten-
oecious, at the areal limit, etc.).
Below are briefly treated the main tools usable
for the evaluation of the environmental quality
based on the presence of insects species.
They, substantially, consist of the European,
national and regional red list drafted according to
IUCN criteria, of the annexes of different interna-
tional convention and European directive, of the
checklists (sometimes georeferenced), and of the
Standard Data Form and the Management Plan of
the Natura 2000 site.
SPECIES INCLUDED IN THE ANNEXES
OF DIRECTIVE 43/92 EEC
Insects species of Annex II to Directive 43/92 EEC
present in Sicily
The taxa listed in Annex II are named as “ Com-
munity interest species whose conservation
requires the designation of special areas of conser-
vation''’ (with an asterisk priority species are
indicated). All these species are very important for
Assessment of Impact because it is mandatory to
take them into account and considering the possible
negative effects induced by the territory's trans-
formation linked to realize a project. Only if you
can exclude negative effects on these species, or in
the presence of negative impacts proposing effect-
ive mitigation measures, it is possible to give a
positive evaluation of the environmental compatib-
ility of the project.
The insect species included in the Annex II to
Directive 43/92 EEC present in Sicily are show in
Table 1 , and briefly commentated on below, em-
phasizing the most important threat factors to
consider for their conservation.
Coenagrion mercuriale (Charpentier, 1 840)
The larvae live in streams, usually on limestone
substrates. Sicilian populations (Fig. 1) are very loc-
alized. The species is rare in Italy and must be con-
sidered vulnerable. The major threats are: river
straightening, water harnessing, swamps and soil
drainage, water table lowering through irrigation,
field destruction or conversion into other agricul-
tural practices, water pollution (IUCN, 2014).
ODONATA
Coenagrionidae
Coenagrion mercuriale (o) - in Italy C. mercuriale castellani
Roberts, 1948
Cordulegastridae
Cordulegaster trinacriae
ORTHOPTERA
Gryllidae
Brachytrupes megacephalus
Myrmecophilus haronii - only Pantelleria island
COLEOPTERA
Lucanidae
Lucanus cerx’us (?) (o)
Geotrupidae
Bolbelasmus unicornis - in Sicily B. romanorum
Cetoniidae
* Osmoderma eremita - in Sicily O. cristinae
Cerambicidae
Cerambyx cerdo
*Rosalia alpina
LEPIDOPTERA
Arctiidae
*Callimorpha quadripunctaria (now Euplagia ) (o)
Satyridae
Melanargia arge
Table 1. Insects species of Annex II to Directive 43/92 EEC present in Sicily. All species listed in Annex II are present also
in Annex IV excluding those followed by symbol (o). The symbol * highlights the priorities species, while the symbol (?)
indicates the uncertain presence of the species in Sicily.
178
Giorgio Sabella etalii
Cordulegaster trinacriae Waterston, 1976
The larvae live in clean streams with sandy
bottom, shaded by tree vegetation. The species is
threatened by chemical and physical water pollution,
by water extraction for human use and by removal of
riparian vegetation. Desiccation due to climate change
is a further threat for this species (IUCN, 2014).
Brachytrupes megacephalus (Lefevre, 1 827)
Large cricket that lives in dune and back-dune
environments, showing strong burrowing habits.
It builds a long one -meter burrow using a spectacular
technique of excavation. The species (Fig. 2) is
threatened by habitat changes due to agricultural
practices and touristic exploitation of beach (see
Petralia et al., 2015).
Myrmecophilus baronii Baccetti, 1966
Endemic species to Pantelleria Island. It is a
mirmecophilous Grillidae generally enfeoffed with
the ants of the genus Lasius Fabricius, 1804.
Lucanus cervus cervus (Linnaeus, 1758)
The presence of this taxon in Sicily is to be con-
firmed. It lives in forests of oak and chestnut,
sometimes, on the trunks and branches of willows
and mulberries. The female lays the eggs at the foot
of the trees; the larvae feed on humus and then
penetrate into the trunk, but generally they dig their
tunnels in the stumps remaining in ground and their
development requires up to 5 years. The species is
threatened by the coppicing of the forests and
cleanliness of the undergrowth. The taxon not yet
assessed for the IUCN Red List. In Sicily is
certainly presents L. tetraodon sicilianus Planet,
1899, showing similar ecological requirements but
is not listed in any Annex to the Habitats Directive.
Bolbelasmus romanorum Amone et Massa, 2010
Bolbelasmus unicornis (Schrank, 1789) is in-
cluded in Annex II of the Habitats Directive and also
B. romanorum, endemic to Sicily, should be inserted
into Annex II. B. romanorum is relatively rare and
localized species. Its biology is still poorly known.
It can be occasionally observed wandering on the
ground or under stones and during crepuscular flight.
The taxon not yet assessed for the IUCN Red List.
*Osmoderma cristinae Sparacio, 1994
The genus Osmoderma Le Peletier de Saint-
Fargeau et Serville, 1828 includes species very
sensitive to environmental changes and everywhere
in rarefaction. O. eremita (Scopoli, 1763) is included
in Annex II of the Habitats Directive as priority
species. Even O. cristinae (Fig. 3), endemic to
northwestern Sicily, and O. italicum Sparacio, 2000,
endemic to Central and Southern Italy, with a
biology entirely comparable with that of O. eremita,
should be inserted into Annex II and regarded as
priority species. O. cristinae is a silvicolous species.
The larvae develop in old rotting trunks of oaks or
maples. The main overall threat is likely to be
degradation or loss of habitat quality, involving
structural changes in the tree populations arising
from changing land use - affecting age structures
and trees density. Exploitation from forestry is often
a key immediate issue, but equally damaging can be
long-term changes towards canopy closure and loss
of old trees as a result of non or minimum interven-
tion management systems which all too often ex-
clude grazing by large herbivores. Fragmentation
and increasing isolation of beetle populations are
also key factors. The restricted area of occupancy
combines limited population size with reduced hab-
itat availability, bird predation, fires, and frequently
unsuitable local techniques of forest management
(Audisio et al., 2007).
Cerambyx cerdo cerdo (Linnaeus, 1758)
Silvicolous species. The adult feeds on leaves,
fruit and lymph and actively flies in the twilight
hours. After mating, which occurs between June and
August, the female lays her eggs in the cracks of the
bark of big oak trees. The saproxylic larva begins to
dig tunnels in the cortical layers and then penetrates
into the wood and its development requires 3-4
years. It is a species threatened by coppicing of oak
trees and by the removal of old decaying plants.
* Rosalia alpina (Linnaeus, 1758)
The species (Fig. 4) lives preferentially in mature
forests with a predominance of beech, especially
The use of the entomofauna in the studies of the Environmental Impact Assessment (E.I.A.) and Assessment of Impact (A.I.) 179
those characterized by very rainy or oceanic climate.
Adults are active during the day on logs felled or
inflorescences of Umbrelliferae. After mating, eggs
are laid in the wood; the larval development takes 3
years, and it is preferably done in dead or decayed
wood of beech exposed to the sun. The larva can
develop also on alder, ash, hawthorn, linden and
maple or conifers. The species is threatened by
excessive cleaning the forest area; perhaps even by
air pollution and by the general contraction of beech
forests, especially mature ones.
*Euplagia quadripunctaria (Poda, 1761)
The only European species of this genus. It can
be found in the cool forests and, in the Mediter-
ranean region, most often in narrow valleys bounded
by mountains with steep slopes with perennial
streams and continuous woodlands, characterized by
a microclimate cooler and wetter than the surround-
ing areas. Adults have primarily nocturnal habits and
spending the day in the dense vegetation. The larvae
emerge after 8-15 days after spawning and feed on
various plants for a short time (like several Ros-
aceae, and other species such as black locust and
eastern plane tree, vines and mulberry trees, honey-
suckle) then they go into hibernation. After the 5th
molt, the caterpillar spins a slight cocoon in the litter.
The pupal stage lasts about 1 month; the imago
emerges between June and August, most often in
July, according to the altitude and the seasons.
Melanargia arge (Sulzer, 1776)
The species is distributed in peninsular Italy and
northern Sicily. Its habitat is represented by arid
steppes with scattered bushes and isolates trees with
outcropping rocks. Most of the sites are located in
the valleys sheltered from the wind or in hilly areas
inland. The fires favored by shepherds and the
Figure 1. Coenagrion mercuriale castellani, Sicily, Palermo, stazione Montemaggiore Belsito, 30.IV.2010. Figure 2.
Brachytrupes megacephalus, Sicily, Trapani, Capo Feto, 1.V.2011. Figure 3. Osmoderma cristinae, Sicily, Madonie
Mountains, Gibilmanna, 2.VII.2014. Figure 4. Rosalia alpina, Nebrodi Mountains, Biviere di Cesaro, 6.VII.2014 (Photos
by C. Muscarella).
180
Giorgio Sabella etalii
ORTHOPTERA
TETTIGONIIDAE
Saga pedo
LEPIDOPTERA
PAPILIONIDAE
Papilio alexanor
Parnassius apollo
Parnassius mnemosyne
Zerynthia polyxena
SPHINGIDAE
Proserpinus proserpina
Table 2. Insects species of Annex IV to Directive 43/92 EEC not listed in Annex II, present in Sicily.
overgrazing can have serious negative effects on
this species along with other habitat alterations.
This species is not believed to face major threats at
the European level.
Insects species of Annex IV to Directive 43/92
EEC not listed in Annex II, present in Sicily
The taxa listed in Annex IV are named as
“Community interest species in need of strict
protection”.
Most of the species listed in Annex II are also
mentioned in Annex IV, so in Table 2 are shown
only the insects species present in Sicily and
listed in Annex IV, but not in Annex II. They are
briefly commentated on below, emphasizing the
most important threat factors to consider for their
conservation.
Saga pedo (Pallas, 1771)
Species distributed from central-southern and
southeastern Europe to central Asia and north-
western China. In Italy it is present in a few areas
of the Alps and Apennines, Sardinia and Sicily.
Saga pedo colonizes areas with more or less open
herbaceous vegetation or shrubs. It can be observed
on the ground or on bushes, where moves rather
slowly. Predator species, feeding mainly on other
Orthoptera (grasshoppers and locusts) that captures
thanks to the long and strong forelegs armed with
spines. Never common in areas where it is present,
is threatened by habitat degradation.
Papilio alexanor Esper, 1800
This butterfly is mostly found on warm and dry
calcareous slopes with flower-rich vegetation and
low-growing bushes. It prefers slopes that are steep
and rocky and it is especially active during the
hottest hours of the day. Different foodplants are
known, all of them umbellifers. Although this
species shows a decline in a part of its European
range, it is not believed to face major threats at the
European scale.
Parnassius apollo (Linnaeus, 1758)
Species widely distributed in the mountains of
Western Europe and Southern and Fennoscandia,
although it is extinct in some areas such as central
Germany, Czechoslovakia and Denmark and it is
absent in Britain. In Sicily, the species is at the
southern limit of its distribution range and is
extremely localized, it is in fact known only from a
few stations on the Madonie Mountains and accor-
ding to some authors belongs to a subspecies P.
apollo siciliae Oberthiir, 1899 (Fig. 5). Parnassius
apollo is linked to stony and mountainous areas
poor of vegetation. It shows a preference for cal-
careous soils and for some plants such as Cardus
spp., Cirsium spp., Origanum spp., Centaurea spp.,
Scabiosa spp. and Knauzia spp.
Parnassius mnemosyne (Linnaeus, 1758)
In Central Europe the species (Fig. 6) lives in
hill and mountain areas up to 1,500 m of altitude,
in Northern Europe in plain areas. In Italy he
attends the clearings and the edges of deciduous
forest (beech, turkey oak). Adults are attracted to
many vegetal species, with a preference for red,
purple and blue flowers as Centaurea spp., Knauzia
spp., Geranium spp. and Lychnis spp. The two main
The use of the entomofauna in the studies of the Environmental Impact Assessment (E.I.A.) and Assessment of Impact (A.I.) 181
causes of its decline are the reforestation and the
changes in traditional agricultural practices, which
have caused the disappearance of many meadow
areas.
Zerynthia polyxena (Denis et Schiffermuller, 1775)
The only Italian species of this genus. It attends
the plain near wetlands, the hilly and mountainous
areas with arid terrain or rocky areas up to 900 m.
It has a single annual generation, usually adults
appear in April-May, but in Sicily may be active
already at the end of February. The caterpillars feed
on various species of Aristolochia L. The disappear-
ance of this species, observed throughout Europe,
is due to the reforestation and the habitat destruc-
tion. Locally may be threatened by excessive
collection.
Proserpinus proserpina (Pallas, 1772)
The only European species of the genus. It lives
from the sea level up to 1 ,500 m in different biotopes
such as valleys, forest edges, clearings and banks of
the streams, in rich sites of Epilobium angustifolium
L. Adults are primarily nocturnal and prefer nectar-
rich flowers, such as the common oregano, several
species of fireweed, wild pink and honeysuckle. The
species has disappeared from many localities in re-
cent times, but the causes are not known. Some po-
pulations seem to disappear for a few years and
reappear suddenly, for no apparent reason.
SPECIES INCLUDED IN THE RED LIST
BASED ON IUCN CRITERIA
One other very useful tool is represented of the
Red Lists based on IUCN criteria (IUCN, 2012).
On the site http://www.iucnredlist.org/ can be check
the Red List of Threatened Species, which are men-
tioned all insect species considered threatened at the
global level. For each of them, informations on tax-
onomy, assessment, geographic range, population,
habitat and ecology, and major threats are provided.
However, there are European red lists among which
are to mention those of saproxylic Coleoptera
(Nieto & Alexander, 2010), of butterfly (Van Swaay
et al., 2010), and of dragonflies (Kalkman et al.,
2010). Also to be mentioned some national red lists
such as those on Italian invertebrates (Cerfolli et al.,
Figure 5. Parnassius apollo, Sicily, Madonie Mountains,
Pizzo Carbonara, 15.VII.2006. Figure 6. Parnassius mnemo-
syne, Sicily, Madonie Mountains, Piano Battaglietta,
12.VI.2012 (Photos by C. Muscarella).
2002), on butterflies (Prola & Prola, 1990) and the
recent red lists of Italian saproxylic Coleoptera
(Audisio et al., 2014) and Italian dragonflies
(Riservato et al., 2014). For Sicily, currently, there
are not regional red lists of insects, the only work,
that concerns only Coleoptera and Lepidoptera, is
a list of species present within the Regional Parks,
in which, for each species, informations on assess-
ment, geographic distribution, and habitat are
provided (Sabella & Sparacio, 2004).
SPECIES LISTED IN THE ENTOMOLEX
A very useful tool, drawn up under the auspices
of the Italian Entomological Society, is represented
182
Giorgio Sabella etalii
by Entomolex (Ballerio, 2004). It is a review that
aims to provide an overview of all the rules concern-
ing the conservation of the Italian insects. For each
mentioned species is considered its inclusion in the
annexes of international conventions (Conventions
of Washington and Bern), of EU legislation
(Directive 43/92 EEC), and of national and regional
(Regions Friuli Venezia-Giulia, Liguria, Lombardy,
Piedmont, Tuscany, and Veneto and the autonomous
provinces of Trento and Bolzano) laws.
CKMAP OF ITALIAN FAUNA, FAUNA
D’lTALIA AND REGIONAL CATALOGUE
The CKmap project (Ruffo & Stoch, 2005) and
its database have made available information on the
punctual distribution in Italy of approximately
10,000 terrestrial and freshwater species selected
from the checklist because protected, threatened,
with scientific or biogeographical interest, or bioin-
dicators. The project represents an important tool
for a correct and scientific management of the biod-
iversity and the natural habitats, as from Checklist
of the Italian fauna (Minelli etal., 1993-1995), that
comprises about 55,000 species. In the CKmap, for
each species, geo-referenced data of the Italian
localities, of the distribution, of the ecology, and its
value as a bioindicator are provided. The analysis
of so large sample of species has allowed to identify
the most important areas in terms of the number of
species, of the concentration of endemic species, of
the species with restricted distribution and/or of
particular biogeographical interest. All that permit
to draw a picture of the overall distribution of
animal biodiversity in Italy with a level of accuracy
and detail unthinkable a few years ago.
The CKmap can provide, therefore, detailed
information on many species of Sicilian entomo-
fauna, and must be integrated with the many mono-
graphes of the series "Fauna d'ltalia" dedicated to
insects, some regional checklist (see for example
Pilato et al., 2007 for Iblean region) and many
regional faunistic catalogs concerning various taxo-
nomic groups such as Ephemeroptera (Belfiore et
al., 1991), Plecoptera (Fochetti & Nicolai, 1987;
Ravizza & Gerecke, 1991), Neuroptera (Pantaleoni,
1986), Coleoptera Cerambycidae (Sanaa &
Schurmann, 1980), Coleoptera Staphylinidae
(Sabella & Zanetti, 1991), Coleoptera Pselaphidae
(Sabella, 1998), Coleoptera Tenebrionidae (Aliquo
& Soldati, 2010; Aliquo & Soldati, 2014), etc. Of
course many other citations of Sicilian insect
species are dispersed in numerous scientific public-
ations, it would be better to know, but the use of the
tools suggested previously can be deemed suffi-
cient to estimate the environmental quality of an
area and assess the impact of the implementation of
a project.
STANDARD DATA FORM AND MANAGE-
MENT PLAN OF NATURA 2000 SITES
At each site Natura 2000 is associated a standard
data form, available, for Italian sites, on the official
website of the Ministry of Environment and Protec-
tion of Land and Sea (http://www.minambiente.it/
pagina/schede-e-cartografie). The standard data
form, which is still required in an Annex when
processing a report of Impact Assessment, listing
all habitats and species of Community interest
whose conservation requires the designation of
special areas of conservation (for the invertebrates
see the section 3.2.f.) and also all other important
species (see the section 3.3), because listed in the
national red list (motivation A), endemics (motiva-
tion B); included in the international conventions
(motivation C) or for other reasons (motivation D).
In regards to the Sicilian Region, on the official
website of the Regional Ministry of Land and
Environment (http://www.artasicilia.eu/old_site/
web/natura2000/index.html) most of the Manage-
ment Plans of the Sicilian Natura 2000 sites are
available and downloadable in pdf format. In these
plans can be find detailed information on the animal
species including their distribution in the site
habitats and their ecological requirements.
BRIEF CONCLUSIVE CONSIDERATIONS
This paper attempts to emphasize the import-
ance of the insect fauna study in environmental
impact assessment and more generally in the ter-
ritory planning and nature conservation.
The numerous problems related to the study of
the entomofauna should not discourage, because by
a reasonable approach, it can get to a list of species
that, far from being exhaustive, may represent a
The use of the entomofauna in the studies of the Environmental Impact Assessment (E.I.A.) and Assessment of Impact (A.I.) 1 83
good basis for the assessment, in terms of fauna, of
the environmental quality and thus to assess any im-
pacts. In another article in this volume, a study case,
in which were used the approach and tools previou-
sly treated, is proposed (Sabella et al., 2015).
ACKNOWLEDGEMENTS
We are grateful to Calogero Muscarella
(Palermo, Italy) for the photos.
REFERENCES
Aliquo V. & Soldati F., 2010. Coleotteri Tenebrionidi di
Sicilia. Edizioni Danaus, Palermo, 176 pp.
Aliquo V. & Soldati F., 2014. Updating the CD-rom on
Coleoptera Tenebrionidae of Italy and check-list of
the same family. Biodiversity Journal, 5: 429^442.
Audisio P., Brustel H., Carpaneto G. M., Coletti G., Man-
cini E., Piattella E., Trizzino M., Dutto M., Antonini
G. & De Biase A., 2007. Updating the taxonomy
and distribution of the European Osmoderma, and
strategies for their conservation (Coleoptera, Scara-
baeidae, Cetoniinae). Fragmenta entomologica, 39:
273-290.
Audisio P., Baviera C., Carpaneto G.M., Biscaccianti
A.B., Battistoni A., Teofili C. & Rondinini C.
(compilatori), 2014. Lista rossa IUCN dei Coleotteri
saproxilici Italiani. Comitato IUCN e Ministero
dell’Ambiente e della tutela del Territorio e del Mare,
Roma, 132 pp.
Adham F.H., Gabre R.M. & Ibrahim I. A., 2009. Some
aquatic insects and invertebrates as bioindicators for
the evaluation of bacteriological pollution in
El-Zomor and El-Mariotya canals, Giza, Egypt.
Egyptian Academy Journal of Biological Sciences, 2:
125-131.
Ballerio A., 2004. La conservazione degli insetti e la
legge.[4° aggiornamento: 30 giugno 2004],
http ://www. societaentomologicaitaliana.it/it/servizi/e
ntomolex.html.
Barman B., 2014. The Importance of Aquatic Insects as
Biomonitors of Freshwater Ecosystem. International
Journal of Environment and Natural Sciences, 1:
82-85.
Bern Convention, 1979 on the Conservation of European
Wildlife and Natural Habitats.
Belfiore C., D’ Antonio C., Audisio P., Scillitani G., 1991.
Analisi faunistiche e biogeografiche sugli Efemerot-
teri della Sicilia (Insecta, Ephemeroptera). Animalia,
18: 3-60.
Burgio G., Baldacchino F, Magarelli A., Masetti A.,
Santorsola S. & Arpaia S. (a cura di), 2013. II
campionamento dell’artropodofauna per il monitor-
aggio ambientale. Applicazioni per la valutazione
dell’impatto ambientale delle Piante Geneticamente
Modificate. ENEA, 137 pp.
Caoduro G., Battiston R., Giachino P.M., Guidolin L. &
Lazzarin G., 2014. Biodiversity indices for the as-
sessment of air, water and soil, quality of the “Biod-
iversity Friend” certification in temperate areas.
Biodiversity Journal, 5: 69-86.
Cerfolli F., Petrassi F. & Petretti F. (a cura di), 2002.
Libro rosso degli animali d’ltalia. Invertebrati. WWF
Italia Onlus, 83 pp.
Directive 79/409 EEC of the European Parliament and
of the Council of 2 April 1979, on the conservation
of wild birds. Official Journal of European Union L
103,25/04/1979.
Directive 92/43/EEC of the European Parliament and of
the Council of 21 May 1992, on the conservation of
natural habitats and of wild fauna and flora. Official
Journal of European Union L 206, 22/07/1992.
Directive 2001 /42/EC of the European Parliament and of
the Council of 27 June 2001, on the assessment of the
effects of certain plans and programmes on the en-
vironment. Official Journal of European Union L
197,21/07/2001.
Directive 2009/ 147/EC of the European Parliament and
of the Council of 30 November 2009, on the conser-
vation of wild birds. Official Journal L 20/7,
26/01/2010.
Directive 2014/52/UE of the European Parliament and of
the Council of 16 April 2014, amending Directive
2011/92/EU on the assessment of the effects of
certain public and private projects on the environ-
ment. Official Journal of European Union L 124/1,
25/04/2014.
Fochetti R. & Nicolai R, 1987. Plecotteri di Sicilia e
Sardegna. Animalia, 14: 169-175.
Gobbi G., 2000 . Gli artropodi terrestri e la tutela degli
ecosistemi in Italia. II Naturalista Siciliano, 24: 1 89—
223.
http://www.minambiente.it/pagina/schede-e-cartografie
http://www.artasicilia.eu/old_site/web/natura2000/index.
html
Kalkman V.J., Boudot J.P., Bernard R., Conze K.J., de
Knijf G., Dyatlova E., Ferreira S., Jovic M., Ott J.,
Riservato E. & Sahlen G. (compilators), 2010.
European Red List of Dragonflies. Luxembourg: Pub-
lications Office of the European Union, vii + 28 pp.
IUCN, 2012. Red List Categories and Criteria: Version
3.1. Second edition. Gland, Switzerland and
Cambridge, UK: IUCN, iv + 32 pp.
IUCN, 2014. Red List of Threatened Species 2014.3.
http://www.iucnredlist.org/.
La Greca M., 1995. II concetto di fauna e le caratteristi-
che della fauna italiana. XX giornata dell'ambiente".
184
Giorgio Sabella etalii
Atti del Convegno dell’Accademia Nazionale dei
Lincei, 118: 13-28.
Legge Provinciale n. 16 del 27 luglio 1973. Norme per
la tutela di alcune specie della fauna inferiore.
Bollettino Ufficiale della Regione Trentino-Alto
Adige n. 34 del 7 agosto 1973.
Legge Provinciale n. 27 del 13 agosto 1973. Norme per
la protezione della fauna. Bollettino Ufficiale della
Regione Trentino-Alto Adige n. 39 dell’ 1 1 settembre
1973.
Legge Regionale n. 53 del 15 novembre 1974. Norme
per la tutela della fauna inferiore e della flora. Bol-
lettino Ufficiale della Regione Veneto n. 47 del 1974.
Legge Regionale n. 33 del 27 luglio 1977. Prowedimenti
in materia di tutela ambientale ed ecologica. Bollet-
tino Ufficiale della Regione Lombardia n. 30,
supplemento ordinario del 29 luglio 1977.
Legge Regionale n. 34 del 3 giugno 1981. Norme per
la tutela della natura e modifiche alia legge re-
gionale 27 dicembre 1979 n. 78. Bollettino Ufficiale
della Regione Friuli Venezia Giulia n. 63 del 3
giugno 1981.
Legge regionale n. 32 del 2 novembre 1982. Norme per
la conservazione del patrimonio naturale e delf
assetto ambientale. Bollettino Ufficiale della Regione
Piemonte n. 45 del 10 novembre 1982.
Legge Regionale n. 36 del 15 dicembre 1992. Modifica
ed integrazione alia Legge Regionale 22 gennaio
1992 n. 4. Bollettino Ufficiale della Regione Liguria
n. 21 del 23 dicembre 1992.
Legge Regionale n. 56 del 6 aprile 2000. Norme per la
conservazione e la tutela degli habitat naturali e
seminaturali, della flora e della fauna selvatiche -
Modifiche alia legge regionale 23 gennaio 1998, n. 7
- Modifiche alia legge regionale 1 1 aprile 1995, n. 49.
Bollettino Ufficiale della Regione Toscana n. 17 del
17 aprile 2000.
Minelli A., Ruffo S„ La Posta S. (Eds.), 1993-1995.
Checklist delle specie della fauna italiana. Calderini.
Minissale P., 2015. The Use of Flora, Vegetation and Ha-
bitats in the Studies of Environmental Impact As-
sessment. The International Congress on speciation
and taxonomy, May 16th- 18th 2014 Cefalu-
Castelbuono (Italy). Biodiversity Journal, 6: 171-174.
Nieto A. & Alexander K.N.A. (compilators), 2010.
European Red List of Saproxylic Beetles. Luxembourg:
Publications Office of the European Union, vii + 45 pp.
Pantaleoni R.A., 1986. Neurotteri dell’Italia meridionale
ed insulare. Animalia, 13: 167-183.
Petralia A., Petralia E., Sabella G., Brogna F. & Bianca
C., 2015. Presence's mapping of Brachytrupes mega-
cep halus (Orthoptera, Gryllidae) within the Natural
Reserve of Vendicari (Noto, SR, Italy). The Interna-
tional Congress on speciation and taxonomy, May
May 16th- 18th 2014 Cefalu-Castelbuono (Italy).
Biodiversity Journal, 6: 323-326.
Pilato G., Sabella G., Turrisi F., Bella S., Scuderi D. &
Lisi O., 2007. La fauna della regione iblea. Atti del
Convegno “L’Uomo negli Iblei”, Sortino 10-12
ottobre 2003: 51-116.
Prola G. & Prola C., 1990. Libro rosso delle farfalle
italiane. Quaderni WWF 13, 71 pp.
Ravizza C. & Gerecke R., 1991. A review of the distri-
bution of Plecoptera on Sicily. Memorie della Societa
Entomologica Italiana, 70, 2: 9-31.
Riservato E., Fabbri R., Festi A., Grieco C., Hardersen
S., Landi F., Utzeri C., Rondinini C., Battistoni A. &
Teofili C. (compilatori), 2014. Lista rossa IUCN delle
libellule Italiane. Comitato IUCN e Ministero
dell’Ambiente e della tutela del Territorio e del Mare,
Roma, 39 pp.
Rosenberg D.M., Danks H.V. & Lehmkul D.M., 1986.
Importance of Insects in Environmental Impact As-
sessment. Environmental Management, 10: 773-783.
Ruffo S. & Stoch F. (Eds.), 2005. Checklist e dis-
tribuzione della fauna italiana. Memorie del Museo
Civico di Storia Naturale di Verona, 2a serie, Sezione
Scienze della Vita, 16.
Sabella G., 1998. Pselafidi di Sicilia. Monografie del Museo
Regionale di Scienze Naturali, Torino, 25: 1M16.
Sabella G. & Petralia E., 2012. Zoological aspects of the
assessment of human impact on environments. 2nd
Djerba International Mediterranean Environments
Sustainability Conference 22-25 April 2012. Atti e
Memorie dell’Ente Fauna Siciliana, 11: 171-181.
Sabella G. & Sparacio I., 2004. II ruolo dei Parchi
siciliani nella conservazione di taxa di insetti di
particolare interesse naturalistico (Insecta Coleoptera
et Lepidoptera Ropalocera). II Naturalista siciliano,
28: 477-508.
Sabella G. & Zanetti A., 1991. Studi sulle comunita di
Coleotteri Stafilinidi dei monti Nebrodi. 1° con-
tributo. Animalia, Catania, 18: 269-297.
Sabella G., Alicata A. & Viglianisi F.M., 2015. A study
case of Assessment of Impact using the invertebrates.
The International Congress on speciation and tax-
onomy, May May 16th- 18th 2014 Cefalu-Castel-
buono (Italy). Biodiversity Journal, 6: 185-192.
Sama G. & Schurmann O., 1980. Coleotteri Cerambicidi
di Sicilia. Animalia, 7: 189-230.
Van Swaay C., Cuttelod A., Collins S., Maes D., Lopez
Munguira M., Sasic M., Settele J., Verovnik R.,
Verstrael T., Warren M., Wiemers M. & Wynhof
I. (compilators), 2010. European Red List of
Butterfies. Luxembourg: Publications Office of the
European Union, viii + 47.
Walters A.W., 2011. Resistance of aquatic insects to low
flow disturbance: exploring a trait-based approach.
Journal of the North American Benthological Society,
30: 346-356.
Washington Convention, 1973 on International Trade in
Endangered Species of Wild Fauna and Flora.
Biodiversity Journal, 2015, 6 (1): 185-192
Monograph
A study case of Assessment of Impact using the invertebrates
Giorgio Sabella", Antonio Alicata & Fabio Massimo Viglianisi
Dipartimento di Scienze Biologiche, Geologiche e Ambientali, sez. Biologia Anim ale, Universita di Catania, via Androne 81,
95124 Catania, Italy; e-mail: sabellag@unict.it; antonioalicata@g mail. com; fabiovgl@unict.it
Corresponding author
ABSTRACT A study case of Assessment of Impact (A. I.) in regards to the project of achieving diaphragm
containment for homogeneous areas T and V of the Gela Refinery is explained. The inver-
tebrates were used to evaluate the environmental quality and also to identify appropriate and
effective mitigation measures and for preparing a post-operam monitoring. Some methodo-
logical proposals and an index of faunistic habitat value have been proposed.
KEY WORDS Assessment of Impact; Invertebrates; Sicily; Faunistic value index.
Received 29.11.2014; accepted 21.02.2015; printed 30.03.2015
Proceedings of the 2nd International Cong re ss “Speciation and Taxonomy”, M ay 1 6 th- 1 8 th 20 14, Cefalit-Castelbuono (Italy)
INTRODUCTION
The article 6 of the Directive 92/43 EEC estab-
lishes the rules, which govern and regulate the con-
servation and management of the Nature 200 0
network sites, and determines the guidelines to be
adopted by the member states for proper relation-
ship between the protection of natural resources and
the land use. In particular, the paragraphs 3 and 4
establish procedures governing the approval of
plans or projects that insist on SCI or SPA, and not
directly related to their management. Essentially,
any transform at ion that interests a Natura 2000 site,
as well as areas adjacent thereto must be subjected
to a procedure for Assessment of Impact, which
excludes negative effects on the site, or, if it recog-
nizes them, proposes corrective measures (mitiga-
tion or compensation).
The realization of a diaphragm containment of
some areas of the Gela Refinery (Sicily), fell back
within the perimeter of the SCI and S PA ITA050001
- Biviere and Macconi of Gela. Therefore, in com-
pliance with the requirements of the aforemen-
tioned legislation, the project proposer has decided
to proceed to the elaboration of the Assessment of
Impact to verify if the project could have the
adverse effects on habitats and species in Annexes
I and II Directive 92/43 EEC and species of Annex
I to Directive 2009/147 EC of the Natura 200 0 site.
The project involved the construction of a bar-
rier to excavation with composite diaphragm (self-
hardening mud and HDPE sheet) associated to a
system of pumping wells of groundwater, already
pre-existing for much, built upstream of the
diaphragm. For its realization the excavation of a
trench, about 1 meter wide, 25 meters on average
deep, and about 2.5 kilometers long was foreseen,
and so effects on soil fauna, which concerns
substantially invertebrates, were expected.
Although invertebrates are little used in environ-
mental impact assessments (Sabella et al., 2015), in
this case, for project evaluation their study was
necessary, given their importance in determining
the composition and structure of the soil fauna. For
this reason, at the study of terrestrial Vertebrates it
is added that of the invertebrates, with particular
attention to the Insects.
186
Giorgio Sabella etalii
MATERIAL AND METHODS
Study area
The study area includes a territory in which it is
believed, on the basis of the project data, are pos-
sible impacts on wildlife induced by its realization.
The area is located in the district of “Piana del
Signore”, in the municipality of Gela, within the
larger territory of "Piana di Gela". It is bordered to
west by the Priolo Channel, to the east by the New
Priolo Channel and is between the coast and the
south side of the Gela Refinery (Fig. 1).
Sampling and analysis
The species list refers to the study area identified
in figure 1. The annotated catalogue of the ter-
restrial Vertebrates was based on the Nature 2000
site's Standard Data Form, and also on literature
references believed to be accurate, on personal
observations and/or on the presence of potentially
suitable habitat for the species. The annotated cata-
log of theArthropods was based on the Natura 20 0 0
site's Standard Data Form, on literature references
believed to be accurate, on a faunistic sampling
campaign, with a monthly intervals, from June to
November, with various techniques (collection on
view, mowing, and sifting). For the purposes of an
biocoenotic investigation on soil fauna, was also
used the method of pit-fall traps, which allowed
to sample many species, not detected by other
sampling methods.
For each species were reported data on: 1) sci-
entific name, author and year, according to the
nomenclature adopted by the check-list of Italian
fauna (Minelli et al., 1993-1 995) and Ckmap of
Italian fauna (Ruffo & Stoch, 2005), considering
Figure 1. Study area. Red line: perimeter of the Natura 2000 site; in green: study area; in yellow: remediation area of basin
A zone 2 of Gela Refinery, Sicily; in gray: affected area in the project of the diaphragm containment for homogeneous
areas T and V of the Gela Refinery.
A study case of Assessment of Impact using the invertebrates
187
the subsequent changes in the nomenclature of the
recent literature; 2) the chorologic category, accord-
ing to Vigna Taglianti et al. (1992, 1 999), while for
birds according to Brichetti (1997); 3) the habitats
potentially utilized by the species in the area; 4) if
known, the phenology of the species; 5) the trend
of European and Italian populations.
Particular attention was given to measures of
protection and conservation of which the species
is the subject, indicating its presence in the fol-
lowing annexes:
- II (strictly protected species of fauna) and III
(protected fauna species) of the Berne Convention,
law 5 August 1981 n. 503, on the Conservation of
European Wildlife and NaturalHabitats in Europe;
- I (endangered migratory species) and II
(migratory species to be the subject of agreements)
of the Bonn Convention, law 25 January 1 983 n.
42, on the Conservation of migratory species of
w ild an im als ;
-A (species threatened with extinction which are
ormay be an action of the trade) and B (species not
necessarily threatened with extinction at the present
time, but that may become so unless trade is not
subject to regulation close) of the Washington
Convention, law 19 December 1975 n. 874, on
international trade in animal and plant species
threatened with extinction (CITES) and subsequent
amendments and additions;
- II (animal and plant species of Community in-
terest whose conservation requires the designation
of special areas of conservation), IV (animal and
plant species of Community interest in need of strict
protection) and V (animal and plant species of
Community interest whose taking in the wild and
exploitation may be subject to management meas-
ures) of EEC Directive 92/4 3, D.P.R. 8 September
1997 n. 357, on the conservation of natural habitats
and of wild fauna and flora in Europe.
As for the birds, for each species was specified
inclusion in the Annexes (I, II/A , II/B , III/A and
II I/C of the Directive EC 2009/147 and the conser-
vation status according the Species of European
Conservation Concern of Birdlife International,
2 0 04 (SPEC1, SPEC2, SPEC3, Non-SPEC E and
Non-SPEC).
For Mammals and Birds species, their possible
protection established by the law 11 February 1992,
n. 157 (rules for the protection of homeotherme
wildlife and for hunting) and their inclusion in
article 2, which provides for such species specific
protective measures, was also considered.
The species conservation status, inferred by the
website IUCN 2014 and by the various national
(Prola & Prola, 1990; C erfo lli et al., 20 0 2;
Rondinini et al., 2013; Audisio et al., 2014; Riser-
vato et al., 2014) and regional (AA.VV., 2008) red
lists, based on IUCN criteria (IUCN, 2012), was
also indicated.
Species of Annex II to Directive 43/92 EEC = 1.5 +0.50 if prior itary
Species of Annexes to international convention, or to national or regional laws — 1
Species included in national red lists based on IUCN criteria
M CR = 1
■ EN = O.SO
*vu = 0.60
■ NT = 0.40
■ LC = 0.20
Species of binge ographical interest
■ Sicilian endemic species = 1+ 0.20 if present only in southern Sicily
■ Sicilian endemic subspecies =0.50 +0.20 if present only in southern Sicily
■ Species with restricted distribution — 0.50
* Species at the northern end of their distribution = 0.70
■ Species which in Italy are present only in Sicily — 0.50 (1.00 if in Europe are present only in Sicily)
■ Species at the southern end of their distribution — 0.50
Species relevant for ecological aspects
■ Stenotops or stenoecious = 0.70
■ Susceptible to human disturbance — 0.70
0 Localized — 0.50
0 Restricted populations =0,50
Table 1. Criteria used for the faunistic value attribution to the invertebrats species.
188
Giorgio Sabella etalii
A faunistic value (see for example M assa &
Canale, 2008) to each species is assigned, for inver-
tebrates it was based on the criteria showed in
Table 1. If a species fell within in more than cat-
egories, the values were summed. Within the study
area, based on the vegetation and the land use
Habitat
Acronym
Shoreline and sandy shore
BAR
Aquatic environment and riparian zone
ACQ
Sand dune
DUN
Juniperus maritimus scrub
MAG
Retama raetam scrub
MAR
Tama fix g ro u p in g s
TAM
Back dune open environment
APR
Saccarum monophytic g ro u p in g s
SAC
Eucalyptus rostrata reforestation
EUC
Acacia saligna reforestation
ACA
Pinus pinea reforestation
PIN
Table 2. Habitat types within the study area and
used acronym s.
maps, the following 11 natural and seminatural
habitat types have been identified (Table 2):
In order to compare the faunistic values of the
habitats aforementioned, considering the specific
biodiversity level in each habitat, and the faunistic
value of each species, an index of the faunistic
I Aj)
/<*) = £— x 100
Z>0)
1(h) = Index of habitat faunistic value
j = species
h = habitat
v(j) = faunistic value of species
nh(j) = number of habitats in which the species is present
S= species of all habitat
Figure 2. Formula used to calculate the faunistic
value of the habitat.
value of the habitat, 1(h), calculated with the for-
mula showed in Fig. 2.
Each species contributes to the ecosystems
functioning, becoming part of the trophic netw orks,
and using, at various levels, the habitat resources,
so none of them can take a null faunistic value.
Therefore, it was considered appropriate to assign
a minimum value of 0.01 to each taxon. This value
has been estimated as half of the minimum value of
W/) found .
nhti)
RESULTS
A total of 273 animal taxa were counted, of
which 198 were Arthropods, and 186 Insects.
The study of the invertebrate fauna of a geo-
graphical area, although of limited size, requires
very long times and the use of many specialists of
different taxonomic groups, in consideration of its
great richness and of its articulation, which allow s
it to occupy most part of habitats, and in any case,
can not be exhaustive (Sabella et al., 2015). Just
remember that the check-list of Italian fauna
(Minelli et al., 1 993-1 995 ) cites for Sicily over
I 2,000 terrestrial taxa, with the Order of the
Coleoptera which includes about 4,400 species
and subspecies.
The study of the invertebrate fauna, therefore,
was aimed to examine only some of the fauna
components considered important to establish the
environmental quality and to identify the potential
impacts related to modifications of the environ-
ment. So, some groups were considered relevant to
the study of the fauna of the soil, and of the sub-
aerial environments. In particular, were considered,
among the Chelicerata, Araneidae, and among the
M andibulata, Crustacea (terrestrials amphipods and
isopods) and Insecta (Odonata, Orthoptera, B lat-
toidea, Heteroptera, Coleoptera, Lepidoptera and
Hymenoptera Form icid ae ). A m on g these 96 species
were Coleoptera, 14 Lepidoptera, and 22 Hymenop-
tera Form icidae.
Among collected Insects taxa, three ( OvthetVWTl
trinacria (Seiys, 1 8 4 1 ) , Ochrilidia sicula Saifi,
1931 and Carabus faminii faminii Dejean, 1826)
have already been proposed for inclusion in Annex
II to Directive 92/43 EEC, while two (CalOTYievCl
littoralis nemoralis (Olivier, 1 7 90), and Eurynebria
COTTiplciVlCltCl ( Linnaeus, 1767) are included in annex
A of regional law 6 April 2000 n. 56 of Tuscany
A study case of Assessment of Impact using the invertebrates
189
Total
BAR
ACQ
DIN
MAG
MAR
TAM
APR
SAC
EUC
ACA
PIN
N of species
273
34
83
12!
105
109
98
135
54
72
57
64
faunistic value (VF)
89.75
9.017
13.623
14.510
7.540
10.115
7.682
12,959
2.811
4.723
3281
3.490
N of species with VF = 0.00!
169
17
45
75
62
64
57
88
29
40
33
38
N of species with VF> 0.001
104
17
38
46
43
45
41
47
25
32
24
26
N of species exclusives of habitat
39
5
16
l
0
2
4
11
0
0
0
1
1(h)
944
J4.59
15.87
8.67
i lit
8,74
14.44
338
549
3.89
4.17
Table 3. Distribution per habitat of species number, faunistic value, and fa unis tic value index. BAR = Shoreline and sandy
shore; ACQ = Aquatic environment and riparian zone; DUN = Sand dune; MAG = JllYlipeVUS YYUJLvitimUS scrub ; MAR =
Retama raetam scrub; tam = Tamarix groupings; a pr = Back dune open environment; sac = Saccarum monophytic
groupings; EUC = Eucalyptus WStrata reforestation; ACA = Acadd SClligna reforestation; PIN = PinUS pined reforestation .
VF = Faunistic value. 1(h) = Habi tat fa unis tic value index.
Region (Ballerio, 2004). DocioStaUYUS minutUS L a
Greca, 1 962 is endemic to southern Sicily, while
five species (Ochrilidia sicula, Isomira paupercula
(Baudi, 1883), NotOXUS siculliS L a Ferte -S en ec tere ,
1 849, Temnothorax laestrygon (Sants chi, 1 9 3 1 ),
and Temnothorax lagrecai (Baroni Urbani, 1964),
and six subspecies ( Euchorthippus albolineatus
siculus Ramme, 1927, ErodiuS Siculus Siculus
Solier, 1 834, TaSgiuS falcifer aliquoi (Bordoni,
1 976), Tasgius globulifer evitendus (Tottenham,
1945), Tasgius pedator siculus (Aube, 1 842), and
Pimelia rugulosa sublaevigata Solier, 1 8 3 6 ) are
endemic to S icily.
Twelve taxa show a distribution restricted to the
Mediterranean basin. Among these, one species,
Pimelia grossa Fabricius, 1792, has a Sardinian-Si-
cilian-M aghrebian geonem y, four species, Ocneridia
nigropunctata (Lucas, 1 849), Platycranus putoni
Reuter, 1 8 79, Bl'OSCUS politUS (Dejean, 1 828), and
Carabus faminii faminii, show a Sicilian-M aghrebian
geonemy, while Cylindera trisignata siciliensis ( w .
Horn, 1891) has a S ic ilian -Tun isian distribution,
and Temnothorax kraussei (Emery, 19 16) shows a
Sicilian-Sardinian-Corsican geonemy. Also, two
taxa, Brachygluta aubei (Tournier, 1 867) and
Plagiolepis schmitzi F orel, 1 895 in Italy are known
only to Sicily, while two other, Orthetrum trinacria
and Hypocacculus elongatulus (Rosenhauer, 1856)
are known only to Sicily and Sardinia.
Twenty four species could be considered steno-
topes and/or stenoecious, sometimes with a strict
and exclusive binding to a particular type of habitat.
They often show populations of a few specimens
and they are very localized and very sensitive to the
an tropic disturbance (e.g. Orthetrum trinacria,
Pterolepis annulata (Fieber, 1853), Ochrilidia sicula ,
Masoreus aegyptiacus Dejean, 1828, and Myrmica
Sabuleti M e inert, 1861). Therefore, the insect fauna
shows remarkable faunistic emergencies, which are
related essentially to the dune and back-dunes eco-
systems and to the open environments.
In Table 3 are shown, for each habitat, its
species number, its faunistic value, and its faunistic
value index, while figure 3 shows, in decreasing
order, the 1(h) values of each habitat.
Figure 3. Histogram of 1(h) values of the different habitats
in the study area. DUN = Sand dune; ACQ = Aquatic envir-
onment and riparian zone; APR = Back dune open environ-
ment; MAR = Retdnid raetam scrub; BAR = Shoreline and
sandy shore; TAM = Tdmdrix gro up ing s ; MAG = JimiperUS
maritimus seru b ; e u c = Eucalyptus rostrata re f o re s ta tio n ;
pin = Pinus pinea reforestation; aca = Acacia saligna
reforestation; SAC = Saccarum monophytic groupings .
190
Giorgio Sabella etalii
The richest habitat in species has been the back
dunes open environment (APR), followed by the
sand dune (DUN), and by the scrub habitats (MAR
and MAG). The reforestations (EUC, ACA, and
PIN) are relatively poor in species, while the
shoreline and sandy shore (BAR) is the habitat with
the least number of taxa.
The faunistic value of each habitat is equivalent
to the sum of the faunistic values (VF) of the
species present in its interior. The highest faunistic
value is found in the sand dune (DUN), followed
by the aquatic environment and riparian zone
(ACQ), by the back dune open environment (APR),
by the RetCUTlO. VCietCllfl scrub (MAR), and by the
shoreline and sandy shore (BAR). These habitats
take a particular value precisely in relation to the
invertebrates fauna component, while the species of
terrestrial Vertebrates not would have highlighted
the importance of these habitat from the wildlife
point of view.
The lowest values are found, instead, in the
Saccarum monophytic groupings (SAC) and in the
reforestations (EUC, ACA, and PIN).
The 1(h) values have been subdivided into 4
classes :
- Class 1 Habitat of low faunistic value for 1(h)
values ranging between 0.01 and 4.
- Class 2 Habitat of medium faunistic value for
1(h) values ranging between 4.01and 8.
- Class 3 Habitat of high faunistic value for 1(h)
values ranging between 8.01 and 12.
- Class 4 Habitat of very high faunistic value for
1(h) values ranging between 12.01 and 16.
Relying on these classes, a map of the faunistic
value of the study area was processed (Fig. 4); for
some useful methods to the identification of
the areas of faunistic interest see also Giunti et
al. (2008), Sabella et al. (2009), Petralia (2010),
Ingegnoli (2011), Petralia (2012).
Figure 4. Map of the faunistic value of the study area (Gela Refinery, Sicily).
A study case of Assessment of Impact using the invertebrates
191
The analysis of this map showed as the most
part of the realization of the containment diaphragm
concerned the areas of low or medium faunistic
interest. On this base, the identification of the
project potential impacts, and the propositions of an
optimal allocation of building sites and of the safe-
guard of the neighboring habitats with high natural-
istic value were also possible.
Were also proposed appropriate and effective
m itigation measures, based on criteria, not aesthetic,
but scientific and naturalistic. Specifically, renatura-
tion actions with the restoration and the extension of
the habitats of particular naturalistic interest, as back
dune open environments and RetCllTlCl TCietCllTl scrubs
in place of reforestations, have been provided.
CONCLUSIONS
The level of knowledge about the ecological
responses of species and communities to environ-
mental changes not still allows an accurate and
precise quantification of their effects.
The study of the invertebrate fauna, in relation
to its great species richness and the various and
articulated ecological requirements of the latter,
allows a more detailed assessment of the environ-
mental quality and a mo re accurate prediction of the
changes that may occur in the structure and in the
dynamics of the zoocoenosis in response to perturb-
ations induced by the realization of a project. So
this study, together with that of the Vertebrates,
enables better the identification of the areas of
faunistic interest and the evaluation of their value.
Then it is possible a more accurate assessment of
potential impacts of the project on wildlife and the
proposal for suitable and effective mitigation
measures and the post-operam monitoring the
actual effectiveness of these latter.
The study highlighted that, in unsuitable en-
vironmental conditions to the stay of the vertebrates
community, in relation to their high levels of
anthropic disturbance and/or to the limited exten-
sion of the territory, the study of invertebrates com-
munities for the environmental quality assessment
from the faunistic point of view can be very useful.
In fact, confined habitats can retain good levels of
animal biodiversity and represent a refuge for many
rare species of invertebrates, and so they have a
relevant importance for the wildlife conservation.
Unfortunately, in impact assessment studies, the
invertebrates are often completely neglected and the
evaluations are based solely on the vertebrate
species. When "Umbrella SpedeS " or habitats of
community interest are lacking, the communities of
invertebrates are, therefore, at risk.
REFERENCES
AA.VV., 2008. Atlante della Biodiversita della Sicilia:
Vertebrati Terrestri. Studi e Ricerche, 6.Arpa Sicilia,
P alerm o , 536 pp .
Audisio P., Baviera C., Carpaneto G.M ., BiscacciantiA.
B., BattistoniA., Teofili C. & Rondinini C. (compil-
atori), 2014. Lista rossa IU C N dei Coleotteri
saproxilici Italian i. Comitato IUCN e Ministero
dell’Ambiente e della Tutela del Territorio e del
Mare, Roma, 132 pp.
Bailer io A., 2004. La conservazione degli insetti e la
legge. [4° aggiornam ento : 3 0 giugno 2 004].
http ://w w w .so cietaen tom o log icaitaliana.it/it/serviz i/e
n tom olex.htm 1.
Bern Convention 1979, on the Conservation of European
Wildlife and Natural Habitats.
BirdLife International. 2004. Birds in Europe. Population
estimates, trends and conservation status. BirdLife
International, Ser. 12, Cambridge, UK.
Bonn Convection 1979, on the Conservation of
Migratory Species of Wild Animals.
Brichetti P., 1 997. Le categorie corologiche d ell'a v ifau n a
italiana. In: Manuale pratico di Ornitologia.
Calderini, Bologna: 223-237.
Cerfolli F., Petrassi F., Petretti F. (a cura di), 2002. Libro
rosso degli animali d ’ italia - In vertebrati. W W F Italia
Onlus, 83 pp.
Directive 92/43/EEC of the European Parliament and of
the Council of 21 May 1992, on the conservation of
natural habitats and of wild fauna and flora. Official
Journal of European Union L 206, 22/07/1992.
Directive 2009/147/EC of the European Parliament
and of the Council of 30 November 2009, on the
conservation of wild birds. Official Journal L 20/7,
26/01/2010.
D.P.R. 8 settembre 1997 n. 357, Regolam ento recan te
l’attuazione della Direttiva 92/43/CEE relativa alia
conservazione degli habitat naturali e sem inaturali,
nonche della flora e della fauna selvatiche. Supple-
mento Ordinario alia G.U.R.I. n. 248 del 23 ottobre
1 997.
Giunti M ., Castelli C ., Colliggiani L., Di Vittorio M.,
Ientile R. & Lastrucci B., 2008. Metodologia per
l’individuazione di aree di i m porta nza faunistica.
Estimo e Territorio, 2: 36-47.
192
Giorgio Sabella etalii
Ingegnoli V., 2011. Bionomia del paesaggio. L'ecologia
del paesaggio b io lo g ic o - in te g ra ta per la form azione
di un “m edico” d e i sis tem i eco logic i. Springer-Verlag
Italia, xix + 346 pp.
IUCN, 2012. Red List Categories and Criteria: Version
3.1. Second edition. Gland. Switzerland and
Cambridge, UK: IU C N , iv + 3 2 pp.
IUCN, 2014. Red List of Threatened Species 2014.3.
http://www.iucnredlist.org/.
Legge 19 dicembre 1975 n. 874. Ratifica ed esecuzione
della convenzione sul commercio internazionale
delle specie animali e vegetali in via di estinzione,
firm ata a Washington il 3 marzo 1973. Supple-
mento Ordinario alia G.U.R.I. n . 49 del 24 febbraio
1 975.
Legge 5 agosto 1981 n. 503, Ratifica ed esecuzione della
convenzione relativa alia conservazione della vita
selvatica e d ell’am b ie n te naturale in Europa, con
allegati. adottata a Bern a il 19 settembre 1979.
Supplem ento Ordinario alia G.U.R.I. n. 250 dell'll
settem bre 19 8 1.
Legge 25 gennaio 1983 n. 42, Ratifica ed esecuzione
della convenzione sulla conservazione delle specie
migratorie appartenenti alia fauna selvatica, con
allegati, adottata a Bonn il 23 giugno 1979. G.U.R.I.
n. 48 del 18 febbraio 1 983.
Legge 11 febbraio 1992 n . 157, Nor me per la protezione
della fauna selvatica omeoterma e per il p relievo
venatorio. Supplem ento Ordinario alia G.U.R.I. n . 46
del 25 febbraio 1 992.
Legge Regionale n. 56 del 6 aprile 2000. Norme per la
conservazione e la tutela degli habitat naturali e
sem in aturali, della flora e della fauna selvatiche -
Modifiche alia legge regionale 23 gennaio 1998, n. 7
- Modifiche alia legge regionale 11 aprile 1995, n . 49.
Bollettino U fficiale della Regione Toscana n. 17 del
1 7 ap rile 2 000.
Massa B. & Canale E.D., 2008. Valutazione della biod-
iversita in Sicilia: 237-248. In:AA.VV.Atlante della
Biodiversita della Sicilia: Vertebrati Terrestri. Studi
e Ricerche, 6.Arpa Sicilia, Palermo, 536 pp.
M inelli A., Ruffo S. & La Posta S. (Eds.), 1993-1995.
Checklist delle specie della fauna italiana. Calderini.
Petralia E., 2010. Strum enti GIS per la p ia n ific a z io n e
nelle aree protette: il caso di Vendicari: 395-406. In:
Petralia A. (a cura di). L'Area protetta di Vendicari.
Atti del Convegno Celebrativo per il 35° Anno
di Fondazione dell’Ente Fauna Siciliana (Case
Cittadella, Ve d ic ari- n o to (SR) 25-26 ottobre 2008).
Collana Ecologia Phoenix, 1 2: 1-432.
Petralia E., 2012. GIS for environmental planning in
protected areas: fauna aspects. 2nd Djerba Interna-
tional Mediterranean Environments Sustainability
Conference 22-25 April 2012. Atti e Memorie
dell'Ente Fauna Siciliana, 11: 109-116.
Prola G. & Prola C ., 1 990. Libro rosso delle farfalle
italiane.QuaderniWWF: 13,71 pp.
Riservato E., Fabbri R., Festi A., Grieco C., Hardersen
S., Landi F., Utzeri C., Ron din ini C., BattistoniA. &
Teofili C. (comp ila tori), 2014. Lista rossa IUCN delle
libellule Italiane. Comitato IUCN e Ministero
dell'Ambiente e della tutela del Territorio e del M are,
Roma, 39 pp.
Ron din ini C., Battistoni A., Peronace V. & Teofili
C. (compilatori), 2013. Lista Rossa dei Vertebrati
italiani. Comitato Italiano IUCN e Ministero
dell'Ambiente e della Tutela del Territorio e del
M are. Roma, 54 pp .
Ruffo S. & Stoch F. (Eds.), 2005. Checklist e dis- tri-
buzione della fauna italiana. Memorie del Museo
Civico di Storia Naturale di Verona, 2a serie, Sezione
Scienze della Vita, 16.
Sabella G., Alicata A., Sorrentino M . & Sorgi G., 2009.
Aree di in ter esse faunistico: 61-81. In: Mancuso C.,
Martinico F., Nigrelli F. C. (a cura di). I Piani
Territoriali Paesistici della provincia di Enna.
Urbanistica Quaderni (collana dell’Istituto Nazionale
d i U rb an is tic a ), 5 3: 1 -1 68.
Sabella G., Lisi O. & Viglianisi F.M., 2015. The use of
the entom ofauna in the studies of the Environmental
Impact Assessment (E.I.A.) and Impact Assessment
(I. A.). The International Congress on speciation and
taxonomy. May 16th -18th 2014 Cefalu-Castelbuono
(Italy). Biodiversity Journal, 6: 1 75-1 84.
Vigna TagliantiA.,Audisio P.A., Belfiore C., Biondi M .,
Bologna M. A., Carpaneto G.M., De Biase A., De
Felici S., Piattella E ., Racheli T., Zappa roli M . & Zoia
S., 1992. Riflessioni di gruppo sui corotipi fon da-
men tali della fauna W-paleartica ed in particolare
italiana. Biogeographia. Lavori della Societa Italiana
di B iogeografia, n. s., 16: 1 59-1 79.
Vigna TagliantiA.,Audisio P.A., Biondi M ., Bologna M .
A., Carpaneto G .M ., De Biase A., Fattorini S .,
Piattella E., Sindaco R., VenchiA. & Zapparoli M.,
1999. A proposal for a chorotype classification of the
Near East fauna, in the framework of the Western
Palearctic region. Biogeographia. Lavori della
Societa Italiana di Biogeografia, n . s . , 20: 3 1- 59.
Washington Convention 1973, on International Trade in
Endangered Species of Wild Fauna and Flora.
Biodiversity Journal, 2015, 6 (1): 193-196
Monograph
Diversity in the population of Brassica incana Ten.(Cruciferae)
in Sicily
Francesco Maria Raimondo &Vivienne Spadaro
Dipartimento STEBICEF, Sezione di Botanica ed Ecologia Vegetale, Universita degli Studi di Palermo, via Archirafi 38, 90123
Palermo, Italy
ABSTRACT Phenotipic diversity in Sicilian populations of Brassica incana Ten. (Cruciferae) is here ana-
lyzed in comparison with the only one known population of B. raimondoi Sciandrello et al.,
taxonomic close species recently described from the coastal relief of eastern Sicily. The analysis
of diagnostic characters of these two taxa does not reveal significant differences that justify a
treatment at species level of the population of B. raimondoi. On this base, the authors deemed
to include this taxon in the infraspecific variability of B. incana and consider most appropriate
the rank of subspecies. Therefore is here proposed the establishment of the trinomial com-
bination B. incana subsp. raimondoi.
KEY WORDS Mediterranean flora; wild cabbage; Brassicaceae; taxonomy.
Received 02.03.2015; accepted 21.03.2015; printed 30.05.2015
Proceedings of the 2nd International Congress “Speciation and Taxonomy”, May 16th- 18th 20 14, Cefalu-Castelbuono (Italy)
INTRODUCTION
The Brassica Sect. Brassica (Cruciferae), is a
taxon represented by numerous forms described
both at specific and infraspecific level. The Sicilian
floristic district (sensu Fenaroli & Giacomini, 1968)
is considered the diversity center of this group,
because its geographical area, including Sicily, its
archipelagos and the islands of Malta and Gozo,
expresses the greatest biodiversity of this Section
seen that 70% of species related to it, not counting
the many infraspecific taxa, is concentrated in this
district. (Raimondo, 1997; Raimondo 2001). The
flora of Sicily includes in this section: B. macro-
carpa Guss., B. insularis Moris, B. rupestris Raf.,
B. villosa Biv., B. trichocarpa Brallo C. et al. and
B. incana Ten. To these, recently, has been added
B. raimondoi Sciandrello et al. (Fig. 1) very close
to B. incana and described from around Castelmola
(Messina), restricted area above the village of
Taormina, locus classicus of the taxon (Sciandrello
et al., 2013), next to one of the most classic coastal
localities known of B. incana (Capo S. Alessio)
(Fig. 2).
As part of the taxonomic review of the different
Sicilian populations of Sect. Brassica (Mazzola &
Raimondo, 1988; Raimondo et al. 1 99 1 ; Raimondo
& Mazzola, 1997), some taxa - previously considered
at specific level as B. tinei Lojac., B. drepanensis
(Caruel) Damanti and B. bivonana Mazzola et Rai-
mondo were included in B. villosa at subspecific
level. Similar treatment was given to some popula-
tions of B. rupestris differing from the type of the
species by phenotypic, ecological and distributive
characters; they were directly assigned the rank of
subspecies ( B . rupestris subsp. hirsuta Raimondo
et Mazzola and B. rupestris subsp. brevisiliqua Rai-
mondo et Mazzola) (Raimondo & Mazzola, 1997).
Based on these premises, and on the discovery
of a new population of B. incana s.str. (Tenore,
194
Francesco Maria Raimondo & Vivienne Spadaro
1812) on the Madonie Mountains (Raimondo in
PAL), we wanted to deepen the analysis of the
variability of this taxon foreseeing to include in it
B. raimondoi, taxon for which we propose the rank
of subspecies within B. incana.
MATERIAL AND METODHS
We studied the topotypical population of B.
incana Ten. and of B. raimondoi (Sciandrello et al.,
2013). Diagnostic characters of the two taxa repor-
ted in Sciandrello et al. (2013) are analyzed and
evalueted. In addition to morphological characters,
the spatial distribution in comparison with the
Sicilian populations of B. incana , in order to
exclude possible genetic interferences between the
populations, spatially but not orographic close,
then, subjected to two different bioclimates.
RESULTS AND DISCUSSION
Based on the analysis and evaluation of morpho-
logical characters and the criteria followed in the
interpretation of the variability occurring in the
other species of the same group previously treated,
the distinctive characters of the taxon are not suffi-
ciently discriminating to interspecific level. The
color of the petals is not a character that is distrib-
uted continuously in B. incana. In Sicily, on the
Tyrrhenian coast, between Capo d'Orlando and
Gioiosa Marea (Messina), there are populations of
this species with individuals with yellow or white
flowers, respectively (Fig. 3), maintaining constant
the other characters. In contrast, the same color of
petals and lenght of siliques - given as discriminant
of B. raimondoi by the authors - on the basis of the
study of the topotypical population are not constant.
In fact, although the white flowered individuals are
prevalent, yellow flowered individuals occur
scattered (Figs. 5, 6). The indumentum of flowering
pedicel and sepals (Fig. 4), of stem and adult leaf
hairless or weakly pubescent, are variable charac-
ters in B. incana and therefore are not considered
stable enough to be discriminating. Also in
Brassica, the different characters of the siliqua,
Mazzola & Raimondo (1988) distinguished B.
bivonana from B. villosa, then reduced to the rank
of subspecies of B. villosa by the same authors (Rai-
mondo & Mazzola, 1997) [B. villosa subsp.
bivonana (Mazzola & Raimondo) Raimondo &
Mazzola].
Similarly, on the same characters and their
variability was based the distinction, within B.
rupestris, of a new subspecies occurring in the
western limit of the distribution end of this species
including the Tyrrhenian coast between the
promontory of Cefalu (Palermo ) to the east, the
promontory of Macari (Trapani) to the west and the
inland of the Madonie and Palermo Mountains,
including Rocca Busambra, to the south; it is the
case of B. rupestris subsp. brevisiliqua Mazzola et
Raimondo.
In light of the above considerations and of the
knowledge of the group, the authors believe that B.
raimondoi is not sufficiently distinctat specific level
and consider the population of Brassica of the cliffs
of Castelmola (Messina) as part of the variability of
B. incana, close and spatially overlapping to this
taxon, present in the underlying Ionian coast, near
Cape S. Alessio. Therefore, we give to it the follow-
ing arrangement:
Brassica incana Ten. subsp. raimondoi
(Sciandr., C. Brullo, Brullo, Giusso, Miniss. et
Salmeri) Raimondo & Spadaro stat. & comb. nov.
Bas. Brassica raimondoi Sciandrello, C. Brullo,
Brullo, Giusso, Minissale & Salmeri in PI. Biosyst.
147(3): 813 (2013).
CHOROLOGICAL AND TAXONOMIC
REMARKS
In the Mediterranean Region, among the species
of Brassica sect. Brassica, there are many endemic
taxa. Two in particular have a distribution almost
specular from north to south. They are B. insularis
Moris and B. incana Ten. The first, to the west,
from the French coast, via Sardinia, goes south to
Pantelleria and Tunisia; the second extends its dis-
tribution throughout the Tyrrhenian and Adriatic
coasts to Sicily, includingin this trajectory the
eastern and western sides of the Italian peninsula to
Sicily where B. incana occupies the eastern sector;
the Madonie Mountains represent the southern-
western limit.
In the south-eastern part of the distribution of
this species, an isolated population of the Ionian
coastal sector of the Island, described sub B.
Diversity in the population of Brassica incana Ten. (Cruciferae) in Sicily
195
Figure 1. Brassica raimondoi in flower (white): locus classicus, Castelmola, Jonian coast of Sicily. Figure 2. Brassica
incana in flower (yellow): Capo S. Alessio, Taormina, Jonian coast of Sicily. Figure 3. Brassica incana at S. Gregorio, Capo
d’Orlando, Tyrrenian coast of Sicily: plants with yellow and white flowers respectively. Figure 4. Comparison between
Brassica raimondoi and B. incana s.str.: a (flower) and b (siliquae) of B. raimondoi ; c (flower) and d (siliquae) of B. incana
(recomposed from Sciandrello et al., 2013).
196
Francesco Maria Raimondo & Vivienne Spadaro
Figure 5. Brassica raimondoi, rarely with yellow flowers, in his locus classicus (Castelmola).
Figure 6. Brassica raimondoi in the locus classicus (Castelmola): plants with white and yellow flowers respectively.
raimondoi , has not significantly discriminant
phenotypic characters which suggest to include the
taxon within B. incana. For the small size of the pop-
ulation and the spatial isolation of the population of
B. raimondoi remains taxonomically distinct and
still subject to subspecific level.
ACKNOWLEDGMENTS
Study carried out as part of a project funded by
the University of Palermo with the financial support
of the Fondazione internazionale pro Herbario
mediterraneo.
REFERENCES
Brullo C., Brullo S., Giusso del Galdo G. & Ilardi V.,
2013. Brassica trichocarpa (Brassicaceae), a new
species from Sicily. Phytotaxa, 122: 45-60.
Fenaroli L. & Giacomini V., 1968. La Flora. Milano,
T.C.I.
Mazzola P. & Raimondo F.M., 1988. A new species of
Brassica from Sicily. Lagascalia, 15 (extra): 249-
251.
Raimondo F.M., 1997. Les membres italiens du com-
plexe de Brassica oleracea: leur distribution et spe-
cificites ecologiques. Bocconea, 7: 103-106.
Raimondo F.M., 2001. Conservazione della flora in Ita-
lia. II caso dei cavoli selvatici della Sicilia. Pp. 229-
241 in: Gomez-Campo C., Conservation de especies
vegetales amenazadas en la region mediterranea oc-
cidental. Madrid, Fondacion Areces.
Raimondo F.M. & Mazzola P, 1997. A new taxonomic
arrangement in the Sicilian member of Brassica L.
sect Brassica. Lagascalia, 19: 831-838.
Raimondo F.M., Mazzola P. & Ottonello D., 1991. On
the taxonomy and distribution of Brassica sect. Bras-
sica (Cruciferae) in Sicily. Flora Mediterranea, 1 : 63-
86 .
Sciandrello S., Brullo C., Brullo S., Giusso del Galdo G.,
Minissale P. & Salmeri C., 2013. A new species of
Brassica sect. Brassica (Brassicaceae) from Sicily.
Plant Biosystems, 147: 812-820.
Tenore M., 1812. Flora Napolitana, Prodromus. Stampe-
ria Reale, Orto Botanico, Napoli.
Biodiversity Journal, 2015, 6 (1): 197-204
Monograph
Taxonomy and conservation in Higher Plants and Bryophytes
in the Mediterranean Area
Gianniantonio Domina*, Giuseppe Bazan, Patrizia Campisi & Werner Greuter
Orto botanico ed Herbarium Mediterraneum dell’Universita di Palermo, via Lincoln 2, 90133 Palermo, Italy
■"Corresponding author
ABSTRACT The Mediterranean Region is among the areas of the world richest in wild and cultivated taxa.
Extinctions in the Mediterranean area are bound to have occurred in historical times but they
are not documented. The probable and documented cases of plant extinction in specific areas
within the Mediterranean are equivalent to 0.25% of total species-by-area records. Species
with a large range are more prone to local population size fluctuations and eventual extinction
than species with a reduced population. Small islands floras are more prone to extinction than
those on large islands and on the mainland. Reliability of our data on Mediterranean plant ex-
tinctions is poor. New emphasis on floristic research is needed to boost our deficient knowledge
of the Mediterranean flora. A closer collaboration between scholars and amateurs can increase
floristic knowledge and also help unravel taxonomic problems.
KEY WORDS vascular plants; mosses; extinctions; nomenclature.
Received 11.01.2015; accepted 02.03.2015; printed 30.03.2015
Proceedings of the 2nd International Congress “Speciation and Taxonomy”, May 1 6th- 1 8th 2014, Cefalu-Castelbuono (Italy)
INTRODUCTION
The Mediterranean Region is among the areas
of the world richest in wild and cultivated taxa. The
vascular flora comprises about 25.000 species. 50%
of the Flora (about 12.500 species) are endemic
(Medail & Quezel, 1999). This richness is not due
to high local species density but to small mean dis-
tributional areas reflected in a remarkable number
of narrow endemics (Greuter, 1991; 2001). In spite
of this high biodiversity, not all Mediterranean
countries have their own Red List of endangered
plants as yet; it is therefore quite difficult to make
between-country comparisons. An overview was
offered by Leon et al. (1985), who summarised the
risk status of endemics in Mediterranean countries.
More recently new lists were prepared for single
countries. The red list of the flora of Greece (Phitos
et al., 2009) is an outstanding example. Global
summaries for Europe have been presented by
Sharrock & Jones (2009), Bilz et al. (2011), and
Heywood (2012), but for the extra-European parts
of the Mediterranean they are still wanting.
MEDITERRANEAN VASCULAR PLANT
EXTINCTIONS
Extinctions in the Mediterranean area are bound
to have occurred in historical times, with the advent
of agriculture and the profound transformation of
the biota it entailed; but they are not documented.
Even the alleged extinction of the famous
“silphium” is not proven with certainty. The plant,
probably a Ferula or other giant umbellifer, was
used in classical antiquity in medicine and was also
198
Gianniantonio Domina etalii
fed to sheep and cattle. It was an essential item of
trade with the ancient North African city of Cyrene
(Fig. 1). By the first century A.C. the species, due
to overgrazing and over-collection, was considered
extinct in nature (Applebaum, 1979). However,
from extant written documents and paintings it is
not possible to name the plant with certainty. Its
identity with several species has been suggested in
lively and long-lasting debates, but it is still not
possible to know for certain whether or not the plant
is indeed extinct (Parejko, 2003).
The probable and documented cases of plant ex-
tinction in specific areas within the Mediterranean,
as recorded in Med-Checklist volumes 1,3, and 4
(Greuter et al., 1984-1989), are 116, equivalent to
0.25% of 47,298 total species-by-area records
(Greuter, 1991). The reported cases of total extinc-
tions of taxa are 22 (0.17 % of 12,886 taxa), of
which: 7 are “mystery cases”, 5 are cases of pos-
sible or actual rescue, and 10 are genuine cases of
(presumed) extinction (Greuter, 1991). Since 1991,
continuing field research has resulted in even more
reassuring figures: the 7 “mystery cases” remain the
same, 4 cases of possible or actual rescue were
added ( Coincya monensis subsp. puberula, Salvia
peyronii, Silene rothmaleri, Silene tomentosa),
bringing the total to 9; and of the genuine cases of
(presumed) extinction, only 6 remain.
The 7 “mystery cases” are: 1) Alyssum panicu-
latum Desf. (Cruciferae), based on a painting by
Aubriet, allegedly representing a Cretan plant found
by Tournefort in 1700 that matches no species
known to grow in that area. 2) Armeria arcuata
Boiss. et Reuter (Plumbaginaceae), once collected
by Welwitsch in Portugal and never again found;
according to Nieto Feliner (1987) it may well have
been an occasional intersectional hybrid. 3) Cam-
panula pyrenaica A. DC. (Campanulaceae), based
on two specimens, one allegedly from the Balearic
islands, the other from the Pyrenees. It has recently
been considered a synonym of Campanula sch-
euchzeri Vill. (Castro viejo et al., 2010). 4) Genista
melia Boiss. (Leguminosae), described from Milos
(Cyclades, Greece) and once doubtfully reported
from the Troad (Anatolia). The origin of the type,
which may well belong to the W Mediterranean
Genista scorpius complex, is in doubt. 5) Lathyrus
allardii Batt. (Leguminosae), described from near
Alger (Algeria) in 1879 and never seen since then.
Its native status has already been doubted by its
author. Perhaps it is only a form of Lathyrus gor-
goni Park, native further east and occasionally in-
troduced. 6) Quercus sicula Lojac. (Fagaceae),
described from a tree cultivated in the Botanical
Garden of Palermo of unknown, probably not Si-
cilian origin. An altogether doubtful taxon, perhaps
a mere variant of the Quercus pubescens complex.
7) Silene vulgaris subsp. aetnensis (Strobl) Pignatti
(Caryophyllaceae), described in 1885, at varietal
rank, from a single spot on Mt. Etna (Sicily).
Considered an enigmatic plant not recently seen
(Giardina et al., 2007).
The 9 cases of possible or actual rescue are: 1)
Coincya monensis subsp. puberula (Pau) Leadlay
(Brassicaceae), described in 1902 from Sanijan in
Galicia (Spain) but looked for unsuccessfully in its
locus classicus by Castroviejo (1982), was reported
from 4 localities in 1995 (Vioque & Pastor, 1995).
2) Diplotaxis siettiana Maire (Brassicaceae), an
endemic of Alboran island (Spain) where its only
population has recently been destroyed. It survives
in cultivation and seed banks, and reintroduction
into its native habitat looks promising (Perez
Latorre & al., 2013). 3) Erodium astragaloides
Boiss. et Reuter (Geraniaceae), described from
Sierra Nevada (Spain) where it has always been rare
and was not again found in this century, was re-
cently rediscovered in the Sierra de Cazorla
(Gomez-Campo, 1987). 4 ) Lysimachia minoricensis
Rodr. (Primulaceae), an endemic of Menorca
(Balearic Islands, Spain) that has disappeared from
its natural habitat but survives in cultivation. Rein-
troduction into its original habitat has been attemp-
ted (see Gomez-Campo, 1987) but so far has not
been successful, although attempts continue
(Galicia Herbada & Fraga Arquimbau, 2011). 5)
Onobrychis aliacmonia Rech. f. (Leguminosae),
described from Greek Macedonia, had its single loc-
ality, on the banks of Aliakmon River, flooded by
an artificial lake in c. 1975. A very similar plant, dis-
covered in Laconia (Peloponnesus), was first iden-
tified with it but later described as a distinct
subspecies then species, O. peloponnesiaca (Iatrou
et Kit Tan) Iatrou et Kit Tan. The genuine O. aliac-
monia was rediscovered in 1985 close to its clas-
sical locality, where it managed to colonise new
habitats (Greuter, 1987). 6) Limonium dufourii
(Girard) Kuntze (Plumbaginaceae), an endemic of
the Albufera de Valencia (Spain), first described in
1842, last seen in 1972, was considered a victim of
Taxonomy and conservation in Higher Plants and Bryophytes in the Mediterranean Area
199
Fig. 1. Sylver Coin of Cyrene dating back late 6th-early 5th centuries BC. depicting the silphium © Trustees of the British
Museum (Reproduced by kind permission of the British Museum of London) on the left and Ferula communis on the right.
reclamation of its native wetland areas. At present,
6 small populations are known to have survived
(Laguna et al., 1994). 7) Salvia peyronii Post
(Lamiaceae), discovered in 1883 on cliffs near
Feitroun (Lebanon) and never seen until recently,
although it is showy and had been looked for re-
peatedly, was found again in the same area (Jabal
Moussa) in 201 1 (Tohme & Tohme, 2011). 8) Silene
rothmaleri Pinto da Silva (Caryophyllaceae), de-
scribed in 1945 from Cabo S. Vicente (Algarve,
Portugal) and not seen since in spite of a thorough
search by Jeanmonod (Greuter & Raus, 1984), was
rediscovered in 2000 (Dinter & Greuter, 2004). 9)
Of Silene tomentosa Otth (Caryophyllaceae) a few
1 9th-Century specimens were known, all from the
E side of the Gibraltar rock (Spain), but until 1984
none had been collected in the 20th Century (Jean-
monod, 1984). In 1994 the species was rediscove-
red growing in the wild and is since cultivated in
the Alameda Botanical Gardens (Linares, 1998).
The 6 remaining cases of (presumed) extinction
are: 1) Cephalaria kesruanica Mouterde (Dip-
sacaceae), discovered in 1939 in Lebanon. Its type
locality was probably destroyed; a record from a
further locality requires confirmation (Mouterde,
1980). 2) Trachelanthus foliosus (Paine) Tristram
(Boraginaceae), discovered in Jordan in 1973 and
found in a second locality in 1886. It has not been
seen again (Feinbrun, 1978). 3 ) Dianthus multinervis
Vis. (Caryophyllaceae) discovered by Botteri on the
isolated islet of Jabuka (Porno, Croatia) where it
was not collected again and has probably disap-
peared (Greuter, 1995). 4) Fibigia heterophylla
Rech. f. (Brassicaceae), discovered in 1911 between
Homs and Palmyra in Syria and never again collec-
ted (Mouterde, 1970). 5) Morina subinermis Boiss.
(Dipsacaceae), described from plants collected in
Bithynia (Anatolia) without exact locality and never
seen since, although it is a showy species. 6) Trifo-
lium acutiflorum Murb. (Leguminosae). Described
from Murbeck’s own gathering made at Marrakesh
(Morocco), but never collected again in spite of
thorough searches (Fennane & Ibn Tattou, 1998).
The above figures do not refer to the entire
Mediterranean vascular flora but only to that part
(ca. 45 %) covered by the first three published
volumes of Med-Checlclist (Greuter et al., 1984-
1989). An attempt has subsequently been made by
Greuter (1994) to produce a similar list for the entire
Mediterranean flora, using various other sources.
The result was a table with 33 names, not taking
into account the “mystery cases” and redeemed
species. Of the taxa in this second list 21 are addi-
tional to the first, of which 12 were included bit not
200
Gianniantonio Domina etalii
considered extinct in Med- Checklist and 9 have not
yet been treated in that work. Focusing on the
former, we find that not all are worthy additions to
the extinct (Ex) category. One represents an inter-
specific hybrid ( Thalictrum simplex subsp. gallicum
(Rouy et Fouc.) Tutin = T.xtimeroyi Jord., see Hand
2001). Limonium dubyi (Gren. et Godr.) Kuntze,
described from France, is currently included in the
synonymy of L . bellidifolium (Gouan) Dumort. T
ephrosia kassasii Boulos, from the borders of the
Nile in Egypt, is not considered as extinct in that
country’s recent floristic and conservational literat-
ure. Thymus oehmianus Ronniger et Soslca obvi-
ously survives in its locus classicus in the Treska
gorge, as its live portrait appears on a recent (2003)
postage stamp of the FYR Makedonija. Of the 7
Turkish endemics listed on the faith of Ekim et al.
(1989), 4 apparently survive according to recent
assessments of that country’s flora (Ekim et al.,
2000, Eken et al., 2006): Campanula oligosperma
Damboldt, Onosma affinis Riedl, Sedum polystri-
atum R.T. Clausen, and Silene oligotricha Hub.-
Mor. This leaves us with a reduced number of 4
genuine, additional presumed extinctions: Local
and Global extinctions
Species with a large range are more prone to
local population size fluctuations and eventual
extinction than species with a reduced population.
Neslia paniculata (L.) Desv. (Brassicaceae) is an
example of a species with large distribution that
registered important local extinctions at the borders
of its range (Fig. 2). Spirodela polyrrhiza (L.)
Schleid. (Lemnaceae), a species distributed almost
world-wide, was considered extinct in Catalonia
but has been found in the lower course of the Ebro
River and in the Vallvidrera reservoir (Curto et al.,
2013). Rhamphidium purpuratum Mitt. (Bryophyta,
Ditrichaceae), known to be widely distributed in
Macaronesia and Crete, had its only mainland site,
known since 1940, in the north of Portugal. The
Fig. 2. Distribution of Neslia paniculata from Jalas et al. (1996). The crosses in the NW part
of the distribution indicate the extinction of the plant in that area.
Taxonomy and conservation in Higher Plants and Bryophytes in the Mediterranean Area
201
species was considered as vulnerable in the
European Red List (www.bio.ntnu.no/ECCB/ RDB
Taxon.php), and later considered extinct in Portugal
by Sergio et al. (1994, 2001). After more than 60
year it was found again in south-west Portugal near
the Monchique mountains (Sergio et al. 2011).
For taxa with a single locality destruction of the
habitat implies its complete loss: Adenostyles
alpina subsp. nebrodensis (Wagenitz et I. Mull.)
Greuter (Asteraceae) is known from a single loc-
ality in a canyon of the Madonie Mountains (Sicily);
the capture of a source, c. 50 years ago, has aridified
the area and brought the taxon to the brink of
extinction, with but a single individual still alive.
Limonium catanense (Lojac.) Brullo (Plumba-
ginaceae), at the beginning of the 20th century, was
only known in an area that now belongs to the
harbour of Catania.
Small islands floras are more prone to extinction
than those on large islands and on the mainland
(Greuter, 1995). This may be due to the greater fra-
gility of island habitats due to their smaller surface
and higher human pressure (Domina & Mazzola,
2011). The population of Daucus rupestris Guss.
(Apiaceae) on Lampione is extremely depleted and
faces im min ent extinction due to high concentration
of nitrogen from gull droppings, of which the plant
is intolerant (Lo Cascio & Pasta, 2012). Limonium
intermedium (Guss.) Brullo (Plumbaginaceae) was
growing on Lampedusa in a salt marsh near the
harbour, now converted to a soccer field (Fig. 3); at
present only some individuals survive in the Bota-
nical Garden of Catania, grown from seed sampled
in the field about 40 years ago (Brullo, pers. comm.).
Cistus xskanbergii Lojac. (Cistaceae) is the
natural hybrid between C. monspeliensis and C.
parviflorum and occurs in scattered localities in the
Mediterranean, wherever the two parents meet. It
was described from the island of Lampedusa (Italy),
where today but a single individual is known and
only one parent still occurs. Efforts to conserve that
individual would be a futile exercise.
There are several cases of old, unconfirmed
records, due perhaps to misidentification, and of
taxa of uncertain taxonomic position, that must be
taken into account in management and conservation
plans for endangered species. Orobanche aegyp-
tiaca Pers. (Orobanchaceae) was reported in error
from Italy (Monte Gallo and Lampedusa). It was re-
corded in Scoppola & Spampinato (2005) as being
Fig. 3. The soccer field in Lampedusa on the place of
the salt marsh near the harbour locus classicus et unicus of
Limonium in termedium .
very rare and endangered. The study of specimens
from both sites showed that they had been misiden-
tified and belong to O. mutelii F.W. Schultz
(Domina et al., 201 1). Euphrasia mendoncae Samp.
(Orobanchaceae) was described by Sampson in
1936 from specimens collected in 1932 by F.
Mendonga and thought to be endemic to Braganga
(Portugal). It was never found again despite extens-
ive searches in 1990 and 1996. In Flora Iberica
(Vitek, 2009) now it is treated as a synonym of E.
minima Jacq. ex DC. The presence of such non-
species in lists of plants requiring protection diverts
attention from others that are really threatened.
Among the mosses there are also cases of taxa
considered as extremely rare or extinct, only to be
later included in other, widely distributed taxa.
Clasmatodon parvulus (Hampe) Sull. (Bra-
chytheciaceae) was believed to occur in North
America and very rarely in Germany and Spain.
Meinunger (1992) deemed it as extinct in Germany
because it had not been found again since 1851, and
it came to be known as one of the rarest mosses of the
European continent (Frey et al., 1995; Dull, 1985),
being so was included as endangered (EN) in the
Red Data Book of European Bryophytes (Schu-
macker & Martiny, 1995). Eleras et al. (2006) found that
both the German and Spanish records were based
on misidentified plants of Pseudoleskeella tectorum
(Funck ex Brid.) Kindb. ex Broth, and must be ex-
cluded from the European and Mediterranean bry-
oflora. Thamnium cossyrense Bott. (Neckeraceae),
described by Bottini (1907), was considered en-
demic to Pantelleria until 2001, when Mastracci
202
Gianniantonio Domina etalii
(2001) included it in Scorpiurium sendtneri
(Schimp.) M. Fleisch., a species widely distributed
in Mediterranea area. Fissidens exiguus Sull. (Fis-
sidentaceae), considered rare in France and Greece
(Schumacker & Martiny, 1995), is now synonymised
with F. bryoides Hedw., a common taxon in
Temperate areas (Pursell, 2007). Likewise, Trichosto-
mopsis aaronis (Lorentz) S. Agnew & C. C. Towns
(Pottiaceae), thought to be a rare taxon of Spain and
Turkey (Schumacker & Martiny, 1995), has recently
been synonymised with Didymodon australasiae
(Hook, et Grev.) R. H. Zander, a common taxon
throughout the Mediterranean area (Ros et al., 2013).
DELIMITATION OF TAXA
Different delimitation bear on the range and the
conservation status of taxa (Lastrucci et al., 2014).
A glaring example is Thymus herba-barona Loisel.
that can either be considered to comprise a single
taxon, occurring in the Balearic islands, Corsica and
Sardinia (Molins et al., 2011), or split into different
taxa based on chromosome number: the diploid
Thymus herba-barona subsp. bivalens Mayol et al.
(2n = 28), endemic to a single locality in Serra
D’Aljabia (Mallorca) with only about 50 mature
individuals; Thymus herba-barona subsp. herba-
barona , tetraploid (2n = 56) growing in Corsica,
and T. catharinae Camarda, hexaploid (2n = 84),
restricted to Sardinia.
Arenaria bolosii (Canig.) L. Saez et Rossello
(Caryophyllaceae), a critically endangered taxon
only known from a single site on the island of Mal-
lorca (Bibiloni & Mus, 2006), was first described
as a variety, Arenaria grandiflora var. bolosii
Canig., then considered a subspecies, A. grandiflora
subsp. bolosii (Canig.) Colom, eventually to be
recognised a as a separate species.
Some taxonomists, called lumpers, tend to fa-
vour a broad taxon concept, whereas others, known
as splitters, emphasize minute differences. These
preferences depend in part on the size of the area on
which a researcher is working. Those who study
plants of a restricted territory will likely search for
differences whereas botanists with a broad geogra-
phical interest may favour a synthetic approach.
Several taxa considered to be narrow endemics can
just as well be interpreted as local expressions of
wide-ranging taxa. However, as long as there can be
reasonable doubt it is better to maintain the local
taxa rather than letting them disappear in synonymy.
As an example, in Pancratium maritimum L., some
populations may not deserve the status of separate
species yet they possess a well-diversified genome
that deserves being preserved (Giovino et al., 2015).
CONSIDERATIONS
New emphasis on floristic research is needed to
boost our deficient knowledge of the Mediterranean
flora. Amateurs, if well directed, can play an impor-
tant role in this endeavour. Many academic scholars
spend much of their time and efforts in the laborat-
ory, to the detriment of field research, yet they may
dispose of funds for networking that allow combin-
ing the efforts of amateurs over a wider geograph-
ical area than they might cover individually. A
closer collaboration between scholars and amateurs
can, by promoting field research by the latter, not
only increase floristic knowledge but also help
unravel taxonomic problems.
Lists of taxa that are part of laws or regulations
and their annexes should be updated at regular
intervals. In order to apply to the originally intended
taxa they must use their currently correct names;
but they should also, in addition, include synonyms
to reflect historical usage and accommodate altern-
ative taxonomic views. For higher plants, syn-
onymic checklists now exist that are widely accep-
ted by the scientific community, notably Euro+Med
Plantbase (http://www.emplantbase.org/), resulting
from a project funded within the V and VII Frame-
work Programme of the European Community.
Likewise, the checklists published by Ros et al.
(2007, 2013) provide a sound basis for channeling
conservation measures of Mediterranean hepatics,
anthocerotes and mosses.
REFERENCES
Applebaum S., 1979. Jews and Greeks in ancient Cyrene.
Leiden, Brill.
Bibiloni G. & Mus M., 2006. Arenaria bolosii. The
IUCN Red List of Threatened Species. Version
2014.3. <www.iucnredlist.org>. [Accessed on 01
January 2015].
Bilz M„ Kell S.P., Maxted N. & Lansdown R.V., 2011.
European Red List of Vascular Plants. Publications
Office of the European Union, Luxembourg.
Taxonomy and conservation in Higher Plants and Bryophytes in the Mediterranean Area
203
Bottini A., 1907: Sulla briologia delle isole italiane.
Webbia 2: 345-402.
Castroviejo S., 1982. Sobre la flora Gallega, IV. Anales
Jardin Botanico de Madrid 39: 157-165.
Castroviejo S., Aldasoro J.J. & Alarcon M., 2010. Cam-
panulaceae. - In: Euro+Med Plantbase - the informa-
tion resource for Euro-Mediterranean plant diversity.
Published on the Internet: http://ww2.bgbm.org/
EuroPlusMed/ [accessed 28/12/2014]
Curto R., Arrufat M., Beltran J., Creix A., Fontanet J. &
Royo F., 2013. Retrobada a Catalunya (NE de la pe-
ninsula Iberica) una poblacio de Spirodela polyrhiza
(Araceae). Orsis 27: 141-150.
Dinter I. & Greuter W., 2004. Silene rothmaleri (Cary-
ophyllaceae), believed extinct, rediscovered at Cabo
de Sao Vicente (Algarve, Portugal). Willdenowia, 34:
371-380.
Domina G. & Mazzola P., 2011. Considerazioni biogeo-
grafiche sulla presenza di specie aliene nella flora
vascolare del Mediterraneo. Biogeographia 30: 269-
276.
Domina G., Marino P. & Castellano G., 2011. The genus
Orobanche (Orobanchaceae) in Sicily. Flora Medi-
terranea, 21: 205-242
Dull R., 1985. Distribution of the European and Maca-
ronesian mosses (Bryophytina). Part II. Bryologische
Beitrage, 5: 110-232.
Eken G., Botdogan M., Isfendiyaroglu S., Kilig D. T. &
Lise Y., 2006. Turkiye’nin onemli doga alanlan. Cilt
2. Ankara: Doga Dernegi.
Ekim T., Koyuncu M., Erik S. & flarslan R., 1989.
Turkiye’nin tehlike altmdaki nadir ve endemic bitki
turleri IUCN red data book kategorilerine gore
hazirlami§tir. Ankara: Turkiye Tabiatim Koruma
Dernegi.
Ekim T., Koyuncu M., Vural M., Duman H., Ay tag Z. &
Adiguzel N., 2000. Turkiye bitkileri kmmizi kitabi
(egrolti ve tohumlu bitkiler). Red Data Book of
Turkish plants (Pteridophyta and Spermatophyta).
A nk ara: Turkiye Tabiatim Koruma Dernegi.
Feinbrun-Dothan N., 1978. Flora Palaestina, 3. Israel
Academy of Sciences and Humanities, Jerusalem.
Fennane M. & Ibn Tattou M., 1998. Catalogue des plan-
tes vasculaires rares, menacees ou endemiques du
Maroc. Bocconea 8: 5-243.
Frey W., Frahm J. P., Fisher E. & Lobin W., 1995. Die
Moos- und Famp Hanzen Europas. [Gams, H. (ed.),
Kleine Kryptogamenflora, 4, ed. 6]. Stuttgart: Fischer.
Galicia Herbada D. & Fraga Arquimbau P., 2011.
Lysimachia minoricensis. The IUCN Red List
of Threatened Species. Version 2014.3. <www.
iucnredlist.org>. [Accessed on 18 January 2015]
Giardina G., Raimondo F.M. & Spadaro V., 2007. A cata-
logue of plants growing in Sicily. Bocconea 20: 5-582.
Giovino A., Domina G., Bazan G., Campisi P. & Scibetta
S., 2015. Genetic and nomenclatural investigation of
Pancratium maritimum (Amaryllidaceae) in the Cen-
tral Mediterranean, from a conservationist’s point of
view. Journal of Coastal Conservation (submitted).
Gomez-Campo C. (Ed.), 1987. Libro Rojo de especies
vegetales amenazadas de Espana peninsular e Islas
Baleares. Madrid: ICONA.
Greuter W., 1979. Mediterranean conservation as viewed
by a plant taxonomist. Webbia, 34: 87-99.
Greuter W., 1987. Onobrychis aliacmonia (Legumino-
sae) - the unusual story of a rediscovery. Plant Sy-
stematics and Evolution, 155:21 5-2 1 7.
Greuter W., 1991. Botanical diversity, endemism, rarity,
and extinction in the Mediterranean area: an analysis
based on the published volumes of Med-Checklist.
Botanika Chronika, 10: 63-79.
Greuter W., 1994. Extinctions in Mediterranean areas.
Philosophical Transactions of the Royal Society, Ser.
B, 344: 41-46.
Greuter W., 1995. Origin and peculiarities of Mediter-
ranean island floras. Ecologia Mediterranea, 21: 1-10.
Greuter W., 2001. Diversity of Mediterranean island
floras. Bocconea 13: 55-64.
Greuter W., Burdet H.M. & Long G., 1984-1989. Med-
Checklist, 1, 3, 4. Geneve & Berlin.
Greuter W. & Raus T., 1984. Med-Checklist Notulae, 9.
Willdenowia 14: 37-54.
Hand R., 2001. Revision der in Europa vorkommenden
Arten von Thalictrum subsect. Thalictrum (Ranun-
culaceae). Frankfurt a.M.: BVNH.
Heras P., Infante M. & Buck W.R., 2006 On the presence
of Clasmatodon parvulus (Bryopsida) in Europe.
Herzogia 19: 317-321.
Heywood V.H., 2012. The impacts of climate change on
plant species in Europe. Pp. 95-244 in: Biodiversity
and climate change: Reports and guidance developed
under the Bern Convention, 2. Strasbourg: Council
of Europe.
Jalas J., Suominen J. & Lampinen R. (Ed.), 1996. Atlas
Florae Europaeae. Distribution of Vascular Plants in
Europe, 11. Commitee for Mapping the Flora of
Europe & Societas Biologica Fennica Vanamo,
Helsinki.
Jeanmonod D., 1984. Revision de la section Siphono-
morpha Otth du genre Silene L. (Caryophyllaceae)
en Mediterranee occidentale II: le groupe du S. mol-
lissima. Candollea39: 195-259.
Laguna E., Aguilella A., Carretero J.L., Crespo M.B.,
Figuerola R. & Gonzalo M., 1994. Libro de la flora
vascular rara, endemica o amenazada de la comunidad
valenciana. Generalitat Valenciana, Valencia.
Lastrucci L., Foggi B., Ferretti G., Guidi T., Geri F. &
Viciani D., 2014. The influence of taxonomic revi-
sions on species distribution assessment: the case of
204
Gianniantonio Domina etalii
three Asplenium species on Tuscan ultramafic soils.
Webbia, 69: 295-300. doi: 10.1080/00837792.2014.
955961
Leon C., Lucas G. & Synge H., 1985. The value of in-
formation in saving threatened Mediterranean plants.
Pp. 177-196 n : Gomez-Campo C., (Ed.): Plant Con-
servation in the Mediterranean area. Dordrecht: W.
Junk (no. 7 in Geobotany series, ed M. J. A. Werger).
Linares L., 1998. Silene tomentosa Otth in DC. Gibraltar
campion. A description based on observations of the
species in the wild, and on plants grown from seed
at the Alameda Botanical Gardens. Almoraima 19:
257-259.
Lo Cascio P. & Pasta S., 2012. Lampione, a paradigmatic
case of Mediterranean island biodiversity. Biod-
iversity Journal, 3: 311-330.
Mastracci M., 2001. Taxonomic status of Thamnium cos-
syrense and T. cossyrense var. melitense (Bryopsida).
Annales Botanici Fennici, 38: 45-46.
Medail F. & Quezel P., 1999. Biodiversity hotspots in the
Mediterranean basin: Setting global conservation
priorities. Conservation Biol. 13: 1510-1513.
Meinunger L., 1992. Endangered bryophytes in the
eastern part of Germany. Biological Conservation,
59: 211-214.
Molins A., Bacchetta G., Rosato M. Rossello J.A. &
Mayol M., 2011. Molecular phylogeography of
Thymus herba-barona (Lamiaceae): Insight into the
evolutionary history of the flora of the western
Mediterranean islands. Taxon 60: 1295-1305.
Mouterde P., 1980. Nouvelle Flore du Liban et de la
Syrie, 3. Impr. catholique, Beyrouth.
Nieto Feliner G., 1987. The genus Armeria (Plumbagi-
naceae) in the Iberian Peninsula: comments and new
data for a review. Anales Jardin Botanico de Madrid,
44: 319-348.
Parejko K., 2003. Pliny the Elder’s Silphium : First recor-
ded species extinction. Conservation Biology, 17:
925-927. doi: 10. 1046/j. 1523- 1739.2003.02067.x
Perez Latorre A.V., Cabezudo B., Mota Poveda J., Penas
J. & Navas P., 2013. Diplotaxis siettiana. The IUCN
Red List of Threatened Species. Version 2014.3.
<www.iucnredlist.org>. [Accessed on 29 December
2014],
Phitos D., Constantinidis T., Kamari G., (Ed.) 2009. The
Red Data Book of rare and threatened plants of
Greece, 1-2. Hellenic Botanical Society, Patras.
Pursell R.A., 2007. Fissidentaceae Schimper. Pp. 331-
343, in: Crosby, M. (Ed.) Flora of North America
north of Mexico, 27. Oxford University Press, New
York.
Ros R.M., Mazimpaka V., Abou-Salama U., Aleffi M.,
Blocked T.L., Brugues M., Cano M.J., Cros R.M.,
Dia M.G., Dirkse G.M., El Saadawi W., Erdag A.,
Ganeva A., Gonzalez- Mancebo J.M., Herrnstadt I.,
Khalil K., Kurschner H., Lanfranco E., Losada-Lima
A., Refai M.S., Rodriguez-Nunez S., Sabovjlevic M.,
Sergio C., Shabbara H., Sim-Sim M. & Soderstrom
L., 2007. Hepatics and Anthocerotes of the Mediter-
ranean, an annotated checklist. Cryptogamie Bryolo-
gie, 28: 351-437.
Ros R.M., Mazimpaka V., Abou-Salama U., Aleffi M.,
Blocked T.L., Brugues M., Cros R.M., Dia M.G.,
Dirkse G.M., Draper I., El-Saadawi W., Erdag A.,
Ganeva A., Gabriel R., Gonzalez-Mancebo J.M.,
Granger C., Herrnstadt I., Hugonnot V., Khalil K.,
Kurschner H., Losada-Lima A., Luis L., Mifsud S.,
Privitera M., Puglisi M., Sabovljevic M., Sergio C.,
Shabbara H.M., Sim-Sim M., Sotiaux A., Tacchi R.,
Vanderpoorten A. & Werner O., 2013: Mosses of the
Mediterranean, an annotated checklist. Cryptogamie
Bryologie, 34: 99-283.
Scoppola A. & Spampinato G., 2005. Atlante delle specie
a rischio di estinzione. - CD-ROM enclosed in:
Conti, F., Abbate, G., Alessandrini, A. & Blasi, C.
2005: An annotated checklist of the Italian vascular
flora. Palombi editore, Roma.
Schumacker R. & Martiny P., 1995. Threatened
bryophytes in Europe including Macaronesia.
In: European Committee for Conservation of
Bryophytes, editor. Red data book of European
bryophytes. Trondheim: University of Trond-
heim, pp. 29-193.
Sergio C., Casas C., Brugues M. & Cros R.M., 1994.
Lista Vermelha dos Briofitos da Peninsula Iberica /
Red List of Bryophytes of the Iberian Peninsula.
ICN, Lisboa.
Sergio C., Brugues M. & Cros R.M., 2001. New data
concerning extinct bryophytes on the Iberian Red
List. Novitates Botanicae Universitatis Carolinae, 15:
95-105.
Sergio C, Rodriguez-Gonzalez P.M., Albuquerque A. &
Garcia C.A., 2011. Rediscovery of Rhamphidium
purpuratum Mitt. (Bryophyta, Ditrichaceae) in Por-
tugal after more than 60 years. Field Bryology, 105:
40-41.
Sharrock S. & Jones M., 2009. Conserving Europe’s
threatened plants: Progress towards Target 8 of the
Global Strategy for Plant Conservation. Botanic
Gardens Conservation International, Richmond, UK.
Tohme G. & Tohme H., 2011. Nouvelles recherches sur
la flore endemique et naturalisee du Liban. Lebanese
Science Journal, 12: 133-141
Vioque J. & Pastor J., 1995. Aportaciones al conoci-
miento cariologico del genero Coincya (Brassicaceae)
en la peninsula iberica. Studia Botanica, 14: 143—
151.
Vitek E., 2009. Euphrasia. In Castroviejo S., Benedi
C., Rico E., Giiemes J. & Herrero A. (Eds.), Flora
Iberica, 13. CSIC, Madrid, 454^173.
Biodiversity Journal, 2015, 6 (1): 205-214
Monograph
Diversity in the genus Hieracium Linnaeus s. str. (Asteraceae)
in Sicily
Emilio Di Gristina 1 ’*, Francesco Maria Raimondo 1 & Pietro Mazzola 2
'Dipartimento STEBICEF, Sezione di Botanica ed Ecologia Vegetale, Universita degli Studi di Palermo, via Archirafi 38, 90123
Palermo, Italy; e-mail: francesco.raimondo@unipa.it
2 Dipartimento di Scienze Agrarie e Forestali, Universita degli Studi di Palermo, via Archirafi 38, 90123 Palermo, Italy; e-mail:
pietro.mazzola@unipa.it
’Corresponding author, e-mail: emilio.digristina@unipa.it
ABSTRACT The present taxonomic and floristic knowledges on Hieracium L. s. str. in Sicily are commen-
ted. In total, 1 1 taxa occur in this island, 10 of which are endemic and 1 has a wider range. For
each of these taxa, biological form, phenology, distribution, ecology, chromosome number,
conservation, and taxonomy are taken in consideration. A key to the taxa is also provided.
KEY WORDS Apomixis; conservation; endemism; taxonomy; vascular plants.
Received 02.11.2014; accepted 13.02.2015; printed 30.03.2015
Proceedings of the 2nd International Congress “Speciation and Taxonomy”, May 1 6th- 1 8th 2014, Cefalu-Castelbuono (Italy)
INTRODUCTION
Hieracium Linnaeus (1753) s. str. (Asteraceae)
is well known as one of the most species-rich plant
group in the world. It includes perennial herbs
distributed predominantly in temperate regions of
Europe, Asia and North America (Chrtek et al.,
2006). Hieracium belongs to a group of genera in
which diplosporous agamospermy and polyploidy
seem to prevail (Chrtek et al., 2006). The great
majority of Hieracium taxa are triploid (2n=27) or
tetraploid (2n=36) apomicts (Mraz et al., 2001).
Sexuality is extremely rare and confined to a few
diploid species, mostly distributed in South Europe
(Merxmuller, 1975; Chrtek et al., 2004).
Hybridization also appears as a very rare
phenomenon and is most likely confined to crosses
between diploid sexual species (Chrtek et al., 2006).
Agamospermy together with sexuality and hybrid-
ization in the past have given rise to a very large
number of variants that have been described as sub-
species, as has traditionally been the case in Central
Europe (Zahn, 1921-1923), or at rank of species
(British Isles, Scandinavia, East Europe) (Mraz et
al., 2001; Chrtek et al., 2006).
The Sicilian taxa (Fig. 1) have recently been
revised as for as taxonomy and distribution are con-
cerned (Raimondo & Di Gristina, 2004, 2007a, b;
Di Gristina et al., 2005, 2006; Geraci et al., 2007;
Di Gristina et al., 2012; Gottschlich et al., 2013; Di
Gristina et al., 2013; Caldarella et al., 2014).
These studies have already resulted in the
description of five new taxa: Hieracium racemosum
subsp. pignattianum (Raimondo & Di Gristina,
2004) Greuter (2007), H. schmidtii subsp. madoni-
ense (Raimondo & Di Gristina, 2007b) Greuter
(2007), H. pallidum subsp. aetnense Gottschlich,
Raimondo & Di Gristina (2013), H hypochoeroides
subsp. montis-scuderii Di Gristina, Gottschlich,
Galesi, Raimondo & Cristaudo (2013) and H.
busambarense Caldarella, Gianguzzi & Gottschlich
(2014).
206
Emilio Di Gristina etalii
Furthermore, the names of four taxa described
by Michele Lojacono (1903), H. cophanense, H.
crinitum var. caulescens , H. crinitum var. eriosta-
chyum and H. nebrodense, have been typified by
Aghababyan et al. (2008). The remaining five
accepted taxa described from Sicily, H. crinitum
Smith (1813), H. lucidum Gussone (1825), H. atro-
virens Froelich (1838), H. pallidum Bivona-
Bernardi (1838), H. symphytifolium Froelich
(1838), and three other names usually treated as
synonyms, H. racemosum subsp. todaroanum Zahn
(1922), H. siculum Gussone (1844) and H. siculum
var. minus Gussone (1844), have been typified by
Di Gristina et al. (2012).
On the whole, at present, several taxonomic and
chorological questions still remain open. Among
these, several issues of biodiversity conservation
are important especially for some strictly local
apomictic endemics, that are often considered of
secondary relevance respect to sexual species (Rich
et al., 2008) and, then overridden as for as conser-
vation is concerned.
Presently, an extensive field survey on the Sici-
lian territory is carried. The programme includes:
(1) field surveys in order to verify the occurrence
of the taxa known only from old reports or herbar-
ium data but not recently observed, (2) the collec-
tion of data of biological, ecological or phyto-
geographical interest for in situ and ex situ con-
servation.
A molecular approach using “DNA barcoding”,
in order to define the phylogenetic and systematic
relationships among the Sicilian taxa, and a cyto-
geographical analyses at population level, are also
in full progress. Waiting for a comprehensive
update account of the genus, the framework of
present knowledge is here summarized for each
taxon.
Figure 1. Distribution of the Sicilian Hieracium taxa. A) Hieracium busambarense; B) H. hypochoeroides subsp. montis-
scuderii; C) H. lucidum ; D) H. lucidum subsp. cophanense ; E) H. murorum subsp. atrovirens; F) H. pallidum', G) H. pallidum
subsp. aetnense; H) H. racemosum subsp. crinitum; I) H. racemosum subsp. pignattianum; L) H. schmidtii subsp. madoni-
ense; M) H. symphytifolium.
Diversity in the genus Hieracium Linnaeus s. str. (Asteraceae) in Sicily
207
MATERIAL AND METHODS
Floristic, herbarium, and literature research
carried out between 1999 and 2014 are surveyed
here. Specimens collected in the respective loci
classici and some other Sicilian localities are stored
in PAL. Zahn’s species and subspecies concept
(Zahn, 1921-1 923) has been adopted for taxonomic
nomenclature. Biological forms, following
Raunkiaer’s classification (1934), are abbreviated
as proposed by Pignatti (1982). Chromosome
numbers come from our karyological analyses and
other literature data. Conservation status follows
the IUCN (2010) criteria.
RESULTS AND DISCUSSION
In Sicily Hieracium s. str. is so far represented
by 1 1 taxa. 1 0 of them ( H . busambarense, H. hy-
pochoeroides subsp. montis-scuderii, H. lucidum,
H. lucidum subsp. cophanense, H. murorum subsp.
atrovirens, H. pallidum , H. pallidum subsp.
aetnense , H. racemosum subsp. pignattianum, H.
schmidtii subsp. madoniense and H. symphytifo-
lium ) are endemic to the island; the remaining (//.
racemosum subsp. crinitum ) has a wider range.
These taxa are well differentiated from morpholo-
gical point of view and belong to sections Bifida
(Arv.-Touv.) Clapham, Grovesiana Gottschl., Italica
(Fr.) Arv.-Touv., Hieracium (Pulmonaria Monnier),
Oreadea (Fr.) Arv.-Touv. Most of them are chasmo-
phytes confined to vertical cliffs or rocky slopes.
Their chorology and ecology testify the relict state
of the genus Hieracium in Sicily.
The island is indeed situated at the southern
border distribution of the genus (see map in
Brautigam, 1992) and its climatic conditions are
suitable for only a few taxa of Hieracium
(Gottschlich et al., 2013). However, among them,
the diploid H. lucidum , according to Pignatti (1979,
1982, 1994), ascribes the interesting role of likely
differentiation centre of the genus to Sicily.
Most of the taxa are endemic to restricted areas
(one population with an estimated area of occu-
pancy less than 1 0 km 2 ) in which periodical wild-
fires occur. Therefore, according to the IUCN
(2010) criteria for the conservation status assess-
ment, they should be classified as “Critically
Endangered” (CR). Their phytogeographical and
taxonomical relevance, together with the extreme
conservation status require special protection meas-
ures. Unfortunately, the current status and priorities
for conservation of the Hieracium taxa, as for many
other Sicilian endemics, are poorly known, and
consequently they are neglected by local admin-
istrations.
Taxonomic list
Hieracium busambarense Caldarella, Gianguzzi et
Gottschl., PL Biosystems, 148: 439. 2014.
H. sect. Grovesiana Gottschl.
Biological form. H ros/ H scap.
Phenology. Flowering from second half of
June to first decade of July; fruiting in July (Cal-
darella et al., 2014).
Distribution and Ecology. Chasmophyte
endemic to Rocca Busambra (PA) (CW-Sicily) (Fig.
2). Calcareous-dolomite vertical cliffs between
1500 and 1600 m a.s.l, in shaded localities (Cal-
darella et al., 2014).
Chromosome number. Unknown.
Conservation status. “Critically Endangered”
(CR): C2ab(i) (Caldarella et al., 2014).
Taxonomical notes. H. busambarense belongs
to the H. Sect. Grovesiana, recently described from
Italy (Gottschlich, 2009). Its distribution area is
located at the extreme southern limit of the
Apennines range of that section, therefore it could
be interpreted as an endemo-vicariant unit, probably
originated after the long geographical isolation of
the population on Rocca Busambra (Caldarella et
al., 2014). Among the taxa of H. Sect. Grovesiana,
H. busambarense appears very close to the
Calabrian endemic H. terraccianoi Di Gristina,
Gottschlich & Raimondo (2014), but it differs from
this species in having no spotted leaves, more acute
involucral bracts and in the bract indumentum (less
stellate hairs and more glandular hairs) (Di Gristina
et al., 2014).
Hieracium hypochoeroides subsp. montis-scuderii
Di Grist., Gottschl., Galesi, Raimondo et Cristaudo,
FI. Mediterr., 23: 49. 2013.
208
Emilio Di Gristina etalii
H. sect. Bifida (Arv.-Touv.) Clapham
Biological form. H ros.
Phenology. Flowering June; fruiting from June
to the first decade of July.
Distribution and Ecology. Chasmophyte
endemic to Mt Scuderi (ME) (NE-Sicily) (Fig. 3).
NW-facing carbonate rocks and vertical cliffs
between 1 145 and 1180m a.s.l.
Chromosome number. Unknown.
Conservation status. “Critically Endangered”
(CR): Bla+2a; C2a(ii).
Taxonomical notes. H. hypochoeroides s.l. is
a young aggregate of apomictic microtaxa with
often local distribution, that have evolved during
the post-glacial period. The map given by
Brautigam (1992, under the name H. wiesbauri-
anum) indicates a very disjunct area. Only in
southern France an extensive closed area exists. In
southern Europe one can only find local popula-
tions, most of them seem to be relict (Di Gristina
et al., 2014). H. hypochoeroides subsp. montis-
scuderii is also such relict endemic taxon.
Hieracium lucidum Guss., Index Sem. Hort.
Boccadifalco 1825: 6. 1825.
FI. sect. Italica (Fr.) Arv.-Touv.
Biological form. Ch suffr.
Phenology. Flowering from October to Novem-
ber; fruiting in November.
Distribution and Ecology. Chasmophyte
endemic to Mt Gallo (PA) (NW- Sicily) (Fig. 4).
NW-facing calcareous rocks and vertical cliffs
between 220 and 310 m a.s.l.
Chromosome number. 2n = 18 (Merxmiiller,
1975; Brullo & Pavone, 1978; Brullo et al., 2004).
Conservation status. “Critically Endangered”
(CR): Bla+2a; C2a(ii).
Taxonomical notes. H. lucidum is one of the
few diploid species in the whole genus. Therefore
it could be considered as a probable ancestor for
many European Hieracium taxa (Pignatti 1979,
1982, 1994).
Hieracium lucidum subsp. cophanense (Lojac.)
Greuter, Willdenowia, 37: 164. 2007.
= H. cophanense Lojac., FI. Sic. 2(1): 218.
1903.
H. sect. Italica (Fr.) Arv.-Touv.
Biological form. Ch suffr.
Phenology. Flowering from October to Novem-
ber; fruiting in November.
Distribution and Ecology. Chasmophyte
endemic to Mt Cofano and Mt Passo del Lupo (TP)
(NW-Sicily) (Fig. 5). NW-facing calcareous rocks
and vertical cliffs between 220-280 and 670-710 m
a.s.l.
Chromosome number. 2n = 18 (Brullo et al.,
2004; Geraci et al., 2007).
Conservation status. “Critically Endangered”
(CR): Bla+2a; C2a(ii).
Taxonomical notes. It differs from H. lucidum
in having few to moderately dense simple hairs on
the stem and on the margin, along the midrib at the
lower surface of the basal and cauline leaves.
Hieracium murorum subsp. atrovirens (Froel.)
Raimondo et Di Grist., Willdenowia, 37: 165. 2007.
= H. atrovirens Froel., in Candolle, Prodr. 7:
231. 1838.
H. sect. Hieracium (Pulmonaria Monnier)
Biological form. H scap/ H ros.
Phenology. Flowering June; fruiting from June
to first decade of July.
Distribution and Ecology. Endemic to the
Madonie Mountains (PA) (N-Sicily) (Fig. 6), along
the NW-facing carbonate rocks and stony slopes of
the Passo della Botte and Rocca di Mele (Petralia
Sottana, PA), in clearings of the beech forest,
between 1350 and 1580 m a.s.l.
Chromosome number. 2n = 3x = 27 (Geraci et
al., 2007).
Conservation status. “Critically Endangered”
(CR): B 1 a+2a; C2a(ii).
Taxonomical notes. In the past, the taxonomic
rank has been rather controversial. It was described
as an species (Froelich, 1838), but it was sub-
Diversity in the genus Hieracium Linnaeus s. str. (Asteraceae) in Sicily
209
Figures 2-7. Blooming individuals in nature of: Fig. 2) Hieracium busambarense (from Caldarella et al., 2014); Fig. 3) H.
hypochoeroicles subsp. montis-scuderii; Fig. 4) H. lucidum; Fig. 5) H. lucidum subsp. cophanense; Fig. 6) H. murorum
subsp. atrovirens; Fig. 7) H. pallidum.
sequently considered as synonym of H. murorum
Linnaeus (1753) (Fries, 1862; Belli, 1904) or of H.
glaucinum Jordan (1848) (Zahn, 1921;Fiori, 1928).
Recently, the Sicilian population has been con-
sidered distinct and treated at subspecific rank of
H. murorum (Raimondo & Di Gristina, 2007).
Hieracium pallidum Biv., in Bivona-Bernardi,
Nuove piante: 11. 1838.
H. sect. Grovesiana Gottschl.
Biological form. H ros/ H scap.
Phenology. Flowering from second half of June
to first decade of July; fruiting in July.
Distribution and Ecology. Chasmophyte
endemic to Mt Etna (CT) (E-Sicily) (Fig. 7).
Shaded volcanic rocks and stony slopes of Mt.
Pomiciaro, Mt Zoccolaro and Serra del Salifizio fa-
cing the Valle del Bove (Zafferana Etnea, CT),
between 1550 and 1900 m a.s.l.
Chromosome number. 2n = 4x = 36 (Brullo et
al., 2004; Di Gristina et al., 2005).
Conservation status. “Critically Endangered”
(CR): B 1 a+2a; C2a(ii).
Taxonomic al notes. According to Greuter
(2008), it should be placed in the “collective
species” (Zahn, 1921-1923) H. schmidtii. Never-
theless, the presence of 2 cauline leaves with
winged petioles (in H. schmidtii s.l. 0-1 not winged
leaf per stalk) allow to treat it as a local endemic
species to Sicily belonging to H. sect. Grovesiana
(Gottschlich et al., 2013).
Hieracium pallidum subsp. aetnense Gottschl.,
Raimondo et Di Grist., PI. Biosystems, 147: 826.
2013.
H. sect. Grovesiana Gottschl.
210
Emilio Di Gristina etalii
Biological form. H scap.
Phenology. Flowering from second half of
June to first decade of July; fruiting in July.
Distribution and Ecology. Endemic to Mt
Etna (CT) (E-Sicily) (Fig. 8). Volcanic soil, on the
border and in clearings of scrubland in a very
restricted area on Mt Pomiciaro (Zafferana Etnea,
CT), between 1580 and 1650 m a.s.l.
Chromosome number. 2n = 4x = 36 (Di Gristina
et al., 2014).
Conservation status. “Critically Endangered”
(CR): B2ab(iii, v); C2a(ii), D.
Taxonomical notes. Closely related to H.
pallidum , but different by morphology of basal
leaves (more lanceolate and dentate), number of
cauline leaves (up to 3) and by peduncles and bracts
indumentum (more simple hairs and stellate hairs
only at the margin of the bracts).
Hieracium racemosum subsp. crinitum (Smith)
Rouy, FI. France, 9: 410. 1905.
= H. crinitum Sm., FI. Graec. Prodr., 2: 134.
1813.
= H. crinitum var. caulescens Lojac., FI. Sic.,
2(1): 219. 1903; H. crinitum var. eriostachyum
Lojac., FI. Sic., 2(1): 219. 1903; H. racemosum
subsp. todaroanum Zahn, in Engler, Pflanzenr., 79:
979. 1922.
H. sect. Italica (Fr.) Arv.-Touv.
Biological form. H scap/ H ros.
Phenology. Flowering from second half of
August to first decade of November; fruiting from
September to second decade of November.
Distribution and Ecology. Corsica, Italy,
Balkan Peninsula and Turkey (Fiori, 1928; Pignatti,
1982). In Sicily (Fig. 9), the taxon frequently occurs
in shaded stony slopes and in clearings of woods on
the main mountains of north-east (Nebrodi,
Peloritani and Etna) and on the islands of Salina and
Lipari (Lojacono, 1903), between 350 and 1750 m
a.s.l.
Chromosome number. 2n = 3x = 27 (Brullo et
al., 1997, 2004; Geraci et al., 2007).
Conservation status. “Least Concern” (LC).
Taxonomical notes. Taxon highly polymorph-
ous. Sicilian and southern Italy populations appear
very differentiated and need critical revision.
Hieracium racemosum subsp. pignattianum (Rai-
mondo et Di Grist.) Greuter, Willdenowia, 37: 171.
2007.
= H. pignattianum Raimondo & Di Grist., PI.
Biosystems, 17: 314. 2004.
H. sect. Italica (Fr.) Arv.-Touv.
Biological form. H scap/ H ros.
Phenology. Flowering from second half of
August to October; fruiting from September to first
decade of November.
Distribution and Ecology. Endemic to the
Madonie Mountains (PA) (N-Sicily) (Fig. 10),
along the NW-facing carbonate rocks and stony
slopes of Mt Mufara (Isnello, Polizzi Generosa,
Petralia Sottana, PA), Mt Quacella (Polizzi Gen-
erosa, PA), Mt Daino, Cozzo del Filatore, Pizzo
dellTnferno and Rocca di Mele (Petralia Sottana,
PA), in clearings of the beech forest, between 1300
and 1700 m a.s.l.
Chromosome number. 2n = 3x = 27 (Raimondo
& Di Gristina, 2004).
Conservation status. “Vulnerable” (VU): Bla
+ 2a.
Taxonomical notes. Similar to subsp. crinitum
but the two subspecies show marked differences
regarding indumentum, leaf morphology and size
of the bracts (Raimondo & Di Gristina, 2004).
Hieracium schmidtii subsp. madoniense (Rai-
mondo & Di Grist.) Greuter, Willdenowia, 37: 173.
2007.
= H. madoniense Raimondo & Di Grist., Boc-
conea, 141: 86. 2007.
H. sect. Oreadea (Fr.) Arv.-Touv.
Biological form. H ros.
Phenology. Flowering second half of June;
fruiting from June to first decade of July.
Distribution and Ecology. Chasmophyte
endemic to the Madonie Mountains (PA) (N-Sicily)
(Fig. 11). NW-facing carbonate rocks and stony
Diversity in the genus Hieracium Linnaeus s. str. (Asteraceae) in Sicily
211
Figure 8-12. Blooming individuals in nature of: Fig. 8) Hieracium palliclum subsp. aetnense; Fig. 9) H. racemosum
subsp. crinitum; Fig. 10) H. racemosum subsp. pignattianum; Fig. 11) H. schmidtii subsp. madoniense; Fig. 12) H.
symphvtifol ium .
slopes of Rocca di Mele (Petralia Sottana, PA), in
clearings of the beech forest, between 1520 and
1700 m a.s.l.
Chromosome number. 2n = 3x = 27 (Di Gristina
et al., 2005).
Conservation status. “Critically Endangered”
(CR): B 1 a+2a; C2a(ii).
Taxonomical notes. For a long time confused
with H. pallidum but easily distinct by leaf, stem,
bract indumentum (short simple crisp hairs) and
more lanceolate and dentate leaves (Raimondo &
Di Gristina, 2007).
Hieracium symphytifolium Froel., in Candolle, 7:
232. 1838.
= H. siculum Guss., FI. Sicul. Syn., 2(1): 404.
1844; H. siculum var. minus Guss., FI. Sicul. Syn.,
2(1): 404. 1844.
H. sect. Italica (Fr.) Arv.-Touv.
Biological form. H ros/ H scap.
Phenology. Flowering from end of June to July;
fruiting in July.
Distribution and Ecology. Chasmophyte
endemic to the Madonie Mountains (PA) (N-Sicily)
(Fig. 12). NW-facing carbonate rocks and stony slopes
of the highest reliefs, between 1250 and 1800 m a.s.l.
Chromosome number. 2n = 4x = 36 (Brullo et
al., 2004; Di Gristina et al., 2006).
Conservation status. “Critically Endangered”
(CR): Bla+2a; C2a(ii).
Taxonomical notes. The status of this plant has
only recently been clarified. According to Zahn
(1921-1923), the taxon represented an “interme-
diate species” between H. lucidum and H. crinitum
{Hieracium racemosum subsp. crinitum ). However,
morphological and genetic studies showed that it
is not a hybrid, but an independent species (Di
Gristina et al., 2006).
212
Emilio Di Gristina etalii
Key of the Sicilian taxa
1. Achenes dark when mature. Flowering June-
July 2
Achenes pale when mature. Flowering end of
August-November 3
2. Bracts with rather dense glandular hairs and
sparse or no simple hairs
H. murorum subsp. atrovirens
- Bracts with few to moderately dense glandular
hairs and moderately dense to rather dense
simple hairs 4
4. Cauline leaves 3-6. Bracts 0.9- 1.3 mm wide
H. symphytifolium
Cauline leaves 0-3. Bracts 0.4-1 mm wide
5
5. Plants with 1-4 mm long, denticulate soft or
crisp simple hairs 6
- Plants with 4-10 mm long, denticulate rigid
simple hairs 7
6. Basal leaves unspotted; cauline leaves 1-2(3)
H. busambarense
- Basal leaves few to intensely dark spotted;
cauline leaves 0-1 8
8. Leaves denticulate above, on the margin and
along the midrib with crisp simple hairs
H. hypochoeroides subsp. montis -scuderii
- Leaves dentate to serrate-dentate only on the
margin and along the midrib with crisp simple
hairs H. schmidtii subsp. madoniense
7. Basal leaves ovate, denticulate, truncate or
cuneate at base; cauline leaves 2 H. pallidum
- Basal leaves lanceolate, denticulate or serrate-
dentate, long attenuate at base; cauline leaves
up to 3 H. pallidum subsp. aetnense
3. Leaves coriaceous, glabrous or with few to
moderately dense simple hairs 9
- Leaves soft with moderately dense to rather
dense simple hairs 10
9. Leaves glabrous H. lucidum
- Leaves with few to moderately dense simple
hairs on the margin and along the midrib
H. lucidum subsp. cophanense
10. Basal leaves with moderately dense stellate
hairs on both surfaces. Bracts 0.7-1 mm wide
H. racemosum subsp. pignattianum
11. Basal leaves without stellate hairs on both
surfaces. Bracts 0.8- 1.3 mm wide
H. racemosum subsp. crinitum
ACKNOWLEDGEMENTS
Financial support by the International Founda-
tion pro Herbario Mediterraneo and by Universita
degli Studi di Palermo (Fondi di Ateneo per la
Ricerca) is acknowledged.
REFERENCES
Aghababyan M., Greuter W., Mazzola P. & Raimondo
F.M., 2008. Typification of names of Compositae
taxa described from Sicily by Michele Lojacono
Pojero. Flora Mediterranea, 18: 513-528.
Belli S., 1904. Hieracium. In: Fiori A. & Paoletti G.
1904. Flora Analitica d’ltalia, Vol. 3. Tipografia del
Seminario, Padova, 442-505.
Bivona-Bernardi A., 1838. Nuove piante inedite.
Lorenzo Dato, Palermo, 23 pp.
Brautigam S., 1992. Hieracium. In: Meusel H. & Jager
E.J. 1992. Vergleichende Chorologie der zentraleur-
opaischen Flora 3. G. Fischer, Jena, Stuttgart, New
York, 325-333, 550-560.
Brallo S., Pavone P, Terrasi M.C. Zizza A., 1977.
Numeri cromosomici per la Flora Italiana: 299-3 14.
Informatore Botanico Italiano, 9: 57-61.
Brallo S. & Pavone P, 1978. Numeri cromosomici per
la Flora Italiana: 464-483. Informatore Botanico
Italiano, 10: 248-265.
Brallo S., Campo G. & Romano S., 2004. Indagini cito-
tassonomiche sul genere Hieracium L. (Asteraceae)
in Sicilia. Informatore Botanico Italiano, 36: 481-
485.
Caldarella O., Gianguzzi L. & Gottschlich G., 2014.
Hieracium busambarense, a new species of the sect.
Grovesiana (Asteraceae) from Sicily (Italy). Plant
Biosystems, 148: 439-443.
Diversity in the genus Hieracium Linnaeus s. str. (Asteraceae) in Sicily
213
Chrtek J. jr., Mraz P. & Severa M., 2004. Chromosome
numbers in selected species of Hieracium s.str.
(. Hieracium subgen. Hieracium) in the Western
Carpathians. Preslia, 76: 119-139.
Chrtek J. jr., Mraz P. & Sennikov A., 2006. Hieracium
xgrofae, a rediscovered diploid hybrid from the
Ukrainian Carpathians. Biologia, section Botany, 61 :
365-373.
Di Gristina E., Geraci A. & Raimondo F.M., 2005.
Osservazioni citotassonomiche su popolazioni
siciliane afferenti a Hieracium pallidum (Asteraceae).
Informatore Botanico Italiano, 37: 26-27 .
Di Gristina E., Geraci A. & Raimondo F.M., 2006.
Biosystematic investigation on Hieracium symphyti-
folium Froel. (Asteraceae). Bocconea, 19: 275-286.
Di Gristina E., Raimondo F.M., Domina G. & Gottschlich
G., 2012. Typification of eight names in Hieracium
(Asteraceae). Taxon 61: 1317-1320.
Di Gristina E., Gottschlich G., Galesi R., Raimondo F.M.
& Cristaudo A., 2013. Hieracium hypochoeroides
subsp. montis-scuderii (Asteraceae), a new endemic
subspecies from Sicily (Italy). Flora Mediterranea,
23:49-55.
Di Gristina E., Gottschlich G. & Raimondo F.M., 2014.
Taxonomic remarks on Hieracium sartorianum var.
lucanicum Arv.-Touv. (Asteraceae), a little known
taxon of Cilento (Campania, Southern Italy). Nordic
Journal of Botany, Doi: 10. Ill 1/njb. 00755.
Di Gristina E., Gottschlich G. & Raimondo F.M., 2014.
Hieracium terraccianoi (Asteraceae), a new species
endemic to the Pollino National Park (Southern
Italy). Phytotaxa, 188: 55-60.
Di Gristina E., Domina G. & Geraci A., 2014. Reports
(1837). In: Kamari G., Blanche C. & Siljak- Yakovlev
S., 2014. Mediterranean chromosome number reports
- 24. Flora Mediterranea, 24: 283-284.
Fiori A., 1928. Nuova Flora Analitica d’ltalia., Vol. 2. M.
Ricci, Firenze, 1120 pp.
Fries E., 1862. Epicrisis Generis Hieraciorum. Edquist
& Berglund, Upsaliae, 159 pp.
Froelich J.A., 1838. Hieracium F. In: Candolle A.P de
1838. Prodromus systematis naturalis regni veget-
abilis, Vol. 7. Treuttel & Wiirz., Paris, 198-240.
Geraci A., Di Gristina E. & Schicchi R., 2007. Reports
(1640-1644). In: Kamari G., Blanche C. & Garbari
F. 2007. Mediterranean chromosome number reports
- 17. Flora Mediterranea, 17: 314-319.
Gottschlich G., 2009. Die Gattung Hieracium F. (Com-
positae) in der Region Abruzzen (Italien). Eine flor-
istisch-taxonomische Studie. Stapfia, 89: 1-328.
Gottschlich G., Raimondo F.M. & Di Gristina E., 2013.
Hieracium pallidum subsp. aetnense (Asteraceae), a
new subspecies from Sicily (Italy), with notes on the
taxonomy of H. pallidum Biv. Plant Biosystems, 147:
826-831.
Greuter W. & Raab-Straube E. Von, 2007. Notulae
ad floram euro-mediterraneam pertinentes No. 25,
Euro+Med Notulae, 3. Willdenowia, 37: 139-189.
Greuter W., 2008. Med-Checklist 2. Dicotyledones
(Compositae). OPTIMA Secretariat, Palermo, Berlin,
798 pp.
Gussone G., 1825. Index seminum anni 1825 que ab
Horto Regio in Boccadifalco pro mutua commuta-
tione exibentur. Palermo, 12 pp.
Gussone G., 1844. Hieracium E. In: Gussone G. 1844.
Florae siculae synopsis, Vol. 2. Tramater, Neapoli,
402-405.
IUCN 2010. The IUCN Red Fist of threatened species,
version 2010.4. IUCN Red Fist Unit, Cambridge,
UK. Available: http://www.iucnredlist.org.
Jordan A., 1848. Catalogue des Graines recoltees au
Jardin botanique de la Ville de Dijon en 1848,
offertes en echange. Dijon (Divionensis), 22 pp.
Finnaeus C., 1753. Species Plantarum. F. Salvius,
Stockholm, 1200 pp.
Fojacono Pojero M., 1903. Hieracium F. In: Fojacono
Pojero M. 1903. Flora Sicula, Vol. 2(1). Stab.
Tipografico Virzi, Palermo, 214-222
Merxmiiller H., 1975. Diploide Hieracien. Anales del
Instituto Botanico A. J. Cavanilles, 32: 189-196.
Mraz P, Chrtek J. jr. & Kirschner J., 2001. Genetic va-
riation in the Hieracium rohacsense Group. {Hiera-
cium sect. Alpina). Phyton, 41: 269-276.
Pignatti S., 1979: Plant geographical and morphological
evidences in the evolution of the Mediterranean flora
(with particular reference to the Italian represent-
atives). Webbia, 34: 243-255.
Pignatti S., 1982. Flora d’ltalia, Vol. 3. Edagricole,
Bologna, 780 pp.
Pignatti S., 1994. Ecologia del Paesaggio. Utet, Torino,
228 pp.
Raimondo F.M. & Di Gristina E., 2004. Hieracium
pignattianum (Asteraceae), a new species from the
Madonie Mountains (N-Sicily). Bocconea, 17: 313-
324.
Raimondo F.M. & Di Gristina E., 2007a: Hieracium
murorum subsp. atrovirens (Froel.) Raimondo & Di
Gristina, comb. & stat. nov. In Greuter W. & Raab-
Straube E. Von 2007. Notulae ad floram euro-
mediterraneam pertinentes No. 25, Euro+Med
Notulae, 3. Willdenowia, 37: 165.
Raimondo F.M. & Di Gristina E., 2007b. Hieracium
madoniense (Asteraceae), a new species from Sicily.
Plant Biosystems, 141: 86-92.
Raunkiaer C., 1934. The life forms of plants and statist-
ical plant geography. The Clarendon Press, Oxford,
632 pp.
Rich T.C.G., Mcdonnel E.J. & Fledo M.D., 2008.
Conservation of Britain's biodiversity: the case of
Hieracium cyathis (Asteraceae) and its relation to
214
Emilio Di Gristina etalii
other apomictic taxa. Botanical Journal of the
Linnean Society, 156: 669-680.
Rouy G., 1905. Flora de France, 9. Asnieres & Paris, 410
pp.
Smith J.E., 1813. Hieracium L. In: Smith J.E. 1813.
Florae Graecae Prodromus, Vol. 2. R. Taylor &
Soc., London, 132-135.
ZahnK.H., 1921-23. Hieracium. In: EnglerA. 1921-23.
Das Pflanzenreich. Engelmann, Leipzig, 75(IV.280):
1-288; 76(IV.280): 289-576, 77(IV.280): 577-864
(1921); 79(IV.280): 865-1146 (1922); 82(IV.280):
1147-1705 (1923).
Biodiversity Journal, 2015, 6 (1): 215-218
Monograph
Lycopodiidae for the “Flora Critica d’ltalia”: material and
methods
Angelo Troia 1 *, Francesco Maria Raimondo 1 & Werner Greuter 2
1 D ip artim e n to STEBICEF, Sezione di B otanica ed Ecologia Vegetale. Universita degli Studi di Palermo, via Archirafi 3 8. 90 1 23
P ale rm o . Italy
2 Herbarium M editerraneum Panorrn itanum . Universita degli Studi di Palermo, Orto Botanico, via Lincoln 2, 90133 Paler mo. Italy
Corresponding author, e-mail: an g e lo .tro ia @ unipa.it
ABSTRACT Procedures are presented that were followed during the preparation of the first pteridophyte
family treatments for the “Flora Critica d’ltalia”: Lycopodiaceae.Isoetaceae, Selaginellaceae.
The work was mainly based on the study of literature and herbarium specimens. In some
cases SEM observation of spores has proved useful. Data collected from herbarium specimens
and other verified sources were loaded into a database, from which a distribution map was
prepared for each taxon. Several preliminary papers have been published, and foreach family
a taxonomic conspectus, with type designations, maps and an identification key, has been
prepared. The treatment of these three families for the “Flora Critica d’ltalia” (in Italian) is
about to be published or (Isoetaceae) has already been published.
KEY WORDS Italy; flora; vascular plants; pteridophytes; lycopodiophytes; herbarium; SEM ; taxonomy.
Received 21.12.2014; accepted 12.03.2 0 15; printed 30.03.2015
Proceedings of the 2nd Interna tional Congress “Speciation and Taxonomy”, M ay 16 th -18th 2014, Cefalu-Castelbuono (Italy)
INTRODUCTION
The need for an up-to-date “Flora Critica
d’ltalia” has long been recognized. About 10 years
ago the Societa Botanica Italiana (Italian Botanical
Society) endorsed the project and prepared a model
(Pignotti, 2006). The project, after the estab-
lishment of the Fondazione per la Flora Italiana
(Foundation for the Italian Flora), has now entered
its active phase of implementation: a few months
ago the first preliminary results have been publi-
shed (Cecchi & Selvi, 2014; Troia & Greuter,
20 14). Procedures are presented that w ere follow ed
during the preparation of the first pteridophyte
family treatments for the “Flora Critica d’ltalia”:
L y c o p o d iac e ae , Isoetaceae, Selaginellaceae.
Accor ding to recent literature, these three famil-
ies (known as lycopodiophytes or “ ly c o p h y te s ” )
constitute the Lycopodiidae, the first of the five
major subclasses of pteridophytes recognized by
Christenhusz et al. (2011) and Christenhusz &
Chase (2014) (Fig. 1).
MATERIAL AND METHODS
The treatment is mainly based on the study of
literature and herbarium specimens. It encompasses
all Lycopodiidae taxa that grow spontaneously in
the National territory, either native or naturalized.
We studied all Italian and selected foreign
Lycopodiidae specimens kept in the Herbarium
Centrale Italicum (FI) and Herbarium M editer-
raneum Panormitanum (PAL, including PAL-Gr);
several specimens, notably original material for
2 1 6
Angelo Troia et alii
relevant names, were supplied by the Herbarium of
the Botanischer Garten und Botanisches Museum
Berlin-Dahlem (B ). Each specimen has been docu-
mented photographically. In addition, we examined
high-resolution digital images, available online or
provided on request, from the following herbaria:
APP, BOLO, CAT, GDOR, MFU, MRSN, MSNM ,
PAD, RO, ROV, SIENA, TO, TR, and K, LINN, P,
PH, UPS (abbreviations according to Thiers, 2014),
and had the presence of selected specimens verified
by colleagues in others (e.g. MI). Specimens con-
served in the private collections of Bonafede
(Bologna, Italy), Selvi (Florence, Italy), Tondi
(Rome, Italy) have also been studied.
For mapping the distribution, reports based on
photographs have been considered only when
species identification was not in doubt; in particu-
lar, data from popular websites such as Acta
Plantarum (www.actaplantarum.org) have been
taken into consideration. However, literature
reports not supported by herbarium vouchers have
been discarded for mapping purposes; comments
have been added for those of special historical or
phy togeographical interest.
Data (both original data and metadata) were
loaded into a specific spreadsheet. Data fields in-
cluded not only the scientific name and geograph-
ical parameters but also biological aspects, so as to
enable future searches on, for example, phenology
or altitudinal range. All specimens with sufficient
locality data have been georeferenced and plotted
on a base map of Italy (Cecchi & Selvi, 2014).
Lycopodiidae
Ophioglosstdae
Equisetidae
Marattiidae
Polypodl idae
Pi n i dae
Gnetidae
Gink goo idae
tycadidae
Magnoliidae
Figure 1 . The main subclasses of living vascular plants, cladogram based on Schuettpelz & Pryer (2 0 0 8), Pryer et al. (2009),
and Grewe et al. (2013); clade names according to Christenhusz et al. (201 1) and Christen husz & Chase (2014) for pterido-
phytes. Chase & Reveal (2009) for sperm atophytes. Figures 2-4. Example of SEM images prepared for the “Flora Critica
d'ltalia”: IsOCtCS gymVlOCQ,rpCl (G ennari) A . Braun, megaspores and microspores from the type specimen in TO (see Troia
& Greuter, 2014). Figure 2: megaspore in proximal view. Figure 3: detail of Fig. 2. Figure 4: microspores.
Lycopodiidae for the “Flora Critica d’ltalia”: materials and methods
217
Figure 5. Example of a distribution map prepared for
the Flora Critica d’ltalia: Isoetes longissima Bory.
Currently, a map is used that shows physiogra-
phical rather than administrative territorial units,
following the model proposed by Cecchi & Selvi
(2014). In a firstphase, only a selection of specimens
have been mapped: 1 to 3 preferably recent speci-
mens for each territorial units, so as to avoid excess-
ive c ro w d in g o f th e d o ts ; territo rial u n its in which th e
species in question is present were shaded.
For some critical taxa, particularly in the genus
Isoetes, scanning electron micrographs were pro-
duced to illustrate and document megaspore and
microspore features (Figs. 2-4).
RESULTS AND CONCLUSIONS
W ith regard to Isoetaceae, a synthetic paper w ith
a taxonomic conspectus, type designations and an
identification key has been published in the journal
Plant Biosystems (Troia & Greuter, 2014), and
similar papers for Lycopodiaceae and S e 1 a -
ginellaceae, including distribution maps (Fig. 5),
are ready for publication. Preliminary results were
presented by Troia et al. (2012, 2014b) and Troia &
Greuter (2013), as well as a paper with a SEM study
of spores of the IsOCtCS longissilflCl group (Troia et
al. 2014a).
The treatment (in Italian) for the Flora Critica
d’ltalia of these three families, following guidelines
prepared by the Editorial Committee, has just been
published (Isoetaceae: Troia & Greuter, 2015) or is
about to be published (Lycopodiaceae and Selagi-
nellaceae: Troia & Greuter, in prep.).
ACKNOWLEDGEMENTS
We thank the staff of the consulted herbaria for
their assistance, Lorenzo Cecchi (FI) for his valu-
able support during the preparation of the distribu-
tion maps, and Carmela Di Liberto (D ip artim en to
STEBICEF, Universita degli Studi di Palermo) for
technical assistance with scanning electron micro-
scopy. This study is part of the “Flora Critica d’
Italia” project and as such was funded by the
Societa Botanica Italian a onlus, the Fondazione per
la Flora Italiana, and the International Foundation
Pro Herbario Mediterraneo.Additional support was
provided by the Universita degli Studi di Palermo
(Fondi di Ateneo per la Ricerca).
REFERENCES
Cecchi L. & Selvi F., 2014. A synopsis of Boraginaceae
subfam . H y d ro p h y 1 1 o id e a e and H e lio tro p io id e a e in
Italy. Plant Biosystems, 148: 2-12.
Chase M .W. & Reveal J.L., 2009. A phylogenetic classi-
fication of the land plants to accompany APG III. Bo-
tanical Journal of the Linnean Society, 161: 122-127.
Christenhusz M.J.M. & Chase M.W., 2014. Trends and
concepts in fern classification. Annals of B otany, 113:
571-594.
Christenhusz M.J.M., Zhang X ,-C . & Schneider H .,
2011. A linear sequence of extant families and genera
of lycophytes and ferns. Phytotaxa, 19: 7-54.
Grewe F., Guo W ., G ubbels E.A., Hansen A.K. & M ower
J.P., 2013. Complete plastid genomes from Ophio-
glossum californicum, Psilotum nudum , and Equis-
etum hyemale reveal an ancestral land plant genome
structure and resolve the position of EqilisetClleS
among monilophytes. BMC Evolutionary Biology,
13: 8.
Pignotti L. (Ed.), 2006. Progetto per una Flora critica
dell’ Italia. Societa Botanica Italiana, Firenze, 147 pp.
Pryer K .M ., Smith A .R . & Rothfels C ., 2009. Polypo-
2 1 8
Angelo Troia et alii
diopsida Cronquist, Takht. & Zimmerm. 1966. Ferns.
Version 14 January 2009 (under construction).
http://tolweb.org/Polypodiopsida/20615/2009. 01. 14
in The Tree ofLife Web Project, http://tolweb.org/
Schuettpelz E. & Pryer K.M ., 2008. Fern phylogeny. In:
Ranker T.A. & Flaufler C.H. (Eds.), Biology and
Evolution of Ferns and Lycophytes. Cambridge
University Press, Cambridge, 395-416.
Thiers B ., 2014. Index herbariorum: A global directory
of public herbaria and associated staff. New York
Botanical Garden’s Virtual Herbarium. h ttp ://s w ee t-
gum.nybg.org/ih/ [continuously updated].
Troia A. & Greuter W., 2013. Towards a Critical Flora of
Italy. Assessing the Lycopodiophyta. In: Abstracts of
the XIV OPTIMA Meeting, Palermo, September 9-
15,2013 (ISBN: 978-88-903108-8-1): 151.
Troia A. & Greuter W., 2014. A critical conspectus of Ita-
lian IsOeteS (Isoetaceae). Plant Biosystems, 148: 13-
20 .
Troia A. & Greuter W ., 2015. Isoetaceae (vers. 1.0). In:
Peruzzi L., Cecchi L., Cristofolini G., Domina G.,
Greuter W., Nardi E . , Raimondo F.M., Selvi F. &
Troia A. (Eds.), Flora critica d’ltalia. Fondazione per
la Flora Italiana, Firenze. Published online on
25 february 2015 at: http://www.floraditalia.it/pdf/
Isoetaceae.pdf
Troia A., Greuter W., Nardi E. & Raimondo F.M ., 2012.
Contributo alia Flora Critica d’ltalia: i generi della
famiglia Lycopodiaceae. In: Riassunti di relazioni,
com unicazioni e poster del 107° Congresso
della Societa Botanica Italiana. Benevento, 18-22
Settembre 2012, 142.
Troia A., Raimondo F.M . & Campisi P., 2014a. The ISOC-
teS longissimCl complex (Isoetaceae) in Italy: obser-
vations on the morphology of spores and leaves, and
taxonomic implications. Phytotaxa, 174: 149-156.
Troia A., Raimondo F.M. & Greuter W., 2014b. On the
presence, distribution and conservation status of
Lycopodium Icigopus (Lycopodiaceae) in Italy. In:
109° Congresso della Societa Botanica Italiana, In-
ternational Plant Science Conference, Firenze, 2-5
September 2014, Proceedings: 64.
Biodiversity Journal, 2015, 6 (1): 219-244
Monograph
Hotspot of new megafauna found in the Central Amazon
(Brazil): the lower RioAripuana Basin
Marc G.M.van Roosmalen
‘MVRS Marc van Roosmalen Stichting, Leiden, The Nether lands; e-mail: marc.mvrs@gmail.com
ABSTRACT Here I announce the discovery of a whole new ecosystem in the central-southern part of the
Brazilian Amazon: the RioAripuana Basin. Overall, it seems to have created mo re ecological
niches than any other river basin in the Amazon, in particular so to aquatic and non-volant
terrestrial mammals. This is plausibly explained for by the unique geo-morphological history
of the region. During the Pliocene and Early Pleistocene the entire area to the southeast of
the Rio Madeira contained one huge clear-water system that was drained toward the south
into the Atlantic Ocean. In the course of several million years a biome quite different from
the rest of Amazonia could evolve in this drainage system. Living relicts from ancient times
that happened to survive in isolation here, are: a dwarf manatee here described as TrichechllS
pygmueUS n. sp., a dolphin locally called “boto roxo” that is suspected to be closer related to
marine Rio Plata dolphins Pontoporici blciiiivillei (Gervais et d'Orbigny, 1 844) than to
Amazonian dolphins of the genus Illici (d'Orbigny, 1834), a black dwarf tapir ( TcipivilS
pygmaeus Van Roosmalen, 2013, with T. k.Clb OinCllli C ozzuol et al., 2013 as junior name), a
dwarf marmoset Ccillibcllci hWTlilis Van Roosmalen et Van Roosmalen, 2003, a new mono-
specific genus of C allitrich id ae that stands at the base of the phylogenetic tree of all extant
m arm o sets (i.e ., Cebuellci Gray, 1866, Mico Lesson, 1840, and Cullithrix Erxleben, 1777), a
giant striped paca here described as AgOllti silvClgCLVCicie n. sp., and an arboreal giant anteater
spotted in the wild but remains to be collected and described (MymieCOphagCl n. sp.). A
number of other, more advanced mammalian species discovered in the Rio Aripuana Basin,
among which a third specie s of brocket here d escribed as MCLZCIJTICI tieJlllOVeni n. sp., evolved
after a dramatic vicariance took place about 1-1.8 MYA (million years ago), the break-through
of the continental watershed by the proto-Madeira River during one of the glacial epochs of
the Middle Pleistocene. It marked the birth of the modern fast- fl owing Rio Madeira, in terms
of total discharge the biggest tributary of the Amazon proper and the second strongest river
barrier in the entire Amazon Basin. Furthermore, current threats to the environment in this
sparsely inhabited and poorly explored river basin will be addressed. We intend to have this
‘lo st w orld ’ preserv ed as a UNESCO N atural W orld H eritage Reserve through the div ulgation
of new, hitherto not yet identified mammals that it appears to harbor.
KEY WORDS Brazilian Amazon; nova species; Rio Aripuana Basin.
Received 15.06.2014; accepted 12.12.2014; printed 30.03.2015
Proceedings of the 2nd Internatio nal C ongre ss “S p ec iatio n and Taxonomy”, May 1 6 th - 1 8 th 2014, Cefalu-Castelbuono (Italy)
INTRODUCTION any other basin in the Amazon, in particular to
aquatic and terrestrial mammals. In terms of spe-
cies evolution and p h y lo g e o g r ap h y the Rio
Aripuana Basin distinguishes itself from Amazo-
Overall, the Rio Aripuana Basin (Fig. 1) seems
to have created more ecological niches than
220
Marc G.M. van Roosmalen
nia west of the Rio Madeira and north of the Rio
Amazonas by harboring:
- Five sympatric species of peccaries (Tayas-
s u id ae : TayClSSU G. Fisher, 1814; PeCClri L inn aeu s ,
1 75 8 ), instead of two species elsewhere in the
Amazon;
- Three sympatric species of brocket deer
(C ervidae: MflZCUflCl Rafinesque, 18 17), including a
new species we here describe as M. tieilhoveilin. sp.,
instead of two species elsewhere in the Amazon;
- Two sympatric species of coati (Procyonidae:
NdSUCl Storr, 1 780), including a newly identified
red-coated pair-living coati we here resurrect as N.
solitaria Schinz (ex Wied, MS), 1821, as Spix &
Martius (1 823-1 83 1 ) refer to it in their account
“Reise in Brasilien in den Jahren 1 8 1 7-1 820”,
instead of only one gregarious species elsewhere in
the A m azon ;
- Two sympatric species of giant anteater
(E den tata: MyVJYieCOphcigCL Linnaeus, 1758),includ-
ing a new species being tree-dwelling and climbing
by its hind feet, instead of only one ground-
dwelling species elsewhere in the Amazon;
- Two sympatric species of lowland tapir (Ta-
piridae: TapiruS Briinnich, 1 772), including a new
species in 2013 described by me as T. pygtnaeUS
(w ith T. kab OVncmi Cozzuol et al., 2013 as a junior
synonym), instead of only one species elsewhere in
lowland Amazonia;
- Two sympatric species of jaguar (Felidae: PcM-
thera. Oken, 1 8 1 6 ) , including a new larger-sized
species reported to hunt in pairs, its coat being all-
black but a white throat, instead of only one species
elsewhere in the Amazon;
- Two sympatric species of paca (Rodentia:
AgOUti Lacepede, 1 799), including a new species
here described as A. silVQgClTcicie n. sp., being
larger-sized, its coat orange-brown with white
stripes instead of dots, instead of only one species
elsewhere in the Amazon;
- Two sympatric species of porcupine (Rodentia-
Erethizontidae: Coendu L acepede, 1 799), includ-
ing a new species described as C. ( SphiggUTUS )
roosmalenorum Voss et Da Silva, 2001 belonging
to the vestltUS gro up of sm all-bodied dwarf porcu-
pines form erly known only from the Andean Moun-
tains in Colombia, instead of only one species
elsewhere in lowland Amazonia;
- Two sympatric species of woolly monkey
(Primates: LagOtHvix FI um b oldt, 1 8 1 2 ), including
L. nigra n. sp. that is all-black, small, and ranging
in atypical small social groups (Van Roosmalen,
2013a; 2014; 2015; Van Roosmalen & Van Roos-
malen, 2014), instead of only one species elsewhere
in lowland Amazonia;
- Two sympatric species of Amazonian mar-
moset (Primates, C allitrichidae), including a new
species, first described as Callithrix huinilis Van
Roosmalen, Van Roosmalen, Mittermeier et De
Fonseca, 1998, and later as a new genus, Callibella
Van Roosmalen et Van Roosmalen, 2003, which is
much smaller, does not show any territorial beha-
vior, and occurs in sympatry with MicO WianicOVen-
SIS Van Roosmalen, Van Roosmalen, Mittermeier
et Rylands, 2000, instead of only one species
elsewhere in the Amazon east of the Rio Madeira;
- Two sympatric species of a large-bodied river
dolphin (D elphinidae ), including a new species
locally called “boto roxo” that we suspect to belong
to the m arine genus P ontopovia G ray, 1 8 7 0 , it being
smaller, having an overall bluish-grey colored skin,
lacking a distinct melon (and therefore maybe
foraging by eye-sight and not by echo location),
living in pairs with a single offspring, and restricted
to the clear-water habitat of the lower Rio A rip u ana,
instead of only one species elsewhere in the
Amazon Basin;
- Two sympatric species of freshwater manatee,
including a new species described in this work as
TrichechuS pygniaeus n. sp., it being less than half
the size and one-fifth of the body weight of com-
mon Amazonian manatees T. inunguis (N atterer,
1 8 8 3 ), and its skin deep black instead of grey,
instead of only one species elsewhere in lowland
Amazonia downstream of rapids and waterfalls;
- A number of newly identified large-fruited,
large-seeded, sy nzoochorically dispersed trees and
lianas that are dem ographically confined to the terra
firm e forests east of the Rio Madeira (Van Roos-
malen, 2013b). These woody plants seem to have
co-evolved with scatter hoarding rodents belonging
to the genera DdSypWCta Illiger, 18 11 (agoutis) and
Myoprocta Thomas, 1903 (ac ouch is), among which
we identified some possibly new species;
- Primate diversity, here defined as the total
number of taxa that occur in sympatry within a
10x10 km quadrant of land overlying both banks of
a river at certain latitudes, is the highest for the Rio
Madeira at the longitude of the mouth of the Rio
Aripuana, reaching at least 25 (!) valid species. That
Hotspot of new megafauna found in the Central Amazon (Brazil): the lower Rio Aripuana Basin
221
Figure 1. Study area. Central Amazon (Brazil): the Rio
Aripuana Basin (shaded area).
exceeds with at least two species the hitherto
highest primate diversity (in total 23 valid species)
found west of the Madeira River, along the Rio
Purus at its confluence with the Rio Tapaua (Van
Roosmalen, 2013a; 2015; Van Roosmalen & Van
Roosmalen, 2014).
- The Rio Aripuana is a clear-water river drain-
ing the area north of the Chapada dos Parecis, a
mountain range that is part of the crystalline
Pre-Cambrian Brazilian Shield. Together with the
clear- water Rios Tap ajo s - Ju r u en a , Teles-Pires,
and Xingu, the Rio Aripuana seems to harbor
relicts of a highly species-rich endemic Miocene
freshwater mollusk (shellfish or bivalve) fauna
with extant shells, oysters and mussels only to
be found east of the Madeira River (Hoorn &
W esselingh , 20 10).
RESULTS
New mammalian species descriptions from
the Rio Aripuana Basin, Brazilian Amazon
1. New species of living brocket deer (Mam-
malia Cervidae) from the Rio Aripuana Basin
Up to recently, only two members of the Neo-
tropical Odocoileinae (brocket deer), a subfamily
of the Cervidae (deer), from lowland Amazonia
were known to science, belonging to the extant
genus Mazama (Wilson & Reeder, 1 993): the red
brocket M. americana (Erxleben, 1777), and the
grey brocket M. nemorivClgCl (F. Cuvier, 1817). The
latter has being recently (Rossi, 2000) distinguished
from M. gouazoupira (G. Fischer, 18 14), which
species is said to range south of Amazonia on the
open savannas and shrub savannas (cerrado) of
Central Brazil, Bolivia, Paraguay, N Argentina and
Uruguay. Although the evolutionary history of
brocket deer dates back almost 20 million years ago
(M YA), Duarte etal. (2008) suggest that in the Fate
Pliocene, approximately 2.5-3 MYA, the uplift of
the Panamanian land bridge allowed deer to spread
south, as participants in the “great American inter-
change” between N orth and S outh A m erica. A ccord-
ing to Duarte et al. (2008), these were the first deer
to enter the S o u th - A m eric an continent, and their
surprising success in South America may be attrib-
uted to the absence of other ruminants (Webb,
2000).
Class Mammalia
OrderArtiodactyla or Cetartiodactyla (if w hales are
to be included)
Family Cervidae Goldfuss, 1820
Subfamily Odocoileinae Pocock, 1923
Genus Mazamci Rafinesque, 1817
Mazama tienhoveni V an Roosmalen et Van Hooft
Examined material. Two skins in possession of
hunters from the village of Tucunare along the
lower Rio Aripuana were examined. Moreover, a
complete skull and mandible still in the flesh from
an adult female specimen, and one spike from an
adult male specimen were obtained from them in
the course of the year 2006. The settlement of
Tucunare is situated along the Parana do Santa
Maria, a shortcut from the community of Santa
Maria to that of Tucunare, along the left bank of
the middle Rio Aripuana, State of Amazonas,
Brazil (05°45'S, 60°15'W). Holotypus: Specimen
MR204, complete head with partly damaged
mandible (Fig. 3), adult female, on May 1 2, 2006
killed for food by a local hunter along the left bank
of the Rio Aripuana near the settlement of
Tucunare, skull, spike (Fig. 4) and skin (Fig. 5). The
type specimen MR204 is deposited as INPA4273,
Mammal Collection of the National Institute for
Amazon Research, Manaus, Amazonas, Brazil.
222
Marc G.M. van Roosmalen
Figures 2-9 . MUZCLTHCL tieflhovevii n . sp . Figure 2 . M. tieflhoveTli n. sp. drawing reconstructed from plate depicting M. TieTTlOVi-
VClgCl (Eisenberg, 1989). Fig. 3. Skinned head of a holotype female fair brocket deer M. tiCTlhoVCTli n . sp. Fig. 4. Two spikes
of M. nemorivaga and one (the smallest) of M. tienhoveni n. sp. Fig. 5. Skin of M. tienhoveni n. sp. from Tucunare village,
Rio Aripuana. Figs. 6-8. skull and mandible of gray brocket deer M. neniorivOgCL (MPEG 1969). Fig. 9. Distribution map for
M. tienhoveni n. sp.
Description of holotypus. Measurements.
Two skins obtained from hunters along the lower
Rio Aripuana were measured. Body weight not
taken but according to local hunters ranges from 20-
25 kg. Skull length 185 mm, mandible length 145
mm. Diastema length in skull 53 mm. Condylobasal
length 167 mm. Palatal length 114 mm. Length of
nasals 55 mm. Interorbital constriction 41 mm. Zy-
gomatic breadth (= breadth across zygomatic ar-
ches) 80 mm. Breadth ofbraincase 55 mm. Length
of upper tooth-row 53 mm. Length oflower tooth-
row 58 mm. Breadth of M 2 12 mm, breadth of M2
8 mm. Dental formula: I 0/3, C ( 1 )/ 1 , P3/3, M 3/3.
Length of spikes (including the coronet) 55 mm.
Variability. No paratypes have been collected
thus far.
Etymology. We would like to name the species
for Dutch lawyer and naturalist Pieter Gerbrand
van Tienhoven (1875-1953), co-founder of a main-
stream conservation organisation in the Netherlands
(N atu u rm o n u m en te n ) and one of the founding
fathers of the International Union for the Conser-
vation of Nature and Natural Resources (IUCN):
Van Tienhoven's fair brocket deer, M. tienhoveni
n . sp .
Hotspot of new megafauna found in the Central Amazon (Brazil): the lower Rio Aripuana Basin
223
Van Tienhoven's fair brocket deer M. tiCYlhoVCVli
n. sp. is locally known as “veado branco”, which
means “white brocket deer”. This way locals distin-
guish it from M. americana commonly known as
“veado vermelho” or “veado capoeira”, which
means “red brocket” or “secondary-growth
brocket”, referring to its overall orange-red color
and preference for edge habitats and forest clear-
ings, and from M. nemorivaga locally known as
“veado roxo”, which means “purplish-grey brocket”.
Distribution. The geographical distribution of
Van Tienhoven's fair brocket is thought to be
restricted to the lower and middle part of the Rio
Aripuana Basin, but it might well be distributed
across the entire interfluvium delineated by the Rio
Madeira in the west, the Rio Tapajos-Juruena in the
east, the Rio Amazonas in the north and the Rio
Guapore in the south. Since it seems to be confined
to terra firme rainforest habitat, we assume that
its real distribution is much smaller and does not
extend into the northern part of the Rios
M adeira/Tap ajo s interfluvium, where many open
savannas and extensive floodplains are found. We
have observed the species in the wild only along
both banks of the Rio Aripuana.
Ecology. Van Tienhoven's fair brocket, M. tien-
hoveni n. sp., seems to be restricted to dense terra
firme (upland) rain forest, where it lives solitary or
in pairs. It occupies rather small territories and
occurs in the Rio Aripuana Basin in sympatry with
the locally much rarer grey brocket .M. nemorivaga,
and the greater red brocket M. americana. The lat-
ter, however, occurs more frequently in disturbed
areas with secondary growth and edge habitats, and
in open areas such as white-sand savannas, which
are common in the region. Nothing is known about
Van Tienhoven’s fair brocket, its ecology, and habits
in the wild. The author has seen it only a few times
in the wild during the dry season, while it was visit-
ing the BaCtris maraja (palm) dominated margins
of muddy ponds, mud pools and saltlicks. These can
be found locally in the middle of the rain forest at
sometimes long distances from any substantial
water supply, such as rivers, streams, lakes and
ponds.
Phylogeny. DNA was extracted from a skin
sample from each of the two brocket species, M.
tienhovenin. sp. and M. nemorivaga, both collected
from the forests along the left bank of the Rio
Aripuana. Partial mitochondrial cytochrome b DNA
sequences of 233 bp (sites 1 33-365) and 295 bp
(sites 108-402) in length were obtained for respect-
ively Mazama tienhovenin. sp. and M. nemorivaga
with the conserved primers L 1484 1 and H15149
(Kocher et al., 1989). DNA extractions, PCR reac-
tions, and DNA sequencing were performed accord-
ing to standard laboratory protocols. The sequences
are deposited in Genbank under the accession
numbers: GQ 268320 {M. tienhoveni n. sp.) and
GQ268321 (M. nemorivaga) . Unfort unate ly, we did
not have a skin sample from a specimen of the third
sympatric brocket M. americana. However, differ-
ent cytochrome b DNA sequences from this species
and all other currently known Amazonian deer
species could be obtained from Genbank (Genbank
accession numbers given in Fig. 10). Most of these
sequences have been used in a recent phylogenetic
study on the South American deer (Duarte et al.,
2008). We generated a m inim um -evolution (ME)
distance tree by adding our two sequences to those
used in Duarte et al., 2008. Furthermore, we in-
cluded Genbank sequences not used in that study,
belonging to various S outh-A m erican deer species,
and excluded those with a large number of missing
data in the 1 33-365 bp region of cytochrome b. The
ME- tree was constructed with MEGA 4 (Tamura et
al., 2007). We used the substitution model K2P
(Kimura, 1 980) with a constant rate applied and
w ith Rangifer tarandus (Linnaeus, 1758)being out-
group, as has also been done in Duarte et al., 2008.
The tree is based on the 1 33-365 bp region of cyto-
chrome b with unresolved nucleotides deleted by
pairwise deletion. Divergence times were estimated
assuming separation between BlastOCemS/ Pudli and
Mazama/ Odocoileus 5 MYA (Duarte et al., 2008).
Mazama tienhoveni n. sp. and Genbank se-
quence AY 886753, not being used in Duarte et al.
(2008) although being from a brocket classified as
M. gOliazOUpira, formed a distinct clade that diver-
ged from the other South American deer species
(average sequence divergence: 8.3%) 5 MYA (Fig.
10). This w o u Id im p ly th at M. tienhoveni n . sp . di-
verged already before the uplift of the Panamanian
land bridge and invaded South America during the
“great A m erican interchange” between both contin-
ents. A distinct clade not only supports the separate
species status of M. tienhovenin. sp., it also indic-
ates that Genbank sequence AY 8 86753 was
wrongly identified as M. gOliazOUpira. The latter
224
Marc G.M. van Roosmalen
observation is not unlikely, as low levels of mor-
phological differentiation in the genus MdZCltflCl
have caused numerous errors in species identifica-
tion in the past (Duarte et al., 2008). Genbank
sequence AY 886753 should either be attributed to
M. tienhovenin. sp. or to a separate species in its
own right, which is very well possible considering
the fact that it diverged 2-3 M YA from the M. tien-
hoveni n. sp. sequence. This divergence seems to
have occurred, more or less coinciding with the
uplift of the Panamanian land bridge.
Remarks. Mazama tienhoveni n. sp. differs
from the two other known Amazonian species, the
ion
GQ
ms
IE
IPV
C
H r
33
90
1Q3
SB
^1
Hr
93
ID
ffl 1
51
£
92 l
96 r
BI L
91
S3
“C
El
□E
99
Mazama gouazoupira DQ7S9181
Mazama gouazoupira DQ789203
Mazama gouazoupira DQ789189
Manama gouazoupira DQ789229
Mazama gouazoupira DQ789188
Mazama gouazoupira DQ789183
Mazama gouazoupira DO7890Q2
Mazama gouazoupira DQ789179
Mazama gouazoupira DQ786GQ0
Mazama gouazoupira DQ789186
Mazama gouazoupira DQ789194
Blastoceros dichotomic AY326234
Blastoceros dichotomus DQ789176
Blastoceros dichotomus DQ789175
Hippocamelus beulcus DQ739173
Hippocamelus be ulcus DG789177
Mazama gouazoupira DQ379303
Mazama nemorrvaga DG789226
Mazama nemorivaga 0Q789205
# Mizamanemorwagafrioaripuana)
Mazama nemorivaga DQ789213
Mazama nemorivaga D Q789206
Pudu pud a DG3793Q9
Pudu pud a L.43435
Hippocamelus antisense DQ379307
Qzotoceros bezoartious L43434
Ozotocercs bezoartious DQ789198
□zotoceros bezoartious DQ789199
Ozotoceros bezoartious D Q7S9193
Ozcdoceros bezoartious DQ789196
Odocoileus hemionus FJ188800
Odocoileus hemionus FJ 188781
Odocoileus hemionus FJ 188768
Odocoileus hemionus FJ 188874
Odocoileus hemionus FJ188805
Odocoileus hemionus FJ188764
Mazama bororo DQ789231
Mazama nana DQ 73C214
Mazama nana DQ789227
Mazama nana DQ789210
Mazama sp. AJ 1X0027
Mazama sp, DQ789180
Mazama americana DG789217
Mazama americana DQ780218
Mazama americana DQ789209
Mazama americana DQ 789211
Mazama americana DQ 789219
Mazama americana DQ7BS220
Mazama americana DQ 789223
MEama americana DQ789221
Mazama americana DQ789230
Mazama americana DQ789215
Mazama americana DQ 78022*4
Mazama americana DQ 78 9208
Mazama gouazoupira AY886753
0 Mazama tienhoveni
Rangifer tarandus NC007703
Rangifer tarandus AJ0CCO2 9
Rangifer tarandus AYD90 106
Rangifer tarandus AY090107
Rangifer tarandus L49403
Rangifer tarandus AF 4941 96
Rangifer tarandus AF 4941 95
7 6 5 4 3 2 1 0
005 J 005 0O4 ' 003 002 001 OQO
Fig are 1 0 . L inearized minimum-evolution tree showing phylogenetic relationships among South American deer derived fro m
a 2 3 3 bp fragm entofthem ito chondrial cy to chrome b. The scale on top correspo nd s to the time scale in millions of years while
the scale below corresponds to the observed mean sequence divergence using the substitution model K2P. Bootstrap values
(1000 re plica tes, > 50%) are denoted above nodes. Numbers behind taxon names c orresp o nd to Genbank accession numbers.
Hotspot of new megafauna found in the Central Amazon (Brazil): the lower Rio Aripuana Basin
225
Greater Red Brocket M. americana and the Gray
Brocket M. nemorivaga, in being intermediary in
size, but with 55 mm total length and coronet dia-
meter 24x30 mm having the shortest but most ro-
bust spikes (mean spike length 74 mm and coronet
diameter 21x22 mm in M. nemorivaga) . M ost of the
body is overall light brown colored, grading toward
almost white on the sides and ventrally, whereas the
dorsal parts of M. Cline ricCMCl are of a (deep) reddish
brown color, grading ventrally into a more rusty
color, and those of M. nemorivaga are dull or pale
yellowish or grayish brown to chestnut brown, grad-
ing ventrally into yellowish or whitish (Husson,
1978). The males of M. tienhoveni n. sp. do not
have the distinct crest of hairs on the forehead as
M. nemorivaga has, neither do the males of
Mazama americana. Head-body length is not
known yet, but M. tieilhoveTli n. sp. is said to be in-
term e diary in size between M. nemorivClgCl, being
760-1015 mm (N = 6) (Rossi, 2000), with shoulder
height 480 mm (Duarte, 1 99 6), and M. americana
being 1120-1135 mm. The short tail is 75 mm long,
dorsally has the same color of the back but shows a
conspicuous white tuft at the end, being predomin-
antly white below, whereas the tail in M. americana
is 1 60-200 mm long, including the tuft, and 60-106
mm in M. nemorivaga. Furthermore, hind foot
length (w ith hoof) in M. americana is 313-318 mm
and ear length is 94-100 mm, and 82-93 mm in M.
nemorivaga (Husson, 1 978; Rossi, 2000). Weight
of adult specimens is reported less than 15 kg in M.
nemorivaga, 25-40 kg in M. americana, and about
20-25 kg in M. tienllOVeni n . sp., according to local
hunters. M. tienhoveni n . sp. can be distinguished
from other brocket deer by its intermediary-sized
head and various other intermediary skull charac-
ters (Table 1 ). O verall, the cranium of M. tienhoveni
seems more related to that of M. nemorivaga than
that of M. americana, but it differs clearly from Ma-
Zama nemorivaga in the following mean cranial
measurements: greatest skull length 185 versus 174
mm; palatal length 114 vs. 105 mm; length of nasals
55 vs. 50 mm; interorbital constriction 41 vs. 39
mm; zygomatic breadth 80 vs. 73 mm; braincase
breadth 55 vs. 53 mm; alveolar breadth of the upper
second molar 12 vs. 11.3 mm; alveolar breadth of
the lower second molar 8 vs. 7.4 mm; and length of
mandible 145 vs. 134 mm.
The divergence time between M. tienhoveni n.
sp. and the two other brocket deer, derived from
partial cytochrome b DNA sequences, is estimated
Skull
Greatest orcondylobusa! length (-length
anterior lip of I 1 to rear ofcondyles)
W,
iimt’rirtiiiii
(n-l 1)
221
M,
nemorivaga
<n=S)
174
M.
tienlwvetil
(n=l)
1K5
Hiisal length (= length anterior lip of l‘
to proximal end afetmdylcs)
210
162
Ib7
Palatal length
127
102
114
Length of nasals
bS
SO
55
Inte [orbital eons met mil
47
39
41
Zygomatic breadth (breadth across
zygomatic arches)
97
73
HO
Breadth oTbra incase
b3
S3
55
Length of diastema
70
52
53
Alveolar length of upper moth -tow
b2
52
53
Alveolar breadth of M 1
14,6
11.3
12
Length of mandible (^length from 1' to
rear of processus condylieus)
172
134
145
Alveolar length of lower tooth •row
72
59
5K
Alveolar breadth of M2
9,7
7,4
S
Length of spikes
Hit (n=5)
74 (n=2)
55
D isla nee between spikes
3b (n=5)
35 (n=2)
Diameter of coronet
21-22 RA
(n=2)
24-30 RA
Table 1. Skull measurements (in mm) of Mazaina ameri-
cana (N = ii; nhml), M. nemorivaga (N = 5; nhml), and
M. tienhoveni ; M. tienhoveni n . sp. is represented by the ho-
lotype - an adult female from Tucunare, Rio Aripuana, State
of A m azonas.
at 5 million years before present, which is well
before the uplift of the Panamanian land bridge. As
in other brocket deer, Van Tienhoven's fair brocket
seems to live solitary or in pairs. In view of recent
developments in the Rio Aripuana Basin where it
lives and due to its limited distribution, we consider
Van Tienhoven's fair brocket highly endangered.
Conservation status. All three brocket species,
occurring in sympatry in the Rio Aripuana Basin,
are favorite game to the locals. Hunting the ‘blond’
or fair brocket M. tienhoveni n. sp. is said to be
more successful. Along the lower Rio Aripuana, it
is said to be the most commonly encountered type
of brocket, at least in terra firm e (upland) rain
forest, whereas greater red brocket are more often
226
Marc G.M. van Roosmalen
found near forest edges and clearings, such as fields
and plantations. A lthough human occupation in this
part of the Amazon is very low nowadays, this
situation might soon change. In the Rio Aripuana
region unprecedented illegal extraction of timber,
gold and gravel is taking place, ironically after the
whole lower Rio Aripuana region was declared a
State of Amazonas Sustainable Development
Reserve (Reserva de D esen volvim ento Sustentavel
- RDS do Baixo Rio Aripuana). Recent road build-
ing through the area has as objective to connect the
town of M anicore on the right bank of the Rio
M adeira with the now booming town ofApui at the
border of the Tenharim Savanna and the State of
Mato Grosso, areas of large-scale industrialized
soybean agriculture. As recently as the year 2006,
gold was found where the road from Novo
Aripuana crosses the Rio Juma, a clear-water tribu-
tary of the right-bank Rio Aripuana. A crowd of
over 10,000 gold diggers then settled in. Locals told
us that ever since commercial hunters flocked into
the area.
They use trained dogs for the hunt on game
species as giant peccary PeCCUl maximUS Van Roos-
malen et al., 2007, Van Tienhoven's fair brocket M.
tienhoveni n. sp., and both dwarf tapir TcipiruS
pygmueUS Van Roosmalen, 2013 (also known as T.
kaboTnani Cozzuol et al., 2013 - ajunior name) and
lowland Brazilian tapir TcipiriiS tewestvis (Linnaeus,
1758) to feed hungry settlers and gold-miners.
Taking increasing hunting pressure and the species's
limited distribution into account, M. tieYlhoV£Yli n.
sp. is considered highly endangered. It is recom-
mended to include this new species in the IUCN
Global Red List, based on criterion D (very small
or restricted population). Besides Van Tienhoven's
fair brocket, the Rio Aripuana region seems to
harbor a number of other faunal elements new, or
possibly new, to science. Identified so far are a new
species of peccary Pecciri JfldXilflUS (Van Roos-
malen et al., 2007), a new species of dwarf porcu-
pine Coendu ( Sphig gurus) roosmalenorum (Voss &
Da Silva, 200 1 ), and at least four new primate
species (Van Roosmalen et al., 1 998; Van Roos-
malen et al., 2000; Van Roosmalen et al., 2002; Van
Roosmalen and Van Roosmalen, 2003). Among
these primates, the black-crowned dwarf marmoset
Callibella humilis Van Roosmalen et al., 199 8
represents a complete new genus first seen and
collected by me in 1 996. Most surprisingly, up to
today not a single area exists in the region that is
effectively protected by Brazilian environmental
law. Given the uniqueness of the region in terms of
biodiversity and its current status of biological terra
incognita, we here suggest UNESCO to urge
the Brazilian Government to declare the entire
lower Aripuana Basin a Natural World Heritage
R eserve.
2. New species of living rodent from the Rio
Aripuana Basin: the giant striped paca
( Mammalia Agoutidae)
Up to recently only two members of the Neo-
tropical family Agoutidae (pacas), synonymous to
Cuniculidae, were known to occur in the Americas
belonging to the extant genus AgOUti (Wilson &
Reeder, 1993): the spotted common paca AgOUti
paCd (Linnaeus, 1 766), which species ranges in
C + S America from San Luis Potosi, SE Mexico, to
Paraguay, the Guianas, and S Brazil (the species
was introduced into Cuba) and occupies suitable
lowland habitats, and the mountain paca A.
taczanowskii (Stolzmann, 1865), a species from
the high cloud forest (altitudes between 2,000-3,000
m) of Andean regions of Peru, Ecuador, Colombia,
and NW Venezuela (Eisenberg, 1 989; Eisenberg
& Redford, 1999).
Class Mammalia Order Roden ti a
Family Agoutidae Gray, 1821
Subfamily Agoutinae or Cuniculinae
Genus AgOUti Lacepede, 1799
Agouti silv agar ciae \ an Roosmalen et Van Hooft
Examined material. Holotypus: adult female,
complete head (Fig. 13), killed for food by a local
hunter on May 28, 2006, along the left bank of the
Rio Aripuana near the settlement of Tucunare.
Paratypus: stuffed specimen found under number
MPEG 22302 in the mammal collection of the
Museu Paraense Emilio Goeldi, Belem, Para,
Brazil, its provenance not given (Fig. 14). Head-
body length 750 mm. The heads were preserved on
spirit (Fig. 13). Grain shot had severely damaged
the skull of the giant paca specimen making it
impossible to take cranial measurements.
Hotspot of new megafauna found in the Central Amazon (Brazil): the lower RioAripuana Basin
227
Figures 11-16. Fig. 11. Plate from Eisenberg (1989) depicting the common spotted paca AgOUti pCLCCL', Fig. 12. The common
spotted paca A. pCICCI as depicted in Emmons & Feer(1990).Fig. 13. Heads offreshly killed common spotted paca A. pOCCl
(at bottom) and Silva Garcia’s striped giant paca A. silvCLgClTCicie n . sp. (at top). The skulls and mandibles of these specimens
are stored at Tucunare village, Rio Aripuana, State of Amazonas, Brazilian Amazon. Figs. 14, 15. Stuffed specimen of
striped giant paca A. silvCLgarcicie n . sp. found by the first author in the collection ofMuseu Paraense Emilio Goeldi under
MPEG 22302 - with out locality and misidentified as A. pCLCCL. Fig. 16. Skull of the common spotted paca A. pCLCCL fo und by
the firstauthor in the collection ofMuseu Goeldi under MPEG 5418, from the locality ofTapirinha. Skull length 147 mm,
skull width 97 mm.
Description of holotypus. The general dorsal
pelage color of the common spotted paca AgOUti
pCICCI is uniformly chestnut or mummy brown to
almost black, usually with a striking pattern of four
horizontal lines of white or light yellowish dots on
each side of the body, the two middle ones at least
extending all the way from the neck to the rump. In
these two middle rows, the spots in the middle part
may be fused to an uninterrupted stripe. The lower
of the rows of spots is only visible in the extreme
anterior and posterior parts, as the middle part is
fused with the white ventral surface of the body.
One or two of the upper rows of spots are shorter
than the other rows; they are visible only in the
posterior half of the body. The hairs are stiff and
shiny. The dark brownish hairs show a lighter
median line. The dorsal surface of the head is of the
same color as the back, but the hairs are shorter and
less stiff. On the snout, there are long stiff whiskers.
The upper whiskers are blackish and the lower are
white, the color difference being quite striking.
Similar stiff whisker-like hairs, though fewer, are
implanted below and slightly in front of the ears;
here too, upper hairs are blackish brown, lower ones
white. The ears are relatively large; a tuft of
blackish and yellowish longer hairs is implanted
before the opening of the ear. The throat and the
cheeks are uniformly cream -colored, as is the entire
ventral surface of the body. The line of demarcation
between the dark dorsal and the whitish ventral
color is distinctly marked. The outside of the legs
is of the same brown color as the dorsal surface of
228
Marc G.M. van Roosmalen
the body; the inside of the legs is yellowish white
basally, brown or brownish distally. The tail is ve-
stigial, very short, and hardly noticeable. There
are four toes with nails both on the fore- and hind-
feet. In the forefeet, the nail of the thumb is very
small, the others are well developed and of equal
size. In the hind-feet, the three middle nails are
large and of about the same size, while the nails of
the inner and outer toes are markedly smaller and
implanted higher, the inner nail being again some-
what smaller than the outer one. In the female there
is on each side one pectoral mamma, at about the
level of the bases of the front legs. Dental formula:
1 1/1, CO/1, P 1/1, M 3/3. The skull of an adult com-
mon paca is immediately characterized by the zy-
gomatic arch that has grown out to an enormously
swollen, distinctively sculpted bony plate, which is
about two-third the length of the palate; this plate
is strongly produced downward, in lateral view ob-
scuring the teeth and the basal part of the mandible.
Anteriorly, this plate encloses a deep and very large
cavity at each side of the very narrow palate in front
of the tooth-rows. In comparison to other rodents,
the teeth are placed far backward. The palate ends
at the line between the last and the penultimate
molars. The infra-orbital opening has become a
narrow canal aim ost entirely enclosed by bone. The
outer surface of the zygomatic arch is covered with
a honey-comb of irregular bony ridges, giving it a
strongly rugose appearance. These rugosities extend
also onto the larger parts of the nasals, frontals and
parietals; in full-grown specimens the sutures
between these bones are not or only partly visible.
Even in newborn and juvenile specimens, the zy-
gomatic arch is relatively high, but still smooth. The
external measurements of three adult specimens of
AgOUti paca from Suriname on which the above
mentioned description is based, are: head-body
length 650; 676; 662 mm; tail length 18; 17; 19
mm; hind foot (including nail) 119; 117; 113 mm;
ear length 46; 41; 48 mm; weight 9.2; 9.1; and 9.5
kg (Husson, 1 978). Eisenberg (1 989) gives head-
body length averages 600-795 mm, vestigial tail
length 19 mm, hind foot 188 mm, ear length 45 mm
and weight 7. 5 kg. He also states that the adult male
is about 15% larger than the adult female. Silva
Garcia’s giant paca A. silvagarciae n . sp. from the
Rio Aripuana Basin has been reported by locals to
weigh between 12-15 kg, its general color is bright
orange brown. Average weight of the sympatric
common paca A. paca in the Rio Aripuana Basin is
reported to be 5-6 kg. The head of the giant paca
shot near Tucunare settlement measures 155 x 80
mm (compared to 115 x 80 mm in the common paca
specimen shot the same night at the same locality
of Tucunare) (Fig. 13). The eight upper whiskers
are black and about 110 mm long, the eight lower
whiskers are white, stronger and stiffer than the
upper ones and 105-110 mm in length (similar to
those in the common paca). Three out of four lateral
rows show the white spots (almost) completely
fused (Figs. 14, 15).
Measurements. Body weightwas not taken from
the holotype specimen of A. silvagarciae n . sp ., but
according to local hunters body weight ranges from
12-15 kg. The head of the giant paca shot near
Tucunare settlement measured 1 55 x 80 mm,
whereas that of the adult common paca shot at the
same locality the same night measured only 115 x
80 mm (Fig. 13).
Variability. Paratype: stuffed specimen of
striped giant paca A. silvagarciac n. sp. found under
numberMPEG 22302 in the mammal collection of
the Museu Paraense Emilio Goeldi, Belem, Para,
Brazil, misidentified as A. paca, its provenance not
given (Figs. 14, 15). Head-body length 750 mm.
Etymology. This paca is named in honor of the
author’s spouse Antonia Vivian Silva Garcia.
During our visit to the community of Tucunare she
heard the villagers talk about the two types of paca
the locals there distinguish, one specimen of each a
local hunter had shot for food that very night.
Undoubtedly, the two species of paca must there-
fore be considered (m icro)-sym patric.
Vernacular name: A. silvagarciae n. sp. is
known locally as “paca concha”, which means
“shellfish paca”. This way, locals distinguish it from
the common paca A. paca that is known as “paca
pintada” (“spotted paca”).
Distribution. Members of the genus AgOUti are
distributed from southern Mexico to northern
Argentina in suitable lowland habitats. The geo-
graphical distribution of the giant paca is thought
to be restricted to the interfluvium confined by the
Rio Amazonas in the north, the Rio Madeira in the
west, the Rio Ji-Parana or Rio Guapore in the south,
and the Rio Tapajos-Juruena in the east (Fig. 1). We
have observed the species in the wild only along
both banks of the Rio Aripuana. Type locality of
Hotspot of new megafauna found in the Central Amazon (Brazil): the lower Rio Aripuana Basin
229
Agouti paca Rg n°
Sex
Greatest length mm
Condylobasal length
Basal length
Palatal length
Length of nasals
Zygomatic breadth
Hght zygomatic arch
Lgth zygomatic arch
Intero rb ital constr,
Braincase breadth
Mastoid breadth
Bullae lgth x bdth
Height of rostrum
Diastema
Alveolar length P-M 3
Breadth of m 2
Length of mandible
Alveolar length P-M 3
Breadth of M3
23902
18233
17756
-
7
-
139
143
143
129
138
144
123
131
136
77
$2
86
45
fused
56
92
96
104
46
50
60
80
87
102
45
42
44
45
43
47
61
57
65
20x15
18x14
18x17
39
40
42
44
48
51
28
30
29
7.1
7.2
7.9
97
87
103
30
31
32
7.5
6.8
8.5
21889
21891
21892
142
148
148
136
145
139
128
138
133
82
86
85
44
50
47
87
100
95
39
56
47
80
98
86
40
45
45
45
49
46
60
65
62
21x17
17x14
20x18
39
41
42
46
51
50
28
32
28
8.0
7.4
7.3
18013
7
18017
7
M
136
141
143
128
137
137
121
129
130
75
82
82
46
50
48
-
92
95
37
47
48
72
82
86
37
40
42
44
43
45
59
55
61
20x14
19x17
19x16
37
40
40
42
46
47
29
30
29
7.0
7.6
7.4
93
96
95
31
33
31
7.2
8.0
7.6
Table 2. Cranial measurements (in mm) of eight specimens of the common spotted paca AgOUti paca from Suriname (zoolo-
gical collect ion of the NHML, Leiden, the Netherlands) on which our description is based. With “height of zygoma tic arc h’ -
is meant the greatest height; with “breadth of the braincase’" is meant the width of the skull at the leveljust above the external
auditory meatus. Also, the total length of the zygomatic arch is noted being the distance between the extreme anterior and
posterior borders. The length of the mandible was measured from the processus angularis. Mean skull length 143 mm, mean
mandible length 95 mm. Diastema length in skull 47mm. Condylobasal length 137 mm. Palatal length 82 mm. Length of
nasals 48 mm. Interorbital constriction 42 mm. Zygomatic breadth (= breadth across zygomatic arches) 95 mm. Breadth of
bra incase 45 mm. Bread th of M 2 7.4 mm, breadth of M ’ 7.6 mm. Dental formula: I 1/1, C 0/0, Pl/1, M 3/3.
A. silvagarciae n . sp. is the Rio Aripuana, close to
the settlement of Tucunare, situated along the
Parana do Santa Maria, a shortcut from the com-
munity of Santa Maria to that of Tucunare, sitting
on the left bank of the middle Rio Aripuana, State
of Amazonas, Brazil (05°4 5'S,60°15'W).
Ecology. Pacas of both species are nocturnal
and have their hiding-places in hollow fallen tree
trunks. They always carefully make two entrances
to their burrows, so that they can escape when
hunted down by dogs. Pacas usually live close to
rivers and creeks. When pursued by dogs, they
frequently take refuge in the water. Notw it h standing
its fat body, it manages to walk on the bottom of
any substantial water body. The formidable teeth
and enormous masticatory muscles of pacas enable
them to break open the hardest fruits and seed
kernels. In contrast to the agouti, the paca digs
burrows that are sometimes interconnected with
others.
The giant paca A. silvagCircicie n. sp. is assumed
to be restricted to dense terra firme upland rain
forest, where it lives solitary or in pairs. It occupies
rather small territories and occurs in the Rio
Aripuana Basin in sympatry with the locally much
more common spotted paca A. pCICCl. The latter,
however, is more frequently found along edges,
such as roadsides, streams and creeks, in disturbed
areas with secondary growth, and in open areas on
white-sand savannas common in the region.
Nothing is known about its ecology and habits in
the wild. I myself have seen it in the wild only a
few times during the dry season while visiting the
Bactris vnarcijci M art (palm) d o m in ated m arg in s o f
muddy ponds, mud pools and saltlicks. These can
be found locally in the middle of the rain forest,
230
Marc G.M. van Roosmalen
often at long distances from any substantial water
body, such as rivers, streams, lakes or ponds.
Phylogeny. One complete mitochondrial D-
loop and two nuclear SINE PRE-1 DNA sequences
of Silva Garcia’s giant paca were carried out and
compared with Genbank sequences of the sympatric
common paca (A. paCO.) . The results (15.5% dif-
ference between species) clearly support the distinc-
tion into valid species. As genetic distances based
on partial mtDNA cytochrome b sequences (283
bp) in Bovidae are estimated 1.25% = 1 MYS, di-
vergence tim e betw een A. pciCCl and A. silvagarciae
n. sp. is estimated at about 10 million years. The
giant paca therefore seems to have derived from an-
cestral pacas in the Late Miocene to Early Pliocene.
Remarks. This second species of paca from the
Brazilian Amazon is distinctly bigger than the
morphologically most related species that occurs in
the Amazon, the common spotted paca A. pacci.
One complete mitochondrial D-loop and two
nuclear SINE PRE-1 DNA sequences of the giant
paca compared with that of the sympatric common
paca {A. paca) supports the distinction. Divergence
time is estimated at 10 million years. As in the com-
mon paca, giant pacas are nocturnal and reported to
live solitary or in pairs. In view of recent develop-
ments in the interfluves where it lives, due to its
limited distribution and for being a prime target to
local hunters, we consider Silva Garcia’s giant paca
on the verge of extinction.
Several specimens of stuffed pacas in the zo-
ological collection of Museu Paraense Emilio
Goeldi, Belem, Para, identified as common pacas,
are suspected to represent giant pacas, hereafter
named A. silvagarciae n. sp. (Figs. 14, 15). This
assumption is based on the orange-brown skin
color, the pattern of horizontal white stripes instead
of spots, total body length, weight, and cranial
measurements. Unfortunately, none of them has
been given a proper geographic locality.
Conservation status. The two paca species oc-
curring sy m p atric ally in the Rio Aripuana Basin are
favorite game to the locals. Although human occu-
pation in this part of the Amazon is very low
nowadays, this situation may change in the near
future. In the Rio Aripuana region unprecedented
extraction of timber and gravel is taking place.
Recent road building through the area is intended
to connect the town of Manicore on the right bank
of the Rio Madeira with the boomtown of Apuf
located at the border of the Tenharim Savanna and
the State of Mato Grosso, areas oflarg e-scale indus-
trialized soybean agriculture. In view of these
developments, we fear that commercial hunters
using trained dogs will focus first on large animals,
such as the giant paca AgOUti silvagavtiaC n . sp ., to
feed hungry settlers and gold-diggers. Taking
increasing hunting pressure and the species's li-
mited distribution into account, I consider A. silvag-
arciae n. sp. on the verge of extinction. It is
recommended to include this new species in the
IUCN Global Red List, based on criterion D (very
small or restricted population). Besides the giant
paca, the Rio Aripuana region is thought to harbor
a number of other mega-faunal elements new to
science. I have identified so far a new species of
peccary Pecari lTiaxilTIUS (Van Roosmalen et al.,
2007), a new species of dwarf porcupine Coendu
( Sphiggurus ) roosmalenorum (Voss & Da Silva,
2001 ) - and seven new primate species, four of
which are already officially described [Van Roos-
malen et al. (1 998); Van Roosmalen et al. (2000);
Van Roosmalen et al. (2002); Van Roosmalen &
Van Roosmalen (2003)]. Among these primates, the
dwarf marmoset Callibclla hlATTlilis represents a new
primate genus first seen and collected by me in
1 996. Most surprisingly, not a single area protected
by Brazilian environmental law exists in the region.
Given the uniqueness of the region in terms of biod-
iversity and its current status of biological terra
incognita, we here suggest UNESCO to urge the
Brazilian Government to declare the entire region
a Natural World Heritage Reserve.
3. New species of living manatee ( Mammalia
Trichechidae) from the Rio Aripuana Basin
- shallow clear -water adapted dwarf manatee
is on the verge of extinction
Manatees (Mammalia Trichechidae) are fully
aquatic mammals of the ancient Order Sirenia.
Worldwide there are two extant genera, TrichechuS
Linnaeus, 1758 and DugOVlg Lacepede, 1 799. The
Amazonian manatee T. inunguis (N atterer in
Pelzeln, 1 8 83 ) is the only species strictly adapted
to fresh-water environments. However, here we
announce the discovery of a second taxon from the
Amazon that is also adapted to fresh-water habitat.
Hotspot of new megafauna found in the Central Amazon (Brazil): the lower Rio Aripuana Basin
23 1
Class M am m alia
O rder Sirenia
Family Trichechidae
Genus TvichechuS Linnaeus, 1758
Trichechus pygmaeus Van Roosmalen et Van der
V list
Examined material. Holotype: skull with lower
jaw of adult male is numbered CCM181, Zo-
ological Collection of the Brazilian Institute for
Amazon Research (INPA), M an a u s- A m azo n as ,
Brazil, collected eight km upstream from the m outh
of the Rio Arauazinho (Fig. 17), a left bank tributary
of the lower Rio Aripuana, State of Amazonas,
Brazil (06°16'94”S, 6 0 ° 2 0 ' 8 7 ” W ), 25. IX. 2002,
M.G.M. van Roosmalen legit.
Description of holotypus. Holotype skull
length 24 cm, greatest width 15 cm. Mandible
length 15.5 cm. Rostrum length 6.5 cm. Frontal
bones convex, greatest width 5.1 cm. Cheek teeth
(fully erupted molars) 4, maxillar molars 0.9 x 0.9
cm (Fig. 18), mandibular molars 0.8 x 0.6 cm.
Skull roof 5 cm wide, lacking parasagittal crests.
Braincase volume approximately 210 cc.
Variability. The species description is based on
two adult males and generalized in accordance with
reports from local hunters. Both adult male dwarf
manatees have been examined, each measuring 130
cm in length, 90 cm in circumference, and weighing
about 60 kg. The skin is overall pitch-black with a
circular to tear-shaped white patch on the abdomen
reported to be so in both the sexes, measuring in the
captive male ca. 52 cm long and 26 cm in diameter.
The flippers measured 32x11 cm and the paddle
36x40 cm. Snout comparatively short, circumf-
erence 46 cm, 19 cm in diameter, beset with long,
stiff bristle hairs. Whole body is thinly beset with
bristle-hairs (see also Fig. 19).
Etymology. Pygmaeus in Fatin means “short”,
“very small” or “dwarf”, reflecting as such the fact
that the new taxon of manatee is a dwarf compared
with the common Amazonian manatee T. iflUTlguis.
Vernacular name: “Dwarf manatee”, or “peixe-
boi anao”. It is locally known as “pretinho”, which
means “little black fellow”. This way, the locals
distinguish it from the common Amazonian fresh-
water manatee T. ifiungliis widely known as
Figure 17. Landsat images of the Rio Arauazinho, bran-
ching off in three directions, one lengthy branch coming
from the south running parallel with the Rio Aripuana, one
short branch coming straight from the north, and one main
branch coming from the northwest. The latter drains the ex-
tensive wetlands along the watershed with the upper Rio
Mariepaua, which harbor the last remaining population of
T- pygmaeus.
Figure 18. (Above) Maxillary molars of a young juvenile
male T. inunguis (Inpa Pb248) compared with those of the
holotype adult male T. pygmaeus n . sp. (below).
232
Marc G.M. van Roosmalen
“peixe-boi comum” (Portuguese for “common fish
cow”).
Distribution. Currently known distribution
restricted to the Rio Arauazinho Basin, a dear-
water tributary of the left-bank Rio Aripuana, State
of Amazonas, Brazil. Dwarf manatees may also
occur in the headwater region of the Rio Mariepaua,
the wetlands of which are interconnected with those
of the northernmost branch of the Rio Arauazinho.
Comparisons. The new taxon is assigned to the
genus TrichechllS, because it possesses a number of
traits in common with the parapatric, though in eco-
logical respect allopatric Amazonian manatee, T.
inunguis, from which it differs by its total adult
body length 130 cm (280-320 cm in T. inunguis),
and weight ca. 60 kg (350-500 kg in T. inunguis )
(Domning & Hayek, 1986). Growth curves of free-
ranging Amazonian manatees ( T. inunguis ) in B razil
are described by Vergara-Parente et al., 2010. Age
estimates and biometrics from 60 Amazonian
manatees captured between 1 993 and 2006 by local
residents of the mid-Solimoes and Pirativa Rivers
in the Brazilian Amazon are given as follows:
length at birth for T. inunguis is estimated at 13 3.2
cm (average = 113.0 cm; SD = 34.4 cm) for males,
and 13 1.0 cm (average = 124.7 cm; SD = 22.0 cm)
for females. A maximum length of 299.4 cm is
given in males, and 256.1 cm in females. Therefore,
both the adult male holotype T. pygmueUS n. sp. and
the adult male dwarf manatee that we kept alive for
over four months in an enclosure by fencing off a
bend in the Rio Arauazinho, had the same total
body length as just-born infants of the common
manatee T. inunguis. Moreover, the skin in T. inun-
gUlS is evenly dark grey colored, with individually
very variable irregular elongated white stripes
on the abdomen in females and only a few small
Figure 19. Adult dwarf manatee male kept for over four months in a fenced-off river bend of the Rio Arauazinho where it
was fed with its local natural food; note the saturated eumelanin black skin, relatively short head, short trunk and flippers, the
bristle hairs on the snout, and the large, tear-shaped albinotic white patch on the abdomen.
Hotspot of new megafauna found in the Central Amazon (Brazil): the lower RioAripuana Basin
233
irregular white blotches in males (Da Silva, pers.
comm.). Dwarf manatees, in contrast, are saturated
eumelanin coal black, the black pigmentation most
likely being an adaptation to its preferred habitat,
fast flowing shallow clear-water streams, protecting
them from skin burn by UV radiation (Fig. 19).
Common Amazonian manatees that are evolu-
tionarily adapted to murky silt-laden w hite-w ater or
low visibility dark-brown stained black-w ater, kept
in clear-water tanks at IN PA , Manaus, have to be
protected from severe skin burn by blocking off any
direct sunlight (Da Silva, pers. comm.). Male T.
pygmaeus n. sp. have a white tear- shaped, ca. 52 cm
long patch on the abdomen, greatest width 26 cm
(Fig. 19). Flippers of the captive male measuring
32 x 11 cm were too short to reach the mouth. In
contrast, the flippers of T. inunguis are proportion-
ately longer in the animals kept at IN PA . They are
used to push floating stems and foliage toward the
trunk and into the mouth. The snout of adult T.
pygmaeUS n. sp. is beset with long bristle hairs (Fig.
19), whereas that of infants T. UlUYlguis kept at
IN PA is smooth lacking bristle hairs. The large
white ventral patches reported in both sexes of
T pygmaeUS n. sp. perhaps have been selectively
evolved as protection against stingray attacks (an
irregular black-and-white belly pattern may deceive
its visual perception) during horizontal browsing of
pastures of Eleocharis minima K unth (C yperaceae)
and Thuvnia spp. (Thurniaceae). These aquatic
herbs grow on the sandy bottom of the mostly
shallow cle ar- w ater A rau az inho River. Those pas-
tures offer ideal hiding places for stingrays of all
sizes (Fig . 2 2).
The holotype skull of T. pygmaeus n. sp. is 24
cm long and 15 cm wide, the rostrum is 6.5 cm in
length and lacks the expanded nasal basin of adult
T. inunguis. The skull of T. inunguis measures 34 x
19 cm and the rostrum 11.5 cm (Figs. 20, 21).
Frontal bones in the holotype skull of an adult male
T. pygmaeus n. sp. are convex and 5.1 cm wide,
whereas in T. inunguis they are concave and only 4
cm wide (Fig. 21 - note that the skull of the juvenile
male T. inunguis , IN PA Pb248, is not damaged, but
falls apart along the fissures). The skull roof lacks
the parasagittal crests of T. inunguis, and the brain-
case volume in both taxa is about equal being ca.
210 cc. Therefore, it is much larger in T. pygmaeus
relative to its total body size. The total number of
cheek teeth (fully erupted molars) in each jaw
quadrant is 4 in the holotype T. pygmaeus n. sp.
(indicating a trend to neotony), but 6 ( - 8 ) in T.
inunguis (Domning & Hayek, 1986). Fu rth erm ore ,
they are much smaller-maxillary molars are 0.9 cm
in diameter in T. pygmaeus, and 1.3 cm in diameter
in T. inunguis (Fig. 2 1 ) . While the 3-4 anterior
molars in juvenile T. inunguis are hardly worn in
comparison to the just erupted posterior molar, this
is strikingly different in the holotype T. pygmaeus
with the 3 anterior molars strongly worn, thus
showing indisputably its adult status (Fig. 18). The
skull of the holotype male T. pygmaeus compared
with a similar-sized skull (23 x 15 cm) of a young
male T. inunguis (IN PA Pb24 8) reveals the
following major differences: 1/ the skull of T.
pygmaeus n. sp. is thick, robust and solid (the
cranial sutures, especially the basisphenoid-
basioccipital one, are fully fused), whereas the skull
bones of the young T. inunguis are thin and not
fused yet, so that its skull falls apart along the
fissures; 2/ the frontal bones in T. pygmaeus n. sp.
are convex and 5.1 cm wide, whereas in T. inunguis
they are concave and only 3.8 cm wide; and 3/ the
3-4 anterior molars in T. pygmaeus are fully worn,
whereas in young T. inunguis only three cheek teeth
are fully erupted .
The latter are sharply crested and do not show
any rate of abrasion (Figs. 18, 21). We have se-
quenced a fragment of 410 bp of the left domain of
the mitochondrial control region (D-loop), using
DNA extracted from a skin sample of a living
specimen. We used the same primers as were used
for T. manatUS and T. inunguis in Garcia-Rodriguez
et al. (1998). The resulting sequence was identical
to the most frequent T. inunguis h ap lo ty p e -h ap lo -
type T, frequency 31% (Garcia-Rodriguez et al.,
1 998). At first sight, this result seems to be dis-
crepant with the valid species status allocated by us
to the dwarf manatee. We first thought this result
could be explained by the relatively slow control
region mutation rate in manatees, being only
1.5%/1 million years (equivalent to 1 point muta-
tion/163,000 years) between lineages as compared
to 8-15%/lmillion years in most terrestrial
mammals (Garcia-Rodriguez et al., 1 998). This
would indicate a maximum divergence time of
485,000 years before present (p = 0 .0 5 ). W ithin such
a long space of time, sub-specific and even specific
dwarfism is possible. For example, episodes of
invasion and subsequent dwarfing affected many
234
Marc G.M. van Roosmalen
tnunguts
pygmaeus
pygmaeus
tnunguts
pygmae
Figure 20. Comparing skull and mandible of TrichechuS ifiunguis adult female Inpa Pbl97; T. pygmaeus adult male holotype
Inpa CCM181; and T. fflClUCltUS ( ill u stratio n taken from Husson, 1978). Note the convex and wider frontals in T. pygmaeus,
and the comparatively greater resemblance of its skull to that of T. lYlClYlCltUS.
Hotspot of new megafauna found in the Central Amazon (Brazil): the lower RioAripuana Basin
235
ad.
inunguis
Figure 21. Comparing skull, mandible and cheek teeth (fully erupted molars) of TrichechuS inungliis adult female Inpa Pbl97;
T. pygmaeus n. sp. adult male holotype Inpa CCM 181; and T. inungliis juvenile male Inpa Pb 24 8 . N o te the strikingly d iff e rent
wearing pattern when comparing the cheek teeth of the adult male T. pygmaeUS n. sp. and the juvenile T. inunguis.
tnvnguis
ad. ,
munguis
ad.
pygmaeus
r ad.
pygmaeus
juv.
inunguis
236
Marc G.M. van Roosmalen
insular fossil Elephantidae species from islands in
the Mediterranean and W allacea Seas. Some of
these have become dwarfs in less than 5,000 years
(Lister, 1 993; Lahr & Foley, 2004). Also, Wrangel
Island mammoths diminished by about 65% in
body size within at most 5,000 years after the
severing of the Late Pleistocene land bridge to
Eurasia (Lister, 1 993). Early Pleistocene large ele-
phants, with which the Sirenians are closely related,
swam and walked from the European mainland to
the island of Crete. There, they evolved into a 90
kg weighing dwarf species ElephciS CTCticUS Bate,
1907 (Caloi et al., 1 996). It could be hypothesized
that common Amazonian manatees once acciden-
tally got trapped in the Rio Arauazinho Basin isol-
ated from the main population in the Rio Aripuana
Basin. Forced to survive in (to the species inappro-
priate) clear-water habitat, these colonizers may
have drastically changed their diet and adopted a
different foraging technique. In addition, they might
have dwarfed under strong selective pressure of
limited food resources. Nowadays, hybridization
between the two extant taxa will not take place
easily, for dwarf manatee’s preferred habitat, diet
and foraging strategy is completely different from
that of the common Amazonian manatee. However,
a local from San Antonio village once reported
having seen twenty years ago a group of seven
dwarf manatees (“Pretinhos”) swimming along the
fishing nets he had put up along a sandy beach at
Prainha, right bank of Rio Aripuana, during the
peak of the dry season. If true, in theory female
dwarf manatees that accidentally drift into the
Aripuana River during the flood season and are not
able to return to the Rio Arauazinho (because its
mouth has fallen dry earlier than expected), may be
fertilized by male T. inilTlguis. The latter are said to
hibernate during the dry season in deep pools in the
main Aripuana River. Even if hybridization would
take place only once in the course of tens of thou-
sands of years, some gene flow between popula-
tions of both taxa would significantly obscure their
true divergence time. Accordingly, we believe that
the dwarf manatee should be placed at the base of
the phylogenetic tree of all fre sh w ate r- ad ap te d
manatees. In this quadrant of the Amazon Basin
south of the Rio Amazonas and east of the Rio
Madeira, it might have adapted to clear-water we-
tland habitat already in Late M iocene to Early Plio-
cene, times in which the landscape east of the
proto-Madeira was predominately drained by clear-
water rivers and streams - until the Late Pleistocene
vicariance that marked the birth of the modern fast-
flowing Rio Madeira.According to this geophysical
scenario, common Amazonian manatees may have
derived from archetypical ancestral dwarf manatees
during the Pliocene by adapting to black- and
white-water floodplain systems. This could have
happened after the Amazon reversed its course
about 8 M YA and began to drain the East-Andean
region into the Atlantic Ocean. At the same time,
this evolutionary scenario would explain for the
horizontal feeding posture of dwarf manatees in
which they stand on their flippers while browsing
on aquatic vegetation growing on the bottom of
fast-flowing clear-water streams. The horizontal
posture of dwarf manatees during feeding and
foraging resembles that of marine manatees from
which they could have derived during Late Miocene
to Early Pliocene, when the Andean uplift was
crucial for the evolution of Amazonian landscapes
and ecosystems reconfiguring drainage patterns,
creating a vast influx of sediments into the Amazon
Basin, and boosting its biodiversity (Hoorn &
W esselingh, 2010).
Ecology and conservation. Along the lower
and middle Rio Aripuana T. inUYlgllis is during the
flood season commonly found in the deep, slow-
moving, rather turbid dark waters of the Rio
Aripuana, its floodplain and deep back-water lakes.
The latter are filled with black-water coming from
local streams that drain nearby alluvial sand savan-
nas. T. pygmaeus n. sp., on the contrary, occurs ex-
clusively in the shallow, fast-flowing, clear waters
of the Rio Arauazinho Basin. During the rainy
season, the water level of the lower Arauazinho
rises over 7 m annually, and the dark rather turbid
waters from the Aripuana River then inundate the
riparian forest (igapo) fringing the lower Araua-
zinho. Dwarf manatee’s preferred food, most
importantly Eleochciris IflinilTlCl R. B r. grass (Cyper-
aceae), dies off for lack of sunlight. Its rhizomes
survive and hibernate in the dark during the flood
season. According to locals living in a small
community at the mouth of the Arauazinho, and
confirmed by us during surveys of the entire Rio
Arauazinho Basin in 2006 (on foot) and 2011 (by
dugout canoe), shortly before the Rio Aripuana
starts flooding the igapos, a number of dwarf
manatees migrate back to the headwater wetlands.
Hotspot of new megafauna found in the Central Amazon (Brazil): the lower RioAripuana Basin
237
Figure 22. The lower Rio Arauazinho showing the dwarf manatee’s pre ferred habitat: shallow fast-flowing transp a rent w aters
w ith Eleocharis minima, meadows growing on arenite white-sand subs tr ate. At places shown above, where the river becomes
shallow and flows faster, abundantly growing meadows ol E. minima attract dwarf manatees from as far upriver as the head-
w ater wetlands in the northwest (see Fig. 17). During the entire dry season (July to January), this amphibian herb belonging
to the family of Cyperaceae provides the dwarf manatee with its preferred staple food. Then, the Rio Arauazinho is teeming
with stingrays in all sizes re presenting a true danger, for dwarf manatees expose their bellies while standing on their flippers
browsing pastures of E. minima. Dwarf manatees are reported to associate with “jaraqui” fish Scmapmchilodus insignis (in
a picture above seen swimming near a PaleOSUchuS caiman). The fish’s sharp eye-sight together with the manatee’s ex-
traordinarily keen sense of hearing seem to provide both species with the perfect audio-visual protection against electric eel
and potential predators, such as anaconda, jaguar and man fishing with bow and arrow or harpoon.
swamps and lakes located in the northwest (Fig.
17). From there, they had wandered down during
the dry season while feeding upon Eleochcifis
minima meadows that grow locally on a thin layer
of arenite sand overlying the pre-Cambrian
sandstone bedrock (Fig. 22). During the wet season
the entire population is believed to hibernate there
while feeding upon Eleocharis minima, two aquatic
coarse-leaved Thuvnia H o ok. f. species belonging
to the aquatic plant family Thurniaceae, one
Cabomba Aubl. (Cabombaceae) species locally
called “camarao”, a wild variety of rice ( Oryza sp.)
and several algae (called “sulape”). All these aquatic
food plants grow abundantly in the narrow but
locally deep river itself and in the wetlands it
drains, that contain numerous clear-water lakes,
ponds and swamps dominated by MauritiafleXUOSa
L.f. palms. During the dry season, some animals
browsing Eleocharis minima meadows may
descend as far as seven km from the mouth of the
Rio Arauazinho. Before it flows into the Rio
Aripuana, the Arauazinho widens into a 0-30 cm
deep lake. During the summer T. inunguis cannot
enter and T. pygmaeilS cannot leave the mouth and
lower course of the Rio Arauazinho. Therefore, the
two parapatric manatee taxa cannot hybridize.
238
Marc G.M. van Roosmalen
Performing above-substrate browsing in a hori-
zontal feeding posture, dwarf manatees appear to
have adapted to feeding on (sem i - ) a q u atic herbs
that grow attached to the sandy bottom of shallow,
fast-flowing clear-water streams. Its seasonally
available preferred staple food is Eleocharis
minima, a Cyperaceae grass that grows in sub-
mersed pastures up to 1 m below the surface on a
shallow arenite -sandy substrate overlying the
sandstone bedrock.
It has edible leaves and rhizomes that the dwarf
manatee easily pulls whole from the sandy substrate
with its trunk and lips. Chewing the entire plant in-
cluding the sand-containing rhizomes seems to be
responsible for the strong abrasion of the molars as
seen in adult T. pygmaeus n. sp. (Figs. 20, 2 1). In
contrast, T. inunguis feeds on floating and sub-
merged plants in deeper waters (>2 m deep), being
consumed in situ or, in the case of floating plants,
taken below the surface and manipulated into the
mouth by the flippers, preferentially if depth allows
in a vertical position. While foraging in shallow
waters, dwarf manatees when detecting people
walking or canoeing along the riverbank immedi-
ately seek seclusion in the deep pools found in river
bends. There, they stay underwater for three
minutes at the most. When on ease, they slowly
come to the surface and take a breath every 30-55
seconds.
TrichechuS inunguis, when persecuted, can stay
underwater up to 20 minutes without breathing.
According to the locals and confirmed by our own
observations, dwarf manatees tend to associate with
schools of “jaraqui” fish (Semaprochilodus insignis
- Prochilodontidae) while browsing on Eleocharis
minima. This polyspecific association helps to
protect them against defensive shocks from electric
eels, and attacks of potential predators such as over
8 m long anacondas and spotted jaguars (Fig. 22).
Dwarf manatees are considered critically en-
dangered due to their most restricted geographical
and ecological range, small population size (we
estimate it to be less than 100 individuals), value as
game, and their extremely vulnerable and delicate
preferred habitat, clear-water streams and wetlands.
The skull of the type specimen is recovered from
game occasionally killed with bow and arrow and
eaten by the locals. Habitat favorable to dwarf
manatees occurs, aside of the Rio Arauazinho, only
in the basins of two other clear-water tributaries of
the lower Rio Aripuana - Rio Aracu and Rio Juma.
Trichechus pygmaeus n. sp., though, is not
reported to exist there. The Rio Aracu Basin has
been completely destroyed after a colonization
scheme was implanted by the local government in
the late 1970s. The Rio Juma Basin has been signi-
ficantly affected after a goldmine was opened in
2006. Over 10,000 people flocked into the area
polluting the Juma and Aripuana Basins using high-
pressure hose-pipes and large amounts of mercury.
Illegal mining of gravel and gold, timber extraction,
commercial hunting and fishing in the Rio Aripuana
Basin pose serious threats to the survival of both
Amazonian manatee species. The discovery of T.
pygmaeUS adds to the uniqueness of the lower
Aripuana Basin and shows once more that it is a
poorly explored hotspot of biodiversity and
endemism. My biodiversity surveys conducted after
the year 2000 indicate that the region harbors at
least seven primates new to science, four of which
being described, including the new genus Callibella
never reported orcollected before (Van Roosmalen
et al., 1998; Van Roosmalen et al.,2000; Van Roos-
malen et al., 2002; Van Roosmalen & Van Roos-
malen, 2003; 2014; Van Roosmalen, 2013b; 2015).
Disturbingly, there is not a single officially protec-
ted area in the entire basin.
Conclusion. Trichechus pygmaeus n . s p . , th e
dwarf manatee, represents a second taxon of living
fresh-water manatees and the smallest (130 cm in
length) of all extant sirenians. The new species
differs from the other known western Atlantic
manatees, T. inunguis and T. manatUS, in being two
to three times smaller, with a more streamlined, less
bloated appearance, a deep black instead of dark
greyish skin, a large symmetrical, circular to tear-
shaped white patch on the abdomen in at least the
males (and reported equally in the females), a
shorter head and shorter flippers, the tips of which
do not reach the mouth (Fig. 19).
In September 2002, the author collected a
complete skull of a recently killed adult male. Two
years later, he could film, photograph, examine, and
study for the first time an adult male dwarfmanatee
while keeping it alive for over four months in its
natural habitat. It then escaped and returned to its
natural environment. Figure 23 shows the fenced-
off river bend along the lower course of the Rio
Arauazinho in which we kept, fed and observed for
Hotspot of new megafauna found in the Central Amazon (Brazil): the lower RioAripuana Basin
239
over four months a solitary adult male dwarf
manatee that was captured by a local from Araua-
zinho while feeding on Eleocharis minima at ab o u t
seven km from the confluence with the Aripuana
River. Floating vegetation was systematically re-
fused. Food plants we brought in from the nearby
river consequently had to be fixed onto the sandy
bottom of his pan in order to be recognized as food,
browsed and eaten in a horizontal feeding posture.
Nine years later, Van Roosmalen and Van der
Vlist conducted an expedition by canoe and found
the last existing population of dwarf manatees in
the wetlands situated along the northern branch
of the upper Rio Arauazinho near the watershed
with the Rios Urua and Mariepaua (Figs. 17, 22).
Dwarf manatees were found to be fully adapted to
foraging in fast-flowing shallow clear-water
streams. Standing on their flippers they browse in
a horizontal position on aquatic grasses and other
non -floating plants that grow on or near the bottom.
In contrast, the three times bigger common fresh-
water manatee T. iviUTlguis is restricted to calm
Figure 23. The fenced-off river bend along the lower course of the Rio Arauazinho. Here we kept, fed, filmed and observed
for over four months a solitary adult male dwarf manatee that was cap tu red at seven km fro m the mouth of the Rio Arauazinho
by a local from Arauazinho village.
240
Marc G.M. van Roosmalen
waters of rivers and lakes of the black- and white-
water types offering limited visibility.
It feeds on floating aquatic plants and sub-
mersed foliage of floodplain (igapo and varzea)
plant species. Mitochondrial control region DNA
sequences revealed a haplotype identical to T.
lYlUYlglllS. We believe that this resulted from some
gene flow that must have taken place in the past,
as the two taxa are parapatric and only allopatric
in ecological respect.
We consider the dwarf manatee at the verge
of extinction, for only the headwaters of the
northernmost branch of Rio Arauazinho, a 120
km long left-bank clear-water tributary of the Rio
Aripuana, are thought to harbor a viable relict
pop u latio n .
DISCUSSION
In phy to-sociological respect, the many scrub
and open savannas on white-sand alluvial soils in
the entire Rio Aripuana Basin are unique and found
nowhere else in the Amazon. Together with the
adjacent low savanna forests their branching pattern
seen on satellite images does indicate that the entire
basin preceding the Late Pleistocene was drained
southward - instead of northward like the Rio
Aripuana and its tributaries nowadays drain the area
into the Madeira River, and through the Rio
M adeira into the Amazonas and eventually into the
Atlantic Ocean. The alluvial sand deposits of
former Pliocene and Early Pleistocene creeks and
rivers show a branching pattern in their headwaters,
mean in g toward their northernmost end. Toward the
southernmost end of the basin, where the Rio
Aripuana later in the Pleistocene originated, is situ-
ated nowadays the Tenharim Savanna, a large con-
tinuous savanna area. It is located east of the city
of Porto Velho, close to the Rio Jl-Parana, a river
that together with the Rio Guapore drains the
western part of the Brazilian Shield into the Rio
Madeira. The huge Tenharim Savanna has been
interpreted by geo-morphologists as the result of
sedimentation in a Quaternary long-lived clear-
water inner lake. If we look at the geo-morpholo-
gical history of the Rio Aripuana Basin, it is
assumed that during part of the Miocene a large
freshwater inland lake existed, called the Beni
Lake.
This lake stretched westward across the
Bolivian Amazon. The brackish-w ater marine
molasses-lakes from the Oligocene might have
turned in the Miocene into the fresh-watermolasses-
lakes, and in the Pliocene into the sub-andine inner
or inland lakes. During the Pliocene and Pleistocene
these lakes filled with rainwater flowing down from
the eastern foothills of the (by then) higher Andes
Mountains (Hoorn & Wesselingh, 2010). In the
Pleistocene era, three main drainage systems or
basins were formed in form er A m azonia: the white-
water basin influenced by the eroding volcanic
Andes in the western part of Am azonia, drained by
the proto-Amazon River flowing toward the At-
lantic Ocean; the clear-water basin draining the
so u th -east A m azonian crystalline Brazilian Shield
with the watershed running across the Chapada dos
Parecis toward the south through the proto-Beni,
proto-M am ore, and proto-G uapore Rivers; thirdly,
the black-water basin draining the northern Amazo-
nian alluvial white-sand area through the proto-Rio
Negro. During the Late Pleistocene oceanic levels
repeatedly have dropped on a global scale and the
sub-andine inner lakes were quickly emptied by the
much stronger eroding power of the proto-Amazon
rushing toward the Atlantic Ocean - its surface
lying 100-120 m lower during the subsequent ice
ages of the Pleistocene.
During the glacial periods of the Late Pleisto-
cene (1-2 M YA) the ancient continental watershed
running across the Chapada dos Parecis has been
broken through by the proto -M adeira River, which
in turn was connected with the mighty Amazon
River. The M adeira /Amazonas drainage system, as
a way of speaking, then ‘sucked’ its way through
the watershed powered by huge water volumes on
their way to the up to 120 m lower lying water table
of the Atlantic Ocean. The vortex holes in what a
geologist would call an “unripe riverbed” - in the
400 km long stretch of the upper M adeira River and
a shorter stretch in the middle Rio Aripuana, as well
as in the Rio Roosevelt - tell the tale about a former
battle over one watershed between two drainage
systems, each draining one side of it. The proto-
Madeira and Amazonas Rivers thus conquered the
clear-water catchment area of the Brazilian Shield.
From then on, they made a connection with what
was left of the former Pantanal/C haco Lake through
the Mamore, Beni, and Abuna Rivers. Thereafter,
these white-water rivers began to leak the sub-
Hotspot of new megafauna found in the Central Amazon (Brazil): the lower Rio Aripuana Basin
241
andine Bolivian drainage system, this time to the
north instead of to the east, connected as they now
were with the Madeira and Amazon Rivers. The
P an tan al/C h ac o Lake was quickly emptied out to
the east through the Madeira flowing into the
Amazon and then into the Atlantic Ocean. In the
northern part of the former Pantanal/C haco Lake
one or more clear- water lakes that had formed there
since the Pliocene, were now also emptied out by
the combined Rios Madeira/Amazonas drainage
sy stem .
One of these large clear- water lakes was situated
exactly where nowadays the Tenharim Savanna is
located, just north of the pre-andine watershed
running across the Chapada dos Parecis. This
Tenharim Lake was so far fed by rivers running in
a north-south direction within what is nowadays the
larger Rios M adeira/A m azonas/Tapajos-Juruena
in te rflu v i- u m (the Tenharim Lake was drained
southward toward the Pantanal through the proto-
Guapore River). After the conquest of the water-
shed by the combined Madeira/Amazonas drainage
system, rivers such as the Ji-Parana, Roosevelt,
Guariba, and Aripuana began to flow north- and
westward, this way draining the entire Aripuana
Basin directly into the Rio Madeira. Clear evidence
that rivers like the Aripuana and Roosevelt origin-
ated in a more recent geological era (the Late Plei-
stocene) is the occurrence of so-called
“ S tru d ello ch e rn ” in the crystalline bedrock of
the middle and upper courses of these rivers. In the
Rio Aripuana, south of Prainha, are nowadays
found the unsurpassable Periquito Falls, among
other extensive stretches of rapids and waterfalls.
Moreover, the very deep deposits of gravel in the
form of small brown rounded-off, polished pebbles
that are laid down in calmer waters downstream of
the rapids, assign to the afore-mentioned geological
(vicariance) event of a Pleistocene watershed
break-through.
Once the complex history of S o u th - A m eric a ’s
continental landscape and river systems, and, in
particular, the relatively recent (Pliocene through
Pleistocene) geo-morphological model of the
Aripuana River’s drainage system is clearly under-
stood, about all demographic and evolutionary odds
of this river basin, that were hitherto considered ‘hit
and miss’ distributions, may now be plausibly
explained for. It seems that during a large part of
the Pliocene and Early Pleistocene eras the entire
pre-Aripuana river drainage system with its pre-
dominantly clear-water habitats was effectively
blocked off from Amazonia west and north of the
proto-Madeira River, for it was drained by rivers
flowing southward toward the eastern part of the
late-Miocene sub-andine Pantanal/C haco Lake, and
from there into the Atlantic Ocean. The continental
watershed built from pre-Cambrian crystalline
rock, together with the (those days) extensive lacus-
trine habitats, effectively isolated this peripheral
drainage system from sub-andine w hite-w ater river
systems, that were drained by the p ro to -M ad e ira
and Amazon Rivers. Over millions of years oppor-
tunities for allopatric divergence were provided, for
no gene flow could take place between non-volant
terra firm e and aquatic fauna of the clear-water
drainage system and the rest of Amazonia, which
was drained to the far northeast into the Atlantic
Ocean. A number of ground- as well as tree-
dwelling vertebrates, but also aquatic (mostly mam-
malian and mollusk) fauna, could therefore evolve
in seclusion. The first vicariance must have taken
place already in the E arly -Plioc en e , about 5 MYA,
when ancestral proto/archetypical forms of a 1 1 -
Amazonian generic groups (i.e., the marmosets,
spider-, woolly-, capuchin-, saki-, tit i- , night- and
howling monkeys, tapirs, anteaters, rodents like
porcupines, pacas, agoutis, and acouchis, manatees,
and ‘botos’) have diverged from closely related
species found in the rest of the Amazon - to the west
and north of the proto-Madeira and Amazon Rivers.
A second, more dramatic vicariance took place
during one or more of the glacial epochs of the
Middle Pleistocene, about 1-1.8 MYA, the break-
through of the continental watershed by the proto-
Madeira, being in turn powered by the modern Rio
Amazonas drainage system in times that ocean
levels had dropped over 120 m. So far, this water-
shed had run across the Serra dos Parecis in the
Brazilian State of Rondonia. This way, the modern
M adeira River originated and, at the same time, the
Rios Aripuana, Ji-Parana, Tapajos-Juruena, and,
perhaps, also the Rios Xingu and Araguaia,
although the headwaters of the latter two rivers are
found in the ‘cerrado’ (white-sand savannas) of
Mato Grosso. These rivers also cleared themselves
a way through the watershed of the former clear-
water north-south directed drainage system and
242
Marc G.M. van Roosmalen
began to empty their waters into the modern
Madeira and Amazonas Rivers. From then on, the
Rio Madeira became the rather straight and fast-
flowing, second largest river barrier in the entire
Amazon Basin, after the Amazon proper.
Ever since, no gene flow of terrestrial mega-
fauna could take place to and from the western and
northern Amazon. The RiosAripuana and Ji-Parana
first emptied out the former Tenharim Lake into the
modern Madeira, there where for a long time lacus-
trine environments and wetlands had deposited
white sand. The former clear-water drainage system
left behind, aside of the Tenharim Savanna, many
smaller patches of white-sand savanna and savanna
forest on alluvial sandy soils deposited by former
Pliocene and Early Pleistocene rivers and streams.
Locally, new rivers arose and began to drain these
areas dotted with white-sand savannas and stretches
of savanna forest. That explains why they are of the
black-water type. To name a few: the Rios Araua,
Mariepaua, Urua, Manicore,Atininga, Canuma, Su-
cunduri, AcarL Some local rivers draining areas that
do not contain alluvial white-sand deposits, but
instead having heavily weathered pre-Cambrian
arenite (sandstone) reaching the surface, remained
of the clear-water type, such as the Rios Aracu,
Arauazinho, and Juma. The entire former (Plio-
cene) clear-water drainage system, from then on,
was intersected by new rivers draining the system
in opposite ( so u th -n o rth ) direction, most import-
antly the Rios Aripuana, Tapajos- Juruena, Xingu,
Teles-Pires, and Araguaia. In the course of several
millions of years, a different biome could develop
in this SE Amazon clear-water drainage system
harboring a mixture of endemics and newcomers.
The latter were ancestral forms of non-volant
terrestrial mammals that, after cros- sing the
Panamanian land bridge formed between 2.5-3
MYA, had migrated into the northern and western
sub-andine regions of the Amazon. Subsequently,
some managed to traverse the proto-M adeira River
and established themselves in most of this ancient
clear-water drainage system.
As such, newcomers such as ancestral collared
and white-lipped peccaries, jaguars, pumas, small
cats, canids, coatis and mustelids, lived side by side
with endemics such as primates, rodents (squirrels,
pacas, agoutis, acouchis, capybaras, spiny rats),
marsupials (D idelphidae), edentates (anteaters.
armadillos, sloths), tapirs, and porcupines. Not
before the last glacial period of the Holocene, about
10,000 YA, some modern mega-fauna elements,
among which the common spotted paca A. pdCd,
that had evolved west and north of the strong
geographic barrier formed by the non-meandering
Rio Madeira, managed to circumvent the head-
waters of the Rio Madeira. Thereafter, they mi-
grated into the Rios Madeira/Tapajos inter fluvium.
They took either the northwestern route following
the western border of the Tenharim Savanna, or the
southeastern route circumventing the Tenharim
Savanna along the southern border, or they mi-
grated into the Rios Madeira/Tapajos interfluvium
along both paths. So, they entered a different eco-
system full of phylogenetically related but for-
merly allopatric species endemic to the region.
Consequently, the new-comers may then have out-
competed closely related species that occupied
similar ecological niches, causing their extinction.
Or, one species may have become genetically
absorbed by the other through cross-breeding. Or,
allopatric species may have diverged that much
from one another in habitat and dietary preferences,
foraging strategy and/or social and sexual behavior,
that they were able to co-exist and live on in sym-
patry. The latter scenario may well explain, for
instance, the sympatric occurrence in the Rio
Aripuana Basin of two different species of brocket
deer (i.e., Mazamci nemorivdgd and M. tienhoveni
n. sp.) and two different species of paca (i.e., the
common spotted paca A. pdCd and Silva Garcia’s
striped giant p ac a A. silvdgdvtide n . sp.).
ACKNOWLEDGEMENTS
Molecular analyses were co-financed by the
Treub Stichting (Society for the Advancement
of Research in the Tropics), and performed at
the Animal Breeding and Genomics Centre of
Wageningen University, The Netherlands. The
author received a grant for conducting biodiversity
fieldwork in the Aripuana region from the Van
Tienhoven Foundation for International Nature
Conservation. We are grateful to Rene Dekker and
Hein van Grouw of The Netherlands Natural
History Museum Naturalis in Leiden, who kindly
provided their museum specimens for our com-
parative research on the genus MdZdlTld.
Hotspot of new megafauna found in the Central Amazon (Brazil): the lower RioAripuana Basin
243
REFERENCES
Caloi L., Kotsakis T., Palombo M.R. & Petronio C
1996. In: Shoshani J. & Tassy P. (Eds.), 1996. The
Proboscidae: evolution and palaeoecology of ele-
phants and their relatives. Oxford University Press,
Oxford, 234-239.
Domning D .P. & Hayek L.C., 1986. Interspecific and
intraspecific morphological variation in manatees
(Sirenia: TrichechuS) . M arine Mammal Science, 2:
87-144.
Duarte J.M.B., 1996. Guia de identificagao de Cervldeos
Brasileiros. FUNEP. Jaboticabal.
Duarte J.M.B., Gonzalez S. & Maldonado J.E., 2008.
The surprising evolutionary history of South Amer-
ican deer. Molecular Phylogenetics and Evolution,
49: 17-22.
Eisenberg J.F., 1989. Mammals of the Neotropics. The
Northern Neotropics, Vol. 1. Panama, Colombia,
Venezuela, Guyana, Suriname, French Guiana. Uni-
versity of Chicago Press, Chicago, 449 pp.
Eisenberg J.F. & Redford K .H ., 1999. Mammals of the
N eo tropics, Vol. 3. The Central Neotropics: Ecuador,
Peru, Bolivia, Brazil. Chicago: University of Chicago
Press, Chicago, 609 pp.
Emmons L .H . & Feer F., 1990. Neotropical rainforest
mammals. A field guide. University of Chicago
Press, Chicago, 281 pp.
G arcia-R odriguez A. I., Bowen B.W., Domning D .,
Mignucci-GiannoniA.A., Marmontel M ., M on toy a-
Ospina R.A., Morales-Vela B ., Ruding M ., Bonde
R .K ., & M cG uire P.M ., 1998. Phylo geography of the
West Indian Manatee (TrichechuS IflCinCltllS) : how
many populations and how many taxa? Molecular
Ecology, 7: 1 137-1 149.
Hoorn C. & Wesselingh F.P., 2010. Introduction: Amazo-
nia, landscape and species evolution - a look into the
past. Wiley-Blackwell, Ox ford, 477 pp.
Husson A.M., 1978. The Mammals of Suriname.
Rijksmuseum van N atuurlijke Historic. Zoologische
M onografieen . No. 2, E.J. Brill, Leiden, The Nether-
lands, xxxiv + 569 pp.
Kimura M ., 1980. A simple method for estimating
evolutionary rates of base substitutions through
compa- rative studies of nucleotide sequences.
Journal of M olecular Evolution, 16: 1-20.
Kocher T.D., Thomas W.K., Meyer A., Edwards S.V.
& Paabo S., 1989. Dynamics of m itochondrial-
DNA evolution in animals - amplification and
sequencing with conserved primers. Proceedings
of the National Academy of Sciences, 86: 6 196-
6200.
Lahr M .M . & Foley R.A., 2004. Palaeo-anthropology :
Human evolution writ small. Nature, 43 1, 1043-
1044.
ListerA.M., 1993. Mammoths in miniature. Nature, 362:
288-289.
Rossi R.V., 2000. Taxonomia de Mazama Rafinesque,
1817 do Brasil ( A rtiod ac ty la, Cervidae). Master's
thesis, Instituto de Biociencias, Universidade de Sao
Paulo, 1-174.
Tamura K ., Dudley J., N ei M. & Kumar S., 2007.
MEGA4: Molecular evolutionary genetics analysis
(MEGA) software version 4.0. Molecular Biology
and Evolution, 24: 1 596-1599.
Van Roosmalen M.G.M., 2013a. Barefoot through the
Amazon - On the path of evolution. Paperback, 500
pp. https://www.createspace.com/ 4177494
Van Roosmalen M.G.M., 2013b. Wild fruits from
the Amazon Vol. I. Paperback, 280 pp. https://www.
ere ate space. com/4 177494
Van Roosmalen M.G.M., 2014. Distributions and phylo-
geography of Neotropical primates - A pictorial
guide to all New World monkeys. Paperback, 71 pp.
https://www. ere a tespace.com / 4596480
Van Roosmalen M.G.M., 2015. Live from the Amazon.
Elliot Editori, Rome, Italy, 608 pp.
Van Roosmalen M.G.M. & Van Roosmalen T., 2003. The
description of a new marmoset genus, Ccillibellci
(C allitrichinae. Primates), including its molecular
phylogenetic status. Neo tropical Prim ates, 11: 1-10.
Van Roosmalen M.G.M. & Van Roosmalen T., 2014. On
the origin of allopatric primate species and the
principle of metachromic bleaching. Paperback, 146
pp. https://www.createspace.com/ 4549738
Van Roosmalen M.G.M., Frenz L., Van Hooft P., De
longh H.H. & Leirs H., 2007. A new species ofliving
peccary (Mammalia:Tayassuidae) from the Brazilian
Amazon. Bonner zoologische Beitrage, 55: 105-112.
Van Roosmalen M.G.M., Van Roosmalen T., Mitterme-
ier R.A. & De Fonseca G.A.B., 1998. A new and
distinctive species of marmoset (C allitrichidae,
Primates) from the lower Rio Aripuana, State of
Amazonas, central Brazilian Amazon. Goeldiana,
Zoologia, 22: 1-27
Van Roosmalen M.G.M.,Van Roosmalen T., Mittermeier
R.A. & Rylands A.B., 2000. Two new species of
marmoset, genus Cdllithvix Erxleben, 1 111 (Cal-
litrichidae. Primates) from the Tapajos/M adeira
interfluve, south central Amazonia, Brazil. Neo-
tropical Primates, 8: 2-18.
Van Roosmalen M.G.M.,Van Roosmalen T. & Mitter-
meier R.A., 2002. A taxonomic review of the titi
monkeys, genus CcillicebuS Thomas, 1903, with the
description of two new species, CcillicebuS bernhciKcli
and CcillicebuS StephennClsJli, from Brazilian Amazo-
nia. Neotropical Primates, 10 (Supplement): 1-52.
Vergara-P a rente J.E., Parente C.L., M arm on tel M ., Silva
J.C.R. & S a F.B., 2010. Growth curve of free -ranging
TrichechuS inunguis. Biota Neotropica, 10: 89-93.
244
Marc G.M. van Roosmalen
Von Spix J.B . & Von M artius C.F.Ph., 1 823-183 1. Reise
in Brasilien in den Jahren 1 8 1 7-1 820. 3 Vols. + 1
Atlas. Verlag M. Lindauer, Munchen, 1 388 pp.
Reprint 1967, Stuttgart.
Voss R.S. & Da Silva M .N .F., 2001. Revisionary notes
on Neotropical porcupines (Rodentia: Erethizon-
tidae). 2. Review of the Coendll VeStitUS Group with
descriptions of two new species from Amazonia.
Novitates, American Museum for Natural Flistory,
New York, 335 1: 1-36.
Webb S.D., 2000. Evolutionary history of new world
deer. In: Vrba E.S. & Schaller G .B . (Eds.), 2000.
Antilopes, Deer, and Relatives. Yale University
Press, London, 38-64.
Wilson D.E. & Reeder D .M . (Eds.), 1993. Mammal
Species of the World. A Taxonomic and Geographic
Reference. John Hopkins University Press, B al-
tim ore, 1206 pp.
Biodiversity Journal, 2015, 6 (1): 245-252
Monograph
The diversity of wild animals at Fezzan Province (Libya)
Mohamed Faisel Ashour Essghaier, Ibrahim MoftahTaboni & Khaled Salem Etayeb*
Zoology Department, Faculty of Science, Tripoli University, P.O.Box:13227, Tripoli, Libya
Corresponding author, e-mail: khaledetayeb@ yahoo.com
ABSTRACT Fezzan province (Libya) is a segment of true Sahara, is characterized by diverse habitats
that are utilized as shelters and feeding ground for many desert wildlife species. Oases with
water table near the surface are the most prominent feature in the Libyan desert. The diversity
in habitats resulted in diversity in wildlife, as well as the plant cover (trees and bushes) is
the most effective factor for the existence and the abundance of wild animals, in particular
bird species. This study observed many species of reptiles, birds and mammals. In the study
is also reported the rock hyrax PWCCIVICI CCipensis Pallas, 1 766 (Hyracoidea Procaviidae) a
rare and endemic species at the area.
KEY WORDS Oases; diversity; endemic; wild animals.
Received 25.06.2014; accepted 30.08.2014; printed 30.03.2015
Proceedings of the 2nd International Cong re ss “Speciation and Taxonomy”, May 1 6 th - 1 8 th 2014, Cefalu-Castelbuono (Italy)
INTRODUCTION
Libya is mostly characterized by arid climatic
conditions, except the coastal strip and the northern
hills toward the east and the west, while the rest of
the country is located under the conditions of desert
and semi-desert because of its geographical location
in terms of latitude. This resulted in the presence of
environments with distinct characteristics in terms of
temperature, humidity and rainfall that reflected on
the biological components of the plants and the
animals that are able to co-exist in various ways with
those difficult environmental conditions (Hufnagel,
1972).
In Libya there are a lot of ecosystems that range
from the coastal environment with all its scattered
salt marshes along the coastline, to green plains in
the northeastern region and northwest highlands
(which include N afusa Mountains), to desert and
semi-desert ecosystem showing its content of oases
and valleys (Toschi, 1 969). The desert is ecologic-
ally sensitive and very important in terms of
wildlife (flora and fauna), which coexist in this
habitat in spite of the harsh living conditions as
much heat, especially during the summer months
in addition to water scarcity and drought. However,
these systems include a few diversity and abund-
ance of species particularly those that have the
capacity to live under these circumstances and
some of them are endemic.
Fezzan province is a segment of true Sahara, is
characterized by many habitats that are utilized as
shelters and feeding ground for many desert wild-
life species (Bundy, 1 976). It is situated in the
southwest of Libya within the desert ecosystem,
which includes desert wadis, oases, palm planta-
tions and irrigated cropland. Studies and reports,
which are relatively scarce since the period of
Italian occupation until the present, concluded that
the wildlife in this region has declined in terms of
number of species and individuals to alarming
situation where some taxa are either subject to
246
M.F.A. Essghaier et alii
extinction or already had disappeared from their
previous range. The reason of it can be attributed
to: (i) urbanization in some areas on the expense
of natural resources and natural vegetation; (ii)
construction of roads leading to the open areas
where the shelters and habitats for wildlife are; (iii)
modern vehicles which facilitated access to rugged
areas; (iv) overhunting; (v) explorations and oil
investments and activities associated with this
industry; (vi) establishment of some sites for the
purpose of various agricultural activities which led
to the presence of human activity with a negative
impact on wildlife communities and, (vii) desert
tourism that led to the emergence of some negative
effects (Sbeta et al., 2006).
In spite of all the mentioned circumstances met
in this region, either due to Nature (high temper-
atures and scarcity of water surfaces and precipita-
tion), or to different activities of humans that have
negative effects on wildlife, especially those onset
early in the sixties, there still are a number of
species inhabiting the desert wadis and oases.
In this study we report some examples of animal
species which were recorded in the region, and
whose presence we hope to continue to record in
some areas of the region, even if they are few in
number. However, wildlife studies in the province
of Fezzan need to focus on areas that may be
susceptible of urbanization and Industrial sprawl,
in order to save (for future generations) what can
be saved of wildlife that still inhabit some of these
sites. This study is based on a review of available
publications and reports since the last decades until
now, supported by field visits to the selected sites
of the region to investigate the species of reptiles,
birds and mammals.
Study area
This study was focused on the diversity of
animals in eight sites (Wadi Al-shati, Sebha,
Traghen, Murzuq, Ubari, Al-awenat, Ghat and
Akakus) at Fezzan region (Fig. 1).
The habitat type is mainly arid (72% of the total
area of the region) whith very harsh environmental
conditions which are unsuitable for the growth of
plants (Sbeta et al., 2006). The rest of the region
can be classified into; irrigated crops area, planta-
tions of palm trees, pastoral land and salt marshes
(Table 1).
MATERIAL AND METHODS
Field visits were conducted in summer 2006 by
the authors of the present paper. Observations,
collection of samples and bird watching started from
dawn to dusk. Opticron binoculars (with magnifica-
tion 10x50) and Optolyth spotting-scope were used
for accounting of birds, as well as for some wild
mammals. Field guides (Heinzel et al., 1 998;
Figure 1. Map of Libya showing the study area.
Habitat type
Area in
hectare
0/
/o
Irrigated agricultural land
1 37.500
0.25
Plantation of palm trees
39.675
0.07
Pastures and dry valleys
80.65 1
0.15
Arid land
40,137.1 85
72.25
Sands and sand dunes
15,109.334
27.20
Salt marshes (Sobkhas)
36.536
0.07
U rban areas
1 1 .949
0.02
Total
55,552.830
100.00
Table 1. The percentage of each habitat type in Fezzan
province. Source: Project of natural resources mapping for
agricultural use and planning (Libya/04).
The diversity of wild animals at Fezzan Province (Libya)
247
Mullarney et al., 2001) were used to identify birds.
However, the status of bird species was assigned by
their frequency of occurrence.
The following categories were adapted from
Bundy (1 976) and Toschi (1 969): MB - Migrant
breeder; PV - Passage visitor; RB - R esident breeder;
W V - Winter visitor. Reptiles were collected by us-
ing rubber bands, while life traps were fitted for
overnight to catch rodents species.
RESULTS AND DISCUSSION
Amphibians and Reptiles
Despite the harsh climatic conditions in the
whole province, which is inappropriate for the
presence of life, the streams and cultivated areas,
including some wetlands easily available, provided
an opportunity for some amphibian species to
inhabit the area.Amphibian diversity in the Mediter-
ranean basin is much lower than reptile diversity.
This being largely a reflection of the extent to
which arid and semi-arid habitats predominate in
large parts of the region (Cox et al., 2006). In this
study two species of amphibians and fourteen
species of reptiles were encountered (Table 2).
The sub desert Toad Amietophrynus xeros,
previously known as Bufo XeWS, was observed in
pools and farms in Sebha and Ghat, this finding is
in accordance with the results of Ibrahim (2008).
We also observed Green Toad Bufo viridis in either
COMMON NAME
SCIENTIFIC NAME
SITE OF OBSERVATION
1
S ub desert Toad
Amietophrynus xeros
(Tandy, Tandy, Keith et
D uff-M acKay, 1976)
S ebha. Ghat
2
Green Toad
Bufo viridis (Laurenti, 1768)
Sebha, Murzuq, Al-awenat
and Ghat
3
B ib ron’s agam a
Agama impalearis b oettger, 18 74
Al-awenat, Akakus
4
D esert agam a
Trapelus mutabilis
(M errem , 1 8 2 0)
Ubari, Al-Awenat
5
B ell’s dabb-lizard
Uromastyx acanthinura
B ell, 1825
Al-shati, Traghen, Al-awenat
6
Ragazzi’s fan-footed gecko
Ptyodactylus ragazzii
A nderson, 1898
T raghen
7
Elegant gecko
Stenodactylus sthenodactylus
(Lichtenstein, 1 823)
T rag hen
8
Moorish gecko
Tarentola mauritanica
Linnaeus, 1758
Al-shati, Sebha, Traghen,
M urzuq
9
Tripoli dwarf gecko
Tropiocolotes tripolitanus
Peters, 1880
Al-Shati, Sebha, Traghen
10
N idua lizard
Acanthodactylus scutellatus
(Audouin, 1827)
Traghen, M urzuq, Ubari
11
Leopard Fringe-fingered Lizard
Acanthodactylus pardalis
(Lichtenstein, 1 823)
M urzuq
12
Red-Spotted Small Lizard/
D esert-Racer
Mesalina rubropunctata
(Lichtenstein, 1 823 )
Traghen, Murzuq
13
O cellated skink
Chalcides ocellatus
Lorsskal, 17 75
Al-Shati, Sebha, M urzuq
14
Sand fish
Scincus scincus (Linnaeus, 1758)
Al-Shati, Traghen, Ubari
15
Schokari Sand Snake
Psammophis schokari
(Forskal, 1775)
Traghen, Ubari
16
Horned viper
Cerastes cerastes Linnaeus, 1758
Al-shati, Sebha, Traghen,
M urzuq
Table 2. Species of amphibians and reptiles recorded in study area.
248
M.F.A. Essghaier et alii
cultivated lands or wetlands in Sebha, M urzuq, Al-
awenat and Ghat. Scortecci (1935) mentioned the
presence of this species in the province of Fezzan.
Studies on reptiles are very rare, but the desert
valleys and some habitats in the region are the most
important areas for some species of lizards, such as:
Desert monitor VcirCMUS griseUS (Daudin, 1803),
Chameleon CHaniaeleO chcilflUeleon Linnaeus, 175 8
and Spiny-tailed lizards UromClStyX ClCanthinura.
Furthermore, the most important species of snakes
that live in this environment is the Florned desert
viper Cerastes cerastes (Bennett, 1 970; Awami,
1976; Ibrahim, 2008). A total of 14 species of rep-
tiles were recorded in the present paper in the
province (Table 2). However, the majority of them
are mentioned in some previous studies (e.g.
Kramer & Schnurrenberger, 1963; Schleich et al.,
1996; Frynta et al., 2000; Ibrahim, 2008), except
the Desert agama TrapelliS YYllltabUis which was
observed in Al-Awenat and Akakus and recorded
for the first time in these sites (Fig. 2). Schleich et
al. (1996) reported the presence of this species in
Cyrenaica (east to Tubruk) and Wagner et al. (2011)
mentioned another record of this species in Tripoli.
Birds
The present study accounted a total of 2975 in-
dividuals belonging to 26 bird species; the majority
of them were non-w aterbirds species with a disparity
in numbers of species and individuals between sites
(Table 3). A total of 12 species were reported as
resident breeders (Bundy, 1976). However, many
previous studies during decades ago reported the
presence of more than 100 species as winter visitors
during their migration from Asia and Europe to
Africa, where they stop for few days and then
continue to the south. While around 20 species we re
recorded as residents along the year seasons such as;
Sandgrouses Pteracles orientalis, Pteracles senegal-
lus, Owls Bubo bubo. Partridge Alectoris barbara
and some species of raptors (Toschi, 1969; Bundy,
1976; Brehme e t al. , 2002 a, b , 2 00 3 a , b , 2004 ).
Fezzan province is composed of many oases,
cultivated areas, irrigated crop sites, urban and res-
idential areas, these may provide roosting sites and
shelters for many bird species, particularly, those
who adapted to live within and adjacent to anthro-
pological environments. This reflects the large num-
bers of sparrows that inhabit the urban areas
(Spanish sp arro w andDesertsparrow;fig.3),whilst
those species were absent in Akakus.
Furthermore, there was a difference in species
diversity among the study sites depending on habitat
types. Five w aterbirds species ( Ardea dtierea,
Egretta garzetta, Ardeola ralloides, Anas quer-
quedula and Gallinula chloropus) were observed in
Sebha (sewage site) and Ubari (oases); while the rest
of species were found on plant covers (bushes,
shrubs and trees; pers. obs.).
Mammals
The province of Fezzan is reasonably character-
ized by good diversity of Mammal species. During
this study a total of 11 species were recorded.
Order Erinaceomorpha and Chiroptera
Two species of hedgehogs belong to the family
Erinaceidae were observed: Long-eared hedgehog,
Hemiechinus auritus S.G. Gmelin, 1770 and desert
hedgehog, ParaechinUS aethiopicus (Ehrenberg,
1 832) close to the farmlands in Traghen and
M urzuq. These two species are common in the area
(H ufnagel, 1 972).
A bat species from family Vespertilionidae ( Pipis -
terellus sp.) was observed just after the sunset at all
visited sites. As all species of mammals in the south
of Libya, bats need to be addressed in a comprehens-
ive study in order to identify the extant species and
their relations to other bats populations in the north.
Order Carnivora
Of this group of mammals, only two species
where recorded, the Jackal CaYlis aureus Linnaeus,
1 758 was only identified by tracks left in sites in
Traghen. It usually inhabits areas with optimum
food and shelter. This species is reported in different
types ofLibya habitats (Hufnagel, 1972). However,
IUCN classified this species as Least Concern, due
to its widespread range, but due to the urbanization
and destruction of natural habitats, these animals
were no longer seen in the nature (pers. observa-
tions). Lurthermore, a Caracas of Lennec, VulpeS
Zmfl Zimmermann, 1 780 was found on the road
between Al-awenat and Ghat. Despite, this species
is very common in the province; especially close to
human dwellings.
The diversity of wild animals at Fezzan Province (Libya)
249
Scientific name
Common
name
Wadi
Al-shati
Sebha
Traghen
Murzuq
Ubari
Al-
awenat
Ghat
Akakus
Status
1
Ardea cinerea
Linnaeus, 1758
Grey heron
-
l
-
-
-
-
-
-
PV
2
Egretta garzetta
(Linnaeus, 1766)
Little egret
-
6
-
8
-
-
-
-
PV
3
Ardeola ralloides
Scopoli, 1769
S quacco
Heron
-
5
-
-
-
-
-
-
PV
4
Ciconia ciconia
Linnaeus, 175 8
W hite stork
-
-
-
-
36
1 died
-
-
PV
5
Anas querquedula
Linnaeus, 1758
G arg aney
-
8
-
-
-
-
-
-
PV
6
Circus aeruginosus
Linnaeus, 1758
M arsh
harrier
-
-
-
-
2
-
-
-
PV
7
Falco biarmicus
Temminck, 18 2 5
Lanner
Falcon
-
-
-
-
5
-
-
-
RB
8
Gallinula chloropus
(Linnaeus, 1758)
M orhen
-
1
-
3
7
-
-
-
RB
9
Pterocles coronatus
Lichtenstein, 1823
Crowned
S andgrouse
-
-
-
-
38
-
-
-
RB
10
Columba livia
G m elin, 1789
Rock dove
-
-
-
-
-
-
77
55
RB
11
S treptopelia turtur
(Linnaeus, 1758)
T urtle dove
-
50
38
20
22
-
-
-
M B
12
Streptopelia
senegalensis
(Linnaeus, 1 7 76)
Laughing
Dove
60
17
70
30
50
42
28
-
M B
13
Apus pallidus
Shelley, 1870
Pallid sw ift
-
65
-
-
30
-
-
-
M B
14
Galerida cristata
(Linnaeus, 1 758)
Crested lark
-
-
-
-
-
1
19
-
RB
15
Ammomanes
deserti
(Lichtenstein, 1 823)
D esert lark
-
-
-
-
23
7
15
-
RB
16
Riparia riparia
(Linnaeus, 1758)
S and m artin
-
-
-
-
-
-
33
14
PV
17
Cercotrichas
galactotes
(Temminck, 1820)
R u fo u s
Bush Robin
18
-
-
10
1 7
14
18
-
PV
18
Oenanthe leucopyga
(Brehm, 1855)
W hite-
cro w ned
W heatear
18
-
17
1 1
22
36
14
2
RB
19
Acrocephalus
scirpaceus
(H erm an n , 1 8 04)
Reed
w arb ler
-
18
-
18
-
-
-
-
PV
20
Iduna pallida
(H em prich et
Ehrenberg, 1 833)
O livaceous
W arb ler
22
-
-
-
1 1
5
17
-
M B
Table 3. Numbers of birds species observed in Fezzan province and their status (continued).
250
M.F.A. Essghaier et alii
Scientific name
Common
name
Wadi
Al-shati
Sebha
Traghen
Murzuq
Ubari
Al-
awenat
Ghat
Akakus
Status
21
Lanius meridionalis
(Temminck, 1 820)
G reat grey
shrike
17
29
-
12
2
-
-
-
RB
22
Turdoides fulva
(D esfontaines, 1789)
Fulvous
Babbler
28
-
14
13
9
-
-
-
RB
23
Corvus ruficollis
Lesson, 1830
B ro w ii-
ii e c k e d
Raven
9
-
-
7
100
12
-
-
RB
24
Passer
hispaniolensis
Temminck, 1820
Spanish
Sparrow
150
200
120
150
150
160
300
-
W V
25
Passer simplex
(Lichtenstein, 1 823)
D esert
Sparrow
-
50
50
100
-
-
90
4
RB
26
Emberiza sahari
Levaillant, 1850
House
B unting
-
-
-
-
-
-
25
-
RB
TOTAL
322
450
309
382
524
277
636
75
2975
Table 3 (continued). Numbers of birds species observed in Fezzan province and their status.
o rder Hyracoidea
One of the most important finding of this study
is the observation of Rock hyrax, PwCdVid Cdpeiisis
Pallas, 1 766. It occurs throughout most of Africa
from the southernmost tip north to a line from
Senegal throughout southern Algeria, Libya and
Egypt into the Middle East, except Congo and
Madagascar (Olds & Shoshani, 1 982). The rock
hyrax is one of the four living species of the order
Hyracoidea, and the only living species in the genus
PrOCClvici Storr, 1780. However, the distribution of
this species in Libya is limited to the far southern
mountains (Hufnagel, 1 972). Study on distribution,
density and biology of this species in Libya is
needed. We had a visit to Akakus mountains, where
usually this species had been found, but we could
not observe any individuals in the area. However,
our observation of this animal is based on four
captives of this species kept in an old rocky house
of a local family (Fig. 4).
o rder Artiodactyla
Even-toed ungulates are more or less rare in the
province. In this study two species of family Bo-
vidae were sighted; Barbary sheep, AnWIOtVdgUS
lervia Pallas, 1 777 and Dorcas gazelle, GdZelld
doTCCIS Linnaeus, 1 75 8 ). Only horns of barbary
sheep were recovered in Al-Awenat (Fig. 5a). It was
a clear evidence of the presence of this species
around the area. Moreover, locals emphasized that
this species still exists in the region. A total of 12
Dorcas gazelle were observed in Al-jaza’a protected
area in Al-Shati (Fig. 5b). This species cover a wide
range in Libya (Bennett, 1 970; Hufnagel, 1 972;
Essghaier, 1980), but population trend has recently
declined due to overhunting and habitat destruction.
Order Lagomorpha
R ab bit, LepUS sp. is the most common widely
distributed species in Libya (Hufnagel, 1 972);
usually inhabits macchia-type vegetation, grass-
land, bushveld, and semi-desert areas. This species
was observed in the most study sites (WadiAl-shati,
Sebha,Traghen,Murzuq and Ubari).
o rder Rodent ia
Two species of rodents were reported by the
present study, Jerboa JdCulliS jdClluluS (Linnaeus,
1758) of family Dipodidae and Gerbil, GerbilluS sp .
of family M uridae, which is in accordance with the
findings ofHufnagel (1972). A total of 5 specimens
The diversity of wild animals at Fezzan Province (Libya)
25 1
Figure 2. Desert agama TrapeluS llUlt-
abilis (Akakus).Figure 3. A female of
Desert sparrow. Figure 4. Rock hyrax
in an old house in Al-Awenat.
Figure 5. Horns ofBarbary sheep. Figure 6. Dorcas gazelle in Aljaza’a protec ted area in Al-Shati.
252
M.F.A. Essghaier et alii
of Gerbil were caught by life traps at the area
between Wadi Al-shati and Sebha. However, the
distribution of these species can be estimated by
their w holes.
CONCLUSIONS
In conclusion, the present study, conducted
during summer 2006 , documented some species
from different orders of vertebrates. It is also high-
lighted the importance of biodiversity in Fezzan
province. Altough the survey was in summer, and
thus few numbers of species and individuals were
observed, nevertheless, it is emphasized the wild
animal diversity and urged the need to implement a
comprehensive study for the province in different
seasons of the year.
ACKNOWLEDGMENTS
The authors of this paper are grateful to the
Engineering Consul ting Bureau,Ministry of Hous-
ing and U tilities.
REFERENCES
AwamiA., 1976. Guide to NaturalHistory Museum. The
Authority ofArchaeology, Libya, 65 pp.
Bennett C., 1970. W ild animals of Libya. Department of
Zoology, Faculty of Sciences. Field notes, 11 pp.
Brehme S., Thiede W. & Borges E., 2002a. Beitrage zur
vogewelt Libyens, II: P o d ic ip ed id ae bis Anatidae.
O rnithologische M itteilungen, 6: 202-2 12.
Brehme S., Thiede W. & Borges E ., 2002b. Beitrage zur
vogewelt Libyens, III: Accipitridae bis C h aradriid ae .
O rnithologische M itteilungen, 2: 54-66.
Brehme S., Thiede W. & Borges E. 2003a. Beitrage zur
vogewelt Libyens, IV: Scolopacidae bis Ptero-
clididae.OrnithologischeMitteilungen, 11: 391-399.
Brehme S ., Thiede W. & Borges E ., 2003b. Beitrage zur
vogewelt Libyens, V: Columbidae bis Hirundinidae.
Ornithologische M itteilungen, 7/8: 277-287.
Brehme S., Thiede W. & Borges E., 2004. Beitrage zur
vogewelt Libyens, VI: Motacillidae bis Turdidae.
Ornithologische M itteilungen, 6/7: 207-219.
Bundy G ., 1 976. The Birds of Libya. British Ornitholo-
gists Union, 1976, London, NW1 4RY, 102 pp.
Cox N ., Chanson J. & Stuart S. (Compilers), 2006. The
Status and Distribution of Reptiles and Amphibians
of the Mediterranean Basin. IUCN, Gland, Switzer-
land and Camb ridge, UK. vii + 51 pp.
Essghaier M .F.A ., 1 980. A plea for Libya's gazelles.
Oryx, 1 5: 384-385.
Frynta D., Kratochvil L., Moravec J., Benda P., Dandova
R., Kaftan M ., Klosova K., Mikulova P., Nova P. &
Schwarzova L., 2000. Amphibians and reptiles re-
cently recorded in Libya. Acta Societatis Zoologicae
Bohemicae, 64: 1 7-26.
Heinzel H., R.S.R. Fitter & Parslow J., 1995. Birds of
Britain and Europe with N orth A frica and the Middle
East. Harper Collins, London, 384 pp.
Hufnagel E., 1 972. Libyan Mammals. The oleander
press, 85 pp.
Ibrahim A .A ., 2008. Contribution to the herpetology of
southern Libya. Acta Herpetologica, 3: 35-49.
Kramer E. & Schnurrenberger H., 1963. Systematik,
Verbreitung und Okologie der Lybischen Schlangen.
Revue suisse de zoologie, 70: 453-568.
Mullarney K., Svensson L., Zetterstrom D. & Peter J.
Grant., 200 1. The most complete field guide to the
birds ofBritain and Europe. Harper Collins, London,
400 pp.
Olds N. & Shoshani J., 1982. ProCClvid Capensis. M am -
malian Species, 171: 1-7.
Schleich H., Kastle W. & Kabisch K. 1996.Amphibians
and Reptiles of North Africa. Koenigstein, Koeltz
Scientific Books, 627 pp.
Sbeta A., Mansour S., Essghaier M. & Al-hamali K .,
2006. Plant cover and wildlife in Fezzan province.
The third generation project plans. Engineering
Consulting Bureau, Ministry of Housing and Utilit-
ies. Unpublished report, 39 pp.
Scortecci G ., 1 935. Cenni sui risultati di una campagna
di ricerche zoologiche nel Fezzan. N atura, 25: 93-
103.
Schleich HH, Kastle W. & Kabisch K., 1 996. Amphi-
bians and Reptiles of North Africa. Koenigstein,
Koeltz Scientific Books, 630 pp.
ToschiA., 1969. Introduzione alia ornitologia della Libia.
Supplemento alle ricerche di zoologia applicata alia
caccia, 6 : 1-381.
Wagner P., Melville J., Wilms T.M . & Schmitz A., 201 1.
Opening a box of cryptic taxa - the first review of the
North African desert lizards in the TrCipelllS TYlUtClbilis
Merrem, 1820 complex (Squama ta: Agamidae) with
descriptions of new taxa. Zoological Journal of the
Linnean Society, 1 63: 884-9 1 2.
Biodiversity Journal, 2015, 6 (1): 253-262
Results of the eighth winter waterbird census in Libya in
January 20 1 2
Khaled Salem Etayeb 1 *, Ali Berbash 2 , Wajeeh Bashimam 3 , Mohamed Bouzainen 2 , Ashrof Galidana 2 ,
Mokhtar Saied 2 ,JaberYahia 2 & Essam Bourass 2
'Tripoli University, Department of Zoology, P.O. Box 13227, Lybia
2 Environment General Authority, (EGA-Libya), Lybia
3 Libyan Society for Birds (LSB), Lybia
‘Corresponding author
ABSTRACT After sporadic observations and reports on Libyan birds during the last century, a regular census
of wintering birds at Libyan coastal wetlands started in January 2005. Results of each winter
census till 2011 have been published. The survey of 2012 was carried out by the authors of
the present paper. The general aim was to continue the census of wintering waterbirds in Libya,
despite the difficulties that faced the team after the War of Liberation, and the fact that certain
areas, very important for birds, have been declared military areas. A total of 29,3 14 individuals
belonging to 69 waterbird species was counted. Comparatively, the number of sites covered in
2012 was less than that in previous years of the survey. The majority of individuals counted
belong to seven gull species. This survey also observed a total of 56 individuals of Ay thy a
nyroca Guldenstadt, 1770, a Near Threatened species, as well as, for the first time, a single
individual of Canada Goose Branta canadensis (Linnaeus, 1758) in eastern Libya.
KEY WORDS Waterbirds; Aythya nyroca; Canada Goose; Libya.
Received 16.07.2014; accepted 15.10.2014; printed 30.03.2015
Proceedings of the 2nd International Congress “Speciation and Taxonomy”, May 16th- 18th 2014, Cefalu-Castelbuono (Italy)
INTRODUCTION
After sporadic observations and reports of
Libyan birds during the last century, a regular
census of wintering birds at Libyan coastal wet-
lands started in January 2005. Results of each winter
census till January 2011 have been published (e.g.
Azafzaf et al., 2005, 2006; Etayeb et al., 2007;
Hamza et al., 2008).
These field surveys resulted in the publication
of the Atlas of Wintering Waterbirds of Libya 2005-
2010. In addition, 2005 and 2006 results were
published in Wildfowl (Smart et al., 2006) and
recently, results of the seventh winter waterbird
census in Libya (January-February 2011) were
published (Bourass et al., 2013).
The Environment General Authority (EGA),
the official Libyan body responsible for the
implementation of international agreements relat-
ing to biodiversity, co-sponsored the previous
ornithological surveys of wetlands in Libya,
under a Memorandum of Agreement with the
RAC/SPA and AEWA, and with support from
Wetlands International, the Instituto Nazionale
per la Fauna Selvatica INFS (Italy) and the Office
National de la Chasse et de la Faune Sauvage
ONCFS (France).
254
K.S. Etayeb et alii
The survey of 2012 was carried out by the
authors of the present paper. The general aim was
to continue the census of wintering waterbirds in
Libya, despite the difficulties that faced the team
after the War of Liberation and the declaration of
certain very important areas for birds as military
areas. The study also aimed to compare the present
results with the previous results (2005-2011) and
to report on whether there were records of any new
species.
MATERIAL AND METHODS
The survey was principally focused on the far
eastern and western regions, and very few sites in
the middle region of the country were covered (Fig.
1). A total of 42 sites was covered (Table 1); the
survey was carried out in two periods, 3rd-8th Jan
and 22nd-31st Jan. Moreover, unlike previous
years, the survey of 2012 excluded some important
bird sites, because of their declaration as military
sites during the Libyan War of Liberation.
Unfortunately, there was no access to the
Tawergha complex (Qaser Ahmed, Tawergha
Spring and A1 Hisha; 32°00T2,9" N; 15°08'41,9"
E) one of the most important sites for waterbirds,
where numerous species and individuals were ob-
served in previous years (2005-2011). In order to
examine the population trend of waterbirds winter-
ing in Libya from 2005 to 2012, the Living Planet
Index (LPI) was used. The use of LPI was started
in 1997 by the World Wide Fund for Nature (WWF)
to investigate the changes of global biodiversity
over time, especially for measuring the average
trends of vertebrate populations (Loh et al., 2005).
In this paper, the Chain method was used to
calculate the index, where the logarithm of the ratio
of the population of each pair of years was calcu-
lated using the formula:
d t = log(N t /N t . 1 )
where N= population size and t= years (time).
The specific values of dt were generated for nt as:
n t
dt = l Id it
n t i-i
Finally, the index for waterbird populations in
Libyan wetlands in a standard year t was calculated
as:
dt
I t = I t _i 10
RESULTS AND DISCUSSION
The overall number of species and individuals
of waterbirds and non-waterbirds was lower than
"i A
MALTA
MEDITERRANEAN SEA
CRETE
TUNISIA ~
♦ • •
i
. . -
• •
/
4
4
A TT TV T A
LIBYA
EGYPT
Figure 1. Sites included in the winter census in January 2012.
Results of the eighth winter waterbird census in Libya in January 2012
255
S.n
site name
N
E
1
Ajdabiyah GMMR reservoir
30.58
20.34694
2
Ajdabiyah sewage farm
30.69472
20.25889
3
Al Labadia
32.50472
20.89306
4
Al Mallahah
32.89972
13.28694
5
Al Maqarin karstic lakes
32.15917
20.13861
6
Assabri beach
32.13667
20.07278
7
Ayn Taqnit
32.125
12.80722
8
Ayn Zayyanah
32.21389
20.15556
9
Bab al Bahr coast
32.89667
13.16417
10
Benghazi harbours
32.10472
20.05778
11
Bin Jawwad dam
30.80028
18.06694
12
Bou Dzira
32.16833
20.13194
13
Coast Abu Kammash to Ras
Ajdir
33.11139
11.63639
14
Farwah Lagoon
33.08806
11.76028
15
Sabkhat Abu Kammash
33.08389
11.59389
16
Sabkhat al Kuz
32.44083
20.43333
17
Sabkhat al Manqub
32.90944
12.12639
18
Sabkhat al Thama and
Sabkhat Esselawi
32.14944
20.10278
19
Sabkhat ash Shuwayrib
30.72361
20.12972
20
Sabkhat at Tamimi
32.35917
23.07528
21
Sabkhat Ayn ash Shaqiqah
32.81444
21.47972
22
Sabkhat Ayn az Zarqa
32.80444
21.45917
23
Sabkhat Fairuz
32.04333
20.02222
24
Sabkhat Julyanah
32.09028
20.05944
25
Sabkhat Karkurah
31.40111
20.055
26
Sabkhat Millitah
32.83083
12.28278
27
Sabkhat Qaminis and
Sabkhat Jaruthah
31.74528
19.93444
28
Sabkhat Qanfudhah
32.00028
19.98861
29
Sabkhat Ras at Tin
32.60917
23.12222
30
Sea off Farwah Island
33.11639
11.74861
31
Tajura coast
32.89583
13.37
32
Tobruk harbour
32.06861
23.98583
33
Tripoli harbour
32.90167
13.19194
34
Umm al Jarami
32.52444
23.09361
35
Wadi al Mujaynin dam
32.29
13.2525
36
Wadi al Qusaybat and Ain al
Wahsh
32.31639
23.09694
37
Wadi at Tut dam
32.11722
12.42083
38
Wadi Ghan dam
32.23778
13.13083
39
Wadi Ka’am dam
32.39667
14.32917
40
Wadi Ka’am mouth
32.52667
14.44639
41
Wadi Zaret dam
32.10611
12.80333
42
Zuwarah harbour
32.92306
12.12139
that in all years between 2005 and 201 1, as well as
the number of sites covered (Table 2). A total of
29,3 14 individuals belonging to 69 species from 20
families of waterbirds and wetland-dependent
raptors was recorded during January 2012 (Table
3). This survey was mainly focused on the eastern
and western regions, but included some sites in the
middle region of the country.
The population index of wintering waterbirds
in Libya showed fluctuations throughout the years
of census (2005-2012), with peaks of up to more
than 50% in January of 2006, 2008 and 2010 (Fig.
2). Furthermore, the Living Planet Index showed a
population decline in January 2012 of up to 0.3%
for the above mentioned reasons.
Family PODICIPEDIDAE
Three species from this family were counted:
Black-necked Grebe Podiceps nigricollis, Little
Grebe Tachybaptus ruficollis and Great Crested
Grebe Podiceps cristatus (Table 3). These species
were reported in the previous surveys from 2005 to
2011 (Azafzaf et al., 2005, 2006; Etayeb et al.,
2007; Hamza et al., 2008; Bourass et al., 2013). The
largest number was of the Black-necked Grebe with
a total of 495 individuals.
Family PROCELL ARIIDAE
A total of fourteen Yelkouan Shearwater
Puffinus yelkouan was counted during this survey.
Since the start of wintering survey in 2005, Yelko-
uan Shearwater was only observed in winters 2005
(EGA-RAC/SPA Waterbird Census Team, 2012),
and 2011 with a total of five individuals (Bourass
et al., 2013).
Family SULIDAE
Six individuals of Gannet Morus bassanus were
observed in winter 2012 (four in Wadi Ka'am, one
at Tajura Coast and one at Farwah Island). The
number of Gannets ranged from 3 to 40 individuals
during the previous surveys 2005-2011, and the
peak was in January 2011.
Table 1 . Number of sites covered in
January 2012, Libya.
256
K.S. Etayeb et alii
Years
2005
2006
2007
2008
2009
2010
2011
2012
No. WB
29,996
51,698
39,303
53,632
40,369
51,652
34,842
29,314
No. WB sp.
79
85
92
79
65
86
81
69
No. NWB
301,60
146,621
39,130
13,378
13,047
60,000
506,155
2,054
No.
NWB sp.
74
60
69
64
55
60
67
23
Covered
sites
65
56
43
50
49
94
84
42
Period of
census
3-17 Jan
19-31 Jan
3-15 Feb
20-31 Jan
26 Jan-
7 Feb
24 Jan-
3 Feb
29 Jan-
13 Feb
3-8 Jan,
22 Jan-1 Feb
Table 2. Numbers of birds (species and individuals) counted during winters 2005 to 2012, Libya.
WB=Waterbirds, NWB= Non-Waterbirds
Family PHALACROCORACIDAE
In January 2012 this family was represented
only by the Cormorant Phalacrocorax carbo, with
a total of 1357 individuals counted in 25 different
sites. The highest numbers were observed in Wadi
Ka'am and Farwah Lagoon (313 and 236, respect-
ively). Since winter 2005 the total has ranged from
987 to 2606, with a peak in 2010 (EGA-RAC/SPA
Waterbird Census Team, 2012).
Family ARDEIDAE
Five species belonging to this family were ob-
served during the current survey: Cattle Egret,
Squacco Heron, Little Egret, Great Egret and Grey
Heron (Table 3). The highest number was of Cattle
Egret, with a total of 61 1 individuals, and the lowest
of Squacco Heron where only two individuals were
observed in Wadi Ka'am. However, from 2005 to
2010 the number of Squacco Heron ranged from 2
to 5 (EGA-RAC/SPA Waterbird Census Team,
2012). Relatively, the numbers of the other species
of this family were at the same levels for the years
2005 to 2011 (Bourass et al., 2013; EGA-RAC/SPA
Waterbird Census Team, 2012).
Family CICONIDAE
Five individuals of White Stork Ciconia ciconia
Results of the eighth winter waterbird census in Libya in January 2012
257
were counted in A1 Labadia in eastern Libya (Table
1). Bourass et al. (2013) reported a total of 86 indi-
viduals in winter 2011. From 2005 to 2010 numbers
ranged from 4 to 50 (EGA-RAC/SPA Waterbird
Census Team, 2012). However, White Storks are
more common in farmland than in coastal wetlands
(Bundy, 1976).
Family THRESKIORNITHIDAE
The current survey counted 61 Eurasian Spoon-
bills Platalea leucorodia, the lowest total so far; the
peak was in 2011 with a total of 145 individuals
(Bourass et al., 2013).
Family PHOENICOPTERIDAE
A total of 219 individuals of Greater Flamingo
Phoenicopterus roseus was counted in six sites (Al
Mallahah, Sabkhat Millitah, Sabkhat Abu Kam-
mash, Sabkhat Qanludhah, Sabkhat al Kuz and Far-
wah Lagoon). This observation is the lowest among
the years from 2005 to 201 1 . Moreover, the highest
number of Flamingos was observed in 2009 with a
total of 3292 individuals (EGA-RAC/SPA Water-
bird Census Team, 2012).
Family ANATIDAE
A total of 1 1 species belonging to this family
was observed in this survey (Table 3). The highest
numbers were of Shoveler Anas clypeata and Teal
Anas crecca, with totals of 747 and 394 individuals,
respectively. Other species of the family Anatidae
numbered from 1 to 193 (Table 3). However, indi-
vidual numbers of these species were the lowest
recorded, in comparison to the numbers in previous
surveys (2005-2011). Unexpectedly, during count-
ing of birds in Al Labadia on 29th Jan 2012, mem-
bers of the census team observed an individual of
Canada Goose Branta canadensis with a flock of
111 Shoveler, seven Pintail Anas acuta and four
individuals of Ferruginous Duck Ay thy a nyroca.
This is the first record of this species in Libya,
although two other species of geese have been
reported in Libya: White-fronted Goose Anser
albifrons (Bundy, 1976) and Greylag Goose Anser
anser (Bundy, 1976; EGA-RAC/SPA Waterbird
Census Team, 2012). Description of Canada Goose:
Larger than all species of duck, long neck, brownish
body, black head and neck and white patches on the
face. This observation was in early morning. The
team was able to observe this species at a distance
of 100-120 m for more than one hour, using
Swarovski Telescope and Svensson et al. (2010)
guide.
Family PANDIONIDAE and ACCIPITRIDAE
A total of 20 Marsh Harriers Circus aeru-
ginosas was observed in different wetlands along
the coastline and inland. This observation is the
lowest so far as the range was 21-74 individuals
from 2005 to 2011 (EGA-RAC/SPA Waterbird
Census Team, 2012; Bourass et al., 2013).
Although it is mentioned as a winter and passage
visitor (Bundy, 1976), and there is no evidence of
breeding, our observations from field visits to dif-
ferent sites in Libya recorded the presence of
Marsh Harrier in all months of the year. Further-
more, this species is reported as a resident breeder
in Tunisia which is the neighbouring country to
Libya (Isenmann et al., 2005). A solitary Osprey
Pandion haliaetus has been observed during winter
2012 in Tajura Coast. However, a total of four
individuals was observed in 2005 (Smart et al.,
2006) and only one in 2008 (Hamza et al., 2008)
and 2010 (EGA-RAC/SPA Waterbird Census
Team, 2012). The Osprey is reported as a winter
and passage visitor in Libya (Bundy, 1976).
Family RALLIDAE
Unlike previous years, Moorhen Gallinula
chloropus was counted at only three sites (Al Mal-
lahah, Wadi Ka'am and Al Labadia), with a total of
297 individuals (Table 3). However, this number
was in the range of Moorhens (38-701) counted
during the previous years 2005-20 11; the peak was
in 2009 (EGA-RAC/SPA Waterbird Census Team,
2012). Coot Fulica atra was observed in 13 sites,
mostly freshwater wetlands. The total of 901 indi-
viduals is the highest among the previous winter
surveys where the range was 211-763.
Family HAEMATOPODIDAE
A total of 22 Eurasian Oystercatcher Haema-
topus ostralegus was recorded, as usual, at the
westernmost wetlands on the Libyan coastline. The
previous annual maximum was 56 in 201 1 (Bourass
et al., 2013).
258
K.S. Etayeb et alii
Family
Scientific name
Common name
Total
ANAT1DAE
Branta canadensis (Linnaeus, 1758)
Canada Goose
1
Tadorna tadorna (Linnaeus, 1758)
Shelduck
64
Anas platyrhynchos Linnaeus, 1758
Mallard
26
Anas strepera (Linnaeus, 1758)
Gadwall
1
Anas acuta Linnaeus, 1758
Pintail
63
Anas clypeata Linnaeus, 1758
Shoveler
747
Marm aron etta angustirostris
(Menetries, 1832)
Marbled Duck
10
Anas crecca Linnaeus, 1758
Teal
394
Aythyaferina (Linnaeus, 1758)
Pochard
193
Ay thy a nyroca Guldenstadt, 1770
Ferruginous Duck
56
Anas sp.
Duck sp.
22
PROCELLARIIDAE
Puffinus yelkouan Acerbi, 1827
Yelkouan Shearwater
14
SUL1DAE
Morus bassanus Linnaeus, 1758
Gannet
6
PHALACROCORAC1DAE
Phalacrocorax carbo (Linnaeus, 1758)
Cormorant
1357
ARDEIDAE
Bubulcus ibis Linnaeus, 1758
Cattle Egret
611
Ardeola ralloides Scopoli, 1769
Squacco Heron
2
Egretta garzetta (Linnaeus, 1776)
Little Egret
116
Casmerodius albus (Linnaeus, 1758)
Great Egret
19
Ardea cinerea Linnaeus, 1758
Grey Heron
83
CICONIDAE
Ciconia ciconia Linnaeus, 1758
White Stork
5
THRESKIORNITHIDAE
Platalea leucorodia Linnaeus, 1758
Spoonbill
61
PHOENICOPTERIDAE
Phoenicopterus roseus Pallas, 1811
Flamingo
219
POD1CIPED1DAE
Podiceps nigricollis Brehm, 1831
Black-necked Grebe
495
Tachybaptus ruficollis (Pallas, 1764)
Little Grebe
88
Podiceps cristatus Linnaeus, 1758
Great Crested Grebe
82
ACCIP1TRIDAE
Circus aeruginosus Linnaeus, 1758
Marsh Harrier
20
PANDIONIDAE
Pandion haliaetus (Linnaeus, 1758)
Osprey
1
RALLIDAE
Gallinula chloropus (Linnaeus, 1758)
Moorhen
297
Fulica atra Linnaeus, 1758
Coot
901
HAEMATOPOD1DAE
Haematopus ostralegus Linnaeus, 1758
Oystercatcher
22
RECURVIROSTR1DAE
Himantopus himantopus
Linnaeus, 1758
Black-winged Stilt
550
Recurvirostra avosetta Linnaeus, 1758
Avocet
8
BURHINIDAE
Burhinus oedicnemus Linnaeus, 1758
Stone Curlew
35
CHARADR11DAE
Charadrius hiaticula Linnaeus, 1758
Ringed Plover
61
Table 3. Number of waterbird species and individuals counted in January 2012, Libya (continued).
Results of the eighth winter waterbird census in Libya in January 2012
259
Family
Scientific name
Common name
Total
CHARADR11DAE
Charadrius alexandrinus
Linnaeus, 1758
Kentish Plover
339
Pluvialis squatarola Linnaeus, 1758
Grey Plover
44
Pluvialis apricaria Linnaeus, 1758
Golden Plover
430
Vanellus vanellus Linnaeus, 1758
Lapwing
263
SCOLOPAC1DAE
Calidris alba Pallas, 1764
Sanderling
12
Arenaria interpres (Linnaeus, 1758)
Turnstone
27
Calidris alpina Linnaeus, 1758
Dunlin
1781
Calidris ferruginea Pontoppidan, 1763
Curlew Sandpiper
3
Calidris minuta Leisler, 1812
Little Stint
231
Tringa glareola Linnaeus, 1758
Wood Sandpiper
8
Tringa ochropus Linnaeus, 1758
Green Sandpiper
16
Actitis hypoleucos Linnaeus, 1758
Common Sandpiper
22
Tringa totanus Linnaeus, 1758
Redshank
696
Tringa erythropus Pallas, 1764
Spotted Redshank
66
Tringa nebularia Gunnerus, 1767
Greenshank
68
Tringa stagnatilis Bechstein, 1 803
Marsh Sandpiper
34
Limosa limosa Linnaeus, 1758
Black-tailed Godwit
3
Limosa lapponica Linnaeus, 1758
Bar-tailed Godwit
2
Numenius arquata Linnaeus, 1758
Curlew
340
Numenius phaeopus Linnaeus, 1758
Whimbrel
1
Gallinago gallinago Linnaeus, 1758
Snipe
110
Philomachus pugnax Linnaeus, 1758
Ruff
13
LAR1DAE
Chroicocephalus ridibundus
Linnaeus, 1776
Black-headed Gull
11981
Chroicocephalus genei Breme, 1839
Slender-billed Gull
804
Larus melanocephalus Temminck, 1820
Mediterranean Gull
1035
Larus argentatus Pontoppidan, 1763
Herring Gull
31
Larus michahellis Naumann, 1840
Yellow-legged Gull
1398
Larus audouinii Payraudeau, 1826
Audouin's Gull
87
Larus fuscus Linnaeus, 1758
Lesser Black-backed Gull
2374
Larus sp.
Gull sp.
9
STERN ID AE
Sterna sandvicensis Latham, 1787
Sandwich Tem
362
Hydroprogne caspia Pallas, 1770
Caspian Tem
40
Sterna bengalensis Lesson, 1 82 1
Lesser Crested Tem
1
Chlidonias hybridus Pallas, 1811
Whiskered Tem
43
Table 3 (continued). Number of waterbird species and individuals counted in January 2012, Libya.
260
K.S. Etayeb et alii
Family RECURVIROSTRIDAE
At eight Libyan coastal wetlands, 550 Black-
winged Stilts Himantopus himantopus were counted
(previous maximum 753 in 2011). This species is
mentioned as a passage visitor (Bundy, 1976), but re-
cently has been recorded as a breeder at A1 Mallahah
wetland (Etayeb et al., 2013). A total of eight Avo-
cets Recurvirostra avosetta was observed in two sites
in eastern Libya (Al Labadia and Ayn Zayyanah).
The previous annual maximum was 193 in 2006
(EGA-RAC/SPA Waterbird Census Team, 2012).
Family BURHINIDAE
Eurasian Stone Curlew Burhinus oedicnemus
was counted in two sites: Wadi Ka’am dam and
Tajura Coast with totals of 1 0 and 25 individuals
respectively. The total of 35 Stone Curlews is the
highest so far, with the range in the previous years
of 1-12 individuals.
Family CHARADRIIDAE
Five species belonging to this family were obser-
ved along the coastline: Ringed Plover 61 individuals
(previous maximum 101 in 2011), Kentish Plover
339 individuals (previous maximum 1797 in 2007),
Grey Plover 44 individuals (previous maximum 195
in 2006), Golden Plover 430 individuals (previous
maximum 645 in 2006) and Lapwing 263 individuals
(previous maximum 96 in 2011) (Table 3; Smart et
al., 2006; Etayeb et al., 2007; EGA-RAC/SPA
Waterbird Census Team, 2012; Bourass et al., 2013).
Family SCOLOPACIDAE
In different sites along the Libyan coast, partic-
ularly those with shallow water, we counted a total
of 18 species belong the family Scolopacidae. This
family was the largest during this survey (see Table
3). The number of individuals varied from species
to species, and the highest was 1781 for Dunlin
Calidris alpina , while the lowest was a solitary
Whimbrel Numenius phaeopus in Farwah Lagoon.
Moreover, Redshank Tringa totanus numbered 696
(previous maximum 1544 in 2010). Only three
Black-tailed Godwits Limosa limosa (previous
maximum 10 in 2005, 2006) and two Bar-tailed
Godwits Limosa lapponica (previous maximum 1 7
in 2011) were observed at the westernmost part of
Libya (Coast Abu Kammash to Ras Ajdir). How-
ever, other species fluctuated in numbers through
the years from 2005 to 201 1 , and showed a relative
decrease in 2012, in relation to the reduced number
of sites covered.
Family LARIDAE
A total of seven species of gull was observed
(Table 3). In comparison to the previous years, the
number of individuals was very low, for instance
Black-headed Gull Chroicocephalus ridibundus in
2012 numbered 1 1,980 individuals, whereas the pre-
vious maximum was 25,352 in 2008. A total of 87 of
the Near Threatened Audouin's Gull Larus audouinii
(IUCN Red List) was counted in seven sites around
Tripoli and Benghazi. However, this number was the
lowest so far (previous maximum 670 in 2006).
Family STERNIDAE
Four species were observed from this family
(Table 3). The highest number was for the Sandwich
Tern Sterna sandvicensis with a total of 362 indi-
viduals. This number was in the range of the pre-
vious counts (83 in 2007 and 395 in 2010). Although
this species existed in good numbers compared to
the other Sterna species, there is no evidence so far
of breeding in Libya. It is reported as a winter visitor
(Bundy, 1976). Caspian Tern Hydroprogne caspia
and Whiskered Tern C hlidonias hybridus were more
or less in the range of previous counts (Table 3). A
solitary individual of Lesser Crested Tern Sterna
bengalensis was observed in Tajura Coast. This
species is a summer breeder in some sites in eastern
Libya. The population of Lesser Crested Tern can
be seen in good numbers in Libya from late April
till August (Hamza & Azafzaf, 2012).
Family ALCEDINIDAE
10 individuals of Kingfisher A leedo atthis were
observed at different sites along the coastline (pre-
vious maximum 19 in 2005).
Non-waterbird species
Although this census did not target non-water-
bird species, some species were occasionally recor-
Results of the eighth winter waterbird census in Libya in January 2012
261
Family
Scientific name
Common name
Total
ACCIPITRIDAE
Buteo rufinus (Cretzschmar, 1827)
Long-legged Buzzard
2
FALCONIDAE
Falco tinnunculus Linnaeus, 1758
Kestrel
3
STRIGIDAE
Bubo ascalaphus (Savigny, 1809)
Pharaoh Eagle Owl
1
UPUPIDAE
Upupa epops Linnaeus, 1758
Hoopoe
7
ALAUDIDAE
Galerida cristata Linnaeus, 1758
Crested Lark
67
Melanocorypha calandra
(Linnaeus, 1766)
Calandra Lark
13
H1RUNDIN1DAE
Riparia riparia (Linnaeus, 1758)
Sand Martin
3
Hirundo fuligula (Lichtenstein, 1 842)
Rock Martin
20
Hirundo rustica Linnaeus, 1758
Bam Swallow
6
TURD1DAE
Phoenicurus ochruros (Gmelin, 1774)
Black Redstart
3
MOTACILLIDAE
Motacilla alba Linnaeus, 1758
White Wagtail
41
TURD1DAE
Erithacus rubecula (Linnaeus, 1758)
Robin
4
Saxicola torquata (Linnaeus, 1766)
Stonechat
26
SYLVIIDAE
Sylvia melanocephala (Gmelin, 1789)
Sardinian Warbler
4
Acrocephalus scirpaceus
(Hermann, 1804)
Reed Warbler
5
Phylloscopus collybita (Vieillot, 1817)
Chiffchaff
18
LANIIDAE
Lanius excubitor Linnaeus, 1758
Great Grey Shrike
11
TIMAL1IDAE
Turdoides fulvus (Desfontaines, 1789)
Fulvous Babbler
5
CORVIDAE
Corvus corax Linnaeus, 1758
Raven
3
STURNIDAE
Stum us vulgaris Linnaeus, 1758
Starling
1725
PASSERIDAE
Passer domesticus Linnaeus, 1758
House Sparrow
79
FRINGILLIDAE
Carduelis carduelis Linnaeus, 1758
Goldfinch
5
Serinus serinus Linnaeus, 1766
Serin
3
Table 4. Number of non-waterbird species and individuals counted in January 2012, Libya.
ded in and around wetlands. A total of 2054 indi-
viduals belonging to 23 species from 16 families
was observed during this survey (Table 4). How-
ever, these numbers were the lowest among the pre-
vious years (2005-2011, see Table 2).
ACKNOWLEDGMENTS
We sincerely acknowledge Mr. Shawki Moamer
for hosting the team in Zwara. Our special thanks
to Mr. Mike Smart for reviewing and proofreading
the manuscript.
REFERENCES
Azafzaf H., Baccetti N., Defos du Rau P., Dlensi H.,
Essghaier M.R, Etayeb K., Hamza A. & Smart M.,
2005. Report on an Ornithological Survey in Libya
from 3 to 17 January 2005. Cyclostyled report to
Regional Activities Centre/Special Protected Areas
262
K.S. Etayeb et alii
(MAP/UNEP), Tunis, Environment General Author-
ity, Libya, and African-Eurasian Waterbird Agree-
ment (UNEP/AEWA).
Azafzaf H., Baccetti N., Defos du Rau P., Dlensi EL,
Essghaier M.F., Etayeb K., Hamza A. & Smart M.,
2006. Report on an Ornithological Survey in Libya
from 19 to 3 1 January 2006. Cyclostyled report to the
Regional Activity Centre/Special Protected Areas
(MAP/UNEP), Environment General Agency, Libya
and to the African-Eurasian Waterbird Agreement
(UNEP/AEWA).
Bourass E., Baccetti N., Bashimam W., Berbash A.,
Bouzainen M., De Faveri A., Galidan A., Saied A.M.,
Yahia J. & Zenatello M., 2013. Results of the seventh
winter waterbird census in Libya, January-February
2011. Bulletin of the African Bird Club, 20: 20-26.
Bundy G., 1976. The Birds of Libya: An Annotated
Check-list. BOU Check-list No. 1. London, UK:
British Ornithologists’ Union.
EGA-RAC/SPA Waterbird Census Team. 2012. Atlas of
Wintering Waterbirds of Libya, 2005-2010. Tunis:
Imprimerie COTIM. Temporarily available at:
http ://ww w. isprambiente . gov. it/files/pubblicazioni/
pubblicazionidipregio/Atlas_of_wintering_waterbird
in_Libya_20052010.pdf.
Etayeb K.S. & Essghaier M.F.A., 2007. Breeding of
marine birds on Farwa Island, western Libya.
Ostrich, 78: 419-421.
Etayeb K., Essghaier M.F., Hamza A., Smart M., Azafzaf
H., Defos du Rau P. & Dlensi H., 2007. Report on an
Ornithological Survey in Libya from 3 to 15 February
2007. Cyclostyled report to the Regional Activities
Centre/ Special Protected Areas (MAP /UNEP) and
Environment General Authority, Libya.
Etayeb K.S., Yahia J., Berbash A. & Essghaier M.F.A.,
2013. Ornithological importance of Mallaha wetland
in Tripoli, Libya. Bulletin de la Societe zoologique
de France, 138: 201-211.
Hamza A. & Azafzaf H., 2012. The Lesser Crested Tern,
Sterna bengalensis, State of knowledge and conser-
vation in the Mediterranean Small Islands. Initiative
PIM, 20 pp.
Hamza A., Saied A., Bourass E.M., Yahya J., Smart M.,
Baccetti N., Defos du Rau P., Dlensi H. & Azafzaf
H., 2008. Report on a fourth winter ornithological
survey in Libya, 20-31 January 2008. Cyclostyled
report to the Regional Activities Centre/ Special
Protected Areas (MAP/UNEP) and Environment
General Authority, Libya.
Isenmann P., Gaultier T., El-Hili A., Azafzaf H., Dlensi
H. & Smart M., 2005. Oiseaux de Tunisie. Societe
d’ Etudes Ornithologiques de France, Museum Na-
tional d’Histoire Naturelle, Paris, France.
Loh J., Green R.E., Ricketts T., Lamoreux J., Jenkins M.,
Kapos V. & Randers J. 2005. The living planet index:
using species population time series to track trends
in biodiversity. Philosophical Transactions of the
Royal Society B, 360: 289-295.
Smart M., Essghaier M.F., Etayeb K., Hamza A., Azafzaf
H., Baccetti N. & Defos Du Rau R, 2006. Wetlands
and wintering waterbirds in Libya, January 2005 and
2006. © Wildfowl & Wetlands Trust, 56: 172-191.
Svensson L., Mullamey K., Zetterstrom D. & Grant P.J.,
2010. Collins Bird Guide. 2nd Revised edition.
Harper Collins Publishers, United Kingdom, 448 pp.
Biodiversity Journal, 2015, 6 (1): 263-270
New knowledge on diet and monitoring of a roost of the long-
eared owl, Asio otus (Linnaeus, 1 758) (Strigiformes Strigidae)
on Mount Etna, Sicily
Agatino Maurizio Siracusa*, Elisa Musumeci, Vera D’Urso & Giorgio Sabella
'Depart ment of Biological, Geological and Environmental Sciences - Section of Animal Biology, University of Catania, via Androne
8 1, 95 1 24 Catania, Italy
Corresponding author, e-mail: ant sir a@unict.it
ABSTRACT A study during autumn and winter in Monte Serra area (Mount Etna) was performed on the
pellets of a roost of long-eared owl, Asio OtUS (Linnaeus, 1 75 8) (Strigiformes Strigidae).
Besides, in order to better understand the feeding habits of this species on Mount Etna, the
data from Monte Serra were integrated with those from Linguaglossa Pin eta (breeding period).
The study was performed through the analysis of 1,724 preys. The species most preyed was
the Mammalia Microtidae MiCTOtllS SCIVii (de Selys-Longchamps, 1838). The average weight
of the preys was 23.48 g, while the average meal was 36.63 g. Besides, the results of the yearly
monitoring of the roost studied are given.
KEY WORDS Asio otus ■ trophic niche; roost; Sicily.
Received 03.11.2014; accepted 0 9.02.2 0 15; printed 30.03.2015
Proceedings of the 2nd Interna tional Congress “Speciation and Taxonomy”, May 1 6 th - 1 8 th 2014, Cefalu-Castelbuono (Italy)
INTRODUCTION
The trop hie n ic he of Asio OtUS (Linnaeus, 1758)
(S trigiform es S trig id ae) during a u turn n and w inter in
a site of Mount Etna was studied in order to better
understand that niche during all the year. The literat-
ure data concern in g Sic ily consistonly on the in form -
ation by Siracusa et al. (1 996) focalized on the diet
during the reproductive period in two localities (Lin-
guaglossa and Roccapalumba). Up to now, no inform-
ation on the roost monitoring in Sicily are known.
MATERIAL AND METHODS
The studied site lies on the “Monte Serra”, one
of the volcanic cones of Mount Etna, at an altitude
of 450 m a.s.l., which originated on the side south-
east during 122 B .C . It has a characteristic shape
of a horseshoe, as a result of the collapse of the
summit of the crater and of the volcano slope.
In recent centuries the landscape, due to human
settlement and agricultural activities, has been
progressively modified and the natural vegetation
was represented just by some residual strips
unevenly distributed. After the abandonment of
cultivation, has started a new and slow recolonisa-
tion of the Mediterranean natural vegetation. This
vegetation consists mainly in the bushes of ever-
green holm oak, QuCVCUS HeXL., wild olive, OleO.
europaea l ., and carob, Ceratonia siliqua l .
The slopes of Monte Serra are covered with a
shrubby in which are present the common broom,
Spartium junceum l., and the Etna broom, Genista
aetnensis (Raf. ex Biv.) DC, as the predominant
sp ec ie s (Fig. 1 ) .
264
Agatino Maurizio Siracusa etalii
At the base of the mountain, lies a forest left in
its natural state, the "Forest of Cyclamen" in which
the essence most represented is the tree oak,
Quercus virgilicinci Ten o re , followed by hornbeam,
Ostrya carpinifolia Scopoli, and flowering ash,
Fraxinus ornus l .
The climatic characteristics of the study area
are derived on the data of the time series of the
Viagrande term oplu viom etric station (Fig. 2),
which is located on the slope mostaffected by rain-
fall (annual rainfall higher than the whole of Etna),
because invested by the moisture deriving from
the Ionian Sea, which it overlooks. The study area
falls within the Mediterranean B io g e o g r ap h ic a 1
Region, in the range of the m eso-M editerranean
climate (Brullo et al., 1 996). The data on diet of
long-eared owl in autumn and win ter were obtained
from the analysis of pellets collected in a roost
located at “Parco di Monte Serra”, in the period
between September 2012 and March 2013.
The pellets were weekly collected. The collec-
ted material was provided on-site of a label re-
porting a detailed tagging, as well as the date and
time of collection, the GPS coordinates of indivi-
dual roost sites, weather conditions, information
on the presence/absence and number of specimens
observed.
The pellets were dried in the open air for a few
days, wrapped in polythene bags containing cam-
phor (to prevent any damage caused by the attack
Figure 1 . Land use in area of “Monte Serra" and neighboring areas (from Angelinietal., 2009, modified). Legend according
to Corine Biotopes Code: 31.81(brown) Middle-European scrubs; 32.215 (orange) low scrub w ith CalicOtOHie sp.; 34.8 1 (light
brown) M editerranean subnitrophilous meadows; 41 .732 (green) peninsular and insular Italy oak deciduous woods; 45.3 1A
(light green) Southern Italy and Sicilian holm oak woods; 82.1 (light yellow) arable intensive and continuous; 82.3 (yellow)
cultures of extensive type and complex agricultural systems; 83.11 (azure) groves; 83.21 (blue) vineyards; 83.3 22 (dark
green) plantations of EuCCllyptllS ; 83.16 (ocher brown) citrus orchards; 86.1 (gray) towns.
New knowledge on diet and monitoring of a roost of the long-eared owl, Asio otus on Mount Etna, Sicily
265
of scavenging arthropods) and then transported to
the laboratory to be analyzed.
The content of each intact pellet was noted
separately and the number of prey items concerned
was taken to equal to the greatest number of identi-
fied fragments of one species (greatest number of
low er jaw s, etc .).
For the study of the pellets was used the pellets
analysis technic (Contoli, 1 9 80). The pellets were
opened by dry technique, with the aid of a tweezers
and of an entomological brush. For those too com-
pact it was preferred the immersion in hot water
for a few minutes in order to more easily separate
th e bones.
Before opening, were taken measures, with a
digital gauge, relating to the length and the width
of pellets. For the sorting of the content was used a
stereoscopic microscope for better visibility of the
alveoli of the molars and of the bones of small
mammals.
To count the preyed specimens is considered
their minimum number (Chaline et al., 1 974). The
jaws of the rats and the synsacrum of birds collec-
ted were measured with a digital caliper, in order to
estimate the weight of the preyed specimens.
The identification of small mammals was based
on the cranium features and dichotomous keys
(Toschi & Lanza, 1 959; Toschi, 1 965; Chaline et
al., 1 974; Amori et al., 2008), while for the larger
prey was used the morphology of the long bones.
The calculation of the biomass was carried out
by assigning to each species an average weight,
relative to the species of small mammals and Cole-
optera in Sicily (Table 1), provided by Di Palma &
Massa, 198 1, and for savi’s pine vole ( MicwtliS
Savii) provided by Catalisano & Massa, 1987.
Using the equation of Di Palma & Massa
(198 1), the calculated average weight of the brown
rat ( RattUS llOrVegicUS B erkenhout, 1 769) results
102.6 j_ 2 7 .0 (SD) g (number of sampled specimens
n = 46), and 94.5 j_26.4 (SD) g (n = 14) regarding
RattUS sp. The weight of 94.5 g, equivalent to that
of RattUS sp., has been assigned also to RattUS VttttUS
(Linnaeus, 1 75 8 ), considering that only 2 speci-
mens were collected and no measurable jaws were
av ailab le .
Using the equation of Di Palma & Massa
(1981), the calculated average weight of the birds
results 14.1 j_ 5 . 2 (SD) gr (n = 187) based on
synsacrum found in the pellets.
Figure 2. Climogramma of Viagrande term o p lu v io m e trie
station (from Zampino et al.. 1 997).
Th*
Weight |g)
References
Aftw domeltfctt*
12.3
IM Palma & Massa. 19*1
Apnkmiit ,n/i«rfrni
2o.h
JJi Palma & Massa. 19*1
flfjrtfr.i
1026
RjuatlonofDi Palma 6! Massa, 19* t
Mil) bp
94.5
Equation of Di Palma & Mava. 19* L
Raft its rtUltis
94.5
Same weight of Ruimx bp.
Mit'rtHm
20
Lalalisani) & Massa, 19*7
C rfH'idtirn j icutu
6.7
Ui Palma Jit Massa. 1 9*1
t’oleopter*
0.)
Di Palma & Massa, 19*1
C hiraplera
10
l/tyrilre/fos sp. Il average wcighl of sampled species
Avti
14,1
EHiiaiion of Di Ftiliru £ Massa, I9SI ( ss n sacrum I
del.
34,1
As erairo wclghl uf all allegories uf pres
1 irscEud iim fttillns mu-region and L'l'ks'plcrj I
Table 1. Values used for the calculation of the biomass.
Besides the study of the trophic niche based on
the collected pellets, the monitoring of the presence
of the owls in the roost from May 2012 to June
2013 was performed (Fig. 3).
RESULTS AND DISCUSSION
A total of entire 875 pellets were collected and
examined in the Monte Serra area. The pellets have
an average lenght of 32.2 +_ 7 .9 (SD) mm and an
average width of 18.8 j^3.2 (SD) mm. A total of
1,421 prey have been identified (Table 2, Fig. 4),
with an average of 1.62 prey/pellet; 1 or 2 prey/
pellet were found in most cases, 3 sometimes, 4
266
Agatino Maurizio Siracusa etalii
occasionally. During the study period, mammals are
the most represented group (about 67% of the prey)
followed by birds (about 32%). MicWtUS SCIVU
represents the most preyed species (50.2% of the
catch), while Apodemus sylvaticus (L innaeus,
Figure 3. Specimen of Asio OtllS o n QuCTCUS sp .
(October 2012, photo by E. Musumeci).
Monte Serra
Category of prey
n
% weight {g>
biomass (g)
%
Microtus savii
714
50.25
20
14280
41,88
Microtidae
714
50.25
14280
41.08
Mus domesticus
19
1.34
12.2
231.8
0.68
Apodemus sylvaticus
100
7.60
20.8
2246.4
6.59
Rattus rattus
2
0.14
94,5
189
0.55
Rattus norvegicus
75
5.28
102,6
7695
22,57
Rattus sp.
26
1,83
94.5
2457
721
Muridae
230
16.19
12819.2
37.6
Rodentia
944
66.43
27099.2
79.48
Croodura sicula
1
0.07
6.7
6.7
0.02
Soricidae
1
0,07
6.7
0.02
Soricomorpha
1
0.07
$.7
0.02
Chiroptera
7
0.49
10
70
0.21
MAMMALIA
952
67
2717S.9
79.66
AVES
452
31.81
14.1
6373.2
18.69
INSECTA Coleoptera
1
0.07
0.1
0.1
0
Mot det.
16
1.13
34.1
545.6
1.6
Total prey
1421
Pellets
875
Prey/peltets
1.62
Total biomass (g>
34094,8
Average weight prey (g) 23.99
Average meal (g) 38 .86
Table 2. Results of the pellets analysis in the study area.
1758), generally the main trophic resource in wooded
areas, shows the frequency of only 7.6% . A 1 though
Monte Serra is a suburban park in a discretely
anthropized area, MllS domeSticUS Linnaeus, 1758
is very little represented (only 1.3%), but about
7.2% of prey (about 30% of biomass) belongs to
RattUS spp. (Table 2): this latter result could be
justified with the energetic advantage obtained by
long-eared owl feed on rats, because these have a
greater weight than other prey and owls could save
energy by reducing the hunting with an equal gain
of biomass. Soricom orpha and Chiroptera, as well
as Insecta, are present in very low percentages of
prey, less than or equal to 0.5%. The only found
specimen of Soricidae, CwcidlirCl siciilci Miller,
1 900, could be due to a selective choice of prey as
well as environmental factors, like the pressure of
the human presence in the area of Monte Serra. It
should be emphasized that, although it is uncom-
mon the predation of birds, in the examined site the
percentage of this prey is significant and this is in
agreement with some studies conducted on winter-
ing sites in Northern Italy and in Spain (Albufera
de Valencia) which recorded a presence of birds
even higher, 50% of the total num her of individuals
preyed (M as trorilli, 2000; Escala et al., 2009). The
discrepancy between these results is likely attrib-
utable to the opportunistic habits of the long-eared
owl that, when possible, implements group hunting
strategies able to ferret out and in some cases cut
off entire dormitories of passerines (Mikkola,
1 983).
The data of the present study were compared
with those of the long-eared owl diet during the
reproductive period, detected always by the pellets
analysis technic, in two Sicilian sites, Pineta di Lin-
guaglossa and R o cc ap alu m b a, characterized by dif-
ferent environmental features (Siracusa et al., 1996):
an old pinewood and a cultivated area respectively.
A total of 191 pellets were collected and ex-
amined in Linguaglossa Pineta station (Table 3).
Mammals are the most represented group (about
94% of the prey) followed by birds (about 6%).
ApodemUS sylvaticus was the most preyed species
(60% of the catch) and with MicrOtUS Savii ( 32.67%
of prey) represent about 93% of the preys. MuS
domesticus is very little represented (only 0.66%),
while no species of RattUS were collected. Sorico-
rnorpha and Chiroptera were present in very low
percentages equal to 0.33% (Siracusa et al., 1996).
New knowledge on diet and monitoring of a roost of the long-eared owl, Asio otus on Mount Etna, Sicily
267
m.js*
Coteopiera
0.07%
Noi del. 1.13%
Chiioptera
1 .49%
Crocitfitm
sicuta
Mus domesticus 1 . 34® »
Roll ns sp,
I 83%
Rama norwgicus
5.28%
0.14%
Figure 4. Graphical results of the pellets analysis
in the study area.
A total of 21 pellets were collected and examin-
ated in Roccapalumba station (Table 3). Mammals
are the most represented group (about 93% of the
prey) followed by birds (more than 3%), arthropods
(more than 2%) and amphibians and reptiles (about
1 . 5 %). Microtus savii was the most preyed species
(89.42% of the catches), while the other species of
mamma is (Apodemus sylvaticus, Mus domestic us,
RattUS rattus and Crocidura sicula ) were present
in very low percentages, less than 2% (Siracusa et
al., 1996).
In order to identify the trophic niche of the
species in the piedmont areas of the Etna eastern
slope, the stations of M onte Serra and Linguaglossa
(a pinewood), were considered as a single sample
MONTE SERRA
LINGUAGLOSSA
ROCCAPALUMBA
Category of
prey
n
%
biomass
(g)
%
n
%
biomass
(g)
%
n
%
biomass
(9 1
%
Microtus savii
714
50.25
14280
41.88
99
32,67
1683
26.34
245
89,42
4165
89.93
Mus
domesticus
19
1,34
231,8
0 68
2
0.66
25
0.39
3
1.09
37.50
0.81
Apodemus
sylvaticus
108
7.60
2246.4
6 59
182
60,07
4277
66.94
1
0.36
23.50
0.51
Rattus rattus
2
0.14
189
0.55
-
-
-
-
1
0.36
118
2.55
Rattus
norvegicus
75
5.28
7695
22.57
Rattus sp.
26
1,83
2457
7.21
-
-
-
-
-
-
-
Crocidura sicula
1
0.07
6.7
0.02
1
0.33
6.5
0.10
5
1.82
32.50
0.70
Chlroptera
7
0,49
70
0.21
1
0.33
20
0.31
-
-
-
-
MAMMALIA
952
67
27175.9
79.66
285
94.06
6011.5
94,08
255
93.06
4376.50
94.50
AVES
452
31.81
$373.2
18.69
18
5.94
378
5.92
9
3.28
189
4.08
AMPHIBIA +
REFT ILIA
-
-
-
*
4
1.46
60
1.30
ARTHROPOD A
1
0.07
0.1
0
*
ifli
-
6
2.19
6
0.13
Not del
16
1.13
545.6
1.6
-
-
-
-
*
-
-
-
Total
875
303
274
LINGUAGLOSSA
ROCCAPALUMBA
Total prey
303 (243 on entire pellets}
Total prey
274 (66 on entire pellets )
Pellets
191
Pellets
21
Prey/peilets
1,27*
Prey/pellets
3.14*
Total biomass (g)
6389.50**
Total biomass (g)
4631.5 **
Average weight prey (g)
21.09**
Average weight prey (g)
16.9**
Average meal (g)
26.78*
Average meal (g)
53.07*
Table 3. Comparison of results of pellets analysis during w inter period (M t. Serra) and during breeding period of long-eared
owl from Pineta di Linguaglossa and from Roccapalumba (* calculated only on prey on en tire pellets; ** calculated on total
prey ) (from Siracusa et al.. 1 996, modified).
268
Agatino Maurizio Siracusa etalii
(Table 4), although the first case concerns the diet
in the autumn and winter, while the second case
regards the trophic niche in the reproductive period.
It must be emphasized that the two sites, with
different vegetations, are located both in the foothill
region of Etna Mountain.
A total of 1,066 pellets were processed (Table
4). Also in this case, mammals are the most rep-
resented group (about 72% of the prey) followed
by birds (about 27%), while arthropods are almost
absent. MicWtUS SClvii is the most preyed species
(more than 47% of the catches), while the
Muridae provide the greatestcontribution in terms
of biomass (42.30% of total). Soricomorpha and
Chiroptera are present in very low percentages
less than or equal to 0.46%. The average weight
of the preys is 23.48 g, while the average meal is
36.63 g.
The roost of Mt Serra was observed by the end
of May 2012, when a young specimen has been
TOTAL MS+L
Category of prey
n
%
biomass fg)
%
Microtus savii
813
47.16
15963
39.43
Microtidae
813
47.16
15963
39.43
Mus domesticus
21
1.22
256 8
0.63
Apodemus sylvaticus
290
16.82
6523,4
16,11
Rattus rattus
2
0.12
189
0.47
Rattus norvegicus
75
4.35
7695
19
Rattus sp.
26
1.51
2457
6.07
Muridae
414
24.01
17121.2
42.30
Roden tia
1227
71.17
33084.2
81.72
Crocidura sicula
2
0.12
13.2
0.03
Sortcidae
2
0.12
13.2
0,03
Soricomorpha
2
0.12
13.2
0.03
Chiroptera
8
0.48
90
0.22
MAMMALIA
1237
71.75
33187.4
81.97
AVES
470
27.26
6751.2
16.68
INSECT A Coleoptera
1
10.06
0.1
0
Notdet.
16
0.93
545.6
1.35
Total prey 1724 {1664 on entire pellets)
Pellets 1066
Preylpellets 1.56*
Total biomass {g} 40484. 30**
Average weight prey (g) 23.48“
Average meal (g) 36.63*
Table 4. Sum of results of the pellets analysis of long-eared
owl during breeding period in Pineta di Linguaglossa and
during autumn in Monte Serra. * calculated only on prey on
entire pellets (n=1664); * * calculated on total prey (n = 1 7 24).
sighted among GcflistCl, at the end of June 2013
(Fig. 5). Although traces of their presence (as
plumage and very few pellets) were evident from
June to August, only in early September 2012 were
observed 7 specimens on an oak near the structure
used by the Butterfly House as Information Point.
This same roost was used by the group for most of
the autumn season. During the sightings were coun-
ted from 1 to a maximum of 7 specimens, with
greater presence during the afternoon hours. The
owls we re quite confident and they tolerated human
presence. In November sporadic observations of
owls were recorded and no pellets were found. The
causes of this absence could, at least partially, be
attributed to human disturbance or it might have
been a time of reorganization of the roost. In late
November (29th), after many days of absence, a
roost of mo re than 11 specimens occupied the pines
located inside the playground for children of the
butterfly house. This roost, that throughout the
winter period was composed of about 20 speci-
mens, was present until the end of February. From
late February to mid-March, were observed no
more than 7 specimens as to restore the situation of
Septem ber-N ovem ber. No specimens were spotted
from middle M arch to the end of June.
This study has allowed us to integrate know-
ledge about the trophic niche of the long-eared owl
in Sicily for which was known a single study that
refers to the diet of this species in the breeding
season; however, were not known data concerning
the trophic niche during the autumn-winter period
and concerning the roost in wintering period. The
data obtained from the pellets analysis of about 20
specimens and the analysis of the characteristics of
the study area, have confirmed the selective beha-
vior o f Asio otus in the choice of prey, specifically
the Microtids (as shown by the high percentage of
MicWtUS SQVii found). It also highlighted a certain
plasticity of the species that, if necessary, takes
advantage of favorable situations such as the
presence of dormitories of birds that are flushed out
with a technique of group hunting. The above
explains the significant number of birds found in
the pellets, which is not a data usually reported in
bibliography. This study is also useful for the
increase ofknowledge on the wintering sites of the
long-eared owl in Italy and can be inserted in the
national register of the roosts, set up by the project:
"Gufiamo: count the long-eared owls wintering in
New knowledge on diet and monitoring of a roost of the long-eared owl, Asio otus on Mount Etna, Sicily
269
Figure 5. Histogram date/number of long-eared owl specimens observed.
Italy", started a few years ago from the collabora-
tion of GIC & EBN Italy, with the Global owl
project that provides for the establishment of a net-
work containing data on the presence of the roosts
in Italy. From the analysis of the monitoring of the
roost of Monte Serra, one might assume the
presence oftwo differentpopulations.The first one,
in the Park throughout the year, although with
changes in the choice of the roost and possibly with
nesting site located not many miles away.
The other population, more numerous, would
take advantage of the Monte Serra Park as a
wintering site. Specimens of this second population
may be resident in the territory of Etna and make
seasonal vertical migrations or could be migratory
specimens that stop to winter. The hypothesis of
two different populations is supported by the owl
attitude observed in the days of collecting pellets.
Whenever the collector approached at the roost,
systematically, part of specimens are alerted and
receding in flight; however, remained always 5-7
specimens, very confident, as if they were already
accustomed to the environment and the presence of
visitors to the park. Because the site is regularly
occupied, this would allow regular long-term mon-
itoring of the roost; furthermore, the use of mo-
lecular studies of feathers collected throughout the
year could also clarify the phenology of the species
in Sicily, whose presence as a nesting species has
been established only recently.
ACKNOWLEDGEMENTS
We thank Pierangela Angelini (ISPRA) for al-
lowing the use of the shape file of Carta N atura
1:50.000 of Sicily for the characterization of the
study area.
REFERENCES
Angelini P., Bianco P., Cardillo A., Francescato C. &
Oriolo G ., 2009. G li habitat in Carta della N atura -
Schede descrittive degli habitat per la cartografia
1:50.000. Manuali e linee guida 49/2009 ISPRA.
Amori G., Contoli L. & N appi A ., 2008. Mammalia II:
Erinaceomorpha, Soricomorpha, Lagomorpha, R o -
dentia. Fauna d 'Italia, 44. C alder ini, Bologna, 754 pp.
Brullo S., ScelsiF., Siracusa G. & Spampinato G., 1996.
C ara tte ris tic h e b io c lim atic h e della Sicilia. Giornale
Botanico Italiano, 1 30: 1 77- 1 8 5.
Catalisano A. & Massa B ., 1 98 7. Considerations on the
structure of the diet of the Barn Owl ( TytO albtt) in
Sicily ( Italy). Bollettino di Zoologia, 54: 69-73.
Chaline J., B audvin H., JammotD. & Saint Giron M.C.,
1974. Les proies des rapaces. Doin ed., Paris, 141
pp.
Contoli L., 1 980. Borre di Strigiformi e ricerca teriolo-
gica in Italia. Natura e Montagna, 3: 73-94.
Di Palma A.M.G. & Massa B., 1981. Contributo m e to -
dologico perlo studio dell’alimentazione deiRapaci.
In: AttidellConvegno italiano diOrnitologia,Aulla:
69-76.
270
Agatino Maurizio Siracusa etalii
Escala C., Alonso D., Mazuelas D., Mendiburu A.,
Vilches A. & Arizaga J., 2009. W inter diet of Long-
eared Owls AsiO OtllS in the Ebro valley (NE Iberia).
Revista Catalana d ’ O rn ito lo g ia , 25: 49-53.
Mastrorilli M., 2000. The importance of birds in the
winter diet of Long-eared Owls ( AsiO OtllS ) in the
Bergamo district in (Lombardy, northern Italy).
Proceedings International Symposium "Ecology and
Conservation of European Wood Owls", Harz: 45.
Mikkola H., 1983. Owls of Europe. Poyser ed., 397 pp.
Siracusa M ., Sara M ., La Mantia T. & Cairone A., 1996.
Alimentazione del Gufo com une ( Asio OtllS) in
Sicilia. II Naturalista siciliano, 20: 3 1 3-320.
Toschi A., 1 965. Mammalia. Lagomorpha, Rodentia,
Carnivora, Ungulata, Cetacea. Fauna d' Italia, 7.
Calderini, Bologna, 647 pp.
Toschi A. & Lanza B ., 1 959. Mammalia. Generality,
Insectivora, Chiroptera. Fauna d'ltalia, 4. Calderini,
Bologna, 485 pp.
Zampino D ., Duro A., Piccione V. & Scalia C ., 1 997.
Fitoclima della Sicilia. Termoudogrammi secondo
Walter e Lieth delle stazioni te rm o p 1 u v io m e trie h e
della Sicilia occidentale. In: Guerrini A. (Ed.), Atti
del 6° Workshop del Progetto Strategico C. N. R.
"Clima Ambiente e Territorio del Mezzogiorno"
(Taormina, 13-15 Dicembre 1995), I: 229-292.
Biodiversity Journal, 2015, 6 (1): 271-284
Monograph
Morphological differences between two subspecies of Spotted
Flycatcher Muscicapa striata (Pallas, 1 764) (Passeriformes
Muscicapidae)
Michele Vigano 1 & Andrea Corso 2
'MISC- Via Ongetta, 5-21010 Germignaga, Varese, Italy; e-mail: mikivigano@yahoo.com
2 MISC- Via Camastra, 10- 96100 Siracusa, Italy.
* Corresponding author
ABSTRACT Four subspecies of Spotted Flycatcher {Muscicapa striata Pallas, 1764) (Passeriformes Mus-
cicapidae) are usually recognized within the Western Palaearctic. We carefully analysed two
of these in order to determine and quantify their morphological differences: M. striata striata
(inhabiting most of continental Europe east to the Ural mountains and a small portion of
north-western Africa) and M. striata tyrrhenica Schiebel, 1910 (breeding on the Tyrrhenian
islands of Corsica, Sardinia and the Tuscan Archipelago). We examined total of 58 Spotted
Flycatcher specimens from Italian museums (of which 18 M. striata tyrrhenica ) and obtained
data about morphological features such as wing point, length and formula, and bill length,
width and depth; furthermore, we investigated plumage colour using a spectrometer. Biomet-
ric measurements and an analysis of plumage streaking confirmed the presence of important
differences between the two taxa; the colorimetric analysis did not produce the expected res-
ults, although it had some interesting implications concerning the preservation of museum
specimens and their use in studies of plumage colour.
KEY WORDS Spotted Flycatcher; Muscicapa striata tyrrhenica', morphology; museum specimens.
Received 26.01.2015; accepted 01.03.2015; printed 30.03.2015
Proceedings of the 2nd International Congress “Speciation and Taxonomy”, May 1 6th- 1 8th 20 1 4, Cefalu-Castelbuono (Italy)
INTRODUCTION
The Spotted Flycatcher {Muscicapa striata Pal-
las, 1764) is a songbird in the family Muscicapidae
and is the only member of its genus in Europe, with
at least twenty more species in Asia and Africa. The
Spotted Flycatcher is found through most of the
Palaearctic, with a continuous distribution from the
Iberian peninsula to the Mongolia-China border. It
is a long-distance, trans-Saharan migrant, and most
of the population winters south of the Equator
(Cramp & Perrins, 1993).
Seven subspecies are currently recognized in
this extensive range (del Hoyo et al., 2006): M.
striata striata (Pallas, 1764) (Figs. 1, 2) breeds in
Europe east to the Ural mountains and in north-
western Africa, and winters south of the Sahara;
M. striata balearica von Jordans, 1913 (Fig. 1),
breeds in the Balearic islands and winters in west-
ern and south-western Africa; M. striata tyrrhe-
nica Schiebel, 1910 (Figs. 1, 2), breeds in Corsica
and Sardinia and presumably winters in Africa; M.
striata inexpectata Dementiev, 1932, breeds in
Crimea and winters in Africa; M. striata neumanni
Poche, 1904, breeds in the islands of the Aegean
Sea east to the Caucasus and northern Iran and
south to Cyprus and the Fevant, in addition to cent-
ral Siberia, and winters in eastern and southern
Africa; M. striata sarudnyi Snigirewski, 1928,
breeds from eastern Iran to northern and western
272
Michele Vigano & Andrea Corso
Pakistan and presumably winters in southern and
eastern Africa; M. striata mongola Portenko,
1955, breeds from the south-eastern Altai moun-
tains to northern Mongolia, and presumably win-
ters in southern and eastern Africa.
Only two (M. striata striata and M. striata
tyrrhenica) of these seven subspecies are regularly
found in Italy, while M. striata neumanni, which
could potentially occur in migration, has not yet
been confirmed (Corso, 2005; Brichetti & Fracasso,
2008). The nominate subspecies breeds throughout
continental Italy and Sicily, where it is considered
common and widespread, although its distribution
is somewhat patchy with gaps in high mountain
areas. The core breeding range of M. striata tyrrhe-
nica comprises Corsica and Sardinia, but contra del
Hoyo et al. (2006) and Cramp & Perrins (1993), it
also breeds in the Tuscan Archipelago (Brichetti &
Fracasso, 2008), while its presence along a narrow
band of the Tyrrhenian coast remains to be con-
firmed (Brichetti & Fracasso, 2008; Tellini et al.,
1997). The authors provide some interesting in-
formation on the abundance of Spotted Flycatcher
subspecies in Italy.
Although the nominate subspecies breeds al-
most throughout continental Italy, it is never abund-
ant, with population densities that rarely exceed 0.2
pairs/hectare. On the other hand, as many as 0.6
pairs/hectare have been found in M. striata tyrrhe-
nica (VV. AA. in Thibault & Bonaccorsi, 1999); so
the species seems to fit the usual pattern on islands
of densisty inflation due to lower species richness
(eg MacArthur & Wilson, 1967; George, 1987;
Blondel et al., 1988). Interestingly, high population
densities have been recorded along the Tyrrhenian
coast in Tuscany; densities are far lower only 50 km
inland (Tellini et al., 1997).
There are currently no reliable data on the win-
tering range of M. striata tyrrhenica (Cramp &
Perrins, 1993, del Hoyo et al., 2006). The
M. striata tyrrhenica subspecies of the Spotted
Flycatcher was described for the first time by
Schiebel (1910) in a paper on the Corsican avi-
fauna and a syntype taken in Aitone, Corsica, on
19 May 1910 is currently held at the Zoologisches
Forschungs institut und Museum Alexander
Koenig in Bonn, Germany. The identification of
this subspecies is generally dealt with very super-
ficially in the ornithological literature, with limited
discussion of its distinguishing characteristics.
Several examples are below:
- Arrigoni degli Oddi (1929): “ similar to the pre-
vious species [ authors ’ note: the subspecies
Figure 1. In Western Europe three subspecies of Spotted Flycatcher are found: Muscicapa striata striata (blue), M. striata
tyrrhenica (red) and M. striata balearica (green). Figure 2. Two subspecies of Spotted Flycatcher breed in Italy: M. striata
striata (blue) and M. striata tyrrhenica (red); M. striata tyrrhenica could be present in the yellow area too, but further
research is needed.
Morphological differences between two subspecies of Muscicapa striata (Passeriformes Muscicapidae)
273
striata]; central spots on the cervix and streaking
on breastless distinct ”;
- Cramp & Perrins (1993): “ more warm brown
on upperparts, distinctly less streaked on breast,
streaks replaced by broader, coalescing spots''’',
- Brichetti & Fracasso (2008): “ upperparts
browner and warmer-toned, and streaking on the
underparts less well-defined and tending to merge
into spots ”;
- van Duivendijk (2010): “Primary -projection
slightly shorter that striata; upperparts warmer
brown, underparts almost unstreaked but with
broad, faint spots'”.
Over the last ten years we have carried out in-
depth field and museum studies on the morpholo-
gical differences between the two taxa in question
(Figs. 3-6). In the field, the immediate impression
given by M. striata tyrrhenica is of a paler bird with
wanner tones to the back and more homogeneous
underparts. The breast markings, which are gener-
ally well defined streaks in M. striata striata, appear
faded and more spot-like. The streaking on the nape
is also less well defined compared to the nominate
subspecies, due to the lower contrast between the
streaks and the nape’s background colour. Primary
projection is one of the most important characters:
while the primary projection beyond the tertials is
longer that the tertials themselves in continental
birds, individuals from Corsica and Sardinia have
a primary projection that is shorter than, or at most
equal to the length of the tertials.
This paper mainly reports the results of our
museum studies, while an article on field identific-
ation criteria is forthcoming (Vigano et al., personal
data).
MATERIAL AND METHODS
Our first observations on the morphological dif-
ferences between the two subspecies were made in
the field: M. striata tyrrhenica was studied in
southern Sardinia near Villasimius (Cagliari) in July
2004, August 2005, July 2006, and May 2011 and
on the island of Elba in July 2014. We have studied
this taxon in Corsica as well, in the area of the Gulf
of Calvi, in July 2007, July 2008, and May 2012.
Our studies of M. striata striata have taken place
continuously during the breeding season since 2005
in northern Italy; additionally, we have studied this
taxon during spring migration on various small
islands off central and southern Italy, especially
Ventotene (Latina) in April 2010 and 2011 and
Linosa (Agrigento) in May 2006, April 2007, and
April 2009, where on good days hundreds or even
thousands of individuals can be seen.
Other observations took place opportunistically
elsewhere in the Western Palaearctic, both during
the breeding season and in migration. Studies of
museums skins complemented our field observa-
tions and were of fundamental importance for this
paper (Figs. 7, 8; Table 1). There are very few spe-
cimens of M. striata tyrrhenica in Italian and
foreign museums; indeed, there are none at all in
the largest bird collection in Europe at the Natural
History Museum at Tring, U.K. We arranged for all
of the M. striata tyrrhenica specimens held at the
Museo Civico di Storia Naturale in Milan, Italy
(MCSM), Museo Civico di Zoologia in Rome, Italy
(MCZR), and Museo di Scienze Naturali in Forli,
Italy (MSNF) to be sent on short-term loan to the
Museum of the Institute for Environmental Protec-
tion and Research (Istituto Superiore per la Pro-
tezione e la Ricerca Ambientale - ISPRA) in
Ozzano dell’ Emilia (Bologna, Italy) so that they
could be studied side-by-side along with the speci-
mens held in the last-named institution.
We took the following measurements: wing
chord, longest primary (P3), distance of each
primary from P3, bill length from the nostrils, bill
height and thickness at the nostrils. Measures that
are generally taken during ringing activities such as
tail, tarsus, and bill-to-cranium length were not
taken since they vary depending on the way the spe-
cimen was prepared (Winker, 1998; Eck et al., 2011;
Kuczynski, 2003). The measurements considered
here are also subject to some degree of variation
depending on specimen preparation; measurements
taken on live animals may add a degree of precision
and some additional information, but we felt that
museum specimens were better suited to taking
biometrical and plumage colour data together.
As concerns wing chord length, one study that
looked at the wings of Rooks ( Corvus frugilegus )
measured upon capture, after 8 weeks, and again
after 144 weeks found a difference in length
between fresh and dried wings of about 1.84%
(Knox, 1980). Measurements were taken using a
stopped ruler (to the nearest 0.5 mm), callipers (to
274
Michele Vigano & Andrea Corso
Figures 3, 4. Spotted Flycatcher ( Muscicapa striata tyrrhenica), Villasimius (Cagliari), Sardinia, May 2011. Note the quite
pale and warm general colour, the subtle head and breast markings and the short primary projection compared to tertials
length. Figures 5, 6. Spotted Flycatcher ( Muscicapa striata striata ), Ventotene, Latina, Italy, April 2011 (Fig. 5) and Pan-
telleria, Trapani, Sicily, May 2009 (Fig. 6, photo by Igor Maiorano). The overall impression is of a colder and less homo-
geneous bird, with bold markings on breast and head; primary projection is longer than tertials length.
Morphological differences between two subspecies of Muscicapa striata (Passeriformes Muscicapidae)
275
the nearest 0.1 mm) and a thin strip of graph paper
(to the nearest 0.5 mm) strengthened by an equally
thin strip of transparent plastic.
The latter tool was necessary to measure P3: this
feather is usually measured using a special ruler, but
due to the specimens’ age, their rigidity, and their
historic value, some are from the prized Arrigoni
degli Oddi collection, we decided to use graph
paper as it is thinner and less invasive. Colour
analysis of the upperparts of Spotted Flycatcher
specimens was undertaken using an Ocean Optics
USB 2000 spectrometer at ISPRA.
Before proceeding with the spectrometer ana-
lysis of Spotted Flycatcher plumage we had to
calibrate the instrument and its associated software,
Ocean Optics Spectrasuite, which is provided by the
manufacturer of the spectrometer and the lamp. The
spectrometer was calibrated by reading and record-
ing on the software two values that were to corres-
pond with white and black. In order to do so we
used Ocean Optics’ WS1 Diffuse Reflectance
Standard for white, while for black we placed the
lighting fibreover the black square on X-Rite’s
Color Checker’s colour scale.
Once the programme was launched, only two
parameters needed to be set. Scan-to-average was
set at 5: for each colour reading of a given point,
five scans are automatically made, and their average
is recorded as the final value. Integration time was
set at 300 in order to prevent peaks in the graph
above the upper margin when the scanner was
placed above the white standard; in other words, to
ensure that reflectance on a white standard would
not return excessively high values that would have
led to a loss of information on the portion of the
graph falling outside the margins.
Once calibration was completed, we sampled
colours on each specimen as follows: three meas-
urements were taken from the mantle (usually two
from the right-hand side and one from the left) and
three more from the rump (by moving the scanner
along a vertical line from the top to the bottom of
the rump).
This means that for each specimen, the data
reported in the Table 2 comprises the averages of
15 measurements on the mantle and 15 on the
rump. In accordance with the instructions reported
by Hill & McGraw (2006) we selected and ranked
the data before analyzing them: we only considered
values with wavelengths between UV and red
(299.74 <X> 700.28), then sub-divided them into
intervals of approximately 10 nm, e.g. from 410nm
to 420nm.
The values we calculated (for both mantle and
rump) are as follows:
- Total Reflectance: the sum of all intervals
- UV Component: the sum of values falling
between 300nm and 400nm
Figure 7. The provenience of the museum specimens analysed. Figure 8. The number of birds collected per decade.
276
Michele Vigano & Andrea Corso
Museum
M. striata
striata
M. striata tyr-
rhenica
MCSM
10
7
MCZR
15
6
MSNF
5
4
ISPRA
8
1
Total
38
18
Table 1 . This table summarizes the number of specimens
studied, the museum they belong and their subspecific iden-
tification.
- UV Chroma: UV Component to Total Reflect-
ance ratio
- RED Component: the sum of values falling
between 600nm and 700nm
- RED Chroma: RED Component to Total Re-
flectance ration
The results are reported in the Tables 2 and 3 in
the following chapter. In order to better investigate
the results obtained with the colorimeter, we made
subsets of the original data (Figs. 9-20): we began
by removing from the sample of M. striata striata
all individuals from the narrow strip of Tyrrhenian
coastline in Tuscany, Latium, and Campania where
M. striata tyrrhenica may be breeding; we did not
remove two individuals captured on Ventotene
Island (Latina) and Capri (Naple) because they
matched M. striata striata in every regard and we
considered them to be spring migrants of M. striata
striata with a reasonable degree of certainty. A
second subset was made comparing the usual
sample of Sardinian specimens with a subset (n=6)
of M. striata striata specimens that show particu-
larly cold plumage tones on visual inspection. We
also divided the sample into old “pre-1960” and
recent “post- 1960” subsets, meaning that ‘recent’
specimens were no more than fifty years old, fol-
lowing Annenta et al. (2008).
In order to evaluate the differences in nape and
breast streaking between the two subspecies, we
compared the specimens visually (see, for example,
Galeotti et al. 2009). After an initial evaluation of
all specimens, we established categories that could
represent in sufficient detail the variability present
in the two taxa. We scored breast streaking on a 0
to 6 scale (0 indicating no streaking and 6 the heav-
iest streaking) and nape streaking on a 0 to 5 scale.
We assigned those values to each specimen; when
necessary, we compared the specimen under obser-
vation directly with the reference specimens.
RESULTS
Biometric analyses found significant differences
in wing morphology. Differences in maximum wing
chord were found to be statistically significant using
a t-test (t = 9.4407, p = 6.079e-12), confirming our
field observations of a shorted primary projection
in M. striata tyrrhenica.
Similar wing measurement data are reported in
the literature (e.g. Cramp & Perrins, 1993; Brichetti
& Fracasso, 2008). On the other hand, in a study of
birds ringed between mid- April and mid-May at
Capo Caccia, Sardinia (Marchetti & Baldaccini,
1995) did not report such a difference, although the
authors themselves suggested that such compar-
isons were better made using birds caught on their
breeding grounds during the reproductive season in
order to ensure correct subspecific identification. In
addition to the wing chord, significant differences
were found in the wing formula as well. The values
calculated for each primary are summarized in
figure 22, which shows wing formula for each
taxon. The most significant difference concerns the
relative distance between the longest primary (P3)
and P2; this characteristic is also depicted in figure
2 1 , which shows the distance (in mm) between P2
and P3 in each taxon. The t-test reveals significant
differences between the two subspecies concerning
this character (t = -5.1674, p = 6.536e-06), as well
as in the distance between P3 and P4 (t = 5.8634, p
= 6.768e-07). Differences in wing shape of this type
and extent are highly interesting. Similar discrepan-
cies have been found between sister species in
which one is a short-distance migrant and the other
a long-distance migrant (Chandler & Mulvihill,
1988;Monkkonen, 1995), or where there is a
gradient between more or less migratory subspecies
of the same species (Arizaga et al., 2006; Winkler
et al. 2010) or again in similar species where one is
migratory and the other sedentary (Chandler &
Mulvihill, 1990; Mila et al., 2008).
Morphological differences between two subspecies of Muscicapa striata (Passeriformes Muscicapidae)
277
Figures 9-14. A value for underpart markings was given to each specimen; seven categories were determined
(one central category not depicted), ranging from least marked (value 0) to boldly marked (value 6).
278
Michele Vigano & Andrea Corso
Figures 15-20. A value for head streakings was given to each specimen; six categories were determined,
ranging from least marked (value 0) to boldly marked (value 5).
Morphological differences between two subspecies of Muscicapa striata (Passeriformes Muscicapidae)
279
This phenomenon is known as “Seebohm’s
rule” and can be summed up as follows: long-
distance migrants have more pointed wings (shorter
inner primaries and longer outer primaries) com-
pared to short-distance migrants or noil-migratory
species, since longer and more pointed wings make
for more powerful flight compared to shorter, more
rounded wings (Seebohm, 1901; Calmaestra &
Moreno, 2001).
We also found differences in bill length meas-
ured from the distal end of the nostrils to the tip of
the bill, with p = 0.01582 and t = 2.5203, with M.
striata striata showing on average a longer bill; we
did not use the commonest bill measurement
method, from the tip of the bill to the base of the
skull, because for museum specimens it is less
reliable than the parameter used in this study
(Winker, 1998; Kuczynski et al., 2003). Our scores
for breast and nape streaking also confimied our field
observations, namely that nape and breast streaking
is less well defined in Sardinian and Corsican birds.
Colour analysis did not reveal any statistically
significant differences except in the sum of X fall-
ing between 300 and 400nm, or within the UV
spectrum. These differences fade away if one con-
siders the UV chroma, namely by dividing the UV
value by total reflectance. In order to better under-
stand the reasons for this, we carried out a number
of tests by modifying the data sample used in the
analysis in an attempt to remove the effect of certain
parameters that may have generated background
noise and muddled the results. The first subset
excludes all individuals from the narrow strip of
Tyrrhenian coastline where M. striata tyrrhenica
may breed: using this sample, differences in mantle
UV are no longer significant, but differences
emerge in terms of total reflectance and the red
component of the mantle, with M. striata tyrrhenica
slightly redder and paler than M. striata striata,
albeit with low statistical significance. However,
further manipulation of the sample for colorimetric
analysis, comparing M. striata tyrrhenica speci-
mens with six particularly cold plumaged M. striata
striata specimens did not find statistically signific-
ant differences for any variable.
This unexpected result, involving striata speci-
mens that showed clear differences in mantle tones
compared to M. striata tyrrhenica on visual inspec-
tyrrhenica
sffiaia rynrttenics « « ” « « « *
striata
Figure 21 . Chord values (in mm) recorded on Muscicapa. striata striata and M. striata tyrrhenica specimens.
Figure 22. Wing formula for both subspecies; note the rounder shape of M. striata tyrrhenica birds.
280
Michele Vigano & Andrea Corso
Muscicava striata striata Muscicava striata txrrhenica
variable
mean + sd
n
range
mean + sd
n
range
t
P
PI
13.75926 ±
1.931572
27
9.3-17.4
12.67500 ±
1.438286
16
10.2-15.1
1.9448
0.05868
P2
59.57778 ±
2.469247
27
55.0-64.4
55.33125 +
1.520841
16
52.9-57.8
6.2002
2.245e-07
P3
62.96429+
1.914509
28
60-67
60.12500 +
1.258306
16
58-63
5.3003
3.996e-06
P4
61.64074 +
2.176670
27
58.0-66.5
60.06250 +
1.223043
16
58.0-62.5
2.6544
0.01126
P5
57.36667 ±
2.300000
27
53.8-62.2
56.26154±
1.413352
13
54.2-59.1
1.5879
0.1206
P6
50.97407±
2.695094
27
47.5-59.5
49.61250±
1.433353
16
47.4-52.4
1.8645
0.06943
P7
45.86667±
1.840568
27
42.9-50.2
45.67333±
1.258381
15
43.7-48.1
0.3616
0.7195
P8
42.66296±
1.784581
27
39.7-46.6
42.35625±
1.175284
16
40.2-44.9
0.6118
0.544
P9
39.69259±
1.836369
27
36.9-43.8
39.73125±
1.151068
16
36.9-43.8
- 0.0757
0.9401
P10
36.84074±
1.727662
27
34.3-40.9
37.56000±
1.136913
15
35.2-40.4
-1.444
0.1565
chord
86.14286+
1.603567
28
83-90
81.50000 +
1.505545
16
79-84
9.4407
6.079e-12
bill L
8.392593+
0.2758566
27
8.0-8.9
8.140000 +
0.3680062
15
7.2-8.6
2.5203
0.01582
bill W
3.496429±
0.1990387
28
3. 1-3.9
3.420000±
0.1373213
15
3.2-3. 7
1.3244
0.1927
billT
4.357143±
0.2379365
28
3. 6-4.8
4.353333±
0.3888934
15
3.6-5. 1
0.0399
0.9683
breast
4.096774+
0.7897189
31
3-6
1.466667 +
0.9904304
15
0-3
9.7384
1.504e-12
head
3.586207+
0.7327659
30
2-5
1.333333 +
0.8164966
15
0-3
9.2998
9.361e-12
f R tot refl
385.2981±
74.04245
36
263.55-
603.68
397.1140±
55.03810
15
316.97-
486.38
-0.556
0.5807
f R UV
59.08778±
15.51324
36
36.00-99.42
61.77267±
13.38254
15
41.97-85.83
-0.5849
0.5613
f R CROM
A UV
0.151982±
0.0151761
36
0.11943-
0.18782
0.154283±
0.0155461
15
0.1324-
0.1910
-0.49
0.6263
f R RED
141.8281±
22.45401
36
100.9-200.3
148.8973±
20.66444
15
114.11-
181.89
-1.0476
0.3
fRCHRO
MA RED
0.370492±
0.0201814
36
0.331798-
0.41221
0.375287±
0.0162703
15
0.35564-
0.4098
-0.815
0.419
f M tot refl
303.9143±
33.40160
36
236.78-
344.78
329.1888±
40.71543
16
290.13-
405.80
-1.5784
0.1302
f_M_UV
44.53722+
6.774240
36
31.52-57.73
49.10250+
7.547965
16
39.27-65.49
-2.1659
0.03512
f M CHRO
MA UV
0.144340±
0.0137475
36
0.11600-
0.171272
0.147015±
0.0119409
16
0.13113-
0.1696
-0.6728
0.5042
f M RED
112.1264±
9.672242
36
92.18-
126.12
120.3400±11
.764493
16
108.76-
141.91
-1.773
0.09145
f M CHRO
MA RED
0.376839±
0.0175326
36
0.338173-
0.40683
0.375442±
0.0206807
16
0.34161-
0.4047
0.2508
0.803
Table 2. All the statistical results from our study are here summarized; the variables highlighted
in boldface are those for which the t-test found values <0.05.
Morphological differences between two subspecies of Muscicapa striata (Passeriformes Muscicapidae)
281
Muscicapa striata striata Muscicava striata txrrhenica
variable
mean + sd
n
range
mean + sd
n
range
t
P
M tot refl
309.2468 ±
2
236.78-365.09
329.1888 ±
16
290.13-405.80
-2.2777
0.0279
31.11327
9
40.71543
45.6875 ±
2
49.10250 ±
M UV
35.20-56.61
16
39.27-65.49
-1.6468
0.1071
6.037829
9
7.547965
M CHROMA
0. 1476630 ±
2
0.123644-
0.147015 ±
16
0.13113-
0.179
0.8588
UV
0.0113407
9
0.171272
0.0119409
0.1696
115.6518 ±
2
120.3400 ±
M RED
92.18-126.12
16
108.76-141.91
-2.5046
0.01623
10.24447
9
11.764493
M CHROMA
0.3748562 ±
2
0.338173-
0.375442 ±
16
0.34161-
-0.0983
0.9222
RED
0.0180199
9
0.399920
0.0206807
0.4047
R tot refl
388.8814 ±
2
263.55-603.68
397.1140 ±
15
316.97-486.38
-0.3589
0.7215
78.95214
9
55.03810
R UV
60.81893 ±
2
36.00-99.42
61.77267 ±
15
41.97-85.83
-0.1947
0.8466
16.22125
9
13.38254
R CHROMA
0.1549453 ±
2
0.121704-
0.154283 ±
15
0.1324-0.1910
0.1413
0.8884
UV
0.0141347
9
0.187822
0.0155461
141.9193 ±
2
148.8973 ±
R RED
100.9-200.3
15
114.11-181.89
-0.9519
0.3468
23.99409
9
20.66444
R CHROMA
0.3674105 ±
2
0.331798-
0.375287 ±
15
0.35564-
-1.3203
0.1941
RED
0.0197631
9
0.408994
0.0162703
0.4098
Table 3. Same colorimetric variables analyzed in the previous table, but with a different subset of data: birds collected from
the narrow strip of Tyrrhenian coastline in Tuscany, Latium, and Campania, where M. striata tyrrhenica could occur, were
removed.
tion, suggests that the method we used for our
colorimetric analysis is not ideal for detecting such
subtle differences in plumage pigmentation.
Additional comparisons looked at the effects of
time on the state of preservation of specimen. In
accordance with other works that tested colour
deterioration in museum specimens (Armenta et al.,
2008; Doucet & Hill, 2009), we found highly
significant differences between old (pre-1960) and
recent (post- 1960) specimens.
We used this data to build a linear model to
identify the variables that most affected colour
variation. As expected, taxon did not have a stat-
istically significant effect, while year of collection
did (F(1.46) = 7, P = 8.408e-05). In other words,
specimens that were more than fifty years old
showed a statistically-significant higher total re-
flectance, and thus appeared paler.
The biometric and colorimetric data collected
in this study is summarized in Table 2, which also
indicates sample size (n), the minimum and max-
imum values recorded (range) and the t and p va-
lues for the t-test as applied to each variable for the
two taxa. The variables highlighted in boldface are
those for which the t-test found values <0.05, mean-
ing that the differences between the two taxa for
282
Michele Vigano & Andrea Corso
the variable in question were statistically signific-
ant. The variables “PI” to “P10” indicate primary
length from the outermost to the innermost; “chord”
indicates the length of the maximum wing chord,
namely the closed wing measured from the carpal
joint; “bill L, H, and T” respectively indicate bill
length, height, and thickness; “breast” and “head”
indicate the amount of streaking in these two areas
scored after a visual examination.
Colorimetric data follows: variables initialed
with an M refer to the mantle, and those with an R
to the rump; tot_refl refers to total reflectance, UV
and RED respectively refer to the sum of X falling
between 300 and 400 and between 600 and 700;
UV CHROMA and REDCHROMA indicate the
ratio between these two variables and total reflec-
tance.
DISCUSSION AND CONCLUSIONS
The objective of this study was to test the dif-
ferences observed in the field between the M.
striata striata and M. striata tyrrhenica subspecies
of Spotted Flycatcher as objectively as possible, by
using methods that would not be influenced by dif-
ferences in perception of colour and proportions on
the part of different observers. The results con-
firmed the morphological differences observed in
the field and cited in the literature, and the different
intensity and extent of streaking on the underparts
and the nape. Nevertheless, to better assess these
parameters a larger sample, in both quantitative and
qualitative terms, would be preferable, and would
ideally include a larger number of birds captured on
their breeding grounds. Differences in wing-shape
are important not only from an identification per-
ceptive, but also in light of the relationship between
wing morphology and migratoiy distance (Baldwin
et al., 2010; Monkkonen, 1995).
The shorter, more rounded wings of M. striata
tyrrhenica suggest that birds breeding in Corsica
and Sardinia may have a shorter migration com-
pared to birds from continental Italy and Europe.
This is all the more interesting given that there is
no solid data in the literature on the non-breeding
range of M. striata tyrrhenica (Cramp & Perrins,
1993; del Hoyo et al., 2006), that should anyway be
sub-saharan, given the absence of evidence of
winter sightings north of the Sahara.On the other
hand, our colorimetric analyses failed to confirm
the differences observed in the field and reported in
the literature. To conclude, biometric measurements
and an analysis of plumage streaking confirmed the
presence of some important differences between the
two taxa, including characters that can be seen in
the field, while the colorimetric analysis did not
produce the expected results, although it had some
interesting implications concerning the preservation
of museum specimens and their use in studies of
plumage colour.
There are several other instances of taxa that
have similar distributions to M. striata tyrrhenica
Spotted Flycatchers and are morphologically very
similar to the taxa breeding elsewhere in Italy and
Europe being recognized as full species after
in-depth analyses of morphology, voice, ecology
and DNA: examples include Corsican Finch
( Carduelis corsicana ) (Cramp & Perrins, 1993;
Sangster, 2000; Forschler & Kalko, 2007, Forschler
et al. 2009) and Moltoni’s Warbler ( Sylvia sub-
alpina ) (Brambilla et al., 2008), both recently re-
cognized as full species; further research on M.
striata tyrrhenica Spotted Flycatcher is needed.
ACKNOWLEDGMENTS
We would like to thank Professor Giuseppe
Bogliani (Pavia, Italy) for his advice and his
constant support throughout this study. We also
thank Giorgio Chiozzi of the Museo Civico di
Storia Naturale of Milan (Italy), Carla Marangoni
and Maurizio Gattabria of the Museo Civico di
Zoologia of Rome (Italy), the staff of the Museo
di Storia Naturale of Forli (Italy), and Lorenzo
Serra, Simone Pirrello and Adriano de Faveri of
the Institute for Environmental Protection and
Research, ISPRA (Italy); without their help this
study would not have been possible. For their help
fulsuggestions and advice we thank Giacomo
Assandri (Turin, Italy), Mattia Brambilla (Milan,
Italy), Monica Clerici (Milan, Italy), Andrea
Galimberti (Milan, Italy), Ottavio Janni (Naples,
Italy), Igor Maiorano (Trieste, Italy), Violetta
Longoni (Milan, Italy), Diego Rubolini (Milan,
Italy), and Roberto Sacchi (Pavia, Italy). We are
very grateful to Ottavio Janni (Naples, Italy) who
translated this paper from Italian to English.
Morphological differences between two subspecies of Muscicapa striata (Passeriformes Muscicapidae)
283
REFERENCES
Arizaga J., Campos F. & Alonso D. 2006. Variations in
wing morphology among subspecies might reflect
different migration distances in Bluethroat. Omis
Fennica, 83: 162-169.
Armenta J.K., Dunn P.O. & Whittingham L.A., 2008.
Effects of specimen age on plumage colour. Auk,
125: 803-808.
Arrigoni degli Oddi E., 1929. Ornitologia italiana.
Milano, Hoepli, 1046 pp.
Baldwin M.W., Winkler H., Organ C.L. & Helm B.,
2010. Wing pointedness associated with migratory
distance in common-garden and comparative studies
of stonechats ( Saxico/a torquatd). Journal of Evolu-
tionary Biology, 23: 1050-1063.
Blondel J., Chessel D. & Frochot B. 1988. Bird species
impoverishment, niche expansion, and density infla-
tion in Mediterranean island habitats. Ecology, 69:
1899-1917.
Brambilla M., Vitulano S., Spina F., Baccetti N., Gar-
gallo G., Fabbri E., Guidali F. & Randi E. 2008. A
molecular phylogeny of the Sylvia cantillans
complex: cryptic species within the Mediterranean
basin. Molecular Phylogenetics and Evolution 48:
461-472.
Brichetti, P. & Fracasso, G. 2008. Ornitologia italiana.
Vol. 5. Turdidae-Cisticolidae. Oasi Alberto Perdisa
Editore, Bologna, 432 pp.
Calmaestra R.G. & Moreno E., 2001. A phylogenetic
analysis on the relationship between wing morpho-
logy and migratory behavior in Passeriformes. Ardea,
89: 407-415.
Chandler C.R. & Mulvihill R.S., 1988. The use of wing
shape indices: an evaluation. Ornis Scandinavica, 19:
212-216.
Chandler C.R. & Mulvihill R.S., 1990. Wing-shape
variation and differential timing of migration in dark-
eyed juncos. Condor 92: 54-61.
Corso A., 2005. Avifauna di Sicilia. L’Epos editore,
Palermo, 324 pp.
Cramp S. & Perrins C.M. (Eds.), 1993. Handbook of the
Birds of Europe and the Middle East and North
Africa. Vol. VII Flycatchers to Shrikes. Oxford
University Press, Oxford, 584 pp.
del Hoyo J., Elliot A. & Sargatal J., 2006. Handbook of
the Birds of the World. Vol. XI. Lynx Edicions,
Barcelona, 800 pp.
Doucet S.M. & Hill G.E., 2009. Do museum specimens
accurately represent wild birds? A case study of ca-
rotenoid, melanin, and structural colours in long-
tailed manakins Chiroxiphia linearis. Journal of
Avian Biology, 40: 146-156.
Eck S., Fiebig J., Fiedler W., Heynen I., Nicolai B.,
Topfer T., van den Elzen R., Winkler R. & Woog F.,
2011. Measuring birds. Vogel vennessen. Deutsche
Ornithologen-Gesellschaft. Wilhelmshaven, 118 pp.
Forschler M.I. & Kalko E.K.V., 2007. Geographical
differentiation, acoustic adaptation and species
boundaries in mainland citril finches and insular
Corsican finches, superspecies Carduelis [ citrinella ].
Journal of Biogeography, 34: 1591-1600.
Forschler M.I., Senar J.C., Perret P. & Bjorklund M.,
2009. The species status of the Corsican Finch
(Carduelis corsicana ) assessed by three genetic
markers with different rates of evolution. Molecular
Phylogenetics and Evolution, 52: 234-240.
Galeotti P., Rubolini D., Sacchi R. & Fasola M., 2009.
Global changes and animal phenotypic responses:
melanin-based plumage redness of scops owls in-
creased with temperature and rainfall during the last
century. Biology Letters, 5: 532-534.
George T.L., 1987. Greater land bird densities on island
vs. mainland: relation to nest predation level.
Ecology, 68: 1393-1400.
Hill G.E. & McGraw K.J., 2006. Bird Coloration,
Volume 1. Mechanisms and Measurements. Harvard
University Press, Cambridge, 41-89.
Knox A., 1980. Post-mortem changes in wing lengths
and wing formulae. Ringing & Migration, 3: 29-31.
Kuczynski L., Tryjanowski P., Antczak M., Skoracki M.
& Hromada M., 2003. Repeatability of measurements
and shrinkage after skinning: the case of the Great
Grey Shrike Laniusexcubitor. Bonner Zoologische
Beitrage, 51: 127-130.
MacArthur R.H. & Wilson E.O., 1967. The theory of
island biogeography. Princeton University Press, 224
pp.
Marchetti C. & Baldaccini N.E., 1995. I Pigliamosche
(Muscicapa striata ) in transito prenuziale su Capo
Caccia: caratteri biometrici e fenologici. Supple-
mento alle Ricerche di Biologia della Selvaggina, 22:
537-538.
Mila B., Wayne R.K. & Smith T.B., 2008. Ecomorpho-
logy of migratory and sedentary populations of the
yellow-rumped warbler (Dendroica coronata ).
Condor, 110: 335-344.
Monkkonen, M. 1995. Do migrant birds have more
pointed wings?: a comparative study. Evolutionary
Ecology, 9: 520-528.
Sangster G., 2000. Genetic distance as a test of species
boundaries in the Citril Finch (Serinus citrinella ): a
critique and taxonomic reinterpretation. Ibis, 142:
487-490.
Seebohm H., 1901. Birds of Siberia. Murray, London,
512 pp.
Schiebel G., 1910. Neue Vogelformenaus Corsica. Or-
nithologisches Jahrbuch, 21: 102-103.
Tellini F.G., Baccetti N., Arcamone E., Meschini A. &
Sposimo P., 1997. Atlante degli uccelli nidificanti e
svernanti in Toscana (1982-1992). Provincia di
284
Michele Vigano & Andrea Corso
Livorno e Centro Ornitologico Italiano. Quademi del
Museo Provinciale di Storia Naturale di Livorno.
Monografie 1.
VV. AA. in Thibault J.-C. & Bonaccorsi G., 1999.
The birds of Corsica. BOU Checklist No. 17. British
Ornithologists’ Union. The Natural History Museum,
Tring, Herts, 171 pp.
vanDuivendijk N., 2010. Advanced bird identification.
New Holland Publishers, London, 308 pp.
Winker K., 1998. Suggestions for measuring external
characters of birds. Ornitologia Neotropical, 9:
23-30.
Biodiversity Journal, 2015, 6 (1): 285-296
Monograph
A quantitative morphological geographical study from a
widely distributed raptor: the Lesser Kestrel Falco naumann
Fleischer, 1818 (Falconiformes Falconidae)
Andrea Corso 1 *, Lorenzo Starnini 2 , Michele Vigano 3 & Justin J.F.J. Jansen 4
'Via Camastra 10, 96100 Siracusa, Sicily, Italy; e-mail: zoologywp@gmail.com
2 Via Cavour 71, 06019 Umbertide, Perugia, Italy; e-mail: lorenzo.stamini@gmail.com
3 Via Ongetta 5, 21010 Germignaga, Varese, Italy; e-mail: mikivigano@yahoo.com
4 c/o Naturalis Biodiversity Center, Darwinweg 2, 2333 CR Leiden, the Netherlands; e-mail: justin.jansen@naturalis.nl
"■Corresponding author
ABSTRACT Lesser Kestrel Falco naumanni Fleischer, 1818 (Falconiformes Falconidae) is considered a
monotypic species. F. naumanni pekinensis Swinhoe, 1870 was described from Beijing, China.
Although considered valid for most of the 20th century, some authors treated F. naumanni
pekinensis as a synonym of F naumanni naumanni, and subsequent authors have since regarded
“ pekinensis ” as an invalid taxon. Recent field observations in Asia and Europe and museum
studies have confirmed diagnosable differences in (fresh) adult males. Comparing morphology
between nominate “ naumanni ” and “ pekinensis ”, with the latter invariably showing more
extensive grey on the wing coverts and darker and more saturated colours on both the under-
parts and upperparts, with all grey areas, including the hood, being a darker, deeper lead-grey.
Females often have more extensive dark markings and a better-defined dark eye-line but
apparently are indistinguishable in most cases. This study aims to re-evaluate F. naumanni
pekinensis and to discuss geographic variation in the subspecies in a widely distributed raptor.
KEY WORDS Lesser Kestrel; pekinensis', naumanni', subspecies; geographical variation.
Received 25.05.2014; accepted 30.06.2014; printed 30.03.2015
Proceedings of the 2nd International Congress “Speciation and Taxonomy”, May 1 6th- 1 8th 20 1 4, Cefalu-Castelbuono (Italy)
INTRODUCTION
Today, Lesser Kestrel Falco naumanni Flei-
scher, 1818 (Falconiformes Falconidae) is conside-
red a monotypic species (cf. Cramp & Simmons,
1980; Snow & Perrins, 1998; Forsman, 1999;
Clark, 1999; Corso, 2000, 2001a; Ferguson-Lees &
Christie, 2001). After a few years, was described F.
chenchris pekinensis Swinhoe, 1870 from two birds
(adult male and immature male) (cf. Swinhoe,
1870; Dresser, 1871-1881). Currently, F. naumanni
pekinensis is regarded as a synonym of F. naumanni
naumanni (cf. Vaurie, 1965; Dickinson & van
Remsen, 2013).
In September-October 2003, two authors (AC,
JJ) were at Chokpak Ornithological Station, Jambyl
Province, Kazakhstan together with Wim Nap and
Arend Wassink, studying raptors and other birds in
collaboration with Andrei and Edward Gavrilov as
Vladimir Kolbinsev. AC was intrigued by the up-
perwing pattern of several adult male Lesser Ke-
strels that were caught in the large Heligoland-traps
onsite (and subsequently ringed) as in birds obser-
ved in the field. They appeared consistently diffe-
rent from birds AC observed within the Western
Palaearctic. The past twelve years, in addition to our
field studies of Lesser Kestrel, we have studied
286
Andrea Corso et alii
skins from museums worldwide and photos from
throughout their breeding range. We have found
that eastern populations, especially the well-isolated
breeding grounds in China, are phenotypically stri-
kingly different from western populations. This
suggests that F. naumanni pekinensis may be a well
identifiable taxon, although it’s breeding and win-
tering distribution remains to be fully elucidated.
This paper reports the preliminary results of our stu-
dies concerning western and eastern populations,
with a focus on the latter, particularly “ pekinensis ”.
In this paper we describe plumage colour and pat-
tern, in special fresh adult males, of both Western
(. F. naumanni naumanni ) and Eastern Lesser Kestrel
(Chinese Lesser Kestrel F. naumanni pekinensis and
intermediate populations). Their field identification
and plumage variability will be discussed separately
(Corso et al., personal data).
MATERIAL AND METHODS
In this study we investigated in detail adult birds
in the field within the borders of the Western Pala-
earctic, and to a more limited extent in Asia. Bet-
ween 2003 and 2014, we studied birds from Africa
(Egypt, Eritrea, Kenya, Morocco, Somalia, Sudan,
Tanzania and Tunisia), Asia (Armenia, Azerbaijan,
Burma, China, Georgia, India, Israel, Kazakhstan,
Laos, Mongolia, Russia, Oman, Turkey, Turkmeni-
stan, Saudi Arabia and Yemen) and Europe (France,
Greece, Portugal, Spain and Italy) in field, museum
or photographs. During fall 2003, AC and JJ studied
tens of Lesser Kestrels in the hand during ringing
operations in Kazakhstan as in the field (up to 1 .000
birds were observed during their stay). Particular
attention was given to the adult males and to a lesser
extent to adult females. Juveniles were not studied
in much detail.
The skins we studied are held in the following
museums and bird collections: American Museum
of Natural History, New York, U.S.A. (AMNH);
Institute of Zoology, Almaty, Kazakhstan (IZA);
Museo Civico di Scienze Naturali “Angelo Priolo”,
Randazzo, Italy (MCR); Museo Civico di Storia
Naturale of Milan, Italy (MCSM); Museo Civico of
Terrasini, Italy (MCT); Museo Civico dell’Univer-
sita di Scienze Naturali of Catania, Italy (MCUCT);
Museo Civico di Zoologia of Rome, Italy (MCZR);
Museum National d'Histoire Naturelle, Paris,
France (MNHN); Museo Regionale di Scienze Na-
turali of Turin, Italy (MRSN); Museo di Storia Na-
turale “Giacomo Doria”, Genoa, Italy (MSNGD);
Museo di Storia Naturale “La Specola”, Florence,
Italy (MSNLS); National Zoological Museum of
China, Beijing, China (NZMC); Naturalis Biodiver-
sity Center, Leiden, the Netherlands (NBC); Natural
History Museum, Tring, England (NHM); Naturhi-
storisches Museum Wien, Vienna, Austria (NMW);
Peabody Yale Museum of Natural History, New
Haven, U.S.A. (PMNH); Museo Civico di Storia-
Naturale di Carmagnola, Italy (SNCa); Museum fur
Naturkunde, Berlin, Germany (ZMB) and in thir-
teen private collections.
Other abbreviations: AC: Andrea Corso; JJ: Ju-
stin J.F.J. Jansen; MV: Michele Vigano.
The list of the specimens (both skins and moun-
ted) examined from the museums and private col-
lections were, after objective examination, divided
into four groups (Figs. 1, 2):
Group A: Falco naumanni pekinensis : one of
two syntypes of the subspecies (Figs. 8, 9); 20 adult
males and 8 adult females (China). For Figs. 3, 4, 5
and 6 we used only fresh breeding plumage males
from China (N=13).
Group B: Falco naumanni ssp.: 28 adult males,
25 females (age combined) (Asia: Mongolia, Altai
Mountains breeding area as well as Burma, India
and Laos wintering area). For Figs. 3, 4, 5 and 6 we
used only fresh breeding plumages males from
Mongolia (N=13).
Group C: Falco naumanni ssp.: 87 adult males;
60 (age combined) females (Asia: Arabian Penin-
sula (unspecified countries), Afghanistan, Azerbai-
jan, Iraq, Kazakhstan, Pakistan, Turkmenistan;
Africa: Eritrea, Kenya, Somalia, Tanzania). For
Figs. 3, 4, 5 and 6 we used only fresh breeding plu-
mage males from Kazakhstan (N=9), Turkmenistan
(N=6), Azerbaijan (N=7) and Afghanistan (N=4).
Group D: Falco naumanni naumanni : 349 adult
males; 172 (age combined) females (Europe: Alba-
nia, Czech Republic, France, Greece, Italy, Mace-
donia, Portugal, Slovenia, Spain; Africa: Algeria,
Angola, Botswana, Egypt, Ethiopia, Libya, Mauri-
tania, Morocco, Niger, Senegal, South Africa, Tan-
zania, Tunisia; Asia: Annenia, Georgia, Iran, Iraq,
Israel, Jordan, Kyrgyzstan, Lebanon, Palestine,
Syria, Turkey). For Figs. 3, 4, 5 and 6 we used only
fresh breeding plumage males from Turkey (N=5),
Greece (N=12), Albania (N=5), France (N=5) and
Spain (N=29).
A quantitative morphological geographical study from a widely distributed raptor: the Lesser Kestrel Falco naumanni 287
Figure 1. Plumage types as indicated in the text: upper 2 birds (group D), ssp. naumanni, among this group, left bird is an
example with least grey extension on upperwing coverts and tertials, and on the right typical nominate naumanni. Note the
grey extension on wing, plumage colour saturation and bare parts colour. Central bird, intermediate bird, plumage group B
and C in the text. Lower 2 birds, classic “ pekinensis ”, plumage group A, from China breeding populations and allegedly
from India, Laos and Burma during winter and on passage. The slight variation found is shown (artwork by Lorenzo Stamini).
Note on pekinensis that the plumage colour, chiefly mantle and grey tones, is darker, more saturated than nominate naumanni,
and that the upperwing coverts are entirely grey as well as the bare parts are more orange-ochre.
288
Andrea Corso et alii
Additionally, to test the reliability of plumage
colour and pattern in discriminating between popu-
lations, we took four photographs of each Lesser
Kestrel skin. We sampled: upperside, underside and
lateral sides (in order to check if there were diffe-
rences between upperwing pattern of both wings)
(Figs. 10-19). We only used indirect sunlight. All
specimens were photographed with the same ca-
mera at a fixed distance, with a Kodak Gray Scale
(Kodak, 2007) in the background (a standard scale
of grey values ranging from 0, white to 19, jet
black) (Fig. 7). This is an objective way of measu-
ring the grey colours in bird plumages (cf. Adriaens
et al., 2010; Bot & Jansen, 2013). The grey card in
the background enabled us to calibrate the colours
in the photograph, using Adobe Photoshop 10.0.1.
We used brightness as a measure to assess the grey
and red parts in adult males. To quantify the bri-
ghtness of the red parts we measured its median
spectral reflectance of red, green and blue (RGB)
by using Photoshop. To assess the amount of grey
and red on the head and upperparts, we measured
the saturation. This was calculated by using the for-
mula: saturation = 1 00 x ((MAX-MIN) / MAX),
where MAX and MIN are the maximum and mini-
mum of the median RGB values as measured by
Photoshop. A low saturation value corresponds to
much grey in the head and upperparts and vice
versa. Forthe analysis of colours we used the plates
in Ridgway (1912: plate II and III). In particular,
we compared the following:
1 . Grey value of rump and tail;
2. Grey value of head (hood);
3. Percentage of grey coloured upperwing coverts.
To sample the upperwing coverts, we measured
the percentage of grey coloured coverts (versus rus-
set-brown coverts), therefore, how many coverts
(median MC+ greater GC+ lesser LC) were grey.
On skins, we considered the coverts visible on clo-
sed wing by using Photoshop to calculate the per-
centages. All statistical analysis was performed
using R version 2.15.1 (R Development Core Team
2013). Numerical variables (Grey hood and % of
grey UPW covers) were tested for normality using
the Shapiro-Wilk test. Tests were significant which
indicates that these variables do not come from a
normally distributed population. Therefore we used
a non-parametric approach (Kruskall- Wallis test) to
test for a significant difference between groups for
these 2 variables. To test for differences between
groups in the other 2 variables (colour_of_upper-
parts and deep_intensity_of_underparts) we used
generalized linear models (GLM).
Regarding the specimens and photos of adult fe-
males under investigation, we focused on the bol-
dness, width and distribution/demarcation of the
dark markings on the head, underwing, tail, mantle
and breast. We arbitrary divided specimens on the
basis of the extent and boldness of these marking
without looking at the label (no geographical pre-
indication), and subsequently checked if these fea-
tures were related to the classified groups (Fig. 2).
After that, we assigned a numerical code to the pre-
sence, definition and boldness of the dark markings
on the face:
1 . no dark eye-line and weak moustache mark;
2. weak eye-line and bolder moustache;
3. obvious and bold dark eye-line as well as moustache.
The Falco naumanni pekinensis syntypes
Swinhoe (1870: 442) described “ pekinensis ”
using an adult male collected on Septemberl8th
1868 at the hills overlooking the Ming Tombs (40°
25’ 05” N, 116° 22’ 41” E) (42 kilometres north-
northwest of Beijing) (Figs. 8, 9) and an juvenile
male collected at the Western Hills (roughly in
Miyun County about 93 kilometres northeast of
Beijing (indicative 40°31’40.8”N, 116°48’02.5”E)
between 10-12 August 1868 (Swinhoe, 1870: 436).
Specimens from Swinhoe became spread, and
the syntype now present in NHM, arrived as part of
a load of 480 Accipitres and Striges in three loads
in 1886 (Sharpe 1906) donated by Henry Seebohm.
Amongst these specimens is the adult male, now la-
belled as BMNH. 1886.3.25.272, and regarded as
syntype (Warren 1966: 222). This specimen has
been collected according to Warren (1966) and it is
labelled as collected on October 18th (sic!) 1868.
Henry E. Dresser noted that both specimens in his
‘A history of the birds of Europe' (1871-82, VI: 135)
were still in Swinhoe’s private collection, and col-
lected in August and September 1868 near Beijing.
A request at electronic Bulletin for European
Avian Curators (EBEAC - September 2014) and re-
quests elsewhere did not help in locating the imma-
ture male. By chance, a juvenile and adult Amur
Falcon Falco amurensis Radde, 1863 were collec-
ted near Beijing by Swinhoe in August 1868 (Tri-
A quantitative morphological geographical study from a widely distributed raptor: the Lesser Kestrel Falco naumanni 289
stram Collection Liverpool T2824 (Tristram, 1889:
67) was reported (Clem Fisher in litteris). Their plu-
mage is notably different, but strangely enough not
reported by Swinhoe (1870).
Description: Swinhoe described the types rather
briefly: “ Large numbers of Kestrels were flying and
hovering about. Their movement struck me as pe-
culiar; and on shooting a male we found the species
to be a race of Falco cenchris, Naumann. We pro-
cured on this occasion an adult male, and in the We-
stern Hills a young male. They agree in size and
form with Falco cenchris of Europe; but the adult
male has all the wing-coverts grey right up to the
scapulars, most of them narrowly edged with ru-
fous. The adult has the inner or short primaries
broadly bordered at their tips with whitish, rufous
in the immature, and wanting in the European bird.
Both adult and immature have the white on the
under quills 3 V 4 inches short of their tips; in the
European bird it advances one inch nearer the tips.
I will note this Eastern race as var. pekinensis. It
will probably be the bird that winters in India.”
RESULTS
We sampled 108 fresh adult males from the
breeding ranges for this analysis. We sampled the
percentage of grey upperwing coverts, grey value
in the hood as for the colouration of the colour of
Falco naumanni naumanni
Group D
Falco naumanni ssp
Group C and B Fako naumanni pekinensis
Group A
Group D
— ?
Group A (and B?)
^5
Group C and B
Figure 2. World distribution (breeding range - red, wintering area - green) of the two subspecies discussed in this paper
(plumage group A and D), and of intermediate populations and pekinensis-type birds (B/C) as found during this study and
according to past literature. Delimitation of the range is roughly indicated and should be considered as solely indicative.
290
Andrea Corso et alii
the upperparts and intensity of underparts. The pro-
portion of Group A is 12 %, group B 12 %, group
C 24 % and for group D 52 %. Although the pro-
portion of group D is well out of proportion we
think the number is sufficiently large to allow good
comparison as shown in the results. The results of
testing four characters among the four defined areas
is summarized in Fig. 2 and also outlined below.
Group A. The adult males collected in China
(N=13) show a mean of 1 1 .46 (min. 10; max. 12) on
the Kodak Grey Scale (Kodak, 2007) in hood colou-
ration (Figs. 3.7). For grey in the upperwing coverts
the mean is 93.5 % (min. 85; max. 99) of grey (Figs.
4, 8, 10, 11, 13, 15, 16). The intensity of underparts
is 100 % Mikado-orange mars yellow (Fig. 5). The
colouration in upperparts varies slightly as show in
Fig. 6, but the mean colouration was Burnt Sienna
(8 out of 13 birds). Breeding adult females: 62.5%
of the specimens from China (N=8) showed a well-
defined dark eye-line (code 3). The remaining (N=3)
showed a weaker marking (code 2). Flowever, juve-
niles of both “ pekinensis ” and “ nanmanni show a
bolder and better-marked moustache and eye-line,
compared to adult female, making any relevant use
in the field to identify “ pekinensis ” from “ nau -
manni ” of these characters extremely hard. Concer-
ning the underwing pattern, the outer primaries
(wing-tip) as well as the trailing edge of the wing
(inner primaries and secondaries) are more extensi-
vely and conspicuously dark in all Chinese females
than in adult female “naumanni” in group D. Also,
although rather variable, on average the black bar-
like marks on the mantle were wider and more con-
spicuous than in typical adult female “naumanni ” .
Birds from photographs shown a higher amount of
variability, with several individuals lacking dark
eye-line thus being almost identical to European fe-
males but a little darker and more patterned. As for
adult male, also in adult female, the cere and the bare
skin of the eye-ring, is brighter, deeper coloured and
more orange-ochreous than in adult female “ nau -
mannF.
Group B. The adult males collected in Mongolia
(N=13) shows a mean of 8.8 (min. 7; max. 10)on
the Kodak Grey Scale (Kodak, 2007) in hood co-
louration (Fig. 3, 7). For the wing coverts the mean
is 71.9 % (min. 50 %; max. 90 %) of grey (Fig. 4)
(Kodak, 2007). The intensity of underparts is
mixed, as 7 birds are mikado-orange-mars-yellow
and 6 are orange -buff (Fig. 5). The colouration in
upperparts is mixed, as seen in Fig. 6.
Group C. The adult males collected in Kaza-
khstan, Turkmenistan, Azerbaijan and Afghanistan
(N=26) show a mean of 8.07 (min. 7; max. 10) on
the Kodak Grey Scale in hood colouration (Fig. 3,
9) (Kodak, 2007). For the wing coverts the mean is
56.5 % (min. 40 %; max. 86 %) of grey (Fig. 4).
The intensity of underparts is mainly orange-buff
(20 out of 26) (Fig. 5). The colouration in upper-
parts varies as shown in Fig. 6.
Group D. The adult males collected Turkey,
Greece, Albania, France and Spain (N=56) show a
mean of 7. 14 (min. 6; max. 1 0) on the Kodak Grey
Scale in hood colouration (Figs. 3, 7) (Kodak,
2007). For the wing coverts the mean is 30.2 %
(min. 15; max. 45) of grey (Figs. 1,4, 17). Notable
is the differentiation in western and eastern birds of
the distribution are of this population. As we found
a certain variability on the extension of the grey co-
loured upperwing coverts. Some as typical birds as
shown in every field guide and handbooks as well
as birds showing almost no grey on coverts or only
some tinged grey (Figs. 1, 17) in the western part
of the distribution area. The intensity of underparts
is mostly buff-yellow or capucine-yellow (46 out of
56) (Fig. 5). The colouration in upperparts varies as
shown in Fig. 6.
For what concern the female, only 5% of adult
females showed a dark eye-line (code 3) (higher
percentage when looking at juvenile; Corso, 2000,
2001a; AC pers. obs.). Dark markings on the mantle
were on average narrower and less striking than in
the most marked adult female pekinensis. However,
we failed to find relevant differences. In juvenile
naumanni we found them being darker and with
bolder/wider dark markings than adult females, ad-
ding to the difficulty to the separate them. Bare
parts were less orange and paler, andalways yello-
wer than “ pekinensis ”.
DISCUSSION
The four groups (adult males in fresh plumage)
were significantly different when considering the
features: hood and % of grey UPW covers (Kru-
skall- Wallis test) (Table 1). This can be seen in Figs.
3, 4. These figures report averages and standard er-
rors (SE) for each group. The four groups were also
significantly different when considering the other
two variables (Table 2). Figures 5 and 6 represent
A quantitative morphological geographical study from a widely distributed raptor: the Lesser Kestrel Falco naumanni 291
Plot of Means
Plot of Means
Figure 3. This figure represent averages
and standard errors (SE) for the four
groups in grey colouration in Lesser Ke-
strel hood. Fig. 4. This figure represent
averages and standard errors (SE) for the
four groups in the % of grey coloured in
Lesser Kestrel upperwing coverts. Fig.
5. Represent the % within the group of
the considered colouration of the upper-
parts within each variable. Fig. 6: repre-
sent the % within the group of the
considered intensity of colouration in the
underparts within each variable. Fig. 7.
Kodak Grey Scale value of the hood in
fresh adult male Lesser Kestrel Falco
naumanni showing the range encounte-
red in the specimens studied: range 10-
12 in “ pekinensis ” (mean 11.46); range
6-10 in nominate “ naumanni ” (mean
7.14). In fact, grey hue and intensity is
on average deeper and darker in Chinese
Lesser Kestrel compared to Western Les-
ser Kestrel.
292
Andrea Corso et alii
Figure 8. Falco naumanni pekinensis, adult male Reg. no. BMNH. 1886.3.25.272. Near Ming Tombs, north of Peking,
18.X. 1868, syntype, R. Swinhoe leg. (H. van Grouw, NHM, Tring). No illustration in any modern field guide is available of
such a plumage, with no description or illustration reporting these characters. Fig. 9. F. naumanni pekinensis, same bird as
plate 5 (H.van Grouw, NF1M, Tring). Note the very richly coloured underparts, much richer than any nominate “naumanni” .
Fig. 10. F. naumanni pekinensis, adult male, Hebei, China, 27.IV. 1937 (He Peng, NZMC). A fresh adult male “ pekinensis ”
from the typical breeding range of the taxon, showing very intense brick-red (Burnt Sienna) mantle and sooty-led grey plumage
areas. Fig. 11. A naumanni pekinensis, adult male, Hebei, China, 27. IV. 1937 (He Peng, NZMC). Same bird of Figs. 10, 12.
Note that the entire upperwing coverts are typically solidly dark sooty led-grey, as never shown by any nominate “naumanni”.
Fig. 12. F. naumanni pekinensis, adult male from Hebei, China, 27. IV. 1937 (He Peng, NZMC). Same bird of Figs. 10, 11
Note the very richly coloured underparts, more saturated and extensively coloured than in typical nominate “naumanni ” .
A quantitative morphological geographical study from a widely distributed raptor: the Lesser Kestrel Falco naumanni 293
Figure 13. Falco naumanni pekinensis, adult male, Bejing, China, 4.IV.1961 (He Peng, NZMC). Note the typically entirely
grey upperwing coverts of this male from the terra typica of the pekinensis taxon. Fig. 14. F. naumanni pekinensis, adult
male, Bejing, China, 4.1V. 1961 (He Peng, NZMC). Same bird of Fig. 15. Note intensely coloured underparts, with very
saturated colour. Fig. 15. Falco naumanni pekinensis, adult male, Hebei, China, 8. X. 1965 (He Peng, NZMC). Note that
the entire upperwing coverts are typically solidly dark sooty led-grey, as never shown by any nominate naumanni. Fig.
16. F. naumanni pekinensis, adult male, Hebei, China, 27.IV. 1937 (He Peng, NZMC). Same bird of Figs. 10,12 to show
a close up view of the upperwing coverts pattern. Compare to Fig. 17. Fig. 17. F. naumanni pekinensis, ad. male, Spain
(A. Corso, NHM, Tring). Plumage type D according to description given in the text. Note that in many European birds
(ca.20%) the grey on wing coverts is very limited and pretty hard to be seen in the field or even in the hands. Note that the
plumage is paler, less intense and less deep in both the grey of hood and wing-coverts and of the mantle and underparts.
294
Andrea Corso et alii
the percentage within group of the considered cate-
gory within each variable. The analyses showed that
the groups are significantly different. The largest
difference was found between group A and D (Ta-
bles 1, 2).
These groups, possibly, connect in winter/mi-
gration areas, but more study is necessary. Also the
wintering areas for the individual groups are un-
known, but in Fig. 2 we displayed the supposed
wintering areas. The differences between group A
and D is large and both phenotypes differ 86.2%
taken into account the four sampled morphological
features. We did not investigate whether there are
genetic differences among “ naumanni ” and “ peki -
nensis ”, something that surely should be the target
of future studies. The mean differences between
other groups are A vs. B 49 %, A vs. C 75 %, B vs.
C 14.7%, B vs. D 67.8% and C vs. D 39.3 %.
According to the criteria to show discrete cha-
racter differences (Rolan- Alvarez & Rolan, 1995;
Corbet, 1997; Johnson et al., 1999; Garnett & Chri-
stidis, 2007; Rising, 2007; Winker et al., 2007; Ci-
cero, 2010; Remsen, 2010) 86.2 % fall well in the
criteria set by Amadon (1949), Simpson & Roe
(1939) and Mayr (1969) (George Sangster in litte-
ris). We advise that the Chinese population known
under the synonym “ pekinensis ” should be conside-
red valid, despite the apparent intermediate zone bet-
ween this and nominate “ naumanni ” . For the Lesser
Kestrel the same applies as for other polytypic spe-
cies of raptor with a wide breeding distribution area
that it has a certain amount of clinal variation (Fer-
guson-Lees & Christie, 2001). Examples are Com-
mon Buzzard Buteo buteo ssp., Black Kite Milvus
migrans ssp., Saker Falco cherrug and Peregrine
Falco peregrinus ssp. (Vaurie, 1961; Ellis & Garat,
1983; Brosset, 1986; Dixon et al., 2012; White et al.,
2013). The distributional areas are often poorly de-
fined and a large variability applies in subspecies
(Dementiev, 1957; Corso, 2001b; Eastham et al.,
2001; Brichetti & Fracasso, 2003; Eastham & Ni-
cholls, 2005; Karyakin & Pfeffer, 2009; Pfeffer,
2009; Zuberogoitia et al., 2009; Karyakin, 2011; Ro-
driguez et al., 2011). To meet the criteria as set by
Amadon (1949), Simpson & Roe (1939) and Mayr
(1969) most currently recognised subspecies fall
short when assessed on the overlap between pheno-
types.
To simplify, we are faced with two choices: 1)
we consider the currently recognized subspecies of
Variable
Chi squared
df
Pr(>Chi
Kodakhood
53,62
3
<0.01***
percentageofgreyUP
Wcoverts
89,2
3
<0.01***
Table 1 . Showing the Kruskall-Wallis test at all four groups,
considering hood and % of grey upperwing coverts.
Variable
Chi squared
df
Pr(>Chi
colourofupperparts
30,73
3
<0.01***
deep_intensity_of_
underparts
94,79
3
<0.01***
Table 2. Showing GLM test, on the colouration of
upperparts and the intensity of underparts.
a forementioned raptor species and other wide-
ranging raptors as representatives of clinal variation
only. And therefore unworthy of taxonomic rank, in
which case we would not consider “ pekinensis ” as
a valid taxon in light of the intermediate birds found;
2) we believe all these taxa, including “pekinensis ” ,
to be worthy of taxonomic rank. In any case, as the
Chinese population of Lesser Kestrel is always iden-
tifiable, and geographically isolated. It is therefore
worthy of taxonomic rank, which will also help
focus attention on its conservation (Patten, 2015).
ACKNOWLEDGMENTS
For this research, we relied heavily on speci-
mens preserved in public museums, and in thirteen
private collections, either by visiting them or on
loan. Our sincere gratitude goes to (in no specific
order): Mark Adams, Hein and Katrina van Grouw,
Robert Prys-Jones (NHM); He Peng (NZMC); An-
drei Gravilov (IZA); Anita Gamauf (NMW); Ste-
ven van der Mije (NBC); Patrick Bousses, Anne
Previato (MNHN); Giorgio Chiozzi (MCSM); the
late Vittorio Emanuele Orlando (MCT); Rosario
Grasso (MCUCT); Carla Marangoni (MCZR);
Claudio Pulcher (MRSN); Enrico Borgo (MSNG);
Marta Poggesi and FaustoBarbagli (MSNF); Gio-
vanni Boano (SNCa); the late Angelo Priolo
(MCR); Paul Sweet, Mary Le Croy, Matthew
Shanley, Tomas Trombone (AMNH), Kristof Zy-
A quantitative morphological geographical study from a widely distributed raptor: the Lesser Kestrel Falco naumanni 295
skowski (PMNH), SylkeFrahnert (ZMB). Henry
McGhie (The Manchester Museum, Manchester,
England) and Clem Fisher (Liverpool Museum, Li-
verpool, England) supplied additional information.
For help with photos and discussions we are grate-
ful to: Mark Andrews, Aurelien Audevard, Arnoud
B. van den Berg, Claudio Carere, Wouter Faveyts,
Peter Kennerly, Fumin Lei, Zhi-Yun Jia, Jonathan
Martinez, Gerald Oreel, Ran Schols, Manuel
Schweizer, Xuky Summer, Terry Townshend, Pirn
Wolf and Arend Wassink. JJ is grateful to the ‘Sti-
chting P.A. Hens Memorial Fund’ for funding his
ZMB visit. AC and MV are indebted to Ottavio
Janni who was pleasant company in the field (as
part of the MISC group) as in museum studiesand
also improved the final manuscript. Finally we
thank Francesco Angeloni for his help with the sta-
tistics and reading the final manuscript. George
Sangster supplied us with information on taxono-
mic difficulties.
REFERENCES
Adriaens R, Bosnian D. & Elst J., 2010. White Wagtail
and Pied Wagtail: a new look. Dutch Birding, 32:
229-250.
Amadon D., 1949. The seventy-five per cent rule for sub-
species. Condor, 51: 250-258.
Bot S. & Jansen J.J.F.J., 2013. Is Peat Partridge a valid
subspecies of Grey Partridge? Dutch Birding, 35:
155-168.
Brichetti P. & Fracasso G., 2003. Ornitologia Italiana 1.
Gaviidae-Falconidae. Alberto Perdisa Editore, Bolo-
gna, 464 pp.
Brosset A., 1986. Les populations du Faucon Pelerin-
Falco peregrinus Gmelin en Afrique du Nord: un
puzzle zoogeographique. Alauda, 54: 1-14.
Cicero C., 2010. The significance of subspecies: A case
study of Sage Sparrows (Emberizidae, Amphispiza
belli). Ornithological Monographs, 67: 103-113.
Clark W.S., 1999. A Field Guide to Raptors of Europe,
The Middle East and North Africa. Oxford University
Press, Oxford, 371 pp.
Corbet G.B., 1997. The species in mammals. Pp. 341-
356. In Species: M. F. Claridge, H. A. Dawah & M.
R. Wilson (eds.). The Units of Biodiversity. Chapman
and Hall, London.
Corso A., 2000. Less is More: British vagrants, Lesser
Kestrel. Birdwatch, 91: 29-33.
Corso A., 2001a. Notes on the moult and plumages of
Lesser Kestrel. British Birds, 94: 409-418.
Corso A., 2001b. Le Faucon de Barbarie Falco pelegri-
noides. Status en Europe et criteres d’ identification.
Omithos, 8: 164-175
Cramp S. & Simmons K.E.L., 1980. The Birds of the We-
stern Palaearctic. Vol. 2: Hawks to Bustards. Oxford
University Press, Oxford, 695 pp.
Dementiev G.P, 1957. On the Shaheen Falco peregrinus
babylonicus. Ibis, 99: 477-482
Dickinson E.C. & Remsen J.V.Jr., 2013. The Howard and
Moore Complete Checklist of the Birds of the World,
4th. edition. Vol. 1. Aves Press Limited, Eastbourne,
461 pp.
Dixon A., Sokolov A. & Sokolov V., 2012. The subspe-
cies and migration of breeding Peregrines in northern
Eurasia. Falco, 39: 4-9.
Dresser H.E., 1871-1881. A history of the birds of Eu-
rope, including all the species inhabiting the Western
Palaearctic region. Part VI. Dresser, London, 708 pp.
Eastham C.P, Nicholls M.K. & Fox N.C., 2001. Mor-
phological variation of the Saker {Falco cherrug ) and
the implications for conservation. Biodiversity and
Conservation, 10: 1-21.
Eastham C.P. & Nicholls M.K., 2005. Morphometric ana-
lysis of large Falco species and their hybrids with im-
plications for conservation. Journal of Raptor
Research, 39: 386-393.
Ellis D.H. & Garat C.P, 1983. The Pallid Falcon Falco
kreyenborgi is a color phase of the Austral Peregrine
Falcon {Falco peregrinus cassini). Auk, 100: 269-27 1 .
Ferguson-Lees J. & Christie D.A., 2001. Raptors of the
World. Christopher Helm, London, 992 pp.
Forsman D., 1999. The Raptors of Europe and the Middle
East. A Handbook of Field Identification. L.T & A.D.
Poyser, London, 589 pp.
Garnett S.T. & Christidis L., 2007. Implications of chan-
ging species definitions for conservation purposes.
Bird Conservation International, 17: 187-195.
Johnson N.K., Remsen J.V. & Cicero C., 1999. Resolu-
tion of the debate over species concepts in ornitho-
logy: a new comprehensive biologic species concept.
Proceedings International Ornithological Congress,
22: 1470-1482.
Karyakin I.V. & Pfeffer R., 2009. About subspecies and
scientific name of the Saker Falcon in North-Western
Middle Asia. Raptors Conservation, 17: 89-92.
Karyakin I.V., 2011. Subspecies population structure of
the Saker Falcon range. Raptors Conservation, 21:
116-171.
Kodak, 2007. Kodak Color Separation Guides and Gray
Scales / Q14. Kodak, Birmingham.
Mayr E., 1969. Principles of systematic zoology.
McGraw-Hill, New York, 434 pp.
Patten M.A., 2015. Subspecies and the philosophy of
science. The (is’s Auk) and behind 485 (a dot.).
Pfeffer R., 2009. About geographical variances of the
Saker Falcon. Raptors Conservation, 16: 68-95.
296
Andrea Corso et alii
Remsen J.V., 2010. Subspecies as a meaningful taxono-
mic rank in avian classification. Omithologische Mo-
natsberichte, 67: 62-78.
Rodriguez B., Siverio F., Siverio M. & Rodriguez A.,
2011. Variable plumage coloration of breeding Bar-
bary Falcons Falco ( peregrinus ) pelegrinoides in the
Canary Islands: do other Peregrine Falcon subspecies
also occur in the archipelago? Bulletin of the British
Ornithologists’ Club, 131: 140-153.
Rolan- Alvarez E. & Rolan E., 1995. The subspecies con-
cept, its applicability in taxonomy and relationship to
speciation. Argonauta, 9: 1-4.
Ridgway R., 1912. Color Standards and Color Nomen-
clature. Ridgway, Washington, 44 pp.
Rising J.D., 2007. Named subspecies and their signifi-
cance in contemporary Ornithology. Ornithological
Monographs, 63: 45-54.
Sharpe R.B., 1906. Birds. In Gunther A. (Ed.), 1904-
1912. The History of the Collections contained in the
Natural History Departments of the British Museum,
vol. 2. British Museum, London: 79-515.
Simpson G.G. & Roe A., 1939. Quantitative Zoology.
McGraw-Hill, New York, 414 pp.
Snow D.W. & Perrins C.M., 1998. Birds of the Western
Palaearctic: concise edition. Vol. 1 - Non-Passerines.
Oxford University Press, Oxford, 1008 pp.
Swinhoe R., 1870. Zoological notes of a journey from
Canton to Peking and Kalgan. Proceedings Zoologi-
cal Society of London, 38: 427-451.
Vaurie C., 1961. Systematic notes on Palearctic birds.
No. 44 Falconidae: the genus Falco (Part 1, Falco pe-
regrinus and Falco peregrinoides ). American Mu-
seum Novitates, 2035: 1-19.
Vaurie C., 1965. The Birds of the Palaearctic Fauna. A
systematic reference. Non-Passeriformes. H.F. & G.
Whiterby Limited, London, 763 pp.
Warren R.L.M., 1966. Type-specimens of birds in the
British Museum (Natural History). Volume 1. The
British Museum (Natural History), London, 320 pp.
Winker K., Rocque D.A., Braile T.M. & Pruett C.L.,
2007. Vainly beating the air: species-concept debates
need not impede progress in science or conservation.
Ornithological Monographs, 63: 30-44.
White C.M., Cade T.J. & Enderson J.H., 2013. Peregrine
Falcons of the World. Lynx Edicions, Barcelona,
379 pp.
Zuberogoitia I., AzkonaA., Zabala J., Astorkia L., Ca-
stillo I., Iraeta A., Martinez J. A. & Martinez J.E.,
2009. Phenotypic variations of Peregrine Falcon in
subspecies distribution border. In Sielicki J. & Mizera
T. (Eds.), Peregrine Falcon populations - status and
perspectives in the 21st century. European Peregrine
Falcon Working Group & Society for the Protection
of Wild Animals, Warsaw, 295-308.
Biodiversity Journal, 2015, 6 (1): 297-304
Monograph
Current knowledge on the Sicilian tardigrade fauna
Oscar Lisi
Department of Biological, Geological and Environmental Sciences, Section of Animal Biology, University of Catania, Via Androne
81, 95124 Catania, Italy; e-mail: olisi@unict.it
ABSTRACT Based on the literature, and adding personal contribution, the author takes stock of the know-
ledge about the species of limno-terrestrial tardigrades present in Sicily and the main small
islands around it (Aeolian Islands, Ustica, Egadi Islands). In total 111 species are reported:
108 from Sicily (main island), 35 from the Aeolian Islands, 17 from Ustica and 11 from the
Egadi Islands. Two species are new records only for the respective islands, 13 are new records
for the whole studied area, four of which are new also for the Italian fauna. A good 1 3 species
(11.7%) are at present endemic for the studied area. The zoogeographic spread of the 111
Sicilian tardigrade species confirms the modern ideas about tardigrade zoogeography.
KEY WORDS Tardigrada; Sicilian fauna; zoogeography; taxonomy.
Received 25.10.2014; accepted 07.01.2015; printed 30.03.2015
Proceedings of the 2nd International Congress “Speciation and Taxonomy”, May 1 6th- 1 8th 2014, Cefalu-Castelbuono (Italy)
INTRODUCTION
In the Surveys on the Sicilian tardigrade fauna
(Ustica, Aeolian and Aegadean archipelagos in-
cluded) until 2009 allowed to recognize 94 terrestrial
and freshwater species (Arcidiacono, 1962; 1964,
Binda, 1969; 1978; Binda & Pilato, 1969a,b, 1971,
1972, 1984, 1985, 1987; Binda et al., 1980; Pilato,
1969, 1971a, b,c, 1973, 1974, 2009; Pilato & Catan-
zaro, 1988, 1989; Pilato et al., 1982, 1989, 2000).
Though that a number may have appeared high,
the variety of Sicilian environments considered, the
composition of Sicilian tardigrade fauna may be
considered far away from being completely known;
for this reason, I have recently carried out, in col-
laboration with G. Pilato and G. Sabella, new stud-
ies that until now have led us to describe four
species new to science (Pilato et al., 2014; Lisi et
al., 2014). The total number of limno-terrestrial
tardigrade species up to now reported from Sicily
(and surrounding small islands) is 98.
MATERIAL AND METHODS
The current study has been based only on the re-
examination of abundant old material, from the
Pilato and Binda collection (Museum of the Depart-
ment of Biological, Geological and Environmental
Sciences, section of Animal Biology “Marcello La
Greca”, University of Catania), collected and
partially identified in the far past but the results had
remained unpublished; in some cases the old
diagnosis had to be updated revealing new records,
and even new species, which is not surprising con-
sidering that past tardigrade taxonomy was based
on less strict criteria and some wrong convictions,
so that little differences between populations, when
noticed, were more easily attributed to individual
variability within a single species rather than con-
sidered as an indication that the two populations be-
longed to distinct species. About the new records
reported in Table 2, all the data about localities and
samples are the only available, due to the fact that
298
Oscar Lisi
at the time of collection it was not in use to take
note of more detailed information.
All the studied specimens were mounted in
polyvinyl lactophenol. Specific diagnosis was based
on the original descriptions and eventual redescrip-
tions (Plate, 1889; Cuenot, 1929; Marcus, 1936;
Binda, 1971; Pilato & Sperlinga, 1975; Dastych,
1984; Binda & Rebecchi, 1992; Bertolani &
Rebecchi, 1993; Pilato & Binda. 1997/1998; Pilato
et al., 2003; Tumanov, 2006; Pilato et al., 2011) and
on the monograph by Ramazzotti e Maucci (1983);
by comparison, specimens of the Pilato and Binda
collection of the following species were examined:
Hypsibius scabropygus Cuenot, 1929, Diphascon
pingue (Marcus, 1936),/). biggins i Binda, 1971 and
D. chilenense Plate, 1889. All observations and
measurements were made under xlOO oil immersion
using a Leica Phase Contrast Microscope equipped
with a micrometer.
RESULTS
The progression of those studies has led to
recognise other 13 species that represent new
records for the studied area, and four of them are
new records for the Italian fauna as well. The total
number of limno-terrestrial tardigrade species for
that area then raises to 111.
The updated checklist of limno-terrestrial tardi-
grade species present in Sicily (and surrounding
small islands) is reported in Table 1, where the
island in which each species was found is reported;
I found it interesting to take into consideration also
the presence of each species in north Africa, for hav-
ing an idea of the faunal affinity with that region.
In total 111 species are reported (98 already
known, plus 13 new findings): 108 from Sicily
(main island), 35 from the Aeolian Islands, 17 from
Ustica and 11 from the Egadi Islands. Two species
are new records only for the respective islands, 13
are new records for the whole studied area, four of
which are new also for the Italian fauna (Table 2).
A special mention has to be made about the tardi-
grade fauna of North Africa, with which a remarkable
affinity has come out: it shares with Sicily 40 species,
representing a good 36.0% of the Sicilian species.
Thirteen species (11.7%) today result to be
endemic for the studied area. Nine “terrestrial” more
or less recently described, and 4 already reported
freshwater species: Carphania fluviatilis Binda,
1979 (the only freshwater species of the class Het-
erotardigrada), Isohypsibius tubereticulatus Pilato et
Catanzaro, 1990, 1. verae Pilato et Catanzaro, 1990,
and Macroversum mirum Pilato et Catanzaro, 1989.
As regards possible biogeographic evaluations,
the geographic distribution of the 111 Sicilian tar-
digrade species seems to confirm the modem ideas
about tardigrade biogeography. It was very hard in
the past to make biogeographic evaluations about
the species of this group, due to wrong convictions
about species individual variability and poorly strict
criteria for specific diagnosis, and an overestimated
effect of passive dispersal; these had great impact
on the believed geographic distribution of the
species creating great confusion and making very
difficult to study tardigrade species from a biogeo-
graphic point of view. However, thanks to a change
in the evaluation of individual variability and the
taxonomic criteria for specific diagnosis (eg. Pilato,
1975; 1979), as well as a reevaluation of the pos-
sibility of passive dispersal (Pilato, 1979) which
reflected into a reconsideration of the geographic
distribution of the species, many old diagnosis
mistakes have been corrected (and this correction
still continues today), and starting with Mclnnes
(1994) and Pilato & Binda (2001), it is today
universally accepted to consider tardigrade species
from the biogeographic point of view.
Limiting myself to use the available data from
the literature, in which old diagnosis mistakes very
probably still hide, it is possible to notice that tardi-
grade species tend to have a limited geographic
distribution, at the level of zoogeographic region,
not cosmopolitan, or nearly such, as believed in the
far past. In Table 3 the zoogeographic spread of the
111 Sicilian tardigrade species is reported and the
data confirms the above expressed idea.
As regards the relatively high number of
species reported from the literature as present in 7
zoogeographic regions, it must be stressed that
those 1 7 species are mostly represented by species
described in the far past (even about a century ago),
so that there had been all the time, before the
“revolution” of the last decades, for various authors
to report the same species from all over the word;
as a matter of fact, the correction of such diagnosis
mistakes has been in the last decade one of the
great goals of tardigrade taxonomists, and much
still remains to be done.
Current knowledge on the Sicilian tardigrade fauna
299
rank
Sicily
Aeolian
Archipel.
Ustica
Aegadean
Archipel.
North
Africa
CARPHANIIDAE
Carphania fluviatilis Binda, 1978
E
*
ECHINISCIDAE
Biyodelphax tatrensis Weglarska, 1959
*
*
*
Biyodelphax weglarskae Pilato, 1972
*
Cornechiniscus lobatus (Ramazzotti, 1943)
*
Echiniscus blumi Richters, 1 903
*
*
*
Echiniscus trisetosus Cuenot, 1932
*
*
*
Echiniscus mediantus Marcus, 1930
*
*
Echiniscus bisetosus Heinis, 1908
*
*
*
Echiniscus canadensis Murray, 1910
*
*
*
Echiniscus testudo (Doyere, 1 840)
*
*
*
Echiniscus merokensis Richters, 1904
*
*
*
*
Echiniscus granulatus (Doyere, 1 840)
*
*
*
Echiniscus qucidrispinosus (Richters, 1902)
*nr
*
Echiniscus carusoi Pilato, 1972
E
*
*
*
Echiniscus ramazzottii Binda et Pilato, 1969
E
*
Parechiniscus chitonides Cuenot, 1926
*
*
Pseudechiniscus pseudoconifer Ramazzotti, 1943
*
MILNESIIDAE
Milnesium almatyense Tumanov, 2006
NRI
*
Milnesium tardigradum Doyere, 1 840
*
*
*
*
*
EOHYPSIBHDAE
Bertolanius weglarskae (Dastych, 1972)
*
HYPSIBIIDAE
Astatumen trinacriae (Arcidiacono, 1962)
*
*
Bindius triquetrus Pilato, 2009
E
*
Diphascon belgicae Richters, 1911
*
*
Diphascon brevipes Marcus, 1936
*
Diphascon carolae Binda et Pilato, 1969
*
Diphascon chilenense Plate, 1888
NRS
*
Diphascon higginsi Binda, 1971
NRS
*
*
Diphascon nelsonae Pilato, Binda, Bertolani et
Lisi, 2005
*
Diphascon nobilei Binda, 1 969
*
*
Diphascon patanei Binda et Pilato, 1971
*
*
Diphascon pingue Marcus, 1936
NRS
*
*
Diphascon procerum Pilato, Sabella et Lisi, 2014
E
*
Diphascon recamieri (Richters, 1911)
*
Diphascon serratum Pilato, Binda, Bertolani et
Lisi, 2005
E
*
Diphascon scoticum Murray, 1905
*
Table 1. Limno-terrestrial tardigrade species from Sicily; Ranks: NRS = new record for the whole studied area (Sicily and
surrounding islands); NRI = new record also for the Italian fauna; E = endemic. In the geographic region column, “nr”
indicates new record only for the single island/arcipelago. Taxonomy according to Bertolani et al. (2014).
300
Oscar Lisi
rank
Sicily
Aeolian
Archipel.
Ustica
Aegadean
Archipel.
North
Africa
HYPSIBIIDAE
Diphascon serratum Pilato, Binda, Bertolani et
Lisi, 2005
E
*
Diphascon scoticum Murray, 1905
*
Diphascon ziliense Lisi, Sabella et Pilato, 2014
E
*
Hypsibius convergens (Urbanowicz, 1925)
*
*
*
*
Hypsibius conifer Mihelcic, 1938
*
Hypsibius dujardini (Doyere, 1840)
*
*
*
Hypsibius microps Thulin, 1928
*
*
*
Hypsibius pallidoides Pilato, Kiosya, Lisi,
Inshina et Biserov, 2011
NRI
*
Hypsibis pallidus Thulin, 1911
*
Hypsibius ragonesei Binda et Pilato, 1985
E
*
Hypsibius scabropygus Cuenot, 1929
NRS
*
Mixibius saracenus (Pilato, 1973)
*
Mixibius parvus Lisi, Sabella et Pilato, 2014
E
*
Platicrista angustata (Murray 1905)
*
*
RAMAZZOTTIIDAE
Ramazzottius oberhaeuseri (Doyere, 1840)
*
*
Ramazzottius thulini (Pilato, 1970)
*
ISOHYPSIBIIDAE
Doiyphoribius dory p horns (Binda et Pilato, 1969)
*
*
*
Doiyphoribius macrodon Binda, Pilato et Dastych,
1980
*
Doiyphoribius zappalai Pilato, 1971
*
Eremobiotus alicatai (Binda, 1969)
*
*
Hexapodibius micronyx Pilato, 1969
*
Isohypsibius arbiter Binda, 1980
NRS
*
Isohypsibius austriacus (Iharos, 1966)
*
*
Isohypsibius dastychi Pilato, Bertolani et Binda, 1982
*
Isohypsibius deconincki Pilato, 1971
*
*
Isohypsibius elegans Binda et Pilato, 1971
*
*
*
*
*
Isohypsibius granulifer Thulin, 1928
*
Isohypsibius kristenseni Pilato, Catanzaro
et Binda, 1989
*
Isohypsibius longi unguis Pilato, 1974
*
Isohypsibius lunulatus (Iharos, 1966)
*
*
*
Isohypsibius marcellinoi Binda et Pilato, 1971
*
Isohypsibius monoicus Bertolani, 1981
*
Isohypsibius nodosus (Murray, 1907)
*
Isohypsibius pappi (Iharos, 1966)
*
*
Isohypsibius prosostomus Thulin, 1928
*
Isohypsibius reticulatus Pilato, 1973
*
Table 1. Limno-terrestrial tardigrade species from Sicily; Ranks: NRS = new record for the whole studied area (Sicily and
surrounding islands); NRI = new record also for the Italian fauna; E = endemic. In the geographic region column, “nr”
indicates new record only for the single island/arcipelago. Taxonomy according to Bertolani et al. (2014).
Current knowledge on the Sicilian tardigrade fauna
301
rank
Siciliy
Aeolian
Archipel.
Ustica
Aegadean
Archipel.
North
Africa
Isohypsibius ronsisvallei Binda et Pilato, 1969
*
Isohypsibius sattleri Richters, 1902
*
*
*
*
Isohypsibius silvicola (Iharos, 1966)
*
Isohypsibius tetractyloides Richters, 1907
*
Isohypsibius tubereticulatus Pilato et Catanzaro,1990
E
*
Isohypsibius verae Pilato et Catanzaro, 1989
E
*
Parhexapodibius lagrecai (Binda et Pilato, 1969)
*
*
*
*
Pseudobiotus matici (Pilato, 1971)
*
Pseudobiotus kathmanae Nelson, Marley et
Bertolani, 1999
*
Thulinius ruffoi (Bertolani, 1982)
*
Thulinius stephaniae (Pilato, 1974)
*
MACROBIOTIDAE
Macrobiotus diffusus Binda et Pilato, 1987
*
*
*
*
*
Macrobiotus echinogenitus Richters, 1904
*
*
*
Macrobiotus harmsworthi Murray, 1907
*
*
*
*
*
Macrobiotus hufelandi Schultze, 1834
*
*
*
*
*
Macrobiotus insuetus Pilato, Sabella et Lisi, 2014
E
*
Macrobiotus islandicus Richters, 1 904
*
*
Macrobiotus macrocalix Bertolani et Rebecchi,1993
NRS
*
Macrobiotus nuragicus Pilato et Sperlinga, 1975
NRS
*
*
Macrobiotus pallarii Maucci, 1954
*
*
*
Macrobiotus patiens Pilato, Binda, Napolitano et
Moncada, 2000
*nr
*
*
Macrobiotus persimilis Binda et Pilato, 1972
*
*
*
*
Macrobiotus pilatoi Binda et Rebecchi, 1992
NRS
*
Macrobiotus polonicus Pilato, Kaczmarek,
Michalczyk et Lisi, 2003
NRI
*
Macrobiotus sapiens Binda et Pilato, 1984
*
Macrobiotus simulans Pilato, Binda, Napolitano et
Moncada, 2000
*
*
Macrobiotus terminalis Bertolani et Rebecchi, 1993
NRS
*
Paramacrobiotus areolatus (Murray, 1907)
*
*
*
*
Paramacrobiotus csotiensis (Iharos, 1966)
*
Paramacrobiotus richtersi (Murray, 1911)
*
*
*
*
*
Minibiuotus furcatus (Ehrenberg, 1859)
*
*
Minibiotus intermedins (Plate, 1889)
*
*
*
*
Minibiotus pseudofurcatus (Pilato, 1972)
*
*
Minibiotus weinerorum (Dastych, 1984)
NRI
*
Richtersius coronifer (Richters, 1903)
*
*
Tenuibiotus tenuis (Binda et Pilato, 1972)
*
Xerobiotus pseudohufelandi (Iharos, 1966)
*
*
*
*
Table 1. Limno-terrestrial tardigrade species from Sicily; Ranks: NRS = new record for the whole studied area (Sicily and
surrounding islands); NRI = new record also for the Italian fauna; E = endemic. In the geographic region column, “nr”
indicates new record only for the single island/arcipelago. Taxonomy according to Bertolani et al. (2014).
302
Oscar Lisi
rank
Siciliy
Aeolian
Archipel.
Ustica
Aegadean
Archipel.
North
Africa
MURRAYIDAE
Dactylobiotus parthenogeneticus Bertolani, 1982
*
Dactylobiotus dispar (Murray, 1907)
*
Macroversum minim Pilato et Catanzaro, 1989
E
*
Murray on pullari (Murray, 1907)
*
NECOPINATIDAE
Necopinatum mirabile Pilato, 1971
*
Table 1. Limno-terrestrial tardigrade species from Sicily; Ranks: NRS = new record for the whole studied area (Sicily and
surrounding islands); NRI = new record also for the Italian fauna; E = endemic. In the geographic region column, “nr”
indicates new record only for the single island/arcipelago. Taxonomy according to Bertolani et al. (2014).
MILNE SIIDAE
Milnesium almatyense Tumanov, 2006 NRI
Cesaro (Messina) “Portella Femmina Morta”; moss sample.
Ramacca (Catania); moss sample.
HYPSIBIIDAE
Diphascon chilenense Plate, 1889
Belpasso (Catania) “Contrada Milia”; chestnut leaf litter
Diphascon higginsi Binda, 1971
Belpasso (Catania) “Contrada Milia”; chestnut leaf litter
Diphascon pingue (Marcus, 1936)
Belpasso (Catania) “Contrada Milia”; chestnut leaf litter
Hypsibius pallidoides Pilato, Kiosya, Lisi, Inshina et Biserov, 2011 NRI
Belpasso (Catania) “Contrada Milia”; chestnut leaf litter
Hypsibius scabropygus Cuenot, 1929
Isnello (Palermo) “Pizzo Antenna” Madonie Mountains; lichens on tree trunk.
ISOHYPSIBIIDAE
Isohypsibius arbiter Binda, 1980
Belpasso (Catania) “Contrada Milia”; chestnut leaf litter.
Bronte (Catania) “Contrada Rinazzo”; moss sample.
MACROBIOTIDAE
Macrobiotus macrocalix Bertolani et Rebecchi, 1993
Contessa Entellina (Palermo), Contrada Mazzaporro, - Nebrodi Mountains; moss sample
Macrobiotus nuragicus Pilato et Sperlinga, 1975
Mandanici (Messina), Madonie Mountains; moss sample
Macrobiotus pilatoi Binda et Rebecchi, 1992
Madonie Mountains; Sphagnum sample
Macrobiotus polonicus Pilato, Kaczmarek, Michalczyk et Lisi, 2003 NRI
Maletto (Catania) “Sciara St. Venera”; moss sample - Catania; moss sample
Macrobiotus terminalis Bertolani et Rebecchi, 1993
Cesaro (Messina), Mt. Soro (Nebrodi Mountains); moss sample
Minibiotus weinerorum D astych, 1984 NRI
Isnello (Palermo) “Pizzo Antenna” Madonie Mountains; lichens on tree trunk.
Table 2. New records of limno-terrestrial tardigrade species for Sicily and surrounding islands.
NRI = new record also for the Italian fauna.
Current knowledge on the Sicilian tardigrade fauna
303
Palearctic
(of which
endemic)
Present in 2
zoogeographic
regions
Present in 3
zoogeographic
regions
Present in 4
zoogeographic
regions
Present in 5
zoogeographic
regions
Present in 6
zoogeographic
regions
Present in 7
zoogeographic
regions
Total
40(13)
19
15
5
11
4
17
111
36.0% (11.7%)
17.1%
13.5%
4.5%
9.9%
3.6%
15.3%
100%
Table 3. Zoogeographic spread of the limno-terrestrial tardigrade species of Sicily.
CONCLUSIONS
As it can be seen, a limited study of old material
may reveal new records and even new species, also
with the change of old diagnoses, not correct in the
light of the modem criteria; this, as well as improv-
ing knowledge in general, allows to go ahead in the
progress of tardigrade taxonomy and biogeography.
The current study, thanks to the promising results
obtained, still goes on and intends to proceed with
the correction of very old literature mistakes (which
affect the correct knowledge of tardigrade biod-
iversity and species geographic distribution), enrich
faunistic and biogeographic knowledge, and put in
evidence the variety and eventual peculiarity of
Sicilian tardigrade fauna, with repercussions on
general knowledge of Sicilian biodiversity.
REFERENCES
Arcidiacono R., 1962. Contribute) alia conoscenza dei
Tardigradi dei Monti Nebrodi e descrizione di una
nuova specie di Itaquascon. Bollettino delle Sedute
dell’Accademia Gioenia di Scienze Naturali di
Catania, S. IV, 3-7: 123-134.
Arcidiacono R, 1964. Secondo contributo alia cono-
scenza dei tardigradi dei Monti Nebrodi. Bollettino
delle Sedute delTAccademia Gioenia di Scienze
Naturali di Catania, S. IV, 3: 187-203.
Bertolani R., Guidetti R., Marchioro T., Altiero T.,
Rebecchi L. & Cesari M., 2014. Phylogeny of Eutar-
digrada: new molecular data and their morphological
support lead to the identification of new evolutionary
lineages. Molecular Phylogenetics and Evolution, 76:
110-126.
Bertolani R. & Rebecchi L., 1993. A revision of the
Macrobiotus hufelandi group (Tardigrada, Macrobio-
tidae), with some observations on the taxonomic
characters of eutardigrades. Zoologica Scripta, 22:
127-152.
Binda M.G., 1969. Nuovi dati su Tardigradi di Sicilia con
descrizione di due nuove specie. Bollettino delle
Sedute dell’Accademia Gioenia di Scienze Naturali
di Catania, S. IV, 9: 623-633.
Binda M.G., 1971. Su alcuni Tardigradi muscicoli del
Nord- Africa. Bollettino delle sedute dell'Accademia
Gioenia di Scienze Naturali in Catania, 10: 759-765.
Binda M.G., 1978. Risistemazione di alcuni Tardigradi
con l’istituzione di un nuovo genere di Oreellidae e
della nuova famiglia Archechiniscidae. Animalia, 5:
307-314.
Binda M.G. & Pilato G., 1969a. Su alcune specie di
Tardigradi muscicoli di Sicilia. Bollettino delle
Sedute dell’Accademia Gioenia di Scienze Naturali
di Catania, S. IV, 10: 159-170.
Binda M.G. & Pilato G., 1969b. Ulteriore contributo alia
conoscenza dei Tardigradi di Sicilia con descrizione
di due nuove specie. Bollettino delle Sedute dell’
Accademia Gioenia di Scienze Naturali di Catania,
S. IV, 10:205-214.
Binda M.G. & Pilato G., 1971. Nuovo contributo alia
conoscenza dei Tardigradi di Sicilia. Bollettino delle
Sedute dell’Accademia Gioenia di Scienze Naturali
di Catania, S. IV. 10: 896-909.
Binda M.G. & Pilato G., 1972. Tardigradi muscicoli
di Sicilia (IV Nota). Bollettino delle Sedute dell’
Accademia Gioenia di Scienze Naturali di Catania,
S. IV, 11:47-60.
Binda M.G. & Pilato G., 1984. Macrobiotus sapiens ,
nuova specie di Eutardigrado di Sicilia. Animalia, 1 1 :
85-90.
Binda M.G. & Pilato G., 1985. Hypsibius ragonesei,
nuova specie di Eutardigrado di Sicilia. Animalia, 12:
245-248.
Binda M.G. & Pilato G., 1987. Tardigradi dell’Africa.
V. Notizie sui Tardigradi del Nordafrica e descrizione
della nuova specie Macrobiotus diffusus. Animalia,
14: 177-191.
Binda M.G., Pilato G. & Dastych H., 1980. Descrizione
di una nuova specie di Eutardigrado: Doryphoribius
macrodon. Animalia, 7: 22-27 .
Binda M.G. & Rebecchi L., 1992. Precisazioni su
Macrobiotus furciger Murray, 1907 e descrizione di
304
Oscar Lisi
Macrobiotus pilatoi n. sp. (Eutardigrada, Macrobio-
tidae). Animalia, 19: 101-109.
Cuenot L., 1929. Description d'un tardigrade nouveau de
la faune francaise. Archives d'anatomie microsco-
pique, 25: 121 -125.
Dastych H., 1984. The Tardigrada from Antarctic with
descriptions of several new species. Acta Zoologica
Cracoviensia, 27: 377-436.
Lisi O., Sabella G. & Pilato G., 2014. Mixibius parvus
sp. nov. and Diphas con ( Diphascon ) ziliense sp. nov.,
two new species of Eutardigrada from Sicily. Zootaxa
3802, 4: 459-468.
Marcus E., 1936. Tardigrada. Das Tierreich, 66: 1-340.
Mclnnes S.J., 1994. Zoogeographic distribution of ter-
restrial/freshwater tardigrades from current literature.
Journal of Natural History, 28: 257-352.
Pilato G., 1969. Su un interessante Tardigrado esapodo
delle dune costiere siciliane: Hexapodibius micronyx
n. gen. n. sp. Bollettino delle Sedute delTAccademia
Gioenia di Scienze Naturali di Catania, S. IV, 9: 619—
622.
Pilato G., 1971a. Necopinatum mirabile n. gen. n. sp.,
interessantissimo Eutardigrado incertae sedis. Bol-
lettino delle Sedute delTAccademia Gioenia di
Scienze Naturali, Catania, S. IV, 10: 861-867.
Pilato G., 1971b. Tardigradi delle acque dolci siciliane.
Nota prima. Bollettino delle Sedute delTAccademia
Gioenia di Scienze Naturali di Catania, S. IV, 11:
126-134.
Pilato G., 1971c. Su una nuova specie di Doryphoribius
(Eutardigrada, Hypsibiidae) e considerazioni sulla
posizione filogenetica del genere. Bollettino di
Zoologia: 38: 145-149.
Pilato G., 1973. Tardigradi delle acque dolci siciliane.
Nota seconda. Bollettino delle Sedute delTAcca-
demia Gioenia di Scienze Naturali di Catania, S. IV,
12: 177-186.
Pilato G., 1974. Tardigradi delle acque dolci siciliane.
Terza nota. Animalia, 1: 135-244.
Pilato G., 1975. On the taxonomic criteria of the Eutar-
digrada. Memorie delTIstituto Italiano di Idrobiolo-
gia Marco De Marchi, Pallanza, 32: 277-303.
Pilato G., 1979. Correlations between cryptobiosis and
other biological characteristics in some soil animals.
Bollettino Zoologico, 46: 319-332.
Pilato G., 2009. Bindius triquetrus gen. nov. sp. nov.
(Eutardigrada, Hypsibiidae) from Sicily (Italy).
Zootaxa, 2058: 62-68.
Pilato G. & Binda M.G., 1997-1998. A comparison of
Diphascon ( D .) alpinum Murray, 1906, D. (D.)
chilenense Plate, 1889 and D. ( D .) pingue Marcus,
1936 (Tardigrada), and a description of a new
species. Zoologischer Anzeiger, 236: 181-185.
Pilato G. & Binda M.G., 2001. Biogeography and limno-
terrestrial tardigrades: Are they truly incompatible
binomials? Zoologischer Anzeiger, 240: 511-516.
Pilato G. & Catanzaro R., 1988. Macroversum mirum
n. gen. n. sp., nuovo Eutardidgrado (Macrobiotidae)
dei Monti Nebrodi (Sicilia). Animalia, 15: 175—
180.
Pilato G. & Catanzaro R., 1989. Tardigradi delle acque
dolci siciliane. IV. lsohypibius tubereticulatus e
Isohypsibius verae , due nuove specie di Eutardigradi
dulcacquicoli di Sicilia. Animalia, 16: 81-88.
Pilato G., Bertolani R. & Binda M.G., 1982. Studio degli
Isohypsibius del gruppo elegans (Eutardigrada,
Hypsibiidae) con descrizione di due nuove specie.
Animalia, 9: 185-198.
Pilato G., Catanzaro R. & Binda M.G., 1989. Tardigradi
delle acque dolci siciliane. V Nota. Animalia, 16:
121-130.
Pilato G., Binda M.G., Napolitano A. & Moncada E.,
2000. The specific value of Macrobiotus coronatus
De Barros, 1942, and description of two new species
of the harmsworthi group (Eutardigrada). Bollettino
delle Sedute delTAccademia Gioenia di Scienze
Naturali di Catania, 13: 103-120.
Pilato G., Sabella G. & Lisi O., 2014. Two new tardi-
grade species from Sicily. Zootaxa, 3754, 2: 173—
184.
Pilato G., Kaczmarek L., Michalczyk L. & Lisi O., 2003.
Macrobiotus polonicus, a new species of Tardigrada
from Polan (Eutardigrada: Macrobiotidae, " hufelandi
group"). Zootaxa, 258: 1-8.
Pilato G. & Sperlinga G., 1975. Tardigradi muscicoli di
Sardegna. Animalia, 2: 79-90.
Pilato G., Kiosya Y., Lisi O., Inshina V. & Biserov V.,
2011. Annotated list of Tardigrada records from
Ukraine with the description of three new species.
Zootaxa 3123: 1-31.
Plate L., 1889. Beitrage zur Naturgeschichte der Tardi-
graden. Zoologische Jahrbiicher. Abteilung fur
Anatomie, 3: 487-550.
Ramazzotti G. & Maucci W., 1983. II Phylum Tardi-
grada. Memorie delTIstituto Italiano di Idrobiologia
Marco De Marchi, 41: 1-1012.
Tumanov D.V., 2006. Five new species of the genus
Milnesium (Tardigrada, Eutardigrada, Milnesiidae).
Zootaxa, 1122: 1-23.
Biodiversity Journal, 2015, 6 (1): 305-308
Monograph
On the presence of Campodea majorica sicula Conde, 1 957
(Diplura Campodeidae) in the "Abisso della Pietra Selvaggia"
cave (Mount Pellegrino, Palermo, Italy)
Alessandro Marietta 1,2 *, Giuseppe Nicolosi 2 &Tiziana Grech 2
'Department of Biological, Geological and Environmental Sciences - section ofAnimal Biology “ M . La Greca”. University of
Catania, via Androne 81, 95124 Catania, Italy; entail: am arlet@ unict.it
2 Centro Speleologico Etneo, via Valdisavoia 3, 95123 Catania, Italy; entail: gnicolosi@hotmail.it, tizianagrech@hotmail.it
* C o rre sp o n d in g author
ABSTRACT W e report for the first tim e the presence of Cdlfipoded UldjoriCd sicillci Conde, 1957 (Insecta,
Diplura, Campodeidae) in the "Abisso della Pietra Selvaggia", a vertical karst cave situated in
the southern slope of Mount Pellegrino, adjacent to the city of Palermo (Sicily). This hypogean
subspecies is considered endemic of Sicily and up to now it was known only forthe “A ddaura
Caprara” cave, located at the opposite slope (north-east) of Mount Pellegrino. During a spe-
leological excursion in the "Abisso della Pietra Selvaggia" cave, organized by “Centro Spe-
leologico Etneo” (Catania, Italy), 14 specimens of this subspecies were collected in the bottom
of the cave, at -170 m. The bottom is one of the few humid areas of the cave, whereas the rest
is very dry, dusty and apparently without Diplura. In addition to C. ITlClj OVicd siculd, currently
are known the following C. fflCljoricd subspecies, all hypogean: C. Hldjoricd llUljoricCl C o n d e ,
1 9 5 5, C. mdjoricd intcrjectd Conde, 1955, both endemic of some caves of Majorca Island
(Balearic Islands, Spain) and C. ffldjoricd VdleYltUld Sendra et Moreno, 2004, found inside 7
caves located in the karstic area of M ount M onduver and Sierra de Corbera (SE of Valencia,
Spain).
KEY WORDS Diplura; b io s p e le o lo g y ; karst cave; hypogean fauna; Sicily.
Received 22.07.2014; accepted 30.10.2014; printed 30.03.20 1 5
Proceedings of the 2nd Interna tional Congress “Speciation and Taxonomy”, M ay 1 6 th - 1 8 th 2014, Cefalu - Caste lb uono (Italy)
INTRODUCTION
Diplura is a poorly investigated order wich con-
tains small and wingless species. They are unpig-
mented and eyeless. The antennae are long and
moniliformis.The abdomen ends with a pair of cerci
that can be long and thin or short and pincer-like.
The majority of the species usually are 2-5 mm long,
although some species can reach 50 mm. There are
about 94 known species in Italy, 19 of which occur
in Sicily (Thibaud, 2013). The most part of the
species belong to the Campodeidae family, that
includes epigean and hypogean species.
Campodea majorica sicula Conde, 1957 wasde-
scribed on 4 specimens collected by P. Strinati on
21 August 1 956 in the A ddaura Caprara III cave
(Conde, 1 957), located in the NE slope of Mount
Pellegrino, near the city of Palermo (Sicily). Until
now this was the only record in Sicily for the taxon.
In the present paper we report for the first time the
306
Alessandro Marletta et alii
presence of this subspecies also in the "Abisso della
Pietra Selvaggia", a vertical karst cave situated in
the southern slope of Mount Pellegrino.
U p to now, in addition to C. ITldjoricd siculd,
other three subspecies are known: C. tl'ldjoricd tTld-
jorica Conde, 1955 , C. mdjoricd interjectd Conde,
1955, both endemic of some caves of M ajorca Island
(Balearic Islands, Spain) and C. lTldjovicd VdleYltUld
Sendra et Moreno, 2004, found inside 7 caves loc-
ated in the karstic area of Mount Monduver and
Sierra de Cor her a (SE of Valencia, Spain). All these
taxa are strictly hypogean (Conde, 1 955b, 1 957;
Sendra, 1985, Sendra & Moreno, 2004) .
MATERIAL AND METHODS
During the sampling in the "Abisso della
Pietra Selvaggia" cave, 14 specimens of this sub-
species were collected by hand and preserved in
7 0 % ethanol.
The “Abisso della Pietra Selvaggia" cave is loc-
ated in the Mount Pellegrino massif, within the
"Riserva Naturale Orientata Regionale Monte Pel-
legrino" (managed by Rangers d'ltalia), in the
North-West Sicily, at the northern side of Palermo
(Figs. 1, 2). The Reserve covers about 1020 ha and
was created in 1996 to protect the Mount Pellegrino
massif and the "Tenuta Reale della Favorita". It is
also a Site of Community Importance (SCI)
ITA020014. Mount Pellegrino is a car bona tic mas-
sif (606 m a.s.l.) made up of rocks originated in
shallow seas from Trias to Eocene. The mount is
subjectto k ars t p h e n o m e n a and counts 134 caves
of both marine and karst origin.
The "Abisso della Pietra Selvaggia", is a ver-
tical karst cave situated in the southern slope of
M ount Pellegrino at an elevation of 425 m a.s.l. It
is 171 m deep (Fig. 3) and itconsists of a sequence
of four shaft, respectevely of 31 m, 6 m, 38 m, 62
m (Mannino, 1 985).
The specimens were collected in the bottom of
the cave (-157/-171 m), on the moist soil, among
the stones and near stalagmites (Fig. 4). The hot to m
is one of the few humid areas of the cave, the rest
is very dry, dusty and apparently without Diplura.
The specimens were examined in laboratory
using a Feica M 205A stereom icroscope equipped
with a Feica DFC 450 digital camera and a multi-
focus image acquisition software (Feica Applica-
tion Suite v.4.2.0). Moreover some macrophotos
have been made on-site using digital SFR camera.
Taxonomical reference are based on the checklist
of "Fauna Europaea", version 2.6 (Thibaud, 2013).
ABBREVIATIONS: ma: medial anterior m a c -
rochaeta; la: lateral anterior macrochaeta; lp : lat-
eral posterior macrochaeta.
Figure 1. Location of Mount Pellegrino (Palermo, Sicily). Figure 2. Satellite image of Mount Pellegrino, with location of
the investigated caves. Red square: Addaura Caprara; yellow square: Abisso della Pietra Selvaggia (from Google Earth)
Campodea majorica sicula (Diplura Campodeidae) in theAbisso della Pietra Selvaggia” cave (Mount Pellegrino, Italy) 307
Figure 3. Longitudinal section and plan of“Abisso della Pietra Selvaggia” cave (M . Panzica survey, from M annino, 1985).
Figure 4. Bottom of “Abisso della Pietra Selvaggia” (Photo by F. Fiorenza).
Figure 5. C. lliajoriCQ. sicula ad ult female from “Abisso della
Pietra Selvaggia”, body length 8 mm (Photo by F. Fiorenza).
Campodea majorica sicula Conde, 1957
Examined material. Italy, Sicily, Palermo,
“Abisso della Pietra Selvaggia” cave,
38°09’33.2”N ; 1 3 °2 1 ’ 2 5 .6 ”E ; 16. III. 2014, 3 fe-
males, 2 males; 11. V. 2014, 5 females and 4 males,
Marietta A., Nicolosi G., Grech T. legit.
Description. Body length: 7mm male; 8 mm fe-
rn ale (Fig. 5). Head (Fig. 6): Antennae with 41-43
antennom eres, cupuliform organ with 9-12 sensilla.
Insertion line of antennae bordered by 3 + 3 macro-
chaetae. Sensillum of third antennomere bacilliform
and in latero -stern al position. Thorax: the typical
notal macrochaetae distribution is similar to the
o ther C. majorica subspecies with 3 + 3 (ma, la, lp)
pronotal macrochaetae, 3 + 3 (ma, la, lp) mesonotal
macrochaetae and 1 + 1 (nr a) nretanotal macro-
chaetae (Fig. 7). Notal macrochaetae and setae
similar to the other C. majorica subspecies, but
slightly longer and thinner. The posterior-marginal
setae are thick and crenellated. Mesonotal macro-
chaetae lp/ma ratio is 1.74-2.21. Abdomen (Fig. 8):
urotergite VI with 1 + 1 la macrochaetae, lp macro-
chaetae absent. Urotergite VII with 1 + 1 la macro-
chaetae and 1 + 1 lp macrochaetae. Urotergite VIII
with la macrochaetae absent and 3 + 3 lp macro-
chaetae. Posterior margin of urosternite I of males
with 2-3 rows of glandular setae. Cerci 1.5 times
longer than body length, with about 20 elongated
articles with long macrochaetae and setae (Conde,
1957; Send r a & Moreno, 2004).
308
Alessandro Marletta et alii
Remarks. According to Conde (1948, 1955a,
1957) and Sendra & Moreno (2 004), C. maj OHCd
subspecies, together with C. CymeCl Conde, 1948
and C. blandinae Conde, 1948 (both hypogean
species endemic of Corsica island), are considered
closely related with C. gVCISSi Silvestri, 1912, an
epigean species w id e spread in the western Mediter-
ranean area (Italy mainland, Corsica, Sicily,
Tunisia, Algeria and north-eastern of Iberian
Peninsula) (Silvestri, 1912). They form a group of
related species that share following common
characters: elongated appendages, hypertrophic
cupuliform organ of antennae, robust and short
notal m ac ro c h ae tae , lateral posterior (lp) macro-
chaetae absent or reduced, short clothing setae,
body surface densely covered with thin micro-
denticles and abdomen with 1 + 1 lateral anterior
(la) macrochaetae from urotergites V or VI, 1 + 1
lp from VII and 3 + 3 lp in VIII. The phylogenetic
history of these species and their adaptation pro-
cess in the hypogean environment are still uncer-
tain, thus more investigation would be necessary,
also with mitochondrial DNA-based analysis.
ACKNOWLEDGEMENTS
We would like to thanks Dr. Alberto Sendra
(Valencia, Spain) for his precious advices on iden-
tification of this taxon; Dr. Salvatore Palascino
(Director of Mount Pellegrino Natural Reserve)
Figure 6. Campodea
majorica sicula-. head
(dorsal view ).
Figure 7. Thorax (dorsal
view ).
Figure 8. Abdomen,
urotergites V-X (for ab-
breviations see text).
for allowing us to conduct this study. And also the
speleologists of “Centro Speleologico Etneo” for
their technical support into the cave.
REFERENCES
Conde B ., 1 948. Campodeides hypoges de Corse. Bul-
letin de la Societe des Sciences de Nancy, N. S. tome
V II, no 3 .
Conde B., 1 955 a. Campodeides cavernicoles des
Baleares. Notes biospeologiques, 9: 121-132.
Conde B ., 1 955b. Sur la faune endogee de Majorique
(P e n ic ilia te s , Protoures, Diploures Campodeides,
P a lp ig ra d e s ) . Bulletin du Museum d' Histoire
naturelle, 2e serie, 26: 674-677.
Conde B., 1957. Campodeides recoltes en Sicile par P.
Strinati. Fragmenta Entomologica, II (14): 137-141.
Mannino G., 1 9 8 5. L e grotte di Monte Pellegrino. Edizioni
Etna Madonie del Club Alpino Siciliano: 212-225.
Sendra A ., 1985. Campodeidos caver nicolas de Baleares.
Endins, 10-11: 33-35.
Sendra A. & Moreno A., 2004. El subgenero CttlTipodcCl
s.str. en la Peninsula Iberica (Hexapoda: Diplura:
C am podeidae). Boletin Sociedad Entomologica
Aragonesa, 35: 1 9-3 8.
Silvestri, F. 1912. Contribuzione alia conoscenza dei
Campodeidae (Thysanura) d’Europa. Bolletino del
Laboratorio di Zoologia generale ed agraria del R.
Istituto superiore agrario di Portici, 6: 110-147.
Thibaud J.M., 2013. Fauna Europaea: Diplura, Cam-
podeidae. Fauna Europaea version 2.6, http:// www.
faunaeur.org
Biodiversity Journal, 2015 , 6 ( 1 ): 309-322
Monograph
Remarks on the composition of the Auchenorrhyncha fauna
in some moist areas in Southern Apulia (Italy)
Adalgisa Guglielmino 1 & Christoph Buckle 2
'Department ofAgriculture, Forests, Nature and Energy, University of Tuscia, Viterbo, Italy
"Neckarhalde 48, D-72070 Tubingen. Germany
Corresponding author, e-mail: guglielm@ unitus.it
ABSTRACT A list of 84 Auchenorrhyncha species collected from field excursions in the province of Lecce
(Southern Apulia) in June 2011 and April 2012 is given. Prevalently three areas were studied:
the Regional Natural Park “Bosco e Paludi di Rauccio”, the Protected Oasis “LaghiAlimini”
and the State Natural Reserve “Le Cesine”. Four species ( DdphciX lYlCridiOYlQlis (Haupt, 1924),
Delphacodes capnodes (Scott, 1 8 7 0), Parapotes reticulatus (Horvath, i 897)and Calamotettix
taeniatus (Horvath, 19 11) are recorded for the first time for Italy, five (Steiiokslisici CUlgUStCl
Ribaut, 1934, Eludes basUmeCl ( Germar, 1821), ChlorionCL glciucescens F ieb er, 1 866, Hecalus
Storai ( Lindberg, 1 936) and Melillaia desbrochersi ( Lethierry, 1 899) are new records for the
Ape n nine Peninsula (“S” in the checklist of the Italian fauna) and 26 new for Apulia. For some
species of special interest their ecology and distribution is discussed. The investigated areas
are of high relevance for nature conservation as they constitute small relics of formerly vastly
extended coastal marshes, where several stenotopic Auchenorrhyncha species occur, associated
particularly with moist vegetation. Interesting is a group of taxa that are known only from the
Balkan region and South Italy. Possibly the isolated occurrence of some other A uchenorrhy ncha
taxa in Apulia is connected rather with the Balkan Peninsula than with Central Europe.
KEY WORDS Faunistics; Ecology; Biogeography.
Received 15.07.2014; accepted 30.09.2014; printed 30.03.2015
Proceedings of the 2nd International Congress “Speciation and Taxonomy”, M ay 1 6 th - 1 8 th 2014, Cefalu-Castelbuono (Italy)
INTRODUCTION
As the knowledge on the distribution of many
species of Auchenorrhyncha in Italy is still rather
fragmentary, and recent data for the southern
regions of the peninsula are almost completely
lacking, it may be useful to publish some data deri-
ving from two sampling trips in June 2011 and April
2012, respectively, in some moist areas in the
province of Lecce in Southern Apulia (Fig. 1).
Especially three zones of notable naturalistic
importance were investigated: the Regional Natural
Park “Bosco e Paludi diRauccio”,the State Natural
Reserve “Le Cesine” and the Protected Oasis
“L ag h i A lim in i” .
The Regional Natural Park “Bosco e Paludi di
Rauccio” (Figs. 2-5) comprises many different
habitats: forest of QueVCUS USX L . (residue of the
“Foresta di Lecce”, a forest area that in the M iddle
Ages extended between Lecce, the Adriatic coast,
Otranto and Brindisi), a swampy area named
Speech i a della Milogna, small ponds and moist
areas, two coastalbasins (Id um e and Fetida), sandy
seashore, som e zones of M editerranean m aquis and
garigue, ruderal areas and pastures. The State Nat-
ural Reserve “Le Cesine” (Figs. 6, 7) is an area of
extreme envir on mental value. Even if prevalently a
3 10
Adalgisa Guglielmino & Christoph Buckle
humid area, it includes in addition a large variety of
habitats and transitional zones, which create a vast
ecological mosaic. A part fro m extended reed areas,
numerous canals, swamps and marshes and the
basins of Pantano Grande and Salapi, there are
many other habitats as pine forest, Mediterranean
maquis, QuerCUS il6X forest and ruderal areas. The
reserve includes 620 ha, defined as “moist zone of
international value” (RAMSAR convention, 197 1);
out of these 620 ha, 34 8 are “natural reserve of ani-
malrepopulation” administrated by the W W F-ltaly.
The Alim ini lakes (Fig. 8) consist of two basins:
Alim ini Grande and Alimini Piccolo, named also
Fontanelle, with the former being a salt water, the
latter a fresh water lake. The oasis includes valuable
areas of M editerranean m aquis and costal retrodunal
lagoons of great naturalistic interest. The protected
area is one of the most important natural sites of the
Salento region, with an ecosystem rich of plant and
animal species. It constitutes a “Zone of Special
Protection” (ZPS), proposed as Site of European
Community Importance (pSIC). The protected
Oasis of Alimini lakes is a very important place
where birds can rest and winter.
MATERIAL AND METHODS
The samplings were carried out in June 2011
and April 2012 at 18 localities (two of them
s am pled tw ice).
Figure 1. Map of the investigated areas in South Apulia. 1 =
Bosco e PaludidiRauccio; 2 = Le Cesine; 3 = LaghiAlimini.
We applied two collection methods: a) by en to -
mological net and aspirator, b) directly by sight of
single specimens by means of the aspirator.
List of collecting sites
In order to facilitate the comparison of data in
our different papers on the Italian A uchenorrhyncha
fauna we maintain the number system of collecting
localities applied already in other publications.
- St. 5 5 2: Torre Chianca, Bosco di Rauccio;
N 40°27’52.4” E 1 8 ° 1 0 ’ 0 0 .4 ” ; 3 m; 1 9/06/20 1 1;
ruderal area with shrubs of PistdCid lentisCUS L . an d
Phillyrea l . (Fig. 2 ).
- St. 5 5 3: Torre Chianca, Bosco di Rauccio;
N40°28’11.8” E18°10’10.7”; 2 m; 20/0 6/2 011;
Phragmites australis (c a v.) T rin . on th e m arg in of a
field and moist areas with JuYlCUS L., BolboSChoenUS
maritimus (L.) Palla, Cyperaceae (Fig. 3).
- St. 554: Torre Chianca, Bosco di Rauccio;
N 4 0 °2 8 ’ 0 2 .0 ” E18°10’19.8”; 6 m; 20/06/20 1 1;
margin of Quercus ilex- forest, Pistacia lentiscus ,
Phillyrea and open areas.
- St. 555: Torre Chianca, south of Bosco di
Rauccio; N40°27’09.0” E 1 8 ° 1 1 ’ 5 7 .6 ” ; 3 m;
20/06/2011; moist area with CdreX L ., JuriCUS,
Cyperaceae, Poaceae (Fig. 4).
- St. 556: Torre Chianca, Bosco di Rauccio;
Speech i a della Milogna; N40°28’09.6”
E 1 8° 1 0’29.9 ”; 3 m; 2 1 /06/20 1 1; moist area with
Bolboschoenus (Asch.) Palla in H allier & Brand,
Carex, Tamarix l . , Phragmites a d a n s , June us. and
dry ruderal area.
- St. 557: Torre Chianca, Bosco di Rauccio;
N40°27’56.5” E 1 8 ° 1 0 ’ 0 7 . 8 ” ; 7 m; 2 1 /06/20 1 1;
forest of Quercus ilex with Pistacia lentiscus ,
Phillyrea , Clematis l ., Hedera l . etc .
St. 558: Torre Chianca, Bacino Idume;
N40°28’08.0” E 1 8 ° 1 1 ’2 1 .8 ”; 5 m; 2 1/06/20 1 1;
vegetation near the sea and the basin with ElymUS
l ., Phragmites , etc.
- St. 5 5 9: L ag h i A lim in i, no rth o f L ag o Grande;
N40°12’31.6” E 1 8°25 ’4 1 .4”; 10 m; 22/06/20 1 1;
moist area with J uncus, Phragmites, Cyperus l .
St. 560: Laghi Alimini, L ago Piccolo;
N40°10’50.9” E 18°27’04.7”; 4 m; 22/06/20 1 1;
m oist area w ith CdreX ( Fig. 8).
- St. 561: Torre Chianca, south of Bosco di
Rauccio; N40°27’08.9” E 1 8 ° 1 1 ’ 5 7 .4 ” ; 6 m;
Remarks on the composition of the Auchenorrhyncha fauna in some moist areas in Southern Apulia (Italy)
3 1 1
Figure 2. Bosco e Paludi di Rauccio: St. 5 52. Figure 3. Bosco e Paludi di Rauccio: St. 55 3. Figure 4. South of Bosco e
Paludi di Rauccio: St. 555. Figure 5. South of Bosco e Paludi di Rauccio: St. 561. Figure 6. Le Cesine: St. 563. Figure 7.
Le Cesine: St. 562. Figure 8. LaghiAlimini: St. 560. Figure 9. Porto Badisco: St. 564.
3 1 2
Adalgisa Guglielmino & Christoph Buckle
22/06/2011; open dry stony area, Phillyrea ,
P o a c e a e , Ca VeX , RubllS L., thistles (Fig. 5).
St. 5 62: Natural Reserve “Le Cesine”;
N40°21’16.7” E 1 8 ° 2 0 ’ 2 6 .2 ” ; 4 m; 23/06/20 1 1;
shore of lagoon, Phragmites , Bolboschoenus ,
Carex, Tamarix (Fig. 6 ) .
St. 563: Natural Reserve “Le Cesine”;
N 4 0 °2 1 ’ 0 3 .9 ” E 1 8°2 1 ’05 .4”; sea level;
23/06/20 1 1; shore of lagoon, Bolboschoenus ,
Carex, Cyperaceae (Fig. 7).
- St. 5 64: between Porto Badisco and Santa
Cesarea; N 4 0 ° 0 4 ’ 0 8 .0 ” E 1 8 °2 8 ’44 .8 ” ; 45 m;
24/0 6/20 1 1; dry rocky area and srn all pine forest w ith
Brachypodium p . B eau v., Carex, Poaceae (Fig. 9).
- St. 622: Natural Reserve “Le Cesine”; sea
level; 1 7/04/20 1 2; forest, shore of lagoon, shrubs,
herbaceous vegetation.
- St. 623: West ofNatural Reserve “Le Cesine”;
N40°20’50.3” E 1 8 ° 1 9 ’3 3 .8 ” ; 20 m; 1 7/04/20 1 2;
olive grove, herbaceous vegetation with prevalently
Fabacae, Poaceae.
St. 624: road S. Cataldo - Frigole;
N 40°23’38.6” E 1 8 ° 1 5 ’ 2 3 .4 ” ; 20 m; 1 7/04/20 1 2;
open dry area with Poaceae and maquis vegetation.
- St. 625: Torre Chian ca, Bosco di Rauccio;
N40°27’23.6” E 1 8 ° 1 0 ’ 0 0 . 7 ” ; 6 m; 1 8/04/20 1 2;
meadow, herbaceous vegetation.
- St. 626: Torre Chianca, Bosco di Rauccio;
N40°27’56,5” E18°10’07.8”; 7 m; 18/0 4/2 012;
forest of Quercus ilex w ith Pistacia lentiscus,
Phillyrea, Clematis, Hedera e tc .
- St. 627: Torre Chianca, south of Bosco di
Rauccio; N40°27’09.0” E 1 8 ° 1 1 ’ 5 7 .6 ” ; 3 m;
1 8/0 4/20 1 2; moist area with Mentha L., CareX,
J UnCUS, Cyperaceae, Poaceae.
- St. 628: coast between Frigole and Torre
Chianca; N40°2 7 , 3 3.8 ” E 1 8 ° 1 2 ’ 5 0 .9 ” ; 2 m;
18/04/2012; open area near seashore with Poaceae,
CareX, herbaceous vegetation.
in the - List of collected specimens" are indicated
for each species: the collection locality and in
parentheses the number of males, females and (if
present) nymphs, separated by semicola, respecti-
vely. For some species brachypterous (b) and
macropterous (m ) specimens are listed separately;
ifboth forms were present they are divided by com-
rnata. New records for Italy are indicated by N I, new
records forpeninsular Italy (“ S ” in D’Urso, 1995a)
by N P I, and new records for A p u lia by N R A .
RESULTS
List of collected specimens
Fam ilia C IX IID A E
Pentastiridius suezensis (Matsumura, 1 9 1 o )
555 (1; 0) 558 (1; 0) 562 (26; 13) 563 (1; 1)
Familia DELPHACIDAE
Asiraca clavicornis (Fabricius, 1 7 9 4 )
552 (0; 1 ) 627 (0; 1 )
Kelisia guttula (G erm ar, 18 18)
556 (1 ; 1 )
Kelisia guttulifera (Kirschbaum, 1 8 6 8 )
552 (1; 0) - NRA
Kelisia gr. ribauti Wagner, 193 8
559 (2; 3) 628 (1 ; 0) - NPI
Stenocranus fuscovittatus (Stai, 1 8 5 8 )
556 (1; 1 ) - N R A
Stenokelisia angusta Ribaut, 1934
554 (1; 0) 555 (7; 3; 9) 556 (0; 1) 560 (6; 0; 2)
Eurysanoides rubripes (Matsumura, 1 9 1 o )
624 (3b; 5b) - N R A
Delphax inermis Ribaut, 1934
555 (0; lb) - N R A
Delphax meridionalis (Haupt, 1924)
556 (1 ; 0) - N I
Euides basilinea (G erm ar, 1 8 2 1 )
5 6 2 ( 1 m ; 0 ) - NPI
Chloriona glaucescens Fieber, 1 8 6 6
558 (2m; 0) 562 (3m; lm) - NPI
Chloriona sicula M atsum ura, 19 10
553 (8m; 3b, 4m) 554 (4m;2b)555 (14m;4b,2m; 1)558
(lm; 0) 559 (11m; 2m) 627 (12m; 2b) - NRA
Laodelphax striatella (Fallen, 1 8 2 6 >
552 (0; 2m) 553 (lm; 0) 559 (2m; 3m) - NRA
Remarks on the composition of the Auchenorrhyncha fauna in some moist areas in Southern Apulia (Italy)
3 1 3
Delphacodes capnodes (Scott, 1 8 70)
555 (1 m ; 0) - N I
Muirodelphax aubei (Perris, 1 857)
556 (2 b , lm;lb,2m),558 (7b; 3b; 2) 624 (0; lb) 628 (0; lb)
Florodelphax leptosoma (Fior, 1 8 6 1 )
556 (0; lb) 559 (4b,2m;4b,2m)-NRA
Toya obtusangula (Linnavuori, 1 95 7)
553 (lm; lm)
Toya propinqua (Fieber, 1 866)
552 (lm; lm) 559 (3m; 2m) 564 (3m; 0) 623 (lm; 0)
N R A
Flastena fumipennis (Fieber, 1 866 )
559 (lb; lb) - N R A
F am ilia TROPIDUCHIDAE
Trypetimorpha sp.
556 (0 ; 0 ; 1 ) - N R A
Figure io. Pentastiridius suezensis. Figure li. Stenokelisia angusta. Figure 12. Delphax meridionalis. Figure 13.
Mocydiopsis oranensis. Figure 14. Melillaia desbrochersi. Figure 15. Adarrus reductus. Photos by m a s s im o Vo llaro .
3 14
Adalgisa Guglielmino & Christoph Buckle
F a m ilia CALISCELIDAE
Caliscelis bonellii (Latreille, 1 8 0 7 )
56 1 (1 ; 0)
Homocnemia albovittata a. Costa, 1 8 5 7
556 (0; 0; 1 )
Peltonotellus quadrivittatus (Fieber, 1 8 7 6 )
564 (0; 1)
Ommatidiotus dissimilis (Fallen, 1 8 o 6 )
556 (0; 0; 1 ) 628 (0; 0; 2) - N R A
Familia ISSIDAE
Agalmatium bilobum (Fieber, 1 8 7 7 )
552 (1 ; 0)
Agalmatium flavescens (Olivier, 1 7 9 1 )
552 (2; 1) 561 (1; 0)
Issus lauri Ahrens, 1818
552 (0; 1)554 (6; 1)557 (2;4)622 (2; 0; 2)
Latissus dilatatus (Fourcroy, 1 78 5 )
55 7 (2; 1)
Familia CERCOPIDAE
Cercopis sanguinolenta (Scopoii, 1 7 6 3 )
624 (2; 0) 625 (3; 1)
Fam ilia APHROPHORIDAE
Lepyronia coleoptrata (Linnaeus, 175 8)
553 (1; 1) 554 (2; 0) 555 (0; 1) 556 (1; 0) 562 (2; 0)
Neophilaenus campestris (Fallen, 1 805)
552 (0; 1 ) 5 5 7 (0; 1 ) 564 (1 ; 1)
Neophilaenus lineatus (Linnaeus, 175 8)
559 (9; 1) 561 (5; 1) 564 (1; 0) 622 (0; 1)
Philaenus spumarius (Linnaeus, 1 758 )
552 (0; 2) 554 (1; 0) 557 (1; 0) 624 (0; 1) 625 (2; 3)
Familia CICADELLIDAE
Agallia consobrina Curtis, 1 8 3 3
552 (0; 1 ) 557 (3; 13)
Anaceratagallia laevis (Ribaut, 1935)
552 (2; 2) 556 (3; 4) 559 (3; 0) 562 (3; 0) 564 (2; 2)
Austroagallia sinuata (M ulsant etRey, 1 855)
552 (0; 2) 553 (1; 1) 558 (8; 3) 564 (0; 5) 625 (0; 2)
Bugraia ocularis (M ulsant et Rey, 1 855)
552 (1; 1) 554 (1; 2) 557 (2; 2; 1) 622 (2; 13) 626 (0; 7)
Hecalus Storai (Lindberg, 1 9 36)
561 (2; 0) - NPI
Stegelytra cf. erythroneura Haupt, 1924
557 (0; 0; 1) - N R A
Empoasca alsiosa Ribaut, 1933
628 (1 ; 2) - N R A
Lindbergina (Youngiada) sp.
557 (0; 9) - N R A
Ribautiana tenerrima (Herrich-s chaffer, 1 8 3 4 )
554 (0; 1 ) - N R A
Eupteryx thoulessi Edwards, 19 26
622 (1 ;0) - N R A
Eupteryx zelleri (Kirschbaum, 1 8 6 8 )
564 (1 ; 6) 622 (0; 1 ) 624 (1 ; 0)
Zyginidia adamczewskii d worakowska, 1970
564 (2; 0) - N R A
Zyginidia gr. ribauti d worakowska, 1970
552 (20; 1 8) 55 3 (2; 4) 554 (2; 2) 556 (1; 2) 561 (0; 1)
559 (7; 13) 562 (2; 1) 564 (9; 0) 624 (1; 1) 628 (7; 3)
Arboridia parvula (B ohem an, 1 845 )
552 (1 ; 1 )
Grypotes staurus ivanoff, 18 85
55 7 (3; 4; 2)
Opsius lethierryi Wagner, 1942
556 (3; 0) 562 (1; 3)
Opsius stactogalus Fieber, 1866
555 (1 ; 5 ) 556 (2; 0)
Neoaliturus fenestratus (Herrich-Schaffer, 1 8 34)
552 (0; 1) 553 (0; 1) 556 (1; 0) 559 (0; 2)
Circulifer sp.
552 (0; 1)
Balclutha nicolasi (Lethierry, 1 8 76 )
559 (6; 10; 1) - NR A
Balclutha rosea (Scott, 1 8 7 6 )
55 3 (0; 1 ) 562 (0; 1) - N R A
Remarks on the composition of the Auchenorrhyncha fauna in some moist areas in Southern Apulia (Italy)
3 15
Macrosteles ossiannilssoni Lindberg, 1954
555 (1 ; 3) - N R A
Macrosteles quadripunctulatus (Kirschbaum, 1 8 6 8 )
623 (2; 1)
Maiestas sp.
557 (0; 2) 559 (0; 1)
Varta rubrostriata (Horvath, 1 9 o 7 )
554 (7; 4) - N R A
Doratura g r. paludosa M elichar, 1 897
556 (1 ; 2) 564 (2; 1 ; 1)
Fieberiella florii (Stai, 1 8 6 4 )
552 (0; 1 ) 554 (0; 1 ) 55 7 (1; 0; 1)
Synophropsis lauri { Horvath, 1 897 )
55 7 (1 ; 0) - N R A
Anoplotettix sp.
552 (0; 1)
Selenocephalus stenopterus signoret, 1 8 8 o
56 1 (1 ; 0)
Cicadula lineatopunctata (m atsumura, 1 9 o 8 )
559 (4; 7) - NRA
Mocydia crocea (Herrich-Schaffer, 1 8 3 7)
627 (0; 1)
Mocydiopsis oranensis (M atsumura, 1 908 )
56 1 (4; 4)
Thamnotettix dilutior ( Kirschbaum, 1 8 68 )
55 7 (1 ; 0)
Thamnotettix zelleri (Kirschbaum , 1 8 68)
623 (7; 8; 7)
Conosanus obsoletus (Kirschbaum, 1 8 5 8 )
552 ( 1 ; 0 ) 55 3 (2;5 ) 555 (2;4) 556 (2 ; 3 ) 559 (0;2)
Euscelis alsius Ribaut, 1952
55 3 (1 ; 0) - N R A
Euscelis lineolatus Bruiie, 1 8 3 2
552 (10; 10)553 (5 ; 0 ) 5 5 4 (2;4)555 ( 1 ; 0 ) 5 5 6 (1 ; 2)
55 7 (5; 6) 559 (7; 5) 564 (3; 1)
Streptanus josifovi Diaboia, 1957
624 (5; 1 1 ) 626 (2; 2)
Artianus manderstjernii (Kirschbaum, 1 8 68)
556 (2; 0)
Melillaia desbrochersi (Lethierry, 1 889)
623 (32; 17) 624 (0; 1) 626 (1; 5) - NPI
Paramesus obtusifrons (Stai, 1 8 5 3 )
553 (9; 3) 555 (1; 2) 556 (2; 3 ) 562 (8; 2; 1) 563 (5; 2)
Parapotes reticulatus (Horvath, 1 8 9 7 )
563 (1 0; 3) - NI
Paralimnus phragmitis (Boheman, 1 847)
555 ( 1 ; 2 ) 556 (1 ;2) 562 (2; 14) - NRA
Psammotettix alienus (D ahibom , 1 8 5 o )
552 (6; 6) 553 (2; 9) 554 (0; 7) 556 (0; 2) 558 (2; 3) 559
(11; 0) 5 62 (2 ; 0 ) 5 6 4 ( 7 ; 8 ) 6 2 2 ( 7 ; 7 ) 6 2 3 (5 ; 2 ) 6 2 4 (3;
2) 625 (3; 1) 627 (1; 0) 628 (18; 18; 18)
Psammotettix confinis (D ahibom, 1 8 5 o )
559 (4; 0)
Adarrus reductus (m elichar, 1 897)
561 (25; 22) 564 (17; 9) 627 (0; 1)
Jassargus latinus (Wagner, 1942)
624 (0; 1)
Arthaldeus striifrons (Kirschbaum, 1 8 68 )
556 (0; 3) - NRA
Calamotettix taeniatus (Horvath, 1 9 1 1 )
562 (2; 14) - NI
The investigated areas
1. Bosco di Rauccio and adjacent areas (St. 5 5 2-
558, 561, 625-627) (Figs. 2-5): 67 taxa collected.
The high number of collected species is due to
the m ajor collecting intensity in relation to the other
two investigated areas. Ten localities with different
ecological features were studied, two of them in two
different seasons. Particular importance have the
reed areas with six species of PhrClginiteS fe e d ers ,
among them Pentastiridius suezensis, Chloriona
glaucescens, Delphax inermis and D. meridionalis.
In other moist areas, characterized by Cyper-
aceae and Juncaceae, further inter esting species
were discovered: Stenokelisia angiista, Delphacodes
capnodes , Ommatidiotus dissimilis (ail on Carex
s p p . ) , Florodelphax leptosoma (on J uncus), Toya
3 1 6
Adalgisa Guglielmino & Christoph Buckle
obtusangula (on Poaceae?) and Eupteryx thoulessi
(on Mentha aquatica l.)- Varta rubrostriata lives
on tussocks of a tall Poaceae species (probable
Erianthus ravennae ) which is present on field
margins west of Specchia della Milogna. In the
central forest area nine (unfortunately female)
specimens of an interesting Ty p h lo c y b in ae species,
Lindbergina ( Youngiada ) sp., were collected on
QuerCUS ilex , and the brachypterous Deltocephal-
in ae Melillaia desbrochersi on the low vegetation
of small clearings. The dry areas south of the Nat-
ural Reserve with a garigue like vegetation furni-
shed very interesting results as well. Among other
species there were found AdarrUS redliCtUS , HecaluS
storai and Mocydiopsis oranensis.
2. Le Cesine and adjacent areas (St. 562, 563, 622,
623 ) (Figs. 6, 7): 22 taxa collected.
Only four localities in this area were investig-
ated. Again, the reed areas along the lagoons are
particularly rich of interesting Auchenorrhyncha:
on PhragUliteS the following species were collec-
ted: Pentastiridius suezensis , Euides basilinea ,
Chloriona glaucescens , Paralimnus phragmitis and
Calamotettix taeniatus. Parapotes reticulatus
was found not far from the PHvagniiteS sites on
Schoenoplectus lacustris , Eupteryx thoulessi on
Mentha aquatica. a rich population of Melillaia
desbrochersi was collected in spring on the herb-
aceous vegetation of an olive grove.
3. Laghi Alim ini (St. 559, 560) (Fig. 8): 18 taxa
collected .
Only two sites were studied in this area. A rich
population of Stenokelisia angusta w as observed on
tall sedges near the reed belt around Lago Piccolo.
Kelisia g r . ribauti ( o n Carex s p . ) , Florodelphax lepto-
soma ( o n June us) . Flastenafumipennis and Balclutha
nicolasi {on Cyperus ) were collected in a moist area
with different small Cyperaceae and Juncaceae.
Observations on some taxa of special interest
Pentastiridius suezensis (M atsumura, 1910)
(St. 555, 558, 562, 563) (Fig. 10)
a ii Pentastiridius specimens collected in
201 1/2012 in Apulia (and a population found some
years before in northern Apulia, province of Foggia,
Lago di Lesina) belong to this taxon. Their aedeagus
shape corresponds to the figures given by Van Stalle
(1991), and by Wagner (1954), who probably had seen
the type material. The species shares apparently the
ecological preferences with P. lepOHUUS (Linnaeus,
1761) and was found in abundance on PhragmiteS
australis in coastal lagoon areas and sim ilar h ab itats .
In D’Urso (1995a) the presence of this species
in Italy is regarded as doubtful with records of
Oliarus pollens (Germar, 1821) possibly referring
to P. suezensis. ah Pentastiridius Kirschbaum,
1868 specimens we collected in other parts of Italy
including Sardinia and all Pentastiridius sp e c im e n s
in the S erv a d e i c o lie c tio n under the name OliarUS
leporinus l. and O. pollens , which were checked
by the authors, belong to P. leporinUS. Thus, it
seems that P. Suezensis is present only in a part of
southeastern Italy, where it replaces P. leporinus,
which is present and common in all other regions
of peninsular Italy. P. SU£Zensis is described from
Egypt, and has a wide distribution primarily in
many parts of southern, southeastern and eastern
Europe, but also in Africa and Asia until India and
Philippines (Van Stalle, 1991).
Until now, there are unresolved taxonom ical
problem s in this species group (see Holzinger et al.,
2003, Webb et al., 2013).
Kelisia gr. ribauti w agner, 193 8
(St. 5 5 9, 62 8 )
There are some doubts about the identity of
Kelisia ribauti in Central Europe and the popula-
tions in the M editerranean regions (see Guglielmino
et al., 2005). Italian populations of this species
group were found in many different habitats from
localities near the seashore until moderately high
mountain areas, always in moist environments
on different small Ca rex species. At least at low
altitude they hibernate in the adult stage.
Steno cr anus fuse ovittatus (Stai, 1 8 5 8 )
(St. 5 5 6) (Fig. 16) - NR A
Species widely distributed in the Palaearctic
region. In Italy it is recorded from Trentino Alto
Adige (Servadei, 1 967), Veneto (Minelli &
Mannucci, 1 979), Lazio (Castellani, 1 95 3 ). The
record for Lazio is doubt ful and may refer rather to
S. major (K irschb aum , 1 8 6 8). In A p u lia th e species
is found in marshes on tall sedges. This is in con-
gruence with the observations in Nickel (2003).
Remarks on the composition of the Auchenorrhyncha fauna in some moist areas in Southern Apulia (Italy)
3 1 7
Figure 16 . Stenocranus fnscovittatus. Figure 17 . Euides basilinea. Figure is. Chloriona glances cens. Figure 19. Delphacodes
capnodes. Figure 20 . Ommatidiotus dissimilis. Figure 21. Varta rubrostriata. Figure 22. Parapotes reticulatus. Figure 23.
Calamotettix taeniatus. p hotos Gemot Kunz.
3 1 8
Adalgisa Guglielmino & Christoph Buckle
Stenokelisia angusta Ribaut, 1934
(St. 554. 555, 556, 560) (Fig. 11) - NPI
The species is recorded from Sicily (A sc he,
1 98 5) and Sardinia (Guglielmino et al., 2000). It is
It is indicated in Della Giustina & Remane (1991)
as therm o-xer op hilous and feeding possibly on
Carex flucca S c h ieb er. Habitat and host plant of th e
populations found in Apulia do not coincide with
this characterization . The host plant in Apulia is a tall
sedge like Carex dCUtifOYTYlis E hrh the habitats are
moist areas in marshes. In Sardinia the species was
found in a spring fen at an altitude of about 1000 m.
Delphax inermis Ribaut, 1934
(St. 5 5 5 ) - NR A
The species is widely distributed in the Mediter-
ranean area. In Italy it seems to be rather rare and is
recorded only from Lazio and Sicily (Servadei, 1968;
D’Urso, 1995a). The record forLazio should be con-
firmed. The host plant is PhmgmiteS australis.
Delphax meridionalis (Haupt, 1924 )
(St. 5 5 6) (Fig. 12) - NI
This species is recorded until now only from
Greece. In Italy it is replaced apparently by the
close related D. vibciUticinUS Asche et Drosopoulos,
1982. The new record for Italy represents one of
several examples in which taxa present on the
Balkan Peninsula occur also in southern or south-
eastern Italy. The specimen in Apulia was collected
o n Phragmites australis in a marsh area.
Euides basilinea (Germar, 1 8 2 1 )
(St. 562) (Fig. 17) - NPI
Also this species is a Phragmites fee A tv. In Italy
it w as recorded until now only from Trentino Alto
Adige (Servadei, 1968) and Veneto-Lom bardia
(O sella, Pagliano-O sella, 1989). The specimen from
Apulia was found on the shore of a lagoon together
w ith Pentastiridius suezensis, Chloriona glauces-
cens and Calamotettix taeniatus.
Chloriona glaucescens Fieber, 18 66
(St. 5 5 8, 5 62) (Fig. 18) - NPI
The species is distributed in Europe (except for
the Iberian Peninsula) and in Central Asia. In Italy
it is recorded by Servadei (1967) from Trentino Alto
Adige. This record is dubious in view of the prefer-
ence of this Chloriona Fieber, 1 866 species for
brackish habitats. The habitats in Apulia were reeds
on the seashore or along the shore of lagoons. Host
plant is Phragmites australis.
Delphacodes capnodes (Scott, 1 8 7 o )
(St. 5 5 5 ) (Fig. 19) - NI
The species is widely distributed in central and
southeastern Europe. Tall sedges are recorded as
host plants. This coincides with our observations in
Apulia.
Trypetimorpha sp
(St. 5 5 6) - N R A
Only one nymph was co lie c te d fro m th is g e n u s ,
the identification of which at species level is at
present impossible. In the past there was some
nom enclatural confusion in this genus (see Huang
& Bourgoin, 1993; Guglielmino et al., 2005). In
Italy, two Trypetimorpha Costa 1 8 6 2 species are
present: T. occidentalis h uang et Bourgoin, 1993
widespread and common in Central Italy, and T.
fenestrata C o sta , 1 862 described from Campania
and recorded also from Basilicata by Servadei
(1 967; as T. piloStt Horvath, 190 7 now a synonym
of T. fenestrata). We checked the s p e c im e n s fro m
Basilicata in the S er v ad e i-c o lie c tio n and confirmed
the identification as T. fenestrata.
Ommatidiotus dissimilis (Fallen, 1 8 o 6 )
(St. 5 5 6, 62 8) (Fig. 20) - NR A
The species is widespread in the Palaea retie
region. In the past it was considered ty rp h o p h ilo u s
and monophagous on Eriophorum vaginatum L .
(Nickel, 2003). However, in the meantime it was
found also on other Eriophorum L. taxa and on
several CareX species in quite diverse habitats. In
Italy it is recorded from Trentino Alto Adige and
Veneto (Servadei, 1967), Toscana (M azzoni, 2005),
Abruzzo and L azio (G uglielm ino e t al., 2 0 0 5 ) . H o s t
plants in Apulia are small sedges in moist areas near
the coast. This coincides with the habitats in Lazio.
In Abruzzo, however, the species was found on dry
mountain pastures at an altitude of 1900 m (on
Carex cf. kitaibeliana Degen ex B e c h . ) . No mor-
phological differences were observed between these
different populations.
Hecalus storai (Lind berg, 1 93 6)
(St. 56 1) - NPI
The species is described from the Canary
Islands and recorded also from France. Our identi-
Remarks on the composition of the Auchenorrhyncha fauna in some moist areas in Southern Apulia (Italy)
3 19
fication is based on Ribaut’s description and fig-
ures. In Italy there is only a record from Sicily (Pan-
teller ia) (D’Urso & Guglielmino, 1995). HecaluS
Stal, 1864 species may be rather variable in size and
vertex shape, whereas there are only slight differ-
ences in the genital morphology. Therefore it is
difficult to define specific characters. Linnavuori
(1975) made an importantcontribution to the know-
ledge of the genus, but many problems are left. The
specimens in Apulia were found in a dry and stony
habitat south of Bosco di Rauccio.
Stegelytra cf. erythroneura Haupt, 1924
(St. 5 5 7 ) - N R A
Un til now this genus was not recorded forApulia.
We found only one nymph (on QueVCUS ileX) .The
authors collected in central and Southern Italy (and
Sardinia) only S. erythroneura (on Quercus ilex an A
Q. Cervis L . ) . Probably also the nymph from Bosco
di R auccio belongs to this taxon. The other Stegelytra
Ghauri 1972 taxon present in Italy, S. pUtOlli (M u Is an t
et Rey, 1875), was collected by the authors in Liguria
(on Q. ilex ) (Guglielmino & Buckle, 2007), and was
later recorded by M azzoni (2005) from Toscana.
Lindbergina (Youngiada) sp.
(St. 5 5 7 ) - N R A
No species of YoUUgiada Dlabola, 1959 was
recorded before from Apulia. We collected only fe-
rn ales, an identification of which on species level is
not possible. They present the sam e colouration as
a fern ale collected in Southern Lazio (Guglielmino
etal.,2005) and were found like the specimen from
Lazio on QueVCUS Hex. In Italy until now two
species of this subgenus are recorded: Lindbergina
loewi (Lethierry, 1884), a doubtful record from
Friuli Venezia Giulia, (see D’Urso, 1 995 a) and L.
chobauti { Ribaut, 1952) (Vidano & Arzone, 1987;
M azzoni, 2005).
Zyginidia adamczewskii Dworakowska, 1970
(St. 564) - N R A
The species is described from Croatia and recor-
ded also from Greece (Drosopoulos et a 1. , 1986).
The first and only record in Italy is from Campania
(Vidano, 1 982). Vidano (1 982) indicates Cynodon
dactylon (L.) Pers., Agropyron repens (L.) p.
Beauv. and other Poaceae as host plants. The
specimens in Apulia were collected in a dry rocky
garigue like habitat.
Zyginidia gr. ribauti Dworakowska, 1970
(St. 552-554, 556, 5559, 561, 562, 564, 624, 628)
Very common taxon throughout peninsular
Italy; it is replaced in Sardinia by Z. SCUtellaris
(H e rr ic h - S c h affer, 1 8 3 8 ), and in Northern Italy
p artly by Z. pullula (Bo hem an, 1845). The relatio n -
ships betw een Z. ribauti, Z. Serpentina (M atsum ura,
1908) and Z. italica (Ribaut, 1947) should be clari-
fied (see also Guglielmino et al., 2005). Z. gr. ri-
bauti displays a remarkable variability in its
aedeagus morphology, not only between different
populations, but also within the same population.
Circulifer s P
(St. 5 5 2)
Only one female of this genus was collected in
a maquis like area of Bosco di Rauccio. The genus
(often inserted in NeoaHturUS D istan t, 19 18) is very
problematic in respect of species discrimination.
Italian populations are quite diverse in colouration
and size. However, no distinct differences in the
genital morphology of males and females were
observed. In males, the shape of the genital plates
corresponds to that given by Ribaut (1 952) for c.
HaematOCepS (Mulsant et Rey, 1855). The habitats
are generally dry places in low and median altitude,
also sandy seashores. The host plants are in some
cases apparently CistUS sp., in others Chenopodi-
aceae. In Germany Circulifer c f . haematOCepS w a s
found on Sedum L. (Crassulaceae) (Nickel, pers.
c o m m . ) .
Maiestas sp
(St. 5 5 7, 5 59)
The females of this genus found during our
study in Apulia belong with great probability to M.
Schmidt geni (Wagner, 1939), which is very wide-
spread and common in dry ruderal lowland places
in peninsular Italy.
Varta rubrostriata (Horvath, 190 7)
(St. 5 54) (Fig. 2 1) - NR A
After the revision of the Varta- StyniphaluS g e n -
eric complex ( V iraktam ath , 2004) the distribution
of v: rubrostriata should be checked. In Italy it is
recorded from Lazio and Basilicata (Servadei,
1967). The presence in both regions was con firm e d
by the authors. The host plants in Italy are appar-
ently Erianthus ravennae (L.) p. Beauv. and Im-
perata cylindrica (L.) p. b eauv., in Bulgaria and
Greece it occurs also on SorghuiTl Halepense (L .) Pers.
320
Adalgisa Guglielmino & Christoph Buckle
Doratura gr paludosa Melichar, 1897
(St. 5 5 6, 5 64)
The group of species close to D. paludoSQ. is in
need of revision. A paper on the topic is in prepara-
tio n . The Doi'CltUVCl J. Sahib erg, 1871 pop u latio n s
fo u n d in southern A p u lia belong to the same species
that is found in other Adriatic parts of peninsular
Italy. In the past those populations were recorded
sometimes as D. pClludoSCl, sometimes as D. veneta
D lab o la , 1 95 9.
Mocydiopsis oranensis (Matsumura, 190 8)
(St. 561) (Fig. 13)
W estm editerranean species, in Italy recorded only
from Apulia (Gargano) and Sicily (Guglielmino,
1993). A small localized population was found
during the recent study in Apulia in a dry and stony
garigue like habitat together with AdarrUS VedllCtUS
(M elichar, 1 89 7).
Melillaia desbrochersi (Lethieny, 1 8 8 9 )
(St. 623, 624, 626) (Fig. 14) - NP1
Mediterranean species, in Italy recorded only
from Sicily (D’Urso, 1995b). In Apulia, we collec-
ted it only in spring. It was found in a olive grove
near the Natural Reserve “Le Cesine”, and in the
Natural Park “Bosco e Paludi di Rauccio” in a
ruderal place and on so me small clearings. Probably
the species is widespread and not uncommon in
southern Italy, but until now it was never found
because of its particular life cycle: adults occur only
in the early (and late?) parts of the year.
Parapotes reticulatus (Horvath, 1 8 9 7 )
(St. 563 ) (Fig. 22) - N I
The discovery of this species in Apulia was
quite unexpected. It is distributed in several coun-
tries of central, northern and southeastern Europe,
including Ex-Yugoslavia. As host plants are recor-
ded Schoenoplectus lacustris (L .) Palla and possibly
S. tabernaemontani (Gmei.) Paiia (Nickel, 2003 ).
A quite abundant population of this species was
found in the lagoon area of the Natural Reserve “Le
Cesine”, on Schoenoplectus lacustris.
Adarrus reductus (M elichar, 1 897)
(St. 56 1, 564, 627) (Fig. 15)
The species is described from Croatia. In Italy it
is recorded only from Apulia (Servadei, 1967). It
was collected in two very dry stony garigue like sites
(south of Bosco diRauccio and near Porto Badisco).
Calamotettix taeniatus (Horvath, 191 1 )
(St. 562) (Fig. 23) - NI
The species is recorded from central and eastern
Europe. In Apulia, it was found on the shore of the
Pantano Grande in the Natural Reserve “Le Cesine”
on its host plant, Phragmites australis, in moder-
ately high abundance.
CONCLUSIONS
During our research in Apulia 84 A uchenor-
rhyncha species were found on the whole. Four
species ( Delphax meridionalis, Delphacodes
capnodes , Parapotes reticulatus and Calamotettix
taeniatus ) are recorded for the first time for Italy,
five ( Stenokelisia angusta , Euides basilinea ,
Chloriona glaucescens, Hecalus storai and Melil-
laia desbrochersi) are new records for the A pennine
Peninsula (“S” in the checklist of the Italian fauna),
and 2 6 are new records for Apulia ( Kelisia gUt-
tulifera , Stenocranus fuscovittatus, Eurysanoides
rubripes , Delphax inermis , Chloriona sicula ,
Laodelphax striatella, Florodelphax leptosoma ,
Toya propinqua, Flastena fumipennis , Trypeti-
morpha sp ., Ommatidiotus dissimilis, Stegelytra cf.
erythroneura, Empoasca alsiosa , Lindbergina
( Youngiada ) sp., Ribautiana tenerrima , Eupteryx
thoulessi , Zyginidia adamczewskii. Balclutha ni-
colasi, B. rosea, Macrosteles ossiannilssoni, Varta
rubrostriata , Synophropsis lauri, Cicadula lineato-
punctata, Euscelis alsius, Paralimnus phragmitis,
Arthaldeus striifrons ) .
The high number of new records for Apulia, and
the fact that some of these records regard species
that are widespread and quite common throughout
Italy, show that the knowledge on this region is
presently very scarce. In addition to the here
presented data, many further research is necessary
to achieve to a sufficient understanding of the
A uchenorrhy ncha fauna in southeastern Italy.
Even if the three studied areas furnished very
im portant results, w e are far from aN approximately
complete knowledge on the Auchenorrhyncha of
these areas. Additional in vestigations should include
more localities, biotopes and collecting seasons.
The distribution of some taxa collected during
our recent study in Apulia is particularly interesting:
Apparently these species are present only in the
Balkan region and in South Italy. Delphax ITieridi-
Onalis was considered an endemic species of
Remarks on the composition of the Auchenorrhyncha fauna in some moist areas in Southern Apulia (Italy)
32 1
Greece before it was discovered in Apulia; AdciVVUS
redliCtliS is recorded only from Croatia and Apulia;
and Zyginidia cidcunczc wskii is known fro m Croatia,
Greece and South Italy (Campania, Apulia). In con-
trast to these three cases, other taxa display a wide
distribution in Europe. In Italy, however, they were
found until now only in Apulia and not in the cent-
ral and northern parts of this country. This group in-
cludes Pentastiridius suezensis , Calamotettix
taeniatus. Parapotes reticulatus and Delphacodes
capnodes. we may add Chloriona glaucescens as
well, a halophilous species, the record of which
from Trentino Alto Adige (Servadei, 1967) is
probably erroneous. A molecular study of these
species in order to clarify the relation ships between
populations from central Europe, southern Italy and
the Balkan region would be very interesting.
Unlike most other regions of Italy, a great part
of Apulia consists of plains and low hills, which
nowadays are almost completely cultivated. Thus,
moist habitats (freshwater lakes or springs and
brackish lagoons), and the dry maquis and garigue
areas, have become extremely rare and harbour the
last relics of a flora and fauna, which in former
times were typical for the whole region, but are now
nearly extinct. The protection at least of the few
natural sites left is therefore of particular impor-
tance. Each of the three investigated areas has
its own special characteristics, each is unique but
fragile and vulnerable.
In the case of the Natural Park of “Bosco e
Paludi di Rauccio” we observed some negative im-
pact of agricultural activity on the protected area.
Whereas the central QuCVCUS UcX forest and the
Specchia della Milogna area in the northeastern
sector of the reserve display more or less safe
conditions, there are other zones around the forest
and above all in the southwestern partof the Natural
Park that seem to be conspicuously compromised.
Apparently, the main problem consists in frequent
arsons of vast extension, easily visible in recently
burnt CareX meadows, but also in green areas where
a glance at the soil between the fresh grasses re-
vealed everywhere the black charred remnants of
the plants burnt in the years before. The almost
completely black colouration of populations of
Lepyronia coleoptrata specimens in these areas
may be interpreted as adaptive character to these
particular conditions (a similar case is documented
in Philaenus spumarius fro nr Great Britain (Wilson,
p e r s . comm.).
Finally we point out the interesting area south
of Bosco di Rauccio, off the Natural Park (St. 555,
561, 627). This site, consisting of quite extended
wet meadows and reeds along a central channel
and adjacent dry garigues, has a great value for
plants, birds and insects. We think it very import-
ant to warrant the conservation of this habitat as
a highly valuable addition to the nearby located
N atural P ark .
ACKNOWLEDGMENTS
We are grateful to Carmine A n n ic c h iaric o for
his help concerning the collecting permission in the
Natural Reserve “Le Cesine”. We thank Vittorio De
Vitis for useful advice and information during our
research in the Natural Park “Bosco e Paludi di
Rauccio”. Many thanks also to Massimo Vollaro
and Gemot Kunz for the photos.
REFERENCES
Asche M ., 1985. Zur Phylogenie der Delphacidae Leach,
1815 (Homoptera Cicadina F u lg o ro m o rp h a ) .
M arburger Entom ologische Publikationen,2: 1-910.
Castellani O ., 1 95 3. Contributo alia conocenza della
fauna em itterologica d’ltalia. Hemiptera Homoptera.
Bollettino dellaAssociazione romana di entom ologia,
7: 15-16.
Della Giustina W. & Remane R., 1991. La Faune de
France des Delphacidae (Homoptera Auchenor-
rhyncha). I. Recoltes d’aout 1 9 89. Cahiers des
N a t u ra lis te s , Bulletin des Naturalistes Parisiens, 47:
33-44.
Drosopoulos S., Asche M. & Hoch H., 1986. A prelimary
list and some notes on the Cicadomorpha (Ho-
moptera - Auchenorrhyncha) collected in Greece.
Proceedings of the 2nd International Congress
Concerning the Rhynchota fauna of Balkan and
adjacent Regions, pp. 8-13.
D’Urso V., 1 995a. Homoptera Auchenorrhyncha. In:
M in e Hi, A ., Ruffo, S. & La Posta, S. (Eds.), Checklist
delle specie della fauna italiana, 42: 1-35.
D’Urso V., 1995b. Contributo alia conoscenza della
distribuzione in Italia di alcune specie di A uchenor-
rinchi (Insecta Rhynchota: Homoptera). N aturalista
siciliano, ser. IV, 1 9: 99-104.
D’Urso V. & Guglielmino A., 1 995. Arthropoda di
Lampedusa, Linosa e Pantelleria (Canale di Sicilia,
Mar M e d ite rra n e o ) . Homoptera Auchenorrhyncha.
Naturalista siciliano, 19 (Suppl.): 279-30 1.
322
Adalgisa Guglielmino & Christoph Buckle
Guglielmino A., 1993. I Cicadellidi dell’Etna. Studio
tassonom ico e note ecologiche e b io g e o g ra fic h e
(Homoptera A u c h e n o rrh y n c h a ) . Memorie della
Societa Entomologica Italiana, 72: 49-1 62.
Guglielmino A. & Buckle C., 2007. Contributo alia
conoscenza della fauna ad A uchenorrhyncha (Hemip-
tera, Fulgorom orpha et C ic a d o m o rp h a) di Liguria e
dell’Italia meridionale. Frustula Entomologica, n.s.
30: 149-159.
Guglielmino A., Buckle C. & Remane R ., 2005. Contri-
bution to the knowledge of the A uchenorrhyncha
fauna of Central Italy (Hemiptera, F u lg o ro m o rp h a
et C ic a d o m o rp h a ) . Marburger E n to m o lo g is c h e
P ub lik atio n e n , 3: 13-98.
Guglielmino A., D’Urso V. & Alma A., 2000. Contri-
bution to the knowledge of A uchenorrhyncha (Insecta
Homoptera) from Sardinia (Italy). Deutsche Entomo-
logische Zeitschrift, 47: 161-172.
Holzinger W., Kammerlander I., Nickel H., 2003. The
A u c h e n o rrh y n c h a of Central Europe. I. Fulgoro-
morpha, C icadom orpha excl. C ic a d e Hid ae . Brill
Leiden Boston 2003, 673 pp.
Huang J. & Bourgoin T., 1 993. The planthopper genus
Trypetimorpha. System a tics and phylogenetic
relationships (Hem ipetra: Fulgoromorpha: Tropi-
duchidae). Journal of Natural History, 27: 609-629.
Linnavuori R., 1975. Revision of the Cicadellidae (Ho-
moptera) of the Ethiopian Region III. Deltocephal-
inae, Hecalini. Acta Zoologica Fennica, 143: 1-37.
Mazzoni V ., 2005. Contribution to the knowledge of the
A u c h e n o rrh y n c h a (Hemiptera Fulgoromorpha and
Cicadomorpha)of Tuscany (Italy). Redia, 88: 85-102.
Minelli A. & Mannucci M.P., 1979. Studi sul popola-
mento animale dell’Alto Trevigiano. I. Faunistica e
sinecologia di alcune cenosi riparie dei Laghi di
Revine. Lavori della Societa Veneziana di Storia
Naturale, 4, 48-60.
Nickel H ., 2003. The Leafhoppers and Planthoppers of
Germany (Hemiptera, A uchenorrhyncha): Patterns
and strategies in a highly diverse group of phyto-
phagous insects. Pensoft, Sofia, 460 pp.
Osella G . & Pagliano-Osella M ., 1989. Studi sulla palude
del Busatello (Veneto-Lombardia) 9. Gli Omotteri
A uchenorr inchi. M em orie del M useo Civico di Storia
Naturale, 7: 89-97.
Ribaut H ., 1 952. Homopteres A uchenorhinques. II.
(Jassidae). Faune de France. Paris, 57, 474 pp.
ServadeiA., 1967. Rhynchota (Heteroptera, Homoptera
A uchenorrhyncha). Fauna d’ltalia, volume IX,
Calderini Editore, Bologna, 85 1 pp.
Servadei A., 1 96 8. Contributo alia corologia dei
Rhynchota Homoptera Auchenorrhyncha d’ltalia.
Annali M useo Civico di Storia Naturale "Giacomo
Doria", 7 7: 1 3 8- 1 8 3.
Van Stalle J., 1991. Taxonomy of Indo-M alayan Penta-
stirini (Homoptera, Cixiidae). Bulletin de l’Institut
Royal des Sciences Naturelles de Belgique, Entomo-
logie, 61 : 5-101 .
Vidano C., 1982. Contributo alia conoscenza delle
Zyginidia d’ltalia. Memorie della Societa Entomolo-
gica Italiana, 60: 343-355.
Vidano C. & Arzone A., 1987. Typhlocybinae of
broadleaved trees and shrubs in Italy. 4. Fagaceae.
Redia, 70: 1 7 1-1 89.
Viraktamath C.A., 2004. A revision of the Varta-
Stymphalus generic complex of the leafhopper tribe
Scaphytopiini (Hemiptera: Cicadellidae) from the
Old World. Zootaxa, 713: 1-47.
Wagner W., 1 954. Die Fulgoroidea der Omer-Cooper-
Expedition in die Lybische W ueste (Hemiptera
Homoptera). Bulletin de la Societe Fouad ler
d’Entomologie, 38: 211-219.
Webb M., Ramsay A.J. & Lemaitre V.A., 2013. Reveal-
ing the identity of some early described European
Cixiidae (Hemiptera: Auchenorrhyncha) - a case of
‘forensic’ taxonomy; two new combinations and a
name change for ReptCllllS pClflZ€ri in Britain. Acta
Musei Moraviae, Scientiae biologicae, 98: 57-95.
Biodiversity Journal, 2015, 6 (1): 323-326
Monograph
Presence's mapping of Brachytrupes megacephalus (Lefebvre,
1827) (Orthoptera Gryllidae) within the Natural Reserve of
Vendicari (Noto, Siracusa, Italy)
Alfredo Petralia 1 *, Ettore Petralia 2 , Giorgio Sabella 3 , Filadelfo Brogna 4 & Corrado Bianca 1
*Ente Fauna Siciliana Onlus, Noto, Italy
2 Sud & Dintomi Onlus, Catania, Italy; Studio Oikos, Catania, Italy
’Dipartimento di Scienze Biologiche, Geologiche e Ambientali, sez. Biologia Animale, Catania University, Italy
4 Regione Siciliana, Dipartimento Azienda Regionale Foreste Demaniali, Siracusa, Italy
’Corresponding author, e-mail: alfredo.petralia@yahoo.it
ABSTRACT Brachytrupes megacephalus (Lefebvre, 1827) (Orthoptera Gryllidae) is a species included in
the Annexes II and IV of EU Directive 92/43 as taxon requiring strict protection. The authors
summarize the researches aimed to recognize the localization of this species within the natural
reserve of Vendicari, protected area along the south eastern Sicilian coast in the territory of
Noto (province of Siracusa). The presence of the specimens was ascertained by detecting its
holes on the soil surface. The holes position was recorded using GPS and utilized for mapping
the presence of the species as tool for its protection management in the reserve territory.
KEY WORDS monitoring; wildlife management; protected areas; mapping.
Received 25.07.2014; accepted 30.11.2014; printed 30.03.2015
Proceedings of the 2nd International Congress “Speciation and Taxonomy”, May 1 6th- 1 8th 2014, Cefalu-Castelbuono (Italy)
INTRODUCTION
The Brachytrupes megacephalus (Lefebvre,
1827) (Orthoptera Gryllidae) (Figs. 1, 2), described
on specimens from Sicily, is a South-Mediterranean
species widespread in the sandy environments of
Sicily, Aeolian Islands, Maltese Islands, southern
Sardinia, Northern Africa (included the Saharan
oases): it is an exclusively sandy adapted cricket.
This species is considered as a biodiversity ele-
ment of particular interest thanks to its complex
eco-biology. Previous works (see Conti et al.,
2014) investigated the biological cycle, swimming
ability and digging technique, reproductive mode,
coupling pattern and more. Due to its current rar-
efaction in Europe, this species has been included
in the Annexes II and IV of EU Directive 92/43 as
a species requiring strict protection.
The reserve of Vendicari lies in the south-
eastern extremity of Sicily and is part of a vast
wetlands system that is one of the most important
of the island; it extends for about 8 km of coastline
and 0.3- 1.5 km inland and includes a series of dif-
ferent environments (sweet-water wetlands, coastal
lagoons, garrigue scrubland, Mediterranean ma-
quis) and a very rich biodiversity: a wide descrip-
tion of the reserve was edited by Petralia (2010).
B. megacephalus , having been already monitored
in Vendicari a little bit more than ten years ago
(Petralia et al., 2003), is one of the main com-
ponent of the artropodological fauna in the reserve
(Petralia & Russo, 2010).
324
Alfredo Petralia etalii
This research was aimed to map the localization
of the species in order to provide basic information
useful to manage the protection of the species itself.
The study was carried out during the breeding
season (from March to early May of 2012) when
the digging activity of the specimens (surface active
again after the winter suspension, for mate) is
particularly intense and easily detectable: the
location of individuals was carried out by detecting
the position of the holes that the animals burrow
into the sand and where the animals remain for most
of their life, also where they die after spawning. The
traces that indicate the presence of the animals are
two (Figs. 3,4): the mouths of the burrows and the
little sandy cones occluding those; also the piercing
sound-calls emitted by the males to attract the
females provide further information about the
presence of individuals.
The survey in the reserve was conducted in the
sandy sectors of the A zone (integral reserve) in the
potential habitats for the presence of B. megacepha-
lus (Figs. 5, 6). The concerned areas are: 1, the
mouth of the Tellaro river (Eloro) at the extreme
north of the reserve; 2, the sandy dunes of Calam-
osche; 3, the sandy south western area of the Vendi-
cari island; 4, the southern dune belt where the GPS
position of each detected burrow was recorded.
RESULTS AND COMMENTS
The presence of the species was ascertained
within the areas marked with numbers 1 and 4
(Fig. 6).
In the latter the GPS burrow records (290 in 18
ha) allowed to obtain the representation of the area
were the species localizes (Fig. 7) using gvSIG
program: the animals dwell exclusively on the
sandy belt and do not intrude both the sandy beach
(seaward) and inland; it is also possible to observe
particular concentrations of burrows in the extreme
north of the sandy belt (Fig. 8).
In the Vendicari island were not found speci-
mens of the monitored species. Probably that is due
Figures 1, 2. Specimens of B. megacephalus photographed before their release: on the male's forewings (Fig. 1) is visible
the stridulatory organ, absent on the female (Fig. 2). Figures 3, 4. Examples of burrow's mouth of B. megacephalus (Fig.
3) and little sandy cone that close the burrow on the soil surface (Fig. 4) (Photos by A. Petralia).
Presence's mapping of Brachytrupes megacephalus within the Natural Reserve of Vendicari (Noto, Siracusa, Italy) 325
Figure 5. Localization (jellow star) of the reserve of Vendicari along the south western Sicilian coast. Figure 6.
In dark-green the A zone of the reserve, in light-green the B zone; 1 to 4 the potential habitats of B. megacephalus.
A i\
VJft I
r marshes
■ v "i t yi y | -i.
sea
Figure 7. Mapping of the B. megacephalus presence along
the dune belt in the southern part of the reserve of Vendicari.
to the not good condition of the sandy habitat: too
narrow extension of the sandy surface in the island,
too windy and not protected because of scarce
vegetation, too brackish.
Also in the area marked with number 2 (Calam-
osche) the species was not detected: it is important
to emphasize that in the previous monitoring carried
out in 2003 (Petralia et al., 2003) the species was
present in these sandy dunes. Probably the disap-
pearance here of this species can be related with the
strong anthropic pressure on the dunes just behind
the beach of Calamosche: in particular the anarchic
trampling on the sand (and the consequent destruc-
tion of nests, eggs and young specimens of B.
megacephalus especially during the early stages of
development) to reach the beach for swimming in
the summer months, which increased over time,
could have acted as a decisive factor in habitat
degradation and, as consequence, in disappearance
of the species.
We can conclude that the protection of B. mega-
cephalus depends on a very severe protection of the
stability of his habitat. The results of the mapping
here described, indicate the areas in which is oppor-
tune to concentrate the actions aimed to ensure the
safeguard of the species: firstly a veiy strict prevention
326
Alfredo Petralia etalii
Figure 8. Northern sector of the
dune belt referred to the Fig. 4 to
show the particular concentra-
tions of burrows highlighted by
red dots.
Figure 9. Calamosche in winter
(photo by www.temioggi.it) and
in summer (Figure 10, photo by
www.itineraricamper.it): the very
heavy human pressure on the
beach in summer could had gen-
erated negative effects on the
conditions for the survival of B.
megacephalus in the back dunes,
from which the species has dis-
appeared.
of the trampling (by humans and by cars) on the
dune, given its destructive effects on the sandy hab-
itat integrity.
The Calamosche situation (Figs. 9, 10) (in par-
ticular the disappearance of the species in this area)
represents in this sense a clear warning signal that
induces a very careful control in the Eloro area (num-
ber 1 in Fig. 6) and along the dune belt also exposed
to trampling in violation of the protection rules
provided for the A area where the dunes are located.
REFERENCES
Conti E., Costa G., Petralia A. & Petralia E., 2014.
Eco-ethology of Brachytrupes megacephalus
(Orthoptera, Gryllidae), protected specie in UE. In:
Petralia A. & Bianca C. (Eds.), 2nd Djerba Interna-
tional Mediterranean Environment Sustainability
Conference, Djerba Tunisia, 22-25. April 2012,
Proceedings. © Atti e Memorie dell'Ente Fauna
Siciliana, 11: 51-56.
Petralia A. (a cura di), 2010. L'area protetta di Vendicari.
Atti del Convegno celebrativo per il 35° anno di
fondazione dell'Ente Fauna Siciliana, "Case Citta-
della", Vendicari-Noto (SR) 25-26 ottobre 2008.
© Ente Fauna Siciliana (supplemento a Grifone),
432 pp.
Petralia A. & Russo C., 2010. Artropodofauna e
biowatching. In: Petralia A. (a cura di) - L'area
protetta di Vendicari. Atti del Convegno celebrativo
per il 35° anno di fondazione dell'Ente Fauna
Siciliana, Vendicari-Noto (SR) 25-26 ottobre 2008,
© Ente Fauna Siciliana (supplemento a Grifone), pp.
209-226.
Petralia A., Russo C. & Cartarrasa S., 2003. Topology
of Brachytrupes megacephalus (Lefebvre, 1827)
(Orthoptera, Gryllidae) in some Sicilian Natural re-
serves. Proceedings Fifth International Symposium
on GIS and Computer Cartography for Coastal Zone
Management, Genova, 2003.
Biodiversity Journal, 2014, 5 (4): 327-340
Monograph
First purposive study of beetles (Coleoptera) from endogean
environments in Bulgaria: collection sites and preliminary
results
Rostislav Bekchiev & Borislav Gueorguiev
National Museum of Natural History, 1 Tsar Osvoboditel Blvd, 1000 Sofia, Bulgaria; e-mails: bekchiev@nmnhs.com;
bobivg@yahoo.com
jj*
Corresponding author
ABSTRACT So far, special attention to the endogean and MSS (Mesovoid Shallow Substratum) fauna was
not paid in Bulgaria, though typical subterranean species of the Coleoptera have been described.
The aim of present study is to put on record the results of a broad-scale study of the coleopteran
fauna from the MSS and lower (euedaphic) soil horizons in the country. We carried out invest-
igations in the period April 2006-July 2014, manly in the Vitosha Mt., Pirin Mt., Stara Planina
Mts., Slavyanka Mt., Belasitsa Mt., Erma and Kresna Gorge, Western Rhodopes Mts., and
Srednagora Mts. For the time being, material from the following families was identified to the
genus and species levels: Anobiidae, Aphodiidae, Carabidae, Clambidae, Corylophidae,
Curculionidae, Endomychidae, Histeridae, Leiodidae, Monotomidae, Scyrtidae, Silvanidae,
Silphidae, Staphylinidae (Pselaphinae) and Zopheridae. We report for the first time the sub-
genus A ntisphodrus Schaufuss, 1865 (Carabidae) and Zustalestus Reitter, 1912 (Curculionidae)
from Bulgaria. Blemus discus discus (Fabricius, 1792) is recorded for the second time from
the country.
KEY WORDS Coleoptera; endogean and MSS fauna; Bulgaria; news records.
Received 13.12.2014; accepted 01.03.2015; printed 30.03.2015
Proceedings of the 2nd International Congress “Speciation and Taxonomy”, May 1 6th- 1 8th 2014, Cefalu-Castelbuono (Italy)
INTRODUCTION
The superficial, cave and hemiedaphic inver-
tebrate fauna in Bulgaria has been an object of
comprehensive investigations for almost 120 years
already. In the same time still very little is known
about invertebrates living in the lower soil layers
(so called euedaphic or endogeic environments) and
especially in the network of fissures and crevices in
the maternal rock below the soil horizon.
The latter environment is usually referred to as
Mesovoid Shallow Substratum (MMS), according
to the works of Juberthie et al. (1980, 1981), or
superficial subterranean habitats (SSHs), according
to Culver & Pipan (2008). In regard to the Cole-
optera, it seems that this specific environment has
been widely discussed by southwest Europe authors
(Ruffo, 1959; Laneyrie, 1960; Coiffait, 1963) prior
to its formal introduction by Juberthie et al. (1980).
At present, at least four basic types of MSS habitats
are discriminated (Juberthie, 2000, Ortuno et al.,
2013), based on different combinations of abiotic
and biotic factors.
Typical endogeic species can be found in most
of the soil-dwelling groups of Arthropoda: the
Lower insects (Japygidae), beetles (Carabidae,
Leiodidae), myriapods (Diplopoda, Chilopoda),
isopods (Isopoda), spiders (Araneae), etc.
328
Rostislav Bekchiev & Borislav Gueorguiev
Undoubtedly, one of the most interesting groups
among them are the beetles represented by a relat-
ively high number of endemic species. Special at-
tention to the endogean and MSS fauna in Bulgaria
has been paid only recently (Deltchev et al., 2011;
Langourov et al., 2014). Typical endogean or
hypogean beetles, excluding those collected in
caves and precipices, were found occasionally
(Knirsh, 1930; Genest & Juberthie, 1983; Genest,
1983; Hurka, 1990; Janak & Moravec, 2008).
The aim of present study is to put on record the
results of a broad-scale study of the coleopteran
fauna inhabiting MSS and lower (euedaphic) soil
horizons in Bulgaria. Here we give a list of the
collecting localities and a register of the taxa found
in the different sites.
MATERIAL AND METHODS
The investigation was carried out in the period
April 2006- July 2014, manly in the Vitosha Mt.,
Pirin Mt., Stara Planina Mts., Slavyanka Mts.,
Belasitsa Mt., Erma and Kresna Gorge, Western
Rhodopes Mts., and Sredna gora Mts (Fig. 1.,
Table 1).
The traps were made from PVC pipe with
diameter of the holes 8 cm and length of 60 and 80
cm. One hundred and eight holes were drilled on
each pipe, at 10 cm distance from its end. Traps
were put into 60 or 80 cm deep hole dug as deep
as the limestone or silicate layer. Ten centimeters
high plastic cup tied to polythene rope, and filled
with solution of ethilenglycol or ethilenglycol with
few drops of formalin was put into the end of the
pipe. Traps were covered by solid plastic covers in
order to avoid penetration of superficial fauna into
the pipe and infiltration of water during heavy
rains. In some cases we also used olphactory
attractant (fish).
The identificationof the taxa has been made as fol-
lows: Curculionidae (Luigi Magnano), Histeridae
(Evgeni Chehlarov), Pselaphinae (first author), and
all other families (second author).
BULGARIA
Plovdiv
MACEDONIA
Vania
BLACK
SEA
TURKEY
Sea of Marmara
Figure 1. Distribution of localities with MSS traps in Bulgaria.
First purposive study of beetles (Coleoptera) from endogean environments in Bulgaria: collection sites and preliminary results 329
TRAP NO.
DATE OF
SETTING
SITE DESCRIPTION
LENGTH
OF TUBE
V-N-l
29.IV.2006
Vitosha Mts., northern slope, above Boyana, Boyanski kamak place, at
the bottom of a 4-5 m deep microcave; dry, alt. 847 m.
70 cm
V-N-2&3
30.IV.2006
Vitosha Mts., northern slope, two traps set ca. 30-35 m above Boyanski
kamak, in a scree in mixed forest of Fagus silvestris and Carpinus
betulus; alt. 847 m.
60 cm
V-E-l
13.V.2006
Vitosha Mts., eastern slope, approx. 28 km south of Sofia, on the road Sofia-
Samokov, Yarema place; forest of Fagus sylvatica, in a brown soil, humid,
close to a small river, alt. 1363 m
80 cm
V-W-l
10.VI.2006
Vitosha Mts., western slope, village of Bosnek, near the cave Duhlata, karst,
stony substrate mixed with clay, alt. 964 m
80 cm
V-W-2
24.VI.2006
Vitosha Mts., western slope, village of Bosnek, near the cave Duhlata,
karst, stony substrate, clay, alt. 992 m
60 cm
V-W-3
24.VI.2006
Vitosha Mts., western slope, village of Bosnek, near the cave Duhlata,
karst, stony substrate, clay, alt. 992 m
70 cm
V-SL-1-2
06.VI.2013
Vitosha Mts., Bosnek Vill., near Akademik cave,
N 42°29'28.28" E 23°11T8.28"
60 cm
V-SL-3
06.V1.20 13
Vitosha Mts., Bosnek Vill., scree on the road to Chuipetlyovo
60 cm
V-SL-4
06.VI.2013
Bosnek Vill., Popov Izvor Karst spring
60 cm
V-SL-5
02.X.2013
Bosnek Vill., near Pepelyankata Cave
60 cm
V-SL-6
02.X.2013
Bosnek Vill., near Duhlata Cave
80 cm
BK-mssl
29.IV.2006
Vitosha Mt., above Boyana, Boyanski kamak place, at the bottom of a
4-5 m deep microcave; dry, alt. 847 m.
80 cm
Du-mss4
24.VI.2006
Vitosha Mt., near Bosnek Village, near Duhlata Cave, karst, stony
substrate, clay, alt. 992 m
80 cm
P-W-2
7.V.2006
Pirin Mts., western slope, above village of Ilindentsi, Zandana Area, karst,
in a scree, dry soil/ sandy substrate, alt. 492 m
70 cm
P-W-4
14.V.2006
Pirin Mts., western slope, village of Gradeshnitsa, near Gradeshnichka
banya, at the base of stony/ sandy cliff, dry, sandy/ stony substrate, alt.
312 m
60 cm
P-N-l
24.V.2006
Pirin Mts., northern slope, approx. 6 km before Predela Area, humid ravine,
Fagus sylvatica forest, at the base of Fagus tree, thick layer of leaf litter,
humid soil mixed with stones, alt. 676 m,
60 cm
P-N-2
24.V.2006
Pirin Mts., northern slope, approx. 6 km before Predela Area, humid
ravine, Fagus silvatica forest, humid soil and gravel, alt. 676 m
80 cm
P-E-1&2
25.V.2006
Pirin Mts., eastern slope, 3 km before village of Gospodintsi, Gotse Deltshev
District, approx. 30 m away of the main road Bansko-Gotse Deltshev and
approx. 5-6 m of a small river; in scree at the base of a limestone rocks,
close to broad-leaf tree; alt. 585 m
60 cm
P-S-l
25.V.2006
Pirin Mts., southern slope, approx. 900 m after Popovi livadi Hut on the
main road Gotse Deltshev-Katuntsi, ca. 40-50 m away of the road, marble
stone debris on a small meadow; alt. 1367 m
50 cm
P-S-2&3
18.VI.2006
Pirin Mts., southern slope, approx. 1700 m away of the main road Gotse
Deltshev-Katuntsi, on the secondary road to Orelyak Peak; in a small
valley, Fagus forest, alt. 1560 m
60 cm
Table 1. Distribution of localities with MSS traps in Bulgaria (continued).
330
Rostislav Bekchiev & Borislav Gueorguiev
TRAP NO.
DATE OF
SETTING
SITE DESCRIPTION
LENGTH
OF TUBE
P-S-4&5
8.VL2006
Pirin Mts., southern slope, St. Iliya Site near village of Kalimantsi; close
to the chapel, under the venerable Quercus coccifera trees, alt. 494 m
60 cm
P-S-6
27.VI.2006
Pirin Mts., southern slope, Peshtemik Site near village of Kalimantsi;
against the large travertine, under the double willow, alt. 380 m
70 cm
P-S-7
27.VI.2006
Pirin Mts., southern slope, Peshtemik Site near village of Kalimantsi;
close to the large travertine, in a smaller travertine, under a hazel bush
60 cm
WR-1
23.IV.2006
West Rhodopes Mts., central parts, approx. 1 100 m after the crossroad to
village of Borovo towards village of Belitsa; on the left side of the road, in
a small rocky valley, overgrown with bushes and Pinus nigra, ca. 50 m of
the road, alt. 657 m
55 cm
WR-2
23.IV.2006
West Rhodopes Mts., central parts, approx. 1100 m after the crossroad to
village of Borovo towards village of Belitsa; on the left side of the road,
in a small rocky valley, overgrown with bushes and Pinus nigra, ca. 100
m of the road, alt. 666 m
80 cm
WR-3&4
23.IV.2006
West Rhodopes Mts., central parts, on the way to village of Belitsa;
narrow valley on the right side of the road, ca. 80 m of the road, Pinus
nigra and deciduous bushes, alt. 666-668 m
60 cm
WR-5
14.VII.2007
West Rhodopes Mts., southern parts, near village of Koshnitsa, below
the cave Uhlovitsa; right slope, above the trek, at the foot of hornbeam
bushes, not far from a old beech tree, humid and shady place, gravels in
the soil, alt. 928 m
80 cm
EG-1&2
11.VI.2006
Rui Mts., Erma Gorge, ca. 30 m before the tunnel, on the slope over-
grown with hazel bush, ash-trees; rocky substrate, at the foot of rocks;
685 m
60 cm
SP-1
6.VI.2006
Stara Planina Mts., Toplya Site near village of Golyama Zhelyazna; ca.
20 m of the entrance of Toplya Cave; ca. 25-30 m of the river; karst
slope overgrown with scarce bushes and deciduous trees; 460 m
70 cm
SP-2
7.VI.2006
Stara Planina Mts., Toplya Site near village of Golyama Zhelyazna; ca. 5
m of the entrance of Yalovitsa Cave; karst slpe inca. 25-30 m of the river;
karst slope in young Quercus forest; 608 m
50 cm
S-l
4.VII.2006
Slavyanka Mts., Livade Site near village of Goleshevo in Alibotush Reserve;
karst slope in Pinus forest; ca. 1700 m, N 41°23'532" E 23°36'307"
60 cm
SG-1
29.IV.2006
Sredna gora Mts., St. Ivan Site near Panagyurishte, abandoned vineyard
overgrown with scattered Prune trees and blackberries in close proximity
to forest of Pinus nigra', deep soil layer, lower horizon mixed with stones,
584 m
100 cm
SG-2
29.IV.2006
Sredna gora Mts., same coordinates and site description; situated ca. 30
m apart of SG- 1 .
60 cm
SG-3
29.IV.2006
Sredna gora Mts., situated ca. 30 m apart of SG-1. Trap set in young
artificial forest of Pinus nigra; brown forest soil mixed with stones; 5-7
cm thick layer of pine needles
80 cm
SG-4
29.IV.2006
Sredna gora Mts., same coordinates and site description; trap is situated
ca. 10 m apart of SG-3. Trap set in young artificial forest of Pinus
nigra; brown forest soil mixed with stones; 5-7 cm thick layer of pine
needles
60 cm
DH-1&2
10.V.2007
Derventsky Heights, village of Dennitsa, crossroad to Stefan Karadzhovo,
Yambol District, sink-hole in Quercus forest, at the base of a big stone;
alt. 365 m
60 cm
Table 1 (continued). Distribution of localities with MSS traps in Bulgaria.
First purposive study of beetles (Coleoptera) from endogean environments in Bulgaria: collection sites and preliminary results 331
RESULTS
Up to now, material from the following families
was identified to genus and/or species level:
Anobiidae, Aphodiidae, Carabidae, Clambidae,
Corylophidae, Curculionidae, Endomychidae,
Histeridae, Leiodidae, Monotomidae, Scyrtidae,
Silvanidae, Silphidae, Staphylinidae (Pselaphinae),
and Zopheridae (Table 2).
DISCUSSION
Carabidae
Thirty one ground-beetle taxa at the species
level were collected in the traps. Eight of them, in-
cluding one undescribed species from the genus
Laemostenus, are Balkan endemic species. The sub-
genus Antisphodrus Schaufuss, 1 865 is a new taxon
to the fauna of Bulgaria. So far, no species of this
group was known from the core area of the Balkan
Peninsula. Antisphodrus display scattered distribu-
tion in the Northern Mediterranean as its species
occur from Spain in the west to Iran in the east.
They have restricted distribution by loci and are
confined to endogean and hypogean, primarily lime-
stone habitats. The only female specimen we col-
lected from this subgenus belongs to a new species
for the science. Currently, the description of this
form is prevented for the lack of enough material.
The ground-beetles collected might be divided
condicionally in three categories in view of their de-
gree of specialization to underground way of life.
The first group includes three true endogean
species. Trechus subacuminatus and Laemostenus
( Antisphodrus ) sp. are hither to found only in the
MSS-niche in Bulgaria. The two species are partly
depigmented, and possess small, but functioning
eyes. With certainty, both are very rare and strictly
localized everywhere since they were not caught
before using the standart methods of collecting. To
the same group belongs also Duvalius regisborisi,
which formerly was found only in caves. It is an
eyeless beetle well-adapted to life in the under-
ground environment. The second group contains
seven species (. Blemus discus discus , Laemostenus
cimmerius weiratheri, L. plasoni, L. terricola
punctatus, Trechus austriacus, T. irenis, and T. sub-
notatus),thQ most of them found repeatedly in caves
but now also caught in MSS-traps. That category
occupies an intermediate position between the eu-
eadiphic (endogean) species and the soil-inhabiting
species.
The separation of this group is evidenced from
their frequency and number of individuals found in
the MSS-traps we put. The third group includes
edaphic (or soil) species, which are primarily forest
dwellers. This species complex is the dominant one
with respect to the number of species-twenty
species from fiftheen genera (Table 2). Most of those
species are forest dwellers, except for Bembidion
dalmatinus and Syntomus pallipes , which are char-
acteristic of open and ecotone habitats. It is worth
noting that the dominant species in the MSS-traps
in the Vitosha Mt. is Aptinus bombarda. We did not
find it in the traps put in other places. Blemus discus
discus is recorded here for the second time for the
country (see Hieke & Wrase, 1988).
Leiodidae
Twenty three taxa of the species level from
Leiodidae have been identified till now. This figure
excludes the species of Colon Herbst, 1797 and
Leiodes Latreille, 1796 which identification is still
unaccomplished. The most typical example of the
MSS -environment is the endogean Guerguievella
petrovi. This very small, blind and depigmented
beetle belongs to a monotypic genus and species that
was discovered not long ago (Giachino & Gueor-
guiev, 2007).The type series of this species includes
three dozens of specimens made available by hand-
collecting in six separate visits of the Kraypatnata
Peshtera” Cave near Smilyan Village. The visits
were carried out in the period 1962-2004.
Recently, we collected Guerguievella petrovi
twice in MSS-traps in a mass, as the samples signi-
ficantly differ to each other in the number of indi-
viduals. The first sample, exposed in the dry
summer-autumn season, contained three specimens,
while the next one, exposed in the wet autumn-
winter season, contained more than 60 specimens.
The cholevine species, like Choleva angusara, C.
glauca, Nargus badius, Ptomaphagus sericatus, and
Sciodrepoides watsoni, are detritophagous. They are
sometimes collected in caves in Bulgaria and now
they were found in MSS-traps. Other species, such
as the leiodines ( Agathidium spp., Hydnobius spp.,
Leiodes spp.), eat fungi and live above the ground
or underground (Newton, 1998).
332
Rostislav Bekchiev & Borislav Gueorguiev
Family
Species and subspecies
Trap No.
Collection date
References
1
Familia
ANOB1IDAE
Ptinus sp.
V-N-l
P-W-2
P-S-4&5
30.4-3.6.2006
7.5.-18.6.2006
7.12.2006-19.4.2007
Present paper
2
Familia
APHODIIDAE
Ataenius horticola
Harold, 1869 -Fig. 2.
P-W-2
14.05.-6.07.2006
Gueorguiev &
Bekchiev, 2009
3
Oxyomus sylvestris
(Scopoli, 1763)
P-W-4
7.05.-18.06.2006
Present paper
4
Familia
CARAB1DAE
Abax ( Abacopercus )
carinatus carinatus
(Duftschmid, 1812)
SP-2; V-N-2
P-N-l;
V-SL-4
6.6.-6.9.2006; 30.4.-
3.6.2006; 7.9.2006;
6.6-02.10.2013
Langourov et al.,
2014;
present paper
5
Amara (s.str.) saphyrea
Dejean, 1828
SG-1
SG-2
29.4.-29.5.2006
28.12.2006-20.04.2007
Present paper
6
Aptinus (s.str.) bombarda
(Illiger, 1800)
V-N-l
V-N-2&3
3.6.-25.7.2006; 30.4.-
3.6.2006; 5.11.2006-
6.6.2007
Langourov et al.,
2014
7
Bembidion ( Peryphanes )
dalmatinum dalmatinum
Dejean, 1831
V-SL-3
6.6-2.10.2013
Langourov et al.,
2014
8
Blemus discus discus
(Fabricius, 1792) - Fig. 3
V-SL-4
6.6-2.10.2013
Langourov et al.,
2014
9
Carabus ( Procrustes )
coriaceus cerisyi
Dejean, 1826
SG-1
6.8.2006-18.11.2006
Present paper
10
Cychrus semigranosus
balcanicus
Hopffgarten, 1881
V-SL-3
6.6-2.10.2013
Langourov et al.,
2014
11
Duvalius ( Paraduvalius )
regisborisi (Buresch, 1926)
SP-1
6.6.-6.9.2006
Present paper
12
Harpalus (s.str.)
atratus Latreille, 1804
SP-2
6.6.-6.9.2006
13
Laemostenus (Actenipus)
plasoni ( Reitter, 1885)
P-N-l
P-S-l
P-S-2&3
7.9.2006- 3.7.2007;
9.2006- 4.7.2007;
4.7.-17.10.2007
Present paper
14
Laemostenus
( Antisphodrus ) sp.
EG-1&2
25.06.-2.12.2006
Present paper; new
subgenus to the
fauna of Bulgaria
15
Laemostenus ( Pristony -
chus) cimmerius weira-
theri J. Muller, 1 932
V-SL-1-2
V-SL-4
6.6- 2.11.2013;
6.6- 2.10.2013
Langourov et al.,
2014
16
Laemostenus ( Pristony -
chus) terricola punctatus
(Dejean, 1828)
V-N-l; V-N-l;
V-W-2; V-W-3;
SG-4; BK-mssl
Du-mss4
3.06. -25.7.2006;
5.11.2006-6.6.2007;
26.8. -3.12.2006;
26.8.2006;
6.8. -18.9.2006;
3.06. -25.07.2006
26.08. -3.12.2006
Langourov et al.,
2014 and present
paper
17
Leistus ( Pogonophorus )
rufomarginatus
(Duftschmid, 1812)
V-SL-4
6.6-2.10.2013
Langourov et al.,
2014
Table 2 (1/6). List of the registered edaphicolous and hypogeicolous Coleoptera from MSS- traps.
First purposive study of beetles (Coleoptera) from endogean environments in Bulgaria: collection sites and preliminary results 333
Family
Species and subspecies
Trap No.
Collection date
References
18
Familia
CARAB1DAE
Leistus ( Pogonophorus )
spinibarbis rufipes
Chaudoir, 1843
V-SL-5
2.11.2013-26.6.2014
Present paper
19
Molops (s.str.)
alpestris rhilensis
Apfelbeck, 1904
P-N-l
P-S-l
WR-2
7.9.2006- 3.7.2007;
9.2006- 4.7.2007;
3.4.-9.6.2006
Present paper
20
Molops (s.str.) dilatatus
dilatatus Chaudoir, 1 868
WR-1
23.4.2006-9.6.2006
Present paper
21
Molops (s.str.) piceus
bulgaricus Maran, 1938
V-N-2&3
5.11.2006-6.6.2007
Langourov et al.,
2014
22
Myas (s.str.) chalybaeus
(Palliardi, 1825)
SP-2
6.6.-6.9.2006
Present paper
23
Platynus proximus
(J. Frivaldszky, 1879)
SP-1
06.06.-06.09.2006
Present paper
24
Pterostichus (s.str.) mer-
klii (J. Frivaldszky, 1879)
SP-1
6.6.-6.9.2006
Present paper
25
Pterostichus (Petrophi-
lus) melanarius melana-
rius (Illiger, 1798)
V-SL-4
V-SL-5
V-SL-6
6.6-2.10.2013;
2.10- 2.11.2013;
2.10- 2.11.2013
Langourov et al.,
2014
26
Pterostichus (Platysma)
niger (Schaller, 1783)
V-N-l
V-N-2&3
3.6.-25.7.2006;
5.11.2006-6.6.2007
Langourov et al.,
2014
27
Syntomus pallipes
(Dejean, 1825)
SG-1
29.4.-29.5.2006
Present paper
28
Synuchus vivalis
(Illiger, 1798)
SG-4
6.8.-18.9.2006
Present paper
29
Tapinopterus (s.str.)
balcanicus
Ganglbauer, 1891
V-N-l
V-N-2
V-N-2 & 3
WR-1
WR-3 & 4
3.6. -25.07.2006;
30.4. -3.6.2006;
5.11.2006-6.6.2007;
9.6. -17.7.2006;
23.4. -9.6.2006
Langourov et al.,
2014: present
paper
30
Tapinopterus (s.str.)
cognatus kalofirensis
Maran, 1933
SP-2
6.6.-6.9.2006
Present paper
31
Trechus (s.str.)
austriacus Dcjean, 1831
V-W-l
P-E-2
SG-1
SG-2
V-SL-1-2
V-SL-4
V-SL-5
24.6.-3.12.2006;
7.9.2006-4.7.2007;
29.4.-29.5.2006;
18.9.-28.12.2006;
6.6-2.11.2013;
2.10- 02.11.2013;
2.10- 02.11.2013
Langourov et al.,
2014; present
paper
32
Trechus (s.str.) irenis
Csiki, 1912
V-SL-4
6.6-2.10.2013
Langourov et al.,
2014
33
Trechus (s. str.)
subacuminatus A. Fleischer,
1898
EG-1&2
11.06.-25.06.2006/
2.12.2006-18.04.2007
Present paper
New species for
Bulgaria.
34
Trechus (s. str.) subnotatus
Dejean, 1831
SG-3
18.11.-28.12.2006
Present paper
Table 2 (2/6). List of the registered edaphicolous and hypogeicolous Coleoptera from MSS- traps.
334
Rostislav Bekchiev & Borislav Gueorguiev
Family
Species and subspecies
Trap No.
Collection date
References
35
Familia
CLAMBIDAE
C Iambus sp.
EG-1&2
P-S-4&5
2.12.2006- 8.4.2007;
7.12.2006- 9.4.2007
Present paper
36
Familia
CORYLOPH I DAE
Sericoderus lateralis
(Gyllenhal, 1827)
SG-3
20.4.-1.5.2007
Langourov et al.,
2014
37
Familia
CURCULIONIDAE
Acalles sp.
P-E-1&2
7.9.2006-4.7.2007
Present paper
38
Brachysomus sp.
WR-1
1.4.-25.11.2007
Present paper
39
Dodecastichus geniculatus
(Germar, 1817)
EG-1&2
25.6.-2.7.2006
Present paper
40
Dodecastichus obsoletus
(Stierlin, 1861)
EG-1&2
25.6.-2.7.2006
Present paper
41
Otiorhynchus (s.str.)
albidus Stierlin, 1861
P-S-4&5
19.8.-15.11.2007;
19.5.-13.7.2007
Present paper
42
Otiorhynchus (s.str.)
balcanicus Stierlin, 1861
V-W-3
P-S-4&5
26.8.2006; 23.6.-
7.7.2006; 7.12.2006-
19.4.2007;
13.7. -19.8.2007;
19.8. -15.11.2007
Langourov et al.,
2014; present
paper
43
Otiorhynchus (s.str.)
bisulcatus (Fabricius, 1781)
V-W-2
EG-1&2
26.7-26.8.2006;
25.6.-2.7.2006
Langourov et al.,
2014;
present paper
44
Otiorhynchus (s.str.)
coarctatus Stierlin, 1861
V-W-2
26.7.2006
Langourov et al.,
2014
45
Otiorhynchus (s.str.)
corneolus Weise, 1906
V-W-l
V-W-2
V-W-3
EG-1&2
24.6. -3.12.2006;
26.7-26.8.2006;
4-16.6.2007;
25.6. -2.7.2006
Langourov et al.,
2014;
present paper
46
Otiorhynchus (s.str.)
crataegi Germar, 1 824
V-W-2
26.7.2006
Langourov et al.,
2014
47
Otiorhynchus (s.str.)
juglandis Apfelbeck, 1 895
V-W-2; V-W-3
SG-1
SG-2
P-E-1&2
P-S-2&3
P-S-4&5
WR-3&4
26.7.2006; 26.8.2006;
5-20.8.2007; 29.5-
17.6.2006; 7.9.2006-
4.7.2007;
4.7.-17.10.2007;
23.6.-7.7.2006;
1.4.-25.11.2007
Langourov et al.,
2014;
present paper
48
Otiorhynchus (s.str.)
ovalipennis Boheman, 1 843
P-S-4 & 5
23.6.-7.07.2006;
7.12.2006-19.4.2007;
19.5.-13.7.2007;
19.8.-15.11.2007
Present paper
49
Otiorhynchus ( Podoro -
pelmus) aff. metsovensis
Magnano, 1999
P-S-2 & 3
4.7.-17.10.2007
Present paper;
probably new
species
50
Otiorhynchus ( Zustalestus )
consobrinus Reitter, 1913
P-S-l
4.7.-17.10.2007
Present paper;
new subgenus
for Bulgaria
Table 2 (3/6). List of the registered edaphicolous and hypogeicolous Coleoptera from MSS- traps.
First purposive study of beetles (Coleoptera) from endogean environments in Bulgaria: collection sites and preliminary results 335
Family
Species and subspecies
Trap No.
Collection date
References
51
Familia
Stomodes rotundicollis
P-S-2&3
4.7.-17.10.2007
Present paper
CURCULIONIDA E
Frivaldszky, 1880
52
Sitophilus ory’zae
(Linnaeus, 1763)
P-S-2&3
4.7.-17.10.2007
Present paper
53
Tychius sp.
P-S-4&5
23.6.-7.7.2006
Present paper
54
Familia
Hylaia reissi
WR-2
9.06.-19.07.2006
Present paper
LNDOMYCHIDAL
Csiki, 1911
EG-1&2
25.06.-2.12.2006
P-S-2&3
4.07.-17.10.2007
V-N-l; V-SL-3
5.11.2006-6.06.2007
02.11.2013-30.07.2014
55
Lycoperdina pulvinata
Reitter, 1884
S-l
9.6.2007
Present paper
56
Familia
Abraeus perpusillus
DH-1&2
10-20.5.2007
Present paper
HISTER1DAE
(Marsham, 1802)
57
Familia
Agathidium ( s.str. )
EG-1&2
25.6.-2.12.2006
Gueorguiev &
LEIOD1DAE
bohemicum Reitter, 1884
Belcchiev, 2009
58
Apocatops nigrita
(Erichson, 1837)
EG-1&2
25.6.-2.12.2006
Present paper
59
Catops chrysomeloides
V-SL-4
6.6-2.10.2013
Langourov et al.,
(Panzer, 1798)
2014
60
Catops fuliginosus
V-W-3
26.8.-3.12.2006;
Langourov et al.,
Erichson, 1837
P-N-2
16.6.2006;
2014;
P-S-4&5
27.06.-7.12.2006;
present paper
EG-1&2
25.6.-2.12.2006;
V-SL-4
2.10-2.11.2013
61
Catops grandicollis
SG-1; SG-2
29.04.-29.05.2006;
Present paper
Erichson, 1837
1.05.-25.05.2007
62
Catops neglectus
P-N-2
7.9.2006-3.7.2007;
Gueorguiev &
Kraatz, 1852
WR-2
23.4.-9.6.2006;
Bekchiev, 2009;
EG-1&2
25.6.-2.12.2006;
Langourov et al.,
SG-3
18.11.-28.12.2006;
2014
V-N-2&3
5.11.2006-6.6.2007
63
Catops picipes
V-SL-4
2.10-02.11.2013
Langourov et al.,
(Fabricius, 1792)
2014
64
Catops subfuscus
P-N-2
24.5.-16.6.2006
Gueorguiev &
Kellner, 1846
Bekchiev, 2009
65
Catops tristis
P-N-l
7.9.2006-3.7.2007;
Present paper
(Panzer, 1794)
P-S-l
9.2006-4.7.2007
66
Choleva (s.str.) agilis
V-SL-4
6.6-2.10.2013
Langourov et al.,
(Illiger, 1798)
2014
67
Choleva (s.str.) angustata
SG-1
29.4.-29.5.2006;
Langourov et al.,
(Fabricius, 1781)
V-SL-4
6.6-2.10.2013
2014;
present paper
68
Choleva (s.str.) glauca
P-S-l
9.2006-4.7.2007;
Langourov et al.,
Britten, 1918
V-SL-4
6.6-2.10.2013
2014 and present
paper
69
Choleva (s.str.)
V-SL-4
6.6-2.10.2013
Langourov et al.,
macedonica
Karaman, 1954 - Fig. 4
2014
Table 2 (4/6). List of the registered edaphicolous and hypogeicolous Coleoptera from MSS- traps.
336
Rostislav Bekchiev & Borislav Gueorguiev
Family
Species and subspecies
Trap No.
Collection date
References
70
Choleva (s.str.) oblonga
Latreille, 1807
SG-2
20.4.-1.5.2007
Present paper
71
Choleva (s.str.) reitteri
EG-1&2
25.6.-2.12.2006;
Langourov et al.,
Petri, 1915
V-SL-4
6.6-2.10.2013
2014 and present
paper
72
Choleva ( Cholevopsis )
P-E-2; P-S-2;
7.09.2006-4.07.2007;
Present paper
paskoviensis
SG-1
11.2006-4.07.2007;
Reitter, 1913
6.08.-18.09.2006
73
Colon sp.
P-E-2;
7.9.2006-4.7.2007;
Present paper
P-S-4&5;
7.12.2006-19.4.2007;
EG-1&2
25.6.-2.12.2006; 18.4.-
17.6.2007
74
Guerguievella petrovi
Giachino et Gueorguiev,
2007
WR-5
14.7.-13.10.2007
Present paper
75
Hydnobius punctatus
EG-1&2
25.6.-2.12.2006
Gueorguiev &
Flampe, 1861
Bekchiev, 2009
76
Leiodes sp.
P-W-2
7.5.-18.6.2006;
Present paper
P-S-4&5
7.12.2006-19.4.2007;
EG-1&2
25.6.-2.12.2006
77
Liocyrtusa nigriclavis
EG-1&2
25.6.-2.12.2006
Gueorguiev &
(Hlisnikovsky, 1967)
Bekchiev, 2009
78
Nargus (s.str.)
V-E-l
16.4.-15.7.2007;
Langourov et al.,
badius rotundus
P-N-l
7.9.2006-3.7.2007;
2014 and present
Karaman, 1954
EG-1&2
17.6.-9.7.2007;
paper
V-SL-4
6.6-2.10.2013
79
Nargus ( Demorchus ) sp.
V-W-3
26.08.-3.12.2006
Present paper
80
Ptomaphagus (s.str.)
P-S-l
11.06.-25.06.2006;
Langourov et al.,
sericatus
EG-1&2
9.2006-4.07.2007;
2014;
(Chaudoir, 1845)
V-SL-4
25.06.-2.12.2006; 17.06.-
9.07.2007; 02.10-
02.11.2013
present paper
81
Sciodrepoides watsoni
EG-1&2
25.6.-2.12.2006; 30.4.-
Langourov et al.,
watsoni (Spence, 1815)
V-N-2
3.6.2006; 7.9.2006-
2014 and present
P-N-l
3.7.2007; 7.9.2006-
paper
P-N-2
3.7.2007; 29.05.-
SG-3
17.6.2006; 17.6.-
6.8.2006
82
Familia
Rhizophagus
DH-1&2
6.09-3.11.2007
Present paper
MONOTOMIDAE
( Rhizophagus )
ferruginous (Paykull, 1 800)
83
Rhizophagus
V-SL-4
6.6-2.10.2013
Langourov et al.,
( Rhizophagus ) perforatus
Erichson 1845
2014
84
Familia
SCIRTIDAE
Cyphon sp.
SG-1
29.04.-29.05.2006
Present paper
85
Familia
Oryzaephilus surinamen-
V-SL-4
6.6-2.10.2013
Langourov et al.,
SILVANIDAE
^(Linnaeus, 1758)
2014
Table 2 (5/6). List of the registered edaphicolous and hypogeicolous beetle (Coleoptera) taxa from MSS- traps.
First purposive study of beetles (Coleoptera) from endogean environments in Bulgaria: collection sites and preliminary results 337
Family
Species and subspecies
Trap No.
Collection date
References
86
Familia
SILPHIDAE
Silpha obscura orientalis
Brulle, 1832
P-S-2&3
4.07.-17.10.2007
Present paper
87
Familia
STAPH YL1NI DAE
(PSELAPH1NAE)
Batrisodes elysius
Reitter, 1884
P-W-4
6.7.2006
Present paper
88
Bryaxis dalmatinus
(Reitter, 1881)
P-S-4&5
V-SL-1-2
27.6.-7.12.2006;
6.6-2.11.2013
Bekchiev, 2008;
Langourov et al.,
2014
89
Bryaxis beroni Karaman,
1969 -Fig. 5
EG-1&2
23.06.2008
Present paper
90
Bryaxis islamitus
(Reitter, 1885)
P-N-2
4.07.-16.11.2007
Present paper
91
Bryaxis roumaniae
Raffray, 1904
P-E-1&2; P-N-2;
V-SL-3
4.07.-17.10.2007;
4.07-16. 11. 2007/P.S;
02.10-02.11.2013
Langourov et al.,
2014;
present paper
92
Bryaxis nodicornis
Aube, 1833
V-SL-4
06.06-02.10.2013
Langourov et al.,
2014
93
Bythinus acutangulus
lunifer Karaman, 1948
P-N-2
4.07-16.11.2007
Present paper
94
Claviger cf. elysius
Reitter, 1884
P-E-l
07.09.2006- 04.07.2007
Present paper
95
Trimium caucasicum
Kolenati, 1846
P-S-4&5
P-S-6
19.04.-19.06.2007
15.11.2007
Present paper
96
Trimium puncticeps
Reitter, 1880
V-SL-6
07.2014
Present paper
97
Trimium expandum
Reitter, 1884
P-S-4&5
27.06.-7.12.2006
Bekchiev, 2008
98
Tychus apfelbecki
Karaman, 1955
P-E-l
7.9.2006- 4.7.2007
Bekchiev, 2008
99
Familia
ZOPHER1DAE
Langelandia sp.
P-W-2
P-E-l &2
P-S-4&5
DH-1&2
6.7.2006
7.9.2006- 4.7.2007
7.12.2006- 19.4.2007
10-20.5.2007
Present paper
Table 2 (6/6)285-296. List of the registered edaphicolous and hypogeicolous Coleoptera from MSS- traps.
Choleva macedonica is worth mentioning. It has
been described by a single male specimen collected
from the cave of Bela Voda (Karaman, 1954), in the
south of Republic of Macedonia. The cave lies on
the left bank of Vardar River, close to the archeolo-
gical site Proselc at the Demir-Kapija Canyon.
Szymczalcowski (1976) expressed doubts about the
status of C. macedonica and listed it as questioned
synonym of C. sturmi Brisout de Bameville, 1863.
Subsequently the species status of the former was
confirmed (Nonveiller et al., 1999) and since then it
is considered distinct species (Perreau, 2004). C.
macedonica was recently announced from Bulgaria
(Langourov et al., 2014). Based on two male speci-
mens (one of them without head and pronotum), the
record at Popov Izvor Karst Spring (see Table 2)
represents the second finding of the species after the
description and the first one out of Republic of
Macedonia. The study of the aedeagus supports the
view of Karaman (ibid.) that it is a distinct species,
not synonym of C. sturmi.
From an ecological point of view, the most
striking fact to us seems the coexistence of five
species of Choleva (s. str.) at the same place and
probably in the same time (Popov Izvor Karst
spring, N42.50275 E23. 15317, 06.VI-02.X.2013):
Choleva agilis, Ch. angustata , Ch. glauca, Ch.
macedonia, and Ch. reitteri.
338
Rostislav Bekchiev & Borislav Gueorguiev
Figure 2. Habitus of Ataenius horticola ; Figure 3. Habitus of Blemus discus ; Figure 4. Habitus of Choleva macedonica
(scale figs. 2-4: 2.5 mm); Figure 5. Habitus of Bryaxis beroni (scale: 1.0 mm).
First purposive study of beetles (Coleoptera) from endogean environments in Bulgaria: collection sites and preliminary results 339
Staphylinidae ( Pselaphinae )
All species that were captured with MSS traps
usually can be found in leaf litter, rotten wood or
under bark of trees and under stones. Apparently
these species penetrate deep in the soil and some of
them ( Bryaxis islamitus, Batrisodes elysius) can be
found also in caves (Besuchet, 1978, 1993;
Bekchiev, 2011). We could suppose that the reason
for this vertical migration is the alteration of appro-
priate microclimatical conditions (temperature,
humidity) on the surface of the soil during the dif-
ferent seasons. Interesting fact is the founding of
Bryaxis beroni in MSS, up to now this species was
known only from caves (Bekchiev, 2008; Hlavac et
al., 2008).
Curculionidae
Seventeen species of weevils have been caught in
the MSS -trap as eleven of them belong to the genus
Otiorhynchus Germar, 1822. The representatives of
this genus are usually known as wingless rhizopha-
gous. It is worth noting the finding of two taxa. The
first of them is Otiorhynchus consobrinus. It be-
longs to the Balkan endemic subgenus Zustalestus
Reitter, 1912 and is new to the Bulgarian fauna. So
far, this species was known only from Croatia. The
second species deserving attention is Otiorhynchus
(. Podoropelmus ) sp. aff. metsovensis Magnano,
1999. This taxon might belong to a new species for
the science, but additional material and works are
needed to prove it.
other families
Besides species of the above discussed four
families, we found in the traps also representatives
of other twelve families (Table 2).
Among the last species, the most characteristic
endogean element seems to be the genus Lan-
gelandia Aube, 1842. The species from this genus
are always blind and partially depigmented, and
they are collected sometimes sifting soil litter. We
have distinguished at least three morphospecies of
Langelandia as only L. anophtalma Aube, 1842
was hitherto reported for Bulgaria. The material
from this genus will be object of a separate study.
The representatives of Hylaia Guerin-Meneville,
1857 and Lycoperdina Latreille, 1807 (both
endomychids) have been collected also in Bulgaria
shifting leaf litter, and rarely they fall in the pitfall
traps “Barber”. These beetles eat fungi and live in
the ground, so their finding in the MSS-traps was
not a surprise. An interesting fact is the collection
of Ataenius hordeola. The only species from sub-
family Euparinae in continental Europe was only
recently recorded from Bulgaria with detailed data
(Gueorguiev & Bekchiev, 2009).
AKNOWLEDGEMENTS
This work was made possible with funding from
the Ministry of Education and Science, The Na-
tional Science Fund project No. B- 1523/05 “First
investigation of the "milieu souterrain superficiel"
(MSS) in Bulgaria: comparative analysis of the
fauna of silicate and limestone regions based on
selected groups of invertebrate animals”and from
an Operational Programme "Environment 2007-
2013” grant No 5103020-C-001 “State of the
cave fauna with a view to protection of the caves,
superterestrial, subterranean karstic forms and relief
in the south part of Natural Park Vitosha
(Bosneshkikarstic region)”.
REFERENCES
Bekchiev R., 2008. The subfamily Pselaphinae (Coleoptera:
Staphylinidae) of Southwestern Bulgaria I. Historia
naturalis bulgarica, 19: 51-71.
Bekchiev R., 201 1 . A study of the Pselaphinae (Coleoptera,
Staphylinidae) in the Rhodopes Mountain (Bulgaria).
In: Biodiversity of Western Rhodopes (Bulgaria and
Greece) II. Pensoft & National Museum of Natural
History, Sofia: 267-278.
Besuchet C., 1978. Un Bythinus cavernicole nouveau ( B .
hauseri) de la Grece (Coleoptera Pselaphidae).
Annales Musei Goulandris, 4: 263-265.
Besuchet C., 1993. Pselaphides cavernicoles de Grece
(Coleoptera). Biologia gallo-hellenica, 20: 223-229.
Coiffait H., 1963. Coleopteres cavernicoles et coleopteres
endoges. Spelunca, Memoires, 3: 174-180.
Culver D.C. & Pipan T., 2008. Superficial subterranean
habitats-gateway to the subterranean realm? Cave
and Karst Science, 35: 5-12.
Deltshev C., Lazarov S., Naumova M. & Stoev P., 2011.
A survey of spiders (Araneae) inhabiting the eueda-
phic soil stratum and the superficial underground
compartment in Bulgaria. Arachnologische Mittei-
lungen, 40: 33-46.
340
Rostislav Bekchiev & Borislav Gueorguiev
Genest L.-C, 1983. Une nouvelle espece de Paraduvalius
(Coleoptere Trechinae) de la Stara planina (Bulgarie.
Memoire de Biospeologie, 10: 315-316.
Genest L.-C. & Juberthie C., 1983. Description de
Paraduvalius rajtchevi n. sp. (Coleopteres Trechinae)
du milieu souterrain superficiel des Rhodopes
centraux (Bulgarie). Memoire de Biospeologie, 10:
311-314.
Giachino RM. & Gueorguiev B., 2006. Gueorguievella
nov. gen .petrovi n. sp. of Leptodirinae from Bulgaria
(Coleoptera, Cholevidae). Fragmenta entomologica,
38: 55-63.
Gueorguiev B., 2011. New and interesting records of
Carabid Beetles from South-East Europe, South-West
and Central Asia, with taxonomic notes on Ptero-
stichini and Zabrini (Coleoptera, Carabidae). Linzer-
biologische Beitrage, 43: 501-547.
Gueorguiev B.V. & Bekchiev R., 2009. A contribution
to the coleopteran fauna of Bulgaria (Insecta:
Coleoptera). Acta zoologica bulgarica, 61: 39-44.
Hieke F. & Wrase D.W., 1988. Faunistik der Laufkafer
Bulgariens (Coleoptera, Carabidae). Deutsche
Entomologische Zeitschrift (N.F.), 35: 1-171.
Hlavac R., Ozimec R. & Pavicevic D. 2008. Catalogue of
the troglobitic Pselaphinae (Coleoptera, Staphylin-
idae) of the Balkan Peninsula, with a key to genera
In: Pavicevic D.M.P. (Eds.). Advances in the studies
of the fauna of the Balkan Peninsula Papers dedicated
to G. Nonveiller. Institute for Nature Conservation of
Serbia, Belgrade: pp. 307-328.
Hurka K., 1990. Duvalius (Paraduvalius) hanae sp. n.
(Coleoptera, Carabidae, Trechini) from the Stara-
planina Mts, Bulgaria. Acta Entomologica Bohemoslo-
vaca, 87: 349-351.
Janak J. & Moravec P, 2008. Drei neue Duvalius-ArtQn
aus Bulgarien und Serbien (Coleoptera: Carabidae:
Trechinae). Klapalekiana, 44: 1-19.
Juberthie C., Delay B. & Bouillon M., 1980. Extension
du milieu souterrain en zone non-calcaire: description
d’un nouveau milieu et de son peuplement par les
Coleopteres troglobies. Memoires de Biospeologie,
7: 19-52.
Juberthie C. & Delay B., 1981. Ecological and Biological
Implications of the Existence of a “Superficial Under-
ground Compartment”. Proceedings of the Eight
International Congress of Speleology, 1: 203-206.
Juberthie C., 2000. The diversity of the karstic and
pseudokarstic hypogean habitats in the world. In:
Wilkens H., Culver D.C. & Humphreys W.F. (Eds.).
Subterranean Ecosistems. Amsterdam: Elsevier, pp.
17-39.
Karaman Z., 1954. Weitere Beitrage zur Kenntnis der
mazedonischen Coleopteren-Fauna. Acta Musei
Macedonici Scientiarum naturalium, 2: 65-91.
Knirsh E., 1930. Novyslepy Carabid z Macedonie. Eine
neue blinde Carabiden art aus Mazedonien. Sbomik
Entomologickeho Oddeleni Narodniho Musea v
Praze, 8: 51-52.
Laneyrie R., 1960. Resume des connaissances actuelles
concernant les coleopteres hypoges de France.
Annales de la Societe entomologique de France, 129:
89-149.
Langourov M., Lazarov S. , Stoev P, Gueorguiev B.,
Deltshev Ch., Petrov S.A. B., Simov N., Bekchiev
R. , Antonova V., Ljubomirov I.D.T. & Georgiev D.,
2014. New and interesting records of the MSS and
cave fauna of Vitosha Mt., Bulgaria: In: Zhalov A.,
Ivanov I., Petrov I. (Eds.). Balkan Speleological
Conference “Sofia’2014”, Sofia, Bulgaria, 28-30
March 2014. Caving Club “Helictite”- Sofia, Sofia,
pp. 66-76.
Newton A. F. Jr., 1998. Phylogenetic problems, current
classification and generic catalog of world Leiodidae
(including Cholevidae). In: Giachino P. M. & Peck
S. B. (Eds.). Phylogenyand Evolution of Subterranean
and Endogean Cholevidae (= Leiodidae Cholevinae).
Proceeding of a Symposium (30 August, 1999,
Florence, Italy) XX International Congress of
Entomology. Atti del Museo Regionale di Scienze
Naturali, Torino, 1998: pp. 41-177.
Nonveiller G., Pavicevic D. & Popovic M., 1999. Les
Cholevinae des territoires de l’ancienne Yugoslavie
(excepte les Leptodorini) (Coleoptera, Staphyl-
inoidea, Leiodidae). Apergu faunistique. Zavod za
zastitu prirode Srbije; Posebnaizdanja br. 18;
Beograd: 128 pp.
Ortuno V.M., Jimenez- Valverde A., Sendra A., Perez-
Suarez G. & Herrero-Borgonon J.J., 2013. The
“Alluvial Mesovoid Shallow Substratum”, a New
Subterranean Habitat. PLoS ONE 8(10): e76311.
doi: 10. 137 1/journal.pone. 00763 1 1
Perreau M., 2004. Leiodidae. In: Lobl I. & Smetana A.
(Eds.). Catalogue of Palaearctic Coleoptera, Vol. 2:
Hy drophilo idea-Histero idea- Staphy lino idea.
Stenstrup: Apollo Books, pp. 133-203.
Ruffo S., 1959. La fauna delle caverne. In: “La Fauna”
(Conosci F Italia), vol. III. Touring Club Italiano,
Milano, pp. 154-164.
Szymczakowski W., 1976. Remarques sur la taxonomie
et la distribution des Catopidae (Coleoptera)
paleartiques. Acta Zoologica Cracoviensia, 21: 45-
71.
Biodiversity Journal, 2015, 6 (1): 341-352
Monograph
Ground beetles (Coleoptera Carabidae) diversity patterns in
forest habitats of high conservation value, Southern Bulgaria
Rumyana Kostova
Sofia University, Faculty of Biology, Department of Zoology and Anthropology, 8 “DraganTzankov” blvd., Sofia 1164, Bulgaria;
e-mail: mmy.kostova@gmail.com
ABSTRACT The study presents a comparison between the diversity of the carabid beetles taxocoenoses and
their spatial distribution in different forest types of high conservation value in Strandzha (8
sites), the Rhodopes (4 sites) and Belasitsa (6 sites) mountains. The diversity indices have
demonstrated the highest species richness and the highest diversity values in the riverside sites
of Strandzha Mountain. The lowest species richness has been found in the tertiary relict forest
of oriental beech with undergrowth of rhododendron (Strandzha Mountain) and in the century-
old sweet chestnut forest (Belasitsa Mountain). The lowest values of diversity and evenness
have been found in the beech forest sites in Strandzha and the Rhodopes due to the prevalence
of the Aptinus species. This low diversity is a natural condition for the studied sites. The
classification of the ground beetles complexes from the studied sites by similarity indices and
TWINSPAN has been made. A high level of dissimilarity among the sites has been found,
showing unique species composition and abundance models in each site. Carabid beetles
taxocoenoses in the forests of Strandzha Mountain have shown a low similarity level by species
composition and abundance even in the range of the same mountain. Indicator species have
been shown. The ordination of the carabid complexes has showed that the sites have been
distributed continuously along two significant gradients. The first gradient has been found to
be the altitude (probably due to the temperature conditions) in a combination with the hydro-
logical regime. The second significant gradient probably has been under the complex influence
of the climate conditions and vegetation type.
KEY WORDS Carabidae; diversity; conservation; Bulgaria.
Received 07.09.2014; accepted 30.11.2014; printed 30.03.2015
Proceedings of the 2nd International Congress “Speciation and Taxonomy”, May 1 6th- 1 8th 2014, Cefalu-Castelbuono (Italy)
INTRODUCTION
The present study is a part of the pilot studies of
some indicator species groups as a basis for a long
term monitoring in different forest types of high
conservation value (Natura 2000 sites) in the
Rhodopes, Belasitsa and Strandzha Mountains.
In order to assess the ecosystems before taking
some management decisions there is a need of basic
knowledge of the species compositions and succes-
sional processes of the species assemblages occup-
ying the habitats (Szyszko et al., 2000).
Ground beetles could be a very useful group as
an indicator of the habitat disturbance as well: they
are abundant in most ecosystems; some species
possess strong habitat preferences; most of the
ground beetle species are associated with specific
landscapes and microclimate conditions; they show
rapid response to environmental changes (Pearsal,
2007). Until this study there was scarce information
342
Rumyana Kostova
about carabid beetles’ fauna of Strandzha and
Belasitsa Mountains (Gueorguiev & Gueorguiev,
1995). The diversity patterns and spatial structure
of the ground beetles communities from these
habitats have been unknown as well.
MATERIAL AND METHODS
Study area and sampling methods
The studied sites have been chosen in order to
be representative habitat types for the Rhodopes,
Belasitsa and Strandzha Mountains. The total num-
ber of the studied sites has been eighteen (Table 1,
Fig.l). The description of the sample sites and their
code according to Habitats Directive (Directive
92/43 EEC, EC, 1992) are given in Table 1.
At each site 10 pitfall traps (diameter = 80 mm,
length =110 mm) were set in a line. The conserving
fluid in the traps was propylene glycol. The material
was collected from May to October in the corres-
ponding years shown in Table 2.
Data Analysis
The species richness-number of collected species
in each sample site (S); Shanon’s (H) and Evenness
indexes have been calculated to compare alfa-di-
versity. Chao 1 procedure has been applied to
calculate the expected species richness in the
studied sites (Chao, 2005).
The dominance of the species has been determ-
ined using Pesenco’s logarithmic scale (Pesenko,
1982) and the categories names have been adapted
to Tischler’s dominance categories, (1949): eudom-
inants (very hight abundance), dominants (high
abundance), subdominants (average abundance),
recedents (low abundance) and subrecedents (single
individuals) (Kostova, 2009). Multidimensional
non-parametric scaling (MDS) has been applied to
visualize is similarity distances between the dom-
inance curves of the studied taxocoenoses (Clarke,
1993). Chi-square test has been used to test the
goodness of fit of the studied taxoconoses’ abun-
dance models to the theoretical ones.
Czekanowski-Sorensen and Bray- Curtis simil-
arity coefficients have been used to calculate simil-
arity between carabid taxocoenoses, by species
composition and by relative abundance of the
species respectively. UPGMA method for clustering
has been applied for constructing the dendrograms
(Krebs, 1999). Two way indicator species analysis
(TWINSPAN) for classification of the carabid
beetle complexes has also been performed.
Figure 1. A map of the location of the study sites, S-Bulgaria (Source: Google Earth, 2014).
Coleoptera Carabidae diversity patterns in forest habitats of high conservation value, S-Bulgaria
343
Mountain
Site
Altitude
Characteristic trees
Code HD 92/4 3
Belasitsa
B PI
450
Pia tanus orien talis L inn ae us
92C0 - Platan us orien talis and Liquidambar
oriental is woods
BPICast
400
Pia tonus orien talis
Linnaeus, Casta nea sativa
Miller
92C0 - Platan us orien talis and Liquidambar
oriental is woods
B_Cast_Pl
400
Pia tanus orien talis
Linnaeus, Casta nea sativa
Miller
92C0- Platan us oriental is and Liquidambar
orien talis woods
B Cast
750
Cos tan ea sativa Miller,
Fa gits sylvatica Linnaeus
9260 Ca sta n ea sativa wood s
BF
700
Fa gits sylvatica Linnaeus
(along waterfall)
9110 Luzulo-F age turn beech forests
B_F2
1500
Fagus sylvatica Linnaeus
9110 Luzulo-Fagetum beech forests
Rhodopes
RhQ
1054
Quercus d alec ham pit
Tenore
9 IM0 Pannoman-Balkanic turkey oak-
sessile oak forests
Rh F
1133
Fagus sylvatica Linnaeus
9130 Asperulo-Fagetum beech forests
Rh_F_Ab
1401
Fagus sylvatica Linnaeus,
single trees Picea a hies
K ar s ten , A hies alba Miller
9 130 A sperulo -Fagetum beech forests
Rh_Pic_Ab
1596
Picea a hies Karsten,
Abies alba Miller
9410 AcidophilousP/cea forests of the
montane to alpine levels
Strandzha
S_Q
324
Quercus h artwissiana
Steven, Quercus cerris
Linnaeus
91 M0 *P anno nian-B alkanic turkey oak-
sessile oak forests
S_Q2
15
Quercus frainetto T enore,
Quercus cerris Linnaeus
91 M0 *P anno nian-B alkanic turkey oak-
sessile oak forests
S_Q _F
271
Quercus polycar pa Schur,
single trees Fagus orinetalis
Lipsky
91 M0 *P anno nian-B alkanic turkey oak-
sessile oak forests
S F
401
Fagus orien talis Lipsky
91 SO * Western Pontic beech forests
S_F_Rhod
183
Fagus orien talis Lipsky,
udergrowth Rhododendron
ponticum Linnaeus
91 SO * Western Pontic beech forests
S_Rip
224
Alnus glutinosa Gaertm,
Quercus cerris Linnaeus
9 1 E0 * Alluvial forests with Alnus glutinosa
and Fraximts excelsior
S_Rip2
35
meadow with single trees
Ain us glutinosa G aertn
Salt's sp . , Uglans regia
Linnaeus, Rubussp. near
Quercus sp. forest
91E0 ^Alluvial forests with Alnus glutinosa
and Fraximts excelsior
SLongoz
6
Fraximts angusti folia subsp.
oxycarpa (M.Bieb, ex
Willd.), Alnus glutinosa
Gaertn.
9 1 FO Riparian mixed forests of Quercus
robur , Vim us laevis and Vlmus minor ,
Fraximts excelsior or Fraximts angustifolia,
along the great rivers
Table 1 . Description of the sample sites, S-Bulgaria.
344
Rumyana Kostova
This method makes classification of the
samples, and then uses this classification to obtain
a classification of the species according to their
ecological preferences. It also makes a dichotomy
based on ordination identifying the direction of
variation. It gives an indicator pseudospecies, i.e.
transforms abundance into pseudospecies (Hill &
Smilauer, 2005).
Detrended correspondence analysis (DCA) has
been applied for ordination of the beetle complexes
by sample sites. Data standardization has been
applied for the analysis due to the different duration
of the collecting time. The relative abundance (pro-
portion of the total number of caught individuals)
of the species from a given sample site has been
used to calculate alfa-diversity indices, two way
indicator species analysis and dominant structure
analysis. Mean number of caught individuals per
100 trap/days has been used for cluster and ordina-
tion analysis. The following statistical softwares
were used: Microsoft Excel (Office 2010), Past 3.01
(Hammer & Harper, 2001), Estimate S9.1.0
(Colwell, 2013), Primer 6 (Clarke & Gorley, 2006),
WinTWINS 2.3 (Hill & Smilauer, 2005).
RESULTS
Eleven thousand eight hundred and seventy-six
individuals belonging to one hundred twenty-eight
species have been collected (Tables 2, 3). Only six
species have been common to the three mountains:
Calosoma sycophanta, Carabus convexus, C. in-
tricatus, C. coriaceus, Pterostichus niger and Myas
chalybaeus (Fig. 2).
The highest species richness of ground beetles
has been shown in the riparian site with meadow
and single trees (Strandzha)- 45 species. Relatively
high species richness has also been demonstrated in
the riparian sites of Strandzha with rich herbaceous
undergrowth. The lowest species number has been
found in the tertiary relict forest of Fagus orientalis
with undergrowth of Rhododendron ponticum
(Strandzha), 8 species and in the centuries-old
forest of Castanea sativa (Belasitsa), 9 species.
Relatively low species richness has also been found
in the carabid taxocoenoses from the sample sites
with altitude above 1400 m (the Rhodopes and
Belasitsa Mountains) (Fig. 3). The species number
of the ground beetles at each site has been actually
Mountain
Year of study N exemplars
NSpecies
Rhodopes
2006,2007 5062
29
Belasitsa
2008,2009 1810
46
Strandzha
2009 5004
92
Total
11876
128
Table 2. A summary table of the collected material, S-Bulgaria.
Figure 2. Species richness (empirical and estimated by Chao 1
procedure) of the ground beetle complexes in the studied sites.
greater, because there have been species that do not
fall into the traps. The estimated species number by
Chaol procedure has been almost the same only for
four of the carabid taxocoenoses with relatively low
species richness. The highest species number has
been estimated for the riparian sites, the oak forests
at the seashore in Strandzha and for the oriental
plane forests in Belasitsa (Fig. 3).
Shanon’s diversity index, fairly sensitive to
actual site differences (Krebs, 1999), has demon-
strated relatively high ground beetles diversity for
all of the studied sites (Figs. 4, 5). An exception has
been the beech forests of the Rhodopes and
Strandzha Mountains due to the prevalence of one
species: Aptinus bombarda and A. cordicollis re-
spectively. The carabid taxocoenose of the century-
old sweet chestnut forest in Belasitsa has shown the
highest value of evenness -0.8. The lowest evenness
has been estimated for the carabid taxocoen-
oses from the beech forest of Strandzha and the
Rhodopes due to the above mentioned prevalence
of the Aptinus species (Fig. 6).
Coleoptera Carabidae diversity patterns in forest habitats of high conservation value, S-Bulgaria
345
O.T
Sites
Figure 3. Diversity of the ground beetle complexes in
the studied sites, estimated by Shanon’s index.
Figure 4. Evenness of the ground beetle complexes
in the studied sites.
E
£
s. Hip:
J_Hfp
S^l
i.f
i_QJ
s_Q
l_F
SL«_Cirt
s_(i_w.a<s
■ EuftcunlrUrtti
■ □osninifltJ
■ Subdtjmina ntS
■ Setedenll
a S*jbr*e*denti
5 10 H> » 49 SO
Numbtrs{l|i«l(i
SO Stress- 0.02
• 5_Kip2
S F*
•S_Longoz
RF« #B_F •S_Q
• BP!
S_F Rh* / 3 -PLCast
B_Casl« •R_Pic_Ab
•B_Q2
_ . #R O
R_F_Ab*
• S_Rip
Figure 5. Dominance structure of the ground beetle
complexes, based on Pesenko’s logarithmic scale.
Figure 6. Dissimilarity distances between the dominance
curves of the studied taxocoenoses, an MDS method.
The dominance structure of the riparian site with
meadow in Strandzha has differed strongly from all
the other with many species represented by single
individuals (Figs. 7, 8). The riparian forest of
Strandzha (S_Rip) has showed a dominance struc-
ture close to the chestnut with oriental plane trees
in Belasitsa (B_Cast_Pl) without eudominants and
more species as dominants and subdominants.
These two sites have one thing in common- through
both of them pass eco-trails. They have demon-
strated Log-series model of the abundance, charac-
teristic for disturbed habitats (B_Cast_Pl: Chi
square = 0.97, p = 0.94; S_Rip: Chi square = 0.98,
p = 0.91). The beech woods with prevalence of the
Aptinus species have also represented a close domin-
ant structure, so as the century-old and the tertiary
relict forests with a small number of species and
high evenness. The classification of the carabid
beetles’taxocoenoses by qualitative and quantitative
similarity coefficients has demonstrated low levels
of similarity for the mountains in general. Four
main clusters have been formed by species compos-
ition (Fig. 9). The similarity by species composition
has been relatively high for the studied carabid as-
semblages from the Rhodopes where they have
formed a separate cluster. A separate cluster, al-
though with low similarity, has been formed by the
periodically flooded riparian sites of Strandzha with
thick herbaceous undergrowth (S_Rip; S JLongoz).
The beech and the chestnut forests of Belasitsa
have also represented a separate cluster. The rest of
the studied ground beetle assemblages have formed
a cluster with low to average similarity between
them. The picture of the clustering based on Bray-
Curtis coefficient has shown more differences bet-
ween the studied carabid assemblages.
346
Rumyana Kostova
oj
G
e
o
0
<J
&
1
£
Ui
c
Si
I
o
CO
4
(A
S
c
3
ffl
8
m
i
(A
trt
tf
3
CD
8
0-r
20 - -
40--
60--
eo--
100 ^-
0t
e 20 +
a
it
QJ
8
>.
40
60
80
100 -L
Samples
Figure 7. A dendrogram for hierarchical clustering of the similarity by species composition of the carabid beetles’ complexes,
an UPGMA method, based on Chekanovski- Sorensen coefficient of similarity. Figure 8. A dendrogram for hierarchical
clustering of the similarity by species abundance of the carabid beetles’ complexes, an UPGMA method, based on Bray-
Curtis coefficient of similarity.
The levels of similarity have been much lower
than by species composition only. There have been
three main clusters: one of the riparian sites of
Strandzha; one of the sites of the Rhodopes and one
of all the other sites. At first level of division TWIN-
SPAN analysis of the ground beetles’taxocoenoses
by sample sites has shown separation of the
Strandzha and Belasitsa low altitude sites from the
other sample sites. The following groups of sites
have been formed at second level of division: 1 .the
Rhodopes sites with altitude above 1000 m and
Belasitsa sites above 700 m; 2. Belasitsa and
Strandzha forest sites up to 450m; 3. the period-
ically flooded riparian sites of Strandzha. The clas-
sification of the species based on their habitat
preferences has also been obtained (Table 3).
The ordination of the carabid assemblages by
DCA has demonstrated two significant gradients
(Eigenvalues: first axis = 0.97, second axis = 0.63,
third axis = 0.34, fourth axis = 0.15). The sample
sites have been arranged along the first axis as
follows: the sites from the Rhodopes (above 1000
m) have been followed by the sites from Belasitsa
in direction higher to lower altitude sites, then the
forest sites from Strandzha and the riparian sites
from the same mountain ending with the period-
Coleoptera Carabidae diversity patterns in forest habitats of high conservation value, S-Bulgaria
347
ically flooded forest along the estuary of Veleka
river with altitude almost at the sea level. The
arrangement along the second axis (gradient) has
separated the Norway spruce forests with altitude
above 1400 meters from all the other sites (Fig. 11).
DISCUSSION
The studied carabid beetles’ taxocoenoses have
demonstrated high species richness and diversity as
a whole. There have been some exceptions like the
low species richness of ground beetles in the old
stable forest ecosystems, which is a natural condi-
tion. The higher species number of carabids in the
open area habitats and cleared forests than in
the old forests is typical for the temperate zone
(Kryzhanovsky, 1983). The low values of diversity
indices and evenness of the beech forests of the
Rhodopes and Strandzha Mountains have been due
to the prevalence of one species: Aptinus bombarda
and A. cordicollis, respectively. This natural condi-
tion had also been found for the beech forests in
Vitosha Mountain, Bulgaria (Popov et al., 1998).
The dominance structure and the abundance mod-
els of the carabid beetles’ associations could be im-
portant indicators for the statement of succession
and disturbance (Hill & Hamer, 1998). Only two of
the studied habitats have shown disturbance by this
estimators, probably due to an anthropogenic disturb-
ance of the often visited by tourists eco-trails in
them. However, the use of the abundance models for
assessment of the ground beetles status, respectively
habitat status, is controversial. One of the reasons is
that there are taxocoenoses with natural conditions
differing from log-normal abundance model, which
is an indicator of natural undisturbed communities.
When chi-square test is used for estimating good-
ness of fit to the theoretical models, there appears
another problem. This test has low power and cannot
be used for small samples (for example sites with
low species number cannot be tested), so as for the
different abundance models it has a different power,
and the results of p - value should not be used for
comparisons between the goodness of fit to the
different models (Hammer et al., 2001). Then
Kolmogorov-Smimov one sample test could also be
used. The classification of the studied sites has
shown unique species composition and abundance
of the ground beetle assemblages even within the
range of one mountain. The unique indicator
carabid species and pseudospecies (with trans-
formed abundance) for the studied sites have been
estimated by TWINSPAN analysis. An indicator
pseudospecies could be those with category above
2 (abundance above 5 %), they have to be abundant
enough to be easily found and collected.
As a result, the following indicator species could
be used for the studied taxocoenoses: Cychrus
semigranosus balcanicus and Carabus hortensis
have been found as indicators for the high altitude
beech and Norway spruce forests, Calathus metal-
400 ^h
350 -
300
CM
</>
250 -
3 200
150
100
50
0
Pic Ab
*kh F Ab
^_F2
% F
Q
*8 p
^_Cast*s_G
f e?2 d
S_Q_F
*^_Rip2
^S_lorgoz
*Rh F
160
320
480
640
Axis 1
800
960
1120
1280
Figure 9. Detrended
correspondence ana-
lysis (DCA) ordina-
tion diagram of the
carabid beetles’ com-
plexes.
348
Rumyana Kostova
Species
' !
j£ f
3C I
5
a
Sample sites
ss
si
si
O'
it.
&
Ll.
tf; 1
^ i- l
O'
c#3
s
54
c/5
Species
division
levels
Platydents ruftts Du ft schmid, 1812
- !: 4
II
Op ho ruts laticollis
Mannerhdm, 1 825
it
Carabus mtricatus Linnaeus, 176 1
Tapinopterus bakanicus
beiasicensis M aran , 1 933
4
tout
Laemostenm t err kola pane fat us
Dei can. 1828
lom
PterostU ch us vecors ( T sch i t seller i ne,
1 897)
101 HI
Pferost kbits truck i Scftaum, 1859
* IOT 10
Pferost ic 7j i ts brevis
(Duftschmid. 1812)
*10110
Plat y nus scrobiculatus
( Fahricius, 1801}
lOtIO
Opium us schaubergerianus
(Pud, 1937)
* 101 to
Mo fops ntfipes beiasicensis
Mlynar, 1977
! 5
*10110
L eistus magn ico 1 1 is
Motschuisky, 1 866
*10110
Lebia cyanocephala
(Linnaeus, 1758)
101 to
Harpalus t riser iat us Fiieseher, 1 897
*10110
Harpalus griseus ( Panzer, 1797)
*10110
Synuchus viva l is (Miger, 1 798)
JOtOll
C ye hr us semigranosus haicankiis
HopfTgartcn, 1881
r m i o 1 1
Pterostkhus oblongopunctatus
( Fabric i us, 1787)
KHOIO
Carabus violaceus azurescens
Dejcan. 1826
5
101010
MVJWOIU I
Carabus ho/ tens is Linnaeus, 1758
Xenion ignition (Kraal/, 1875)
1 Ik J I -■» If 1 i r 3 I » ■ J ri / M i. 11 I .-I ■ f mn
101010
m ■ lb- p i 1 m * r v w ■ pit ■ «p ■ ■ Y K ® .■ is,
Aphmpterus ha lean kus
Ganglbauer, 1891
5 j
— r -
101001
101001
Notioph i fits b iguttati is
(Fabricius, 1779)
JOIdOi
Mo lops rhodopens is
Apfclbeck, 1904
loioot
Mo lops diktat us Chaudoir. 1 868
5 ! -
101001
Molops alpestris (Dejean. 1 828)
101001
Mkro/esfes ininufu/us (Goeze, 1777)
101001
Laemostenas t err kola l lerbsl. 1 784
101001
Clivina fbssor (Linnaeus, 1758)
101001
Carabus montivagus bid gar tens
Csiki, 1927
2 ! -
101001
Cola thus mollis (Marsha m. 1802)
10 1 001
Calathus met all kus Dejean, 1828
5 !
101001
Apt inns bombarda (llliger. 1800)
10(001
A box oralis (Dnfischmid. 1812)
101001
Table 3.TWINSPAN analysis’ table of the studied ground beetles’taxocoenoses. Species abundance has been represented
by pseudospecies. Doubled line has shown the first level of division, dotted line has shown the second level of division
(continued).
Coleoptera Carabidae diversity patterns in forest habitats of high conservation value, S-Bulgaria
349
Species
<
it 1
«
Q _
£
Sam pie sites
£2
£2
Vi
O'
1
---
O'
c^ 1
i
i ^
|
I
»
r (
Species
division
levels
Pterostidms wjgc-r (Schaller. 1930)
hi moo
Amaru communis (Panzer, [ 797)
'to 1000
Harpalus rufipes (De Geer 1 774)
* i 3
'too
Ca la thus fuscipes (Gocze, 1777)
2 ; 5
*011
Anisodactil i is hi > lot a fits
(Fabricius, 1787)
*011
Amara aenea (De Geer, 1774)
*011
Trechus quadristriatus
(Schrank, 1781)
2 i 2
*0101
Myas chcilybaetts (Palliardi, 1825)
*0101
Cara bus con vex us Fabric ins, 1775
5
3 ! 2
r
*0101
A box ccirinatm ( Du Itsehtn i d, 1812)
* 010 )
Not ioph Hus nip pcs Curl is, 1829
*0100
Cara hits coriaceus Linnaeus, 1758
*0100
Cal osoma sycophant a
(Linnaeus, 1758)
Harpalus atratus Latreille, 1804
-* — *
4
*0100
*001
Amara saphyreu Dejean, 1828
*001
Amara convex tor Stephens. 1 828
*001
Trechus crucifer B ruled e, 1875
*0010
Pterostich i is properans
(Chaudoir, 1868)
2 ;
*0010
Harpai us calceafi is
(Duftschmid, 1812)
*0010
Molops p teens byz a minus
Apfelbeck, 1902
*0010
Lie jnu.s eass ideas (Fabricius. 1792)
*0010
Laemostenus^ venus fits ( Dejean, 1828)
- ! l
*0010
Laemostenus cimmerius
( F isc her- Wald he i m , 1 82 3 }
*0010
Harpalus sulphitripes Germar, 1824
*0010
Harpai us smaragdimts
(Duftschmid, 1812)
*0010
Harpalus honest us
(Duftschmid, 1812)
*0010
Harpalus froelichi Sturm, 1818
*0010
Chlaenius aenocephctlm Dejean,
1826
*0010
Carabus marietti
Cri stofori et J an , 1837
*0010
Carabus scahrosus Olivier, 1795
*0010
Calosoma inquisitor (Linnaeus,
1758)
*0010
Ca la thus longico/l is M ol sehu I sky,
1864
*0010
Amara tricuspuiata tricuspidata
Dejean. 183 1
*0010
Pterostichus nigrita ( Paykull, 1790)
*0010
Harpalus tardus (Panzer, 1797)
*0010
Dram itts q uadrimaculatus
(Linnaeus, 1758)
*0010
Table 3. TWINSPAN analysis’ table of the studied ground beetles ’taxocoenoses. Species abundance has been represented
by pseudospecies. Doubled line has shown the first level of division, dotted line has shown the second level of division
(continued).
350
Rumyana Kostova
Species
O'
A
cc
<
Lb'
*
i5
3C
Sample sites
lx,
a
02
c.
s.
M5
1
JS
si
Lb'
v; 1
B
,E-'%
2£ i V
o
Species
division
levels
Acupalpus suturalis Dejean, 1829
*001010
i'rcchus sp, {suhnoUitus group}
3 ;
*001001
Parophor ms n mci il icornis
(Duftschroid, 1812)
2 !
*001001
Ophonus simi/is (Dejean, 1 829)
*001001
Ophnnus nithint i ^Stephens, IK2S
2 !
*001001
Notioph i tus patustris
(Duiftschmid, 1812)
2 ;
*00100!
Lets tus rt t fomargina tus L>u ft sc h in i d,
1812
4 ;
*001001
Harpalus jlavicornis Dejean, 1829
*001001
Aptinus cord tool l is Chaudoir. 1 843
*001001
Amara ant hob la Villa, 1833
2 ;
*001001
Harpalus ntbripes
(Duftschmid. 1812
"001000
Ncbria brcvhoUis ( Fabric i us, 1 792)
* 000 ]
Cara bus wiedenun mi
Mcnclrics, 1836
i i 3
*0001
Amara ova fa (habrieius, 1 792)
*0001
/ larpulus serripes (Qucnsel, 1 806)
*(>0001
Harpalus ditniduuus (Rossi. 1790)
- 1 4
*00001
■Si atom us pal iipes ( Dejean, 1825)
2 - 2
*000010
Harpalus alhqnicus Re i tier, 1 900
I ! t
*000010
Bembidion lampros (Herbs!, 1 784)
4 ; 5
* 0000)0
Agon i mi ass ini i la ( Pay kull, 1790)
*000010
Asaphidion jlavipes (Linnaeus. 1761)
4 ! S
*00000 I
Agon um dorsalis
(Pontopippidian, 1763)
4 ; 5
*000001
Trechus obtusus thracicus
Pawiowski, 1973
*000000
Tachys bis trig tus (Duftschmid. 1812)
*000000
Symomus obscurogmtatus
(Duftshmid, 1812)
*000000
Stenolophu smixtus(\ lerhst., 1784)
*000000
Pterostichus stromas (Panzer, 1797)
| 2
*000000
Pterostichus wc/mfCreirtzer, 1799)
*000000
Ft ci -os tick i is melai ran us bulgaric us
Lutshnik, 1915
*000000
Pterostichus icon is i Apfelbeek, 1 904
*000000
Pterostich i ts anthravin us
(llliger, 1798)
*000000
Poecilus cupreus (Linnaeus, 1758)
*000000
Parophot ms com pi at latus
(Dejean, 1829)
! 2
*000000
Fanagaeus entxmajor
(Linnaeus, 1758)
*000000
Ophonus sabulicoia (Panzer, 1 796)
I 5
*000000
Ophon us me/ let i ( H ee r , 1837)
*000000
Oodes gi acilis Villa, 1833
*000000
Harpalus tcnchrasus Dejean, 1829
*000000
Table 3. TWINSPAN analysis’ table of the studied ground beetles ’taxocoenoses. Species abundance has been represented
by pseudospecies. Doubled line has shown the first level of division, dotted line has shown the second level of division
(continued).
Coleoptera Carabidae diversity patterns in forest habitats of high conservation value, S-Bulgaria
351
Species
Sai
np
e sites
Species
division
levels
3
a
Lb
1
■C
X>
2'
£
i
JZ
“l
ifi
ta
3
*
53
S'
u
fis'
s
se‘
35
u
sd
3
Lb
®i
&
U.
*3
a
5
lJ
■P
rm
3
=.
£
a.
be
V.
N
O
SI
c
-3
Harpalus cupreus Dejean, 1 829
5
-
*000000
Harpal i is mi turn mil is
{Duftsehmid, 1812)
1
-
*000000
Harpalus off in is ( Schrank, 1781)
1
-
*000000
Gynandromorphus etruscus
(Quensel, 1806)
i
-
*000000
Dyschirius globosus { i icrbst, 1 783)
5
*000000
Diachrom t is germartus
(Linnaeus, 1758)
-
2
*000000
Chlaenius mgricorn is
(Fabricius, 1787)
-
A
*0000101
Carabus granulatus Linnaeus, 1758
5
*000000
Calathus melanocephalus
(Linnaeus, 1758)
5
-
*000000
Brack inns elegans Chaudoir, 1 842
1
2
*000000
Brachinus crepitans
(Linnaeus, 1758)
2
-
*000000
Bembidion inoptatum Schaum, 1 857
1
4
*000000
Ben i b id ion eiongan t m
t
-
*000000
Bern b id ion andreae ( F a br i e i us 1787)
4
-
*000000
Bern b id ion tethys N eto 1 i tzk y 1926
1
*000000
Bad is ter bipustulatus (Fabricius,
1 792)
2
1
*000000
A n isodactil i is signal! is
(Panzer, 1797)
1
-
*000000
Agomun viduuni (Panzer, 1797)
5
*000000
Agon n in nigrum Dejean, 1 828
2
*000000
Agomun nut Her i Herbst. 1 785
2
-
*000000
Division levels of the sites
s
*
o
*
s
*
o
e
*
s
■*
•fc
s
n
5
*
g
*
g
*
e
E
*
®
in
g
*
s
g
*
s
g
*
g
*
o
*
-*
Table 3. TWINSPAN analysis’ table of the studied ground beetles ’taxocoenoses. Species abundance has been represented
by pseudospecies. Doubled line has shown the first level of division, dotted line has shown the second level of division.
licus has been an indicator for the Norway spruce
forest above 1500 m, Molops rhodopensis has been
found as an indicator species onlyfor the high
altitude Norway spruce forest of the Rhodopes,
Pterostichus brucki, for the high altitude beech
forest of Belasitsa, Platyderus rufus has been uni-
que for the low altitude oriental plane woods,
Pterostichus melanarius bulgaricus, Bembidion
andreae, Calathus melanocephalus, Harpalus
cupreus and Ophonus sabulicola have been an
indicator species for the open area grassy habitats
(S_Rip2), Bembidion andreae has also been an
indicator species only for the riparian meadow,
Poecilus cupreus has also been found as an indic-
ator species for wet grassy habitats like the period-
ically flooded riparian sites of Strandzha, Leistus
rufomarginatus and Trechus sp. (subnotatus group)
have been indicator species for the riparian forest
of Strandzha (S_Rip), Carabus granulatus,
Chlaenius nigricornis, Dyschirius globosus and
Oodes gracilis have been found as indicator species
for the periodically flooded estuary forest of
Strandzha (S_Longoz), Calathus longicollis has
been an indicator species for the Black sea coastal
oak forest, Carabus scabrosus, for the oriental
beech woods of Strandzha.
352
Rumyana Kostova
The ordination of the carabid beetles’ taxocoen-
oses has demonstrated continuous arrangement of
the sites along the first axis (the first gradient). The
first gradient has been found to be the altitude
(probably due to the temperature conditions) in
combination with the hydrological regime
(for example, the periodically flooding of the last
two sites). On this gradient, probably there is a
complex influence of the climate conditions and the
vegetation type. Continuous arrangement according
the temperature conditions had also been found for
the carabid associations of different altitude in
Vitosha Mountain by Popov et al. (1998).
The high conservation value of the studied sites
in the Rhodopes, Belasitsa and Strandzha Moun-
tains has also to be concerned due to the great
diversity of the ground beetles that should be
preserved and monitored. Only the Rhodopes sites
have been under high level of protection as a part
of natural reserves, so as two of the sites in
Strandzha as a part of protected localities. The rest
of the studied habitats from Strandzha and Belasitsa
Mountains have been with low protection status and
therefore threatened by logging.
ACKNOWLEDGMENTS
Field work has been supported by research pro-
jects of the Bulgarian Ministry of Education and
Science (BY-E-8/05, BM-6/2007, DO 02-159/
2008). I would like to thank Dr. Borislav Georgiev
(National Museum of Natural History, Sofia) for
collaboration on the collecting, identification and
counting the ground beetles from Belasitsa Moun-
tain. Thanks also to Dr. Rostislav Bekchiev and Dr.
Elena Tasheva for their company and help in collect-
ing the material. Thanks to Maria Gargova (New
Bulgarian University) for the language revision.
REFERENCES
Chao A., 2005. Species richness estimation. In: Balakrish-
nan N., Read C.B. & Vidakovic B. (Eds.), 2005.
Encyclopedia of Statistical Sciences, Vol. 12, 2nd
edn. Wiley Press, New York, 7909-7916.
Clarke K., 1993. Non-parametric multivariate analyses
of changes incommunity stmcture. Australian Journal
ofEcology, 18: 117-143.
Clarke K. & Gorley R., 2006. PRIMER v6: User
Manual/Tutorial. PRIMER-E, Plymouth.
Colwell R., 2013. Estimate S: Statistical estimation of
species richness and shared species from samples.
Version 9. Available at: http://purl.oclc.org/estimates
European Commission, 1992. Council Directive 92/43
EEC of 21 May 1992 on the conservation of natural
habitats and of wild fauna and flora. Official Journal
L 206, 22/07/1992: 7-50.
Gueorguiev V. & Gueorguiev B., 1995. Catalogue
of theground-beetles of Bulgaria (Coleoptera: Cara-
bidae). Sofia, Pensoft, 279 pp.
Hammer 0., Harper A. & Ryan R, 2001. PAST: Paleon-
tological statistics software package for education
and data analysis. Palaeontologia Electronica 4: 9 pp.
Available at: http://palaeo-electronica.org/2001_l/
past/issue 10 1 .htm
Hill J. & Hamer K., 1998. Using species abundance
models as indicators of habitat disturbance in trop-
ical forests. Journal of Applied Entomology, 35:
458-460.
Hill O. & Smilauer R, 2005. TWINSPAN for Windows
version 2.3. Centre for Ecology and Hydrology &
Universityof South Bohemia, Huntingdon & Ceske
Budejovice
Kostova R., 2009. The Ground Beetles (Coleoptera,
Carabidae) in Two Biosphere Reserves in Rhodope
Mountains, Bulgaria. Acta Zoologica Bulgarica, 61:
187-196.
Krebs Ch., 1999. Ecological Methodology, 2nd ed.
Addison- Welsey Educational Publishers, Inc., Menlo
Park, CA. 620 pp.
Kryzhanovsky O., 1983. Beetles of suborder Adephaga:
families Rhysodidae, Trachypachidae; family Cara-
bidae (introduction, survey of the fauna of SSSR).
Fauna SSSR, Coleoptera (Volume I, Ed. 2). Nauka
publishers, Leningrad, 342 pp. (in Russian).
Pearsal A., 2007. Carabid beetles as Ecological indicat-
ors. Paper presented at the “Effectiveness of Biolo-
gical Conservation” conference, 2-4 November
2004, Richmond, BC.
Pesenko J., 1982. Principles and methods of quantitative
analysis in faunistic studies. Nauka publishers,
Moscow, 287 pp. (In Russian).
Popov V., Krusteva I. & Sakalian V., 1998. Some aspects
of coexistence pattern in forest carabid guilds
(Coleoptera: Carabidae) on Vitosha mountain. Acta
Zoologica Bulgarica, 50: 79-88.
Szyszko J., Vermeulen H., Klimaszweski K., Abs M. &
Schwerk A., 2000. Mean Individual Biomass (MIB)
of ground beetles (Carabidae) as an indicator of the
state of the environment. In: Brandmayr R, Lovey G.,
Zetto Brandmayr T., Casale A. & Vigna Taglianti A.
(Eds.), 2000 Natural History and Applied biology of
Carabid beetles. Pensoft, Sofia-Moscow, 289-294.
TischlerW., 1949. GrundzugederterrestrischenTieroko-
logie. Friedr.Vieweg & Sohn, Braunschweig, 220 pp.
Biodiversity Journal, 2015, 6 (1): 353-364
Monograph
Mollusc assemblages of hard bottom subtidal fringe: a com-
parison between two coastal typologies
Andrea Cosentino & Salvatore Giacobbe*
Department of Biological and Environmental Sciences, Viale Ferdinando Stagno d’Alcontres 31, 98166 S. Agata-Messina, Italy
^Corresponding author, email: sgiacobbe@unime.it
ABSTRACT The mollusc assemblages of subtidal fringe from two different coastal typologies are described
in their qualitative and quantitative features. The large-scale spatial investigation has been
carried out in the lava cliffs of Catania and the conglomerate “beach-rocks” of Capo Peloro
(Messina), whose assemblages have been compared by fourteen shallow sampling stations,
spaced out hundred/thousand meters apart. The similarity/dissimilarity levels of the two
assemblages have been evaluated throughout a set of eighty-six species, exclusive or common
between the two areas. Both the assemblages were characteristic of an impoverished and
highly variable photophilic taxocoenosis. The area was the main discriminating factor that
determined the highest richness and abundance in the rough lava surface. The Catania as-
semblage was more constant in species composition, with presence of exclusive bivalves, cue
of a micro-sedimentary environment. The Messina assemblage was very variable in species
composition, and its structure, dominated by motile gastropods, was evidence of a high energy
environment. Differences in the structure and micro-topography of the natural substratum
from the two areas, besides possible secondary influence of freshwater inputs and wave
exposure, were factors mainly responsible for the observed patterns. The whole data set, with
dominant and accessory taxa, involves a relevant contribution from the deeper subtidal as-
semblage; despite of their ephemeral character, these assemblages contribute to maintain the
local biodiversity on a broader spatial scale.
KEY WORDS Biodiversity; Geographical trend; Mediterranean Sea; Molluscs; Rocky shores.
Received 21.02.2015; accepted 20.03.2015; printed 30.03.2015
Proceedings of the 2nd International Congress “Speciation and Taxonomy”, May 1 6th- 1 8th 2014, Cefalu-Castelbuono (Italy)
INTRODUCTION
It is known that species distribution and related
biodiversity levels are stressed by the interaction
between biotic and abiotic factors, which detennine
a hierarchy of processes that operate at different
spatio-temporal scales (Underwood & Chapman,
1996). Spatial patterns of intertidal hard bottom
assemblages have been widely investigated in the
past, both on a broad geographic (Blanchette et al.,
2008) and local scale (Reichert et al., 2008). In the
Mediterranean basin, despite of the prevalent
microtidal regime, the intertidal zone has received
much attention (Benedetti-Cecchi, 2001; Fraschetti
et al., 2001). By contrast, rocky subtidal as-
semblages have been investigated to a lesser extent,
except for the impacted sessile communities
(Fraschetti et al., 2001) and few groups of vagile
invertebrates, such as polychaetes (Giangrande,
1988; Giangrande et al., 2003).
The mollusc taxocoene, notwithstanding its
relevant diversification and wide geographic and
354
Andrea Cosentino & Salvatore Giacobbe
ecological distribution, has been poorly investigated
in its quantitative aspects, and patterns of spatial
distribution have been rarely described (Chemello
& Milazzo, 2002; Terlizzi et al., 2003). Mollusc
spatial patterns and relationships with substratum
complexity have been locally investigated, for
example in the Aegean Sea (Antoniadou et al.,
2005). Within the subtidal zone, the upper level (the
fringe) has received a scanty interest in the past, and
the associated mollusc assemblages are probably the
less known from the Mediterranean phytal zones.
The subtidal fringe, characterized by strong
environmental constrains and high levels of envir-
onmental disturbance, shows different degrees of
substratum complexity which might affect the
spatial patterns of flora and fauna (Guichard et al.,
2001). Furthermore, the ephemeral character of the
algal covering might accentuate the spatial and
temporal dynamics of the associated vagile fauna,
as proved for shallower as well as deeper subtidal
assemblages (Benedetti-Cecchi & Cinelli, 1992). In
this respect, the Ionian coasts of Sicily might rep-
resent an appropriate case-study, due to rocky cliff
typologies that are quite different from the northern
coastline (of metamorphic and sedimentary origin)
to the central (of mainly volcanic origin) and
southern (carbonatic origin) coastlines. In this paper
mollusc assemblages of subtidal fringe are investig-
ated from two rocky coasts of different origin,
volcanic and sedimentary, respectively located in
the Strait of Messina and in the northern side of the
Gulf of Catania.
Aims of the present investigation are: i) to de-
scribe the mollusc assemblages, which characterise
the upper subtidal fringe from two different coastal
typologies; ii) to investigate their similarity/
dissimilarity at different spatial scales (kilometers,
hundreds of kilometers); iii) to highlight the main
(a)biotic constrains which may affect the as-
semblage composition and structure.
MATERIAL AND METHODS
Study areas
The study area, which corresponds to the
northern segment of the Ionian coast of Sicily (Fig.
1), has a regular N 30°-trending shoreline, extend-
ing for a total length of 1 07 km from Capo Peloro
(North) to Catania (South). On the basis of geolo-
gical and morphological characters, two sub-
provinces can be distinguished. The northern seg-
ment, consisting of the Ionian side of the Peloritani
chain from Capo Peloro to the city of Riposto (first
75 km), is characterized by Kabilo-Calabride
terraces; the southern segment, made up of the vol-
canites from the eastern flank of Mt Etna, reaches
the city of Catania (Longhitano & Zanini, 2006).
Since it represents a microtidal oceanographic
framework, coastal dynamics are mainly influenced
by waves that approach the coast obliquely, and by
long-shore southward currents, controlled by the
complex hydrological dynamics of the Messina
Strait. Such hydrodynamics interact with clockwise,
offshore circulation of the Ionian Sea.
Wave energy affects the coastline differently,
since northwards it is mitigated by the action of the
Messina Strait tidal currents, whilst southwards the
near shore circulation is often diffracted and inhib-
ited by the great complexity of the volcanic
shoreline, marked by small coastal promontories
and indentations (Figs. 2, 3). In the Sicily side of
the Messina Strait the coastline is almost homogen-
eous with a mid-Pleistocene conglomerate out-
crops, along almost two kilometers of shoreline
(Bottari et al., 2005). Such so-called “beach-rock”,
which represents the sole hard substratum of natural
origin, is frequently connected to artificial break-
waters and other concrete structures (Figs. 4, 5).
Rivers, as the main points of sedimentary input,
are localized in the southern part and don’t affect
the Messina Strait. Freshwater inputs are mostly of
phreatic origin in Catania, whilst in Messina sta-
tions they are mediated by the Capo Peloro Lagoon,
throughout the two canals “Faro” and “Due Torri”.
Sampling and analysis
The sampling strategy has been based on two
levels at different spatial scale (Fig. 1). The first
level (100 km scale) distinguished the two areas of
Messina (Capo Peloro) and Catania (Ognina). At
the second level, seven stations per area have been
located along 1.9 km (Messina) and 5 km of coast
(Catania) respectively, according to the two main
substratum typologies (natural vs. artificial), wave
exposures (exposed vs. sheltered), slope (vertical
vs. horizontal). The presence or absence of fresh-
water inputs was also considered. In spring 2002
Mollusc assemblages of hard bottom subtidal fringe: a comparison between two coastal typologies
355
Figure 1. Study area. Messina (upper pane) and Catania (lower pane) coastlines with sampling distribution.
Figures 2-5. Study site. Figs. 2, 3. Messina shoreline with conglomeratic “beach-rocks”.
Figs. 4, 5. Catania shoreline with basaltic rocks.
356
Andrea Cosentino & Salvatore Giacobbe
two random replicates of 25x25 cm scraped surface
were carried out for each station, ten meters spaced
out (pooled data), in a shallow subtidal fringe from
0 to 0.3 m depth. Substratum typology and algal
covering were preliminarily recorded on field. In
laboratory, samples were washed throughout a
0.250 mm mesh sieve and the retained mac-
robenthic fauna was separated from algae throu-
ghout a manual centrifuge. Particles smaller than
0.250 mm were considered as “sediment” and their
amount evaluated as volume and dry weight
(80°C/24 h). Algae were investigated in their
structure by dominant taxa, fresh and dried total
biomass, total fresh volume and degree of branching,
in accordance with Edgar (1983). Besides, the charac-
terising algae were distinguished in the main func-
tional groups of encrusting, thread-like and
branched thallii, according to Littre & Arnold (1982).
The macro-zoobenthos (>0.250 mm sieved frac-
tion) has been sorted out under the stereomicro-
scope at the Phylum/Classis/Ordo levels. Molluscs
have been determined at the species level, and
respective abundances evaluated. The univariate
and multivariate statistical parameters have been
elaborated by means of PRIMER 6.0 software
package. Main factors potentially affecting as-
semblage composition and structure were selected
a priori and tested by the analysis of similarity
procedure (ANOSIM) for one way and two way
crossed designs. The selected abiotic factors were
the sampling area (two fixed levels), the sampling
station (six random levels), the site exposure (two
fixed levels), the substratum typology (two fixed
levels), slope (two fixed levels), freshwater inputs
(two random levels), entrapped sediment (two
random levels).
The selected biotic factors were the algal cover-
ing (three fixed levels), algal volume (three random
levels), dominant algal taxa (seven random levels)
and the algal functional groups (three random
levels). The similarity percentage analysis (SIMPER)
highlighted for those species that were more
responsible for dissimilarity between areas.
RESULTS
The whole examined sample set provided a total
of 86 species, 46 of which were exclusively collec-
ted in Catania and 2 1 were exclusively recorded in
Messina, while 22 species were common to the two
areas (Table 1). Gastropod species were the most
numerous, with 63 species, 34 of which were col-
lected only in Catania and 18 only in Messina, plus
11 shared species; half of the 18 bivalve species
were exclusively found in Catania, with respect to
the two species exclusively recorded in Messina,
whilst other 7 species were collected in both areas.
Polyplacophora accounted one shared species plus
three taxa exclusively found in Catania and one in
Messina (Fig. 6).
The number of species found in each station
ranged from 7 (MEB) to 24 (MEE1) in Messina
with a tendential north-to-south increase; such trend
was more irregular in Catania, with 15 species in
CT12 up to 37 species in CT6. Likewise the number
of species, the abundances per station were higher
in Catania (min 244, max 1960 individuals) than in
Messina (min 27, max 1349), but they were irregu-
larly distributed and not clearly related to the
number of species, except for MEE1 where the
peaks of the two parameters matched (Fig. 7).
The trend of Margalef’s richness agrees with the
number of species. Univariate diversity indexes
showed different trends between the two areas.
Shannon diversity and species equitability had more
remarkable fluctuations in Messina, ranging from
0.52 (MEE1) to 2.04 (MEC) and from 0.16 to 0.89,
respectively. Diversity in Catania was meanly 1.5
in most stations, except for values 2.1 and 2.2 in
■ Gastropoda BivaMa Polyplacophora
Figure 6. Numbers of Polyplacophora, Bivalvia and Gastro-
poda species exclusively recorded in Messina or Catania,
and shared between the two areas.
Mollusc assemblages of hard bottom subtidal fringe: a comparison between two coastal typologies
357
CD U (N
LU IU LI Q Q LU LU
S S S LU LU UJ HI
j; f I- I- H h
h h K Q U U U
O U O
CD <
LU UJ
U X
LU Q
s tu
CM t- CM
□ UJ UJ
111 LU Ui
O CM "fr CO <£> ^ T-
I- H H u o o O
O O O
d
m < O N T- CM
UJ LU LU Q 0 UJ UJ
S S 5 LU LU liJ lil
o cm oq to 'r
^ ^ H *“
h i- i- o o o o
o o o
CD < G CM r- (\|
LU UJ UJ □ Q uu UJ
S 3 S LU LU UJ LU
O CM ^ 00 CD "t
T" V V" j— |— ]— f—
hhhOOQO
QUO
Figure 7. Trends of species number, abundance, Margalef richness, Shannon diversity, Pielou equitability observed in Ca-
tania and Messina. For each area, sampling stations are ordered from North (left) to South (right).
Figure 8. Nm-MDS ordination plot with superimposed cluster
classification of Catania and Messina stations. Presence (Y)
and absence (N) of freshwater inputs is also showed.
CT6 and CT4 respectively; equitability was meanly
low, likewise in Messina, but with a more regular
trend (Fig. 7).
The multivariate analysis highlighted further
differences in the mollusc assemblages from the
two areas. The Bray-Curtis similarity index and the
related cluster analysis (square root transformed
data, average linkage) discriminated, at lower 28%
level, a first group A of all Catania stations plus two
Messina stations, from a second homogeneous
group B of five stations of Messina (Fig. 8). At a
higher level of 37%, the former group was consti-
tuted by a further sub-group A1 of six Catania
stations, which were separated from a small sub-
group A2 of stations from both areas. Such indica-
tion of a different composition/structure of the
Messina and Catania mollusc assemblages was sup-
ported by the ANOSIM test, which indicated such
area-related discrimination as a statistically signifi-
cant factor (Global E 0.78, p-level 0.1%; number of
permuted statistics greater than or equal to Global
R: 0). Among the other factors that potentially
affected the mollusc taxocoenosis within each area
group, the occurrence/absence of freshwater inputs
(Fig. 8) and “exposure” (exposed/sheltered), were
the most significant.
The ANOSIM test (two way crossed) for differ-
ences between areas across all stations with fresh
water inputs, also resulted statistically significant,
358
Andrea Cosentino & Salvatore Giacobbe
with a Global R of 0.87 (p-level 0.4%; number of
permuted statistics greater than or equal to Global
R: 3). A 2D multi-dimensional scaling better
clustered the stations submitted or not to such con-
straint inside the Messina area, with respect to a
weaker separation inside the other clusters (Fig. 8).
Similarly, test for differences between the
factor “area” across the factor “exposure” provided
a Global R of 0.82, but with a lower significance
level of 1.3% (number of permuted statistics
greater than or equal to Global R: 1). The hypo-
thesis of a possible interaction of the two local
affecting factors was less strictly supported by test
for differences between “freshwater inputs” across
all “exposure”, which provided a Global R of 0.79,
but with p-level 4% (number of permuted statistics
greater than or equal to Global R:l); in this
respect, the general absence of freshwater inputs
in the medium and high exposed stations should
be noted. All the other abiotic and biotic factors,
tested with ANOSIM, did not produce significant
differences among the selected levels.
The Messina assemblage, with a lower average
similarity of 32.9% (Table 1), was characterized by
a small number of species, nine of which accounted
for 9 1 .4% intra-group similarity. Most of similarity
(49.1%) was due to two sole species, Pisinna
glabrata (Megerle von Miihlfeld, 1 824) and Setia
amabilis (Locard, 1886), with 33.7% and 15.4%
respectively. The contribution of other species r api-
dly declined down to 3.1% for Columbella rustica
(Linnaeus, 1758). A residual 8.6% cumulative sim-
ilarity was due to 3 1 rare or occasional species. The
Catania assemblage showed a higher 41.5% sim-
ilarity and was due to a small group of frequent
species, eleven of which accounted for 91.2 cumu-
lative percentage. Most of such a cumulative con-
tribution was due to the species Cardita calyculata
(Linnaeus, 1758), Pisinna glabrata and Barleeia
unifasciata (Montagu, 1803), with an average con-
tribution of 25.7%, 16.6% and 10.8%, respectively.
Differently from Messina, the contribution of other
species slowly declined, down to a minimum of
1.9% of Crisilla galvagni (Aradas et Maggiore,
1844). A consistent group of 54 less common
species covered the residual 8.8%.
The same three species that were main respons-
ible for the internal similarity of the Catania area,
(Cardita calyculata , Pisinna glabrata, Barleeia
unifasciata ) had a primary role to determine the
Figure 9. Nm-MDS ordination bubble-plots for some abundant and rare species which characterized the Messina
and Catania mollusc taxocoenosis with superimposed cluster classification of the two areas.
Mollusc assemblages of hard bottom subtidal fringe: a comparison between two coastal typologies
359
inter-group dissimilarity, although at lower extent,
accounting respectively for 13.5%, 8.9% and 9.5%
of the 79.3% average dissimilarity. Over a total of
86 species, just 40 accounted for 90% cumulative
dissimilarity. It is of interest to note that a small
group of species, Setia amabilis, Crisilla semis-
triata (Montagu, 1808), Lasaea aclansoni (Gmelin,
1791), that weakly contributed to the intra-group
similarity, played a significant role in determining
inter-group dissimilarity (Table 1). Such repartition
of species per area is well represented in the bubble
plots of figure 9, showing the abundances of Setia
amabilis and Barleeia unifasciata, which are highly
characterizing species for the Messina and Catania
assemblages, respectively. Other two less abundant
species, which might be linked to a particular area,
were Crisilla galvagni, exclusively found in
Catania, and Gibbula philberti (Recluz, 1843)
which characterized the Messina area.
More in general, qualitative differences were
recognized in some less frequent taxa that were
exclusively or prevalently found in a single area
(Table 2). Between the Rissoacea, for example,
the genus Alvania Risso, 1826 and Crisilla
Monterosato, 1917 best characterized the area of
Catania (with eight and three species respectively)
with respect to Messina (with two and none species
respectively), whilst the genus Rissoa Desmarest,
1814 (two species) and Setia Adams H. & A., 1854
(two species) were exclusively found in Messina.
Similarly, the genera Granulina Jousseaume, 1888
(three exclusive species) and Gibbula Risso, 1826
(five exclusive species) best characterised Catania
and Messina shorelines, respectively. A possible
vicariant distribution between some congeneric
species was also noted, such as for Tricolia
deshampsi Gofas, 1993 and T. miniata (Monterosato,
1884) sampled only in Messina, with respect to T.
landinii Bogi et Campani, 2007 collected exclus-
ively in Catania.
DISCUSSION
In this investigation, which provided the first
quantitative data on mollusc assemblages from the
Ionian subtidal fringe, a comparison between two
areas, Messina and Catania, quite different in the
typology of the natural substrata, has been carried
out. Their spatial separation (almost 100 Km) and
station distribution (less than 1 km spaced), replic-
ated similar investigations on intertidal communit-
ies which have put in evidence a highest grade of
variability on a ten meter spatial scales (Kelaher et
al., 2001) and even among replicates (Reichert et
al., 2008). Such a local variability, in the present in-
vestigation has been considered in terms of
stochastic patchiness and resolved by replicate
pooling. Such procedure allowed a better discrim-
ination of assemblages per stations and areas.
The fringe, as a peculiar aspect of photophilic
habitat submitted to high levels of environmental
stress (e.g. hydrodynamism, insulation, desiccation,
freshwater inputs), was expected to be characterized
by impoverished assemblages; in contrast, the total
number of recorded species was high, in compar-
ison with deeper subtidal assemblages both from
western (Poulicek, 1985), central (Richards, 1983)
and eastern Mediterranean (Antoniadou et al.,
2005). Number of species and abundance were
markedly higher in Catania with respect to Messina,
probably due to the rough lavic substratum and a
more irregular/uneven shoreline, which increases
space availability, habitat complexity and shelter,
with respect to the smooth conglomeratic beach-
rock and connected concrete blocks.
Although richer in species number and indi-
viduals, the lava cliff did not substantially overlie
the conglomeratic beach-rock in terms of mollusc
diversity, that was moderately high in both substrata
typologies. By contrast, equitability was low, thus
testifying for a generalised de-structured condition
of both the assemblages. In general, the studied
mollusc assemblages showed a high grade of
stochastic variability, especially in Messina, accord-
ing to the wide fluctuation of univariate diversity
indices and to the low internal similarity of each
sample group. Nevertheless, statistically significant
differences between the two areas, mainly due to a
small number of dominant species (Kelaher et al.,
2001) were found. Such differentiation of the two
mollusc assemblages was moderately altered by the
“intrusion” of two Messina stations in the Catania
group, which might be viewed as an evidence of a
cenotic affinity, rather than as an ecological trans-
ition. In this respect, we note that most of the recor-
ded molluscs are known to be ecologically related
to the Mediterranean photophilic algal assemblage
complex, with diversified preference in terms of
depth, light, exposure.
360
Andrea Cosentino & Salvatore Giacobbe
ME
sh. CA
ME
sh. CA
Polyp! acophora
Gastropoda
Cadochiton calculus Dell'Angelo & Palazzi,
1994
Haminoea hydatis (Linnaeus, 1758)
9
Acanthochitana err nit a (Pennant, 1 777)
A ca n t ft oc hi ton a fascicu la to (Linnaeus,
*
Haminoea navicula (da Costa, 1778)
*
1 767)
Lepidochrtona monterosaioi Kaas & Van Belle.
* OK as to mia imp roba hi Its O be r li ng , 1970
m
1981
Leptochiton cimicoides (di Monterosato,
* Mi tra co rnicuta (Linnaeus, 1758)
1879)
* Nat tea rius h ebra ms ( Mart yn . 1786)
Gastropoda
Ocinebrina hispidula ( Pall ary, 1 904)
*
A Ivans a ca need ata (da Costa, 1 778)
* OK as to nidi a Koliofuni (Philippi, 1844)
9
Ah 'an ia cimex ( Linnaeus, 1758)
* Omatogyra atom us (Philippi, 1841 )
9
Alvania clathrellti (Segucnza L., 1903)
*
Partitions i ndeco ra (Bergh, 1881)
*
A l vania lanciae (Cal car a, 1845)
9
Part hen in a c lath rata (Jeffreys, 1848)
Pisinna glabra tit (Megerle von Miihlfcld,
9
Alvania sea bru (Philippi. 1844)
*
- 1824)
9
Alvania simulans Locard, 1 886
* Pho reus rich ardi (Payraudeau, 1826)
Alvania subcrenula fa (B.D.D., 1884)
* Pusil Una margin ata (Michaud, 1830)
Alvania Zetland tea ( Montagu. 1815)
Ammonicera fischerima (Monterosato,
* Rissoa similis Scacchi, 1836
1869)
*
li issoa i r arl ah i lis (Von M uh 1 fc Id l , 1824)
*
Aplysia parvula Mdrch, 1 863 *
Setia a ma hi lis (Locard, 1886)
*
Aplysia fasciat a Poiret, 1789
* Setia sc id tie (A rad as & Benoit, 1 876)
#
Ba fleet a un ifasd ata (Montagu, 1803)
* Sinezona cingulata (O.G. Costa, 1861)
Bit Hum lacteurn (Philippi, 1836)
* Tri col ia ties 1 7? ampsi G ofas , 1993
*
Bit titan reticula turn (da Costa, 1778)
*
Tricvlia miniata (Montcrosato, 1884)
*
Bulla striata Bruguicre, 1 792
* Tricolia lan Kind Bogi & Campani, 2007
Centhiopsis nofronii Amati. 1 987
* Vexidum ebenus (Lamarck, 1811)
Cerithium vulgatum Rruguiere, 1792
Vitreolina incurva ( B, D.D.. 1883)
Vitreolina philippi (de Rayneval & Ponzi.
Coiumbeda rustica (Linnaeus, 1758)
Conus veutri cos its G me 1 in , 1791
1854)
* Wi Ilia mi a gussoni (Costa O. G. , I 829)
Crisilla ben lamina (Monterosato, 1884)
* Bjvajvia
Crisilla gafaagni (Aradas & Maggiorc, 1 844)
* Anonu’a ephippium Linnaeus, 1758
CrisiUa semi striata (Montagu, 1808)
* Area noae Linnaeus, 1 758
Eaton in a pumila (Monterosato, 1884)
m
Barb alia barhatu (Linnaeus, 1758)
Epi torn u in p niched lutii ( B i v on a , 1832 )
* Bruch id antes pharaonis (P. Fischer, 1870)
*
Fissured a mt bead a (Linnaeus, 1758)
* Card it a cahvuluta (Linnaeus, 1 758)
*
Fossa rus am hi git us (Linnaeus, 1758)
* Cha ma grypho id es Linnac u s, 1 75 8
*
Gibber til a Janssen i van Aa risen ct al., 1984
* Hiateda arctica (Linnaeus, 1767)
#
Gib hula ad an son ii (Payraudeau, 1826)
9
Hiatefla rugosa (Linnaeus , 1767)
*
Gihbuta ardens (Salts Marschlins, 1 793) *
bus irus (Linnaeus, 1758)
*
G ib h ul a ph ilberti (Recluz, 1843) *
La sue a adansoni (Gmelin, 1791)
G ib b ula racket t i ( Pa yr aud eau , 1826) *
Lima lima (Linnaeus, 1 758)
m
Gib bid a turbi unities ( Dcshay cs, 1 835)
9
Muscat us cos tula tus ( Risso, 1 826)
9
Gib b ula um hi hearts ( Lin nae us. 1 758) *
My ti taster minimus (Poli, 1 795)
Gib but a varia (Linnaeus, 1758) *
Myfi luster sol i dus M ontcro sato , 1883
9
Gran a lino haucheti Gofas. 1 992
* Mvtilus gal lap ravine ia lis Lamarck, 1819
9
G ran 1 1 li rut m arg in ata ( B i vona , 1832)
Grunulina vanhareni (van Aartsen et al..
* Ost t ea edufis Linnaeus, 1758
*
1984)
* Ostrea stentina Payraudeau. 1826
*
Gy rosea la lamellosa (Lamarck, 1822)
9
St ri urea lav tea (Linnaeus, 1758)
Table 1. Similarity Percentage analysis for species contribution to Bray-Curtis similarity (5”) and dissimilarity ( ' ) within
each area and between the two areas respectively. Av., average; SD, standard deviation; N, abundance of individuals.
Mollusc assemblages of hard bottom subtidal fringe: a comparison between two coastal typologies
361
SIMPER
withm/between
Area-Groups
Group
ME
av. S'
Group
ME
S’
Croup
CT
av. S'
Group
CT
S’
av. 6
5/SD
5
contr.%
32.92
41.53
av. N
av, N
Cardita ca /yen lata
2.51
2.73*
15.64
10.68*
10,65
1,48
13.47
Barleeia un if a sc iata
0.00
9.96
4.48*
7.53
1,11
9.51
Pisinna g! a brat a
10.90
1 1 .09*
11.12
6.89*
7.09
1 .37
8.96
A mmonicera fischerian a
0,25
6.53
3.68*
5.22
0,94
6.59
Acantho chiton a crin i ta
0,29
4,53
3.79*
3.29
2,57
4.16
Myti /aster solidus
1,69
0.14
5.24
2,4*
3.21
0.84
4.06
Gib hula turb in aides
3,54
0.15
0.47
2.73
0,87
3.45
Setia amabilis
3.23
5.07*
0.00
2.71
0.87
3.43
Cris i lia sem is tri at a
0.00
3.79
2.62
0.49
3.32
Lasaea adansoni
0.00
2.57
2.29
0.46
2.89
Bittium reticulatum
0,34
2.62
1.43*
1.93
1.11
2.44
Eaton ina pit mi /a
1.05
0.07
3.37
1.35*
1.90
1.07
2.40
Cris ilia ga/vagni
0.00
2.71
0.06
1.81
0.97
2.29
Cris ilia beniamina
0.00
2.30
1.80
0.83
2.27
C a lum bell a n tsti ca
1.69
0.04
2.58
1.3*
1.73
l .36
2.19
Si n ez ana c i ngu la ta
0.00
1.94
1.12
0.51
1.41
Gib hula adamomi
1,1 1
0.40
1.03
0.77
1.30
Fissure! la it ubecula
0.00
1.45
0,98
1.01
1.24
Mitscu las cost it la (us
1.06
0.07
1.68
1.06*
0.93
1,17
1.18
Area noae
0.74
0.98
0.90
0.95
1.14
Gib hula ardetis
1.10
0.00
0.81
0.75
1.02
Bra chid antes ph ara onis
0.14
1.10
-
0.77
1,05
0.98
Myti las ga l lopro v inc 7 a l is
0,00
0,99
0.75
0,50
0.95
Aplysia panada
0.90
0.05
0.00
0.68
1.08
0.86
Alvania sc ah r a
0.00
1.01
0.65
0.68
0,83
Cham a gryph aides
0,29
-
0.57
0.56
0,74
0.71
Alvania lanciae
0.29
0.64
0.54
0.70
0.68
Gy rose ala lamellosa
0.29
0.52
0.46
0.84
0.58
A ca nthochi ton a fast * icu laris
0.00
0.69
0.45
0.40
0.57
G ih bul a ph Uberti
0.49
0.00
0.44
0.79
0.56
Omalogyra atom us
0.00
0.78
0.42
0.40
0.53
Aplysia fascia ta
0.00
0.53
0.42
0.40
0.53
Trie alia landinii
0.00
0.39
0.39
0.58
0.50
Alvania Simula ns
0.00
0.73
0.39
0.40
0.50
Inis irus
0.00
0.39
-
0.39
0.59
0.49
Gib hula racke d i
0.55
0.00
0.37
0.61
0.47
Ostrea edulis
0.00
0.40
0.36
0,60
0.45
Tricolia m in iata
0.49
-
0.00
0.35
0.60
0.44
Alvania can cel lata
0.00
0.49
0.35
0.81
0.44
Gran id ina v an ha re n i
0.00
0.52
0.34
0,60
0,43
Table 2. Distribution of mollusc species in the shallow sublittoral zones of Messina (ME), Catania (CA)
and shared (sh.) between the two areas.
362
Andrea Cosentino & Salvatore Giacobbe
Such melange of species tied to different envir-
onmental conditions is explicated in literature as a
trapping effect exerted by the branching algae
towards the settling larvae and early juveniles
(Poulicek, 1985). This supposed “branching effect”
did not significantly affect the composition and
structure of the mollusc assemblages, which
clustered independently from typology and extent
of algal covering. Such evidences do not agree with
literature data that indicate a significant effect of
algal architecture on the mollusc assemblage discrim-
ination (Chemello & Milazzo, 2002; Pitacco et al.,
2014; Antoniadou et al., 2005). By contrast, the
hypothesis that redundant algal-associated assem-
blages can play a key role for the maintenance of
biodiversity of the broader geographical area
(Antoniadou et al., 2005) is here supported.
Moreover, contrasting effects of algal covering
towards zoobenthic larvae should be considered,
since the wave exposed fringe is submitted to a
highest sediment/nutrient resuspension that is
known to favour spores with respect to benthic lar-
vae in space competition (Richmond & Seed, 1991;
Oigman-Pszezol et al., 2004). Once developed,
algae might favour larval recruitment and
juvenile/adults surviving, providing food and pro-
tection from predators and desiccation (Poulicek,
1985; Antoniadou et al., 2005), but limitedly to a
short time period, due to their ephemeral character.
Mollusc assemblages that were dominated by
small sized species with a short life span reflected
such an irregular and transient availability of
resources in their high densities and low or-
ganization levels. The algal assemblage, in turn,
is driven by the substratum type and its (micro)to-
pographic complexity, both of which directly and
indirectly affect the settlement and persistence of
benthic organisms. In this respect, the geo-litho-
logical structure of the natural substrata and the
related texture might be explicative for the ob-
served patterns of diversity among the two areas
far apart. The conglomeratic rocks of the Messina
Strait is more even and less porous with respect
to the basaltic surface of Catania cliffs, which is
more irregular and richer in hollows and crevices.
The more uneven surface, promoting phyto-
benthic colonization, in turn improves shelter and
sediment trapping, that is in accordance with the
higher richness of the bivalve species observed in
the Catania area.
The less fluctuating Shannon H’ and equitab-
ility of species/abundance of the Catania mollusc
assemblage with respect to the Messina area, also
support the hypothesis that substratum roughness
acts as structuring factor. Other local factors, both
related or not with the natural substratum typo-
logy, do not play a recognizable role, except for
freshwater inputs combined with shelter expos-
ure. The effects of geographical distance, that
might be more important than substratum type or
roughness in determining assemblage structure
(Guarnieri et al., 2009), might be also considered,
in reason of the latitudinal gradient between the
two areas.
Such a gradient determines a southward temper-
ature increase that near Catania marks the crucial
15°C seawater winter isotherm (Bianchi et al.,
2012). In our opinion, the climatic gradient was not
directly responsible for the quantitative differences
between the two areas, but may partially explicate
the different species composition. Some of the
forty-six (Catania) and twenty-one (Messina) not
shared species have a limited Mediterranean distri-
bution, and some of these species are known from
a restricted area, as the recently “rediscovered”
Crisilla galvagni (Scuderi & Amati, 2012).
CONCLUSIONS
The mollusc assemblages from two subtidal
fringes of eastern Sicily configure as an impover-
ished aspect of the photophilic associated fauna,
submitted to some strong environmental constraints
that limit the number of characteristic species but
allow the transient recruitment of opportunistic and
occasional taxa.
The investigated areas, although submitted to
similar climatic and edaphic conditions, have dif-
ferently structured substrata which result more or
less favourable to species settlement and survival
in accordance with high (Catania) or low (Messina)
cliff roughness. The respective assemblages,
although characterized by a high local variability,
show some common traits that allow to recognize
a real taxocoenosis.
Some scarcely known species, mainly localized
in the lava cliff, may be preferentially tied to such
peculiar environment. The accessory taxa that are
partially supplied by other nearby communities
Mollusc assemblages of hard bottom subtidal fringe: a comparison between two coastal typologies
363
testify for the role of shallow fringe assemblages to
maintain the biodiversity at local and at broader
geographical scale.
REFERENCES
Antoniadou C., Koutsoubas D. & Chintiroglou C.C.,
2005. Mollusca fauna from infralittoral hard substrate
assemblages in the North Aegean Sea. Belgian
Journal of Zoology, 135: 119-126.
Benedetti-Cecchi L., 2001. Variability in abundance of
algae and invertebrates at different spatial scales on
rocky seashores. Marine Ecology Progress Series,
215: 79-92.
Benedetti-Cecchi L. & Cinelli F., 1992. Effects of canopy
cover, herbivores and substratum type on patterns of
Cystoseira spp. Settlement and recruitment in littoral
rockpools. Marine Ecology Progress Series, 90: 183—
191.
Bianchi C.N., Morri C., Chiantore M., Montefalcone
M. , Parravicini V. & Rovere A., 2012. Mediter-
ranean sea biodiversity between the legacy from the
past and a future of change. In: Stambler N. (Ed.),
Life in the Mediterranean Sea: A Look at Habitat
Changes: 1-55.
Blanchette C.A., Miner C.M., Raimondi P.T., Lohse D.,
Heady K.E.K. & Boitman B.R., 2008. Biogeogra-
phical patterns of rocky intertidal communities along
the Pacific coast of North America. Journal of
Biogeography, 35: 1593-1607.
Bottari A., Bottari C., Carveni P, Giacobbe S. & Spano
N. , 2005. Genesis and geomorphological and
ecological evolution of the Ganzirri salt marsh
(Messina, Italy). Quaternary International, 140-141:
150-158.
Chemello R. & Milazzo M., 2002. Effect of algal archi-
tecture on associated fauna: some evidence from
phytal molluscs. Marine Biology, 140: 981-990.
Edgar G.J., 1983. The ecology of south-east Tasmanian
phytal animal communities. I. Spatial organization
on a local scale. Journal of Experimental Biology and
Ecology, 70: 1319-1329.
Fraschetti S., Bianchi C.N., Terlizzi A., Fanelli G., Morri
C. & Boero F., 2001. Spatial variability and human
disturbance in shallow subtidal hard substrate as-
semblages: a regional approach. Marine Ecology
Progress Series, 212: 1-12.
Giangrande A., 1988. Polychaete zonation and its relation
to algal distribution down a vertical cliff in the
western Mediterranean (Italy): a structural analysis.
Journal of Experimental Marine Biology and
Ecology, 120: 263-276.
Giangrande A., Delos A.L., Fraschetti S., Musco L.,
Licciano M. & Terlizzi A., 2003. Polychaete as-
semblages along a rocky shore on the South Adriatic
coast (Mediterranean Sea): patterns of spatial distri-
bution. Marine Biology, 143: 1109-1116.
Guarnieri G., Terlizzi A., Bevilacqua S. & Fraschetti S.,
2009. Local vs regional effects of substratum on early
colonization stages of sessile assemblages. Biofoul-
ing: The Journal of Bioadhesion and Biofilm
Research, 25: 593-604.
Guichard F., Bourget E. & Robert J.-L., 2001. Scaling
the influence of topographic heterogeneity on inter-
tidal benthic communities: alternate trajectories
mediated by hydrodynamics and shading. Marine
Ecology Progress Series, 217: 27-41.
Kelaher B.P., Chapman M.G. & Underwood A. J., 2001.
Spatial patterns of diverse macrofaunal assemblages
in coralline turf and their association with environ-
mental variables. Journal of Marine Biological
Associationof United Kingdom, 81: 917-930.
Littre M.M. & Arnold K.E., 1982. Primary productivity
of marine macroalgal functional-form groups from
southwestern North America. Journal of Phycology,
18: 307-311.
Longhitano S. & Zanini A., 2006. Coastal models and
beach types in ne Sicily: how does coastal uplift
influence beach morphology? Italian Journal of
Quaternary Sciences, 19: 103-117.
Oigman-Pszczol S.S., de O. Figueiredo M.A. & Creed
J.C., 2004. Distribution of benthic communities
on the tropical rocky subtidal of Armagao dos
Buzios, Southeastern Brazil. Marine Ecology, 25:
173-190.
Pitacco V., Orlando-Bonaca M., Mavric B., Popovic A.
& Lipej L., 2014. Mediterranean Marine Science, 15:
225-238.
Poulicek M., 1985. Les mollusques des biocenoses a
algues photophiles en Mediterranee: II. - Analyse
du peuplement.Cahier de Biologie Marine, 26: 127—
136.
Reichert C., Buccholz F., Bartsch I., Kersten T. & Gimenez
L., 2008. Scale-dependent patterns of variability in
species assemblages of the rocky intertidal at Helgo-
land (German Bight, North Sea). Journal of the
Marine Biological Association of the U.K., 88: 13 19—
1329.
Richards G.W., 1983. Molluscan Zonation on Rocky
Shoresin Malta. Journal of Conchology, 3 1 : 207-24.
Richmond M.D. & Seed R., 1991. A review of marine
macrofouling communities with special reference to
animal fouling. Bio fouling: The Journal of Bioadhe-
sion andBiofilm, 3: 151-168.
Scuderi D. & Amati B., 2012. Rediscovery and re-eval-
uation of a“ ghost” taxon:t he case of Rissoa galvagni
Aradas et Maggiore, 1844 (Caenogastropoda
Rissoidae). Biodiversity Journal, 3: 511-520.
Terlizzi A., Scuderi D., Fraschetti S., Guidetti P. & Boero
364
Andrea Cosentino & Salvatore Giacobbe
F., 2003. Molluscs on subtidal cliffs: patterns of
spatial distribution. Journal of the Marine Biological
Association of the U.K., 83: 165-172.
Underwood A.J. & Chapman M.G., 1996. Scales of
spatial patterns of distribution of intertidal inverteb-
rates. Oecologia, 107: 212-224.
Biodiversity Journal, 2015, 6 (1): 365-370
Monograph
On the rediscovery of the vermetid “ Siphonium ” gaederop
Morch, 1861 (Gastropoda Vermetidae) with systematic and
ecological observations on the early juveniles stages
Danilo Scuderi
Via Mauro de Mauro 15b, 95032 Catania, Italy; e-mail: danscu@tin.it
ABSTRACT Some specimens of a not identified Dendropoma Morch, 1862 were collected in the Mediter-
ranean. Further taxonomical studies allowed to identify this material as “ Siphonium ” gaederopi
(Morch, 1861), a species never recorded again after its first description. It is here redescribed
and figured on the basis of the mentioned collected material and after the study of the type
material of Morch’s collection, among which the syntype is here selected. This species is
assigned to Dendropoma , according to the morphological characters of the shell, radula, ex-
ternal soft parts and operculum. The shell, the soft parts and the juvenile stage of D. gaederopi
are here figured for the first time and compared to congeners and to Vermetus granulatus
(Gravenhorst, 1831), similar only in shell morphology. The new findings of this species rep-
resent the first certain record, after the doubtful locality of the original description.
KEY WORDS Mollusca; Vermetidae; Siphonium ; rediscovery; Dendropoma', ecology; juveniles.
Received 03.11.2014; accepted 21.01.2015; printed 30.03.2015
Proceedings of the 2nd International Congress “Speciation and Taxonomy”, May 16th- 18th 20 14, Cefalu-Castelbuono (Italy)
INTRODUCTION
The genus Siphonium Morch, 1859 [not Link,
1807] was synonymysed by Keen (1961), who
restored Dendropoma Morch 1861 as a good name
for the genus, with D. lituella Morch, 1861 as type
species.
Scuderi (1995) had been recently reviewed the
systematyc position of the two known Mediter-
ranean species of the genus Dendropoma : D. pet-
raeum (Monterosato, 1878) is synonymysed with
D. glomeratum Bivona, 1832, while the studied
type material of D. anguliferum (Monterosato,
1884) in the ZMR was not enough to establish
whether this latter is a good species, so it still
continues to have an uncertain systematyc position
because of the lacking of recent material. Some spe-
cimens of a third species of Dendropoma, markedly
different from the two congeneric, has been recently
recorded in the Mediterranean sea. This species
corresponded to the description of “ Siphonium ”
gaederopi Morch, 1861, which was described on
material doubtfully reported from Spain and never
cited in the recent time. The comparison of the spe-
cimens found with the type material of this latter
species, housed in ZMUC, confirmed the previous
diagnosis. A syntype is here selected among the
material of the lot GAS-216.
The shell of adult specimens of D. gaederopi
has a close morphological resemblance with that of
another Mediterranean species, Vermetus granu-
latus (Gravenhorst, 1831).
366
Danilo Scuderi
In particular the former is very similar to a
morphotype of the latter, called “form A” (Scuderi,
1999), but has different protoconch and operculum
and the colour pattern of external soft parts is
peculiar too. Differences on the shell sculpture of
sub-adult shells of the two species (see remarks) are
discriminating too.
MATERIAL AND METHODS
Living samples of D. gaederopi were collected
by undermining the shells from hard substrates at
-4/18 m depth; empty shells and juveniles stages
were collected among the shell grit drawn at -35 m
depth collected handly with ARA. Pictures of the
external soft parts were obteined by observing the
living animals in aquarium.
Type material of “ Siphonium ” gaederopi were
examined from the Morch’s collection stored in
ZMUC.
ACRONYMS. AGC: Alfio Germana collection,
Catania, Italy; ARC: Agatino Reitano collection,
Catania, Italy; DISTEBA: Department of Biolo-
gical and Environmental Science and Technologies,
University of Salento, Lecce, Italy; ISMAR-CNR:
Istituto di Scienze Marine, Consiglio Nazionale
delle Ricerche, Genoa, Italy; ZMR: Zoological
Museum, Rome, Italy; ZMUC: Zoological Museum,
University of Copenhagen, Denmark.
SYSTEMATICS
Dendropoma gaederopi (Morch, 1861)
Examined material. Type material. Four lots
in ZMUC labelled GAS 215/216/217/218, with
two labels, probably in Morch’s handwriting
(Figs. 13, 14) and the label of the Museum (Fig.
15), were constituted by several tens of speci-
mens; one specimen on Spondylus gaederopus
Linnaeus 1758 is here selected as syntype among
the lot GAS 216 (Fig. 1).
Other examined material. Agrigento, SW-Sicily,
Italy: Linosa Is., “Cala Pozzolana di Levante”, -4/6
m, on ancient earthenwares; same locality, 7 living
specimens, -18m, on the shell of a living Charonia
tritonis variegata (Lamarck, 1816); “Faro”, -35 m,
shell-grit, 2 shells; “Punta Arena Bianca”, infralit-
toral shell-grit, 1 shell. Siracusa, SE-Sicily, Italy:
Vendicari, beached shell-grit, 1 shell on the upper
valve of Spondylus gaederopus Linnaeus, 1758;
Marzamemi, 3 living specimens on calcareous
stones, -5/6m (ARC and AGC). Lecce, S-Apulia,
Italy: Otranto, 1 specimen with operculum and 4
protoconchs, on calcareous stones, -5/1 Om.
Description. Shell solid, generally funnel
involved, with the last tele-whorl rounded and
equal to the half of the entire shell (Figs. 2-5).
Sculpture constituted by dorsal longitudinal keels,
variable in number (often 2), which could produce
striking spiny formations in large specimens.
Between the keels, numerous and concentric
lamellae cover the surface of the shell, particularly
in young specimens (Figs. 3, 4). The mouth is
rounded and has a diameter of 2-2.5 mm in adult
specimens (Fig. 5): in the syntype here selected
the external diameter of the aperture is 5.8 mm.
Like in many species of Dendropoma , an eroded
scar is often visible on the substratum around the
apertural opening. The basal portion of the ex-
ternal side of the tube forms a second labial lip,
that leans on the substratum. Dense mass of indi-
viduals were not observed, except one little cluster
among type material (GAS 2 1 6) of about 20 shells
of various size on a fragment of a large Bivalve,
maybe a Spondylus Linnaeus, 1758.
The living animal (Fig. 8) is yellow-cream in
colour, becoming red-orange on the anterior part of
the cephalo-thorax, metapodium and foot, around
the operculum; very small black spots are visible
too. Black shade are present on the dorsal part and
around the base of the cephalic tentacles and the
mouth, wich are both yellow in colour; the mantle
edge is yellow with black alternate lines.
The operculum (Figs. 6, 7) is large as the aper-
ture and is quite different from that of all the
congeners, being reverse-cone shaped, with the
concave part upward, relatively deep and brown
in color, often occupied by encrusting calcareous
algae. The convex downward part is glossy and
red-chestnut in color and do not present, unlike
the two congeners, any depression around
the central large button, but only a thin, almost
undetectable scar.
Protoconch (Figs. 9-11) 0.5 mm high and 0.6
mm wide, swollen, but compressed superiorly and
inferiorly, constituted by 1 and U smooth whorls.
Rediscovery of t/ie“Siphonium” gaederopi (Gastropoda Vermetidae) with observations on the early juveniles stages 367
A first embrional shell, separated from the rest of
the protoconch, is distinguishable (Fig. 10).
Early developmental stages. Usually D.
gaederopi has a not polygirate protoconch: the
presence of a first embryonic nucleus, separated
from the rest of the protoconch shell, suggest that
this species could have a planctonic stage, which
allow the diffusion of the species. Few juveniles
found among shell grit and undoubtedly belong-
ing to D. gaederopi , however, show one or two
additional protoconch whorls (Fig. 12), confirm-
ing the possibility of some vermetids to produce
more than one type of larval stage depending on
seasons (Scuderi & Cantone, 2007). Moreover the
first protoconch whorl seem to have a different
coiling axis, differing of about 45° from that of
the rest of the protoconch. This character, ob-
served on other species of the same genus, could
have phylogenetic implications. This changing of
axis coiling in the protoconch seem to anticipate
the subsequent changing of 90° of the first tele-
whorl, which allow vermetids to settle on the sub-
stratum.
Distribution. Except for the doubtful locality
reported in the original description (“...probably
from Spain ”), this species is known only from the
material here examined, from S-Italy to Pelagie
Is., between Sicily andN-African coasts of Libya.
Not full grown specimens are easily mistakable
with other congeners or with V. granulatus and this
could reduce the real geographical distribution of
the species along the Mediterranean and the E-
Atlantic.
Remarks. Dendropoma gaederopi was collec-
ted from the very shallow to the deeper fringe of
infralittoral, 4/35 m depth, often on big shells, like
Spondylus or Charonia Gistel, 1847: this habit ap-
pear not similar to the Mediterranean congeneric
species, which are present on rocks in intertidal
and upper infralittoral zones. Further findings of
this species in S-Apulian, cited as Dendropoma
sp. (Terlizzi et al., 2003; Scuderi & Terlizzi,
2012), seem to suggest that it could be present
in a wider geographical area, mainly in South
Mediterranean, but it probably still remain unre-
cognized, due to taxonomical difficulty in its
identification.
Morch well described D. gaederopi , clearly
distinguishing it from the other species of
“ Siphonium ” (= Dendropoma ). The Mediterranean
congeneric D. cristatum (Biondi, 1857) results
quite different on the basis of teleoconch’s shell
sculpture mainly constituted by more dense and
thin axial lamellae and only one spiral chord, the
not smooth protoconch, the colour pattern of ex-
ternal soft parts (Fig. 20) and the feature of the
operculum (Figs. 16, 17) (see Scuderi, 1995 for
further details). The analysis of the type material
of the second Mediterranean species of Dendro-
poma, D. anguliferum Monterosato, 1884, housed
in ZMR (n. 21295), have stated the differences
between this latter and gaederopi, even if the
question of the validity of the Monterosato’s
species remain opened (Scuderi, 2002).
Moreover, D. gaederopi seem not to be a
gregarious vermetid, like some congeneric
species, which could produces wide “trottoir” in
some localities (Hadfield et al., 1972; Safriel,
1975; Barash & Zenziber, 1985; Chemello et al.,
1990; Scuderi et al., 1998), and seem to prefere
deeper waters.
Another Mediterranean vermetid is close sim-
ilar to this species: except for the protoconch and
the external characters of the soft parts, D. gae-
deropi differs from V granulatus “form A” (Scuderi,
1999) by having a shell ambrate in color, with
spiral sculpture constitued by only two (rarely
more) axial ribs, which, in adult shells, produce
spiny excrescences. In V. granulatus the basal
portion of the external side of the tube never forms
a second lip and no lamellae between the keels, nor
scar eroded into the substratum are present.
Moreover, as could be argued by pictures here
presented, the operculum (Figs. 18, 19) is smaller
and thin, the protoconch (Fig. 22) and external soft
parts (Fig. 21) are different.
DISCUSSION
Dendropoma gaederopi is not reported in any
recent checklist of the Mediterranean malacofauna
(Bruschi et al., 1985; Sabelli et al., 1990-92; Bodon
et al., 1995), even if it was cited as valid species by
Monterosato (1892) fide Morch (1861-1862).
The syntype here selected among the type series
from ZMUC carries a black cross on the upper part
of the shell, maybe to marck the semple from which
368
Danilo Scuderi
k , ,
ZOOLOGISK MUSEUM, K0BENHAVN.
1 C% ^p r C ' rjcc ' 1
LoC ' /&&■/ f,. /9
14
Datum: svyb*
cc^/ /3?
13
15
Fig. 1-14. Dendropoma gaederopi. Fig. 1. Syntype (ZMUC), on Spondylus gaederopus. Fig. 2. Shell, Linosa, aperture 0 1mm.
Fig. 3. Shell, Linosa, aperture 0 1.5 mm. Fig. 4. Shell, not full grown specimen, Linosa, aperture 0 0.5 mm. Fig. 5. Shell, Linosa,
aperture 0 3 mm. Figs. 6, 7. Operculum in downward and lateral (a) view, 0 1.5 mm. Fig. 8. Drawing of the animal 0 1.5 mm.
Fig. 9. Protoconch and first tele-whorl, upward view, Linosa, 0.5 mm x 0.6 mm. Fig. 10. Same, detail of the nucleus. Fig. 1 1 . Pro-
toconch and first tele- whorl, side view, Linosa, 0.5 mm x 0.6 mm. Fig. 12. Multispiral protoconch and first tele- whorl, side view,
Linosa, 1.5 mm x 0.85 mm. Fig. 13, 14. Original labels in Morch’s handwriting (ZMUC). Fig. 15. Label of ZMUC. Fig. 16, 17.
D. cristatum. Operculum in downward and lateral view 03.5 mm. Fig. 1 8 , 1 9. Vermetus granulatus. Operculum in downward and
lateral (a) view 0 0.25 mm. Fig. 20. Drawing of the animal 03.5 mm. Fig. 2 1 . Drawing of the animal 01.5 mm. Fig. 22. Protoconch
and first tele-whorl, lateral view, Vendicari, 0.6 mm x 0.5 mm.
Rediscovery of t/ie“Siphonium” gaederopi (Gastropoda Vermetidae) with observations on the early juveniles stages 369
the operculum, preserved in a separate glass-tube,
was obtained (the shell is breacked probably to
draw out the soft parts). All the material was accom-
panied by two series of labels: one original, prob-
ably in Morch’s hand-writing; the second label of
the ZMUC. An additional label reports: “ Following
Bieler (1996) possible type(s) of Siphonium gae-
deropi March, 1861”.
With D. gaederopi the number of known
Mediterranean species of this genus increase to
three, but further and more exaustive studies
should regard D. anguliferum to ascertain if it is
really a good species.
All vermetid species have normally gastropod-
like coiled early developmental stages: some spown
as free swiming larvae; others are crowling juven-
iles at hatching. Hadfield et al. (1972) stated that
nurse yolk assumption by the embryos influences
the larval mode of life and dimensions, but only in
Vermetus rugulosus both type of larvae could be sim-
ultaneously produced (Scuderi & Cantone, 2007).
The finding of more than one type of juveniles
suggest that this species could take advantage from
the planktonic lifestyle to settle on islands rocky
environments and from the direct development to
ensure specimen’s enlargement to the established
population.
ACKNOWLEDGEMENTS
I am grateful to Tom Schioette (ZMUC) for the
kind loan of Morch’s type material; my friend
Marco Faimali (ISMAR-CNR, Genoa, Italy) fur-
nished logistic accommodation in Linosa Island;
Agatino Reitano and Alfio Germana loaned their
material and foumished accommodation in Mar-
zamemi, Sicily. I am grateful to Antonio Terlizzi
(DISTEBA, Lecce, Italy), who allowed the study of
the Apulian materials. Finally, I am indebted to Jose
Templado (Museo Nacional de Ciencias Naturales,
Madrid) and Alberto Villari, for their hepful sug-
gestions and the critical review of the text.
REFERENCES
Barash A. & ZenziberZ., 1985. Structural and biological
adaptations of Vermetidae (Gastropoda). Bollettino
Malacologico, 21: 145-176.
Bieler R. 1996. Morch's worm-snail taxa (Caenogastro-
poda: Vermetidae, Siliquariidae, Turitellidae).
American Malacological Bullettin, 13: 23-35.
Bodon M., Favilli L., Giannuzzi-Savelli R., Giovine F.,
Giusti F., Manganelli G., Melone G., Oliverio M.,
Sabelli B. & Spada G., 1995. Gastropoda Proso-
branchia, Heterobranchia Heterostropha. In: Minelli
A., Ruffo S. & La Posta S. (eds.), Checklist delle
specie della fauna italiana, 14. Calderini, Bologna,
60 pp.
Bruschi A., Cepodomo I., Galli C. & Piani P., 1985.
Catalogo dei molluschi conchiferi viventi nel
Mediterraneo. ENEA, Collana di studi ambientali:
XII + 111 pp.
Chemello R., Pandolfo A. & Riggio S., 1990. Le
biocostruzioni a Molluschi Vermetidi nella Sicilia
Nord-Occidentale. Atti 53° Congresso UZI, Palermo:
88 .
Hadfield M.G., Kay E.A., Gillette M.U. & Lloyd M.C.,
1972. The Vermetidae (Mollusca Gastropoda) of the
Hawaiian Islands. Marine Biology, 12: 81-98.
Keen A.M., 1961. A proposed reclassification of the
Gasteropod family Vermetidae. Bulletin of the British
Museum (Natural History). Zoology, 7: 181-213.
Morch O.A.L., 1 861-62. Review of the Vermetidae (Part
I-II-III), Proceedings of the Zoological Society of
London (1860): 145-181; (1861): 326-365; (1862):
54-83.
Monterosato A.T., 1892. Monografia dei Vermeti del
Mediterraneo. Bullettino della Societa di malacologia
italiana, 17: 7-48.
Sabelli B., Giannuzzi-Savelli R. & Bedulli D., 1990-92.
Catalogo annotato dei Molluschi marini del Mediter-
raneo, vol. I— III, Libreria Naturalistica Bolognese,
Bologna, 781 pp.
Safriel U., 1974. Vermetid gastropods and Interdital
Reefs in Israel and Bermuda. Science, 186: 1 1 13—
1115.
Scuderi D., 1995. II genere Dendropoma (Gastropoda:
Vermetidae) nel Mediterraneo. Bollettino Malacolo-
gico, 31: 1-6.
Scuderi D., 1999. Contributo alia conoscenza dei Ver-
metidae mediterranei: Vermetus granulatus (Graven-
horst, 1831) e suoi principali morfotipi. Bollettino
Malacologico, 35: 45-48.
Scuderi D., 2002. La famiglia Vermetidae Rafinesque,
1815 (Mollusca: Gastropoda): i tipi della collezione
Monterosato serviti per la compilazione della
“Monografia dei vermeti del Mediterraneo”. 63°
Congresso Unione Zoologica Italiana. Rende (CS).
Scuderi D., Terlizzi A. & Faimali M., 1998. Osservazioni
su alcuni tratti della biologia riproduttiva di vermeti
biocostruttori e loro ruolo nella edificazione dei
“trottoir”. Biologia Marina Mediterranea, Atti XXVIII
Congr. SIBM, 5: 284-289.
370
Danilo Scuderi
Scuderi D. & Cantone G., 2007. Simultaneous ambival-
ent reproductive strategy in the sessile gastropod
Vermetus rugulosus Monterosato, 1878 (Gastropoda:
Prosobranchia). 16th World Congress of Malacology,
20 July 2007- Antwerp, Belgium, 15.
Scuderi D. & Terlizzi A., 2012. I molluschi dell’ Alto
Jonio. Guida teorico-pratica di malacologia mediter-
ranea. Ed. Grifo, Lecce. 108 pp., 41 Tavole.
Terlizzi A., Scuderi D., Fraschetti S., Guidetti P. & Boero
F., 2003. Molluscs on subtidal cliffs: patterns of spa-
tial distribution. Journal of The Marine Biological
Association of The United Kingdom, 83: 165-172.
Biodiversity Journal, 2015, 6 (1): 371-376
Monograph
First assessment of the vermetid reefs along the coasts of
Favignana Island (Southern Tyrrhenian Sea)
Paolo Balistreri 1 *, Renato Chemello 2 & Anna Maria Mannino 1
1 D ip artim e n to di Scienze e Tecnologie Biologiche Chimiche e Farm aceutiche, Universita di Palermo, Via Archirafi 38. 90 1 23
Palermo, Italy; e-mail: requin.blanc@hotmail.it; anna m aria, m annino@unipa.it
2 D ip artim e n to di Scienze della Terra e del Mare, Universita di Palermo, Via Archirafi 28, 90 1 23 Palermo, Italy; e-mail: renato.
chemello@ unipa.it
Corresponding author
ABSTRACT Intertidal vermetid reefs, particularly vulnerable to environmental changes and human
activities, are now experiencing high mortality in several areas of the M editerranean Sea. Since
the increase of knowledge on this habitat is important for conservation purposes, we provide
a first baseline assessment of the vermetid reefs along the coasts of the Favignana Island
(Marine Protected Area “Egadi Islands”). Preliminary results showed the presence of a true
reef, similar to a fringing reef, displaying at least three local patterns, distinguishable for width
(from 2.3 to 15.5 m), height of the outer and of the inner marg in (from 5.6 to 18 cm and from
8.3 to 26 cm, respectively) and number, width and depth of cuvettes. Moreover, significant
differences in topographic complexity among the areas were evidenced whereas no correlation
between coastal exposure and topographic complexity was found.
KEY WORDS Bioconstruction; Favignana Island; habitat and topographic complexity; vermetid reef.
Received 25.08.2014; accepted 30.01.2015; printed 30.03.2 0 15
Proceedings of the 2nd Interna tional Congress “Speciation and Taxonomy”, May 1 6 th - 1 8 th 2014, Cefalu - Caste lb uono (Italy)
INTRODUCTION
Vermetid reefs are b io c o n s tru c tio n s built up by
the gastropod mollusc Dendropoma petmeum
(Monterosato, 1884) in association with some coral-
line algae such as Neogoniolithon brass ica-florida
(Harvey) Setchell & M ason. These bioconstructions
are unique and highly diverse systems that play a
fundamental structural role, as they protect coasts
from erosion, regulate sediment transport and
accum ulation, serve as carbon sinks, make the habitat
more complex and heterogeneous and provide
numerous habitats for animal and vegetal species
thus increasing intertidal biodiversity (Pandolfo et
al., 1 992; Pandolfo et al., 1996; Badalamenti et al.,
1998). In the M editerranean Sea their distribution is
restricted to the warmest part of the basin with the
largest formations generally found off the coasts of
Israel and Lebanon, but they have also been reported
in Turkey, Crete, continental Spain and Baleari
Islands, A lgeria, Morocco, along Maltese and Italian
shores (Peres & Picard, 1 952; Molinier & Picard,
1 953; Molinier, 1 955; Safriel, 1975; Boudouresque
& Cinelli, 1976; Dalongeville, 1977; Kelletat, 1979;
Richards, 1 983; Laborel, 1 987; Azzopardi, 1992;
Garcia-Raso et al., 1992; Templado et al., 1992;
Bitar & Bitar-Kouli, 1 995a, 1 955b; Azzopardi &
Schembri, 1997).
In Sicily, large and mo re or less continuous ver-
metid reefs are present along the north/northwestern
372
Paolo Balistreri et alii
coasts between Zafferano Cape and Trapani and
within the Marine Protected Area (MPA) “Egadi
Islands” (Chemello, 1 989; Chemello et al., 1990a,
1990b; Badalamentietal., 1992a, 1992b; Chemello
et al., 2000; Dieli et al., 200 1; Chemello, 2009).
Isolated reefs are found at M ilazzo Cape and only
small reefs are found around Taormina and Syra-
cuse, on the eastern coast of Sicily, and on the
Islands of Lampedusa and U stica, th at represent the
limit of distribution respectively on the south and
on the north of the Sicilian coasts (Chemello et al.,
1990a; Chemello et al., 2000; Dieli et al., 2001;
Consoli et al., 2008; Chemello, 2009). These
biogenic constructions, enclosed in the SPA/BIO
Protocol (B arcelona C onvention) are now threatened
by environmental changes and human activities
(e.g. pollution, climate change, ocean acidification)
thus experiencing high mortality in several areas of
the Mediterranean Sea (D i Franco et al., 2011;
G alii, 2013; M ilazzo et al., 2014).
Due to the high vulnerability of these habitats,
action plans for their conservation should be a
priority. We know that the increase of know ledge is
essential for the conservation and protection of this
highly valuable habitat. Since only a low percentage
of Sicilian vermetid reefs are subjected to conser-
vation and many of them are not yet investigated
(Chemello, 2009; Chemello & Silenzi, 2011), with
this study we provide a first baseline assessment
of the vermetid reefs present along the coasts of
Favignana Island (MPA “Egadi Islands”).
The aim s o f the p re sen t stu dy were: i) to p ro vid e a
first description of the reef typology and ii) to test the
effect of the coastal exposure on the topographic
complexity of the reefs.
MATERIAL AND METHODS
Study area
The study was carried out at Favignana Island
(MPA “Egadi Islands”), located approximately five
kilometersfrom the west coast of Sicily. The Island,
part of the Aegadian Archipelago, represent an
example of a lower Pleistocene bioclastic cal-
car enite, characterized by a typic association known
as foramol (K il, 2010). The west side is character-
ized by the presence of the calcareous Monte Santa
Caterina (300 metres high), flanked by areas with
lower relief. The mechanical and chemical erosion
Figure 1. Location of the study areas: Favignana Island.
produced detritic deposits that partially mask the
abrasion platform. The eastern side of the mountain
shows a high cliff in the northern part, that
gradually dips to the south. The coastline is highly
rugged and not very high. The south dipping surface
might be the result of either depositional or
erosional processes. From a geological point of
view, two main tectonic units can be recognized:
the Monte Santa Caterina and the Punta Faraglione.
Vermetid reef analysis
A preliminary survey allowed us to locate six
study areas characterized by the presence of a
vermetid reef: Faraglione, Pozzo, A rre Turinu,
Grotta Perciata, Cala Rotonda and Stornello (Fig. 1).
The areas were chosen in such a way to also test
the effects of the coastal exposure on the vermetid
reef topographic complexity. Three along the
northern side: Faraglione, Pozzo and Arre Turinu,
and three along the southern side: Grotta Perciata,
Cala Rotonda and Stornello. In each area the reef
topographic complexity was measured using a lm
x lm quadrat (three random replicates). The 4 sides
and the 2 diagonals of the quadrat were measured
using a meter with a resolution of 0.01m. Topo-
graphic complexity was calculated as the ratio
between the registered measures (real measure, Xi)
and the known measures of the used quadrat (Xn):
X i/X n (Graziano et al., 2009 ).
The more the ratio is far from 1, the more com-
plex is the substrate. To describe the reef typology
the following variables were considered: the reef
width from the inshore towards the open sea (meas-
ured using a meter with a resolution of 0.01m), the
height of the inner and the outer marg in and the slopes
of the margins (measured using a goniometer).
First assessment of the vermetid reefs along the coasts ofFavignana Island (Southern Tyrrhenian Sea)
373
Data analysis
Differences in reef topographic complexity
were analysed using per mutational multivariate
analysis of variance (PERMANOVA, Anderson,
200 1). For the topographic complexity, the design
consisted of three factors: Coastal exposure (Sd;
two levels, fixed factor), Area (Ar; three levels,
random, nested in Sd) and Site (St; three levels,
random, nested in ArxSd).All multivariate analyses
were based on Bray-Curtis dissimilarities of log(x
+ 1) transformed data and each term in the analyses
was tested using 9999 random permutations of the
appropriate units. The analyses were performed
using the software package PRIMER 6 (Clarke &
G orley, 2006).
RESULTS
Reef typology
All the vermetid reefs are consistent with a true
reef (according to Antonioli et al., 1999), displaying
at least three local patterns, distinguishable for
width, height of the outer and of the inner margin
and number, width and depth of cuvettes. A descrip-
tion of the different patterns are reported below.
Pattern 1: Pozzo and Faraglione (northern side,
Fig s. 2-5 ).
Outer margin: wide, flattened and irregular. In
the inner side, crevices were also present. Some-
times at Faraglione are present two outer margins.
Inner margin: Dendropoma petraeum is absent.
Cuvettes: not many, not deep and with a variable
width. At Faraglione they are mainly present near
the outer margin. At Pozzo some of them are
fullfilled of sediment.
Study
Width
Height of
the inner
Height of
the outer
Slope of
the inner
Slope of
the outer
Areas
(m)
margin
(cm)
margin
(cm)
margin
(°)
margin
(°)
Fara-
7.03 ±
8.66 ±
1 8 ±
38.3 ±
45 ±
glione
0.23
0.23
0.23
0.23
0.2 3
Pozzo
15.46 ±
1 7.33 ±
1 5.33 ±
45 ±
45 ±
0.23
0.23
0.23
0.23
0.23
Pattern 2: Grotta Perciata and Stornello (south-
ern sid e, Figs. 6 -9 ) .
Outer margin: thin and not continuously ar-
ranged. inner margin: Dendropoma petraeum is
absent. Cuvettes: not many and not deep.
Study
Areas
Width
(m)
Height of
the inner
margin
(cm)
Height of
the outer
margin
(cm)
Slope of
the inner
margin
(°)
Slope of
the outer
margin
(°)
Grotta
7.3 1 ±
1 0.33 ±
1 5.66 ±
27.5 ±
42.5 ±
Perciata
0.23
0.23
0.23
0.23
0.23
Stor-
5.10 ±
8.33 ±
8 ±
33.3 ±
29.16 ±
nello
0.23
0.23
0.23
0.23
0.23
Pattern 3: A rre Turinu (northern side) and C ala
Rotonda (southern side, Figs. 10-13).
The reef is damaged. Outer margin: it has a
variable height and sometimes it is absent. Some
crevices can also be present togetherwith regrow th
areas. Inner margin: Dendropoma petraeum is
absent. Cuvettes: many and sometimes very deep.
Study
Areas
Width
(m)
Height of
the inner
margin
(cm)
Height of
the outer
margin
(cm)
Slope of
the inner
margin
(°)
Slope of
the outer
margin
(°)
Arre
6.38 ±
26 ±
1 7 ±
26.6 ±
45 ±
Turinu
0.23
0.23
0.23
0.23
0.23
CalaRo-
2.30 ±
12 ±
7.6 ±
4 1 .6 ±
38.3 ±
tonda
0.23
0.23
0.23
0.23
0.2 3
Reef topographic complexity
The PERMANOVA on the reef topographic
complexity provided an evidence of significant
differences in topographic complexity among the
areas whereas no differences were recorded between
the two coastal exposures (Fig. 14; Table 1).
Source
df
ss
MS
Pseudo-F
P(MC)
Sd
i
8 627
8627
0.349
0.77 1
Ar(Sd)
4
9 8876
247 1 9
9.1486
0.000 1
St [Ar
(Sd)]
1 2
32423
270 1 .9
1 .8084
0.0098
Res
36
5 3 789
1494.1
Total
53
1 .9 3 7 2E 5
Table 1. PERMANOVA on the topographic
com p lex ity data.
374
Paolo Balistreri et alii
Figures 2-5.
Scheme of the
pattern 1 (Fig .
2 ) , F arag lio n e
(Fig. 3), Pozzo
(Fig. 4), Pozzo:
outer margin
(Fig. 5).
Figures 6-9.
Scheme of the
p attern 2 (Fig.
6), Grotta Per-
ciata (Fig . 7 ),
S to rn ello (Fig.
8), Grotta Per-
ciata: outer m ar-
gin (Fig. 9).
Figures 10-13.
Scheme of the
p a ttern 3 (Fig.
1 0), A rre T urinu
(Fig. 11), C ala
Rotonda (Fig.
1 2 ), A rre T urinu :
outer margin
(Fig. 13).
First assessment of the vermetid reefs along the coasts of Favignana Island (Southern Tyrrhenian Sea)
37 5
Hprtlwm £rff $airth#f n Side
NT HP NRT HT Ht
Figure 14. Topographic complexity in the studied areas.
DISCUSSION AND CONCLUSION
The vermetid reefs along the coasts of Fav-
ignana are consistent with a true reef described
along the north-western Sicilian coasts (Antonioli
et al., 1999). The reef distribution around Favignana
confirms the need of carbonatic substrates and of
an abrasion platform for the form ation of true reefs
(Dieli et al., 2001 ). All the reefs are characterized,
along a transect from the inshore towards the open
sea, by the tipical patches recognized for other
Sicilian reefs (Chemello et al., 2000; Dieli et al.,
200 1). The reefs width were in agreement with the
values reported for other Sicilian reefs whereas the
height values were lower (Dieli et al., 2001). Some
differences were highlighted locally in the con-
sidered variables, in particular in the m argins, in the
depth and in the number of cuvettes. Data on topo-
graphic complexity showed significant differences
among the areas (small scale) but no relationship
between the coastal exposure (large scale) and the
reef topographic complexity was evidenced.
Vermetid reefs play an important role as modu-
lators of morphological coastal processes and as
ecological “engineers”, making the habitat more
complex and tridimensional and promoting marine
biodiversity (Pandolfo et al., 1996; Chemello et al.,
2000; Bressan et al., 2009). Therefore, much more
attention should be paid to the study of the reef
morphology and distribution together with the
associated communities and the trophic processes
within associated species.
Moreover, since the easy accessibility of ver-
metid reef makes it highly vulnerable to coastal
human activity (Franzitta et al., 2006; Graziano et
al., 2007), a correct planning in the areas where
reefs are present, in order to minimize all potential
environmental threats, should be a priority.
REFERENCES
Anderson M .J., 2001. Permutation tests for univariate or
multivariate analysis of variance and regression.
Canadian Journal of Fisheries and Aquatic Sciences,
58: 626-639.
Antonioli F., Chemello R.,Improta S.& Riggio S., 1999.
Dendropoma lower intertidal reef formations and
their p a la e o c lim a to lo g ic a 1 significance, NW Sicily.
Marine Geology, 161: 155-170.
Azzopardi L., 1 992. Aspect of the ecology of Vermetid
Gastropods on Maltese rocky shores. PhD Thesis,
University of Malta, 163 pp.
Azzopardi L. & Schembri P.J., 1997. Vermetid crusts
from the Maltese Islands (Central Mediterranean).
Marine Life, 7: 7-16.
Badalamenti F., Chemello R., Gristina M . Riggio S. &
Toccaceli M ., 1 992a. C ara tte riz z az io n e delle piatta-
forme a M olluschi Vermetidi nella costa della Riserva
Naturale dello Zingaro (TP). Oebalia, Suppl. 17:
543-545 .
Badalamenti F., Chemello R., Gristina M . Riggio S. &
Toccaceli M ., 1 992b. C ara tteriz z a z io n e delle piatta-
forme a Molluschi Vermetidi nella costa tra Capo
Gallo e Isola delle Femmine (PA): area proposta
come riserva naturale marina. Oebalia, Suppl. 17:
547-549.
Badalamenti F., Chemello R., D’Anna G. & Riggio S.,
1 998. Diversity of the Polychaete assemblage in the
hard bottom mediolittoral along the north-western
Sicilian coast: the role played by the vermetid bioco-
struction. Atti 1° Convegno Nazionale delle Scienze
del Mare, “ Diversita e Cambiamento”, Ischia: p. 14.
Bitar G. & Bitar- Kouli S., 1995a. Apergu de bionomie
bentique et repartition des differents facies de la
roche littorale a Hannouch (Liban, Mediterranee
orientale). Rapport Commission International Mer
Mediterranee, 34: 19.
Bitar G. & Bitar-Kouli S., 1995b. Impact de la pollution
sur la repartition des peuplem ents de substrat dur a
Beyrouth (Liban, Mediterranee orientale). Rapport
Commission International M er M editerranee, 34: 19.
Boudouresque C.F. & Cinelli F., 1976. Le peuplement
algal des biotopes sciaphiles superficiels de mode
battu en Mediterranee occidentale. Pubblicazioni
della Stazione Zoologica di Napoli, 40: 433-459.
Bressan G., Chemello R, Gravina M.F, Gambi M.C,
Peirano A., Cocito S., Rosso A. & Tursi A., 2009.
A ltre principali b io c o stru z io n i. In: Relini G. (Ed.)
2009. Quaderni Habitat - Biocostruzioni marine:
376
Paolo Balistreri et alii
elementi di architettura naturale. Societa Italiana di
Biologia Marina, Genova, 22: 89-114.
Chemello R., 1 989. La bionomia bentonica ed i Mollu-
schi. 5. II piano In fralito rale : il marciapiede a ver-
meti. Notiziario SIM, 7: 167-170.
Chemello R., 2009. Le biocostruzioni marine in Medi-
terraneo.Lo stato delle conoscenze suireefaVermeti.
Biologia Marina Mediterranea, 16: 2-18.
Chemello R., Dieli T. & Antonioli F., 2000. II ruolo dei
“reef” a Molluschi vermetidi nella valutazione della
b io d i v e rs ita . “Mare e cambiamenti globali”. Qua-
dernilC RAM, Roma: 105-118.
Chemello R., Gristina M ., Toccaceli M ., Badalamenti F.
& Riggio S., 1990a. D is trib u z io n e delle formazioni
a Molluschi Vermetidi lungo le coste siciliane. A tti
538 Congresso UZI, Palermo, p. 60.
Chemello R., Pandolfo A. & Riggio S., 1990b. Le bio-
costruzioni a Molluschi Vermetidi nella sicilia Nord-
Occidentale.Atti 538 Congresso UZI, Palermo, p. 88.
Clarke K.R. & Gorley R.N., 2006. PRIMER v6: user m a-
nual/tutorial. Plymouth, MA: PRIMER-E.
Consoli P., Romeo T., Giongrandi U. & Andaloro F.,
2008. Differences among fish assemblanges associa-
ted with a neashore vermetid reef and two other rocky
habitats along the shores of Cape Milazzo (northern
Sicily, central M editerranean Sea). Journal of the M a-
rine Biological Association of the UK, 88: 401-410.
Dalongeville R., 1977. Formes littorales de corrosion
dans le roches carbonatees du Liban: etude morpho-
logique. M e d ite rra n e e , 3: 21-33.
Dieli T., Chemello R. & Riggio S ., 2001. Eterogeneita
strutturale delle formazioni a vermeti (Mollusca:
Caenogastropoda) in Sicilia. Biologia Marina Medi-
terranea, 8: 223-228.
Di Franco, A., Graziano M .,Franzitta G ., F ellin e S ., Che-
mello R. & Milazzo M., 2011. Do small marinas
drive habitat specific impacts? A case study from M e-
diterranean Sea. Marine Pollution Bulletin, 62: 926-
93 3.
Franzitta G., Graziano M ., Di Franco A., Milazzo M . &
Chemello R., 2006. Valutazione dell’impatto di u n
piccolo porto sui popolam enti bentonici di substrato
duro. Biologia Marina Mediterranea, 13: 70-71.
G alii B .S ., 2013. Going going gone: the loss of a reef
building gastropod (Mollusca: Caenogastropoda:
Verm etidae) in the southeast Mediterranean Sea.
Zoology in the Middle East, 59: 179-182.
G arcia-R aso J.E., Luque A.L., Templado J., Salas C.,
H ergueta E., M oreno D.& Calvo M., 1992. Faun ay
flora marinas del Parque Natural de Cabo de Gata-
N ijar. M ad rid , 2 8 8 pp .
Graziano M., D i Franco A., Franzitta G., Milazzo M. &
Chemello R., 2007. Effetti di differenti tipologie di
impatto antropico sui reef a vermeti. B iologia M arina
Mediterranea, 14: 306-307.
Graziano M, Milazzo M. & Chemello R., 2009. Effetti
della protezione e della complessita topografica sui
popolamenti bentonici dei reef a vermeti. Biologia
Marina Mediterranea, 16: 40-41.
KelletatD., 1979. Geomorphologishe Studien an den Kti-
sten Kretas. Abhandlungen der Akademie der
W issenschaften in Gottingen. M a th e m a tis c h -P h y s i-
kalische Klasse, 3° Folge, 32.
K il R.A., 2010. S edim entology and 3D architecture of a
bioclastic calcarenite complex on Favignana, sou-
thern Italy: Implications for reservoir modelling. M Sc
Thesis, University of Delft, The Netherlands.
Laborel J., 1987. Marine biogenic constructions in the
M editerranean. A review. Scientific Reports of Port-
C ro s National Park, France, 1 3: 97- 1 26.
M ilazzo M ., Rodolfo-M etalpa R., Bin San Chan V., Fine
M ., A le ssi C., Thiyagarajan V., Hall-Spencer J.M . &
Chemello R ., 2014. Ocean acidification impairs ver-
metid reef recruitment. Scientific Reports, 4 DOI:
1 0.1 038/srep04 1 89
Molinier R., 1955. Les platformes et cor niches recifales
de Vermets (VeYTIWtUS CYistdtUS Biondi) en Mediter-
ranee occidentale. Comptes Rendus de l'Academie
des sciences Paris, 240: 36 1-363.
Molinier R. & Picard J., 1953. Notes biologiques a pro-
pos d’un voyage d ’ etude sur les cotes de Sicile. A n -
nales de l’Institut O ceanographique, 28: 1 63- 1 88.
Pandolfo A., Chemello R. & Riggio S ., 1 992. Prime note
sui popolamenti associati ai“trottoir” a vermetidi
delle coste siciliane: i Molluschi. Oebalia, 1 7: 379-
3 82.
Pandolfo A., Chemello R., Ciuna I., Lo Valvo M . & Rig-
gio S., 1996.Analisi della distribuzione dei molluschi
nella zona di transizione tra mesolitorale ed infrali-
toral e superiore lungo le coste della Sicilia. Biologia
Marina Mediterranea, 3: 78-87.
Peres J.M. & Picard J., 1952. Les cor niches calcaires
d’origine biologique en Mediterranee occidentale.
Recueil des Travaux de la Station Marine d’En-
doume, 4: 2-34.
Richards G.W., 1983.Molluscan zonation on rocky sho-
res in Malta. Journal of Conchology, 3 1: 207-224.
Safriel U., 1975. The Role of Vermetid Gastropods in
the Formation of M editerranean and Atlantic Reefs.
Oecologia, 20: 85-101.
Templado J., Templado D. & Calvo M ., 1 992. The for-
mations of vermetid gastropod DefldwpOtnCl petVCl-
eum (Monterosato, 1 884) on the coasts of the Iberian
Peninsula (Western Mediterranean). In: Giusti F. &
Manganelli G. (Eds.) 1 992. Abstracts 11th Interna-
tional Malacological Congress, Siena, 514-515.
Biodiversity Journal, 2015, 6 (1): 377-392
A preliminary checklist of the species of non-marine Molluscs
from theAlbumi Mountains, Campania, Southern Italy (Mol-
lusca Gastropoda Bivalvia)
Agnese Petraccioli 1 , Paolo Crovato 2 , Ivano Niero 3 , Laura De Riso 4 , Camillo Pignataro 5 , Gaetano Odierna 1
& Nicola Maio 1 *
'Dipartimento di Biologia, Complesso Universitario di Monte S. Angelo, Universita degli Studi di Napoli Federico II, Edificio 7,
via Cinthia 21, 80126 Napoli, Italy
2 Via S. Liborio 1, 80134 Napoli, Italy
3 Via Cici 17/1, 30038 Spinea (Venezia), Italy
4 Ente Parco Nazionale del Cilento, Vallo di Diano e Alburni, Via Montesani, 84078 Vallo Della Lucania (Salerno), Italy
5 Fondazione I.RI.DI.A. - Museo Naturalistico, via Forese 16, 84020 Corleto Monforte (Salerno), Italy
‘Corresponding author, e-mail: nicomaio@unina.it
ABSTRACT An annotated checklist of the species of non-marine molluscs from the Alburni Mountains
(Salerno Province, Campania, Southern Italy) is reported. The research was carried out
from 2010 to 2013 inside a Site of Community Importance (SCI) and a Special Protection
Area (SPA), of the Cilento, Vallo di Diano and Alburni National Park. The non-marine
molluscs sampled on the field were compared with data available from the literature and
malacological collections. Up to now, only 12 non-marine Mollusc species were known
from the Alburni Mountains through bibliographical data. In all, the malacofauna of Alburni
Mountains is composed by 83 non-marine Mollusc species (73 species of land snails, and
10 species of freshwater molluscs). The presence of nine species (six species of land snails
and three species of freshwater snails) was confirmed by our field investigation, four species
(3 species of land snails and 1 species of allochthonous freshwater snails) were recorded
only by bibliographical data and were not yet found. Our analysis identifies 70 species of
non-marine Molluscs (64 species of land snails, 6 species of freshwater molluscs) recorded
on the basis of field data which were not previously recorded from the study area. At least
1 1 species are new records for the Campania Region. Extremely interesting is the record
of Vertigo angustior Jeffreys, 1830 a species protected in European Union by the Annex
II of the “Habitats Directive” and listed as “Vulnerable” at the European level. A Red List
of Threatened Species is proposed and the species were classified with the code of I.U.C.N.
(Version 2014.3). Five allochthonous species were surveyed for the first time in the study
area: 3 land snails: Lucilla scintilla (Lowe, 1852), Lucilla singleyana (Pilsbry, 1829) and
Paralaoma servilis (Shuttleworth, 1852), and 2 freshwater snails: Potamopyrgus anti-
podarum (J.E. Gray, 1843) and Ferrissia fragilis (Tryon, 1863). Four species are known
exclusively from the literature: Vertigo ( Vertigo ) moulinsiana (Dupuy, 1849), Macrogastra
(Pyrostoma) plicatula (Draparnaud, 1801), Cernuella virgata (Da Costa, 1778), and Haitia
acuta (Draparnaud, 1805).
KEY WORDS Non -marine Molluscs; Alburni Mountains, Campania; faunistics; conservation.
Received 18.07.2014; accepted 29.11.2014; printed 30.03.2015
Proceedings of the 2nd International Congress “Speciation and Taxonomy”, May 1 6th- 1 8th 2014, Cefalu-Castelbuono (Italy)
378
Agnese Petraccioli etalii
INTRODUCTION
The knowledge of the malacofauna of Campania
(about 150 species, personal data; 13,595 sq Km)
is far below than that of other regions. This is even
more evident when we consider the protect areas of
this territory as those of the Alburni Mountains
whose malacofauna is virtually unknown (Fig. 1).
In all Campania Region, only papers on check-
list of Capri Island and Vesuvius National Park are
known in the last 10 years (Petraccioli et al., 2005a,
2005b, 2006a, 2006b, 2007; Picariello et al., 2011);
moreover there are only historical reports, nearly
100 years old, concerning the most common species
and quoted with obsolete names. The purpose of
this paper is, therefore, to help bridge this gap by
increasing the malacofauna knowledge of this
group in an important area of the Campania Region
as the Alburni Mountains, with particular reference
to the species listed in the Annex of the Habitats
Directive, in the Index of the protected fauna of
Italy and in the various red lists (Manganelli et al.,
2000a; Cerfolli et al., 2002; I.U.C.N., 2014).
MATERIAL AND METHODS
The Alburni Mountains are a calcareous massif
in the Salerno Province (Campania Region) belong-
ing to the Lucan Sub Apennines chain, located in the
Eastern area of Cilento, near the borders between
Campania and Basilicata. In North-East the range de-
grades into the plain of Vallo di Diano between the
valleys of the Calore Lucano, Tanagro and Sele
rivers. The massif extends for about 250 km 2 . The
study area (SCI IT8050033 named: “Monti Alburni”,
and SPA IT8050055 named: “Alburni”) is included
in Cilento, Vallo di Diano and Alburni National Park
and covers 14 administrative municipalities (Aquara,
Auletta, Castelcivita, Controne, Corleto Monforte,
Ottati, Petina, Polla, Postiglione, San Pietro al Tana-
gro, San Rufo, Sant' Angelo a Fasanella, Sanf Arsenio,
and Sicignano degli Alburni) (see TEMI, 2010).
A detailed investigation on the historical and cur-
rent literature and a comprehensive study of Neapol-
itan public and private molluscan collections were
preliminarly performed. We also inspected the
original sites reached by Costa (1874) 140 years ago.
From 2010 to 2013 the field surveys were con-
ducted in 127 sampling points (stations or plots)
between 100 and 1742 meters above sea level (sum-
mit of Monte Albumo/Panormo) in all suitable hab-
itats present on the territory of the Alburni Moun-
tains in accordance with the vegetation types
reported in the land use map (1 : 25.000) available by
the “Ente Parco”. Adult specimens and shells of non-
marine molluscs were hand-collected through visual
search, leaf litter and soil collecting and sorting.
Samples were then air dried and sieved down to 0.5
mm mesh. Samples of sediment were screened with
calibrated sieves. The cleaned up material was
examined under lens and/or stereo microscope to
sort the smallest fraction, namely. Fractions above 1
cm were searched by a Leica EZ4 stereo microscope
(Leica Microsystems GmbH, Wetzlar, Germany),
both incident and transmitted, and then photo-
graphed with a digital camera. The specimens for ana-
tomical exams were drowned in water and fixed in
75% ethanol. The reproductive apparatus was extrac-
ted by means of scalpel, scissors and forceps. The
illustrations of genitalia were sketched using a camera
lucida mounted on the above stereomicroscope.
The sampled specimens were collected with
permission of the “Ente Parco Nazionale del Ci-
lento e Vallo di Diano” (Permit no. 16341/
19.10.2010). Two sampled specimens for species
were deposited in the Museo Naturalistico of
Corleto Monforte (Salerno Province), a museum
acknowledged as an “institution of regional in-
terest” (Decreto dalla Giunta Regionale Campania
n. 2010 del 29/12/2008). In addition, when other
specimen/species were collected, we preserved
them in the private collection of the authors. The
species identification was based on qualified dicho-
tomickeys (Giusti & Pezzoli, 1980; Girod et al.,
1980; Bech, 1990; Giusti et al., 1995; Kerney &
Cameron, 1999). The taxonomic order and nomen-
clatural arrangement of the list follow: Bodon et al.
(1995), Manganelli et al. (1995, 1998, 2000b),
Castagnolo (1995), Ponder & Lindberg (1996),
Nordsieck (2002) and Bank (2011); the common
names were based on Janus (1982) and on the web
site: http://media.eol.org.
For each species a brief note on the abundance
in the study area is reported according to the fol-
lowing classification: Very rare (sampled in 1-5
stations), Rare (sampled in 6-10 stations), Uncom-
mon (sampled in 11-19 stations), Common (
sampled in 20-35 stations), Widespread (sampled
in over 35 stations). The bibliographical and
museological data were then reported. If present in
the I.U.C.N. Red List, each species is classified
A preliminary checklist of non-marine Molluscs from the Alburni Mountains, Campania, Southern Italy
379
with the code of Red List of Threatened Species
(I.U.C.N., 2014) and Cuttelod et al. (2011).
data from 1986. Classified as “EC” by Cuttelod et
al. (2011) and by I.U.C.N. (2014).
RESULTS
Species surveyed on the field
Phylum MOLLUSCA Cuvier, 1795
Classis GASTROPODA Cuvier, 1795
Subclassis ORTHOGASTROPODA Ponder et
Lindberg, 1996
Ordo ARCHITAENIOGLOSSA Haller, 1890
Familia COCHFOSTOMATIDAE Kobelt, 1902
Cochlostoma montanum (Issel, 1866)
Cochlostoma montanum cassiniacum (Saint-Simon,
1878)
Common, 34 plots, locally abundant. Museal
Familia ACICULIDAE J.E. Gray, 1850
Platyla talentii Bodon et Cianfanelli, 2008
Rare, 10 plots, locally abundant (Figs. 2, 3). En-
demic of Southern Apennine. Bodon & Cianfanelli,
2008. Classified as “NT” by Cuttelod et al. (2011)
and “NT” by I.U.C.N. (2014).
Ordo NEOTAENIOGFOSSA Haller, 1892
Familia POMATIIDAE Newton, 1891 (1828)
Pomatias elegans (O.F. Muller, 1774)
Round-mounted Snail
Common, 28 plots. Museal data from 1986.
Figure 1. The study area: Albumi Mountains, S-Italy.
380
Agnese Petraccioli etalii
Familia HYDROBIIDAE Stimpson, 1865
Mud snails
Pseudamnicola ( P. ) cfr. moussonii (Calcara, 1841)
Very rare, 1 plot. Classified as “LC” by Cuttelod
et al. (2011) and by I.U.C.N. (2014). Endemic
species in Europe (Cuttelod et al., 2011).
Belgrandia minus cula (Paulucci, 1881)
Very rare, 2 plots. Bodon et al. (2005). Classi-
fied as “DD” and Endemic species in Europe by
Cuttelod et al. (2011) and “DD” by I.U.C.N. (2014).
Potamopyrgus antipodarum (J.E. Gray, 1843)
New Zealand mud snail, Jenkins' Spire Snail
Very rare, 2 plots (Fig. 4). Allochthonous species,
introduced from New Zealand (Lori et al., 2005;
Lori & Cianfanelli, 2007; Cianfanelli, 2009; Cutte-
lod et al., 2011). Classified as “LC” by I.U.C.N.
(2014).
Bythinella opaca (M. von Gallenstein, 1848)
Bythinella schmidtii (Kiister, 1852)
Very rare, 4 plots. Bodon et al. (1999) in an
adjacent locality; Bodon et al., 2005. Classified as
“LC” by Cuttelod et al. (2011) and by I.U.C.N.
(2014). Endemic species in Europe (Cuttelod et al.,
2011 ).
Familia ELLOBIIDAE Pfeiffer, 1854
Carychium tridentatum (Risso, 1826)
Long-toothed Herald Snail
Uncommon, 14 plots, locally abundant.
Familia LYMNAEIDAE Rafinesque, 1815
Galba truncatula (O.F. Muller, 1774)
Dwarf Pond Snail
Very rare, 2 plots. Museal data from 1986. Clas-
sified as “LC” by Cuttelod et al. (2011) and by
I.U.C.N. (2014).
Radix labiata (Rossmassler, 1835)
Lymnaea {Radix) peregra (O.F. Muller, 1774)
Radix peregra (O. F. Muller, 1774)
Wandering Snail
Very rare, 3 plots. Costa (1874): sub Limnaeus
Gibilmannicus (see O. G. Costa, 1839). Museal data
from 1986. Radix peregra is classified by I.U.C.N.
(2014) as synonym of Radix balthica (Linnaeus,
1758). Radix labiata is classified as “LC” by
Cuttelod et al. (2011).
Ordo PULMONATA Cuvier in Blainville, 1814
Subordo BASOMMATOPHORA Keferstein, 1864
Familia ANCYLIDAE
Ancylus fluviatilis O.F. Muller, 1774
River Limpet
Rare, 7 plots. Classified as “LC” by Cuttelod et
al. (2011) and by I.U.C.N. (2014).
Ferrissia fragilis (Tryon, 1863)
Ferrissia wautieri (Mirolli, 1960)
Fragile Ancylid
Very rare, 1 plot (Fig. 5). Classified as “LC” by
I.U.C.N. (2014). Cryptic invader of Italian fresh-
water ecosystems from North America (Cianfanelli
et al., 2007; Lori & Cianfanelli, 2007).
Subordo STYLOMMATOPHORA A. Schmidt, 1855
Familia PYRAMIDULIDAE Kennard et B.B.
Woodward, 1914
Pyramidula pusilla (Vallot, 1801)
Rock Snail
Common, 24 plots, locally abundant.
Pyramidula rupestris (Drapamaud, 1801)
Rock Snail
Very rare, 1 plot. Costa (1874).
Familia VERTIGINIDAE Fitzinger, 1833
A preliminary checklist of non-marine Molluscs from the Alburni Mountains, Campania, Southern Italy
381
Vertigo ( Vertigo ) pygmaea (Drapamaud, 1801)
Common Whorl Snail, Crested vertigo
Very rare, 1 plot. Classified as “LC” by Cuttelod
et al. (2011)
Vertigo ( Vertilla ) angustior Jeffreys, 1830
Vertigo sinistrorso minore, Narrow-mounthed
Whorl Snail
Very rare, 1 plot (Fig. 6). Species protected in
European Union by the Annex II of the “Habitats
Directive”, and in Italy by the D.P.R. n. 357/1997
than modified by D.P.R. n. 120/2003. In Europe,
this species is listed as Vulnerable (VU) (criteria:
A2ac+3c) at the European level and at the level
of the 27 member States of the European Union
(Cuttelod et al., 2011). The species is regionally
protected in Tuscany, Umbria and Emilia-Ro-
magna. The species has been regarded in Italy as
“NT” by Manganelli et al. (2000a). Classified as
“LR” by Cerfolli et al. (2002), and “NT” by
I.U.C.N. (2014).
Columella edentula (Drapamaud, 1805)
Toothless Chrysalis Snail
Very rare, 2 plots. Classified as “LC” by Cutte-
lod et al. (2011) and by I.U.C.N. (2014). Endemic
species in Europe (Cuttelod et al., 2011).
Truncatellina callicratis (Scacchi, 1833)
Rare, 10 plots. Endemic species in Europe clas-
sified as “LC” by Cuttelod et al. (2011).
Familia ORCULIDAE Pilsbry, 1918
Sphy radium doliolum (Bruguiere, 1792)
Uncommon, 12 plots. Classified as “LC” by
Cuttelod et al. (2011).
Pagodulina pagodula (des Moulins, 1830)
Very rare, 1 plot. Classified as “LC” by Cuttelod
et al. (2011) and by I.U.C.N. (2014). Endemic
species in Europe (Cuttelod et al., 2011).
Familia CHONDRINIDAE Steenberg, 1925
Rupestrella philippii (Cantraine, 1840)
Very rare, 4 plots. Classified as “LC” by Cutte-
lod et al. (2011).
Chondrina avenacea (Bruguiere, 1792)
Widespread, 36 plots. Museal data from 1874.
(Costa, 1874). Classified as “LC” by Cuttelod et al.
(2011) and by I.U.C.N. (2014). Endemic species in
Europe (Cuttelod et al., 2011).
Familia LAURIIDAE Steenberg, 1925
Lauria sempronii (Charpentier, 1837)
Rare, 8 plots.
Familia ARGNIDAE Hudec, 1965
Argna biplicata (Michaud, 1831)
Very rare, 4 plots. Classified as “LC” by Cutte-
lod et al. (2011) and by I.U.C.N. (2014). Endemic
species in Europe (Cuttelod et al., 2011).
Familia VALLONIIDAE Morse, 1864
Acanthinula aculeata (O.F. Muller, 1774)
Pricly Snail
Common, 21 plots. Classified as “LC” by Cutte-
lod et al. (2011).
Gittenbergia sororcula (Benoit, 1859)
Rare, 9 plots. Locally very abundant. Classified
as “LC” by Cuttelod et al. (2011).
Familia ENIDAE B.B. Woodward, 1903 (1880)
Chondrula tridens (O.F. Muller, 1774)
Very rare, 2 plots. Classified as “NT” by Cutte-
lod et al. (2011).
382
Agnese Petraccioli etalii
Jaminia quadridens (O.F. Muller, 1774)
Uncommon, 17 plots. Locally very abundant.
Museal data from 1986. Classified as “LC” by
Cuttelod et al. (2011) and by I.U.C.N. (2014).
Merdigera obscura (O.F. Muller, 1774)
Ena obscura (O.F. Muller, 1774)
Lesser Bulin
Uncommon, 11 plots. Classified as “LC” by
Cuttelod et al. (2011) and by I.U.C.N. (2014).
Endemic species in Europe (Cuttelod et al., 2011).
Familia PUNCTIDAE Morse, 1864
Pune turn pygmaeum (Drapamaud, 1801)
Dwarf Snail
Common, 33 plots.
Paralaoma servilis (Shuttleworth, 1852)
Paralaoma caputspinulae (Reeve, 1852)
Pinhead Spot
Very rare, 2 plots. Allochthonous species, intro-
duced from New Zealand (Lori et al., 2005; Lori &
Cianfanelli, 2007; Cianfanelli, 2009; Christensen,
2012 ).
Familia DISCIDAE Thiele, 1931 (1866)
Discus rotundatus (Muller, 1774)
Discus Snail, Rounded Snail
Rare, 7 plots.
Familia HELICODISCIDAE H.B. Baker, 1927
Lucilla scintilla (Lowe, 1852)
Oldfield Coil
Very rare, 1 plot (Fig. 7). Allochthonous species.
The indigenous distribution for this species includes
North America (Lori et al., 2005; Lori & Cianfan-
elli, 2007; Cianfanelli, 2009).
Lucilla singleyana (Pilsbry, 1889)
Smooth Coil
Very rare, 1 plot (Fig. 8). Allochthonous species.
Originally probably from North America, intro-
duced to Europe (Lori et al., 2005; Lori & Cianfan-
elli, 2007; Cianfanelli, 2009).
Familia VITRINIDAE Fitzinger, 1833
Vitrina cfr. pellucida (O.F. Muller, 1774)
Pellucid Glass Snail
Uncommon, 11 plots. Classified as “LC” by
Cuttelod et al. (2011).
Familia PRISTILOMATIDAE T. Cockerell, 1891
Vitrea subrimata (Reinhardt, 1871)
Common, 26 plots.
Vitrea etrusca (Paulucci, 1878)
Very rare, 4 plots.
Vitrea contracta (Westerlund, 1871)
Milky Crystal Snail
Common, 26 plots.
Familia ZONITIDAE Morch, 1864
Aegopis verticillus (Ferussac, 1822)
Very rare, 1 plot (Fig. 9).
Familia OXYCHILIDAE P. Hesse, 1927 (1879)
Retinella olivetorum (Gmelin, 1791)
Retinella olivetorum olivetorum (Gmelin, 1791)
Common, 33 plots. Museal data from 1986.
Oxychilus ( Oxychilus ) cfr. draparnaudi (Beck, 1837)
Drapamaud's Glass Snail, Dark-bodied Glass snail
Uncommon, 16 plots. Museal data from 1986.
A preliminary checklist of non-marine Molluscs from the Alburni Mountains, Campania, Southern Italy
383
Mediterranea hydatina (Rossmassler, 1838)
Very rare, 1 plot.
Daudebardia rufa (Drapamaud, 1805)
Widespread, 42 plots. Locally very abundant
(Figs. 10, 11, 12, 13).
Familia MILACIDAE Ellis, 1926
Tandonia sowerbyi (A. Ferussac, 1823)
Keeled Slug, Sowerby's Slug
Uncommon, 11 plots.
Familia LIMAC1DAE Lamarck, 1801
Limax maximus Linnaeus, 1758
Leopard Slug, Great Grey Slug, Giant Garde Slug
Rare, 6 plots.
Lehmannia marginata (O. F. Muller, 1774)
Tree slug
Very rare, 1 plot.
Limacus flavus (Linnaeus, 1758)
Yellow Slug, Tawny Garden Slug
Very rare, 2 plots.
Familia AGRIOLIMACIDAE H. Wagner, 1935
Deroceras reticulatum (O.F. Muller, 1774)
Netted Slug, Gray Fieldslug
Rare, 6 plots.
Deroceras invadens Reise, Hutchinson, Schun-
ach et Schlitt, 2011
Chestnut Slug, Brown Field Slug, Longneck
Fieldslug, Widespread Pest Slug
Familia EUCONULIDAE Baker, 1928
Euconulus fulvus (O.F. Muller, 1774)
Tawny Glass Snail, Brown Hive
Rare, 8 plots.
Familia FERUSSACIIDAE Bourguignat, 1883
Cecilioides acicula (O.F. Muller, 1774)
Blind Snail
Very rare, 5 plots.
Cecilioides ( Cecilioides ) veneta (Strobel, 1855)
Cecilioides janii (De Betta et Martinati, 1855)
Very rare, 5 plots.
Familia SUBULINIDAE P. Fischer et Crosse, 1877
Rumina decollata (Linnaeus, 1758)
Decollate Snail
Very rare, 4 plots.
Familia OLEACINIDAL H. Adams et A. Adams, 1855
Poiretia dilatata (Philippi, 1836)
Common, 31 plots. Museal data from 1986.
Familia TE S TACELLID AE J.E. Gray, 1840
Testacella scutulum G.B. Sowerbyi, 1821
Shield Shelled Slug
Very rare, 4 plots.
Familia CLAUSILIIDAE J.E. Gray, 1855
Door Snails
Medora sp.
Very rare, 4 plots.
Very rare, 3 plots.
384
Agnese Petraccioli etalii
Cochlodina ( Cochlodina)laminata (Montagu, 1803)
Plaited Door Snail
Rare, 6 plots. (Costa, 1874).
Charpentieria ( Stigmatica ) paestana (Philippi,
1836)
Siciliaria paestana (Philippi, 1836)
Widespread, 42 plots. Locally abundant.
Charpentieria ( Stigmatica ) cfr. ernae (Fauer, 1978)
Very rare, 2 plots. Museal data from 1970.
Endemic of Southern Appennine. (Fauer, 1978;
Welter- Schultes, 2012; Nordsieck, 2013).
Macrogastra ( Pyrostoma ) attenuata (Ross-
massler, 1835)
Macrogastra ( Pyrostoma ) attenuata iriana (Pol-
lonera, 1885)
Lined door snail
Very rare, 2 plots.
Clausilia cruciata (S. Studer, 1820)
Clausilia cruciata bonellii E. Von Martens, 1873
Very rare, 3 plots. Museal data before 1950.
Familia HYGROMIIDAE Tiyon, 1866
Xerotricha conspurcata (Drapamaud, 1801)
Very rare, 2 plots. Museal data from 1986.
Classified as “LC” by Cuttelod et al. (2011).
Hygromia cinctella (Drapamaud, 1801)
Girdled Snail
Rare, 8 plots. Classified as “LC” by Cuttelod et
al. (201 1) and by I.U.C.N. (2014). Endemic species
in Europe (Cuttelod et al., 2011).
Cernuella ( Cernuella ) cisalpina (Rossmassler,
1837)
Common, 21 plots. Locally abundant. Museal
data from 1986. Classified as “LC” by Cuttelod et
al. (2011) and by I.U.C.N. (2014). Endemic species
in Europe (Cuttelod et al., 2011).
Cernuella (Xerocincta) neglecta (Drapamaud, 1 805)
Luddesdown Snail, Neglected dune snail
Very rare, 3 plots. Museal data from 1986.
Classified as “LC” by Cuttelod et al. (2011) and by
I.U.C.N. (2014). Endemic species in Europe (Cutte-
lod et al., 2011).
Cernuellopsis ghisottii Manganelli etGiusti, 1988
Very rare, 3 plots. Hallgas, com. pers., 2013.
This species is endemic to Italy. Classified as “VU”
by Cuttelod et al. (2011) and by I.U.C.N. (2014)
(IUCN Criteria (Europe) (version 3.1):
Blab(iii)+2ab(iii). Endemic species in Europe
(Cuttelod et al., 2011).
Trochoidea ( Trochoidea ) pyramidata (Dra-
parnaud, 1805)
Very rare, 2 plots. Classified as “LC” by Cutte-
lod et al. (2011).
Trochoidea ( Trochoidea ) trochoides (Poiret,
1789)
Very rare, 2 plots. Museal data from 1985. Clas-
sified as “LC” by Cuttelod et al. (2011).
Monacha ( Monacha ) cfr. cartusiana (O.F.
Muller, 1774)
Chartreuse Snail, Carthusian Snail
Common, 35 plots. Museal data from 1986.
Classified as “LC”by Cuttelod et al. (2011) and by
I.U.C.N. (2014). Endemic species in Europe (Cutte-
lod et al., 2011). Regarded as “Edible species” in
Tuscany and Umbria Region.
Monacha ( Eutheba ) cfr. parumcincta (Menke,
1828)
Common, 26 plots. Classified as “LC” by Cutte-
lod et al. (2011) and by I.U.C.N. (2014). Endemic
species in Europe (Cuttelod et al., 2011).
Familia HELICODONTIDAE Kobelt, 1904
Helicodonta obvoluta (O.F. Muller, 1774)
Helicodonta obvoluta obvoluta (O.F. Muller, 1774)
Cheese Snail
A preliminary checklist of non-marine Molluscs from the Alburni Mountains, Campania, Southern Italy
385
Figures 2-9. Non-marine Molluscs from the Albumi Mountains, Campania, Southern Italy. Figure 2. Platyla talentii alive
(Photo by N. Maio). Figure 3. Shell of Platyla talentii'. view. 3A. Particular of dorsal view. 3B. Particular of the mouth
(Photos by N. Maio). Figure 4. Potamopyrgus antipodarum alive (Photo by N. Maio). Figure 5. Shell of F errissia fragilis'.
dorsal, lateral and ventral view (Photo by I. Niero). Figure 6. Shell of Vertigo ( Vertilla ) angustior : apertural view (Photo by
I. Niero). Figure 7. Shell of Lucilla scintilla', umbilical, dorsal and apertural view (Photo by N. Maio). Figure 8. Shell of
Lucilla singleyana : dorsal, apertural and umbilical view (Photo by N. Maio). Figure 9. Shell of Aegopis verticillus dorsal,
umbilical and apertural view (Photo by N. Maio).
386
Agnese Petraccioli etalii
Rare, 7 plots. Classified as “LC” by Cuttelod et
al. (2011) and by I.U.C.N. (2014). Endemic species
in Europe (Cuttelod et al., 2011).
Familia HELICIDAE Rafmesque, 1815
Chilostoma ( Campylea ) cfr. planospira (Lamarck,
1822)
Uncommon, 18 plots. Museal data from 1986.
Classified as “DD” by Cuttelod et al. (2011) and by
I.U.C.N. (2014). Endemic species in Europe (Cutte-
lod et al., 2011).
Marmorana ( Ambigua ) fuscolabiata fuscola-
biata (Rossmassler, 1842)
Widespread, 27 plots. Costa (1874), Degner
(1927). Museal data from 1874. Classified as
“DD” by Cuttelod et al. (2011) and by I.U.C.N.
(2014). Endemic species in Europe (Cuttelod et
al., 2011).
Marmorana {Ambigua) cfr. fuscolabiata wullei
Kobelt, 1903
Common, 25 plots. Kobelt (1903a; 1903b), De-
gner (1927), Bacci (1951), Alzona (1971). Museal
data from 1903.
Eobania vermiculata (O.F. Muller, 1774)
Chocolate-band snail
Very rare, 3 plots. Museal data from 2006. Re-
garded as “Edible species” in Tuscany and Umbria
Region.
Cantareus apertus (von Bom, 1778)
Green Garden Snail
Very rare, 2 plots. Museal data from 1986. Re-
garded as “Edible species” in Tuscany and Umbria
Region.
Cornu aspersum (O.F. Muller, 1774)
Cantareus aspersus (O.F. Muller, 1774)
Garden Snail, Common Snail, Brown Garden Snail
Very rare, 4 plots. Museal data from 1986. Re-
garded as “Edible species” in Tuscany and Umbria
Region.
Helix (Helix) cfr. delpretiana Paulucci, 1878
Rare, 9 plots. Museal data from 1986. This
species is endemic to the Central Appenines in Italy.
Classified as “DD” by Cuttelod et al. (2011) and by
I.U.C.N. (2014). Endemic species in Europe (Cutte-
lod et al., 2011).
Helix (Helix) cfr. ligata O.F. Muller, 1774
Ligate Snail
Uncommon, 11 plots. Museal data from 1986.
Classified as “LC” by Cuttelod et al. (2011) and by
I.U.C.N. (2014). Endemic species in Europe (Cutte-
lod et al., 2011).
Classis BIVALVI A Linnaeus, 1758
Ordo VENEROIDA H. et A. Adams, 1857
Familia SPHAERIIDAE Deshayes, 1855 (1820)
Pisidium casertanum (Poli, 1791)
Caserta Pea Mussel
Very rare, 5 plots. Classified as “LC” by Cutte-
lod et al. (2011) and by I.U.C.N. (2014).
Species exclusively known from the literature
Familia VERTIGINIDAE Fitzinger, 1833
Vertigo (Vertigo) moulinsiana (Dupuy, 1849)
Vertigo of Demoulins, Demoulins’ Whorl Snail
Manganelli et al. (2001), Bodon et al. (2005).
Find only in debris of Fiume Calore, near Grotta di
Caste lcivita, Salerno Province, by S. Cianfanelli
and E. Talenti on 1994. Species protect in European
Union by the Annex II of the “Habitats Directive”
and in Italy by the D.P.R. n. 357/1997 than modified
by D.P.R. n. 120/2003. The species is regionally
protect in Tuscany, Umbria and Emilia-Romagna.
The species has been initially classified as
“LRcd”(= Lower risk, conservation dependant) by
Bouchet et al. (1999), then the species has been re-
garded in Italy as “VU”(criteria: B2a, B2b) by
Manganelli et al. (2000b, 2001). Classified as “LR”
by Cerfolli et al. (2002), “VU” (criteria: A2ac) by
Cuttelod et al. (2011) and by I.U.C.N. (2014).
A preliminary checklist of non-marine Molluscs from the Alburni Mountains, Campania, Southern Italy
387
Figure 10. Shell of Daudebarclia rufa: dorsal, apertural and umbilical view (Photo by I. Niero). Figure II. D. rufa alive
(Photo by N. Maio). Figure 12. Genitalia of D. rufa (Sant’Angelo a Fasanella (SA), 1 160 m, 17.0V.20 13, N. Maio, P. Crovato
& I. Niero legit. Figure 13. Internal structure of the penis. Acronyms in figures: AG= albumen gland; DBC= duct of the
bursa copulatrix; E= epiphallus; EP= epiphallic pore; FO= free oviduct; GA= genital atrium; HD= hermaphroditic duct;
F1G= hermaphroditic gland; P= penis; PPL= penis pleats; PR= penial retractor; Pro= prostate; PVG= perivaginal gland; PS
= penis sheath; RS = reservoir of spermatheca; T= talon; UOS= uterine ovispermiduct; Va= vagina, VD= vas deferens.
(Drawn by I. Niero).
388
Agnese Petraccioli etalii
Familia CLAUSILIIDAE J.E. Gray, 1855
Macrogastra cfr. plicatula (Drapamaud, 1801)
Costa (1874: sub Clausilia plicatula Drap.).
Museal data before 1950.
From the South of Italy are cited 3 subspecies,
in addition to M. plicatula plicatula : M. p. amiaten-
sis Nordsieck, 2006; M. p. apennina Gentiluomo
1868, and M. p. aprutica Nordsieck, 2006 (Nord-
sieck, 2006).
Familia HYGROMIIDAE Tryon, 1866
Cernuella cfr. virgata (Da Costa, 1778)
Banded Snail
Kobelt (1907). Museal data from 1986. Regarded
as “Edible species” in Tuscany and Umbria Region.
Familia OXYCHILIDAE P. Hesse, 1927 (1879)
Oxychilus sp.
Capasso (1958), Capolongo & Cantilena (1974).
Ordo HYGROPHILA A. Ferussac 1822
Familia PHYSIDAE Fitzinger, 1833
Haitia acuta (Drapamaud, 1805)
Acute Bladder Snail, European physa, Tadpole
Snail, Bladder Snail, Pewter physa
Sacchi (1964): sub Physa acuta. Allochthonous
species. The oldest alien species of Italy probably
native to northeastern North America (Lori et al.,
2005; Lori & Cianfanelli, 2007; Cianfanelli, 2009).
DISCUSSION
Up to now, only 13 non-marine Mollusc species
were known from the Albumi Mountains through
bibliographical data (nine species of land snails and
four species of freshwater snails), to which we add
the new species listed in this paper for the study
area. The presence of nine species (six species of
land snails and three species of freshwater
molluscs) are confirmed by our field investigation.
Only four species (the land snails Macrogastra
plicatula , Vertigo moulinsiana, Cernuella virgata
and the allochthonous freshwater snails Haitia
acuta ) were documented exclusively by biblio-
graphical data and have not been confirmed by the
field surveys yet (Fiorentino et al., 2008; Reise et
al., 2011). Our analysis identifies 79 species of non-
marine molluscs (69 species of land snails, ten
species of freshwater molluscs) recorded on the
basis of field data. In total the occurrence of 83
species of non-marine molluscs (73 species of land
snails, 10 species of freshwater molluscs) was
herein attested in the survey area representing
approximately the 56% of the estimated fauna of
Campania Region (about 150 species, personal
data).
At least 1 1 species are new records for the Cam-
pania Region ( Aegopis verticillus, Cernuellopsis
ghisottii, Lucilla scintilla, Argna hiplicata,
Cernuella neglecta, Daudebardia rufa, Helix
delpretiana, Pagodulina pagodula, Vitrina pellu-
cida, Vitrea etrusca and Macrogastra attenuata). M.
attenuata (sub M. lineolata ) and D. rufa were
generically recorded from Matese Mountains
(probably Molise) by Giusti et al. (1985). 70 species
of non-marine molluscs (64 species of land snails,
six species of freshwater molluscs), recorded on the
basis of field data, have not been previously re-
corded from the study area.
Extremely interesting is the finding of samples
of Medora sp.: it seems to be the second record of
this genus for the region. The systematics of the
genus Medora Adams, 1855 is in fact complex and,
in many respects, still controversial. Regarding
Italy, Nordsieck (1970) considered M. italiana
(Kiister, 1 847) of the Central-Southern Apennines
distinct from M. albescens (Menke, 1830) of the
Balkan peninsula. In addition, he assigned to M.
italiana various subspecies: only one from Cam-
pania: M. i. italiana (Kiister, 1847) (locus typicus:
Piedimonte d'Alife (= Piedimonte Matese, Caserta,
Campania). Giusti et al. (1986) suggested that it
was not possible to distinguish M. italiana from M.
albescens with the subspecies M. a. italiana in the
central part of Italy. The populations reported for
Italy as M. dalmatina (Manganelli et al., 1995) were
described by Nordsieck (2012) as a distinct sub-
A preliminary checklist of non-marine Molluscs from the Alburni Mountains, Campania, Southern Italy
389
species. Preliminary data by Colomba et al. (2012)
suggest that the genus Medora shows a much more
complex and articulate differentiation than hitherto
hypothesized by morphological surveys so far. An
attempt to clarify its organization and internal struc-
ture, at various taxonomic levels, a more detailed
analysis including a higher number of molecular
markers and additional Medora populations from
Italy are required.
Other interesting records are: Vertigo angustior,
a species protected in the European Union by the
Annex II of the Council Directive 92/43/EEC of
May 2 1th 1992 on the conservation of natural
habitats and of wild fauna and flora known as
“Habitats Directive”, that includes “animal and
plant species of community interest whose conser-
vation requires the designation of special areas of
conservation (SPA)” and listed as “Vulnerable” at
the European level and Platyla talentii, an endemic
species of Southern Apennine, recently described
by Bodon & Cianfanelli (2008), classified as “Near
Threatned” by Cuttelod et al. (2011) and by
I.U.C.N. (2014).
Five allochthonous species were surveyed for
the first time in the study area: three land snails
(Lucilla scintilla, L. singleyana and Paralaoma
servilis ) and two freshwater snails ( Potamopyrgus
antipodarum and Ferris siafragilis). L. scintilla and
L. singleyana are native in North America; they
were probably introduced into Europe in the second
half of the 20th century (Horsak et al., 2009). L.
singleyana, P. servilis and F.fragilis are the second
records for the Campania Region, the first is Bodon
et al. (2004) sub Flelicodiscus singleyanus (Pilsbry,
1890), Bodon et al. (2004) sub P. caputspinulae
(Reeve, 1852) and D' Antonio & Bravi (1990) sub
F. wautieri (Mirolli, 1960).
In the past, three endemic taxa had been de-
scribed as new for the Alburni area: Helix ( Iberus )
wullei first described by Kobelt (1903a: 14-15, taw.
1766-1768) from “Monte Alburno circa vicum
Postiglione prov. Salernitanae” and later as Iberus
wullei (Kobelt, 1903b: pag. 4-5, fig. without num-
ber); Xerophila ( Xerolauta ) peninsularis forma
“ alburnF described by Kobelt (1907: pp. 59-60,
Tafel 357 Fig. 2221) from “Monte Postiglione, des
alten Albumus” and Siciliaria ernae described by
Fauer (1978: 265, abb. 1) from “Passo Sentinella
im SO der Monti Abumi” [municipality of Corleto
Monforte], and “6 km West of San Rufo”. Today X.
peninsularis is considered a junior synonym of
Cernuella virgata (Da Costa, 1778) but the taxo-
nomic status of Iberus (= Marmorana ) wullei and
of S. ernae need to be confirmed. The Demoulins’
Whorl Snail, listed in the Standard Data Form of
the SCI of “Monti Alburni” is not confirmed by our
field surveys.
A Red List of Threatened Species is also pro-
posed and the species were classified with the code
of I.U.C.N. (Version 2014.3).
ACKNOWLEDGEMENTS
This research was funded with the support of the
Cilento, Vallo di Diano and Alburni National Park.
Authors wish to thank: the National Park Authority
of the Cilento, Vallo di Diano and Alburni for
collaboration as well as advising on technical and
administrative aspects. In particular: Amilcare
Troiano (President), Corrado Matera (Vice-President)
and Angelo De Vita (Director). A special thank goes
to Pasquale and Pietro Cappelli (Ottati, Salerno) for
the trekking guide. We want to thank: Mario
Cuomo, Sergio Duraccio, Gianni D’Anna, Franco
Izzillo, and Massimo Cretella (Naples) and Aless-
andro Hallgass (Rome) for the unpublished data
given; Antonio Pietro Ariani for the malacological
collection consultation of the Zoological Museum
of Naples; Monica Leonardi (Conservator) for the
malacological collection consultation of the Museo
Civico di Storia Naturale di Milano, Maria Tavano
(Conservator) and Giacomo Doria (Director) for the
malacological collection consultation of the Museo
Civico di Storia Naturale di Genova “Giacomo
Doria”. Furthermore we thank: Domenico Davolos
(Dipartimento di Biologia, Universita di Napoli
Federico II), Antonietta Mordente (Banca Monte
Pruno, Credito Cooperative di Roscigno e Laurino)
and Nicola Auricchio (President of Fondazione
I.RI.DI.A. - Museo Naturalistico of Corleto
Monforte) and Antonio Sicilia (Mayor of Municip-
ality of Corleto Monforte) for administrative sup-
port and cooperation.
REFERENCES
Alzona C., 1971. Malacofauna italica. Catalogo e biblio-
grafia dei molluschi viventi, terrestri e d’acqua dolce.
390
Agnese Petraccioli etalii
Parte I. Atti della Societa Italiana di Scienze Naturali
e del Museo Civico di Storia Naturale di Milano,
Milano, 111: 1^433.
Bacci G., 1951. Le razze di Ambigua fuscolabiata
(Rossm.) (Polmonata - Helicidae): un problema di
sistematica e di genetica. Annuario dell’Istituto e
Museo di Zoologia delPUniversita di Napoli (N. S.),
3: 10-24.
Bank R. A., 2011. Fauna Europaea: Mollusca Gastro-
poda. Fauna Europaea version 2.4. Checklist of the
land and freshwater Gastropoda of Italy. Fauna
Europaea Project: 1-49. http://www.nmbe.unibe.ch/
sites/default/files/uploads/pubinv/fauna europaea -
gastropoda of italy.pdf. Last update: November
24th, 2013.
Bech M., 1990. Fauna malacoldgica de Catalunya. Mol-
luscs terrestres i d’aigua dolqa. Treballs de la Institu-
cio Catalana d’Histdria Natural. N. 12. Barcelona.
Bodon M. & Cianfanelli S., 2008. Una nuova specie di
Platyla per il sud Italia (Gastropoda: Prosobranchia:
Aciculidae). Bollettino Malacologico, 44: 27-37.
Bodon M., Favilli L., Giannuzzi Savelli R., Giovine F.,
Giusti F., Manganelli G., Melone G., Oliverio M.,
Sabelli B. & Spada G., 1995. Gastropoda; Prosobran-
chia, Heterobranchia Heterostropha. In: Minelli A.,
Ruffo S. & La Posta S. (Eds.), Checklist delle specie
della fauna italiana. Calderini, Bologna, 14, 60 pp.
Bodon M., Cianfanelli S., Talenti E., Manganelli G. &
Giusti F., 1999. Litthabitella chilodia (Westerlund,
1886) in Italy (Gastropoda Prosobranchia: Hy-
drobiidae). Hydrobiologia, 4 1 1 : 175-189.
Bodon M., Lori E. & Cianfanelli S., 2004. Un'altra specie
aliena per la malacofauna italiana: Hawaiia minus-
cula (Binney, 1840) (Pulmonata: Zonitidae). Bol-
lettino Malacologico, 40: 11-14.
Bodon M., Cianfanelli S., Manganelli G., Pezzoli E. &
Giusti F., 2005. Gastropoda Prosobranchia e Hetero-
branchia Heterostropha d’acqua dolce. In: Ruffo S.
& Stoch F. (Eds.), Checklist e distribuzione della
fauna italiana. Memorie del Museo Civico di Storia
naturale di Verona, 2 Serie Sez. Scienze della Vita,
16: 79-81 + Cdrom. http: //ckmap.faunaitalia. it.
Bouchet P., Falkner G., Seddon M. B., 1999. Lists of
protected land and freshwater molluscs in the Bern
Convention and European Habitats Directive: are
they relevant to conservation? Biological Conserva-
tion, 90: 21-31.
Castagnolo L., 1995. Bivalvia (specie d’acqua dolce:
generi 064-066, 128, 132-134). In: Minelli A., Ruffo
S. & La Posta S. [a cura di], Checklist delle specie
della fauna d’ltalia. Calderini, Bologna. 17 (Bivalvia,
Scaphopoda), 21 pp.
Capasso P., 1958. Speleologia termale. Notapreliminare
sulla localita di Contursi (Salerno) e le sue mofete.
Studia Spelaeologica, 3 (giugno): 95-102.
Capolongo D. & Cantilena S., 1974. Specie cavernicole
di Campania. Annuario dell’Istituto e Museo di
Zoologia delPUniversita di Napoli, 20: 1-198.
Cerfolli F., Petrassi F. & Petretti F., 2002. II Libro Rosso
degli Animali Invertebrati d’ltalia. WWF Italia -
ONLUS - Roma.
Christensen C., 2012. First Records of Paralaoma sennlis
(Shuttleworth, 1852) (Gastropoda: Pulmonata:
Punctidae) in the Hawaiian Islands. Records of the
Hawaii Biological Survey for 2011. Bishop Museum
Occasional Papers, 112: 3-7.
Cianfanelli S., 2009. I Molluschi della Provincia di
Pistoia: le specie da tutelare e quelle da combattere.
Quaderni del Padule di Fucecchio n. 6. Centro di
Ricerca, Documentazione e Promozione del Padule
di Fucecchio, 112 pp.
Cianfanelli S., Lori E. & Bodon M., 2007. Non-
indigenous Alien freshwater molluscs in Italy and
their distribution. In: Gherardi F (Ed.). Biological
invaders in inland waters: profiles, distribution, and
threats. Springer, Dordrecht, The Netherlands.
Chapter 5, pp 103-121, http://dx.doi.org/10.1007/
978-1-4020-6029-8 5
Colomba M.S., Liberto F., Reitano A., Renda W., Poca-
terra G., Gregorini A. & Sparacio I., 2012. Molecular
studies on the genus Medora H. et A. Adams, 1855
from Italy (Gastropoda Pulmonata Clausiliidae).
Biodiversity Journal, 3: 571-582.
Costa O. G., 1839. Cenni sulla Fauna Siciliana. In: Costa
O. G. (a cura di), Corrispondenza zoologica destinata
a diffondere nel Regno delle Due Sicilie tutto cio che
si va discuoprendo entro e fuori Europa (e vice-versa)
riguardante la zoologia in generale. Tip. Azzolino &
Compagno, Napoli. Anno I (11-12): 159-184.
Costa A., 1874. Una peregrinazione zoologica su’ monti
dell’Alburno. Rendiconti della Reale Accademia
delle scienze fisiche e matematiche di Napoli (serie
1), 13: 129-135.
Cuttelod A., Seddon M. & Neubert E., 2011. European
Red List of Non-marine Molluscs. Luxembourg:
Publications Office of the European Union, 98 pp.
D' Antonio C. & Bravi S., 1990. Ricerche naturalistiche
nella tenuta Astroni (Napoli). I. La malacofauna
acquatica. Lavori S.I.M., Atti Congresso di Sorrento
23-31 maggio 1987, 23: 483-493.
Degner E., 1927. Zur Molluskenfauna Unteritaliens.
Mitteilungen aus dem Zoologischen Staatsinstitut
und Zoologischen Museum in Hamburg, 43: 19-1 14.
Fauer W., 1978. Siciliaria ernae n. sp., eine neue rezente
Clausilie aus Italien. Archiv fiir Molluskenkunde,
108: 263-265.
Fiorentino V., Salomone N., Manganelli G. & Giusti F.
2008. Phylogeography and morphological variability
in land snails: the Sicilian Marmorana (Pulmonata,
Helicidae). Biological Journal of the Linnean Society,
94: 809-823.
A preliminary checklist of non-marine Molluscs from the Alburni Mountains, Campania, Southern Italy
391
Girod A., Bianchi I. & Mariani M., 1980. Gasteropodi, 1
(Gastropoda: Pulmonata, Prosobranchia: Neritidae,
Bithyniidae, Valvatidae). Guida per il riconoscimento
delle specie animali delle acque interne italiane.
C.N.R., Roma, 86 pp.
Giusti F. & Pezzoli E., 1980. Gasteropodi, 2 (Gastropoda:
Prosobranchia: Hydrobioidea, Pyrguloidea). Guida
per il riconoscimento delle specie animali delle acque
interne italiane. C.N.R., Roma, 67 pp.
Giusti F., Grappelli C., Manganelli G., Fondi R. & Bulli
L., 1986. An attempt of natural classification of the
genus Medora in Italy and Yugoslavia, on the basis
of conchological, anatomical and allozymic charac-
ters (Pulmonata: Clausiliidae). Lavori della Societa
Italiana di Malacologia, 22: 259-341.
Giusti F., Castagnolo L. & Manganelli G. 1985. La fauna
malacologica delle faggete Italiane: brevi cenni di
ecologia, elenco delle specie e chiavi per il ricono-
scimento dei generi e delle entita piu comuni. Bol-
lettino Malacologico, 21: 69-144.
Giusti F., Manganelli G. & Schembri P.J., 1995. The non-
marine molluscs of the Maltese Island. Museo
Regionale di Scienze Naturali, Torino, 607 pp.
Horsak M., Steffek J., Cejka T., Lozek V. & Jurickova
L., 2009. Occurrence of Lucilla scintilla (R.T. Lowe,
1852) and Lucilla singleyana (Pilsbry, 1890) in the
Czech and Slovak Republics - with remarks how to
distinguish these two non-native minute snails.
Malacologica Bohemoslovaca, 8: 24-27.
I.U.C.N., 2014. IUCN Red List of Threatened Species.
Version 2014.3. Disponibile su www.iucnredlist.org.
Downloaded on 22 June 2014.
Janus H., 1982. The illustrated Guide to Molluscs. Harold
Starke Limited, London, 180 pp.
Kerney M.P. & Cameron R.A. D., 1999. Guide des
escargots et limaces d’Europe identification et
biologie de plus de 300 especes. Delachaux et
Niestle, Paris, 370 pp.
Kobelt W., 1903a. N. 1766-1768. Helix ( Iberus ) wullei
n. Iconographie der Land- und Susswasser-
Molluscken mit vorzuglicher berucksichtigung der
Europaischen noch nicht abgebildeten arten von E.
A. Rossmassler. Neue Folge, vol. 10. C.W. Kreidel’s
Verlag, Wiesbaden, 107 pp.
Kobelt W., 1903b. Diagnoses Heliceorum novorum in
Italia collectorum. Annuario del Museo Zoologico
della Reale Universita di Napoli (N. S.), 1: 1-5.
Kobelt W., 1907. Iconographie der Land- & Siiss-
wasser-Mollusken mit worzuglicher Berucksichtigung
der Europaischen noch nicht abgebildeten Arten
von E.A. Rossmassler. C.W. Kreidel's Verlag.
Wiesbaiden, n.f. [n.s.], tomo [band] 13, pp. 1-65,
taw. 331-360.
Lori E. & Cianfanelli S., 2007. Studio sulla presenza e
distribuzione di Molluschi terrestri e d’acqua dolce
alieni nel territorio della Provincia di Pistoia.
Relazione finale 2007. Museo di Storia Naturale
dell’ Universita degli Studi di Firenze, 97 pp.
Lori E., Bodon M. & Cianfanelli S., 2005. Molluschi
continentali alieni in Italia: presenza e distribuzione.
Notiziario S.I.M., 23: 71.
Manganelli G., Bodon M., Favilli L. & Giusti F., 1995.
Gastropoda Pulmonata. In: Minelli A., Ruffo S., La
Posta S. [a cura di], Checklist delle specie della fauna
d'ltalia, Bologna, 16, 60 pp.
Manganelli G., Bodon M., Favilli L., Castagnolo L. &
Giusti F., 1998. Checklist delle specie della fauna
d’ltalia, molluschi terrestri e d’acqua dolce. Errata ed
addenda, 1. Bollettino Malacologico, 33: 151-156.
Manganelli G., Bodon M. & Giusti F., 2000a. Checklist
delle specie della fauna d’ltalia, molluschi terrestri
e d’acqua dolce. Errata e addenda, 2. Bollettino
Malacologico, 36: 125-130.
Manganelli G., Bodon M., Cianfanelli S., Favilli L.,
Talenti E. & Giusti F., 2000b. Conoscenza e conser-
vazione dei molluschi non marini italiani: lo stato
delle ricerche. Bollettino Malacologico, 36: 5-42.
Manganelli G., Cianfanelli S., Brezzi M. & Favilli L.,
2001. The distribution and Taxonomy of Vertigo
moulinsiana (Dupuy, 1849) in Italy (Gastropoda:
Pulonata: Vertiginidae). Journal of Conchology, 37:
267-280.
Nordsieck H. 1970. Zur Anatomie und Systematik der
Clausilien, VIII. Dinarische Clausiliidae, II: Das
Genus Medora. Archiv fur Molluskenkunde, 100:
23-75.
Nordsieck H., 2002. Contributions to the knowledge of
the Delimini (Gastropoda): Stylommatophora:
Clausiliidae. Mitteilungen der Deutschen malakozo-
ologischen. Gesellschaft, 67: 27-39.
Nordsieck H., 2006. Systematics of genera Macrogastra
Hartmann 1841 and Julica Nordsieck 1963, with the
description of new taxa (Gastropoda: Stylommato-
phora: Clausiliidae). Archiv fur Molluskenkunde,
135: 49-71.
Nordsieck H., 2012. Erganzung der Revision der Gattung
Medora H. & A. ADAMS: Die Medora- Arten Italiens
(Gastropoda, Stylommatophora, Clausiliidae, Alopi-
inae), mit Beschreibung einer neuen Unterart von
Medora dalmatina Rossmassler. Conchylia, 42: 75-81.
Nordsieck H., 2013. Delimini (Gastropoda, Pulmonata,
Clausiliidae) from Apennine Italy, with the descrip-
tion of three new subspecies from Calabria. Conchy-
lia, 44:3-14.
Petraccioli A., Barattolo F., Crovato P., Cretella M., Maio
N. & Aprea G., 2005a. Guida pratica al riconosci-
mento dei macro-gasteropodi terrestri attuali e fossili
dell’Isola di Capri. Bolletino della Sezione Campania
ANISN (N. S.), 16(29): 19-48.
392
Agnese Petraccioli etalii
Petraccioli A., Crovato P., Cretella M., Maio N., Aprea
G. & Barattolo F., 2005b. I Gasteropodi continentali
recenti dell’Isola di Capri. In: Crovato P. & Maio N.
(Eds.). IV International Congress of the European
Malacological Societies. October 10-14 2005, Naples
(Italy). Abstracts of Oral Communications and
Posters. Notiziario S.I.M., 23: 28.
Petraccioli A., Crovato P., Cretella M., Maio N., Aprea
G. & Barattolo F., 2006a. I Gasteropodi continentali
dell'Isola di Capri: risultati preliminari. In: Gugliemi
R. & Nappi A. (Eds.), Atti del Convegno: “La Natura
in Campania: aspetti biotici e abiotici”. Napoli, 18
novembre 2004. Gruppo Attivo Campano A.R.C.A.
- Onlus, pp. 77-86.
Petraccioli A., Maio N., Crovato P. & Picariello O.,
2006b. I Gasteropodi terrestri del Parco Nazionale
del Vesuvio. Risultati preliminari. 67° Congresso
Nazionale Unione Zoologica Italiana. Napoli, 12-15
settembre 2006. Riassunti dei contributi scientifici:
84-85.
Petraccioli A., Crovato P., Cretella M., Maio N., Aprea
G. & Barattolo F., 2007. The fossil land Gastropods
from Capri Island. IV International Congress of the
European Malacological Societies. October 10-14
2005, Naples (Italy). Bollettino Malacologico, 43:
51-56.
Picariello O., Maio N., Petraccioli A., Crovato P. &
Guarino F.M., 2011. Emergenze Faunistiche del
Parco Nazionale del Vesuvio. Atti XV Convegno
Nazionale A.N.I.S.N.: “La Natura, l'Uomo, il
Tempo”. Napoli, 7-12 settembre 2010. Le Scienze
Naturali nella Scuola. Periodico A.N.I.S.N., 20 (44-
III): 81-93.
Ponder W.F. & Lindberg D.R., 1996. Gastropod philo-
geny: Challenges for the 90s. In: Taylor J. (Ed.).
Origin and evolutionary radiation of the Mollusca.
Oxford University press., Oxford, pp. 135-154.
Reise H., Hutchinson J.M.C., Schunack S. & Schlitt B.,
2011. Deroceras panormitanum and congeners from
Malta and Sicily, with a redescription of the wide-
spread pest slug as Deroceras invadens n. sp. Folia
Malacologica, 19: 201-233.
Sacchi C.F., 1964. Origini ed evoluzione della malaco-
fauna appenninica meridionale. Annuario dell'Istituto
e Museo di Zoologia delfUniversita di Napoli,
15/1963 (7): 1-85.
TEMI s.r.l. (a cura di) (2010). Piano di gestione del Sito
di Importanza Comunitaria "Monti Alburni"
(IT8050033) e della Zona di Protezione Speciale
"Alburni" (IT805055). Progetto Life Natura
"LIFE06NAT/IT/000053 " "Gestione della Rete di
SIC/ZPS nel PN del Cilento e Vallo di Diano"
(Cilento in Rete), 294 pp.
Welter-Schultes F.W., 2012. European non-marine
molluscs, a guide for species identification. Planet
Poster Editions, Gottingen, 679 pp.
Biodiversity Journal, 2015, 6 (1): 393-400
Monograph
Distribution of two Amphiope L. Agassiz, 1 840 (Echinoidea
Clypeasteroida) morphotypes in the Western-Proto-Mediter-
ranean Sea
Paolo Stara 1 *, Federico Marini 2 , Giuseppe Carone 3 & Enrico Borghi 4
'Centro Studi di Storia Naturale del Mediterraneo, Museo di Storia Naturale Aquilegia, Via Italia 63, Pirri-Cagliari and Geomuseo
Monte Arci, Masullas, Oristano, Sardegna, Italy; e-mail: paolostara@yahoo.it
2 University of Florence, Earth Sciences Department, Via la Pira 4, 50121 Florence, Italy; e-mail:
federico.marini.87@gmail.com
3 Civico Museo Paleontologico di Ricadi, Palazzo Fazzari, Via Strada Provinciale, 89866 Santa Domenica di Ricadi (W),
Italy; e-mail: p.carone@libero.it
4 Societa Reggiana di Scienze Naturali, Via Tosti 1, 42124 Reggio Emilia, Italy; e-mail: enrico.borghi20@teletu.it
^Corresponding author
ABSTRACT Several species belonging to the genus Amphiope L. Agassiz, 1 840 (Echinoidea Astriclypeidae)
from the Mediterranean Oligo-Miocene have been synonymised with A. bioculata (Des
Moulins, 1835), the type-species of the genus, based on the interpretation given by Philippe
(1998) as a taxon characterized by a large amount of morphological variability. A recent study
introduced the characters of the internal test structure and the plating patterns as taxonomic
tools in this genus. That paper indicated the occurrence of at least five different species in the
examined sample from the Oligo-Miocene of Sardinia, thus pointing to a previous over-
estimation of the variability-range of the type-species and to the need of a review of the largely
unresolved taxonomy of Amphiope. According to a recent study, Amphiope is considered as a
shallow-water echinoid, inhabiting sandy bottoms with high hydrodynamic energy; so it
represents a coastline marker, useful for the study of the paleo-geographic changes occurred
in the Proto-Western-Mediterranean during the Miocene. The diffusion and speciation of
Amphiope were highly influenced by those changes. In particular, the speciation rate of this
genus was likely favored by the occurrence of isolated populations created when islands (e.g.:
Baleares, Calabria, Corse, Kabylies, Sardinia) separate from the mainland, above all in the we-
stern part of that Basin, because of the opening of the Balearic Basin during the Late Oligo-
cene-Early Miocene and of the Tyrrhenian Sea during the Burdigalian-Tortonian (references
in this work). Two main morphotypes of Amphiope sensu Stara & Sanciu (2014), developed
in the Western Mediterranean from the late Oligocene to the late Miocene. They are herein
called the “ bioculata ” group, characterized by roundish to broad elliptical lunules with major
diameter/minor diameter ratio (SI) < 1.59, and the "nuragica" group, with more or less narrow
lunules and SI > 1.6. According to this authors, most Miocene forms with narrow elliptical
lunules would derive from A. nuragica (Comaschi Caria, 1955), late Oligocene-early Miocene
of Sardinia, the most archaic form so far known of this genus. The forms belonging to the
“ bioculata ” group likely derived from a different common ancestor bearing round to broad
ovoidal lunules. “A. bioculata ” described by Cottreau (1914), from the Burdigalian (Philippe,
1998) of Saint Cristol (Nissan, Herault, France), is so far the most ancient known form belong-
ing to this group. This work proposes a possible speciation sequence of the “ nuragica ” group.
KEY WORDS Amphiope ; Western-Proto-Mediterranean Sea; Paleogeography.
Received 12.07.2014; accepted 09.11.2014; printed 30.03.2015
Proceedings of the 2nd International Congress “Speciation and Taxonomy”, May 1 6th- 1 8th 2014, Cefalu-Castelbuono (Italy)
394
Paolo Stara et alii
INTRODUCTION
Paleogeography and paleoecology
Amphiope L. Agassiz, 1840 (Echinoidea Ast-
riclypeidae) is considered as a shallow- water echin-
oid, typical of sandy settings characterized by high
hydrodynamic energy (Stara et al., 2012). According
to Stara & Rizzo (2013, 2014) and Stara & Sanciu
(2014) it represents also a valid coastline marker.
On the basis of the fossil record and the avail-
able paleoecological data it is herein hypothesized
that the diffusion of Amphiope was highly influ-
enced by the paleogeographic (Doglioni et al.,
1998; Rosenbaum et al., 2002; Carminati et al.,
2012, Stara & Rizzo, 2014) and paleoecological
(Popescu, 2009) changes occurred in the Western
Proto-Mediterranean during the Miocene. In partic-
ular the opening of the Balearic and Ligurian Basins
during the Late Oligocene-Early Miocene and
of the Tyrrhenian Sea during the Burdigalian-
Messinian (Doglioni et al., 1998; Rosembaum et al.,
2002; Carminati et al., 2012) originated islands
(e.g.: Baleares, Calabria, Corse, Kabylies, Sardinia)
separated by deep water, thus leading to the occur-
rence of isolated populations and favoring speci-
ation within this Astriclypeid genus.
The orogenetic trend in the Mediterranean area
mainly derived from the differential movement
between the Adria microplate, belonging to the
African plate, and the European one.
The geodynamic and paleogeographic evolution
of the Western Mediterranean may be divided into
two distinct phases: the first occurred during the
Chattian-Burdigalian, the second started in the
Burdigalian and it is still active today.
First phase - The migration of the Sardinia-
Corsica microplate and the Calabrian block, with
respect to the more stable European plate, likely
began 25-23 My ago, with a general translation
towards SE. This drift was accompanied by a 45°
counterclockwise rotation of the Sardinia-Corsica
microplate between 20.5 and 15 Ma, with a broad
oceanic domain in the Li guro- Provencal basin (up
to 400 km in te southern part) between 20.5 and 18
Ma (Gattacceca et al., 2007). These evidences
firstly improved the presence of a connection
between the Liguro-Pro venial Basin and the Valencia
trough before 20.5 Ma, then the connection between
the Alboran and the Algerian basins. Based on the
available paleontological data it is here hypothes-
ized that the transcurrent belt located to the north
of the Sardinia-Corsica microplate and the
Calabrian block led to the formation of a neritic sea,
occasionally connecting for short periods the
Ligurian-Provencal Sea to the Po Basin.
Second phase - Further translation towards SE,
with a rotation of about 15° of the Calabrian block
(Gueguen, 1995), led to the opening of the Tyrrhe-
nian Basin, in the Late Miocene. As a result of these
changes the Mediterranean began to take on an
appearance more similar to the current one.
Doglioni et al. (1998) has affirmed that the Apen-
nine orogenetic front kept on migrating towards E
leading to the emersion of the Apennine Chain, thus
separating the Tyrrhenian Sea from the Adriatic after
the Burdigalian. On the other hand, based on the
opinion by Rosembaum et al. (2002) and on the
available macro-paleontological data (Stara &
Rizzo, 2013; 2014), it seems likely that the connec-
tion between the Tyrrhenian and the Adriatic basins
had been realized in the Plio-Pleistocene when the
Calabrian block reached the Apennine Arc, with the
exception of the Ligurian Channel (the Val Bormida
Channel of Stara & Rizzo, 2013). The crustal
thinning of the back-arc areas located W to the
Sardinia-Corsica microplate led to extensive flows
of basaltic lava (indicated by magnetic field anom-
alies) in the Balearic and the Tyrrhenian basins; both
of them were deep water seas with a maximum
depth of 3000 and 3700 m, respectively.
Kotsakis et al. (2004) prospected the occurrence
of a Sardinia-Tuscany bio-paleoprovince during the
Serravallian, on the basis of the close similarity of
the vertebrate fauna present in these areas. This
would imply the existence of landmass bridges or
shallow water basins separating lands, important
factors conditioning the diffusion of Amphiope
between the two sides of the Tyrrhenian Basin.
MATERIAL AND METHODS
The studied material consists of 78 Amphiope
specimens, preserved as whole coronas deprived of
the spines, from 5 Oligo-Miocene localities of
Western-Mediterranean Basin. 44 A. lovisatoi
Cotteau, 1895 (inventory code: (PL1301-03,
PL1317, PL1413, PL1418- 20, PE1422-24, PL1427,
PL1429, PL1567-70, PL1572-80, PL1583, PL1585-
Distribution of two Amphiope (Echinoidea Clypea steroid a) morphotypes in the Western-Proto-Mediterranean Sea 395
87, PL1692-99, PL1700-07, PL1709-14, PL1715-
18, PL 1720- 23, PL 1726) from Chiaramonti
(Sassari province); 1 Amphiope sp. from Capo
Frasca (Medio Campidano Province); 19 A. nur-
agica from Cuccuru Tuvullao (Cagliari Province)
MAC (PL1590-91, PL1678-80, PL1684, PL1727,
PL 1820, PL 1829; PL 1835-44); 1 A. montezemoloi
Lovisato 1901 from Bonnanaro, Sassari; 5 Am-
phiope sp. from Calabria (Vibo Valentia Province,
Italy), no code; 2 from Alicante (unknown locality);
1 from Torrent, Valencia Province (Spain) are
housed at the Museo di Storia Naturale “Aquilegia”
(MAC code) of Cagliari; the Holotype of A. nur-
agica, at the University of Cagliari Sardinia (Italy),
UNICA code, inventory 9CC.8- 10504. 10 Am-
phiope specimens from Calabria (Vibo Valentia
Province, Italy) number 104/E 101-110, are housed
at the Civico Museo Paleontologico di Ricadi (Vibo
Valentia province), Calabria, Italy. 3 specimens
from Torrent, Valencia Province (Spain) were stud-
ied in private collections. One sintype of A.
bioculata from Sure-Pres-Bollene today “Suze-la-
Rousse” near “Bollene”, France, housed at the
Museum d’Histoire Naturelle of Bordeaux (code
MFINB), France, inventory number MFINBx
2014.6.317.
The plate pattern of the sintype of A. bioculata ,
variety A of Des Moulins (1835), is not visible, but
the two lunules are clearly rounded, as described by
the author (“foraminibus subrotundis
Morphological abbreviations (Fig. 1) TL = test
length; TW = test width; TH = test height; PL =
petalodium length; LI = lunule length; L2 = lunule
width. The measure of TL is reported in mm; other
data in % TL; SI= lunule shape index (L2/L1);WI=
lunule width index (Ll+L2)/2.
Biometric analyses were carried out and data
analyzed using the software PAST-version 3.2
(2014) (Hammer, 2014), to help the interpretation
of the samples collected from Sardinia and
Calabria. Systematic palaeontology follows Kroh
& Smith (2010).
TWO MAIN AMPHIOPE MORPHOTYPES
The genus Amphiope sensu Stara & Sanciu
(2014) developed from the end of the Oligocene to
the late Tortonian- early Messinian in the Proto-
Westem-Mediterranean Sea.
A. nuragica (Comaschi Caria, 1955), from the
Oligo-Miocene of Sardinia, is the more ancient
species so far known belonging to this genus (Stara
& Borghi, 2014), though the genus Amphiope looked
like already well differentiated in the Aquitanian.
Based on the phylogenetic hypothesis proposed
by Stara & Borghi (2014) most forms with narrow
transversely elongate lunules derived from A. nur-
agica , whereas those with sub-rounded to broad
elliptical lunules as A. bioculata (including “A.
TW
Figure 1. Set of morphometric measurements used in this work.
396
Paolo Stara et alii
bioculatcT from the Aquitanian of Carry, France, as
inteipreted by Cottreau, 1914 and Philippe, 1998)
originated from a different common ancestor.
Two main morphotypes of the genus Amphiope
are here proposed: the “ nuragica ” group (PL 1 Figs.
3-4), characterized by narrow transversely elongate
lunules with SI > 1.6, and the “ bioculata ” group (PL
1 Figs. 1-2), with roundish to broad ovoid lunules
and SKI .59. Both these groups are well represented
in the study area, however in this paper we’ll go deep
into the “ nuragica ” forms only, since clear structural
data are so far available only for this group.
DISTRIBUTION OF THE TWO MAIN
MORPHOTYPES IN THE WESTERN ME-
DITERRANEAN
Amphiope has been recorded from more than 30
localities both in Sardinia (Comaschi Caria, 1955,
1972; Stara et al, 2012) and in the Rhone Basin
(France) (Cottreau, 1914; Philippe, 1998). This ech-
inoid has been cited in Italy also in Tuscany (Gian-
nini, 1957; present paper), Campania (Barbera &
Tavernier, 1989), Calabria (Cottreau, 1914; Carone
& Domning, 2007), Sicily (Garilli et al., 2010). In
Spain Amphiope was recorded in Catalogna (Lam-
bert, 1928), Mallorca (Llompart, 1983), Valencia
(personal communication of Bajo Campos, July
2012) and Alicante (present paper). Amphiope has
also been recorded from Algeria (Pomel, 1887-88
and Cotteau et al., 1891) and Corse (Cotteau, 1877).
The finding localities corresponding to these
records are reported in Table 1 and figure 2, with
the attribution to the “ nuragica' ’ or to the “biocu-
lata” group. The asterisc marks the species not
directly examined by the authors.
The syntype of A. bioculata from the Bollene
area has the SI = 1; the sample of A. bioculata
described by Cottreau (1914) has a mean value of
PL = 53 and SI ranges from 0.95 to 1.47, with a
mean of 1.22. The studied sample of A. nuragica
from Sardinia has a similar mean value of PL (51),
however SI ranges from 2 to 3 with a much higher
mean value (2.4) than that of A. bioculata sensu
Cottreau (1914).
Based on its small sized petalodium (PL = 40-
47, with a mean value of 43.3) the sample from the
Tortonian of Calabria clearly differs from all the
others belonging to the “ nuragica ” group (Fig. 3),
with the exception of the few specimens from the
Tortonian of Valencia (mean PL = 44).
In the sample from Calabria SI ranges from 2.4
to 4.3, with a mean value of 3.
A specimen from Sicily shows the lowest value
for the “ nuragica ” group (SI = 1.6), whereas a
specimen of A. hollandei Cotteau, 1877(the holo-
type) from Corse has the highest value (SI = 6.5).
Plate 1. Main Amphiope morphotypes. Figure 1-2, “ bioculata ” group; Figure 1 : A. bioculata syntype MHNBx 2014.6.317;
Figure 2: A. montezemoloi MAC.PL1676. Figure 3-4, “ nuragica ” group; Figure 3: A. nuragica MAC.PL. 1680; Figure 4.
Amphiope sp. from Calabria, MAC.PL 1672.
Distribution of two Amphiope (Echinoidea Clypea steroid a) morphotypes in the Western-Proto-Mediterranean Sea 397
TYPE
PL % TL
LI % TL
L2 % TL
WI
SI
Amphiope bioculata ,
sintype MHNB20I4.6
10
10
10
1
A cf. bioculata (in
Cottreau. 1914)*
53
10.2 (9-12.6)
1.22 (0.95-1.47)
A montezemoloi, San
Giorgio
-
16.8
16
16.4
0.95
A w/rogie^holotype
51
13.5 (11.5-15)
2.4 (2-3)
A. sp., Sicilia
56
9
15
12
1.6
A deydieri, France*
? 47
5.5
15.5
10.5
2.8
A. sp. 1, Calabria
44
5
13
9
2.6
41
5.5
14
9.7
2.5
45
5.5
18
11.7
3.3
44
6
17
11.5
2.8
47
5
16
11.5
3.2
A sp. 2, Calabria
44
4
14.5
9.2
3.6
43
4.5
14
9.2
3.1
40
5
12
8.5
2.4
42
3.5
15
9.2
4.3
A sp. 1, Valencia
42
5
13
9
2.6
44
4.5
11
7.7
2.4
A depressa, Algeria*
44
5
10
7.5
2
A sp., Capo Frasca
54
7.5
16
11.7
2.1
A. sp. 2, Valencia
55
5
14
9.5
2.8
A. sp., Alicante
52
4
17
10.5
4.5
A hollandei , Corsica*
54
3
20
11.5
6.6
A sarasini , France*
50
6
13
9.5
2.1
A. palpebrata, Algeria*
52
6.5
14
10.2
2.1
Table 1. Data PL, LI, L2, WI and SI of species included in the “ nuragica " group, present in the area under study.
For comparison, in the first three rows are reported the data of syntype of A. bioculata and two other forms of the group.
398
Paolo Stara et alii
Figure 2. Distribution of two main moiphotypes in the Western Mediterranean basin.
Red dot = “ bioculata ” group; blue dot = “ nuragica ” group; square dot = insufficient data.
However the last two are border-line cases rep-
resented by single specimens; additionally the spe-
cimen from Corse is poorly preserved (fide Cottreau,
1914) and the drawing may be not reliable.
The shape index of lunules is not discriminant
between the forms of the “ nuragiccT group, how-
ever comparison based on the values of LI and L2
is more significant. In the large sample from
Sardinia (Stara & Borghi, 2014) the range of LI is
6-16.7 (6-11.2 in A. nuragica), that of L2 is 11-23.5
(14.1-23.5 in A. nuragica). The samples from
Calabria and Valencia show much lower values of
the lunule length (LI ranges from 3.5 to a maxi-
mum of 6), L2 ranges from 11 to 18.
Based on these observations the sample from
Calabria is characterized by:
1 . the smallest petalodium so far known for the
genus Amphiope; only some specimens from Valen-
cia and A. depressa Pomel, 1887, from Algeria
show similar values of PL (see Fig. 3)
2. lower values of L2, WI and above all LI
when compared to the species described from
Sardinia by Stara & Borghi (2014): Amphiope
lovisatoi and A. nuragica (Fig. 4). A larger sample
is needed to confirm the same results also for the
examined specimens from Valencia.
CONCLUSIONS
This preliminary study indicates clear morpho-
logical differences within the “ nuragica ” group,
with respect to the size of the petalodium (PL) and
of the lunules. In particular the samples from the
late Miocene of Calabria (Southern Italy) and
Valencia (Spain) show much smaller petalodium
and smaller lunules when compared to the other
known species belonging to this group.
It is presumable that in some nearshore areas of
the Mediterranean (e.g. along the coasts of Calabria,
Balearic, Kabilyes, Sardinia, Corsica), separated by
deep water, different species of Amphiope de-
veloped independently, adapting to the environ-
mental changes occurred through the Miocene,
mainly climate, due also to the latitudinal migration
of lands, and ecology.
On the other hand, during the Miocene
Amphiope showed also structural modifications
Distribution of two Amphiope (Echinoidea Clypea steroid a) morphotypes in the Western-Proto-Mediterranean Sea 399
Figure 3. Petalodium comparison data on some species be-
longing to the “ nuragiccT group (PL in% TL). A = A. lovi-
satoi; B = A. nuragica; C = Amphiope sp. Calabria.
l? s 5-
1 M-
ii.&-
iaa-
7 . 5 '
M-
2 . 5 -
Figure 4. Size of lunules (WI) comparison on some species
belonging to the “ nuragica ” group. A = A. lovisatov, B = A.
nuragica ; C = Amphiope sp. Calabria.
common to almost all of the Mediterranean forms,
such as the decreasing number of plates and a
progressive lightening of the test- structure (Stara
& Borghi, 2014; Stara & Sanciu, 2014).
Future studies focusing on paleoecology will be
probably able to explain these dynamics and in
particular why some populations underwent signi-
ficant morphological modifications (e.g. the de-
creasing in the petalodium size seen in the samples
from Calabria and Valencia).
The results of this study indicate the occurrence
of different species in the “ nuragica ” group, that is
Amphiope characterized by narrow and transversely
elongate lunules. All these forms likely derived from
a common ancestor living in the Archipelago formed
during the Oligocene-Miocene boundary between
the Provencal and the Sardinian-Corsica coasts.
A similar differentiation is expected to be also
in the “ bioculata ” group and also this argument
will be the object of future studies.
AKNOWLEDGEMENTS
We thank Ildefonso Bajo Campos (Museo de
Alcala de Guadaira, Seccion de Paleontologia,
Sevilla, Spain), Fabio Ciappelli (Calenzano) and
Piero Frediani (Castelfiorentino, Firenze), for in-
formation on specimens from Tortonian of Valencia
(Spain) and Tuscany respectively; Paolo Cutuli and
the other people associated to the Gruppo Paleon-
tologico Tropeano (Vibo Valentia, Calabria), for
information about the A mph /opc-beari ng localities
in the Vibo Valentia Province. We warmly thank
Roberto Rizzo (Parco Geominerario, Storico e
Ambientale della Sardegna) for critical reading of
the geologic-structural evolution of the Western
Mediterranean. We also warmly thank Charles
Laurent (Museum d’Histoire Naturelle de Bor-
deaux, France) for providinge us information and
photos of the syntypes of A. bioculata.
REFERENCES
Barbera C. & Tavernier A., 1989. II Miocene del circond-
ario di Baselice (Benevento), significato paleoecolo-
gico e paleogeografico. In Di Geronimo I. (Ed.): Atti
del 3° Simposio di ecologia e paleoecologia
delle comunita bentoniche, Catania - Taormina, 12-
lb Ottobre 1985: pp. 745-772.
Carminati E., Lustrino M. & Doglioni C., 2012. Geody-
namic evolution of the central western Mediterranean
Tectonics vs. Igneus petrology constraints. Tectono-
physics (2012). Elsevier B.V., 20 pp.
Carone G. & Domning P.D., 2007. Metaxytherium ser-
resii (Mammalia: Sirenia): new pre-Pliocene record,
and implications for Mediterranean paleoecology be-
fore and after the Messinian salinity crisis. Bollettino
della Societa Paleontologica Italiana, 46: 55-92.
Comaschi Caria I., 1955. 11 sottogenere Amphiope in
Sardegna. Bollettino della Societa Geologica Italiana,
74: 183-194.
400
Paolo Stara et alii
Comaschi Caria I., 1972. Gli echinidi del Miocene della
Sardegna. Stabilimento Tipografico Ed. Fossataro,
Cagliari, 96 pp.
Cotteau G., 1877. Description des Echinides. In Locard
A. (Ed.): Description des Faunes des terrains Terti-
aires moyen de la Corse. Annales de la Societe
d’Agriculture, Histoire Naturelle et arts utiles de
Lyon, Paris-Geneve, pp. 227-335.
Cotteau G.., Peron P & Gauthier V., 1876-1891. Echin-
ides fossiles de EAlgerie. Etage Miocene et Pliocene,
Paris, 10, 1891, 273 pp.
Cottreau J., 1914. Les echinides neogenes du Bassin
mediterraneen. Annales de l’lnstitut Oceanograph-
ique, Monaco, 6: 1-193.
Des Moulins C., 1837. Troisieme Memoire sur les echin-
ides. Synonymie general. Actes Societe Linneenne,
Bordeaux, 9: 45-364.
Doglioni C., Gueguen E., Harabaglia P. & Mongelli F.,
1998. On the origin of the W-directed subduction
zones and applications to the Western Mediterranean.
In: Durand B., Jolivet L., Horvath F. & Seranne M.
(Eds.), The Mediterranean Basins: Tertiary Extension
within Alpine Orogen. Geological Society London
Special Publications.
Garilli V., Borghi E., Galletti L. & Pollina F., 2010. First
record of the Oligo-Miocene sand dollar Amphiope
Agassiz, 1840 (Echinoidea: Astriclypeidae) from the
Miocene of Sicily. Bollettino della Societa Paleonto-
logica Italiana, 49: 89-96.
Gattacceca J., Deino A., Rizzo R., Jones D.S., Henry B.,
Beaudoin F. & Vadeboin F., 2007. Miocene rotation
of Sardinia: new paleomagnetic and geochronolo-
gical constraints and geodynamic implication. Earth
and Planetary Science Letters, 258: 359-377.
Giannini E., 1957. I fossili delEarenaria di Manciano
(Grosseto). Paleontografia Italica, 51 pp.
Gueguen E., 1995. Le bassin Liguro-Provengal, un
veritable ocean. Exemple de segmentation des marges
et des hiatus cinematiques. Implications sur les
processus d'amincissement crustal. Ph.D Thesis,
Brest University, 315 pp.
Hammer 0., 2014. PAST 3.2, Paleontological Statistical
Software Package for Educational and Data Ana-
lysis. Natural History Museum, University of Oslo
[phammer(at)nhm.uio.no]
Kotsakis T., Delfino M. & Piras P., 2004. Italian Ceno-
zoic crocodilians: taxa, timing and paleobiogeo-
graphic implications. Paleogeography, Paleoclimato-
logy, Paleoecology, 210: 67-87.
Kroh A. & Smith A.B., 2010. The phylogeny and classi-
fication of post-Palaeozoic echinoids. Journal of
Sy- stematic Palaeontology, 8: 147-212.
Lambert J., 1928. Revision des Echinides fossiles
de Catalogne. Museo de Ciencias Naturales de Bar-
celona, Memorias Serie Geologia, 1: 1-62.
Llompart C., 1983. Amphiope bioculata (Desm.) del
Mioceno de Port de Mao (Menorca). Boletin de la
Real Sociedad Espanola de Historia Natural, Seccion
geologica, 81: 67-79.
Philippe M., 1998. Les echinides miocenes du Bassin du
Rhone: revision systematique. Nouvelles Archives
du Museum d’Histoire Naturelle de Lyon, 36: 3-241,
249-441.
Pomel A., 1887-1888. Paleontologie ou Description des
animaux fossiles de EAlgerie. 2, Zoophites, Echino-
dermes. A E Explication de la Carte Geologique de
EAlgerie. Alger 1887-1888, 344 pp.
Popescu S., 2009. Continental and marine environmental
changes in Europe induced by global climate
variability and regional paleogeography changes.
Memoire presente an vue de Eobtention du Diplome
d’Habilitation a Diriger des Recherches. Speciality
Sciences de la Terre e de EUnivers. Universite
Claude Bernard Lyon 1. web publication:
http://hal.archives-ouvertes.fr/docs/00/35/01/16/
PDF/Popescu_HDR.pdf
Rosenbaum G., Lister G. S. & Duboz C., 2002. Recon-
struction of the tectonic evolution of the Western
Mediterranean since the Oligocene In: Reconstruc-
tion of the evolution of the Alpine-Himalayan
Orogen. Rosenbaum G. & Lister G. S. (Eds.) 2002.
Journal of the Virtual Explorer, World Wide Web
electronic publication (http://virtualexplorer.com.au),
8: 107-130.
Stara P. & Borghi E., 2014. The echinoid genus
Amphiope L. Agassiz, 1840 (Astriclypeidae) in the
Oligo-Miocene of Sardinia (Italy). In: Paolo Stara
(Ed.). Studies on some astriclypeids (Echinoidea,
Clypeasteroida), pp. 225-358. Biodiversity Journal,
5:245-268.
Stara P., Rizzo R., Sanciu L. & Fois D., 2012. Note di
geologia e paleoecologia relative ad alcuni siti ad
Amphiope (Echinoidea: Clypeasteroida) in Sardegna.
Parva Naturalia, 9: 121-171.
Stara P. & Rizzo R., 2013. Diffusion of Amphiope
Agassiz, 1840 (Astriclypeidae, Clypeasteroida) from
the Western Proto-Mediterranean Sea, towards the
Ea- stern Neotethys. XIII Giornate di Paleontologia.
Pe- rugia, May 23-25, 2013, Volume dei riassunti,
pp. 119-120, sessione poster.
Stara P. & Rizzo R., 2014. Paleogeography and diffusion
of astriclypeids from Proto-Mediterranean basins. In:
Paolo Stara (Ed.). Studies on some astriclypeids
(Echinoidea, Clypeasteroida), pp. 225-358. Biod-
iversity Journal, 5: 233-244.
Stara P. & Sanciu L., 2014. Analysis of some astriclyp-
eids echinoids (Echinoidea, Clypeasteroida). In:
Paolo Stara (Ed.). Studies on some astriclypeids
(Echinoidea, Clypeasteroida), pp. 225-358. Biod-
iversity Journal, 5: 291-358.
Biodiversity Journal, 2015, 6 (1): 401-411
The genus Erctella Monterosato, 1 894: new molecular evid-
ence (Pulmonata Stylommatophora Helicidae)
Maria Stella Colomba 1 *, Armando Gregorini 1 , Fabio Liberto 2 , Agatino Reitano 3 , Salvatore Giglio 4 & Ignazio
Sparacio 5
'Universita di Urbino, Dipartimento di Scienze Biomolecolari (DiSB), via Maggetti 22, loc. Sasso, 61029 Urbino, Pesaro-Urbino, Italy;
e-mail: mariastella.colomba@uniurb.it, armando.gregorini@uniurb.it
2 Strada Provinciale Cefalu-Gibilmanna n° 93, 90015 Cefalu, Palermo, Italy; email: fabioliberto@yahoo.it
3 Via Gravina 77, 95030 Tremestieri Etneo, Catania, Italy; email: tinohawk@yahoo.it
4 Contrada Settefrati, 90015 Cefalu, Palermo, Italy; email: hallucigenia@tiscali.it
5 Via E. Notarbartolo 54 int. 13, 90145 Palermo, Italy; email: isparacio@inwind.it
’Corresponding author, email: mariasteha.colomba@uniurb.it
ABSTRACT In this paper we report on new molecular data (COI sequences) of different and represent-
ative populations of Erctella mazzullii (De Cristofori et Jan, 1832), E. cephalaeditana Gian-
nuzzi-Savelli, Oliva et Sparacio, 2012 and E. insolida (Monterosato, 1892) (Pulmonata,
Stylommatophora, Helicidae). Present results are compared with those from recent literature
and the current knowledge on phylogenetic relationships among Helicidae pulmonate
gastropods is reviewed. Obtained results suggest that: i) Cornu Born, 1778 and Cantareus
Risso, 1826 are separate and well distinct from Helix Linnaeus, 1758; ii) Erctella
Monterosato, 1894 is a valid and independent genus rather than a subgenus of Cornu; iii)
Cornu aspersion (O.F. Muller, 1774) is a group of species (i.e. " aspersum" group) whose
taxonomic status needs to be defin further studies; iv) Cornu , Cantareus and Erctella
might belong to the same tribe that, still, remains to be defined.
KEY WORDS Erctella; Helicidae; mitochondrial markers; phylogenetic reconstruction.
Received 11.02.2015; accepted 18.03.2015; printed 30.03.2015
Proceedings of the 2nd International Congress “Speciation and Taxonomy”, May 1 6th- 1 8th 2014, Cefalu-Castelbuono (Italy)
INTRODUCTION
Colomba et al. (2011) reported on a multidiscip-
linary study based on genital morphology, DNA
analysis, distribution, ecology and fossil records of
Cornu mazzullii (De Cristofori et Jan, 1832), a
species endemic to North-Western Sicily. Obtained
results supported the hypothesis that C mazzul-
lii should be attributed to the genus Erctella
Monterosato, 1894 and that this genus was probably
structured in three discrete clades (i.e., the mazzullii
group) recognized as species including: (i) the
populations living in Monte Pellegrino (Palermo)
and nearby mountains, E. mazzullii s. str., (ii) the
endemic population of Cefalu, La Rocca, E. ceph-
alaeditana Giannuzzi-Savelli, Oliva et Sparacio,
2012, and (iii) the populations living in the
mountains of Trapani surroundings, E. insolida
(Monterosato, 1892).
Based on the phylogenetic reconstruction
obtained by the multigenic analysis of nuclear
(ITS2) and mitochondrial (16S rDNA, 12S rDNA)
molecular markers, Colomba et al. (2011) strongly
suggested that the genus Erctella should be kept
distinct from the closely related genera Cornu
Born, 1778 and Cantareus Risso, 1826. In the
402
M.S. COLOMBA ET ALII
same paper, this hypothesis was also corroborated
by the analysis of several 16S rDNA partial
sequences downloaded from GenBank for other
genera representatives of Western Palaearctic
Helicidae taxa; noteworthy, the phylogenetic tree
topology clearly showed Cornu and Cantareus
distinct from Helix Linnaeus, 1758 (see Colomba
et al., 20 11, fig. 42).
Cornu Bom, 1778 (type species: Cornu copiae
Bom, 1778) was reintroduced as distinct genus by
Walden (1976) with Cryptomphalus De Charpen-
tier, 1837 (type species: Cryptomphalus aspersum
O.F. Muller, 1774) as junior synonim; it was some-
times considered as subgenus of Helix Linnaeus,
1758 (type species: Helix pomatia Linnaeus, 1758)
and sometimes as a distinct genus. The description
of Cornu copiae was based on a teratological
specimen of 'Helix” aspersa; due to different
interpretations of the Article 1.3.2 of the Code, a
request for conservation of the name Cornu is still
pending a mling of the International Commission
on Zoological Nomenclature.
Cantareus Risso, 1826 (type species: Cantareus
apertus Born, 1778) was sometimes considered as
subgenus of Helix and sometimes as a distinct
genus.
Schileyko (1978) was the first one who de-
scribed the internal structure of male sexual organs
of "Helix" aspersa characterized by a penial papilla
and a prominent semicircular fold in the distal part
of the penis (see also Nordsieck, 2013). Because of
these anatomical differences, the Author attributed
this species to the genus Cryptomphalus.
Giusti et al. (1995) showed a close similarity
between genitalia of "Helix" aperta and "Helix"
aspersa and, therefore, attributed these two
species to the same genus, Cantareus , morpholo-
gically well distinct from Helix. Moreover, they
reported that Helix has a real penial papilla inside
the penis and, distally, an accessory penial papilla,
whereas Cantareus shows a system of a real penial
papilla, a false penial papilla and, distally, an
"annular pad".
Neubert & Bank (2006) mainly confirmed these
morphological differences and concluded in con-
sidering Cornu and Cantareus as related but distinct
genera. One year later, similar observations were
reported by Alonso & Ibanez (2007).
At the same time, findings of scientific studies
based on molecular data were in line with the taxo-
nomic frame showing Helix distinct from Canta-
reus and Cornu , the latter two considered the same
genus (Manganelli et al., 2005; Koene & Schu-
lenburg, 2005; Wade et al., 2006, 2007).
Nevertheless, despite all these anatomical and
molecular evidence, recently Welter-Schultes et al.
(2011) and Welter-Schultes & Audibert (2012) con-
sidered Cornu and Cantareus to belong to the genus
Helix. Bank (2012) argued that such a systematic
position is wrong, and, above all, it does not take
into account a number of studies (cited above)
suggesting a taxonomic choice closer to the real af-
finities among these taxa. Welter-Schultes et al.
(2012), however, reaffirmed their beliefs and,
besides, Welter-Schultes (2012) reported Erctella
as synonym of Helix.
Nordsieck (2013), reviewing the papers, pub-
lished in the last decades, dealing with anatomical
and molecular data, concluded, in summary, that:
“ According to genital morphology and DNA ana-
lysis, ‘‘Helix “ aspersa and relatives are not more
related to Helix than Eobania and other genera of
the Helicinae [. . .] These species must therefore be
generically separated from Helix. The shell and the
genital differences, especially those of the penis
(Giusti et al. 1995, Neubert & Bank 2006,
Colomba et al. 2011), are sufficient for the generic
separation of Cantareus and Cornu (or Cryptom-
phalus, if the name Cornu is not valid because of
Art. 1.3.2 ICZN, cf. Giusti et al. 1995: 491).
Erctella is regarded as a subgenus of Comu instead
of a genus, because it is more closely related to
Comu than to Cantareus”.
More recently, detailed molecular genetics
studies (Korabek et al., 2014; 2015; Razkin et al.,
2015) confirmed Cornu and Cantareus as two
distinct genera forming a group with no sign of a
close relationship with Helix. In addition, Erctella
DNA sequences, when included in such analysis
(see Korabek et al., 2015), confirmed this item, in
line with Colomba et al. (2011).
At present there seems to be broad agreement in
considering Cornu and Cantareus distinct genera,
while on the position of Erctella opinions are still
diverging. In order to be able to further test the
“genus hypothesis” ( Erctella as a distinct genus,
Colomba et al., 2011) versus the “subgenus hypo-
thesis” ( Erctella as a Cornu subgenus, Nordsieck,
2013), we performed an additional molecular ana-
lysis to characterize and define even better, from a
The genus Erctella Monterosato, 1 894: new molecular evidence (Pulmonata Stylommatophora Helicidae)
403
molecular standpoint, the identity and reliability of
Erctella.
In particular, phylogenetic relationships among
taxa under study were analysed by comparing par-
tial sequences of the gene encoding for the cyto-
chrome oxidase subunit I (COI) - which is one of
the most commonly used mitochondrial markers in
molecular evolution and molecular phylogeny.
Besides, to provide a little contribute in sheding
some more light on Helicidae systematics, the ana-
lysis was extended to hundreds of specimens of the
family Helicidae whose COI sequences were down-
loaded from GenBank database. A similar analysis
was carried out including 16S rDNA partial
sequences of the same taxa. Molecular analyses
have been performed either with single (16S or
COI) or combined (16S+COI) molecular datasets.
MATERIAL AND METHODS
Specimens and Collection sites
For each population, 2-5 Sicilian Erctella spe-
cimens were analysed. Please note that each loc-
ality and/or collection site is named in the original
language (Italian). Collected samples were iden-
tified and [labelled] as follows: Erctella insolida
(from Trapani province: Custonaci, Trapani [CU],
M.te Cofano, Trapani [COF]; San Vito lo Capo:
cala Mancina, Trapani [SV]); Erctella mazzullii
(from W-Palermo surroundings: M.te Pellegrino
[MP]; Sferracavallo, Palermo [CMS]; Carini:
M.te Columbrina, Palermo [COL]; Cinisi: M.te
Pecoraro, Palermo [PEC]); Erctella cephalaed-
itana from Cefalu: la Rocca, Palermo [CM];
Cornu aspersum (= H. aspersa ) [CA] from Ce-
falu, Palermo, Sicily; and Cantareus apertus
[CAP] from Cefalu, Palermo, Sicily and Assoro,
Enna, Sicily.
DNA extraction, amplification and sequencing
Samples were stored separately at -20 °C in test
tubes. Of each individual, a piece of foot tissue was
used for total DNA extraction (by Wizard Genomic
DNA Purification Kit, Promega). COI fragments
(581-663 bp) were amplified using LCO_1490 (5’-
GGT C AACAAAT C ATAA AG ATATT GG-3 ’ ) andHCO_
2198 (5 ’-TAAACTTCAGGGTGACCAAAATCAG ’)
(Folmer et al., 1994). PCR cycles were as follows:
95 °C for 5 min; 95 °C for 1 min, 42°C for 1 min,
72°C for 1 min (35 cycles); 72°C for 5 min. To
remove primers and unincorporated nucleotides,
amplified products were purified with the Wizard
SV gel and PCR Clean-up kit (Promega). Sequen-
cing of purified PCR products was carried out using
automated DNA sequencers at Eurofins MWG
Operon (Germany). All COI sequences generated
in this study were uploaded in GenBank (accession
numbers: KR921883-KR921914).
Phylogenetic analyses
The analysis was conducted on two partial gene
sequences: COI and 16S rDNA, integrating our data
with those obtained from GenBank database. In
particular, in addition to the sequences obtained
from specimens tested directly in this study
(KR921883-KR921914), to further expand the ana-
lysis and refine its resolving power, we included
16S rDNA sequences of Erctella mazzullii , E. insol-
ida, E. cephalaeditana. Cornu aspersum and Can-
tareus apertus previously generated by our research
group (GQ402393-GQ402396, GQ402398-
GQ402402, GQ402403-GQ402405, GQ402407-
GQ402409, GQ402410-GQ402411, GQ402412-
GQ402414, GQ4024 1 7-GQ4024 1 9, GQ402420-
GQ402422, GQ402387-GQ402389, GQ402390-
GQ402392, see Colomba et al., 2011), joined to
both COI and 16S rDNA sequences downloaded
from GenBank of the following taxa: Eobania
vermiculata (O.F. Muller, 1774) (KJ458509,
KJ458510, KJ458511, JF277395, JF277393,
JF277391), Theba geminata Mousson, 1857
(KJ458559, HM034468), T. subdentata (Ferussac,
1821) (KJ458562, HM034496), T. pisana
(O.F. Muller, 1774) (KJ458561, JX911311), T. an-
dalusica Gittenberger et Ripken, 1987 (KJ458558,
KF582631), Murella muralis (O.F. Muller, 1774)
(GU391399, JX827154), Helix lucorum Linnaeus,
1758 (AF 126 144, GU784803), Helix pomatia Lin-
naeus, 1758 (AF208297, JX91 1304), Helix secern-
enda Rossmassler 1847 (KP072386, KP072387,
KP072388, KP072086, KP072087, KP072088),
Helix vladika Kobelt, 1898 (KP072303,
KF823104), Helix melanostoma Drapamaud 1801
(KJ458524, KP072 1 07), Iberus gualtierianus (Lin-
naeus, 1758) (AY928605,AY928606, DQ822123,
DQ822165, DQ822166, AY546285), Hemicycla
404
M.S. COLOMBA ET ALII
bidentalis (Lamarck, 1822) (KJ45 8528, HM 147 180),
Pseudotachea splendida (Draparnaud, 1801)
(KJ458552, AY546292), Levantina caesareana
(Mousson, 1854) (KP072332, KP072181) Otala
lactea (O.F. Muller, 1774) (AY937264, AY937263),
O. punctata (O.F. Muller, 1774) (JF717823,
JF717824, KJ458545, JF717805, JF717806,
JF717807), Helix aspersa (AF126139, AF126135,
AF 126 134, AF126140, AF126136, JN701926,
JN701927, GU598217, AY546283, HQ203051,
HQ203052, JX911287), Cantareus apertus
(KJ458491, JX911286). Finally, Limax maximus
Linnaeus, 1758 (Family Limacidae) (KF894386),
L. cinereoniger Wolf, 1803 (KF894380), Limacus
flavus (Linnaeus, 1758) (FJ896815), Muticaria
syracusana (Philippi, 1836) (Family Clausiliidae)
(HQ696868, AY425597) and M. neuteboomi
Beckmann, 1990 (FIQ696866, HQ696867) were
employed as outgroups.
All sequences were visualized with BioEdit
Sequence Alignment Editor 7 (Hall, 1999), aligned
with the ClustalW option included in this software
and refined by eye. As far as concerns single (COI
or 16S rDNA) molecular data sets, phylogenetic
analyses were conducted in MEGA 5 (Tamura et al.,
2011) by Maximum Likelihood algorithm. Substi-
tution models, selected according to the “Find Best
DNA model” option included in the software, were:
HKY+G (COI) and GTR+G (16S rDNA); support
for the intemodes was assessed by bootstrap
percentages (BP) (1000 replicates). For the com-
bined (COI+16S rDNA) datasets, phylogenetic ana-
lyses were conducted in BEAST 1.6.1 (Drummond
& Rambaut, 2007) using the *BEAST implement-
ation (Heled & Drummond, 2010). A series of
initial runs were performed to optimize priors and
runtime parameter choice to obtain effective
sampling sizes (ESS) above 500 for all estimated
parameters. The best-fit evolution models of
nucleotide substitution were: HKY+G (COI) and
GTR+G (16S rDNA) with empirical base com-
position; the Yule Process tree prior for mitochon-
drial data with piecewise linear population size
model was applied with a UPGMA-generated tree
as starting point. Trees from all mns were combined
to produce an ultrametric consensus tree using
TreeAnnotator 1.6.1. The first 10 3 trees were dis-
carded as bumin. Support for nodes was expressed
as posterior probabilities.
RESULTS AND DISCUSSION
COI and 16S rDNA consensus trees and the
multi-genic (COI+16S rDNA) tree included 69
molecular sequences, each. Obtained results al-
lowed to make a few observations of some interest.
In particular, COI consensus tree (Fig. 1), showed
three separate clusters for (A) Erctella (discussed
in detail below), (B) Cantareus apertus and (C)
Cornu aspersum clearly distinct. Similarly, (D)
Eobania vermiculata , (E) Levantina caesareana,
(F) Helix spp. (including several species), (G)
Otala spp. (O. punctata and O. lactea), (H)
Murella muralis, (I) Hemicycla bidentalis,
Pseudotachea splendida, Iberus gualtierianus and
(L) Theba spp. ( T. geminata, T. subdentata, T.
pisana, T. andalusica ) are separated. With regard
to Erctella, the three taxa are clearly distinct and
separated as E. insolida (SV1-SV3, CU4-CU5,
COF2-COF4, from Trapani province), E. mazzullii
(CMS1-CMS5, COL1-COL3, PEC1-PEC3, MP1-
MP3, comprising specimens sampled on M.te
Pellegrino and the nearby mountains of sur-
roundings of Palermo), and E. cephalaeditana
(CM1-CM4, from Cefalu, La Rocca).
The 16S rDNA consensus tree topology (Fig. 2)
is similar to that shown in figure 1 . In fact, also in
this case, Erctella is clearly distinct and well struc-
tured in three taxa, Erctella insolida , E. cephalaed-
itana and E. mazzullii. Once again, it is confirmed
a distinction between the (closely related) genera
Erctella, Cornu and Cantareus ; based on 16S
rDNA sequences analysis, Erctella appears closer
to Cornu, while in the COI tree Cornu is closer to
Cantareus.
Mean molecular distances among the three taxa
of Erctella (assessed by the maximun composite
likelihood method), range from nearly 6 to 10%
(16S rDNA) and about 4 to 7.5% (COI). These
values, despite the issues of using mean molecular
distances (see Meier et al., 2008), nevertheless,
compared with those reported for other species,
including Pulmonata (e.g. Hebert et al., 2003a,
2003b; Steinke et al., 2005; Nekola et al., 2009)
can, in our opinion, justify the separation of
Erctella populations into three species.
Genetic distances between different species
within various animal groups, especially inverteb-
rates, are variable (see for example Meier et al.,
2008 and references therein). This is because they
The genus Erctella Monterosato, 1 894: new molecular evidence (Pulmonata Stylommatophora Helicidae)
405
Figure 1. COI consensus tree. The evolutionary history was inferred by using the Maximum Likelihood method based on
HKY model. The percentage of replicate trees in which the associated taxa clustered together in the bootstrap test (1000 re-
plicates) are shown next to the branches. A discrete Gamma distribution was used to model evolutionary rate differences
among sites [5 categories (+G, parameter = 0.6175)].
406
M.S. COLOMBA ET ALII
Figure 2. 1 6S rDNA consensus tree. The evolutionary history was inferred by using the Maximum Likelihood method based
on GTR model. The percentage of replicate trees in which the associated taxa clustered together in the bootstrap test (1000
replicates) are shown next to the branches. A discrete Gamma distribution was used to model evolutionary rate differences
among sites [5 categories (+G, parameter = 0.7920)].
The genus Erctella Monterosato, 1 894: new molecular evidence (Pulmonata Stylommatophora Helicidae)
407
are function of different parameters among which:
different rates of nucleotide substitution, different
types of environmental pressure, or different types
of mutation which the nucleotide sequences are sub-
ject to, which at times (e.g. for retro-mutations)
cannot be detected a-posteriori. Moreover, generally
speaking, genetic distances per se are not sufficient
to discriminate between different species and, for
pulmonates, a few cases have been documented
where distances turned out to be misleading, neces-
sitating to be integrated with additional data (see
Davison et al., 2009; Sauer & Hausdorf, 2012).
In Erctella , molecular data combined with other
significant data such as morphological, biological,
ecological and paleontological features allow us to
consider it a genus with three different species,
endemic to northwestern Sicily (Liberto et al.,
2010; Colomba et al., 2011).
Concatenated-gene analysis was better resolved
than single-gene analysis and thus represents, prob-
ably, more accurately present relationships among
taxa. It resulted in a tree topology (Fig. 3) which is
quite superimposable to that of the ML trees (Figs.
1, 2) and, for the most part, in line with a recent
review of the molecular phylogeny of the Western
Palaearctic Helicoidea by Razkin et al. (2015). In
particular, it is visible the group including Eobania
vermiculata , Otala lactea and O. punctata (tribe
Otalini, in pink), the group including Iberus gual-
tierianus, Pseudotachea splendida and Hemicycla
bidentalis (Allognathini, in red), Theba species
(Thebini, in yellow), and several species of Helix
and Levantina caesareana (Helicini, in lilac).
In the concatenated-gene analysis, Erctella and
Cornu , considered two distinct genera, are sister
groups.
<3 0 (*
S3, US
Muticaria syracusana
Muticaria neuteboomi
Limacus flavus
Eobania vermiculata (n= 26)
Otala lactea (n=26)
Otala punctata (n=26)
Iberus gualtierianus (n=22)
Pseudotachea splendida (n=22)
Hemicycla bidentalis
Theba geminata
Theba pisana (n=30)
Theba andalusica
Theba subdentata
Can tareus apertus ( n =2 7)
Erctella mazzullii (n=27)
Erctella cephalaeditana
Erctella insolida
Cornu aspersum (n=27)
Murella muralis
Helix melanostoma (n=27)
Helix vladika
Helix lucorum
Helix pomatia (n=27)
Helix secernendo
Levantina caesareana
Otalini
Allognathini
Thebini
Helicini
Figure 3. Phylogenetic annotated tree based on Bayesian inference analysis of the concatenated data set including 16S rRNA
and COI sequences. Numbers correspond to BI posterior probabilities (in %).
408
M.S. COLOMBA ET ALII
Regarding relationships within the group
Cantareus-Erctella-Cornu our data differ from
Razkin et al. (2015). In fact, while for Erctella it is
not possible to make a comparison because the
Authors did not include this taxon in their analysis,
on the other hand, in our tree, neither Cornu nor
Cantareus can be considered Otalini, rather belong-
ing to a distinct cluster (attributable to the “tribe”
level) including Erctella.
Therefore, although Cornu and Cantareus show
a certain degree of affinity particularly with Eo-
bania for genitalia architecture (see Giusti et al.,
1995) and share with Otalini similar biogeographic,
ecological and evolutionary items typical of
Western Mediterranean areas where these terrestrial
molluscs differentiated (see Colomba et al., 2011),
nevertheless, the consideration of Cornu , Cantareus
and Erctella as a separate tribe, which still remains
to be defined, is suggested. Furthermore, Cornu ,
Cantareus and Erctella share the same chromosome
number (n = 27) (Vitturi et al., 1982; Vitturi et al.,
2005) (see Fig. 3), while Eobania and other Otalini
examined up to now have n = 26 (Burch, 1965;
Thiriot-Quievreux, 2003). Finally, Otalini show in
genital organs a relatively little dart sac and well-
developed digit-like appendiges, Cornu-Cantareus-
Erctella, instead, show a massive dart sac and two
groups of digitiform glands with short base and
numerous and short digit-like appendiges.
On the other hand, the separation between
Cornu-Cantareus-Erctella and Helix is supported
by: (i) the different geographical distribution of the
genera: Cornu and Cantareus are widespread in
North Africa and Southern Europe, with Erctella
endemic to Northwestern Sicily, while Helix is
mainly distributed in Central and Eastern Europe
and, to a lesser extent, North Africa; (ii) the dif-
ferent morphology of genital organs (Schileylco,
1978; Giusti et al.,1995; Neubert & Bank, 2006,
Alonso & Ibanez, 2007) showing in Cornu-Can-
tareus-Erctella a different form of dart sac and of
digitiform glands (see above); and (iii) molecular
data (see Korabek et al., 2015 and quotes therein).
Comparing the three phylogenetic trees an inter-
esting consideration about Cornu aspersum can be
made. In fact, in line with other studies (Guiller et
al., 2001; Guiller & Madec, 2010), in our study as
well, this taxon seems to be not a single species but
rather a species group (ie " aspersum " group)
showing a taxonomic situation more complex and
heterogeneous than previously hypothesized within
its area of origin and diversification (Southern
Italy, Sicily and NW Africa). This result is further
confirmed by personal unpublished morphological
and molecular data of numerous Italian, Maltese
and North African C. aspersum populations.
Finally, the position of Murella muralis remains
to be clarified. In fact, it is not only different in all
phylognetic trees but, above all, discordant with
what reported in other papers. This issue, which is
beyond the aim of the present paper, requires further
study and investigation, possibly increasing the
number of specimens (joining to sequences down-
loaded from the database also sequences obtained
from new samples collected directly in the field),
increasing the number of genes analyzed and, above
all, including in the analysis other taxa represent-
atives of subfamilies more closely related to Murel-
linae, such as Ariantinae.
Overall, present results correspond well to
several previous molecular studies carried out by
nuclear and mitochondrial markers (Koene &
Schulenburg, 2005; Colomba et al., 2011; Korabek
et al., 2014; Razkin et al., 2015) and confirm that
Erctella species lie always outside the clusters of
Cornu and Cantareus.
CONCLUSIONS
New molecular evidence provided in this study
suggested also several comments on Erctella
closely related genera. Hence, on this basis, despite
the difficulties that the argument implies, some con-
clusions can be drawn.
The groups comprising Cornu-Cantareus-
Erctella on one hand, and Helix on the other hand,
appear separate and distinct from each other. In line
with most of the papers reporting on anatomical and
molecular characteristics observed in these animals,
there seems to be no evidence that "aperta",
"aspersa" and / or "mazzullii" may belong to the
genus Helix.
Considering Cornu and Cantareus as Otalini, as
assumed by Razkin et al. (2015) is not confirmed
in our analysis. However, as mentioned above, the
issue certainly needs further study in view of their
aforementioned anatomical and biogeographical
affinities.
The genus Erctella Monterosato, 1 894: new molecular evidence (Pulmonata Stylommatophora Helicidae)
409
We suggest considering Cornu , Cantareus and
Erctella as related but distinct genera belonging to
independent lineages; as hypothesized, they might
be included into a new tribe (between Otalini and
Helicini).
Cornu aspersum complex is in need of a thor-
ough taxonomic revision in its area of origin.
Finally, it is appropriate to reiterate that our de-
cision to consider Erctella a distinct genus including
three different species (Colomba et al., 2011) was not
only made on the basis of some, although import-
ant, molecular evidence, but also by the analysis of
many other data that allowed us to assign to the
various Erctella populations morphological, biolo-
gical, paleontological and biogeographical peculiar
characters, amplified by the particular distribution
of the taxon, endemic to Northwestern Sicily. In this
regard it is worth remembering that in the charac-
terization of a taxon, at different levels, while
gathering as many informations as possible (in-
cluding morfological, ecological, molecular data,
etc ...) is necessary, taxonomic reconstmctions ob-
tained with a methodology not always correspond
to the ones obtained with another method (see
Schileyko, 2013); for Erctella, instead, all (numer-
ous) data are consistent with the hypothesis of dif-
ferentiating it from other (similar, closely related)
genera.
So that it seems appropriate to conclude with the
words reported by A. Schmidt (1868) who claimed
that, in taxonomy : “ Kilnstliche Systeme entstehen
durch consequentes Geltendmachen eines einzelnen
Princips ” [“the application of a single criterion
produces artificial classifications”].
In more contemporary terms, we could say with
Poins et al. (2014): “Molecular phylogenetics is an
irreplaceable tool for taxonomists, but interpreta-
tion of the results must be based on clear taxonomic
concepts corroborated by all available resources -
that is, the primary reference, the subsequent taxo-
nomic literature and the type specimens of the
organisms of interest. Otherwise, molecular phylo-
genetics can cause confusion with detrimental con-
sequences to follow-up studies (e.g. ecological and
evolutionary )”.
ACKNOWLEDGEMENTS
We are grateful to Eike Neubert, Research
Institute Senckenberg, Senckenberganlage, Frank-
furt am Main, Germany; Giuseppe Pocaterra, San
Pietro in Casale, Bologna, Italy; Fernando Scarlas-
sara, Melego di Sarego, Vicenza, Italy.
REFERENCES
Alonso M.R. & Ibanez M., 2007. Anatomy and function
of the penial twin papillae system of the Helicinae
(Gastropoda: Helicoidea: Helicidae) and description
of two new, small Hemicycla species from the laurel
forest of the Canary Islands. Zootaxa, 1482: 1-23.
Bank R.A., 2012. Comment on Cornu Born, 1778 (Mol-
lusca, Gastropoda, Pulmonata, HELICIDAE): re-
quest for a ruling on the availability of the generic
name (Case 3518). http://iczn.org/node/40276
Burch J.B., 1965. Chromosome numbers and systematics
in euthyneuran snails. Proceedings of the first
european malacological congress, London, 1962,
215-241.
Colomba M.S., Gregorini A., Liberto F., Reitano A.,
Giglio S. & Sparacio I., 2011. Monographic revision
of the endemic Helix mazzullii De Cristofori & Jan,
1832 complex from Sicily and re-introduction of the
genus Erctella Monterosato, 1894 (Pulmonata,
Stylommatophora, Helicidae). Zootaxa, 3134: 1-42.
Davison A., Blackie R.L.E. & Scothem G.P., 2009. DNA
barcoding of stylommatophoran land snails: a test of
existing sequences. Molecular Ecology Resources, 9:
1092-1101.
Drummond A. J. & Rambaut A., 2007. BEAST: Bayesian
evolutionary analysis by sampling trees. BMC Evol-
utionary Biology, 7 : 1214-1257.
Hall T.A., 1999. BioEdit: a user-friendly biological
sequence alignment editor and analysis program for
Windows 95/98/NT. Nucleic Acids Symposium
Series, 41: 95-98.
Hebert P.D.N., Cywinska A., Ball S.L. & deWaard J.R.,
2003a. Biological identifications through DNA bar-
codes. Proceedings of the Royal Society of London
B, 270: 313-321. DOI: 10.1098/rspb.2002.2218
Hebert P.D.N., Ratnasingham S. & deWaard J.R., 2003b.
Barcoding animal life: cytochrome c oxidase subunit
1 divergences among closely related species. Pro-
ceedings of the Royal Society of London B (Supple-
ment), 270: S96-S99. DOI 10. 1098/rsbl. 2003. 0025
Heled J. & Drummond A. J., 2010. Bayesian inference of
species trees from multilocus data. Molecular
Biology and Evolution, 27: 570-580.
Folmer O., Black M., Hoeh W., Lutz R. & Vrijenhoek R.,
1994. DNA primers for amplification of mitochon-
drial cytochrome c oxidase subunit I from diverse
metazoan invertebrates. Molecular Marine Biology
and Biotechnology, 3: 294-299.
410
M.S. COLOMBA ET ALII
Giusti F., Manganelli G. & Schembri P.J., 1995. The non-
marine molluscs of the Maltese Islands. Museo
Regionale di Scienze Naturali, Torino, Monografie,
15: 1-607.
Guiller A. & Madec L., 2010. Historical biogeography of
the land snail Cornu aspersum : a new scenario in-
ferred from haplotype ditribution in the Westen
Medierranean basin. BMC Evolutionay Biology, 10:
1 - 20 .
Guiller A., Coutellec-Vreto M.A., Madec L. & Deunff J.,
2001. Evolutionary history of the land snail Helix
aspersa in the Western Mediterranen: preliminary
results inferred from mitochondral DNA sequences.
Molecular Ecology, 10: 81-87.
Koene J.M. & Schulenburg H., 2005. Shooting darts:
co-evolution and counteradaption in hermaphroditic
snails. BMC Evolutionay Biology, 5: 25.
Korabek O., Jurickova L. & Petrusek A., 2014. Resur-
recting Helix straminea, a forgotten escargot with
trans- Adriatic distribution: first insights into the gen-
eric variation within the genus Helix (Gastropoda:
Pulmonata). Zoological Journal of the Linnean
Society, 171: 72-91.
Korabek O., Petrusek A., Neubert E. & Jurickova L.
2015. Molecular phylogeny of the genus Helix
(Pulmonata: Helicidae). Zoologica Scripta. DOI:
10. 1111/zsc. 12101
Liberto, F., Giglio, S., Reitano, A., Colomba, M.S. &
Sparacio I., 2010. Molluschi terrestri e dulciacquicoli
di Sicilia della collezione F. Mina Palumbo di Castel-
buono. Monografie Naturalistiche, 2. Edizioni
Danaus, Palermo, 134 pp.
Manganelli G., Salomone N. & Giusti F., 2005. A mo-
lecular approach to the phylogenetic relationships of
the western Palaearctic Helicoidea (Gastropda:
Stylommatophora). Biological Journal of the Linnean
Society, 85: 501-512.
Meier R., Zhang G. & Ali F., 2008. The Use of mean
Instead of Smallest Interspecific Distances Ex-
aggerates the Size of the “Barcoding Gap” and Leads
to Misidentification. Systematic Biology, 57: 809-
813.
Nekola J.C., Coles B.F. & Bergthorsson U., 2009.
Evolutionary Pattern and Process within the Vertigo
gouldii (Mollusca: Pulmonata, Pupillidae) group of
minute North American Land Snails. Molecular
Phylogenetics and Evolution, 53: 1010-1024.
DOI: 10.1 01 6/j. ympev.2009.09.012
Neubert E. & Bank R.A., 2006. Notes on the species of
Caucasotachea C. Boettger 1909 and Lindholmia P.
Hesse 1919, with annotations to the Helicidae
(Gastropoda: Stylommatophora: Helicidae). Archiv
fur Molluskenkunde, 135: 101-132, 6 pis.
Nordsieck H., 2013. The systematic position of Helix
aspersa Muller 1774. Available at.:
http :// www. hnords . de/5 3 5 6429 d6b 1 1 7 F602/5 3 5 642a
15712fcf01/index.html - last access: 10.11.2015
Poins N., Vardinoyannis K., Mylonas M. & Poulakakis
N., 2014. Evaluation of the taxonomy of Helix cincta
(Muller, 1774) and Helix nucula (Mousson, 1854);
insights using mitochondrial DNA sequence data.
Journal of Natural History, DOI: 10.1080/00222933.
2013.825023
Razkin O., Gomez-Moliner B.J., Prieto C.E., Martinez-
Orti A., Arrebola J. R., Munoz B., Chueca L.J.,
Madeira M.J., 2015. Molecular phylogeny of
the western Palaearctic Helicoidea (Gastropoda,
Stylommatophora). Molecular Phylogenetics and
Evolution, 83: 99-117.
Sauer J. & Hausdorf B., 2012. A comparison of DNA-
based methods for delimiting species in a Cretan land
snail radiation reveals shortcomings of exclusively
molecular taxonomy. Cladistics, 28: 300-316. DOI:
10. 1 1 1 1/j. 1096-003 1 .201 1 .00382.x
Schileyko A. A., 1978. Nazemnye molljusk inadse-
mejstva Helicoidea. In: Fauna SSSR, Molljuski, III,
Leningrad, Nauka, 6: 384 pp.
Schileyko A.A., 2013. Family Helicidae excluding
Helicinae (Gastropoda Pulmonata): morphology,
taxonomy, and a catalogue of taxa. Ruthenica, 23:
127-162.
Schmidt A., 1868. System der europaischenClausilien
und ihremachstenVerwandten. Th Fischer, Cassel, [1-
2], 176 pp.
Steinke D., Vences M., Salzburger W. & Meyer A., 2005.
Taxi: a software tool for DNA barcoding using
distance methods. Philosophical Transactions of the
Royal Society of London B, 360: 1975-1980. DOI:
10.1098/rstb.2005.1729
Tamura K., Peterson D., Peterson N., Stecher G., Nei M.
& Kumar S., 2011. MEGA5: molecular evolutionary
genetics analysis using Maximum Likelihood,
Evolutionary Distance, and Maximum Parsimony
methods. Molecular Biology and Evolution, 28:
2731-2739.
Thiriot-Quievreux C., 2003. Advances in chromosomal
studies of gastropod molluscs. Journal of Molluscan
Studies, 69: 187-201.
Vitturi R., Rasotto M.B. & Farinella-Ferruzza N.,
1982. The chromosomes of 16 molluscan species.
Bolletino di zoologia, 49: 1-2, 61-71.
DOE10. 1080/11250008209439373
Vitturi R, Libertini A., Sineo L., Sparacio I, Lannino A.,
Gregorini A. & Colomba M.S., 2005. Cytogenetics
of the land snails Cantareus aspersus and C. mazzul-
lii (Mollusca: Gastropoda: Pulmonata). Micron, 36:
351-357
Wade C.M., Mordan P. B. & Naggs F., 2006. Evolutionary
relationships among the Pulmonate land snails
and slugs (Pulmonata, Stylommatophora). Biological
Journal of the Linnean Society, 87: 593-610.
The genus Erctella Monterosato, 1 894: new molecular evidence (Pulmonata Stylommatophora Helicidae)
411
Wade C.M., Hudelot C., Davison A., Naggs F. & Mordan
P.B., 2007. Molecular phylogeny of the helicoid land
snails (Pulmonata: Stylommatophora: Helicoidea),
with special emphasis on the Camaenidae. Journal of
Molluscan Studies, 73: 411-415.
Walden H.W., 1976. A nomenclatural list of the land
Mollusca of the British Isles. Journal of Conchology,
29:21-25 .
Welter- Schultes F., Audibert C. & Bertrand A., 2011.
Liste des mollusques terrestres et dulcicoles de
France continentale (excl. hydrobioides). Folia
Conchyliologica, 12: 4-44.
Welter- Schultes F., 2012. European non-marine mol-
luscs, a guide for species identification. Planet Poster
Editions, Gottingen, 760 pp.
Welter-Schultes F., Altaba C.R. & Audibert C., 2012.
Comment on Cornu Born, 1778 (Mollusca,
Gastropoda, Pulmonata, HELICIDAE): request for
a ruling on the availability of the generic name
(Case 3518). Bulletin of Zoological Nomenclature,
70: 41-42.
Welter-Schultes F. & Audibert C., 2012. Comment on
Cornu Born, 1778 (Mollusca, Gastropoda, Pul-
monata, Helicidae): request for a ruling on the avail-
ability of the generic name. Bulletin of Zoological
Nomenclature, 69: 124-127.
412
Biodiversity Journal, 2015, 6 (1): 415-430
Monograph
Coen’s Pyramidellidae (Gastropoda Heterobranchia): a
revision of types
Pasquale Micali 1 *, Italo Nofroni 2 , Riccardo Giannuzzi Savelli 3 , Francesco Pusateri 4 & Stefano Bartolini 5
'Via Papiria 17, 61032 Fano, Pesaro-Urbino, Italy; e-mail: lino.micali@virgilio.it
2 Via B. Croce 97, 00142 Roma, Italy; e-mail: italo.nofroni@uniromal.it
3 Via Mater Dolorosa 54, 90146 Palermo, Italy; e-mail: malakos@tin.it
4 Via Caste liana 64, 901 35 P alerm o, Italy; e-mail: francesco@pusate ri.it
5 Via E. Zacconi 1 6, 50 1 37 Florence, Italy; e-mail: stefmaria.bartolini@libero.it
Corresponding author
ABSTRACT Coen introduced several new nominal taxa in the Pyramidellidae and in most Mollusca
families. The Coen types, now at the Hebrew University of Jerusalem, have been examined;
most of them are holotypes or lectotypes. Some lectotypes were already selected by van
Aartsen. as stated in the label, therefore we have not done any further selection of types. The
new pyramidellid species have been practically identified and named by Monterosato, and
were all found in shell grit collected on the beach of Lido (a small island in front of Venice).
None of the Coen’s new species seems to be valid.
KEY WORDS Coen collection; Pyramidellidae;Adriatic Sea; Mediterranean Sea.
Received 29.10.2014; accepted 20.12.2014; printed 30.03.20 1 5
Proceedings of the Eighth Malacological Pontine Meeting, October 4 th- 5th, 2014 - San Felice Circeo, Italy
INTRODUCTION
Giorgio Silvio Coen (1 873-1 95 1 ) was born in
Venice, graduated as a civil engineer in the presti-
gious University of Padua and spent his life in
Venice. The malacology was for him a hobby,
because his main work was as civil engineer. Any-
way he had contacts with most of the eminent ma-
lacologists of the period, wrote several works and
arranged a rich collection of molluscs. He was
a victim of the anti-Semitic laws during fascist
age and being banned from publishing in Italian
magazines, during the years 1 939-1 944 he pub-
lished on theActa PontificiaAcademia Scientiarum,
Civitate Vaticana. After the II World War he re-
turned in Venice, where he died. More informations
on Coen’s life are in Piani et al. (1 990).
The first Coen’s work is dated 1914, when he
was about 40 years old, but his name was already
present in the malacological world, because there
was a Turb onilla coeni Preston, 1 905 possibly
(because the Author did not indicate the origin of
the name) dedicated to him. When Coen was initi-
ating his malacological activity, Tommaso di M aria
(1 84 1 -1927), universally known as Monterosato,
was one of the best known Italian malacologists and
Coen used to send material to him for determina-
tion. Since the first publication, Coen uses names
with indication “Monterosato ms”, to indic-
ate the name, normally specific, was assigned by
M o n tero sa to .
Really the aged Monterosato was quite a “split-
ter”, with an attitude to create new species and
varieties, at the same time it is surprising that all the
416
Pasquale Micali et alii
new Pyramidellidae names, based on Monterosato’s
determ ination are synonyms of well known species.
We may suppose that the bad quality of Coen’s spe-
cimens examined by Monterosato, as well as
possible age-related problems of the Author are
cause of this. The names dictated by the aged
Monterosato, together with Coen’s attitude, some-
time referred as B ourguignat’s “Nouvelle Ecole”,
resulted in the creation of hundreds specific and
varieties names. Possibly Coen, interested to civil
architecture, was attracted by the details and loved
to collect, and name, the various forms of shells.
In some cases Coen understands that even if
Monterosato considered that shell as a variety, and
assigned a variety name, the differences are so
small that should not be the case. This is the case
of Pyrgulina b re vie u la v ar. rejecta M o n tero s a to ms.
for which the Author states “ Forma poco diversa
da lla tipica, cost nominata p ere he trovata nel
detrito [Form slightly different from the typical, so
named because found in shell grit]”. In the case of
Turbonilla ( Tragula ) fen e strata var. turbifacta
M onterosato ms. the A u thor states “ Q uesta fo rm a ,
dal Monterosato consider ata come specie,
turbifacta, mi par piuttosto varieta della fenestrata
[This form, considered by Monterosato a valid
species, turbifacta, looks to me a variety of fe ties -
trata ]”, practically he considers as a variety what
Monterosato considered a separate species. A final
consideration on the pyramidellid species: it is
strange that notwithstanding the Coen’s research in
the northern Adriatic, and the obvious contacts with
local fishermen, all the described species have been
found in the shell grit from Venice-Lido. Possibly
he was not much interested in micro molluscs and
all the species have been found in a sample of shell
grit given by to M onterosato.
Coen published more than sixty m alacological
works (Piani et al., 1 990), the most extensive
and interesting is the “Saggio di una Sylloge Mol-
luscorum A driaticorum ” published on 1933, a com-
mented list of north Adriatic molluscs, with descrip-
tion of several species and varieties. The work was
enriched and published again on 1 937, always as a
“Memoria” of the “Real Comitato Talassografico
Italiano” with the title “Nuovo Saggio di una
Sylloge Molluscorum A driaticorum ”.
The names created for varieties are to be con-
sidered subspecific, according to art. 45.6.4 of
IC ZN (4th. Edition) .
Coen’s collection was very rich, with material
obtained by various m alacologists (Mienis, 2012),
because he established a Museum in Venice, which
attracted the donations. Possibly due to the memory
of the anti-Semitic persecution in Italy, Coen
decided that after his death the collection had to be
donated to the Hebrew University of Jerusalem,
where it was shipped on 1953. Piani’s opin-
ion (Piani, 1 9 83 ) about the presence of many
Monterosato’s types in Coen collection is disputed
by M ien is (2012).
MATERIAL AND METHODS
The Coen’s collection is preserved at the
Hebrew University of Jerusalem and has been
loaned thanks to the courtesy of Dr. Henk K.
Mienis, Hebrew University of Jerusalem. Most of
the type specimens are worn and difficult to de-
termine without a previous knowledge of the local
forms and variability. Photos has been taken using
a DigitalSLR Canon EOS 400D.
Sometimes in the labels the term “co-typus” is
abbreviated in “c-t”. The term “co-typus” is used
by Monterosato and Coen to indicate even the
types, anyway it is no more accepted by ICZN.
RESULTS
The species are listed in the same order of
description, to be clear, in case of synonymy, which
name has precedence. Plate IV, showing the pyr-
amidellid is the same in both works (Coen, 1 933,
1 937), therefore the comments are the same. The
locality “Lido” is the name of the long and narrow
island in front of Venice, that is exposed to open sea
and where is possible to collect fresh shell grit. For
each bibliographic reference it is indicated the used
name, because this is sometime different.
Quality of Coen’s drawing is very bad, much
worse than drawing of shells having some dimen-
sion, done a century before by other Authors
(Philippi, Hoernes, Wood, Deahayes, etc.). All the
drawings show a pointed apex, while some of the
described species have a blunt apex!
In addition to the species dealt with in the
present work, there is a Pyrgulina praecisa Coen,
1914 ex Monterosato ms, of which theAuthorgives
Coen’s Pyramidellidae (Gastropoda Heterobranchia): a revision of types
417
only a very bad photo at pi. IV, fig. 19. This name
is not mentioned in the following w orks . A ccording
to art. 12.2.7 ofICZN, a species name based on an
illustration of the taxon being named to be treated
as not having been published, therefore this is a
nomen nudum.
Tiberia ( Tiberiella ) pretiosa Coen, 1933
ex Monterosato ms.
Tiberia ( Monterosato ) pretiosa Monterosato (sic) -
Coen, 1933: 5 1 (n° 336), 1 64 (note 97), pi. IV,
fig. 3 2
Tiberia ( Tiberiella ) pretiosa Monterosato - Coen,
1 937: 3 8 (n° 261), 148 (note 92 bis), pi. IV, fig.
32
Labels. • Coen’s label with number 730 1.
• M onterosato’s handwritten label “ Tiberiella ,
Mont. ms. T. pretiosa, Mont. Lido!!! ombelicata!”
• M onterosato’s handwritten label “ Tiberia
Mont. S ezio n e di Pyra m idella , p ub blic ata” [ T ib e ria
Mont. S ection of P y ram id e l la , published]
• Museum’s label with register numbers “HU J
53798” and “Coen 7301”.
Remarks. Coen (1 933, 1 64) describes the new
species as: “ Vale la pena di rich iam are V attenzione
su questa rarissima, minuta specie, di cui il mio
esemplare fu determinato dall’Autore medesimo.
La specie tipo di un sotto genere Tiberiella, non
pub blic a to, del Monterosato stesso, e ombilicata
(fig. 3 2)”. [It is worthwhile to call the attention on
this very rare, minute species, of which, my only
specimen was determined by the Author himself.
It is the type species of the subgenus Tiberiella, not
published, of M onterosato’s himself, and is umbil-
ic ate d ] .
Really the species is listed as Tiberia pretiosa
and the new genus Tiberiella is mentioned only in
the note 97. This overlook will be rectified later on
(Coen, 1 937) and the species listed as Tiberia
( Tiberiella ) pretiosa. Author draws one specimen
whose height, derived by the indicated scale factor
is 2.5 mm. Note is unchanged in the second work
(Coen, 1 937). Therefore both the species and the
genus were named by Monterosato, but published
by Coen. The holotype is a worn specimen, 2.4 mm
high (Fig. 1). The Coen’s label indicates as locality
“Lido”. The conical shape, and weak axial sculpture
of the holotype suggest that could be P arthenina
m onte rosatii (Clessin, 1900), that is not rare in the
area. This specimen is the same mentioned by
Aartsen et al. (1998: 7), who refers it to P arthenina
obtusa (Brown, 1 827).
Genus Tiberiella Coen, 1 933 is considered by
Aartsen et al. (1998: 7) and Schander et al. (1999:
151) a junior synonym of P a rth enina Bucquoy,
Dautzenberg etDollfus, 1883, because both have as
type species P arthenina obtusa. Based on our
determination of holotype, the two genera do not
have the same type species, but we agree that shall
be considered synonyms.
The specific name Tiberia ( Tiberiella ) pretiosa
Coen, 1933 ex Monterosato ms. shall be con-
sidered junior synonym of P arthenina m o nte ro s a tii
(C less in , 1 9 00).
Odostomia litoris Coen, 1933
Odostomia litoris Coen - Coen, 1933: 52 (n° 345),
164 (note 98), pi. IV, fig. 33
Odostomia litoris Coen - Coen, 1937: 40 (n° 293),
149 (note 103), pi. IV, fig. 33
Labels. • Coen’s label “N° 7311 Odostomia
acuta veil eta typus (ms). Lido!”.
• M o n tero s ato ’s handwritten label “Odostom ia
gruppo dell ’acuta”
• Museum’s label with register numbers “HUJ
20846” for lectotype and 53797 for paralectotype,
both with “Coen 73 11”.
• A label stating that “lectotype chosen
AARTSEN, 1 982 - middle one with embryonic
whorls preserved”
Remarks. Coen (1933) describes the new
species as: “La nuova specie, da me trovata nel
detrito di Lido, e sottoposta a l M o nte ro s ato , fu da
Lui riconosciuta come una nuova “Odostomia del
gruppo dell’ acuta (sic). Analog a a questa, lie
differisce perd per i giri piatti anziche convessi; per
la sutura line a re fortemente impress a; infill e per
una carena, ottusa ma ben p ro nunziata , che appare
suU’ultimo anfratto (fig. 33). Venezia-Lido” . [The
new species, found by me in the shell grit from
Lido, and submitted to Monterosato, was con-
sidered by him as a new “Odostomia of the acuta
418
Pasquale Micali et alii
group”. It is similar to this, but differs for the flat
instead of convex whorls; for the lineas suture
strongly impressed; at last for a keel, obtuse but
well pronunced, present on the last whorl (fig. 33).
Venice-Lido].
Note is unchanged in the second work (Coen,
1 937). Author draws one specimen whose height,
derived by the indicated scale factor is about 2.9
mm. In this case the specific name was not assigned
by M on te ro s ato .
The lectotype is a worn specimen, 2.3 mm high
(Fig. 6), fixed with the p arale c to ty p e on a black
paper strip. The Coen’s label indicates as locality
“Lido”. The specimen is Odostomia unidentata
(Montagu, 1 803 ), a species not rare in the area.
Odostomia acuta Jeffreys, 1 848 is normally ombel-
icated, has rounded, instead of angulated peri-
phery, oblique instead of angulated columella,
lower first teleoconch whorl and slightly rounded
w horls.
Pyrgulina denticulus Coen, 1933
ex Monterosato ms.
Pyrgulina denticulus Monts. - Coen, 1914: 12, pi.
IV, fig. 17 nomen nudum
Pyrgulina denticulus Monterosato mss. - Coen,
1 933: 52 (n° 355), 1 64 (note 99), pi. IV, fig. 34
Chrysallida ( Babella ) denticulus Monterosato-
Coen, 1 937: 3 8 (n° 274), 149 (note 99), pi. IV,
fig. 34
Labels. • Coen’s label “N 0 7 3 1 8 Chrysallida
(Pyrgulina) denticulus Monts, c. t. Ven ezia -L ido” .
• M onterosato’s handwritten label “Pyrgulina
denticulus Mont. Lido!”
• M onterosato’s handwritten label “La piu
comune tra le Pyrgulina del Lido” [The most com-
mon Pyrgulina at Lido].
• Museum’s label with register numbers “HUJ
53792” for lectotype and 53793 for paralectotype,
both with “Coen 7318”.
• A label stating that “lectotype selected by van
Aartsen in glass vid”
Remarks. Coen (1933, 164) describes the new
spec ie s as : “ A ssai sim He ad una Turbo n ilia, e m olto
vicina alia P. interstincta Montagu per la forma
nettamente turriculato -conica, a profilo quasi
rettilineo. Le pieghe oblique degli anfratti sono
molto accentuate; ognuna di esse porta un tuber-
colo sotto la sutura eel e solcata alia base, onde la
conchig lia tie appare clatrata, in sen so inverso,
sub suturalm ente eel al posto della carena (fig. 34).
Venezia-L ido” . [Very similar to a Turbonilla, it is
very close to P. interstincta Montagu for the tur-
riculate-conical shape, with an almost straight
profile. The oblique costae are very strong; each
one has a tubercle below the suture and is sulcate
at the base, giving to the shell a clathrate appear-
ance in reverse way, subsuturally and at the
place of the keel (fig. 34). Venice-Lido]. The boi
ded part of translation is doubtful because meaning
of the sentence in original description is not clear.
Note is unchanged in the second work (Coen,
1 937). Author draws one specimen, whose height,
derived by the indicated scale factor is about 3 mm.
The lectotype is a 3 mm high (Fig. 11), a little
worn and is P arthenina tereb ellum (Philippi, 1 844),
a species very common in the shell grit of Venice
are a .
Pyrgulina alabastrum Coen, 1933
ex Monterosato ms.
P y rg ulin a alab a strum M o n ts . - C o en , 1 9 1 4 : 12, pi.
IV, fig. 18 nomen nudum
Pyrgulina alabastrum Monterosato mss. - Coen,
1 933: 52 (n° 35 8), 1 64 (note 99), pi. IV, fig. 36
Chrysallida (P arthenina) alabastrum Monterosato
mss. - Coen, 1937: 3 8 (n° 275), 149 (note 96),
pi. IV, fig. 36
Labels. • Coen’s label “N° 732 0 Chrysallida
(Pyrgulina) alabastrum Monts, c -t Ve n ezia Lido”.
• M onterosato’s handwritten label “ Pyrgulina
alabastrum M o n ts . L id o ! ! ”
• Museum’s label with register numbers “HUJ
53786” for holotype, with “Coen 7320”.
Remarks. Coen (1933, 164) describes the new
species as: “ Unico esemplare di una forma vera-
mente turb onillo ide . Conchiglia esilissim a, turricu-
lata, ottusa all’ a pice, composta di 6-7 giri
pianeg g ianti, un poco allargata verso la base, or-
nata di pieghe longituclinali non oblique, solcate
sp ir aim ente sotto la sutura, che e fortemente
Coen’s Pyramidellidae (Gastropoda Heterobranchia): a revision of types
419
impressa (fig. 36). Detrito di Lido”. [A single spe-
cimen of a maredly turbonilloid shape. Shell very
thin, turriculate, blunt at the apex, consisting of 6-
7 flat whorls, a little enlarged toward the base,
sculptured by longitudinal ribs, not oblique, spirally
furrowed below the suture, that is strongly im-
pressed (fig. 36). Shell grit from Lido].
Note is unchanged in the second work (Coen,
1 937). Author draws one specimen, whose height,
derived by the indicated scale factor is about 2.8
m m .
The lectotype is a fresh specimen, 2.7 mm high
(Fig. 17); the presence of two spiral cords on upper
whorls and the lack of columellar plica, indic-
ates that the specimen is a P arthenina indistincta
(Montagu, 1 808), a species quite frequent in the
area, even at low depth.
Pyrgulina ordita Coen, 1933 ex M onterosato ms.
P yrgulina ordita M onterosato - Coen, 1 933: 52 (n 0
360), 1 65 (note 100), pi. IV, fig. 41
C hrysallida ( Trabecula ) ordita M on tero sato - Coen,
1 937: 3 8 (n° 280), 149 (note 1 0 2 ) , p 1. IV, fig . 4 1
Labels. • Coen’s label “N° 7321 C hrysallida
( Pyrgulina ) ordita Mont. Co-types Venezia Lido”.
• M onterosato ’s handwritten label “ Pyrgulina
ordita Monts. Lido!!” and “una delle piu merav-
igliose per la scultura”
• Museum’s label with register numbers “HUJ
53783” for holotype, with “Coen 732 1 ” number.
Remarks. Coen (1 933: 1 65) describes the new
species as: “ Conchiglia rissoiforme, tenuissim a ,
jalina; ultimo giro grande; scultura consistente in
strie long itudinali sottili, estese a tutta la superficie,
anche has ale; peristoma norm ale; fessura ombel-
icale apparente (fig. 41). Detrito di Lido”. [Shell
ris so id -like , very thin, translucent; last whorl large;
sculpture consisting of thin spiral striae, covering
the whole surface, including the base; peristoma
norm al; false ombelical slit (fig. 41). Shell grit from
L id o ] .
Note is unchanged in the second work (Coen,
1937). Author draws one specimen, whose height,
derived by the indicated scale factor is about 1.6
m m .
The holotype is a broken specimen having the
last three whorls, 1.6 mm high (Fig. 21). The
flexuous and crowded axial ribs and the numerous
spiral cords clearly indicates that the holotype is a
P arthenina juliae (d e F o lin , 1 872), a sp ec ie s com-
mon in the area at low depth.
Pyrgulina coeni Coen, 1933 ex M onterosato ms.
Pyrgulina coeni Monts. - Coen, 1914: 1 2 , p 1. IV, fig .
21 nomen nudum
Pyrgulina coeni M onterosato mss. - Coen, 19 3 3: 52
(n° 36 1), 1 64 (note 99), pi. IV, fig. 37
C hrysallida (P artulida) coeni M onterosato - Coen,
1937: 38 (n° 278), 149 (note 98), pi. IV, fig. 37
Labels. • Coen’s label “N° 7322 C hrysallida
(P y rgulina) Coeni M o n ts . ty p e s Ven ezia-L ido ” .
• M onterosato’s handwritten label “ Pyrgulina
Coeni , Monts, ms. Lido!! Non esti (unclear) P.
turhonilloides , Brusin, apice etc.!”
• Museum’s label with register numbers “HUJ
53 7 87” for lectotype and 5 37 88 for paralectotype,
both with “Coen 7 3 22”.
• A label stating that “two species! Left one was
selected by van Aartsen as the lectotype of P. coeni,
right one is too badly preserved to allow identi-
fication”
Remarks. In the first mention of the name
(Coen, 1914: 12) add the note: “non = turbonil-
loides Brus.”, meaning that it is similar, but dif-
ferent from turhonilloides (Brusina, 1 869).
Coen (1933: 164) describes the new species as:
“Essa si differenzia dalle precedents per essere
ovoide, di habitus veram ente rissoiforme, conser-
vando perd tutti i c a ratte ri del gruppo. Le pie g he
longitudinals, non oblique, ne tub ercolate , ne
solcate, cessano b ruscam ente nella regione car-
enale, cost che la base dell’ultim o giro e per- fet-
tamente liscia e lucente. B ian co -latte a (fig. 37).
Detrito di Lido”. [This species differs from the
formers for the ovoid shape, of really rissoid-like
habitus, maintaining anyway all the characters of
the group. The longitudinal ribs, not oblique,
neither tuberculate nor sulcate, abruptely ending at
the periphery of last whorl, so the base of the last
whorl is perfectly smooth and bright. Milky-white
(fig. 37). Shell grit from Lido],
420
Pasquale Micali et alii
M
tS
N
kem&uZ
ft i iJMia
(j*c£
s&hmn/- £? 8 l
AiFpFjpt- ,**4^*#^
* U jl* Aa itt'd
/v. /J/S
*2Z?
(X-C-COCoc- z,'*e4*Jtdj6u <%
■■■■
^Z~~Q~
/ X,'&
CT*
THE HEBREW UNIVERSITY OF JEhUSALE
ZOOLOGICAL MUSEUM
HUJ S’ 5^1
t-ijr g nl ~i nft tU.v\Vi £,«.'! uvl
CoCmAO?
Locality
; Vfci4ktt , \.*A*
» + »«■*»•►••■•■ D®m «+*++■*■»*»«
Coll Oti. ^
J-liS
THE HE SHEW UNIVERSITY OF JERUSALEM
ZOOLOGICAL MUSEUM
HUJ f ■
0^o3.bB WA
C*L4 HW
LoeaJiitv | . . • -
Y*t* * 4*. . L> A 0
o*i# . , . . ..
Coll. Oat ti i
Coax Mil
Figure 1. Tib eria (Tib e rie l la) pretio sa Coen’s collection, holotype HUJ 53798, H: 2.4 mm, Venice-Lido. Figs. 2-4. Original
labels. Fig. 5 . Museum’s label. Fig. 6 . P arthenina m ontero satii (C le s sin , 1900). Figs. 7 , 8 . O dostom ia lit or is Coen’s collec-
tion, lectotype and paralectotype, HUJ 20846, H: 2.3 mm, Venice-Lido. Figs. 9-10. Original labels. Fig. 11. Museum’s label.
Fig. 12. O do sto m ia un id entata (Montagu, 1803). Fig. 13. P yrgulina den tic ulus Coen’s collection, lectotype HUJ 53792, H:
3 mm, Venice-Lido. Figs. 14, 15. Original labels. Fig. 16. Museum’s label. Fig. 17. P arthenina te re b e llum (Philippi, 1844).
Coen’s Pyramidellidae (Gastropoda Heterobranchia): a revision of types
421
. c.-rA
JCj&+
THE HEBREW UNIVERSITY OF JERUSALEM
ZOOLOGICAL MUSEUM
HUJ 5?f fl6 . . .
Py ' hflL
Cu<-rt
Locality ^ _ , . . ■ -
V*tCi^4.
Date
Coll D«
N ./
THE HEBREW UNIVERSITY OF JERUSALEM
ZOOLOGICAL MUSEUM
huj£*H3 ......
Orditu
r , . ..'&** i*j33
Locality .......
Oate .............
Coll Oet &&£****
C At* fill
4
■*
//
THE HEBREW UNIVERSITY OF JERUSALEM
ZOOLOGICAL MUSEUM
HUJ 5" 3} #3 ^
■Bata.'AlJ , . .
X
//
f**4f
Locality
1 t||l I ¥c*Uc - 4*
........... Date .... +
Coll * , Det. M
-K ./
t ui 4 5 r 1 * ' ■
Wfr**.* ki*^
A.rVj#^ FO
4 p -***“ :
rijliF tn. >' *- F " , * ,
— 7
Figure 18. Pyrgulina a lab a strum Coen’s collection, lectotype, HUJ 53786, H: 2.7 mm, Venice-Lido. Figs. 19, 20. Original
labels. Fig. 2 1. M useum’s label. Fig . 2 2. P a rthen in a in d is tin eta (Montagu, 1 808). Fig . 23. Pyrgulina ordita Coen’s collection,
holotype, HUJ 53783, H: 1.6 mm, Venice-Lido. Figs. 24, 25. Original labels. Fig. 26. Museum’s label. Fig. 27. P a rthen in a
juliae (de Folin, 1 872). Fig. 28. Pyrgulina coeni Coen’s collection, lectotype, HUJ 5 3 787, H: 1.9 mm, Venice-Lido. Figs.
29, 30. Original labels. Fig. 31. Museum’s label. Fig. 32. P artulid a incerta (Milaschewitsch, 1916)
422
Pasquale Micali et alii
Note is unchanged in the second work (Coen,
1 937). Author draws one specimen, whose height,
derived by the indicated scale factor is about 2 mm.
The lectotype is a worn specimen, 1.9 mm high
(Fig. 25). Its shape leaves no doubt that it is a
P artulid a incerta (Milaschewitsch, 1916) of which
are synonyms P. turb o nillo id e s (Brusina, 1869)
and P. brusinai (Cossmann, 1921). This species is
frequent in the area, at low depth.
Pyrgulina pyrgulella Coen, 1933
ex Monterosato ms.
Pyrgulina pyrgulella Monterosato - Coen, 1933: 52
(n° 363), 1 65 (note 101), pi. IV, fig. 42
C hrysallida (O do s to m e l la) pyrgulella M onterosato
mss. - Coen, 1 937: 3 8 (n° 269), 148 (note 94),
pi. IV, fig. 42
Labels
• Coen’s label “N 0 7 3 2 3 C hrysallida (P yrgulina)
pyrgulella Monts, (unclear) Venezia Lido”.
• M onterosato’s handwritten label “ Pyrgulina
pyrgulella Monts. Lido!!”
• Museum’s label with register numbers “HUJ
5 3 79 1” for holo type with “Coen 7323”.
Remarks. Coen (1933: 164) describes the new
species as: “ Conchiglia tu rricu lata , ottusa alia
sommita, com posta di 6-7 giri, lisci gli apicali, gli
altri solcati lo ng itudinalm ente , salvo una zona sub-
suturale di ogni giro e la base dell’ultimo, die
rim angono liscie; columella obliquam ente e forte-
mente ritorta; peristoma sinuoso alia base. Bianca
(fig. 42). Detrito di Lido”. [Shell turriculate, blunt
at the apex, consisting of 6-7 whorls, the apicals are
smooth, the others longitudinally costate, except
one subsutural zone of each whorl and the base of
the last whorl, that are smooth; columella markedly
and obliquely folded; peristome sinuose at the base.
White (fig. 42). Shell grit from Lido].
Note is unchanged in the second work (Coen,
1937). Author draws one specimen, whose height,
derived by the indicated scale factor is about 2 mm.
The holotype is 2.1 mm high (Fig. 30), worn and
partially broken. The first teleo conch whorl appears
smooth and this character suggests that specimen
could be P arthenina terebellum (Philippi, 1 844)
(Clessin, 1900), a species quite frequent in the area,
even at low depth.
Pyrgulina vixstriata Coen, 1933
ex Monterosato ms.
Pyrgulina vixstriata Monterosato mss. - Coen,
1 933: 52 (n° 364), 1 65 (note 99), tav. IV, fig. 38
C hrysallida (O do stom ella) vixstriata Monterosato
mss. - Coen, 1 937: 3 8 (n° 267), 148 (note 93),
tav. IV, fig . 3 8
Labels
• Coen’s label “N 0 7 3 24 C hrysallida ( Pyrgulina )
vix striata M onts. c-t Venezia Lido”.
• M onterosato’s handwritten label “Pyrgulina
vix-striata, Mont, buon tipo Lido!! N . B. la gomma
non fa apparire le strie ” [ Pyrgulina vix-striata ,
Mont, valid type Lido!! N. B. the glue prevents the
observation of the striae].
• Museum’s label with Lectotype register num-
bers “HUJ 5 378 1 ” and Paralectotype “HUJ 53782”,
both with “Coen 7324”.
• A label stating that: “the single shell was
selected as lectotyp e by van A arisen, 25.8.1987”.
Remarks. Coen (1933: 148) describes the new
species as: “Ha un habitus ov o ide-ris soifo rm e ;
molto ottusa all’ a pice, const a di 4-5 giri convessi,
ornati di pieghe longitudinali ondulate, che sono
solcate trasv ersalm ente sopra la sutura, eel evanes-
centi (non troncate) sulla base dell’ultimo giro.
B ianca (fig . 3 8). D etrito di Lido” . [It has an o v o id ,
rissoid-like profile; very blunt at the apex, with 4-5
convex whorls, sculptured by longitudinal ondu-
lated ribs, that are transversally sulcate above the
suture and evanescent (not truncated) at the base of
the last whorl. White (fig. 38). Shell grit from Lido]
Note is unchanged in the second work (Coen,
1 937). Author draws one specimen from Venice,
whose height, derived by the indicated scale factor
is 1.5 mm.”
The lectotype is a specimen 1.3 mm high (Fig.
35). The flexuous and crowded axial ribs and the
numerous spiral cords clearly indicates that both the
specimens are P arthenina juliae (de Folin, 1 872),
a species common in the area at low depth.
It is intersting to note that the subgenus has been
corrected in Coen’s label to Pyrgulina , that is not
the one used in the second publication. Unfortu-
nately it is not possible to identify the erased word.
Coen’s Pyramidellidae (Gastropoda Heterobranchia): a revision of types
423
Pyrgulina brevicula var. rejecta Coen, 1933
ex Monterosato ms.
Pyrgulina brevicula var. rejecta Monterosato -
Coen, 1 933: 54 (n° 367), 1 65 (note 99), pi. IV,
fig- 39
Chrysallida ( Partulida ) brevicula rejecta Monterosato
mss.- Coen, 1 937: 3 8 (n° 279), 149 (note 98),
pi. IV, fig. 39
Labels. • Coen’s label “N° 7326 Chrysallida
( Pyrgulina ) tu rb o nillo id e s rejecta Monterosato c-t
Venezia Lido!”.
• M onterosato’s handwritten label “Pyrgulina
rejecta, Monts. Lido!! Gruppo della turbonilloides”
• Museum’s label with register numbers “HUJ
53779” for lectotype and 53780 for paralectotype,
both with “Coen 7326”.
• A label stating that “Specimen to the extreme
right was selected as the lectotype by van Aartsen
25.8.1 987”.
Remarks. Coen (1933: 164) describes the new
species as: “ Forma poco diversa da lla tipica, cost
nominata perche trovata nel detrito: carattere, id
solco spirale alia base delle pieghe longitudinali,
molto p rofo nd am ente impresso. Bianca (fig. 3 9).
Detrito di Lido”. [Shape a little different from the
typical, so called because found in the shell grit:
characterised by very deeply impressed spiral
groove at the base oflongitudinal ribs. White (fig.
39). Shell grit from Lido],
Note is unchanged in the second work (Coen,
1937). Author draws one specimen, whose height,
derived by the indicated scale factor is about 1.6 mm.
The lectotype is a worn specimen, 1.8 mm high
(Fig. 41). Its shape leaves no doubt that it is a
Partulida incerta (Milaschewitsch, 1916) of which
are synonyms turbonilloides (Brusina, 1 869) and
b rusinai (Cossmann, 1 9 2 1 ) . T his specie s is freq uen t
in the area, at low depth.
The lectotype has been selected by Aartsen (see
above its label) from a set of four specimens fixed
on a black paper strip (Fig. 42). The specimen at
extreme left of the strip seems to be a P arthenina
inclistincta (Montagu, 1 808).
It is surprising to note that both M onterosato’s
and Coen’s label correctly refer the variety
“ rejecta ” to turbonilloides, while in both publica-
tions Coen uses a different specific name.
The real identity of Chrysallida brevicula
(Jeff re ys, 1883), originally described as O dostomia
brevicula, has been clarified by Giannuzzi Savelli
et al. (2011), who studied the type material with the
result that the species was based on an immature
specimen of Turbonilla amoena (Monterosato,
1 878).
Pyrgulina cylindracea Coen, 1933
ex Monterosato ms.
Pyrgulina cylindracea Monterosato mss. - Coen,
1933: 54 (n° 368), 1 65 (note 1 02), p 1. IV, fig . 4 3
Chrysallida (O dostom elda) cylindracea Monterosato
mss. - Coen, 1937: 38 (n° 271), 149 (note 101),
pi. IV, fig. 43
Labels. • Coen’s label “N° 7327 Chrysallida
(Pyrgulina) cylindracea Monts, c - t Ven ezia-L ido ! ” .
• M onterosato’s handwritten label “ Pyrgulina
cylindracea Monts. Lido!!”
• Museum’s label with register numbers “HUJ
5 3 784 ” for le c to typ e and 53785 for paralectotype,
both with “Coen 73 27”.
Remarks. Coen (1933: 165) describes the new
species as: “C onchiglia turriculata, m elaniifo rm e ,
vitrea, con 5-6 giri poco, ma re g o la rm e rite ,
convessi, de i quali i p rim i so no glob ulo si e lis c i, g Id
altri deb olm ente solcati. Base liscia. Apertura alia
base a cu tarn e rite svasata, come in una Melania (fig.
43). Detrito di Lido”. [Shell turriculate, melania-
like, vitreous, with 5-6 whorls, slightly and regu-
larly convex, the formers are globose and smooth,
the latters weakly sulcate. Base sm ooth. A perture
acutely expanded at the base, like in a M elan ia (fig.
43). Shell grit from Lido].
Note is unchanged in the second work (Coen,
1937). Author draws one specimen, whose height,
derived by the indicated scale factor is about 1.8 mm.
The lectotype is a worn and partially broken
specimen, 1.9 mm high (Fig. 48) and seems to be
P arthenina te reb ellurn (Philippi, 1 844), a species
very common in the shell grit of Venice area. The
figured specimen is fixed to a black paper strip (Fig.
49) with the final portion (about 2.5 whorls) of a
much larger specimen that seems to belong to same
species.
424
Pasquale Micali et alii
'’*** A* if. /
■fa?
'T' LJL
fa t > I m/if * U
r
THE KERRCVV UNIVERSITY Of JERUSALEM
zoological museum
hujJ^SI
FnitOuIIaa VkAitr.'ntji
FHtA t*V- n * ifer >
C»*y
Xt-^. A*^°
^ v:
■ . Dftt t
Coll. Oer
AilJS*
THE HEBREW UNIVERSITY Of JERUSALEM
ZOOLOGICAL MUSEUM
HUJ OtA 1
Co*l'T
Dili
0*1 t ^ f*C*h
con
e.
Figure 33. P yrgulin a pyrgulella Coen’s collection, holotype, HUJ 53791, H: 2.1 mm, Venice-Lido. Figures 34, 35. Ori-
ginal labels. Fig. 36. Museum’s label. Fig. 37. Parthenina terebellum (Philippi, 1 844). Fig. 38. Pyrgulina vixstriata
Coen’s collection, lectotype, HUJ 5 3 7 8 1, H: 1.3 mm, Venice-Lido. Fig. 39. P. vixstriata , paralectotype HUJ 5 3 7 82,
H: 1.5 mm, Venice-Lido. Figures 40, 41. Original labels. Fig. 42. Museum’s label. Fig. 43. Parthenina juliae (de Folin,
1 872).
[22
Coen’s Pyramidellidae (Gastropoda Heterobranchia): a revision of types
425
to fcl*. 1
W*J
bL<- 1 *-<-X i t-Jpt t
THE HEBREW UNIVERSITY OF JERUSALEM
ZOOLOGICAL MUSEUM
Tyrjulbin
h
A V«r- TCjAttA
C**€Yl T^l 53
Locality ,
Vc,a n* , L*4«
Due ,,,,..
Coll D*t & „i .to 1*1
£♦»* TStfc
THE HEBREW UNIVERSITY OF JERUSALEM
ZOOLOGICAL MUSEUM
hujJHS 0
rwr**J*H* tfivl tnU '»*/.
— - — c^u l \sr~~
L«ihiv . . . , ,
C4 . Li J o
......... Dll* , , . . ,
Coll Dot U 3 Coe*,
Co*m *)/i
£4+4*_ /
Py £ i'i i ^*r r ' e\^
..4k*t6.-.c -4*
THE HEBREW UNIVERSITY OF JERUSALEM
ZOOLOGICAL MUSEUM
r thrift Viy,gliY;t4
Cue.* |Jj i J
Localitv
i (A. L,'.
........... Dale , .
Coir. . . Dei
Co**l 1 it*
THE HEBREW UNIVERSITY OF JERUSALEM
ZOOLOGICAL MUSEUM
muj n>0.£
P^r US a - aMi*
Cjh*^ ij|>3
Locality
X fe#A J ,Vm\u . WT A?
Data , .
Call. ....... Dei tSOtH ...
£*•»
Figures 44, 45. Pyrgulina b revicula var. rejecta Coen's collection, lectotype, HUJ 5 3779, H: 1.8 mm, Venice-Lido and
paralectotypes HUJ 53780, the specimen applied on paper strip at extreme left is a P a rthen in a indistincta (M ontagu, 1808).
Figs. 46, 47. Original labels. Fig. 48. Museum's label. Fig. 49. Partulida incerta (M ilasche w itsch, 1916). Figs. 50, 51.
Pyrgulina cylind racea Coen’s collection, lectotype, HUJ 53784, H: 1.9 mm, Venice-Lido and paralectotype HUJ 5 3785.
Figs. 52, 53. Original labels. Fig. 54. Museum’s labels. Fig. 55. P a rthen in a tereb ellum (Philippi, 1844).
426
Pasquale Micali et alii
Pyrgulina canaliculata Coen, 1933 ex Monterosato
m s .
Pyrgulina canaliculata Monterosato ms. - Coen,
1 933: 54 (n° 369), 1 65 (note 99), pi. IV, fig. 40
C hrysallida (P arthenina) canaliculata Monterosato
mss. - Coen, 1 937: 3 8 (n° 276), 149 (note 97),
pi. IV, fig . 4 0
Labels. • Coen’s label “N° 7328 C hrysallida
{Pyrgulina) canaliculata co-types. Venezia Lido”.
• M onterosato’s handwritten label “ Pyrgulina
canaliculata M onts. Lido ! !”
• Museum’s label with register numbers “HUJ
5 3 7 94 ” fo r holo typ e , with “Coen 7328”.
Remarks. Coen (1933: 165) describes the new
species as: “Conchiglia turricu lato -m etafo rm e ,
con giri piani, di cui i primi lisci, gli altri con
pieghe longituclinali profonde, tagliate sopra la
sutura da un solco trav ersale , e sull’ultimo giro
clatrate fino alia meta; base liscia, apertura
normale. II nome viene clalla sutura, p ro fonda-
nt ente imp ressa . Bianca {fig. 40). Detrito di Lido”.
[Shell turriculate, whorls flat, the formers smooth,
the following with deep longitudinal ribs, crossed
above the suture by a spiral groove and, on the
last whorl, clathrate up to the middle; base smooth,
aperture normal. The name derives from the
deeply impressed suture. White (fig. 40). Shell grit
fro m Lido].
Note is unchanged in the second work (Coen,
1937). Author draws one specimen, whose
height, derived by the indicated scale factor is
about 1.9 mm.
The holotype is a worn specimen, 1.8 mm high
(Fig. 5 1). The presence oftwo spiral cords on upper
whorls and the lack of columellar plica, indic-
ates that the specimen is a P arthenina indistincta
(Montagu, 1 808), a species quite frequent in the
area, even at low depth.
Pyrgulina mitis Coen, 1 933 ex Monterosato ms.
Pyrgulina mitis Monterosato ms. - Coen, 1933:
54 (n° 370), 1 65 (note 103), pi. IV, fig. 44
C hrysallida (O d o s to m e lla) mitis Monterosato
mss. - Coen, 1937: 38 (n° 270), 149 (note 100), pi.
IV, fig. 44
Labels. • Coen’s label “N° 7329 C hrysallida
( Pyrgulina ) mitis Mont, c-t Venezia - Lido”.
• M onterosato’s handwritten label “ Pyrgulina
m itis Monts. Lido!!”
• Museum’s label with register numbers “ HUJ
5 3 7 8 9 ” fo r lectotyp e and 53 790 for pa ralecto typ e ,
both with “Coen 73 2 9”.
• A label stating that “ most right hand specimen
se lected as lectotyp e by van Aartsen 25.08.1989 ”
Remarks. Coen (1933: 165) describes the new
species as: “ Conchiglia turriculata, ad apice ottuso,
giri piani, lisci i primi, gli altri con forth solchi per
lungo, cess anti alia meta dell’ ultimo giro eel ivi
clatrati; sutura p rofo ndis sim a ; apertura allungata,
svasata acutam ente alia base; la columella porta
una piega dentiform e mediana molto pronunziata
eel una p rofo n d a fe s s u r a ombilicale lungo la cal-
losita colum e Ida re . B ianca (fig . 44 )” . [Shell turricu-
late, apex blunt, whorls flat, the initials are smooth,
the following with strong ribs, ending at the middle
of the last whorl, where are clathrate; Suture very
deep; aperture elongate, acutely expanded at the
base; on the central part of the columella there is a
strong tooth-like fold and a deep umbilical chink
along the columellar callus. White (fig. 44).].
Note is unchanged in the second work (Coen,
1937). Author draws one specimen, whose height,
derived by the indicated scale factor is about 1.8 mm.
The lectotype is 1.8 mm high (Fig. 57); it is
P arthenina te re b e llum (Philippi, 1 844), a species
very common in the shell grit of Venice area. The
figured lectotype (the one selected by van Aartsen)
is fixed to a black paper strip (Fig. 58) with other
three specim ens.
Eulimella curtata Coen, 1 93 3 ex Monterosato
m s .
Eulimella curtata Monterosato ms. - Coen, 1933:
54 (n° 375), 1 65 (note 104), pi. IV, fig. 45
Eulimella curtata Monterosato mss. - Coen, 1937:
40 (n° 300), 149 (note 104), pi. IV, fig. 45
Labels. • Coen’s label “N° 7 3 3 1 Eulimella
curtata Monts, c-t Venezia Lido”.
• M onterosato ’s handwritten labels “Eulimella
com m utata = acicula Lido!!” and “ non e commut-
ata m a curtata a Lido anche la non comm utata”.
Coen’s Pyramidellidae (Gastropoda Heterobranchia): a revision of types
427
Figure 56. P yrgulina ca n a lieu la ta Coen’s collection, holotype, HUJ 52794, H: 1.8 mm, Venice-Lido. Figures 57, 58. Original
labels. Fig. 59. Museum’s label. Fig. 60. P arthenina indistincta (Montagu, 1 808). Figs. 61, 62. Pyrgulina mitis Coen’s
collection, lectotype, HUJ 5 3789, H: 1.8 mm, Venice-Lido and paralectotypes HUJ 5 3790. Figs. 63, 64. Original labels.
Fig. 6 5. Museum’s label. Fig. 66. P arthenina terebellum (Philippi, 1 844).
428
Pasquale Micali et alii
Figure 67. Eulimella curtata Coen’s collection, holotype, HUJ 20847, H: 2.7 mm, Venice-Lido. Figs. 68, 69. Original
labels. Fig. 70. Museum’s label. Fig. 71. Eulimella flagellum Coen’s collection, holotype, HUJ 20848, H: 2.7 mm,
Venice-Lido Figs. 72, 73. Original labels. Fig. 74. Museum's label. Fig. 75. Eulimella acicula (Philippi, 1836).
M eaning of the second label not clear, therefore not
translated.
• Museum’s label with register numbers “ HUJ
Remarks. Coen (1933: 165) describes the new
species as: “Poco lontana clalla E. acicula Philippi,
se ne distingue per I’habitus piii m etafo rm e ed ot-
tuso all’ ap ice , per i giri piii convessi, e sop rattutto
2 08 47” for ho lo ty p e , w ith “Coen 7331 ”
Coen’s Pyramidellidae (Gastropoda Heterobranchia): a revision of types
429
per I’apertura che, anziche allungata e regolar-
m ente curva alia base, vi e abbreviata e di forma
trap ezo idale . J a lina (fig . 4 5) . D etrito di Lido" . [Not
far from E. acicula Philippi, may be separated for
the profile more regular and blunt at the apex, the
more convex whorls, and mainly for the aperture,
which instead of elongate and regularly curved at
the base, is abbreviated and trapezoidal. Jaline (fig.
45). Shell grit of Lido],
Note is unchanged in the second work (Coen,
1937).Author draws one specimen, whose height,
derived by the indicated scale factor is about 2.3
m m .
The holotype is a little worn specimen, 2.7 mm
high (Fig. 63), the protoconch is coiled at right
angle with respect to teleoconch axis, the profile is
slightly cyrtoconoid, whorls are flat, suture inclined
and columellar plica is very weak or absent. The
holotype seems to be Eulimella acicula (Philippi,
1 836).
Eulimella flagellum Coen, 1933
ex Monterosato ms.
E u Urn ella flag ellu m Monterosato ms. - Coen, 1933:
54 (n° 377), 1 65 (note 105), pi. IV, fig. 46
Eu Urn ella flagellum M on tero sato ms. - Coen, 1937:
40 (n° 30 1 ), 149 (note 105), pi. IV, fig. 46
Labels. • Coen’s label “N° 7333 Eulimella
flag ellu m Monts, co - ty p u s (ms). Venezia L id o ! ” .
• M onterosato’s handwritten label “ Eulimella
flagellum Monts, oppure: (Unclear)! Lido ”
• Museum’s label with register numbers “ H U J
2 0 8 4 8 ” fo r ho lo typ e , w ith “Coen 7333”.
Remarks. Coen (1 933: 1 65) describes the new
species as: “ Forma stretta, allun g atiss im a , con 10
giri piatti lisci lucenti, sutura profonda, apertura
piriforme stretta ed allungata (fig. 46). D etrito di
Lido". [Shell narrow, very elongate, with 10 flat,
smooth and bright whorls, deep suture, aperture
pyriform, narrow and elongate (fig. 46). Shell grit
fro m L id o ] .
Note is unchanged in the second work (Coen,
1937). Author draw one specimen, whose height,
derived by the indicated scale factor is about 2.4
m m .
The holotype is 2.7 mm high (Fig. 67), the
protoconch is coiled at right angle with respect to
teleoconch axis, the profile is slightly cyrto-
conoid, whorls are flat, suture inclined and
columellar plica is very weak or absent. The
holotype could be a freak of Eulimella acicula
(Philippi, 1836), or an E u Urn e l la c lav a tula Sacco,
1 892, a pliocenic species (see Chirli & Micali,
2 0 11). The presence of a fossil specimen in the
shell grit collected on the beach of the Lido is
however unlikely.
ACKNOWLEDGEMENTS
We thank Dr. Henk K. Mienis, TelAviv Univer-
sity, Department of Zoology, Israel, manager of
m ollu skscollection for the loan of C oen ’sm aterial.
REFERENCES
A artsen J. J. van, Gittenberger E. & Goud J., 1998.
(Mollusca, Gastropoda, Heterobranchia) collected
during the Dutch CANCAP and Mauritania expedi-
tions in the south-eastern part of the North Atlantic
Ocean (part 1). Zoologische Verhandelingen, 321:
3-57.
Clessin S., 1 899-1902. Die Familie der Eulimidae. (Vol.
1, part 28: [4] + 273 pp., 41 pis.), in: Kiister H.C. (ed.),
1837-1919 [1918]. System atisches Conchylien-
Cabinet von Martini und Chemnitz. Bauer & Raspe,
N urn berg (the page concerning P. monterosati was
published in 1900).
Chirli C. & Micali P., 2011. Malacofauna Pliocenica
Toscana. Vol. 8. Pyram idelloidea. X + 131 pp, 40 pi.
Coen G., 1914. Contributo alio studio della Fauna mala-
cologica Adriatica. Regio Comitato Talassografico
Italian o . Memoria XLVI, Venezia, 34 pp., 7 pi.
Coen G ., 1 93 3. Saggio di una Sylloge Molluscorum
A driaticorum . Consiglio Nazionale delle Ricerche.
Regio Comitato Talassografico Italiano. Memoria
C X C II, Venezia, vii+ 186 pp., 10 pi.
Coen G ., 1 93 7. N uovo Saggio di una Sylloge Mol-
luscorum A driaticorum. Consiglio Nazionale delle
Ricerche. Regio Comitato Talassografico Italiano.
M e m oria CCXL, Venezia: vii+ 173 pp., 10 pi.
Giannuzzi Savelli R., Micali P., Nofroni I. & Pusateri F.,
2011. Odostomia b re vie it la Jeffreys, 1 883 junior
synonym of Turbonilla am oen a (Monterosato,
1 878 ) (Gastropoda, Heterobranchia, Pyramidellidae).
Biodiversity Journal, 2: 2 17-220.
430
Pasquale Micali et alii
MienisH.K., 2012. Four important contributed mollusc
collections their histories and contents. l.Giorgio S.
Coen (1873-1951) and his mollusc collection.
Haasiana, 6: 11-37.
P ian i P. 1 9 8 3. Della <<collezione M o n tero sa to > > , di G .S .
Coen e di altre cose ancora. Bollettino Malacologico,
19: 273-278.
Piani P.. Bouchet Ph. & Ghisotti F., 1990. Lavori Mala-
cologici di G.S. Coen. Bollettino Malacologico, 26:
148-152.
Schander C . , A arts e n J. J. van & Corgan J., 1999. Famil-
ies and genera of the Pyramidelloidea (Mollusca:
Gastropoda). Bollettino Malacologico, 34: 145-
166.
Biodiversity Journal, 2015, 6 (1): 431-440
Monograph
Additional notes on the systematics and new records of East
Atlantic species of the genus Sorgenfreispira Moroni, 1979
(Gastropoda Mangeliidae)
Paolo Mariottini 1 , Andrea Di Giulio 1 , Carlo Smriglio 1 * & Marco Oliverio 2
1 Dipartimento di Scienze, Universita di “Roma Tre”, Viale Marconi 446, 00146 Rome, Italy; e-mail: paolo.mariottini@uniroma3.it;
andrea.digiulio@uniroma3 .it
2 Dipartimento di Biologia e Biotecnologie “Charles Darwin”, Universita di Roma “La Sapienza”, Viale delLUniversita 32, 00185
Rome, Italy; e-mail: marco.oliverio@uniromal.it
^Corresponding author, e-mail: csmriglio@alice.it
ABSTRACT The Recent species currently ascribed to the Bela brachystoma-complQx, Gastropoda
Mangeliidae, (i.e.: Bela brachystoma (Philippi, 1844); Bela africana Ardovini, 2004; Bela
ardovinii Mariottini et Oliverio, 2008; Bela exilis (Ardovini, 2004) should better be allocated
in the genus Sorgenfreispira Moroni, 1979. Based on numerous samples, the distribution of
the Recent species is summarised. Sorgenfreispira brachystoma (Philippi, 1844) comb. nov.
ranges from Scandinavia to southern Morocco. Sorgenfreispira africana (Ardovini, 2004)
comb. nov. is first recorded from Western Sahara, Ivory Coast, Angola and Ghana; Sorgen-
freispira ardovinii (Mariottini et Oliverio, 2008) comb. nov. is first recorded from Ivory
Coast; S. exilis (Ardovini, 2004) comb. nov. is first recorded from Mauritania, Western
Sahara, Ivory Coast, Angola. Based on the study of the type material, Bela brachystoma
apicalis Nordsieck, 1977, was actually based on specimens of B. taprurensis Pallary, 1904.
Bela taprurensis is here first recorded from Libya.
KEY WORDS Gastropoda; Mangeliidae; Bela; Sorgenfreispira; Recent; first records; new combinations.
Received 12.11.2014; accepted 19.01.2015; printed 30.03.2015
Proceedings of the Eighth Malacological Pontine Meeting, October 4th- 5th, 2014 - San Felice Circeo, Italy
INTRODUCTION
After the revision by Mariottini et al. (2008,
2009), four morphologically similar Recent species
are included in the Bela brachystoma-complex: Bela
brachystoma (Philippi, 1 844), originally described
from the Mediterranean Sea, and three eastern
Atlantic species, namely B. africana Ardovini, 2004,
B. ardovinii Mariottini et Oliverio, 2008 and B. exilis
Ardovini, 2004, all described, and so far known only
from the type locality in Senegal. The species of this
complex have been so far conservatively included
in the genus Bela Gray, 1847, despite their very
peculiar sculpture of both protoconch and teleo-
conch as compared to the other species traditionally
ascribed to Bela [e.g. Bela zonata (Locard, 1892) or
B. menkhorsti van Aartsen, 1988; see Scarponi et al.,
2014]. However, they certainly belong to a morpho-
logically very homogeneous group, to which also
several fossil species belong. Among them the Mio-
cene Cythara ( Mangelia ) moronii Venzo et Pelosio,
1964, is the type species of the genus Sorgenfre-
ispira Moroni, 1979, which was proposed to allocate
those fossills, and is veiy similar to B. exilis. There-
fore, we propose hereby to include Pleurotoma bra-
chystomum Philippi, 1844 and the three eastern
432
Paolo Mariottini etalii
Atlantic species of the complex in Sorgenfreispira.
We have examined numerous samples of this com-
plex from the East Atlantic, mostly at the Museum
National d’Histoire Naturelle (Paris, France), and
summarized the known geographic ranges of each
species, with new records for most species. For
detailed morphological comparisons among the
species see Mariottini et al. (2008).
ABBREVIATIONS AND ACRONYMS. CS-
PM: Carlo Smriglio and Paolo Mariottini collec-
tion (Rome, Italy); lv: live collected specimen(s);
MCZR: Museo Civico di Zoologia di Roma
(Rome, Italy); MMP: Museo Malacologico Piceno
(Cupra Marittima, Italy); MNHN: Museum Na-
tional d’Histoire Naturelle (Paris, France); MO:
Marco Oliverio collection (Rome, Italy); RA:
Roberto Ardovini collection (Rome, Italy); Scan-
ning Electron Microscopy (SEM); sh: empty
shell(s); SMF: Senckenberg Museum (Frankfurt,
Germany); sta: station.
SYSTEMATICS
Sorgenfreispira Moroni, 1979: 2
Type species: Cythara ( Mangelia ) moronii Venzo
et Pelosio, 1964, by original designation
Description. Shell very small for the genus,
height 3.4-3. 8 mm, width 1.5-1. 6 mm, biconical,
turriculate elongate, solid. Protoconch multispiral,
dome shaped, of2.6-2.7 convex whorls. Protoconch-
I (embryonic shell) of 0.4 whorls, separated by a de-
marcation from protoconch-II (larval shell). First
1 .7-1 .8 apical whorls apparently smooth, the nucleus
with very fine striae, the remaining with reticulated
sculpture of 5-6 granulose spirals (3 major, 1-2
smaller subsutural, 1 smaller suprasutural), crossed
by oblique axial riblets. Maximum diameter of proto-
conch 760-780 pm. Protoconch-teleoconch trans-
ition not well marked. Teleoconch of 2.5-3 whorls,
rounded, sutural ramp convex, whorl sides gently
convex. Fast whorl about 3/5 of shell length. Axial
sculpture of 8-9 prominent, narrowly rounded axial
ribs fading out at the base, regularly spaced, with
equally sized interspaces. Spiral sculpture of 17-18
granulose cords, regularly spaces, with larger inter-
spaces. Smaller granulose cordlets in most inter-
spaces. Entire surface covered by microgranules.
Aperture narrow, ovate, about 2/5 of the shell height.
Siphonal canal short, broad and open, very slightly
deviating on the left. Inner lip with a weak parietal
callus. Outer lip not varicose. Anal sinus marked, ar-
cuate on shoulder slope. Colour yellowish with white
axial ribs, darker brown band in the middle of teleo-
conch whorl, base milk white, parietal callus brown.
Remarks. Moroni (1979) introduced this genus
level taxon for a species of the Italian Miocene,
comprising also a group of species of the Jutland
Miocene, that Sorgenfrei (1958) had ascribed to the
genus Neoguraleus Powell, 1939 (type species
Drillia sinclairi Gillies, 882, Recent, New
Zealand): Pleurotoma tenella Mayer, 1858,
Daphnella Calais Kautsky, 1925, and Mangelia
gurichi Kautsky, 1925. Although the actual sys-
tematics of the three latter species may be debated,
Sorgenfreispira moronii is undoubtedly related to
B. exilis. Therefore, we propose hereby the transfer
of B. brachystoma, B. africana, B. ardovinii and B.
exilis to the genus Sorgenfreispira (for the distribu-
tion of this species see Fig. 1).
Sorgenfreispira africana (Ardovini, 2004) comb,
nov. (Figs. 1, 2-5)
Bela brachystoma africana Ardovini, 2004: 7, Fig.
unnumbered.
Type locality. South of Dakar, Senegal. Type Ma-
terial. Holotype (MMP) and 1 paratype (RA).
Examined material. Western Sahara: sta.
12385-3, 22°33.9’N 16°54’E, 54-58 m 8 sh
(MNHN); sta. 12[3]88-3, 22°30.5’N 16°53.8’E, 56-
57 m 3 sh (MNHN); sta. 12381-1, 22°32.2’N
17°04’E, 58 m 42 sh (MNHN).
Mauritania. R/V N’Diago sta. 204, 17°30’N
16°24’W, 88 m 1 sh (MNHN); sta. 218, 17°36’N
16°26’W, 99 m 2 sh (MNHN); sta. 244, 17°54’N
16°32’W, 200 m 1 sh (MNHN); sta. 245, 17°54’N
16°29’W, 145 m 1 sh (MNHN); sta. 289, 18°54’N
16°32’W, 60 m 1 sh (MNHN); 365, 19°30’N
16°55’W, 78 m 1 sh (MNHN); Miss. P. Etienne 1965
sta. 19, 20°20’N 16°22’W, 10 m 5 sh (MNHN).
Senegal. Region of Dakar: Goree, 95 m 4 sh
(MNHN), 95-110 m 1 sh (MNHN), 100 m 9 sh
(MNHN); 14°51’N 17°30’W, 180-165 m 2 sh
(MNHN); 14°32’N 17°25’W, 50 m 11 sh (MNHN);
off Saloum, 50 m 9 sh (MNHN); 30 miles South of
Dakar, Senegal, 45 m, 12 lv in the gut content of
Astropecten cfr. auranciacus (CS-PM coll).
On the systematics and new records of East Atlantic species of the genus Sorgenfreispira (Gastropoda Mangeliidae) 433
Ivory Cost, unknown locality, plateau contin-
ental [no further data], 107 sh (MNHN).
Ghana. R/V Calypsol953 sta. 25, 4°36.5’N
1°31’W, 4 sh (MNHN).
Angola. Corimba, Luanda, 10-20 m, 3 sh (MNHN).
Distribution. Western Sahara, Mauritania,
Senegal, Ivory Coast, Ghana, Angola (Fig. 1).
Remarks. The valid introduction of this taxon
is by Ardovini (2004), although subsequently
(Ardovini, 2008) its author redescribed it as a new
species, having realised that it was worth of species
rank. Sorgenfreispira africana is not uncommon in
East Africa, and it is here first recorded from
Western Sahara, Mauritania, Ivory Coast, Ghana
and Angola.
Sorgenfreispira ardovinii (Mariottini et Oliv-
erio, 2008) comb. nov. (Figs. 1, 6-9)
Bela ardovinii Mariottini et Oliverio, 2008: 8, Figs.
97-99, 102, 119-126, 149-150, 166
Type locality. South of Dakar, Senegal. Type mate-
rial.
Figure 1. Map of the records of Sorgenfreispira africana,
S. ardovinii and S. exilis. The area of their type locality and
of records in Mariottini et al. (2008), off Senegal, is enclosed
in the grey circle.
Holotype (MNHN 21321) and paratype K (MNHN
21322); paratypes A-F (RA); paratypes G-J, L
(CS-PM); paratypes M-N (MO).
Examined material. The type material, all from
30 miles South of Dakar, -45 m: Ivory Cost: un-
known locality, plateau continental [no further
data], 8 sh (MNHN).
Description. Shell very small for the genus,
height 3.7 mm, width 1.4 mm, biconical, turriculate
elongate, solid. Protoconch multispiral, dome
shaped, of 3. 2-3. 3 convex whorls. Protoconch-I
(embryonic shell) of 0.8 whorls, separated by a
demarcation from protoconch-II (larval shell). First
1. 0-1.1 apical whorls densely covered by micro-
granules, next 0.7-0. 8 whorl apparently smooth, the
remaining with reticulated sculpture of 5 spiral
series of tubercles (3 major, 1 smaller subsutural, 1
smaller suprasutural), crossed by weak opisthocline
axial riblets, more evident subsuturally. Spiral cords
corresponding to each spiral series of tubercles
gradually appearing on the last protoconch whorl.
Maximum diameter of protoconch 690-710 pm.
Protoconch-teleoconch transition not well marked.
Teleoconch of 2. 5-2. 7 whorls, rounded, sutural
ramp convex, whorl sides very gently convex. Last
whorl about 3/5 of shell length. Axial sculpture of
8-10 rounded axial ribs fading at base, regularly
spaced, with narrower interspaces. Spiral sculpture
of one major granulose cord, and 20-28 granulose
cordlets, irregularly spaced. Smaller granulose
threads in most interspaces. Entire surface covered
by microgranules. Aperture narrow, ovate, about 2/5
of shell height. Siphonal canal short, broad and
open, slightly deviating to the left. Inner lip with a
weak parietal callus. Outer lip not varicose. Anal
sinus marked, arcuate on shoulder slope. Colour
uniformly reddish-brown.
Distribution. Senegal, Ivory Coast (Fig. 1).
Remarks. Sorgenfreispira ardovinii remains the
least common among the species of this complex.
The 8 shells from Ivory Coast represent a remark-
able range extension for the species, which was
found there syntopic with S. exilis and S. africana.
Sorgenfreispira exilis (Ardovini, 2004)
comb. nov. (Figs. 1, 10-13)
Bela exilis Ardovini, 2004: 8, Figs, unnumbered
434
Paolo Mariottini etalii
Type locality. South of Dakar, Senegal. Type
material. Holotype (MMP); paratypes 1-3 (RA).
Examined material. Western Sahara: sta.
12[3]88-3, 22°30.5’N 16°53.8’E, 56-57 m 1 sh
(MNHN); sta. 12381-1, 22°32.2’N 17°04’E, 58 m
2 sh (MNHN).
Mauritania. R/V N’Diago sta. 229, 17°42’N
16°131’W, 40 m 1 sh (MNHN); sta. 309, 19°06’N
16°31’W, 24 m 1 sh (MNHN).
Senegal. Region of Dakar: 30 miles South of
Dakar, 45 m, (in the gut content of Astropecten cfr.
auranciacus ) 18 lv; Goree, 95 m 9 sh (MNHN); off
Saloum, 50 m 5 sh (MNHN).
Ivory Cost. Unknown locality, plateau contin-
ental [no further data], 38 sh (MNHN).
Angola. Corimba, Luanda, 10-20 m 15 sh
(MNHN).
Description. Shell very small for the genus,
height 3. 4-3. 6 mm, width 1.3-1. 5 mm, biconical,
turriculate elongate, solid. Protoconch multispiral,
dome shaped, of 2. 8-2. 9 convex whorls. Proto-
conch-I (embryonic shell) of 0.7-0. 8 whorls, sep-
arated by a demarcation from protoconch-II (larval
shell). First 1.6-1. 7 apical whorls apparently
smooth, covered with microgranules, the remaining
with reticulated sculpture of 4 granulose spirals (3
major, 1 smaller subsutural), crossed by oblique
axial riblets. Maximum diameter of protoconch
710-720 pm. Protoconch-teleoconch transition not
well marked. Teleoconch of 2.5-3 whorls, rounded,
sutural ramp convex, whorl sides gently convex.
Last whorl about 3/5 of shell length. Axial sculpture
of 10-11 prominent, flexuous and narrowly
rounded axial ribs, regularly spaced, with broader
interspaces. Spiral sculpture of 2 major granulose
cords, with 25-36 irregularly alternating smaller
granulose cordlets and interspaces of variable size.
Each cordlet actually consisting of a rows of
densely packed rounded granules. Aperture narrow,
ovate, about 2/5 of the shell height. Siphonal canal
moderately long, broad and open, deviating on the
left. Inner lip with a moderately developed parietal
callus. Outer lip not varicose. Anal sinus marked,
arcuate on shoulder slope. Colour yellowish-
brownish with two dark brown bands, one subsu-
tural and the second on the middle of the last whorl;
parietal callus brownish with siphonal canal white.
Distribution. Western Sahara, Mauritania,
Senegal, Ivory Coast, Angola (Fig. 1).
Remarks. Present records represent a remark-
able range extension for the species, which was so
far known only from the type locality (Senegal). It
is here first recorded for Western Sahara, Maurit-
ania, Ivory Coast and Angola.
Sorgenfreispira br achy stoma (Philippi, 1844)
comb. nov. (Figs. 14-17)
Pleurotoma brachystomum Philippi, 1844: 169,
176, pi. XXVI, Fig. 10
Type locality. P. brachystomum , Naples, Central
Tyrrhenian Sea, Italy. Type material. Type ma-
terial of Pleurotoma brachystomum is probably
housed in the National Museum of Natural
History (Santiago del Chile).
Examined material. Recent. [Atlantic] France:
Gulf of Gascogne, CAPBRETON 88 sta. DE-01,
43°39.99’N 1°48.11’W, -134 m, 25 sh (MNHN);
idem, sta. DR-29, 43°46.51’N 2°00.58’W, -165
m, 3 sh (MNHN); idem, DR-11, 43°22.77’N
1°59.18’W, -94 m, 16 sh (MNHN); idem, sta. DE-
05, 43°57.42’N 2°05.16’W, -164 m, 8 sh (MNHN);
Arcachon, [no further data], 3 sh (MNHN), 9 sh
(Locard coll., MNHN); Brest, [no further data] 5 sh
(Locard coll., MNHN); Capbreton, [no further
data], 3 sh (Locard coll., MNHN).
Morocco. Agadir, R/V Vanneau 1923-1929 sta.
10, 29°54’N 9°58’W, -110 m, 16 sh (MNHN);
idem, sta. 32, 34°01’N 7°32’W, -145 m, 3 sh
(MNHN); idem, sta. 101, 30°39’N 10°03’W, 129
m, 3 sh (MNHN); sta. 39, 33°44’N 7°45’W, -85 m
1 sh (MNHN); idem, sta. 9, 30°05’N 09°50’W, -110
m 2 sh (MNHN); idem, sta. 30, 33°55’N 7°34’W, -
75 m 17 sh (MNHN); Tangier, 5-10 m, 1 sh
(MNHN). Mauritanie - R/V N.Diago sta. 239,
17°48’N 16°2UW, -79 m,. 1 lv (MNHN); Mission
Gravel 25.03.08 sac 406, 13 sh (MNHN).
Sweden, [no further data], 1 sh (coll. Jousseaume,
MNHN).
England, [no further data], 3 sh (coll Jousseaume,
MNHN).
[Mediterranean] France. Gulf of Lion, IFREMER/
DEPRO 96 (R/V Europe) sta. chalut-10, 42°24.6’N
3°16.2’E, -100/151 m, 40 sh (MNHN); idem, sta.
chalut- 1 1 , 42°09.4’N 3°22.5’E, -350 m, 5 sh
(MNHN). Cap Bear, ECOMARGE 1984 sta. A61,
42°29.30’N 3°10.30’E, -42 m, 5 sh (MNHN); off
Rhone delta, -50/100 m, 2 sh (MNHN). St Raphael,
On the systematics and new records of East Atlantic species of the genus Sorgenfreispira (Gastropoda Mangeliidae) 435
Figures 2, 3. Sorgenfreispira africana. 6 x 1.7 mm, Corimba, Luanda, Angola, West Africa, 10-20 m (MNHN). Figures
4, 5. S. africana. 5.7 x 1.9 mm, Corimba, Luanda, Angola, West Africa, 10-20 m (MNHN). Figures 6, 7. S. ardovinii.
4.6 x 1.7 mm, Corimba, Luanda, Angola, West Africa, 20 m (MNHN). Figures 8, 9. S. ardovinii. 5.5 x 1.6 mm, Corimba,
Luanda, Angola, West Africa, 20 m (MNHN). Figures 10, 11. S. exilis. 4.4 x 1.3 mm, Corimba, Luanda, Angola, West
Africa, 10-20 m (MNHN). Figures 12, 13. S. exilis. 4.5 x 1.4 mm, Corimba, Luanda, Angola, West Africa, 10-20 m
(MNHN).
436
Paolo Mariottini etalii
Scnckfiiberg-Mus 332619 1 2 . Franfcfurt/M
Fehria taprurensis (PalJary 1904)
syntype* Bela brae hystoma apicalis F. Nordsieck I <57 7
Tunesien: Sfax
Slg, F. Nordsieck cx SMF
Figure 14. Drawing of “Bela brachystoma brachy stoma” by Nordsieck (1977: pi. XI, fig. 85). Figs. 15-17. Sorgenfreispira
brachystoma. 5.2 x 1.8 mm, San Vincenzo, Leghorn, Italy, 43°05’N 10°24'E, 34 m (CS-PM) (Fig. 17 SEM photograph). Fig.
18. Drawing of B. brachystoma apicalis by Nordsieck ( 1 977: pi. XI, fig. 86). Figs. 19-21.5. brachystoma apicalis. Syntype A,
SMF33269/2. 5.2 x 1.9 mm, Sfax, Tunisia, 34°47’N 10°53’E, 15 m (SMF) (Fig. 21 SEM photograph). Figs. 22, 23. Details of
the protoconch of Syntype A, SMF33269/2, SEM photographs. Figs. 24, 25. B. brachystoma apicalis. Syntype B, SMF33269/2.
5.2 x 1.9 mm, Sfax, Tunisia, 34°47’N 10°53’E, 15 m (SMF). Fig. 26. SMF label of B. brachystoma apicalis syntypes.
On the systematics and new records of East Atlantic species of the genus Sorgenfreispira (Gastropoda Mangeliidae) 437
Figures 27, 28. Ginannia taprurensis. Lectotype (MNHN-IM-2000-32699), 7.3 mm, Sfax, Tunisia (MNHN). Figures 29,
30. G. taprurensis. 6.6 x 2.2 mm, Sfax, Tunisia (MCZR). Figures 31, 32. G. taprurensis. 6.6 x 2.3 mm, Sfax, Tunisia
(MCZR). Figures 33, 34. G. taprurensis. 6.1 x 2.3 mm, A1 Khums, Libya, 15 m (CS-PM).
[no further data], 3 sh (Couturier coll., MNHN), 4
sh (Locard coll., MNHN). Marseille, [no further
data], 5 sh (Locard coll., MNHN).
Spain. Estepona, 36°25’N 5°09’W, -150 m, 2 sh
(SR); Baleares, [no further data], 1 sh (MNHN);
Alboran, BALGIM sta. 143, 35 0 57’N 3°07’W, -252
m 1 sh (MNHN); Malaga, beach nourishment [from
-20/40 m] 1 sh (MNHN).
Italy. Off San Vincenzo, 43°05’N 10°24’E, -34
m, 78 sh (CS-PM); off S. Marinella, -150/200 m,
sediment in an old Roman dolium, 41°54.00’N,
011°47.66’E, 1 sh (MO); off Fiumicino, 41°43’N
12°06’W, -80 m, 38 sh (CS-PM); off Fiumicino,
41°38’N 12°11’W, -140 m, 27 sh (CS-PM); off
Fiumicino, [no further data] (in the gut content of
Astropecten irregularis ), 16 juveniles (MO); 5 nm
South of Fiumicino, -25 m (in mud), 2 lv (MO); Tor
Patemo shoal, -150 m, 5 lv, lsh (MO); Ponza Is.,
40°51’N 12°55’W, -40 m, 24 sh (CS-PM); off
Civitanova Marche, 43°18’N 13°46’E, -45 m, 11 sh
(CS-PM); off Pescara, 42°31’N 14 0 12’E, -50 m, 24
sh (CS-PM); Sicily, [no further data], 2 sh (coll.
Letellier, MNHN).
Croatia. Brae Island, 43°24’N 16°30’E, -50 m,
5 sh (CS-PM).
Libya. Unknown locality, -110/150 m, 19 sh
(CS-PM).
Fossil. Italy: Guidonia, 42°00’N 12°43’E
(Pliocene), 2 sh (CS-PM); Gallina, 38°05’N
15°41’E (Pliocene), 3 sh (CS-PM); Ficarazzi,
38°04 , N 13°29’E (Upper Pliocene-Lower Pleisto-
cene), 125 sh labelled "P. granuliferum var. parva"
(coll. Monterosato, ex coll. Brugnone, MCZR);
Monte Pellegrino, 38°04’N 13°29’E (Upper
Pliocene-Lower Pleistocene), 48 sh labelled "var.
striiselevatioribus" (coll. Monterosato, ex coll.
Brugnone, MCZR).
Description. Shell small for the genus, height
4.5-7 mm, width 1.7-2. 5 mm, biconical, turricu-
late elongate, solid. Protoconch multispiral, dome
shaped, of 2. 3-2. 4 convex whorls. First 1.6-1. 9
apical whorls smooth, the remaining with reticu-
438
Paolo Mariottini etalii
lated sculpture of 4-5 granulose spirals (3 major
spirals in the middle of whorl, 1 smaller subsutural
and 1 smaller above the teleoconch suture) crossed
by oblique axial riblets. Maximum diameter of
protoconch 510-650 pm. Protoconch-teleoconch
transition not well marked. Teleoconch of 5-6
whorls, rounded, sutural ramp straight or very
slightly convex, whorl sides gently convex. Last
whorl about 2/5 of shell length. Axial sculpture of
8-9 prominent, slightly opisthocline, flexuous and
narrowly rounded axial ribs, regularly spaced, with
broader interspaces. Spiral sculpture of 9-15 major
cordlets, with irregularly alternating smaller
cordlets and interspaces of variable size. Each
cordlet consists of a rows of densely packed roun-
ded granules. Aperture narrow, ovate, about 1/3 of
the shell height. Siphonal canal short, narrow and
open, deviating on the left. Inner lip with a mod-
erately developed parietal callus. Outer lip varicose.
Anal sinus marked, arcuate on shoulder slope.
Animal with short head and two short tentacles.
Black eyes on the external, thickened basal part of
the tentacles, located on the distal third of their
total height. Foot broad and long, slightly lobate
anteriorly, tapering posteriorly. Background colour
of the head-foot pinkish, semi-transparent, with
light yellow spots, and light yellow speckles on the
proximal part of the tentacles. Siphon pinkish,
semi-transparent, with light yellow spots bordered
by orange.
Distribution. Sorgenfreispira brachystoma is
known from the northeastern Atlantic and from the
entire Mediterranean Sea. Based on literature data
and on the material we have examined, it ranges
from Norway (Hoisseter, 2009), Sweden (Dyntaxa,
2013), United Kingdom and British Isles (Hayward
& Ryland, 1990), to southern Morocco (Lat 34° N),
and the entire Mediterranean Sea. Fossil shells are
known from several Plio-Pleistocene European
outcrops (England, France, Spain, Italy: see Chirli
& Richard, 2008).
Remarks. Sorgenfreispira brachystoma is a
continental shelf species, easily distinguishable
from all other members of the group by its very
distinct shell sculpture (Mariottini et al., 2008). It
has a multispiral protoconch with characteristic
densely granulated spiral ribs. Bela brachystoma
apicalis Nordsieck, 1977 is a synonym of B.
taprurensis (Pallary, 1904) (see below).
Bela Gray, 1847: 270
Type species: Murex nebula Montagu, 1803, by
subsequent designation (Gray, 1847).
=Fehria van Aartsen, 1988 (type species: Ginnania
taprurensis Pallary, 1904, by original designa-
tion)
Bela taprurensis (Pallary, 1904) (Figs. 18-46)
Ginnania taprurensis Pallary, 1904: 218, pi. VII,
Fig. 1
Bela brachystoma apicalis Nordsieck, 1977: 44, pi.
11, Fig. 86
Type locality. Ginnania taprurensis Pallary: Sfax,
Tunisia, Mediterranean Sea. Bela brachystoma
apicalis Nordsieck: Sfax, Tunisia, Mediterranean
Sea. Type material. Ginnania taprurensis Pallary:
Lectotype (MNHN-IM-2000-3269).
Bela brachystoma apicalis Nordsieck: 2 syntypes
(SMF33269/2)
Examined material. The type material and:
Tunisia: Sfax, 34°47’N 10°53’E, 15 m, 25 sh
(CS-PM), 2 sh (coll. Monterosato, MCZR).
Libya: Al Khums, 32°43’N 14°18’E, 15 m, 4 sh
(CS-PM).
Distribution. Southern Mediterranean Sea,
Gulf of Gabes (Pallary, 1 904) and Libya; Aegean
Sea (Manousis, 2012: 169) and Levant Sea (Bogi
et al., 1989).
Remarks. Nordsieck (1977: 45, pi. XII, fig.
90) redescribed Ginnania taprurensis Pallary and
depicted a shell from Karpathos (Greece). Proto-
conch (2 convex whorls, rather blunt) and teleo-
conch description match the species as
represented by the lectotype (Figs. 27, 28) and the
two specimens in coll. Monterosato, presumably
ex Pallary (Figs. 29-32). Nordsieck (1977) also
described B. brachystoma apicalis, differing from
the nominal species mainly for its paucispiral
“protoconch [of] 11/2 very inflated whorls, which
leads to the conclusion of a quite other life of
larves” (Nordsieck, 1977: 44, pi. 11, Fig. 86). The
examination of two syntypes (Figs. 19-23) of B.
brachystoma apicalis revealed that this taxon was
actually based on shells of B. taprurensis (Pallary,
1904) (Figs. 27-46). Present sample from Libya
(Figs. 33, 34) is the first record for the waters of
that country.
On the systematics and new records of East Atlantic species of the genus Sorgenfreispira (Gastropoda Mangeliidae) 439
Figures 35-38. Bela taprurensis. 4.8 x 1.8 mm, Sfax, Tunisia, 34°47’N 10°53’E, 15 m (CS-PM) (Figs. 37, 38 SEM
photographs). Figures 39-46. Details of the shell of figs. 35-38 (SEM photographs).
440
Paolo Mariottini etalii
ACKNOWLEDGMENTS
We would like to express our gratitude to Vir-
ginie Heros and Philippe Maestrati (MNHN),
Ronald Janssen (SMF) and Massimo Appolloni
(MCZR) for the help in the study of the materials
under their care, and Cesare Tabanelli with Cesare
Bogi for the bibliographic assistance.
REFERENCES
Ardovini R., 2004. Due nuove specie e una nuova sotto-
specie di Turridae dal Senegal, West Africa. Malaco-
logia, 43: 7-9.
Ardovini R., 2008. Bela africana n. sp. (Gastropoda
Turridae) West Africa, Senegal. Malacologia, 58: 12-
13.
Bogi C., Cianfanelli S. & Talenti E., 1989. Contributo
alia conoscenza della malacofauna dell'isola di Cipro.
Atti prima giornata Studi Malacologici C.I.S.Ma.,
187-214.
Chirli C. & Richard C., 2008. Les mollusques plaisan-
ciens de la Cote d’Azur. Devaye Imprimeurs, Z.I. la
Frayere, Cannes, 128 pp.
Dyntaxa, 2013. Swedish Taxonomic Database. Ac-
cessed at www.dyntaxa.se [5-02-2015], at http://www.
dyntaxa.se
Gray J.E. 1847. A list of the genera of recent Mollusca,
their synonymia and types. Proceedings of the Zo-
ological Society of London, 15, 129-206.
Hayward P.J. & Ryland J.S. (Eds.), 1990. The marine
fauna of the British Isles and North-West Europe: 1.
Introduction and protozoans to arthropods. Clarendon
Press, Oxford, 627 pp.
Hoisaster T., 2009. Distribution of marine, benthic, shell
bearing gastropods along the Norwegian coast. Fauna
norvegica, 28: 5-106.
Manousis T., 2012. The seashells of Greece. Publishing
House Kyriakidis Brothers S.A., Thessaloniki, 381 pp.
Mariottini P., Di Giulio A., Smriglio C. & Oliverio M.,
2008. Notes on the Bela brachystoma complex, with
description of a new species (Mollusca, Gastropoda:
Conidae). Aldrovandia, 4: 3-20.
Mariottini P., Smriglio C., Di Giulio A. & Oliverio M.,
2009. A new fossil conoidean from the pliocene of
italy, with comments on the Bela menkhorsti complex
(Gastropoda; Conidae). Journal of Conchology, 40:
5-14.
Moroni M.A., 1979. Sorgenfreispira , nuovo genere di
Turridae (Gastropoda, Prosobranchia) del Miocene
europeo. Lavori dell’Istituto di Geologia della
Universita di Palermo, 16: 1-11.
Nordsieck F., 1977. The Turridae of the European Seas.
La Conchiglia Ed., Roma, 131 pp.
Pallary P., 1904. Addition a la faune malacologique du
Golfe de Gabes. Journal de Conchyliologie, 52: 2 12—
248, pi. 7.
Scarponi D., Landau B., Janssen R., Morgenroth H. &
Della Bella G., 2014. Lectotype designation for
Murex nebula Montagu 1803 (Mangeliidae) and its
implications for Bela Leach in Gray 1847. Zootaxa,
3884: 045-054
Sorgenfrei Th., 1958. Molluscan assemblages from the
marine Middle Miocene of South Jutland and their
environments. Danmarks Geologiske Undersogelse,
Series 2, 79: vol. 1 1-355, text figs. 1-7; vol. 2 356—
503, pis. 1-76. Kobenhaven.
Biodiversity Journal, 2015, 6 (1): 441-448
Monograph
Description of a new species of the genus Trophonopsis Bucquoy
et Dautzenberg, 1 882 (Gastropoda Muricidae Pagodulinae)
from the Mediterranean Sea
Carlo Smriglio*, Paolo Mariottini & Andrea Di Giulio
1 D ip artim e n to di Scienze, Universita di “Roma Tre”, Viale Marconi 446. 00146 Rome. Italy; e-mail: csmriglio@alice.it; paolo.
mariottini@ uniroma3.it; andrea.digiulio@ unirom a3.it
Corresponding author
ABSTRACT Based on shell characters, a new species of the gastropod family Muricidae, Trophonopsis
spciracioi n. sp., from Mediterranean Sea is described. Shells of the new taxon were collected
from bathyal bottoms, in the Tyrrhenian Sea. The new taxon is compared with others species
of the genus Trophonopsis Bucquoy et Dautzenberg, 1882, occurring in northeastern Atlantic
and Mediterranean Sea.
KEYWORDS Trophonopsis spciracioi n . sp.; M uricidae; Pagodulinae; M editerranean Sea.
Received 17.0 9.2014; accepted 12.03.2 0 15; printed 30.03.2015
Proceedings of the Eighth Malacological Pontine Meeting, October 4th- 5th, 2014 - San Felice Circeo, Italy
INTRODUCTION
The genus Trophonopsis B u c q u o y et Dautzen-
berg, 1 8 82 has been traditionally included in the
subfamily Trophoninae (Muricidae), while recently
Barco et al. (2012) have included this genus in their
newly erected subfamily Pagodulinae, based on
clear evidence from the radular morphology.
Five Recent species of the genus TrophoYlopsis
were so far recognised in northeastern Atlantic and
Mediterranean Sea according to Houart(2001) and
CLEMAM (G o fas & Le Renard, 2014): T. alboran-
ensis (Smriglio e al., 1 997) (Figs. 1-5, 39, 40), T.
barvicensis (Johnston, 1825) (Figs. 6-10, 41, 42),
T. breviatus ( J e ffre y s , 1882 ) (Figs. 11-15,43,44),
T. droueti (Dautzenberg, 1 8 8 9 ) and T. muricatUS
(Montagu, 1803) (Figs. 16-20, 45, 46). These taxa
have been reviewed by H o u a r t (2001), who con-
sidered T. alboranensis in the original genus
Houartiella S m rig lio , M ario ttin i et B o n fitto , 1 9 9 7,
later synonym ized with Trophonopsis by Penas et.
al. (2006). In particular, two Trophonopsis species
have an Atlantic distribution, being T. droueti
endemic to the bathyal bottoms of the Azores
(Bouchet & Waren, 1 985; Houart, 2001 ), while T.
barvicensis , which occurs at 50-1.000 m depth, is
distributed from Morocco and the Azores, to the
British Isles and West Scandinavia (Houart, 200 1;
Segers et al., 2009 ). Trophonopsis barvicensis w a s
recently reported in the Mediterranean Sea from El
Garraf (Spain), as a W iirmian fossil, by Giribet &
Penas (1997) and from the Djibuti bank (Spain),
which could represent the extreme limit of its
distribution into the M editerranean, by Gofas et al.
( 2011 ). Furthermore, T. alboranensis and T. brevi-
atUS are endemic to A lb ora n and Black Sea, respect-
ively (Smriglio et al., 1997; Houart, 2001; Gofas et
al., 20 11). Trophonopsis breviatus has been also re-
corded from (Tanakkale and Bozcaada Isle, Turkey
(Panayotis O valis, pers. comm.). Only T. muricatUS
442
Carlo Smriglio etalii
displays both a wide distribution, occurring in the
Mediterranean and in the northeastern Atlantic up
to the northern Great Britain, and a wide batimetric
range (0.5-300 m) (Rolan, 1983; Houart, 2001;
Gofas et al., 2011).
Recently, we had the chance to examine shells
of Trophonopsis from a spot located in the Tyrrhe-
nian Sea, inhabited by rich bathyal benthic inver-
tebrate communities (molluscan assemblages have
been partially characterized in the past by Smriglio
et al., 1 989; Smriglio & Mariottini, 1 996, 2000,
200 1; Smriglio et al., 1 999). After morphological
comparison with the species of Trophonopsis
occurring in northeastern Atlantic and Mediter-
ranean, the studied shells have been regarded as
belonging to a distinct, unnamed species, which is
here described as new to science: Trophonopsis
sparacioi n. sp.
ABBREVIATIONS AND ACRONYMS. The
materials used for this study are deposited in the
following private and Museum collections, BA:
Bruno Arnati collection, Rome, Italy; CS-PM : Carlo
Smriglio-Paolo Mariottini collection, Rome, Italy;
H: height; MO: Marco Oliverio collection, Rome,
Italy; MTC: Monterosato collection; MCZR:
Museum of Zoology of Rome (section collections
of Malacology); MNHN: Museum National
d’Histoire N aturelle, Paris, France; M ZB : M useum
of Zoology Bologna (collection of the Laboratory
of Malacology, University of Bologna, Italy); sh:
empty shell(s); W : width.
MATERIAL AND METHODS
Samples consisted mainly of empty shells, in a
few cases with dried softparts, from CS-PM private
collection and material stored in the MTC at the
MCZR.
Sediment sampling was collected by fishermen
trawlers from m u d d y - b a th y a 1 bottoms located off
the coasts of Latiurn (C entral Tyrrhenian Sea). Sedi-
ment samples were sieved through a 1 mm mesh
and sorted under a s te r e o m ic r o s c o p e . Scanning
Electron Microscopy (SEM) observations were car-
ried out by a Philips XL30 at the Interdepartmental
Laboratory of Elec tr on Microscopy (LIM E, U niver-
sity of “Roma Tre”, Rome, Italy). Current system a-
tics is based on WoRMS (2013), that for
Trophonopsis species treated in this work is in
accordance with CLEMAM (Gofas & Le Renard,
2014). Sculpture of the teleoconch was described
according to the notation ofMerle (2001, 2005).
SYSTEMATICS
Family MURICIDEA Rafinesque, 1815
Subfamily Pagodulinae B arc o , Schiaparelli, Houart
et O liv erio ,2012
type genus Pogodlilci Monterosato, 1 884 (by ori-
ginal designation)
Genus Trophonopsis Bucquoy, Dautzenberg et
D ollfus, 1882
type species Mlirex nturicCltllS M on tag u, 1 8 03 (by
original designation)
Trophonopsis sparacioi n . s p .
Examined material. The type material (Figs. 21-
33, 47, 48) consists of 100 shells, 27 of them with
dry soft parts, from the Central Tyrrhenian Sea, off
coasts of Latium, 500/600 m (4 1 °5 1 ’ N 1 1 ° 2 8 ’ E ) .
Holotype, MNHN IM-2000-27897; paratype 1,
MNHN IM -2000-27898; paratypes 2, MZB 60093
and 3, M ZB60094; paratypes 4, M CZR00222a and
5, M CZR00222b; paratypes 6 and 7, M O ; par a types
8 and 9,BA;paratypes 10-99, CS-PM.
Other examined material. Trophonopsis Cllbor-
ancnsis: from CS-PM collection (Rom e): paratypes
“A-B-D” and 13 sh,Alboran Sea (type locality), 80-
150 m; 1 sh,Alboran Island, 180 m.
Trophonopsis barvicensis-. from MTC collec-
tion: 2 sh, Bergen, Norway; 2 sh, Oban, Scotland,
25 fathoms; 4 sh. She tl and , England; 3 sh, England;
42 sh,NorthAtlantic Ocean, 226 m;60 sh. Paler mo,
Italy. From CS-PM collection (Rome): 4 sh, Aber-
deen Bank (57°13’N 0 1 ° 0 5 ’ W ) , Scotland, 59-68 m.
Trophonopsis breviatus-. f r o m cs-pm collection
(Rome): 2 sh Bozcaada Island, Turkey 85 m; 4 sh
from Marmara Island, Marmara Sea.
Trophonopsis muricatUS: from MTC collection:
2 sh, Northumberland coast, Scotland; 5 sh,
England; 10 sh,Le Croisic (Saint-Naza ire), France;
4 sh, Villefranche sur M er (Nige), France; 2 sh,
Minorca, Baleares, Spain; 3 sh, Corsica; 4 sh,
Sardinia, Italy; 4 sh, Positano (Naples), Italy; 3 sh,
Golfo di Napoli, Italy; 1 sh, Naples, Italy; 192 sh,
Palermo, Italy 1 sh, Algeria; 15 sh, Ficarazzi
Description of a new species of the genus Trophonopsis (Gastropoda Muricidae) from the Mediterranean Sea 443
(Palermo), Italy, fossil; 3 sh, Ficarazzi (Palermo),
Italy, fossil; 3 sh, Giannettilla (C altanissetta), Italy;
3 sh,Babbaurra(Caltanissetta),Italy; 1 sh,Magnisi
(Siracusa), Italy; 3 sh, Sciacca (Agrigento), Italy; 1
sh, Morocco. From CS-PM collection (Rome): 3 sh,
Algeciras, Spain, 20-35 m; 4 sh, Capo Corso,
Corsica, France, 70 m; 8 sh, Capraia Island
(Leghorn), Italy, 80-200 m ; 1 sh, Capraia Island
(Leghorn), Italy, 400 m; 14 sh, Elba Island,
Tuscany, Italy, 300 m ; 3 sh, Elba Island (Leghorn),
Italy, 50 m; 2 sh, Civitavecchia (Rome), Italy, 40
m; 3 sh,Fiumicino (Rome), Italy, 160 m; 8 sh,Circeo
(Latina), Italy, 90 m; 108 sh, Ponza Island (Latina),
Italy, 125-165 m; 1 sh, C apo Portiere (La tina), Italy,
unspecified depth; 3 sh, Golf of Carini (Palermo),
Italy, 120 m ; 48 sh, 60 miles offshore Sfax, Tunisia,
100 m; 11 sh, Libyan coasts, 110-150 m .
Pagodula echinata (Kiener, 1 8 4 0 ) : from CS-
PM collection (Rome): about 600 sh, offshore
Fiumicino (Rome), Central Tyrrhenian Sea
(4 1 °5 1 ’N 1 1 °2 8 ’E), Italy, 5 00-600 m .
Description of the holotype. Shell of small size
forthe genus, H = 5.6, W = 2.9 mm, fusiform, elong-
ate, with high spire and siphonal canal open and
moderately long, last whorl about three quarter of
entire shell length. Protoconch paucispiral, with a
diameter of 5 80 pm and 1.5 rounded whorls, orna-
mented with narrow, irregular spiral threads. Teleo-
conch with 3.5 whorls, axial sculpture consisting of
11 lamellate ribs, slightly spiny at the shoulder.
Infrasutural ramp without cords (cords 1 and 2
absent), convex part of the last w horl w ith 6 prim ary
cords (cords 3-8).Aperture small, ovate with a thin,
knife-edge outer lip, to some extent undulate.
Columellar lip narrow, smooth and adherent.
Siphonal canal narrow, with evident growth ridges.
Shell uniformly white or g re y is h - w h ite , vitreous.
Operculum corneous, ovoid, planispiral with lateral
nucleus.
Variability. Shell height ranging from 5.8 to 6.3
mm with an average of 6.10 mm (50 sh measured),
while the width is ranging from 2.9 to 3.1 mm (50
sh measured), with an average of 2.95 mm. Proto-
conch diameter from 550 to 650 pm, with an
average of 589 mm (11 sh measured by SEM
analyses). Teleoconch always comprising 3.5
whorls, which can be considered as a diagnostic
character, and 6 (rarely 7) primary cords in the
convex part of the last whorl. Number of lamellate
ribs of the axial sculpture ranging from 1 1 to 14,
with an average of 12.24.
Etimology. This species is dedicated to Ignazio
Sparacio (Paler mo, Italy), for his great contribution
to scientific research and his editorial work for the
biodiversity of the Mediterranean region.
Distribution and biology. Locus typicus: Central
Tyrrhenian Sea off the coasts of Latiurn (4 1 0 5 1 ’ N
11°28’E). Habitat: Biocoenosis CB (sensu Peres &
Picard, 1964), 360-600 m depth.
Remarks. All shells of T. spcircidoi n. sp. (Figs.
21-33, 47, 48), some of them with soft parts, were
sorted out from sediment samples collected at
bathyal depths. In particular, empty shells from
muddy bottoms (biocoenosis VB, sensu Peres &
Picard, 1 964), while shells with soft parts from
deep-sea coral banks (biocoenosis CB, sensu Peres
& Picard, 1 964). The benthic communities of this
Tyrrhenian Sea area have been investigated since
the late eighties, and have been partly characterized
(Smriglio etal., 1989; Smriglio & Mariottini, 1996,
2000, 200 1; Smriglio et al., 1999). The analysis of
the accompanying dredged organogenic sediment
revealed many fragments of alive a zo o x an te Hate
corals like DeSlflOphyllum cristagalli Milne
Edwards etHaime, 1848 and MadvcpOVCl OCUlatO.
(Linne, 1 758), indicating that this species belongs
to the biocoenosis CB. Furthermore, together with
the empty shells of the new taxon, we collected
another abundant pagoduline, Pdgodulci echinata
(Kiener, 1 840) (Figs. 34-38, 49, 50), which typic-
ally inhabits c ir c alitto r a 1/b a th y a 1 muddy bottoms
(Smriglio et al., 1 989; Houart, 200 1; Gofas et al.,
2011 ). For P. echinata , it is interesting to recall that
the fossil P. Vaginata (De Cristofori et Ian, 1 832)
differs by presenting “a distinct difference in the
larval shell, which in the recent form consist of
about 1.5 whorls while the Pliocene form has more
than 2.5 whorls, and possibly planktotrophic larval
development” (Bouchet & Waren, 1985 p. 138).
Moreover, La Perna (1996) remarked “at that time,
the two species lived in ecologically segregated
populations, P. Vaginata b ein g linked to deep-shelf
and upper- slope bottoms, and P. echinata to deeper
bathyal bottom s”. Possibly, this difference in habitat
has enabled P. echinata to be protected during the
Quaternary climatic cooling, which instead caused
444
Carlo Smriglio etalii
Figures 1-5. Twphonopsis ulboVUVLCnsis . ParatypeD,Fl 4.7 xW 2.8 mm. Alboran Sea, 80-150 m depth. CS-PM. Figures
6-10. T. barvicensis. H 7. 4 x W 3.7 mm. Aberdeen Bank, E. Scotland, 59-68 m depth, 57°13'N-01°05'W. CS-PM.
Figures 11, 12. T. bvevitttUS . H 7.6 x W 4.2 mm. Marmara Island, Marmara Sea. CS-PM. Figuresl3-15. T. bvevicitUS.
H 9.5 x W 5.2 mm. Bozcaada Island, Turkey, 85 m depth. CS-PM. Figures 16-20. T. niUVicdtUS . H 7.6 x W. 3.7 mm.
Circeo, Italy, 90 m depth. CS-PM .
Description of a new species of the genus Trophonopsis (Gastropoda Muricidae) from the Mediterranean Sea 445
Figures 2 1- 25. Tvophoviopsis Spavadoi n. sp. Paratype 10, H 6.2 x W 3.3 mm. Central Tyrrhenian Sea. CS-PM. Figures 26,
2 7.7] Spavadoi n. sp. Paratype 2, FI 5.8 x W 2.9 mm. Central Ty rrh en ian Sea. MZB60093. Figures 28, 29. T. Spavadoi n . sp .
Holotype,H 5.6 x W 2.9 mm. Central Tyrrhenian Sea. MNHN IM-2000-27897.Figures30,31.7] Spavadoi n . sp. Paratype 4,
H4.5xW 2.9 mm.Cen tral Ty rrh en ian Sea. M CZR00222a. Figures 32, 33. T. Spavadoi n . sp. Paratype 11, FI 6.1 xW 2.8 mm.
CentralTyrrhenian Sea. CS-PM . Figures 34-38. Pagodllla Cchifiata. FI 10.7 x W 5.4 m m ; C entral Tyrrhenian Sea. CS-PM .
446
Carlo Smriglio etalii
Figures 39, 40. T. alboranensis . Same as Fig. 1. Figures 41, 42. T. barvicensis. S ame as Fig. 6. Figures 43, 44. T.
breviatus. Same as Fig. 11. Figures 45, 46. T. YYlUriCdtUS . Same as Fig. 16. Figures 47, 48. T. SpUYUCioi n . sp . Same
as Fig. 34. Figures 49, 50. Pagodllla echinata. S ame as Fig. 11.
the extinction of P. VClginCltd. We think that P. Vd~
gillCltCl represents the sister species of P. echinCltd.
that has lost the planktotrophic larval stage showing
a different successful adaptive strategy, as described
for other Recent couples of sibling species (Pusateri
et al., 2012; Pusateri et al., 2013). Trophonopsis
sparacioi n. sp. clearly differs from all other
Trophonopsis occurring in Northeastern Atlantic
and Mediterranean Sea mainly for its small size and
shell sculpture. Only T. dlbOYCinensis has s i m i 1 a r
Description of a new species of the genus Trophonopsis (Gastropoda Muricidae) from the Mediterranean Sea
447
dimensions, but this species shows a totally dif-
ferent shell sculpture consisting of nodulose axial
ribs with a higher number of spiral cords (compare
Figs. 1-5 and 39, 40 to Figs. 21-25 and 47, 48)
(Smriglio et al., 1997; Houart, 200 1; Gofas et al.,
2011 ). Trophonopsis barvicensis possesses both
protoconch and teleoconch of bigger size and its
shell sculpture shows less axial ribs and spiral
cords, also less spiny at intersections (compare
Figs. 6-10 and 41, 42 to Figs. 21-25 and 47, 48).
Trophonopsis sparacioi n . sp. clearly differs from
T. breviatus in many respects, the latter having a
more convex shell outline, being less sculptured,
and having a bigger protoconch (compare Figs.
11-15 and 43, 44 to Figs. 21-25 and 47, 48).
Trophonopsis muricatus is bigger in size, shows a
more convex shell outline, a more reticulated
sculpture, less spiny at the intersections, and a
longer siphonal canal. The protoconch of this species
is similar in size, but shows less spiral threads,
coarser in the last part of the whorl (compare Figs.
16-20 and 45, 46 to Figs 21-25 and 47,48).
ACKNOWLEDGMENTS
We would like to express our gratitude to
Massimo Appolloni (MCZR) for the examination
of the Pagodulinae material stored in the MTC.
Sincere acknowledgments are due to Marco Oliv-
erio (University of Rome “La Sapienza”, Rome,
Italy) and RafaelLa Perna (University ofBari, Bari,
Italy) for their critical suggestions. We would like
to thank the reviewers for their helpful comments
and the resulting improvements in the paper.
REFERENCES
Barco A, Schiaparelli S., Houart R. & Oliverio M . 2012.
Cenozoic evolution of Muricidae (Mollusca, Neo-
gastropoda) in the Southern Ocean, with the de-
scription of a new subfamily. Zoologica Scripta 41:
596-6 16.
Bouchet P. & W are n A., 1985. Revision of the Northeast
Atlantic bathyal and abyssal N e o g a s tro p o d a exclud-
ing Turridae (Mollusca, Gastropoda). Bollettino
M alacologico, Supplemento 1: 1 23-296 pp.
Giribet G . & Penas A., 1997. M alacological marine fauna
from Garraf coast (NE Iberian Peninsula). Iberus, 15:
41-93.
Gofas S. & Le Renard J., 2014. CLEMAM: Check List
of European Marine Mollusca. Available at
h tip :// w w w .som ali.asso.fr/clem am / index, clem am .
h tm 1 . A c c e s se d 2014-oct-01.
Gofas S., Moreno D. & Salas C ., 2011. Moluscos m ari-
nos de Andalucia - I. Universidad de Malaga, Junta
de Andalucia, Consejeria de medio ambiente, 342 pp.
Houart R ., 200 1. A review of the Recent Mediterranean
and Northeastern Atlantic Species of Muricidae.
Evolver. Roma, 227 pp.
La Perna R., 1 996. Phyletic relationships and ecological
implications between P dgoduld VdgUldtd (De Cristo-
fori & Jan, 1 83 2) and Pcigodlllci ecilUldtd (Kiener.
1840) (Gastropoda, M uricidae). Bollettino della
Societa Paleontologica Italiana, 35: 8 1 -92.
Merle D., 2001. The spiral cords and the internal
denticles of the outer lip in the Muricidae: termin-
ology and methodological comments. Novapex, 2:
69-91.
M erle D ., 2005. The spiral cords of the M uricidae (Gast-
ropoda, N e o g a s tro p o d a ) : importance of ontogenetic
and topological correspondences for delineating
structural homologies. Lethaia, 3 8: 367-3 79.
Peres J. M. & Picard J., 1 964. Nouveau Manuel de
Bionomie Benthique de la Mer Mediterranee. Recueil
des Travaux de la Station Marine d'Endoume, 31: 1 -
137.
Pusateri F., Giannuzzi-Savelli R. & Oliverio M ., 2012. A
revision of the Mediterranean Raphitomidae 1: on the
sibling species RciphitomCl COlltiglld Monterosato,
1 8 84 and RaphitOniCl SpCldianCt n . sp. (Gastropoda,
Co no idea). Iberus, 30: 41-52.
Pusateri F., Giannuzzi-Savelli R. & Oliverio M ., 2013. A
revision of the Mediterranean Raphitomidae 2: on the
sibling species RciphitomCl Ulieoldtd (B D D . 1 883) and
Raphitoma smriglioi n . sp. (Gastropoda, Conoidea).
Iberus, 3 1 : 11 -20 .
Penas A., Rolan E., Luque A .A ., Templado J., Moreno
D ., Rubio F., Salas C ., Sierra A. & Gofas S ., 2006.
Moluscos marinos de la isla de Alboran. Iberus, 24:
23-151.
Rolan E., 1983. Moluscos de la Ria de Vigo, 1. Gastro-
podos. Privately published, Vigo, 3 83 pp.
Segers W., Swinnen F. & de Prins R., 2009. Marine mol-
luscs of Madeira. Snoeck publishers, 612 pp.
Smriglio C. & Mariottini P., 1996. Molluschi del Mar
Tirreno Centrale: Contributo XII. Descrizione di una
nuova specie di Cystiscidae Stimpson, 1 865, per
il Mar Mediterraneo: Grdnulitld gofcisi n. sp. La
Conchiglia, 28 1: 54-56.
Smriglio C. & M ariottini P., 2000. Oliobd oliverioi n . s p .
(P ro sob ran c h ia , Rissoidae), a new gastropod from the
Mediterranean. Iberus, 18: 15-19.
Smriglio C. & Mariottini P., 2001 . Emcirginidci bonflttoi
sp. nov. (Gastropoda, Pro sobranchia, F is s ure Hid ae ) ,
448
Carlo Smriglio etalii
a new bathyal species from the Mediterranean Sea.
Basteria, 65: 1 39-143.
Smriglio C ., Mariottini P. & Bonfitto A., 1997. Descrip-
tion of Houartiella n. gen., Trophoninae Cossmann,
1 903, and Houartiella alboranensis n. sp. from the
Mediterranean Sea. Bollettino Malacologico, 32:
27-34.
Smriglio C ., Mariottini P. & Calascibetta S., 1999. De-
scription of a new species of Conidae Fleming, 1822
from the Mediterranean Sea: CoilOplcUTQ, ttlidltt n.
sp. Bollettino Malacologico, 34: 27-32.
Smriglio C., Mariottini P. & Gravina F., 1989. Molluschi
del M ar Tirreno Centrale: ritrovamento di PutZ6ysicl
wiseri (Calcara, 1 842), IschflOChitOfl vanbellei Kaas,
1 9 85 e NeOpililVl zogvafi (D autzenberg & Fischer.
1 896). Contributo VI. Bollettino Malacologico, 25:
125-1 32.
WoRMS Editorial Board, 2013. World Register of
Marine Species. Available from h ttp ://w w w .m arin e
species.org at VLIZ. Accessed 10 September 2014.
Biodiversity Journal, 2015, 6 (1): 449-466
Monograph
The family Cypraeidae (Gastropoda Cypraeoidea) an unex-
pected case of neglected animals
Marco Passamonti
Dipartimento di Scienze Biologiche Geologiche e Ambientali (BiGeA), Via Selmi 3, 40126 Bologna, Italy; e-mail: marco.
passant onti@ unibo.it
ABSTRACT The family Cypraeidae Rafinesque, 1815 (Gastropoda Cypraeoidea), common ly called
Cowries, are particularly well-known among shell collectors, because of their beauty and
relative availability. W hile most species are common in shallow reef environments, some other
are quite hard to find, because they may come from remote or hardly accessible habitats, or
they are in fact just rarely found. Because of this rarity and beauty, several cowries get high
market values among collectable shells. This relevant economic interest produced two kind of
outcomes: a proliferation of taxonomic complexity, and a very detailed knowledge of every
variation of a given species, making cowries collection one of the most specialized ones.
Notwithstanding this, it is quite remarkable that cowries had attracted very little interest by
biologists and professional m alacologists. Few scientific studies are available to date. This
review attempts to overview some of the major biological highlights of the Family, to pro mote
future researches in this diverse group of gastropods.
KEYWORDS Cypraeid ae; Evolution; Biogeography; Speciation.
Received 18.02.2015; ac c ep ted 1 4 .0 3 .2 0 1 5 ; printed 30.03.2015
Proceedings of the Eighth Malacological Pontine Meeting, October 4th- 5th, 2014 - San Felice Circeo, Italy
INTRODUCTION
The family Cypraeidae Rafinesque, 1815,
cowries (Gastropoda Cypraeoidea) comprises about
220 species of marine gastropods (but this figure
may vary with different taxonomies, see f.i. Moretz-
sohn, 2014), widespread along the tropical and sub-
tropical seas. Many species are commonly found in
tropical shallow water reefs, although others are
adapted to temperate waters and/or deep water en-
vironments. Most species are herbivorous grazers,
but some are carnivorous, being more commonly
sponge eaters.
The main characteristic of cowries (which is
however shared by some other gastropods) is the
presence of a retractable mantle that covers the
entire shell, when in full extension (Figs. 1-6). This
makes the cowrie shell particularly shiny, because,
at variance to most gastropods, shell layers are
continuously deposited outside the shell itself, rather
than in the aperture and inside. The mantle is seldom
richly branched, with protrusions known as papillae
(Figs. 1, 2, 4-6), that may have both respiratory and
mimetic functions. When disturbed, the animal can
quickly retract the mantle showing off the brilliant
shell.
The amazing richness of shell and mollusk colors
in cowries (as in many other mollusks as well) has
been always an evolutionary puzzle.
450
Marco Passamonti
Cowries are not toxic animals, as far as we
know, so the brilliant colors of shells and mantles
could not be considered as warning signals for
predators (apo sem atism ). Sometimes, the mantle,
when fully expanded, may camouflage the shell
itself (see f.i. the genus Novia Broderip, 1837, with
fully branched mantle that m ay resemble algae tufts
or coelenterate colonies) (Fig. 7).
In some other cases, the mantle is quite thin and
almost transparent, so the shell is easily visible
below (see f.i. the genus Zoilo Jousseaume, 1884).
Cowries are commonly cryptic (i.e. they hide) and
nocturnal, and this is of course a clear adaptation to
reduce predation. Nevertheless, some species, in
their adulthood, graze freely in the open during
the day (see f.i. the genera Zoilci, Bo.rycypTQ.eQ.
Schilder, 1927; and Cyproea tigris Linnaeus, 1 758).
Such cowries tend to have heavy big shells, which
is a clear adaptation to avoid predation by fishes
and/or crustaceans. Some may also have deltoid
shells, with a flat base, another clear adaptation to
stick to rocky surfaces (thanks to a foot acting as a
sucker) and prevent easy predation [see f.i. Mauri-
tia mauritiana (Linnaeus, 1758), Monetaria caput-
Serpentis (Linnaeus, 1758), etc.J.
Another environmental factor affecting shell
structure is sea current and/or waves action. Gener-
ally cowries with thick and heavy shells are typical
of turbulent waters. This feature is also variable
among individuals of the same species, since
lightweight shells tend to be more common in calm
lagoons or in deeper waters, while heavy calloused
shells are more easily found in the open ocean or in
high surf w aters.
THE UNUSUAL DEVELOPMENT OF A CO-
WRIE SHELL
The cowrie shell follows a developmental
pattern that is quite different from most mollusks.
The first shell to be produced is the larval shell of
the veliger (Fig. 8). While most species spend their
larval time in the plankton, others have a direct
intracap sular development (direct developers).
Once metamorphosed, the shell keeps growing by
adding whorls around its columella (Fig. 9). Even-
tually, during this growth, the shell may first re-
semble an Olivo shell (Fig. 10; i.e. ‘oliva stage’), or
a Bulla (i.e. ‘bulla stage’).
In both such stages, the spire is well visible and
the shell is very different from the adult one, both
for its structure and color. The shell is very thin
and all cowries are cryptic at this stage. This is
easily interpreted as an adaptation to prevent
predation. Although no secure data are available,
many personal observations and info obtained
from shell divers may point to the fact that cowries
get to adulthood very fast, perhaps within few
months from birth.
At the end of the juvenile stage, cowries
undergo a deep change in their shells: the last whorl
usually covers the entire shell, so the spire gets
included in it, and it eventually bends over the
columellar side to tighten the shell aperture (Fig. 1).
This tightening is even more pronounced by the
deposition of shell teeth, one of the most typical
features of cowries (Fig. 12).
Cowries have no operculum, so teeth are an
alternative strategy to make the aperture as narrow
as possible, to prevent access to soft parts when the
mollusk is retracted. Soon after teeth formation, the
shell stops growing, and it starts thickening by
deposition of shell and glaze layers, ending up into
the typical thick and glossy shell (Fig. 12). The fact
that cowries stop growing at adulthood is quite un-
usual among gastropods, which rather tend to have
an undetermined growing pattern. Moreover, the
growing rate and/or time to adulthood seem to be
quite variable, even among the same species, end-
ing up with a remarkable variability in adult shell
sizes (see f.i. Okon, 2013a, b; 2014).
The shell of a cowrie mollusk is therefore quite
different from most gastropod shells, as it evolved
several unusual characteristics, including a relevant
thickness, a very glossy surface, and a very narrow
aperture. Moreover, many cowrie shells are brightly
colored, making them quite visible to predators.
Notwithstanding this, cowries are among the most
successful gastropods in coral reefs, and they
perform quite well in many other marine environ-
ments. As mentioned, some do not even hide at
adulthood (see f.i. Fig. 13).
In the attempt to understand the peculiar adapt-
ations of cowries, we first have to consider that the
shell of Cypraeidae is generally thick and very com-
pact, hard to brake, with a very narrow aperture, and
the mollusk is usually very mucous, which makes
the cowrie quite slimy. These joined characteristics
are likely a good adaptation against predation.
The family Cypraeidae: an unexpected case of neglected animals
45 1
because the thickness of the shell, the absence of
possible holds, the slimy surface, and the relative
unreachability of soft parts, may discourage most
predators. In fact, such characteristics make preda-
tion by small fishes and crabs very difficult.
Actually, cowries are often wholly swallowed
by big fishes, since they cannot easily crack them,
although it sometimes happens with strongly
beaked fishes. Another important source of preda-
tion are octopuses, that drill the cowrie shell and
Figures 1-6. Examples of man ties and papillae in Cypraeidae (all from Hawaii). Fig. 1. Lyncind Cdmeold pVOpUlCjUQ. (G arrett,
1 879); Fig. 2. Tdlpciria talpa (Linnaeus, 1758); Fig. 3. Lurid tessellata (S w ainson, 1 822); Fig. 4. Lyncind fyriX (Linnaeus,
1 758); Fig. 5. Ndrid pOTdvid (Linnaeus, 1 758); Fig. 6. OvdtipSd chinensi dmiges ( M elvill et S tanden, 1915). Photos
courtesy David Lum.
452
Marco Passamonti
Fig u re 7 . Narici efOSa (Linnaeus, 1758) showing its extended
mantle resembling an algae tuft (Zanzibar, Tanzania). The
shell is barely visible in the middle of the dorsum, since the
mantle is not fully extended.
Figures 8-12. Developmental stages of Ndvid Spurca
(Mediterranean Sea). Fig. 8. Veliger shell; Fig. 9. Young
shell (just metamorphosed); Fig. 10. L OlivCf stage; Fig. 11.
Subadult; Fig. 12. Adult. Photos courtesy And re a Nappo and
Dario Marcello Soldan.
digest the mollusk, and eventually use cowrie shells
(and others) to adorn their dens. Predations by shell
drilling gastropods (e.g. Naticidae or Muricidae)
seem to be much more rare.
COWRIES’ REPRODUCTION
Cowries reproduction is quite remarkable too.
Females lay eggs in clusters of capsules on hard
surfaces and, at variance to many other gastropods,
they hatch eggs by covering them with the foot
(Fig. 14). Abandoned eggs may dye quite soon.
This commitment in parental cares is unusual in
marine gastropods, and it may be another reason
for the success of this Family. When intracapsular
development ends, planktotrophic larvae hatch and
swim in the water column until they metamorph-
ose. The length of larval stages may be different,
and could be somehow related to the capacity of a
given species to undergo local genetic diversifica-
tion (i.e. subspecies and/or geographic races).
Some species seem to have quite few divergent
races, while others have a much higher geo-
graphically structured pattern.
On the other hand, direct developers seem to
show different morphologies in different areas. In
direct developers, young mollusks undergo intra-
capsular development and, since they feed at the
expenses of accessory eggs in the capsule, they
keep growing until they hatch as crawling snails.
This development, evolved many times in cowries,
especially in some temperate water genera ( Zoila ,
Cypraeovula Gray, 1 824, Notocypraea Schiider,
1 927), has been often considered as an adaptation
to improve the chances of larvae to find specific
foods, like sponges they feed on. In fact, most
direct developers are fully depending over limited
food supplies, and they need to hatch as close
as possible to their food, to increase chances of
reaching adulthood.
DISTRIBUTION AND BIOGEOGRAPHY
Cowries are subtropical marine animals (Fig.
15), so most of them live in the oceans between the
two tropics. The highest number of species is found
in the Indo-Pacific Ocean, and far less species live
in theAtlantic (and the Mediterranean). Paulay and
The family Cypraeidae: an unexpected case of neglected animals
453
Figure 13. Two Zoila friendH jeaniana (Cate, 1 968) (f.
sherylae L. Raybaudi M assilia, 1990) grazing their host
sponge in the open (Point Qobba, W - Australia).
Photo courtesy Daniel Edinger.
Figure 14. Erronea caurica cfr. quinquefasciata (p.f. Ro-
ding, 1798) on eggs (Oman, Marisah Island).
Photo courtesy Massimo Scali & Beautifulcowries Maga-
zine.
Figure 15. Distribution map of the living species of cowries. Photo courtesy Mirco Bergonzoni.
Meyer (2006) proposed a species richness map of
the Indo-Pacific cowries. The highest species
richness is in the region going from the Philippines
to M elanesia, especially along the boundary between
East Indian Ocean and West Pacific. Species
richness significantly decreases going westtowards
Africa (although it locally increases again there), or
east along the Pacific Ocean towards Polynesia and
West Am erica. Quite significantly, similar species-
richness patters have been found in reef-building
corals, as in other reef-related organisms (see f.i.
Malay & Paulay, 2010). In fact, the region comp ris-
ing the ocean territories of Indonesia, the Philip-
pines, Malaysia (Sabah), East Timor, Papua New
Guinea and the Solomon Islands, is known as ‘the
coral triangle’, hosting more than 500 species of
reef-building corals (Veron, 1995). The reasons for
these similar species distribution patterns could be
both environmental and historical. Apparently the
thousand of islands and reefs in South EastAsia, as
well as their highly diverse habitats, were central to
a rich species radiation in cowries and other reef-
related animals. So that, probably, most of the wide-
spread Indo-Pacific cowries have once originated
in this area, and migrated (with different success)
outwards by larval dispersion.
As mentioned, the duration of veliger stages has
been again related to dispersion and speciation rates
454
Marco Passamonti
Figure 1 6 . G eographic diversification of the Erronea O YiyX species group. From left to right: Erronea adllSta adusta ( East
Africa); Erronea adusta nymphae (M auritius; Chagos); Erronea adusta persica (India, Oman, Persian Gulf); Erronea adusta
andamanensis (East Indian Ocean); Erronea ClduStCl melatiesiae { South West Pacific); Erronea onyx (N orth West Pacific).
Photo courtesy Mirco Bergonzoni.
Indian Ocean
West pacific
,‘rPuY.
'
fa
- ^ ft
East pacific
Figure 17. Examples of the variability of the Leporicypraea mappa species complex from different basins.
Photo courtesy Mirco Bergonzoni & Cypraea.net.
The family Cypraeidae: an unexpected case of neglected animals
455
Figures 18-27. Examples of the variability of the Zondrid pyVWfl species complex. Figs. 18-21. Z. pyruifl pyvwn (G m elin,
1791): M editerranean Sea to M au ri tan ia -Senegal (N orth of Dakar); Fig. 22. Z. pyrUYYl inSuldTWTl Schilder, 1928:Algarve (Por-
tugal), Cadiz (Spa in), Morocco, Canary Is.; Figs. 23-25. Z. pyVWTl petiticiHCl (Crosse, 1872): South of Dakar (Senegal), Ivory
Coast, Gabon, C. Verde; Fig. 26. Z. pyrUYYl angelicas (Clover, 1974): N orth Gabon, Guinea Gulf (?); Fig. 27. Z. Cltlgolensis
(Odhner, 1923): South Gabon, Luanda area (Angola). Photos c o urte sy Mirco Bergonzoni & Beautifulcowries Magazine.
456
Marco Passamonti
Figures 28-39. Examples of close relative couples of taxa of Cypraeidae. The one on the left is always the one with a wide
distribution range, the right one is the endemic relative with its range limited to peripheral locations. Above the line, the
couples considered as subspecies; below the line, the couples considered as different species, from left to right. Figs. 28,
29: Lurid lurida (Mediterranean Sea) and L. luridd OCeanica (Ascension Is.); Figs. 30, 31: Ndrid helvold and N. helvold
hawaiiensis (Hawaii); Figs. 32, 33: Ndrid cernicd and N. cernicd leforti (Easter Island); Figs. 34, 35: Ndrid dCWuldris and
N. dciculdris Sdnctdhelende (Ascension and Saint Helena); Figs. 36, 37: Moiietdrid Cdputserpentis and M. CdpiltdrdCOnis
(Easter Island and Sala Y Gomez); Figs. 38, 39: Cribrdruld dStdryi (f. lefditi ) and C. gdrcidi (Easter Island). Photos courtesy
B eautifulco w ries Magazine.
The family Cypraeidae: an unexpected case of neglected animals
457
by Paulay and Meyer (2006). Although duration of
veliger stages is only weakly correlated to species
range, it is significantly related to the diversification
of cowries along the Indo-Pacific basin: i.e. the
lower the veliger time is, the most is the
geographic/taxonomic diversification. In fact, it is
quite evident that some Indo-Pacific cowrie species
show very little geographic variation, see f.i. Monet-
aria caputserpentis, Monetaria annulus (Linnaeus,
175 8), while others are much more prone to produce
local races/subspecies. Only because they have been
recently analyzed in detail, I can mention here the
Erronea onyx (Linnaeus, 1 75 8) species complex
(Bergonzoni, 2013a), which appeared to differen-
tiate in allopatric races (Fig. 16), and the Lepori-
Cypraea niappa (Linnaeus, 1758) species group
(Fig. 17), one of the most biogeographically and
evolutionary complex cases among cowries (Ber-
gonzoni & Passamonti, 2014). By analyzing such
complexes in detail, very interesting cases on evol-
utionary history of marine organisms became
evident, including allopatric speciation events,
incipient speciation, relevance of genetic flow for
morphological diversification, etc. More case studies
are really interesting, but they still have to be ana-
lyzed in detail, as f.i. the genera Cribvarula Strand,
1929 and TaloStolida Ire dale, 1931, and the Bistolida
Stolida (Linnaeus, 1 75 8) and Erronea CUUrica ( Lin-
naeus, 1758) species complexes,justto name some.
As mentioned, cowries also colonized the At-
lantic, although their species richness is much lower
in this Ocean. Most likely, this colonization was
rather old, since at present the cold currents of
South West America (Humboldt Current) and South
West Africa (Benguela Current) are evidently a
strong barrier for larval dispersion and cowries set-
tling. At present, no species seems to be able to
spread from the Indo-Pacific to the Atlantic Ocean
north of Namibia, or in South America. Of course
no one lives in the Arctic Ocean as well. Neverthe-
less, Atlantic cowries have evident affinities with
Indo-Pacific ones. For instance, the genus MaCW-
Cypraea Schilder, 1930 is present at both sides of
Figure 40. Represen tatives of the South African genus CypraeOVulu. Photo courtesy Goncalo Rosa and Mirco Bergonzoni.
458
Marco Passamonti
the Isthmus of Panama, as it likely originated loc-
ally before the Isthmus was closed. On the other
side of the Atlantic, we have examples of couples
of allied species found in the Mediterranean and the
Red Sea/North West Indian Ocean [Lliria luvida
(Linnaeus 17 58)/Luria pulchva (Gray, 1 824)],
maybe a Thetyan residue. Some other genera are
end em ic to this reg io n ( ZonClria Jousseaume, 1884;
Schilderia Tomlin, 1930). Finally, a peculiar distri-
bution is given by the Nana spUVCO. complex of
species: this com prises three species, Nciria SpUTCCl
(Linnaeus 1 7 5 8), Narid aticularis ( G m e lin , 1791),
and Nciria cernica (Sowerby, 1 870). The first is
distributed in the Mediterranean/WestAtlantic, the
second along the East American coastlines, the lat-
ter is one of the most widely dispersed indopacific
cowries. Although the exact colonization tempos
and modes of Atlantic cowries are hard to speculate,
Figures 41-45. Ext re me variability in Zoila from Western and South Australia. Figs. 41-43: Zoila friendii subspecies /forms.
Fig . 44 : Z. ketyana sub spec ie s/form s . Fig . 45 : Z. VenUStCl sub spec ie s/form s . P ho to s co urtesy M ire o B ergo nzo ni & C ypraea.net.
The family Cypraeidae: an unexpected case of neglected animals
459
their phylogenetic relatedness to Indo-Pacific ones
is evident. Detailed phy logeographic analyses are
potentially of great interest to reconstruct the geo-
logical history of the Atlantic basin, its past connec-
tions and/or geological changes.
Again, as for Indo-Pacific cowries,in theAtlantic
as well the duration of cowries larval stages has
been correlated by the rate of morphological diver-
sification. One paradigmatic example comes from
the Zonaria pyrwn (Gmelin, 1 7 9 1 ) species complex
(Figs. 18-27), spreads in the Mediterranean and
West Africa region. Again, a short larval stage has
been related to the extreme capacity to differentiate
geographically, with different taxa spreading along
the African coast (Bergonzoni, 2013b).
Another interesting case of evolution comes
from cases of species/subspecies couples in which
one has a wide range, and the allied one has a
Figures 46-47. The genus Umbilici (South and East A ustralia). Fig. 46: Umbilici hesitCltCl Species complex.
F ig . 46 : U. armeniaca. Photos courtesy Mirco Bergonzoni & Cypraea.net.
460
Marco Passamonti
Figure 48. Barycypraea teulerei (Oman). Two males ap- Figure 49. Barycypraea fultoni fultoni (Natal, S. Africa),
proaching a female hatching eggs into a empty bivalve shell. Photo courtesy Felix Lorenz & B eautifulco w ries Magazine.
peripheral endemic distribution. Figures 28-39
show some cases. Most of them are isolated
endemics, likely arisen p arap h y le tic ally . Quite
remarkably, they are treated much differently in
established taxonomy: some are actually con-
sidered as full species, some other as subspecies,
although no evident reason (besides A uthors’ opin-
ion) has been produced so far.
As mentioned, cowries were able to colonize
temperate waters as well. This is particularly evid-
ent for South Africa and West/South Australia, in
which endemic genera evolved. In South Africa, the
most striking evolutionary radiation is the genus
Cypraeovula (Fig. 40), including different closely
related species that sometimes hybridize too. On the
other side of the Indian Ocean, in Western Australia,
another striking example of colonization of tempe-
rate waters is the genus Zoila (Lorenz, 2001;
W ilson and Clarkson, 2004) (Figs. 41-45). The Zoila
cowries are sponge eaters and direct developers,
and this caused a flourishing of local races, making
them one of the most taxonom ically complex
groups of marine organisms. Along with Zoila,
another similar case is the genus NotOCypraea,
which is however much less known in detail. Fi-
nally the genus UlYlbili(X Jousseaume, 1884 (W ilson
and Clarkson, 2004) is another sticking Australian
endemism of temperate waters, as it is distributed
along the East and South coast (Figs. 46, 47).
Other cases of direct developers are found as
well, such as the genera Barycypraea and Mura-
cypraea Woodring, 1 957, and few others. In all
cases, these species have a very limited range. F.i.
Barycypraea teulerei ( Cazenavette, 1 846) (Fig. 48),
a shallow water direct developer (Scali, 2013;
2014), seems to be found in a limited area of Oman
only, while the deep water relative Barycypraea
fultOlli (Sowerby III, 1 903) is found between
Mozambique and SouthAfrica (Bergonzoni, 2012)
(Fig. 49). Another case of direct developer with
little dispersal capacity is Muracypraea 171US
(Linnaeus, 1758), limited to the Gulf of Venezuela
coasts and Guajira Peninsula in Colombia.
THE MOLECULAR PHYLOGENETICS
OF COWRIES AND THEIR TAXONOMY
This is probably the field of cowries biology that
has been more thoughtfully investigated. In fact, a
huge phylogenetic reconstruction, based on DNA,
have been proposed by Meyer (2003; 2004). The
primary outcome of this pivotal work is an ultimate
tuning of the so vrasp ecific taxonomy of the family.
Quite remarkably, most of the subfamilies and ge-
nera proposed by older A uthors on morphology (see
f.i. S childer & Schilder, 1 938; Schilder, 1 939;
1 966;) have been confirmed by DNA.
The family Cypraeidae is now subdivided
into 7 subfamilies ( A rch ic y p raein ae , Erosariinae,
Umbiliinae, Cypraeinae, Bernayinae, Luriinae and
C y p raeo v u lin ae ) and 48 genera (Moretzsohn,
2014), and this arrangementhas gained a very good
agreement among cowrie experts.
The family Cypraeidae: an unexpected case of neglected animals
461
Even if a relatively stable sovraspecific tax-
onomy has been reached in the Family, this is
certainly untrue for species level and, even more,
below it. Most of the proposed taxonomies are
based on morphological analyses, as well as on
Authors’ opinions. Only few DNA data and/or
detailed evolutionary studies are available to date.
Moreover, a certain degree oftaxonomic prolifera-
tion has been certainly triggered by economic
factors: in fact, many cowrie collectors want new
names, so that a new named cowrie gets a much
higher value in the marked. This cause what I’d call
‘economic speciation’, with some humor, of course!
This approach should be strongly stigmatized for
two reasons: 1st, it produces an unnecessary prolif-
eration of taxonomic names; 2nd, it has no biolo-
gical bases in most cases.
Another problem comes from the rules of tax-
onomy, and this is particularly evident for species
and subspecies names, which are under the provi-
sions of the International Code of Zoological
Nomenclature. In my opinion, new species and sub-
species names should not be introduced in tax-
onomy if not based on rigorous biological and
evolutionary analyses. Nevertheless, specialized
collectors need to have names to refer to morpho-
logies that are just not so important for evolutionary
biologists, like local variants, unstable morphs,
aberrations, etc. The use of ‘forma’ names should
be a good compromise, because they, one side, meet
collectors’ needs for names, and, the other do not
increase taxomonic complexity (i.e., infrasub-
specific names are not under the provisions of the
Code). This approach is not without problems, of
course, but it seems to me the only possible
compromise between two different, and sometimes
contrasting, needs.
MUTATIONS AND ABERRATIONS
Albinism and rufinism
As it happens in all living beings, cowries may
show some interesting mutations and aberrations.
Some of them, being rare, may produce some of the
most sought-after and priced cowries, so all collect-
ors know them very well. On the other hand, their
biological causes are quite unknown or neglected.
I try to highlight some of them here.
F ig ure 5 0. ZoUa dedpieUS from Broome area, W-Australia. From left to right: black (normal), alb in o and rufinistic shells.
Photo courtesy Drew Strickland.
462
Marco Passamonti
The first example comes from rare recessive
mutations, such as cowries’ albinism (producing
white shells) or rufinism (producing orange shells)
(Fig. 50). These phenotypes are evidently due to
rare mutations affecting the genes for shell color. It
is quite remarkable that white or orange shells are
not necessary associated to white and orange
animals, respectively. This clearly points out to the
observation that genes for shell color are different
from the ones of the soft parts. For this reason, it
would be inaccurate to call such specimens ‘albino’
or ‘rufinistic ’ , but I use these terms here for simpli-
city. Both rufinistic and albino cowries have been
proposed to be the result of mutations over the same
metabolic pathways producing brown/black
pigments (melanin?). For a detailed discussion see
Passamonti and Hiscock (2013).
Besides their high collecting value, the appear-
ance of rare mutants within a population represents
an interesting case to study the dynamics of allele
frequencies, and the effects that collecting pressure
may have on the variability of natural populations.
A paradigmatic example is that of Zoilci WSSelli
SCltiatCl Lorenz, 2002, from Fitzroy Reef, Quobba
Point, North West Australia. This once quite large
population was an important source of rosselli spe-
cimens, and many hundreds have been collected
over the years. Among normal shells, around 40
rufinistic specimens were found (f. edingeri
Raybaudi M assilia, 1 990) (Beals, 2013) (Fig. 51).
These shells were collected over a limited time-
lapse, as the first ones were collected in 1 988, and
they disappeared soon after 1997. Why the mutant
disappeared so fast? One may think that this is
because all orange shells were collected, so they
Figure 51. Zoila rosselli satiata and Z. rosselli satiata
f. edingeri (rufinistic). Both found at Point Quobba,
W-Australia. Photo courtesy Daniel Edinger and Beautiful-
cowries Magazine.
could not produce orange progeny anymore. Howe-
ver, this is not fully the case: since rufinistic muta-
tions are likely recessive (i.e. they may ‘hide’ in
eterozygous individuals), two heterozygous black
mates may well produce 14 of orange shells, accord-
ing to M endelian proportions. So, the overall col-
lecting pressure (on both black and orange
specimens) is rather the reason of this disappear-
ance: by reducing dramatically the number of
individuals, the population underwent a strong
‘bottleneck’, which is well known in evolutionary
biology to reduce genetic variability. Because chances
for rare alleles to pass throughout a population bot-
tleneck are very scarce, the rufinistic allele was
soon lost from the population, and no edingeri was
found since then. This also means that the chances
that this allele will appear again in Quobba are quite
low, and the edingeri rufinistic mutation is simply
no longer existing. However, rufinistic shells are
found within many other Zoila species, as well as
in some other cowries. So rufinism is likely a case
of recurrent mutation. F.i. an independent rufinistic
morph has been recently evidenced in another Zoilci
WSSelli population (see Lorenz, 2011; 2014).
Albinos are much more rare among cowries.
Although many cowries may be white or whitish,
the rare albino mutants are only known for very few
cowries [f.i. in Cyproeo tigris (Fig. 5 2), and Zoila
dedpienS Smith, 1 8 8 0 (Fig. 5 0 ) J . What is quite
interesting is that albino morphs may became fixed
in some populations (i.e. all the shells are albinos),
hence they are not rare mutants anymore: two
paradigmatic examples are Ncirici ebume Cl B a rnes ,
1 824 (Fig. 53) and Erronea nymphae J ay , 1 8 5 0 (see
Fig. 16). Both are clearly related to non-albino
relatives, Narici miliaris (Gm eiin, 1791) and Erronea
adllSta, (Lamarck, 1810), respectively. These are
likely cases in which the ‘albino’ allele was fixed
into a new population because of a ‘founder effect’,
i.e. when a new population had established in a new
area the albino allele become by chance the unique
one (i.e. it was fixed).
Niger and rostrated cowries
This is another interesting feature, which ap-
pears to be unique to some cowries and only one
ovulid species, CalpumuS VerriiCOSUS (Linnaeus,
1758). It is evident that these two characteristics are
the outcome of the unusual developmental scheme
The family Cypraeidae: an unexpected case of neglected animals
463
Figure 52. Two normal CypraeCL tigris along with an albino
one (Madagascar and Philippines). Photo courtesy Mirco
B ergonzoni & Cypraea.net.
Figure 53. Two species of the genus Naria. N avid miliavis
from China (left) and N. eblimeci from New Caledonia
(right).
of cowries’ shells. As mentioned, adult cowries stop
growing, since the deposition of shell layers and
pigments stops or strongly reduces. However, in
some specimens, the signal to stop seems not to
work properly, and the shell keeps growing by
adding layers of shell and/or pigment. Hence such
cowries quickly get a ‘gondola shape’ (rostrated),
as well as a deep black color (melanic or ‘niger’
cowries). Both phenomena may appear together, or
not, depending on species. Some species become
melanistic and rostrated altogether, others may only
be rostrated (these are the ones that do not have
brown/black colors in normal adults). Also, the
degree of rostration and melanism may vary among
individuals (Figs. 54, 55 ).
Even if such phenotypes are occasionally
found over the entire range of some cowrie
species, it is quite remarkable that they get much
more common in two specific areas: the southern
reefs of New Caledonia, and the Keppel Bay area
in Queensland. The biological causes of such aber-
rations are still unknown, and some have linked
these phenomena to the presence on heavy metals
(nickel?) in the water (see, f.i., Pierson and
Pierson, 1975). What I think it is interesting is the
presumable genetic base of melanism and rostra-
tion: as mentioned, both could easily be inter-
preted as a malfunctioning of genetic regulation of
shell development. The pattern of expression of
developmental genes is somehow affected (by
metals? by other environmental factors? by muta-
tions?), and the genes for deposition of shell color
and layers just fail to stop at adulthood, as it hap-
pens in normal cowries. Needless to say that we
have no clue on which genes are involved in such
processes, and this would certainly be a nice case
study for developmental biologists.
CONCLUSIONS
In this paper I tried to highlight some biological
peculiarities of cowries, making them interesting
case studies to many aspects of evolutionary bio-
logy, not only for taxonomy. Cowries are very inter-
esting marine organisms, and, even if they have
been studied by very few professional biologists,
they are well known by amateurs, and a huge
amount of ‘first-hand’ data are available. This
manuscript is far to be complete, and many other
interesting cases could be highlighted; nevertheless
I hope that this short review has attracted your
interest to this amazing group of animals, that
certainly deserves more studies. The collection of
cowries, which is unfortunately so deeply money-
driven, is certainly a restraint to biological studies,
since some species are hardly accessible and col-
lecting data are often vague (to preserve a relevant
464
Marco Passamonti
Figures 54, 55. Examples of different degrees of melanism and rostration in New Caledonian cowries.
Fig. 54: Mauritia ( Arabica ) eglantina-. Fig. 55 . Bistolida stolida.
The family Cypraeidae: an unexpected case of neglected animals
465
source of income for divers/dealers). On the other
hand, an important collecting effort is a precious
help for biologists, since maybe for no other group
of gastropods we have such a huge amount of know-
ledge ‘in the field’. We should therefore try to build
a ‘bridge’ between the two words (cowries amateurs
and biologists) that both may benefit: collectors will
start to consider cowries not as just precious and
beautiful objects, and evolutionary biologists/
professional m alacologists as interesting animals to
study. Only this way, the preconception that cowries
are just pretty but uninteresting animals will be
definitively overcome.
ACKNOWLEDGMENTS
This paper is the written transposition of my
invited talk at the ‘VII Pontine M alacological
Congress’, held in San Felice Circeo (Latina,
Italy). I wish to thank you very much the organ-
izers, Silvia Alfinito and Bruno Fumanti, as well
as the M alakos 2002 association and members, for
this nice opportunity, their lovely welcome and the
beautiful conference location. A special thanks
goes to friends who provided some of the images
of this paper (in alphabetical order): Mirco Ber-
gonzoni (Calderara, Italy), Daniel Edinger (Man-
durah, Western Australia), David Lum (Honolulu,
Hawaii, U.S.A.), Andrea Nappo (Quartu Sant’
Elena, Italy), Goncalo Rosa (Lisboa, Portugal),
M assimo Scali (Imola, Italy), D ario Marcello So Id an
(Milan, Italy), Drew Strickland (Geraldton,
Western Australia).A special thanks goes to Mirco
Bergonzoniforthe numerous evening spent talking
about our beloved cowries.
REFERENCES
Beals M ., 2013. A second review of Zoilci WSSelli
edillgeri Raybaudi, 1990. B eautifulco w ries M agazine,
1 : 4-11.
Bergonzoni M ., 2013a. Let’s make some order in the
Erronea onyx species complex. B eautifulcow ries
Magazine, 4: 31-59.
Bergonzoni M ., 2012. Barycypraeci fllltoni, a tale of a
fallen star. B eautifulco w ries Magazine, 2: 4-19.
Bergonzoni M., 2013b. The ZoilCiria pyruni complex.
B eautifulcow ries Magazine, 3: 35-56.
Bergonzoni M . & Passamonti M ., 2014. A monograph on
Leporycypraea mappa-. a taxonomic and evolutionary
puzzle. B eautifulcow ries Magazine, 6: in press.
Lorenz F., 2001. Monograph of the living Zoilci.
Conch books, Hackenheim (Germany), 188 pp.
Lorenz F., 2011. A new species of Zoilci from SW
Australia (Gastropoda: Cypraeidae). S chriften zur
Malakozoologie (Cismar),26: 11-14.
Lorenz F., 2014. New findings of Zoilci rciyWCllkeri Lo-
renz, 2013. B eautifulcow ries Magazine, 5: 36-37.
Malay M .C . & Paulay G., 2010. Peripatric speciation
drives diversification and distributional pattern of
reef herm it crabs (Decapoda: Diogenidae: CcildnuS) .
Evolution, 64-3: 634-662.
Meyer C.P., 2003. Molecular systematics of cowries
(Gastropoda: Cypraeidae) and diversification patterns
in the tropics. Biological Journal of the Linnaean
Society, 79: 401-459.
Meyer C .P., 2004. Towards c o m p ren siv e n e s s : in-
creased molecular sampling with Cypraeidae and its
phylogenetic implications. Malacologia, 46: 127-
156.
Moretzsohn F., 2014. Cypraeidae: how well-inventoried
is the best-known seashell family? American
M alacological Bullettin, 32(2): 278-289.
Okon M.E., 2013a. On dwarfs and giants. Part 1. Beau-
tifulcowries Magazine, 3: 61-62.
Okon M.E., 2013b. On dwarfs and giants. Part 2. Beau-
tifulcowries Magazine, 4: 60-62.
Okon M .E ., 2014. On dwarfs and giants. Part 3. Beauti-
fulcowries Magazine, 5: 56-58.
Passamonti M . & Hiscock M ., 2013. A closer look at rufin-
istic and albino Zoilci. B eautifulcow ries M agazine, 1 :
8-18.
Paulay G. & Meyer C., 2006. Dispersal and divergence
across the greatest ocean region: do larvae matter?
Integrative and Comparative Biology, 46: 269-
28 1.
Pierson R. & Pierson G., 1975. Porcelaines mysterieuses
de Nouvelle-C aledonie. Self printed, Noumea. 122
pp.
Scali M., 2013. Barycypraeci teulerei (C azenavette,
1846). The rediscovery. B eautifulcow ries M agazine,
3:4-11.
Scali m., 2014. Barycypraeci teulerei, going back to the
recently discovered new population. Beautiful-
cowries Magazine, 5: 31-35.
Schilder F.A . & Schilder M ., 1 93 8. Prodrome of a
monograph on living Cypraeidae. Proceedings of
the M alacological Society of London, 22-23: 1 19 —
23 1.
Schilder F.A., 1939. Die Genera der Cypraeacea.Archiv
fur Molluskenkunde, 71: 165-201.
Schilder F.A., 1966. The higher taxa of cowries and
466
Marco Passamonti
their allies. The Veliger, 9: 31-35.
Veron J.E.N., 1995. Corals in space and time: biogeo-
graphy and evolution of the Scleractinia. UNSW
Press, Sydney (Australia), 334 pp.
Wilson B. & Clarkson P., 2004. Australia’s spectacular
cowries. A review and field study of two endemic
Genera: Zoilci and Umbilici. Odyssey Publishing, El
Cajon (California), 396 pp.
Biodiversity Journal, 2015, 6 (1): 467-480
Monograph
The Recent Rissoidae of the Mediterranean Sea. Notes on the
genus Onoba s.s. H. Adams et A. Adams, 1 852 (Gastropoda
Prosobranchia)
Bruno Amati 1 & Italo Nofroni 2
'Largo Giuseppe Veratti 37/D, 00146 Rome, Italy; e-mail: bmno_amati@yahoo.it
2 Via Benedetto Croce 97, 00142 Rome, Italy; e-mail: italo.nofroni@uniromal.it
"■Corresponding author
ABSTRACT The Mediterranean species belonging to the genus Onoba H. Adams et A. Adams, 1852 as
currently conceived, are reviewed. With the exception of O. semicostata (Montagu, 1803)
and O. aculeus (Gould, 1841) that range mostly in the European North-Eastern Atlantic and
are rarely found in the Western Mediterranean, this genus is represented by six species
with rather limited ranges: O. dimassai Amati et Nofroni, 1991; O. josae Moolenbeek et
Hoenselaar, 1987; O. guzmani Hoenselaar et Moolenbeek, 1987; O. tarifensis Hoenselaar
et Moolenbeek, 1987; O. gianninii (Nordsieck, 1974) and O. oliverioi Smriglio et Mariottini,
2000. A further possibly undescribed species is figured. For all species comparative
morphometries are provided. Onoba josae Moolenbeek et Hoenselaar, 1987 is here recorded
for the first time in Italy, with the easternmost locality in this range.
KEY WORDS taxonomy; Rissoidae; Onoba; Recent; Mediterranean Sea; first record.
Received 21.02.2015; accepted 23.03.2015; printed 30.03.2015
Proceedings of the Eighth Malacological Pontine Meeting, October 4th- 5th, 2014 - San Felice Circeo, Italy
INTRODUCTION
The genus Onoba H. Adams et A. Adams, 1852
has been frequently discussed in the malacological
literature (e.g. H. & A. Adams, 1852: 358; Jeffreys,
1867: 37; Watson, 1873: 387; Verril, 1884: 182;
Friele, 1886: 28; Dautzenberg, 1889: 52; Waren,
1973: 4; Waren, 1974: 130; Rolan, 1983: 139;
Ponder, 1985: 54; Templado & Rolan, 1986: 117;
Bouchet & Waren, 1993: 659; Ponder & Worsfold,
1994: 26; Rolan, 2008: 233; Nekhaev et al., 2014:
269) and has a global distribution, ranging in both
hemispheres from the poles to at least the subtropics
(Ponder, 1985: 55; Rolan, 2008: 233; Avila et al.,
2012:4).
It is currently subdivided into some few sub-
genera: Onoba (type species Turbo striatus J.
Adams, 1797), Ovirissoa Hedley, 1916 (type
species Rissoa adarensis Smith, 1902), Subestea
Cotton, 1944 (type species Alvania seminodosa
May, 1915) and Manawatawhia Powell, 1937 (type
species M. analoga Powell, 1937).
Seven central-western Mediterranean species,
most of which have been described during the last
forty years, are currently ascribed to the nominal
subgenus (Rolan, 1983: 139; Aartsen et al., 1984:
20; Templado & Rolan, 1986: 117; Oliverio et al.,
1986: 35; Moolenbeek & Hoenselaar, 1987: 153;
Hoenselaar & Moolenbeek, 1987: 17; Amati &
Nofroni, 1991: 30; Smriglio & Mariottini, 2000: 15;
468
Bruno Amati & Italo Nofroni
Giannuzzi-Savelli et al., 2002: 80; Rolan, 2008:
233; Gofas et al., 2011: 193; Avila et al., 2012: 5;
Bouchet, 2014); these species are: Onoba semicost-
ata (Montagu, 1803), O. gianninii (Nordsieck,
1974), O. tarifensis Hoenselaar et Moolenbeek,
1987, O. guzmani Hoenselaar et Moolenbeek, 1987,
O.josae Moolenbeek et Hoenselaar, 1987, O. dimas-
sai Amati et Nofroni, 1991 and O. oliverioi Smriglio
et Mariottini, 2000. Another species, O. aculeus
(Gould, 1841), geographically ranging typically on
both sides of northern Atlantic including Greenland,
has been reported only once from the Mediter-
ranean Sea (Giannuzzi-Savelli et al., 2002: 80).
The missing of new further records convinced
some Authors to exclude this species from the main
Mediterranean check-lists (Rolan, 2008: 241;
Nekhaev et al., 2014: 272). A further possibly
undescribed species has been recorded ( Onoba sp.:
Amati & Nofroni, 1991: 34), but never formally
named. The anatomy of the genus Onoba has been
studied by Ponder (1985: 56). Here we utilize the only
shell morphology for the description and comparisons
of the Mediterranean species. The most important
iconographic references are reported for each species.
ABBREVIATIONS AND ACRONYMS. BA:
Bruno Amati collection, Rome, Italy. CS: Carlo
Smriglio collection, Rome, Italy. IN: Italo Nofroni
collection, Rome, Italy, lv: live collected specimen.
MCZR: Museo Civico di Zoologia, Rome, Italy.
MNHN: Museum National d'Histoire Naturelle,
Paris, France. MO: Marco Oliverio collection,
Rome, Italy. MZB: ‘Museo di Zoologia’ of the Uni-
versity of Bologna, Italy. PM: Paolo Mariottini col-
lection, Rome, Italy. RAMM: Exeter’s Royal Albert
Memorial Museum & Art Gallery, Exeter, Devon,
UK. SB-MS: Stefano Bartolini-Maria Scaperotta
collection, Florence, Italy. SEM: Scanning Electron
Microscope, sh: empty shell, v.: versus. ZMA: Zo-
ological Museum, Amsterdam, The Netherlands.
SYSTEMATICS
Family Rissoidae Gray, 1847: 152 (as Rissoaina)
Genus Onoba H. Adams et A. Adams, 1852: 358
Type-species: Turbo striatus J. Adams, 1797 non
Da Costa, 1778 = Onoba semicostata (Montagu,
1803: 326 (by monotypy)
Morphology. Diagnosis shell of genus Onoba
(from Ponder, 1985: 54): “Shell: minute to small,
ovate-conic to elongate-ovate, non-umbilicate to
narrowly umbilicate, smooth or with weak to strong
spiral sculpture, with a few spiral keels. Axial
sculpture usually rather weak to very weak; some-
times axial ribs present but do not extend over base;
sculpture rarely clathrate. Aperture with simple
peristome, oval, weakly angled and channelled
posteriorly, simple and rounded anteriorly; outer
lip opisthocline, varix weak to heavy. Protoconch
dome-shaped, sometimes with 1 or more spiral
keels; smooth (Ovirissoa) or with microsculpture of
granules, anastomosing or spirally aligned raised
threads or, sometimes, wavy, spirally arranged
rows of granules. Periostracum very thin to well
developed ’ ’.
Diagnosis shell of subgenus Onoba : (from
Ponder, 1985: 56): “Shell: broadly ovate-conic to
elongate ovate, rather solid, non-umbilicate,
usually with many well developed spiral cords and,
sometimes, weak axial ribs; microsculpture of fine
spiral lirae usually present. Strong spiral cords in
a few species and, in some species, surface smooth.
Aperture subcircular, subangled and weakly chan-
nelled posteriorly, varix on outer lip strong to mod-
erate. Protoconch domeshaped of about 11/2
whorls in nearly all species, rarely up to 2 2/4
whorls (as in O. 'semicostata '); sculptured vari-
ously, for example, with exceedingly weak to moder-
ately strong spiral lines with either parallel to
oblique wrinkles or granules between, as in O.
aculea (Gould) and O. moreleti Dautzenberg and in
Fretter & Graham's (1978) figure of O. 'semi-
costata'; with irregular, raised, wavy threads, as in
O. foveauxiana. (Suter); with scattered granules, as
in O. fumata, O. kermadecensis (Powell) and sev-
eral other southern species, as well as O. n. sp.from
the Eocene of France; (see also Thiriot Quievreux
& Babio, 1975; Fretter & Graham, 1978 ).”
Onoba semicostata (Montagu, 1803) (Figs. 1-6)
Turbo striatus J. Adams, 1797: 66 non da Costa,
1778: 86
Turbo semicostatus Montagu, 1803: 326, pi. XXI,
fig. 5
Rissoa ecostata Michaud, 1830 (WoRMS: Bouchet,
2014)
Rissoa minutissima Michaud, 1830 (WoRMS:
Bouchet, 2014)
Rissoa peticularis Menke, 1830 (WoRMS: Bouchet,
2014)
The Recent Rissoidae of the Mediterranean Sea. Notes on the genus Onoba s.s. (Gastropoda Prosobranchia)
469
Onoba Candida (Brown, 1827) (Giannuzzi-Savelli,
2002: 80)
Iconographic references. Montagu (1803:
326, pi. XXI, fig. 5); Reeve (1878: pi. V fig. 40 as
Rissoa striata)', Rolan (1983: 139, 140, 3 un-
nambered figures as Onoba aculeus and 5 un-
numbered figures as Onoba striata)', Rolan (2008:
234, figs. 1-12); Giannuzzi-Savelli et al. (2002: 80,
fig. 255 as Onoba Candida (Brown, 1827));
Nekhaev et al. (2014: 269, figs. 1 A-B, 4 A, D).
Type locality. Atlantic Ocean, south coast of
Devonshire, United Kingdom.
Type material. Not seen. Probable syntypes in
Montagu collection (RAMM)
Examined material. Norway: Grande, Viken, -
100/200 m, 07.1974, 1 sh (BA); Spain: Vigo-
Baiona, North-West Atlantic, beached, 08.1982, 1
sh (IN); Vigo Bay (Atlantic) -15 m (legit Palazzi,
1982), 11 juv. sh (IN); Fuengirola, (Malaga)
beached (ex coll. Bogi), 1 sh (IN); France: Carteret,
Normandy, (Atlantic, 1976), beached, 1 sh (IN);
Binard (Atlantic, 1975) among littoral seaweeds, 2
lv (IN); no locality, 1 sh (IN).
Original description. Montagu, 1803: “Z with
a short, conic, white shell, obtusely pointed: volu-
tions four or five, rounded, well defined by the sep-
arating line, and wrought with fait ribs, and fine
obsolete transverse striae on the body whorl, both
of which are inconspicuous on the superior spires:
the ribs do not extend to the lower part even of the
body, where the spiral transverse striae become
most conspicuous: aperture suborbicular; pillar lip
a little reflexed, Columella smooth. Length half a
line; breadth one half its length. Found in sand on
the south coast of Devonshire, but very rare. This
at first sight might be confounded with Turbo
Spiralis, but differs in the volutions being more
rounded, in the ribs being coarser, and in being des-
titute of the tooth-like plication of the columella.'’'’
Distribution and habitat. Eastern Atlantic,
Madera (Avila et al., 2012), Spain (Rolan, 2008),
British Isles (Jeffreys, 1867: 37; Fretter & Graham,
1978), Faroe Islands (Waren, 1996; Sneli et al.,
2005), Iceland (Waren, 1996), Norway (Hoisseter,
2009). Barents Sea, Kola Peninsula (Golikov
& Kussakin, 1978; Nekhaev et al., 2014: 271).
r
Mediterranean Sea (Avila et al., 2012), Alboran
Sea, Fuengirola (Giannuzzi-Savelli et al., 2002: 80).
Common and abundant under rocks and among
algae, from the intertidal to -1000 m depth (Tem-
plado & Rolan, 1986: 120); common on rocks in
-8 m, less common in -80 m, rare under -200 m in
the Zelenetskaya Bay, Barents Sea (Nekhaev et al.,
2014: 272).
Remarks. Onoba semicostata is the only Mediter-
ranean Onoba with a planktotrophic larval develop-
ment, and is therefore easy to identify (Rolan, 2008:
35, figs. 3-6; Nekhaev et al., 2014: 276, figs. 4 A,
D). Shells tend to be curved (var. distorta Marshall
fide Jeffreys, 1887: 35) and occasionally may have
an additional labial varix. Shells collected in the
central Mediterranean are probably fossils (Wurm).
Onoba aculeus differs from O. semicostata in
having a paucispiral protoconch (indicating a non
planktotrophic development), a slightly scalariform
suture with more convex whorls without subsutural
axial ribs. Onoba breogani Rolan, 2008, known, at
moment, for Galicia (Spain, Atlantic), is very
similar to O. semicostata in shell morphology,
having also subsutural axial ribs, but differs in its
paucispiral protoconch.
Onoba aculeus (Gould, 1841) (Figs. 7, 8)
Cingula aculeus Gould, 1841: 266, fig. 172
Rissoa saxatilis Moller, 1842: 9
Rissoa artica Foven, 1846: 156
Rissoa multilineata Stimpson, 1851: 14
Onoba aculeus (Gould, 1841) (Giannuzzi-Savelli
et al., 2002: 80)
Iconographic references. Gould, (1841: 172,
fig. 172) (not a good picture); Bouchet & Waren
(1993: 660, fig. 1507); Delongueville & Scaillet
(2001: 12, fig. 10); Giannuzzi-Savelli et al. (2002:
80, fig. 254); Nekhaev et al. (2014: 272, figs. 2
C-D, 4 B, E).
Type locality. East Boston, Massachusetts
(USA).
Type material. Not seen. Originally deposited
at the Boston Society of Natural History (BSNH:
State Coll., No. 32. Soc. Cab., No. 2359. Gould,
1841: vi, 266).
Examined material. Bergen (Norway, At-
lantic), -1 m, 2 lv (IN).
470
Bruno Amati & Italo Nofroni
Original description. Gould, 1841: “ Shell
minute, sub- cylindrical; whorls convex, covered
with regular, microscopic revolving lines; aperture
ovate; umbilicus partial. Shell minute, ovate-cyl-
indrical, elongated, light yellowish horn-color;
whorls six, convex, and separated by a deep sutural
region; the two upper ones forming a blunt apex,
the lowest rather more than half the length of the
shell; the whole covered with regular, crowded, mi-
croscopic revolving lines; aperture one third the
length of the shell, oval, oblique, angular behind,
the margin simple and entire, barely touching the
preceding whorl, somewhat expanded, and on the
left side elevated, and slightly turned over an
Figures 1-6. Onoba semicostata (Montagu, 1803): Figures 1, 2. Grande, Viken (Norway, Atlantic), height 2.9 mm (BA).
Figures 3, 4. Binard (France, Atlantic), height 2.3 mm (IN). Figures 5, 6. Fuengirola, (Spain, Mediterranean Sea), height
2.7 mm (IN).Figures 7, 8. Onoba aculeus (Gould, 1841), Bergen (Norway, Atlantic), height 3.05 mm (IN).
The Recent Rissoidae of the Mediterranean Sea. Notes on the genus Onoba s.s. (Gastropoda Prosobranchia)
471
umbilical depression or chink; operculum horny.
Length 3/20 inch, breadth 1/15 inch, divergence
23°. Found sparingly on the partially decayed
timbers of an old wharf, and plentifully on stones,
about low-water mark, at East Boston .”
Distribution and habitat. Western Atlantic
(Gould, 1841), Greenland (Moller, 1842; Schiotte
& Waren, 1992), Eastern Atlantic, Faroe Islands
(Sneli et al., 2005), Iceland (Ingolfsson, 1996;
Waren, 1996), British Isles (Fretter & Graham,
1978), Northern Norway (Hoisseter, 2009). Barents
Sea, Kola Peninsula and White Sea (Golikov,
1987), Galicia (Templado & Rolan, 1986: 121).
Mediterranean Sea, Alboran Sea (Giannuzzi-Savelli
et al., 2002: 80). Very common in the Barents Sea
in 0/-3 m on sandy bottoms (Nekhaev et al., 2014:
272). The species seems to prefer shallow waters
with algae, and can tolerate brackish waters (Tem-
plado & Rolan, 1986: 121).
Remarks. The record from Ria de Vigo (Galicia:
Templado & Rolan, 1986: 121) is the southernmost
occurrence in the Atlantic Ocean, whilst the Al-
boran Sea record (Giannuzzi-Savelli et al., 2002:
80) should represent the southern limit overall.
Shells tend to be curved. O. aculeus is very similar
to O. galaica Rolan, 2008, from Galicia (Spain).
Whilst some measurements of teleoconchs (e.g.
number of spirals cords on the penultimate and the
body whorl) and protoconchs (maximum diameter)
are similar in the two species (Rolan, 2008), the
different protoconch sculpture (with fine spiral
cords in O. aculeus and almost smooth in O.
galaica) (Fretter & Graham, 1978; Waren, 1996;
Rolan, 2008) and the less marked teleconch micros-
culpture, along with other minor differences (e.g.
deeper suture, larger size according to Waren, 1996)
allow an easy separation of O. aculeus and O.
galaica. See below under Onoba semicostata for
the differences from Onoba aculeus.
Onoba dimassai Amati et Nofroni, 1991 (Figs. 9-12)
Onoba dimassai Amati & Nofroni, 1991: 30, figs.
1M
Iconographic references. Amati & Nofroni
(1991: 30, figs. 1-4); Giannuzzi-Savelli et al.
(2002: 82, 83, fig. 256)
Type locality. San Felice Circeo, Central
Tyrrhenian Sea , Italy -30/50 m.
Type material. Holotype (MCZR), 9 paratypes
(loc. type) (BA), 2 paratypes (type loc.) (IN), 1
paratype (type loc.) (coll. Di Massa, Trieste), 2
paratypes Ventotene Is., Central Tyrrhenian Sea -25
m (MCZR ex coll. Pizzini), 7 paratypes Ventotene
Is., Fe Sconciglie Shoal, Central Tyrrhenian Sea,
-41m (MO), 3 paratypes Ponza Is., Central Tyrrhe-
nian Sea, bioclastic sand sample Posidonia ocean-
ica -15 m, 04.1979 (coll. A. Fugli, MO), 1
paratype Ponza Is., Central Tyrrhenian Sea, -35 m,
05.1983 (coll. Di Massa TS), 1 paratype S. Stefano
Is., Central Tyrrhenian Sea, -40 m (MZCR ex coll.
Pizzini), 1 paratype Giannutri Is., Central Tyrrhe-
nian Sea, -27 m (MZCR ex coll. Pizzini), 1 paratype
Giglio Is., Central Tyrrhenian Sea, -30 m, 06.1983
(coll. Di Massa, Trieste).
Examined material. Type material; Italy: Ponza
Is., Central Tyrrhenian Sea, -35 m, 1982-83, 3 sh
(BA); Giglio Is., Central Tyrrhenian Sea, -30 m,
05.1983, 1 sh (BA); Ventotene Is., Central Tyrrhenian
Sea, -40 m, Summer 2000, 9 sh (BA); 3 sh (topo-
types) (BA); Zannone Is., Central Tyrrhenian Sea,
-36.5 m, about 60 sh (IN). Egypt: Port Said, 1 sh (IN).
Original description. Amati & Nofroni 1991:
“ Conchiglia di piccole dimensioni, ovato-conica,
elongata, fragile, semitrasparente, non ombelicata.
Protoconca ottusa di 1,20-1,25 giri convessi, lisci;
dimensioni: diametro del nucleo mm 0,13-0,18,
diametro del primo mezzo giro mm 0,25-0,28;
diametro massimo mm 0,30-0,38. Teleoconca di 2-
3 giri convessi, separati da una linea di sutura evi-
dente e leggermente canalicolata. Ultimo giro
abbastanza ampio, pari a circa i 2/3 dell ’altezza
totale. Apertura ovale, angolata posteriormente,
arrotondata e leggermente svasata anteriormente;
labbro ortoclino semplice, tagliente, liscio, legger-
mente inspessito esternamente. Scultura costituita
da numerosi cordoncini spirali (24-30 sull ’ultimo
giro); a forte ingrandimento tutta la superficie, sia
i cordoncini che lo spazio tra gli stessi, appare
percorsa da strie spirali filiformi. Sono presenti
deboli strie di accrescimento ortocline. Colore
biancastro, ma gli esemplari piu freschi appaiono
leggermente giallastri. Opercolo e parti modi
sconosciuti: Dimensioni: h. mm 1,40-2,10; d.mm
0,90-1,15; Rapporto d/2h 0,273-0,343.”
472
Bruno Amati & Italo Nofroni
Distribution and habitat. Central Mediter-
ranean Sea in the infralittoral zone in algal facies
-15/50 m, also reported for Port Said (Egypt).
Remarks. Onoba dimassai may have occasion-
ally an additional labial varix on teleoconch. Com-
pared to that of O. dimassai , the shell of O. josae is
larger and stronger (H 2. 2-3. 2 mm v. H 1.4-2. 2
mm in O. dimassai), deeper suture v. canaliculate
in O. dimassai ; stronger and more spaced spiral
sculpture than in O. dimassai ; outer lip slightly
opisthocline v. orthocline in O. dimassai ; proto-
conch sculptured with 8 thin and irregular spiral
cordlets v. an apparently smooth protoconch (also
at SEM) in O. dimassai. Onoba tarifensis has a
more slender shell with a more cylindrical outline
and a finer, less incised sculpture, consisting in a
higher number of spiral cordlets both on the penul-
timate and on the body whorl (18-24 and 31-38,
respectively v. 8-15 and 18-30 in O. dimassai)', a
protoconch sculptured with 7 thin and irregular
spiral cordlets v. an apparently smooth protoconch
(also at SEM) in O. dimassai. Onoba gianninii has
a larger shell (H 2. 2-2. 6 mm v. H 1 .4-2.2 mm in O.
dimassai), is usually collected at greater depths
(-93/500 m v. -15/50 m for O. dimassai), has a finer
teleoconch sculpture, with a higher number of spiral
cordlets on the body whorl (30-40 v. 18-30 in O.
dimassai), and finally differs in having a clear
umbilical chink, absent in O. dimassai.
Onoba josae Moolenbeek et Hoenselaar, 1987
(Figs. 13-15,27)
Onoba moreleti sensu van Aartsen et al. (1984: 20
fig. 81), not Dautzenberg, 1889
Onoba josae Moolenbeek & Hoenselaar (1987: 153
figs. 6-8)
Iconographic references, van Aartsen et al.,
1984: 20, fig. 81; Moolenbeek & Hoenselaar, 1987:
153, figs. 6-8; Giannuzzi-Savelli et al., 2002: 82,
83, figs. 260-261; Gofas et al., 2011: 193, two un-
numbered figures; Scaperrotta et al., 2013: 62, five
unnumbered figures.
Type locality. Getares, Bay of Algeciras, Spain.
Type material. Not seen. Holotype (ZMA
Moll. no. 3.87.034), 40 paratypes (ZMA Moll. no.
3.87.035), 40 paratypes (coll. H.J. Hoenselaar), 1
paratype (MNHN of Parigi), 1 paratype (IRScNB),
4 juv. paratypes Spain, Getares, 3 paratypes Geta-
res, 28 paratypes Getares (coll. H.J. Hoenselaar),
19 paratypes Getares (ZMA no. 3.87.036 and coll.
H.P.M.G. Menkliorst).
Examined material. Italy: S. Felice Circeo,
Central Tyrrhenian Sea, -30/50 m, 07/1982, 1 sh
(BA); Spain: North of Getares (Cadiz - Mediter-
raneo), legit Gubbioli, 09/1987, 3 sh and 9 frag-
ments, beached (IN); Tarifa -30 m, 1 sh (SB-MS).
Original description. Moolenbeek & Hoense-
laar, 1987: “ Description of the holotype. - Length
2.5 mm, width 1.3 mm (fig. 6 ). Shell oval-conical,
semitransparent with some gloss on the surface,
umbilicum closed. Protoconch dome-shaped, with
about 1 V 4 whorls and with 8 weak and irregular
spirals, protruding very little. Teleoconch about 3
Z 4 whorls with smooth spiral cords. The interstices
are broader than the spiral cords (ratio 1 : 2 ) and
are covered with 7-8 very fine, somewhat undulat-
ing spiral striae. Penultimate whorl with about 9
spiral cords. The upper half of the penultimate
whorl with very weak costae. Body whorl somewhat
convex, with about 22-24 spiral cords. Aperture
subcircular below and rather angular above
(angle about 90°), weakly channeled posteriorly.
Peristome simple, sharp and continuous. Outer lip
clearly opisthocline. Colour white. Operculum,
periostracum and soft parts unknown.’’'’
Distribution and habitat. Strait of Gibraltar,
-30 m. One specimen without soft parts from Latial
coast (Italy), in bioclastic sediment, -30/50 m.
Remarks, van Aartsen et al. (1984) erroneously
identified specimens from Getares (Spain) with O.
moreleti Dautzenberg, 1889 (Ponder, 1985: 162,
figs. 113c, d; Moolenbeek & Hoenselaar, 1987:
155, figs. 1-5), currently considered endemic to the
Azores (originally reported as living at great depths,
but later collected also in shallower waters: Gofas,
1990: 125). So far, O. josae was never reported
from outside the area of Gibraltar Strait. The speci-
men collected in the Central Tyrrhenian and herein
reported is the first record from outside that area.
The record is based on a single, partly broken and
empty adult shell (Fig. 13) so it does not provide
information on the local population viability. The
shell was sorted out from a sample collected by fish-
ing nets residuals from -30/50 m depth, along with
many specimens of O. dimassai. Onoba moreleti
differs from O. josae in having a more slender and
smaller shell, (1.7-1. 9 mm v. 2. 2-3. 2 in O. josae).
The Recent Rissoidae of the Mediterranean Sea. Notes on the genus Onoba s.s. (Gastropoda Prosobranchia)
473
less convex spire, smaller aperture, more or less
dark yellowish colour v. white colour in O. josae,
and a lower number of spiral cordlets both on the
penultimate whorl and on the body whorl (respect-
ively 8-9 and 16-17 v. 9-14 and 22-26 in O. josae).
Onoba josae may have thin subsutural axial ribs
and, very rarely, an additional labial varix.
Onoba guzntani Hoenselaar et Moolenbeek, 1987
(Figs. 21, 22)
Onoba guzmani Hoenselaar & Moolenbeek 1987:
19, figs. 7-12
Iconographic references. Hoenselaar &
Moolenbeek (1987: 19, figs. 7-12); Giannuzzi-
Figures 9-12. Onoba dimassai Amati et Nofroni, 1991: Figures 9, 10. San Felice Circeo, Central Tyrrhenian Sea (Italy) pa-
ratype, height 2.05 mm (BA). Figure. 1 1 . San Felice Circeo, Central Tyrrhenian Sea (Italy) paratype, height 1.85 mm (BA).
Figure. 12. Port Said (Egypt), height 1.7 mm (IN). Figures 13-15. O. josae Moolenbeek et Hoenselaar, 1987: Figure. 13.
San Felice Circeo, Central Tyrrhenian Sea (Italy), height 2.57 mm (BA). Figures 14, 15. Getares Nord, Cadiz (Spain, Me-
diterranean Sea), height 2.6 mm (IN).
474
Bruno Amati & Italo Nofroni
Savelli et al. (2002: 82, 83, fig. 258); Gofas et al.
(2011: 193, 1 unnumbered figure).
Type locality: Tarifa, Spain.
Type material. Not seen. Holotype (ZMA
Moll. no. 3.87.003), 10 paratypes (ZMA Moll. no.
3.87.004) , 25 paratypes (coll. Hoenselaar), 3 para-
types, Spain, Tarifa, IV. 1985 (ZMA Moll. no.
3.87.005) .
Examined material. Spain: Getares North,
Cadiz (Mediterranean) legit Nofroni, 08/1985, 1 sh,
beach (IN); Tarifa -30 m, 1 sh (SB-MS).
Original description. Hoenselaar & Moolen-
beek, 1987: “ Description of the holotype. - Length
1.8 mm, width 0.80 mm (fig. 7). Shell minute, elong-
ate-conic, non-umbilicate, fragile, semitransparent
with some gloss on its surface. Protoconch dome-
shaped, 1 'A whorls, smooth. Teleoconch with 2 3 A
whorls with microscopical pit-marks more or less
forming spirals (fig. 9). Suture deep; whorls
concave. On the base 4 shallow spirals (fig. 10).
Aperture ovate or drop-shaped, with an opistho-
cline outer lip, varix small or lacking, peristome
simple (figs. 10, 12). Operculum, periostracum and
soft parts of the animal unknown.’’''
Distribution and habitat. Reported for the
Strait of Gibraltar and Tangier (Atlantic Morocco)
and Tarifa (Spain) -30 m.
Remarks. Onoba guzmani is very similar to O.
lincta (Watson, 1873), endemic to Madeira (At-
lantic) (Watson, 1873: 387), which has a different
teleoconch sculpture of fine spiral threads and some
strong cords on the base (v. numerous series of
microtubercles spirally arranged, and 4 spiral
cordlets on the base) and the suture more incised,
canaliculated. The protoconch of O. tarifensis it is
sculpted by 7 weak spiral cordlets (v. smooth in O.
guzmani ), a different teleoconch sculpture of 3 1-38
fine spiral cordlets on the last whorl (v. numerous
series of microtubercles spirally arranged, and 4
spiral cordlets on the base) (Hoenselaar & Moolen-
beelc, 1987, figs 3 and 9) and a stronger labial varix.
Onoba tarifensis Hoenselaar et Moolenbeek, 1987
(Figs. 23, 24)
Onoba tarifensis Hoenselaar & Moolenbeek, 1987:
17, figs. 1-6
Iconographic references. Hoenselaar &
Moolenbeek (1987: 17, figs. 1-6); Giannuzzi-
Savelli et al. (2002: 82, 83, fig. 259); Gofas et al.
(2011: 193, 1 unnumbered figure).
Type locality. Tarifa, Spain.
Type material. Not seen. Holotype (ZMA
Moll. no. 3.87.001), 8 paratypes (ZMA Moll. no.
3.87.002), 15 paratypes (coll. Hoenselaar), 1 para-
type, Tarifa, IV. 1985 (coll. Hoenselaar).
Examined material. Spain: Tarifa, Cadiz, legit
Gubbioli, 1988, beach, 1 sh (IN); Tarifa -30 m, 1 sh
(SB-MS); Punta Camero, Getares, 1 sh (CS); Cala
Cica, Getares, 1 sh (CS).
Original description. Hoenselaar & Moolen-
beek, 1987: “ Description of the holotype. - Length
1.55 mm, width 0. 76 mm (figs. 1-4, 6). Shell minute,
elongate-conic, non-umbilicate, fragile and semi-
transparent, some gloss on its surface. Protoconch
dome-shaped, a little less than 1 V 2 whorls with
about 7 smooth spiral cords (fig. 6). Between these
cords there is a microsculpture of rows of exceed-
ingly minute irregular pits, except for the first V 2
whorl which looks smooth. Teleoconch with 2 V 2
whorls, with a very fine spiral sculpture of more or
less smooth spiral cords/ribs; in between these
cords a spongy sculpture of irregular pits (fig. 3).
Penultimate whorl with about 20 spiral cords, body
whorl with about 38 spiral cords. Suture deep, whorls
concave. On the base a strong spiral columel-
lar twist (fig. 4). Aperture ovate with an opistho-
cline outer lip and a strong varix (fig. 5), peristome
simple, weakly angled and channeled posteriorly,
simple and rounded anteriorly. Operculum, peri-
ostracum and soft parts of the animal unknown .”
Distribution and habitat. Reported for the
Strait of Gibraltar (Spain) 0/-30 m.
Remarks. Onoba josae compared to O. tarifen-
sis , has a stronger and larger shell with fewer spiral
cordlets both on the penultimate whorl and on the
body whorl (9-14 and 22-26 respectively v. 18-24
and 31-38 respectively in O. tarifensis). See under
O. guzmani for distinction from O. tarifensis.
Onoba gianninii (Nordsieck, 1974) (Figs. 18-20)
Setia (Crisillosetia) gianninii Nordsieck, 1974: 11,
% 4
The Recent Rissoidae of the Mediterranean Sea. Notes on the genus Onoba s.s. (Gastropoda Prosobranchia)
475
Cingula gianninii (Nordsieck, 1974) (See Verduin,
1984: 61, fig. 25)
Setia gianninii Nordsieck, 1974 (See Amati &
Nofroni, 1991: 32)
Iconographic references. Nordsieck (1974:
11, fig. 4); Verduin (1984: 61, fig. 25); Oliverio
(1988: 113, fig. 1 (operculum and radula)); Amati
& Nofroni (1991 : 32, figs. 6-10); Bouchet & Waren
(1993: 662, figs. 1518, 1519); Ardovini & Cossig-
nani (1999: 38, fig. 035); Smriglio & Mariottini
(2000: 17, figs. 7, 8); Giannuzzi-Savelli et al. (2002:
82, 83, fig. 257); Scaperrotta et al. (2012: 63, 5 un-
numbered figures).
Type locality. Strait of Bonifacio, Corsica,
‘station Kl’, -200/220 m.
Type material. Lectotype (designated by Amati
& Nofroni, 1991) MCZR, 1 paralectotype (coll.
Giannini, Empoli). Bouchet & Waren (1993: 662)
reported some “paratypes” in coll. Carrozza, coll,
van Aartsen and coll. SMNH (not listed in the
original work), which should be more correctly
defined as “paralectotypes”.
Examined material. Lectotype (MCZR);
France: Bastia, Corsica, depth (unprecised) bio-
clastic sands sample, 1 sh (BA); Italy: Capraia Is.,
Northern Tyrrhenian Sea , -400 m, 1 sh (BA); off
Fiumicino, Central Tyrrhenian Sea, -300 m, 4 sh
(BA); Capraia Is., Northern Tyrrhenian Sea, -350
m, 3 sh (IN); Capraia Is., Northern Tyrrhenian Sea,
1 sh (Bogi collection, Livorno).
Original description. Nordsieck, 1974: “Setia
(Crisillosetiaj gianninii n. sp. 3/1,7 mm. Olotipo
nella collezione Giannini. Pallida, semitrasparente;
5 giri molto convessi, il primo (protoconca) at-
tenuate). Sutura profonda. Circa 30/40 strie spirali
sull ’ultimo giro, 15 sul penultimo. Sottile plica
ombelicale. Comparando questa indubbiamente
nuova specie con tutte le altre del sottogenere (v.
tavola R IV del Vol. Ill) ci si avvede che non esiste
alcuna altra specie ad essa avvicinabile sia per la
convessitd dei giri, il numero delle spirali e le
misure della conchiglia.”
Distribution and habitat. Central Mediter-
ranean Sea: Corsica (France), Sardinia, Tuscany
and Latium (Italy), Algeria -93 m. (Bouchet &
Waren, 1993: 663). In bioclastic sediments from
-93/500 m depth.
Remarks. O. gianninii may sometimes have an
additional labial varix. O. oliverioi and O. gianninii
have been found sympatric in the Central Tyrrhe-
nian Sea, in the deepest bathymetric range of O.
gianninii (-200/600 m O. oliverioi v. -93/500 m of
O. gianninii). The shells of these two species are
very similar; O. oliverioi differs mainly for the
smaller size (H 1.6-2. 3 mm at 2.5-3 whorls v. H
2. 5-2. 6 mm at 2.5-3.25 whorls in O. gianninii), the
flatter more sculpted and slightly smaller proto-
conch, (maximum diameter 0.40-0.44 mm v. 0.46
mm (fide Bouchet & Waren, 1993: 663 in O. gian-
ninii), the less slender outline (H/W = 1.44/1.65 v.
H/W = 1.66-1.80 in O. gianninii), and the larger
aperture (H/Ha = 1 .84-2. 16 v. FI/Ha = 2.1 8-2.22 in
O. gianninii). See under O. dimassai for distinction
from O. gianninii.
Onoba oliverioi Smriglio et Mariottini, 2000
(Figs 16, 17)
Onoba oliverioi Smriglio & Mariottini, 2000: 16,
figs. 1-6
Iconographic references. Bouchet & Waren
(1993: 663, figs. 1520, 1521) (sub nomine Onoba
gianninii)', Smriglio & Mariottini (2000: 16, figs.
1 - 6 ).
Type locality. Central Tyrrhenian Sea (41° 5 1 ’
N, 11° 28’ E) off coast of Latium -350/600 m.
Type material. Holotype (MZB 14000); 1
paratype, type locality (MCZR); 9 paratypes, type
locality (CS); 1 paratype, type locality (MO); 1
paratype, type locality (PM).
Examined material. Type material partly ex-
amined: type locality, 9 paratypes (CS), type loc-
ality, 1 paratype (MCZR).
Original description. Smriglio & Mariottini,
2000: “ Shell small (from 1.61 to 2.32 mm in height),
conical-ovate, with a large aperture, blunt apex.
Protoconch dome-shaped consisting of about 1.5
whorls, with a diameter of 400-440 um, sculptured
with 6-8 fine and irregular spiral cordlets. Among
them, several other interrupted fine furrows create
a sort of micro-tuberculated sculpture. Teleoconch
of about 3. 0 rounded convex whorls, the last one is
about 2/3 of the entire length, average ratio H/W =
1.55, average ratio H/Ha = 1.99. Suture pronounced
and shallowly channeled, axial growing lines evid-
476
Bruno Amati & Italo Nofroni
ent, spiral sculpture consisting of about 27 evenly
spaced ribs, with about 2-3 much smaller furrows in
the interspaces. Aperture ovoid, umbilical crevice
slightly visible. Colour milky-white or yellowish
translucent. Operculum and animal unknown
Distribution and habitat. Italy: Central Tyrrhe-
nian Sea: Latium and Sardinia. France: Corsica. On
muddy bottom in a deep-sea coral biocoenosis (bio-
coenosis VB and CB sensu Peres & Picard, 1964)
at a depth of -200/600 m (Bouchet & Waren, 1993:
663; Smriglio & Mariottini, 2000: 16).
Remarks. Onoba oliverioi is characterized by
having a shell with a low H/W ratio and to live at a
maximum depth of -600 m. O. oliveriori differs
Figures 16 17. Onoba oliverioi Smriglio et Mariottini, 2000, Central Tyrrhenian Sea, Latium (Italy) paratype H, height 1.8
mm (CS). Figure 1 8. O. gianninii (Nordsieck, 1 974): Fiumicino, Central Tyrrhenian Sea (Italy), height 2.6 mm (BA). Figures
19, 20. O. gianninii : Bastia, Corsica (France), height 2.5 mm (BA).
The Recent Rissoidae of the Mediterranean Sea. Notes on the genus Onoba s.s. (Gastropoda Prosobranchia)
All
from O. dimassai for the deeper habitat (-200/600
m v. -15/50 m in O. dimassai ), by its higher number
of protoconch whorls (about 1.5 v. 1.2-1.25 in O.
dimassai ), the protoconch sculptured with 6-8 fine
irregulars spiral cordlets v. an apparently smooth
protoconch (also at SEM) in O. dimassai and a larger
maximum diameter of the protoconch (0.40-0.44
mm v. 0.30-0.38 mm in O. dimassai). The number
of the spiral cordlets on the teleoconch is boradly
similar in the two species (about 25-30). See under
O. gianninii for distinction from O. oliverioi.
Onoba sp. (Figs. 25, 26)
Onoba sp. A. Amati & Nofroni, 1991: 34, fig. 5
Figures 21, 22. Onoba guzmani Hoenselaar et Moolenbeek, 1987, Tarifa (Spain), height 2.1 mm (SB-MS). Figures 23, 24.
O. tarifensis Hoenselaar et Moolenbeek, 1987, Tarifa (Spain), height 1.6 mm (SB-MS). Figures 25, 26. Onoba sp. San
Felice Circeo, Central Tyrrhenian Sea (Italy), height 1.7 mm (BA). Figure 27. O.josae Moolenbeek et Hoenselaar, 1987,
Tarifa (Spain), height 2.8 mm (SB-MS).
478
Bruno Amati & Italo Nofroni
Character
Onoba
semicostata
Onoba
aculeus
Onoba
dimassai
Onoba
josae
Onoba
guzmani
Onoba
tarifensis
Onoba
gianninii
Onoba
oliverioi
H
1. 8-3.5
2. 0-4.5
1. 4-2.2
22-3.2
1.4-2. 1
1.45-1.75
2. 5-2. 6
1.61-2.32
W
1.15-1.35
1.35-2.0
0.9-1.15
1.3-1. 5
0.7-1. 1
0.75-0.82
1.5-1.55
1.08-1.4
Ha
1.0-1.15
1.08-1.1
0.75-0.95
1. 1-1.2
0.59-0.95
0.72-0.75
1 .2-1.3
0.85-1.11
R.H/W
2.0-2.59
2.25-2.36
1.56-1.82
1.63-1.81
1.9-1.98
2.0-2.06
1.66-1.8
1.44-1.65
R.H/Ha
2.30-3.04
2-77-2.96
1.88-2.21
2.0-2.22
2.21-2.37
2.22-2.39
2.18-2.22
1.84-2.16
Tcs
yes
yes
no
no
no
no
no
no
St
deep, and
channeled
deep, slightly
to scalanform
slightly
channeled
deep
deep
deep
deep
pronounced
and shallow
channeled
Nw
2.8-3. 8
(5/5.5)*
3.8 (4.5/5. 6)*
2-3
3.25-3.5
2.75-3
2.5-2.75
2.5-3.25
2.5-3
Nspw
12-15
10-14
8-15
9-14
microscopical
pit-marks more
or less forming
spirals
18-24
15-17
11-12
Nslw
25-29
22-24
18-30
22-26
4 shallow spi-
rals on the base
31-38
30-40
23-31
Asc
yes
occasionally,
pronounced
striae of growth
no
occasionally
no
no
no
no
Table I. Ranges of morphometric characters of the teleoconch in Mediterranean species of the genus Onoba. Measurements
in mm. H: height; W: width; Ha: height aperture; R.H/W: ratio height/width; R.H/Ha: ratio height/height aperture; Tcs:
Tendency to curved shells; St: Suture; Nw: number of teleoconch whorls; Nspw: Spiral cords on the penultimate whorl;
Nslw: number of spirals cords on the last whorl; Asc: Axial subsutural cords. *( ) Da Nekhaevet al., 2014. Probably also
include the whorls of the protoconch.
Iconography References. Amati & Nofroni
(1991: 34, fig. 5)
Examined material. Italy: San Felice Circeo,
Central Tyrrhenian Sea, -30/50 m, VIII. 1982, legit
Angelo Amati, 1 sh (BA).
Description. Shell small, fragile, ovate-conical
shape, semi-transparent, non umbilicated. Proto-
conch dome-shaped, paucispiral, with sligthly
twisted nucleus, consisting of just over one whorl
(estimate uncertain, protoconch-teleoconch bound-
ary not clearly visible), 0.25 mm high, with a nuc-
leus diameter of 0. 1 3 mm and a maximum diameter
of 0.32 mm without microsculpture as seen at a
magnification of 90x. Teleoconch of 2.8 convex
whorls with deep suture. Outer lip not tickened
(probably the specimen was not fully adult) ortho-
cline. Sculpture of 24 fine and flat spiral cordlets
on the body whorl, 12 of which above the aperture.
Finer threads covering the entire surface, visible at
a magnification of 90x. Color white. Operculum
and soft parts unknown.
Dimensions: H = 1.7 mm, W =1.05 mm, Ha =
0.84 mm, H/W ratio = 1.619; H/Ha ratio = 2.023.
Distribution and habitat. San Felice Circeo,
Central Tyrrhenian Sea, Italy, a single shell in or-
ganogenic detritus in the infralittoral at -30/50m.
Found sympatric with O. dimassai and O.josae.
Remarks. The single shell, so far known, is pe-
culiar among the European Onoba , in its particular
apex, with a paucispiral protoconch and a twisted
nucleus. It is easily recognizable from all other
species. Onoba dimassai is similar in the fragile
shell, the white colour, the orthocline outer lip and
the teleoconch spiral sculpture. It differs, however,
in the different (not twisted) apex and the wider and
more spaced teleoconch spiral cordlets. O. nunezi
Rolan et Hernandez, 2004, endemic to the Canary
Islands, is slightly smaller (about H 1 .3 mm v. H
1.7 mm in Onoba sp.), is more slender, has a teleo-
conch spiral sculpture of about 10 weak well-
spaced cordlets and the whole teleoconch surface is
covered with finer and more numerous threads
(Rolan & Hernandez, 2004: 174). Manzonia vigoen-
sis (Rolan, 1983) was described as belonging to the
genus Onoba but later Moolenbeelc & Faber (1987)
The Recent Rissoidae of the Mediterranean Sea. Notes on the genus Onoba s.s. (Gastropoda Prosobranchia)
479
and Moolenbeek & Hoenselaar (1992) assigned it
to the genus Manzonia Brasina, 1870; it resembles
Onoba sp. for the general shape of the shell and the
paucispiral protoconch with a twisted nucleus; but
differs for the different sculpture of the teleoconch,
with aligned micro-perforations a thickened outer
lip both typical of the genus Manzonia.
ACKNOWLEDGMENTS
We wish to thank our friends Stefano Bartolini
and Maria Scaperrotta (Florence, Italy) and Carlo
Smriglio (Rome, Italy) for the loan of some material
of their collections and Emilio Rolan (Vigo, Spain)
and Ermanno Quaggiotto (Longare, Vicenza, Italy)
for the bibliographic help. Stefano Bartolini
(Florence, Italy) made some of the light photo-
graphs (Figs. 21-24 and Fig. 27). Marco Oliverio
(La Sapienza Rome University, Italy) commented
an early draft of the manuscript.
REFERENCES
Aartsen J. J. van, Menkhorst H.P.M.G. & Gittenberger E.,
1984. The marine Mollusca of the Bay of Algeciras,
Spain, with general notes on Mitrella, Marginellidae
and Turridae. Basteria, supplement No. 2, 135 pp.
Adams H. & Adams A. 1852. On a new arrangement of
British Rissoae. Annals and Magazine of Natural
History, (2) 10: 358-359.
Adams J., 1797. The specific character of some minute
shells discovered on the coast of Pembrokeshire,
with an account of a new marine animal. Transaction
of the Linnean Society of London, 3: 64-69.
Amati B. & Nofroni I., 1991. Designazione del lectotipo
di Setia gianninii F. Nordsieck, 1974 e descrizione
di Onoba dimassai nuova specie (Prosobranchia:
Rissoidae). Notiziario del C.I.S.Ma, 12: 30-37.
Ardovini R. & Cossignani T., 1999. Atlante delle conchiglie
di profondita del Mediterraneo. L’lnformatore Piceno
Ed. Ancona, Italy, 104 pp.
Avila S.P., Goud J. & Frias Martins de A.M., 2012.
Patterns of Diversity of the Rissoidae (Mollusca:
Gastropoda) in the Atlantic and the Mediterranean
Region. The Scientific World Journal, 30 pp.
Bouchet P, 2014. Onoba H. Adams & A. Adams, 1852.
Accessed through: World Register of Marine Species
at http://marinespecies.org/aphia.php7pMaxlist on
2014-12-12
Bouchet P. & Waren A., 1993. Revision of the Northeast
Atlantic Bathyal and Abyssal Mesogastropoda.
Bollettino Malacologico, suppl. 3: 579-849.
Brasina S., 1870. Bibliotheca malacologica II. Ipsa
Chiereghinii Conchylia. Pisa 280 pp.
Costa (da) E.M., 1778. Hiftoria Naturalis Teftaceoram
Britanniae, or, The British Conchology... Londres,
Imprime pour l’Auteur.254 pp.+ i-vii + pi. xvii.
Dautzenberg P, 1889. Contribution a la faune malacolo-
gique des iles Acores. Resultats des campagnes
scientifiques... Albert Ier, 1: 1-112.
Delongueville C. & Scaillet R., 2001. Faune marine
littorale Svalbard. Novapex, 2: 9-19. 20.
Fretter V. & Graham A., 1978. The prosobranch mol-
luscs of Britain and Denmark. Part 4 - marine
Rissoacea. Journal of Molluscan Studies, Suppl. 6:
153-241.
Friele H., 1886. Mollusca II. The Norwegian North
Atlantic Expedition 1876-1878, 3: 1-44.
Giannuzzi-Savelli R., Pusateri F., Palmeri A. & Ebreo C.,
2002. Atlante delle conchiglie marine del Mediter-
raneo. Edizioni Evolver, Vol. 2, 258 pp.
Gofas S., 1990. The littoral Rissoidae and Anabathridae
of Sao Miguel, Azores. Acoreana, Supplement:
97-134.
Gofas S., Moreno D. & Salas C., 2011. Mollucos marinos
de Andalucia. Universidad de Malaga servicio de
Publicaciones e Intercambio Cientifico. Malaga,
Vol.l, 342 pp.
Golikov A.N., 1987. Class Gastropoda. In: Starobogatov
Ya.I. and Naumov A. D. (Eds). Molluscs of the White
Sea. Nauka, Leningrad: 41-148.
Golikov A.N. & Kussakin O.G., 1978. Shell-bearing
gastropods of the intertidal zone of the seas of the
USSR. Leningrad, Nauka, 292 pp.
Gould A. A., 1841. Report on the Invertebrate of Mas-
sachusetts comprising the Mollusca, Crustacean,
Annelida, and Radiate. Published agreeably to an
order of The Legislature, by the commissioners on
the zoological and botanical survey on the state.
Cambridge, 373 pp.
Gray J.E., 1847. A list of the genera of Recent Mollusca,
their synonyma and types. Proceedings of the
Zoological Society of London 1847: 129-242.
Hoenselaar H.J. & Moolenbeek R.G., 1987. Two new
species of Onoba from southern Spain (Gastropoda:
Rissoidae). Basteria, 31: 17-20.
Hoisaster T., 2009. Distribution of marine, benthic,
shellbearing gastropods along the Norwegian coast.
Fauna norvegica, 28: 5-106.
Ingolfsson A., 1996. The distribution of intertidal macro-
fauna on the coasts of Iceland in relation to temper-
ature. Sarsia, 81: 29-44.
Jeffreys J.G., 1867. British Conchology or an account of
The Mollusca which now inhabit the beitish isles and
the surkounding seas. Vol. IV. Marine Shells, London,
486 pp.
Jeffreys J.G., 1887. A complete Catalogue of British
Mollusca. Gloucester: Herbert W. Marsden, 53 pp.
480
Bruno Amati & Italo Nofroni
Loven S., 1846. Nordens Hafs-Mollusker [list of species
bears subtitle “Index Molluscoram litora Scand-
inaviae occidentalia habitantium”] Ofversigt af
Kongl. Vetenskaps-Akademiens Forhandlingar 3:
134-160,3: 182-204.
Moller H.P.C., 1842. Index Molluscoram Groenlandiae.
1-24. Typis expressit I.G.Salomon. Hafniae.
Montagu G., 1803. Testacea Britannica, part 2. White,
London, pp. 293-610.
Moolenbeek R.G. & Faber M.J., 1987. The Macarone-
sian species of the genus Manzonia (Gastropoda:
Rissoidae). Part III. De Kreukel, 10: 166-179, pis. 2, 3.
Moolenbeek R.G. & Hoenselaar H.J., 1987. On the
identity of Onoba moreleti Dautzenberg, 1889
(Gastropoda: Rissoidae) with the description of
Onoba josae n. sp. Basteria 51: 153-157.
Moolenbeek R.G. & Hoenselaar H.J., 1992. New addi-
tions to the Manzonia fauna of the Canary Islands
(Gastropoda: Rissoidae). Pulicacoes Ocasionais da
Sociedad Portuguesa de Malacologia, 16: 13-16.
Nekhaev I.O, Deart Yu. V. & Lubin P.A., 2014. Molluscs
of the genus Onoba H. Adams & A. Adams,
1852 from the Barents Sea and adjacent waters
(Gastropoda: Rissoidae). Proceedings of the Zoolo-
gical Institute RAS. Vol. 318, No. 3: 268-279.
Nordsieck F., 1974. Molluschi dei fondali della platea
continentale fra la Corsica e la Sardegna. La
Conchiglia, 61: 11-14.
Oliverio M., 1988. On the sistematic of "Setia" gianninii
(Gastropoda: Prosobranchia). Bollettino Malacolo-
gico, 24: 112-114.
Oliverio M., Amati B. & Nofroni I., 1986. Proposta di
adeguamento sistematico dei Rissoidaea (sensu
Ponder) del Mar Mediterraneo. Parte I: famiglia Ris-
soidae Gray, 1847 (Gastropoda: Prosobranchia).
Notiziario C.I.S.Ma. VII- VIII (8-9): 35-52 [1985-86],
Ponder W.F., 1985. A Review of the Genera of the
Rissoidae (Mollusca: Mesogastropoda: Rissoacea).
Records of the Australian Museum. Suplement 4: 1-
221 [1984],
Ponder W.F. & Worsfold T.M., 1994. A Review of the
Rissoiform Gastropods of Southwestern South
America (Mollusca, Gastropoda). Contributions in
Science. Natural History Museum of Los Angeles
County, 445: 1-62.
Reeve L.A., 1878. Conchologia Iconica: or illustrations
of the Shells Molluscous Animals. Vol. XX Mono-
graph of the genus Rissoa, PI. XIII. London.
Rolan E., 1983. Moluscos de la Ria de Vigo. I Gastero-
podos. Thalassas, Santiago de Compostela; 1, Anexo
1, 383 pp.
Rolan E., 2008. The genus Onoba (Mollusca, Caeno-
gastropoda, Rissoidae) from NW Spain, with the
description of two new species. Zoosymposia, 1:
233-245.
Rolan E. & Hernandez J.M., 2004. Descripcion de
una nueva especie de Onoba (Mollusca, Rissooidea)
de las Islas Canarias, con comentarios sobre otras
especies proximas. Iberas, 22: 173-179.
Scaperrotta M., Bartolini S. & Bogi C., 2012. Accresci-
menti. Stadi di accrescimento dei molluschi marini
del Mediterraneo. Volume IV, L’lnformatore Piceno,
184 pp.
Scaperrotta M., Bartolini S. & Bogi C., 2013. Accresci-
menti. Stadi di accrescimento dei molluschi marini
del Mediterraneo. Volume V, L’lnformatore Piceno,
192 pp.
Schiotte T. & Waren A., 1992. An annotated and illus-
trated list of the types of Mollusca described by
H.P.C. Moller from West Greenland. Meddelelser om
Gronland. Bioscience, 35: 1-33.
Smriglio C. & Mariottini P., 2000. Onoba oliverioi n. sp.
(Prosobranchia, Rissoidae) a new gastropod from the
Mediterranean. Iberas, 18: 15-19.
Sneli J.-A., Schiotte T., Jensen K.R., Wikander P.B.,
Stokland 0. & Sorensen J., 2005. Marine Mollusca
of the Faroes. Annales Societatis Scientiarum
Faeroensis, Suppl., 32: 1-190.
Stimpson W., 1851. On several new species of shells
from the northern coast of New England. Proceedings
of the Boston Society of Natural History, 4: 12-18
(1851-1854).
Templado J. & Rolan E., 1986. El genero Onoba H. &
A. Adams, 1854 (Gastropoda, Rissoidea) en las
costas europeas (1). Iberas, 6: 117-124.
Verduin A., 1984. On the taxonomy of some recent
European marine species of the genus Cingula s.l.
Basteria, 48: 37-87.
VerrilA.E., 1884. Second catalogue of Mollusca recently
added to the fauna of the New England coast and the
adjacent parts of the Atlantic, consisting mainly of
deep-sea species with notes on others previously re-
corder. Transactions of the Connecticut Accademy,
6: 139-294.
Waren A., 1973. Revision of the Rissoidae of the
Norwegian North Atlantic Expedition 1876-1878.
Sarsia, 53: 1-14.
Waren A., 1974. Revision of the Artie- Atlantic Rissoidae
(Gastropoda, Prosobranchia). Zoologica Scripta, 3:
121-135.
Waren A., 1996. New and little known Mollusca from
Iceland and Scandinavia. Part. 3. Sarsia, 81: 197-245.
Watson R.B., 1873. On some marine Mollusca from
Madeira, including a new Eulima, and the whole of
the Rissoae of the Group of Islands. Zoological
Society of London, pp. 361-391.
Biodiversity Journal, 2015, 6 (1): 481-490
Monograph
The endemic door snail of Marettimo (Egadi Islands, Sicily,
Italy): Siciliaria ( Siciliaria ) scarificata (L. Pfeiffer, 1 856) (Pulmo-
nata Clausiliidae)
Fabio Liberto 1 , Maria Stella Colomba 2 ,Agatino Reitano 3 , Salvatore Giglio 4 & Ignazio Sparacio 5
1 S tra d a Provinciale Celafu-Gibil manna, 93 - 90015 Cefalu, Palermo, Italy; email: fabioliberto@ yahoo.it
"Universita di Urbino, DiSB, via Maggetti 22 (loc. Sasso), 61029 Urbino, Italy; e-mail: mariastella.colomba@ uniurb.it
'ViaGravina, 77 - 95030 Tremestieri Etneo, Catania, Italy; e-mail: tinohawk@yahoo.it
4 C ontrada Settefrati - 90015 Cefalu, Palermo. Italy; e-mail: hallucigenia@ tiscali.it
'via E. Notarbartolo, 54 int. 13 - 90145 Palermo. Italy; e-mail: isparacio@ inwind.it
ABSTRACT The door snail Sicilicirici ( Siciliaria ) scarificata (L. Pfeiffer, 1856) (Pulmonata Clausiliidae)
is re d e s c rib e d . The species is endemic to Marettimo (Egadi Islands, Sicily, Italy) and it is
the only one of the genus Siciliaria V e st, 1 867 living in this island. Siciliaria Scarificata c an
be morphologically identified by the presence of a high columellar lamella, ascending in a
double “S” curve, a wide anterior upper palatal plica, long basal plica, sulcalis present;
clausilium plate distally less narrowed; genitalia are characterized by very short bursa
copulatrix duct; short diverticulum of bursa copulatrix; penial papilla conic and short. Notes
about its taxonomy, biology and conservation status are also provided.
KEY WORDS Door snail; Siciliaria', island endemism; taxonomy; conservation status.
Received 21.02.2015; accepted 23.03.2 0 15; printed 30.03.2015
Proceedings of the Eighth Malacological Pontine Meeting, October 4th- 5th, 2014 - San Felice Circeo, Italy
INTRODUCTION
Vest (1867) d e sc rib ed the genus Sicilicirici for a
group of door snail species from Sicily with S.
grohmanniana Rossmiissler, 1 836 type species.
Adolf Schmidt (1 86 8 ) classified the seven Si-
cilian species known so far, into two groups which
mainly differ by the formation of the clausilium
plate and by sculpture,developmentoflamellae (in-
serta, inferior lamella) and palatal plicae. The first
group is reported as: “Form enkreis of Septem-
plicata ” which includes ClausUia grohmanniana,
C. septemplicata Philippi, 1 8 3 6 , C. calcarae Phil-
ippi. 1 844, C. confinata Benoit, 1 859 (= scarificata
L . P fe iffer, 1 8 5 6), and C. tiberii A. Schmidt, 1868;
the second group as “Form enkreis of CrdSSicOStdtd”
with C. CrdSSicOStdtd L . Pfeiffer, 1 85 6 and C. Yiobilis
L . P fe iffer, 1 8 4 8.
O. Boettger (1877) named as Sicilidrid Vest,
1 867 sensu stricto the “Form enkreise of Septeifl-
plicdtd" , and as TrindCrid O. Boettger, 1 8 77 the
“Form enkreise of CrdSSWOStdtd” . Nordsieck (1979)
listed the same species as O. Boettger (1 877) and
reunited the species groups (Siciliaria s . str., TrindC-
rid (preoccupied) = Sicddid To m lin ) because S. Cdl-
CdCdC has an intermediate morphological position.
Nordsieck (2002) listed 12 species of Sicilidrid
s.str., sub Chdrpentierid ( Sicilidrid ), and classified
them in two species groups, based on some shell
characters.
Nordsieck (2007), in his catalog on world
Clausiliidae, listed 12 species with 7 subspecies of
Sicilidrid s.str., even as Chdrpentierid ( Sicilidrid ) ■.
Sicilidrid {Sicilidrid) cdlcdrae cdlcdrae, S. cdlcdme
belliemii (Brandt, 1 9 6 1 ) , S. crdssicostdtd, S.
eminens (A. Schimdt, 1 8 6 8 ) , S. ferrox (Brandt,
482
Fabio Liberto et alii
196 1 ), S. grohmanniana , S. leucophryna (L.
Pfeiffer, 1 8 62), S. YlobiUs, S. ribewthi (Brandt,
196 1), S. scarificata (L . Pfeiffer, 1 85 6), S. septem-
plicata septemplicata , S. septemplicata alcamoensis
(Brandt, 1961 ), S. septemplicata hemmeni Beck-
m ann, 2004, S. spezialensis (H . N ordsieck, 1 984), S.
tiberii tiberii, S. tiberii scalettensis Beckmann, 2004 .
This checklist is confirmed by Bank (2011) and
by N ordsieck (2013).
The genus Sidliarias. str. is endemic to Western
Sicily, from Caccamo in the East to the island of
M arettim o in the West, and from San Vito lo Capo
in the North, to C a s te 1 v e tr an o and Ribera in the
South. The hot spot of biodiversity are the moun-
tains in the northern part, whereas in the central and
southern area (Sicani Mountains) the presence of
Siciliaria is discontinuous. The genus Sidliarias. str.
is reported in Quaternary deposits of Palermo (D e
Gregorio, 1 886: Monte Pellegrino; 1 927: Pietrazzi,
Bellolampo; our personal data: Mount Catalfano)
and in the Qua ternary deposit ofWied tal-Bahrija in
the Island of M alta (Giusti et al., 1 995).
Siciliaria scarificata w as discovered by the Si-
cilian naturalistLuigiBenoit (1804-1890) who dis-
tributed shells of this door snail to his m alacologists
colleagues under the name of Clausilia COnfinata.
Luis Pfeiffer (1856) published the first valid d e scrip -
tion w ith th e name C. Scarificata (Fig. 1) reporting
its distribution as “Habitatin Sicilia”. Subsequently,
L. Pfeiffer ( 1 859) specified the distribution as “ in
insula Maretima Siciliae” . h owever in the course of
1800s and up to about the 1970s most authors used
the nam e C. COllfinata B enoit or the incorrect spelling
C. Sacrificata Benoit, 1875 (see below). The original
description and all subsequent descriptions were
based on shell features; while genitalia were never
described and illustrated.
This paper is intended to redescribe this species
in detail (shell and genitalia) and also provided notes
about its taxonomy, biology and conservation status.
MATERIAL AND METHODS
All living specimens were relaxed in water and
then preserved in 80% ethanol. Five specimens
were anatomically investigated under a Leica
MZ12.5 s tere o m ic r o s c o p e using scalpel, scissors
and needles. Empty shells were kept dry, and have
been measured with a digital gauge. The plicae and
lamellae were studied breaking the shells with a
scalpel. The method of calculating the number of
whorls by Kerney & Cameron (1979) was used.
Shell measures were based on the study of 20 spe-
cimens. Photos were carried out with a Panasonic
Lurnix DMC-FZ20 digital camera. Anatomical
details were drawn using a W ild camera lucida. The
collection data are listed as follows: State, region,
municipality, locality, altitude, dates, collection and
number of specimens in parentheses. Toponyms
(place-names) are reported following M ap “IGM 1 :
25000, Isola di Marettimo, sheet 256 IV - N.O.”.
Each locality and/or collection site is named in the
original language (italian).
Voucher specimens were stored in the following
Museums and private collections: F. Liberto,
Cefalu, Italy (LC); Museo N aturalistico F. Mina
Palumbo, Castelbuono, Italy (MNMP);A. Reitano,
Tremestieri Etneo, Italy (RC); I. Sparacio, Palermo,
Italy (SC).
CONCHOLOGICAL ACRONYMS. AUPP:
Anterior upper palatal plica; CL: columellar
lamella; D: shell width; H: shell height; L: lunella;
LPP: lower palatal plica (basal plica); PL: parietal
lamella; PLL: parallel lamella; PP: principal plica;
PUPP: posterior upper palatal plica; SCL: sub-
columellar lamella; SL: spiral lamella; SUL:
sulcalis; SP: sutural plica. ANATOMICAL AC-
RONYMS. BC: bursa copulatrix; BCD: divertic-
ulum of bursa copulatrix; DBC: duct of the bursa
copulatrix; E: epiphallus; FO: free oviduct; G:
penial papilla; GA: genital atrium; P: penis; PR:
penial retractor muscle; V: vagina; VD : vas deferens.
SYSTEMATICS
Family CLAUSILIIDAE J.E. Gray, 1855
Genus Siciliaria vest, 1867
Type species: Clausilia grohmanniana Ross-
m assler, 18 3 6
Siciliaria ( Siciliaria ) scarificata (l . p feiffer, 1 8 5 6 )
Clausilia scarificata, L. Pfeiffer, 1856: 1 8 5 , PI. 2,
figs. 20-22 - Habitat in Sicilia
Clausilia scarificata, L. Pfeiffer, 1 8 5 9 : 765-766 -
Habitat in insula Maretima Siciliae
Clausilia confinata, Benoit, 1 8 5 9 : PI. 6, fig. 6
Clausilia scarificata, Kuster, 1860-1861: 298, pi.
34, figs. 1-3 - Insel M aretima
The endemic door snail of Marettimo (Egadi Islands, Sicily, Italy): Siciliaria scarificata (Pulmonata Clausiliidae)
483
1
ir
Marettimo
Island
/ J\
is* »
Mediterranean Sea
y , m
\ 1
,W
/
^ ° r
Egadi r
\rehipelagoS^
Sicily
2
a
V — S
Figure i. Reproduction of original drawing of SiciHcirici ( Siciliaria ) Scarificata (L . Pfeiffer, 1 856). Figure 2. Map of
Western Sicily, the arrow shows the position of Marettimo. Figures 3, 4. Siciliaria SCa.rifiCQ.ta in natural habitat. Figures 5,
6. Landscape of Marettimo, slope with Mediterranean maquis.
Claus ilia confmata, vest, 1 8 6 7 : 1 6 7
Clausilia confinata, A . Schmidt, 1 8 6 8 : 40-42
Claus ilia confinata, Appeiius, 1 8 6 9 : 1 7 3
Medora scarificata, Kobe it, 1 8 7 1 : 39
Clausilia sacrificata, Benoit, 1 8 7 5 : 1 5 2 - isola di
M are tim o
Siciliaria confinata, m oiiendorff, 1 875: 17
Clausilia confinata, L. Pfeiffer, 1 8 77: 523 - Ins.
M a re tim a Siciliae
Clausilia {Siciliaria) confinata, o. Boettger, 1 8 7 7 :
33, “gruppe Siciliaria ” - Sicilien
Clausilia confinata, W esterlund, 1 8 7 8 : 2 0 - Sicilia
Clausilia sacrificata, O. Boettger, 1 8 7 9 : 89, PI. 172,
fig . 1731 -InselM are tim o im Wes ten von S icilien
Clausilia ( Siciliaria ) sacrificata, Kobe it, 1 8 8 1 : 78 -
M aretim o
Clausilia confinata, Benoit, 1 8 8 2 : 1 05 - isola di
M are tim o
484
Fabio Liberto et alii
Clausilia confinata, Westerlund, 1 8 84: 46 - Mare-
tim o b e i S ic ilie n
Clausilia ( Siciliaria ) confinata, Monterosato, 1 8 9 2 :
28 - Isola di Maretimo
Clausilia ( Siciliaria ) confinata, westerlund, 1892:48
Clausilia (Siciliaria) confinata, westerlund, 1 9 0 1 :
39-40, 180 - I. Maretimo
Delima ( Siciliaria ) scarificata, Wagner, 1924: 124
- Insel M aretim o im w . Von Sizilien
Clausilia (Siciliaria) confinata, Sacchi, 1 9 5 5 : 23
Siciliaria confinata, Sacchi, 1956 : 8-9
Siciliaria Confinata, Sacchi, 1957: 673
Delima (Siciliaria) confinata , Aizona, 1 97 1 : 91 ,
sectio Siciliaria - Is. M arettimo
Siciliaria (Siciliaria) scarificata, n o rd s ieck , 1979 :
259
Siciliaria (Siciliaria) scarificata, Manganeiiietai.,
1995: 24,47 - Isola diMarettimo (Egadi)
Charpentieria (Siciliaria) scarificata, Nordsieck,
2002: 33-34
Charpentieria (Siciliaria) scarificata, Beckmann,
2004: 186, 188 - Insel M are ttimo
Siciliaria scarificata , Fiorentino et ai., 2004 -
M arettim o
Charpentieria (Siciliaria) scarificata, Nordsieck,
2007: 54
Charpentieria (Siciliaria) scarificata, Bank, 20 1 1 :
2 3 - S ic ily
Siciliaria scarificata, Weiter-Shuites, 2012 : 342 -
S. Italy, M arettim o island
Siciliaria (Siciliaria) scarificata, n 01 -dsieck, 2013 :
1-14
Type locality. M arettimo (Egadi Islands,
Sicily, Italy). This species was comunicated by the
naturalist Luigi Benoit to L. Pfeiffer who published
the first valid description reporting its distribution
as “Habitat in Sicilia” (L. Pfeiffer, 1 85 6).
Examined material. Italy, Sicily, Favignana,
Island of Marettimo (Egadi Islands), Punta Troia,
50 m, VIII. 1997, 3 shells (RC); idem, Case
Romane, 200 m, VI. 2005, 23 shells (RC); idem,
VI. 2005, 4 shells (RC); idem, from Case Romane
to Monte Falcone 300-680 m, 30. V. 2010, 11 spe-
cimens, 60 shells, (LC 802 1-8096); idem,
VI. 2005, 3 shells (MNMP); idem, contrada Pelosa,
60 m, 30. V. 2010, 16 specimens and 42 shells (SC);
idem, Case Romane, 200 m, 30. V. 2010, 18 shells
(SC).
Original description. L. Pfeiffer (1 85 6): “ T.
rimata,fusiformis, truncata, solida, confertim plicato-
costulata, purpurascenti-fusca; spira ventrosa,
sublate decollata; sutura albo-papillata; anfr.
superst. 7 1/2 convexiusculi, ultimus basi breviter
cristatus; apertura piriformis; lamella supera
exigua, marginem non attingens, infera valida,
arcuatim ascendens; lunella distincta, angusta,
flexuosa; plicae palatales 3, suprema elongata,
secunda brevior, antice callosa, tertia infera, sub-
columellari parallela; perist. hepaticum, con-
tinuum, breviter solutum, undique expansum et re-
flexiusculum. - Long, (trunc.) 17, diam. 5 mill. Ap.
5 mill, longa, 4 lata ” .
Diagnosis. Terrestrial pulmonate snail with
shell sinistral, fusiform, b r o w n -p u rp lis h in color;
aperture with five lamellae (on parietum and
columellar side) and lunella and five plicae (on
palatum); in particular high columellar (lower
parietal) lamella, ascending in a double “S” curve;
a wide anterior upper palatal plica; long basal plica;
short sulcalis; genitalia are characterized by very
short bursa copulatrix duct, short diverticulum of
bursa copulatrix; penial papilla conic and short.
Description (Figs. 7-14). Shell sinistral, fusi-
form, elongated, generally decollated, rather thick
and robust, b ro w n -p u rp lis h in color, with apertural
margin light brownish; obtuse apex; external sur-
face with transverse ribs, 8.5 ribs per 2 mm of the
penultimate whorl (10 specimens); spire with 9-10
slightly convex whorls (7 in decollate shells),
slowly and regularly growing; sutures shallow, with
slightly evident papillae (papillae more numerous
along sutures from level with first 3-7 whorls);
basal keel little distinct; umbilicus closed; aperture
oval, with five lamellae (on parietum and columel-
lar side) and lunella and five plicae (on palatum).
On palatum there is a short lunella and starting from
suture: a thin sutural plica very close to suture; a
well raised principal plica; a wide anterior upper pal-
atal plica, separated from or connected with upper
palatal plica; long basal plica, internal beginning of
which is joined to the base of lunella; a short sul-
calis (Figs. 15, 16). A relatively conspicuous callos-
ity on the upper external border of palatum
embedding external apexes of upper palatal plica
and principal plica. On parietum, starting from su-
ture; there are: parallel lamella very thin or absent;
non emergent spiral lamella in the centre of pari-
The endemic door snail of Marettimo (Egadi Islands, Sicily, Italy): Siciliaria scarificata (Pulmonata Clausiliidae)
485
Figures 7-10. Shells of SiciUcirici (SiciliCLfici) SCCLrifiCQ-tCl (L . Pfeiffer, 1 856), Island of Marettimo, S icily, Italy
(CL 8032), H: 17.8 m m , D : 5 mm. Figures 11-14. idem, (CL 8033), H: 18.1 mm, D : 4.6 mm.
486
Fabio Liberto et alii
AUPP
17
CL
PL
PLL
SL
■ta
X
u
18
L
21
SCL
Figures 15-2 1. SiciUarid (SiciUariO) scarificata (L . Pfeiffer, 1 8 5 6), Island of Marettimo, S icily, Italy. Figures 15, 16
palatum (CL 8083, 8084). Figures 17, 18: parietum (CL 8085, 8088), Figures 19-21: clausilium (CL 8094-8096).
The endemic door snail of Marettimo (Egadi Islands, Sicily, Italy): Siciliaria scarificata (Pulmonata Clausiliidae)
487
Figure 22. Genitalia o f Siciliaria ( Siciliaria > scarificata (L. Pfeiffer, 1 856), island of m arettimo, Sicily, Italy. Figure 23. Internal
ornam entation of penis, w ith penial papilla (same specimen of Fig. 22). Figure 24. Internal ornamentation of epiphallus (CL 8022).
488
Fabio Liberto et alii
etum ; tooth-like (upper) parietal lamella; high
columellar (lower parietal) lamella, ascending in a
double “ S ” curve; scarcely emergent subcolumellar
lamella (Figs. 17, 18). Peristome continuous,
thickened,reflected, fused above to last who rl w all.
Normal type clausilial apparatus, with palatal edge
of clausilium plate somewhat bent up, outer corner
more or less pointed, sutural angle bent up (Figs.
19-21). The outer edge of the clausilium plate rests
against the lunella and the sulcalis in the closed
p o sitio n .
Body. Animal narrow, posteriorly pointed, skin
yellowish in color with bro w n-greysh tubercles;
foot narrow with sole paler than body, bipartite by
an indistinct longitudinal central groove and with
margins divided by small parallel radial groove (5
specim. in alcohol preserved).
Genitalia (Figs. 22-24). General scheme of
semidiaulic monotrematic type. Gen it alia consisting
of large ovotestis with many close acini; long thin,
convoluted hermaphrodite duct; very large, albumen
gland; well developed o v isp e rrn id u c t, formed by
female portion externally regularly subdivided and
spaced by annular constrictions, large prostatic
portion and seminal groove externally not visible;
slender free oviduct (2.6 mm); bursa copulatrix
comp lex consist of slender copula to ry duct (2.9 mm)
which branches in very short bursa copulatrix duct
with leaf-like bursa copulatrix (2 mm), and slight
longer diverticulum of bursa copulatrix (3.6 mm in
length); vagina (1.8 mm in length) uniform in
diameter for almost its entire length; vas deferens
long and slender, entering epiphallus; epiphallus (3.9
mm) divided by point of insertion of robust penial
retractor muscle into conical proximal portion and
shorter cylindrical distal portion; a swelling is
present at the transition p e n is-ep ip h allu s ; cylindrical
penis (2.4 mm in length) slightly wider than vagina.
Internal walls of penis with two furrows; relatively
short, conic penial papilla with rounded apex (Fig.
23). Internal walls of epiphallus covered with small
papille and crossed by two low pleats (Fig. 24).
Distribution and Biology. Siciliaria SCari-
ficata is endemic of the Island of Marettimo, the
wes tern most of the Egadi Islands, in Wes tern Sicily
(Fig. 2). It lives in limes to ne habitat with Mediter-
ranean maquis, on walls and in the crevices of cal-
careous rocks, under stones, in conoids of debris
and at the base of cliffs (Figs. 3-6).
Remarks. Nordsieck (2002) classified the 12
species of Siciliaria s. str. in two groups, based on
some shell characters. The first group is named
“ flobiliS-CCllcCirCie” and is characterized by:
columellar lamella (inferior lamella) low to mod-
erately high, mostly only one anterior upper palatal
plica present, clausilium plate distally not m arkedly
narrowed, outer corner blunt to pointed. This group
is further divided into two subgroups: “ tlobiUs” sub-
group (S. nobilis, S. spezialensis , S. crassicostata ,
and S. eminens) has palatal edge of clausilium plate
not upbent; “ CCllcGrCl6” subgroup ( S . CCllcClVCie, S.
tiberii, and S. leucophryna) has palatal edge of
clausilium plate more or less upbent. The second
principal group is named “ grohmanniana” ( S .
grohmanniana , S. septemplicata , and S. scari-
ficata), it is characterized by columellar lamella
high, two anterior upper palatal plicae present,
clausilium plate distally narrow ed with outer corner
more or less pointed. Sidlicirici ferrOX and S.
riberothi were not included in none of these groups
because of ambiguous character combinations.
Siciliaria scarificata is considered transitional
to the two principal groups, because it has inferior
lamella less high, second anterior upper palatal
plica missing, clausilium plate distally less nar-
rowed. Nordsieck (2013) re affirm s S. Scarificata is
closely related to the other species of the “ grohman-
niana ” group.
At present it is difficult to establish the real
affinity between these species using only morpho-
logical observations. Consequently, discussion of
the relationships o f S. Scarificata is postponed to
when more data (molecular data in particular) will
b e a v ailab le .
Here we add some morphological data for the
“ grohmanniana ” group not considered by
Nordsieck (2002; 2013 ). Siciliaria grohmanniana
has a small "inserta lamella" (0.7 mm) placed
between the columellar lamella and the spiral
lamella, running from the point of arrest of clausil-
ium outward. This lamella, reported by A. Schmidt
(1868) for S. grohmanniana and also for S. septem-
plicata, is absent in S. scarificata. Siciliaria
grohmanniana and S. septemplicata have a shorter
sulcalis compared with S. Scarificata. Welter-
Shultes (2012) reports on a su b c lau s tr alis , which we
don’t recognize.
The genitalia of S. Scarificata are similar to
those of other species of Siciliaria s. str. known: S.
The endemic door snail of Marettimo (Egadi Islands, Sicily, Italy): Siciliaria scarificata (Pulmonata Clausiliidae)
489
septemplicata { Wagner, 1 9 1 3 , pi. 572, fig. 14), S.
grohmanniana (Wagner, 1925 , pi. 1 , fig. 8) S.
calcarae (w agner, 1925 , pi. 3 , fig. 25 ), S. ferrox
(Brandt, 1961, p. 7, 13, fig. 1). The duct of the
bursa copulatrix is very short, and the diverticulum
is slightly longer of the duct of the bursa copulatrix
+ bursa copulatrix; cylindricalpenis; slight swelling
at the conjunction p e n is -ep ip h allu s .
Westerlund (1 892) described two varieties of S.
scarificata (sub confinata )■. C. confinata merens
Westerlund, 1 8 92, locus typicus “Sicilien, in der
Provinz Palermo”, which is a synonym of S.
leucophryna (see Nordsieck, 2013), syntype in
Goteborg Natural History Museum n° 2638, and C.
confinata commeata westerlund, 1 892, locus
tipicus “Sicilien, bei Trabia” which is a probable
older synonym of S. fCTYOX B randt, 1961 (Reitano
et al., 2007, Nordsieck, 2013).
Conservation status. Although S. scarificata
has a scattered distribution over the whole island,
its limited distribution to Marettimo justified an
assessment as Lower Risk (Near Threatened) [NT,
nt]. The Island of Marettimo is included in the
SICp “Isola diMarettimo” (ITA 010002) and in the
ZPS “Arcipelago delle Egadi - area marina e ter-
restre” (ITA 0 1 0027), however S. Scarificata is not
protected by any specific regulam entation or law,
although it should be strongly reco mended. Sug-
gested measures include sympathetic habitat man-
agement and population monitoring.
REFERENCES
Alzona C ., 1971. Malacofauna italica. Catalogo e biblio-
grafia dei molluschi viventi, terrestri e d'acqua dolce.
Atti della Societa Italiana di Scienze Naturali e del
M useo Civico diStoria Naturale di Milano, 111: 1-433.
Appelius F.L., 1869. Bibliografia. Monographia Heli-
ceorum viventium, Auctore Ludovico Pfeiffer, D.
Cassellano. Vol. V e VI, (S upplem entum tertium I,
II). Bullettino malacologico Italiano, 2: 1 70- 1 73.
Bank R.A., 2011 . Fauna Europaea: M ollusca Gastropoda.
Fauna Europaea version 2.4. Checklist of the land
and freshwater Gastropoda of Italy. Fauna Europaea
Project: 1-49.
Beckmann K.H., 2004. Zur Verbreitung der endem ischen
n o rd w e s ts iz ilian is ch en Clausiliidae der Untergattung
Clmrpenteria (Siciliaria) mit Beschreibung von zwei
neuen Unterarten (Gastropoda: S ty lo m m ato p ho ra :
Claus iliidae). Archiv fir rMolluskenk unde, 133: 185-191.
Benoit L., 1 85 7-1 862. 1 1 1 u s tra z io n e sistematica critica
iconografica de’ testacei estramarini della Sicilia
Ulteriore e delle isole cir costa nti. Gaetano Nobile,
Napoli, 248 pp., 8 pis. [Quaderno l.o: pp. i-xvi, 1-52,
pls.l, 2 (1 857); Quaderno 2.o: pp. 53-116, pis. 3, 4
(1857); Quaderno 3.o: pp. 117-180, pis. 5, 6 (1859);
Quaderno 4o: pp. 181-248, pis. 7, 8 (1862). The
publication date of the Table 9, 11, 12 is unknown.
Benoit L., 1875. Catalogo delle conchiglie terrestri e flu -
viatili della Sicilia e delle Isole circostanti. B ullettino
della Societa Malacologica italiana, 1: 1 29-1 63.
Benoit L., 1882.Nuovo catalogo delle conchiglie terrestri
e fluviatili della Sicilia o continuazione alia illus-
trazione sistematica critica iconografica de’ testacei
estramarini della Sicilia Ulteriore e delle isole cir-
costanti. D'Amico, Messina, VI+176 pp.
BoettgerO., 1877. Clausilienstudien. - Palaeontographica
(Neue Folge) Supplement 3: [1-2], 1-222, Taf. 1-4.
Boettger O., 1879. Gattung ClciUSiliCL Drap. In: Ross-
massler, Iconographie der Land- & Siisswasser-
Mollusken mit vorzirglicher Beriicksichtigung der
europaischen noch nicht abgebildeten Arten von E.
A. Rossmassler fortgesetzt von Dr. W. Kobelt.
W iesbaden, Kreidel, 6 (4/6): 52-153, Taf. 167-178.
Brandt R. A., 1961. Diagnose n neuerClausiliiden. Archiv
furMolluskenkunde,90: 1-23,2 Pis.
De Gregorio A., 1886. Into mo a un deposito di roditori e
di carnivori sulla vetta di Monte Pellegrino con uno
schizzo s in c ro n o g ra fic o del calcare p o s tp lio c e n ic o
della vallata di Palermo. Atti della Societa Toscana
di Scienze Naturali, 8: 1-39.
De Gregorio A., 1927. Molluschi terrestri e fluviali
quaternari di Sicilia. Annales de geologie et de
p ale o n to lo g ie , 44: 1-22, 4 Pis.
Fiore ntino V., Cianfanelli S., Manganelli G. & Giusti F.
2004. I molluschi non marini delle isole Egadi
(Canale di Sicilia): biodiversita e conservazione.
Poster XIV congresso nazionale Societa Italiana di
E c o lo g ia, Siena.
G iusti F., M anganelli G . & Schembri P.J., 1995. The non-
marine molluscs of the Maltese Islands. Museo
Regionale di Scienze Naturali, Torino, Monografie,
1 5: 1-607.
Kerney M.P. & Cameron R.A.D., 1979. A field guide to
the land snails of Britain and North-west Europe.
Collins, London, 288 pp.
Kobelt W., 1871. Catalog der im europaischen Faunen-
gebiet lebenden Binnenconchylien. Mit besonderer
Beriicksichtigung der in Rossmassler' s Sammlung
enthaltenen Arten. Fisher, Cassel, [1-16], 150 pp.
Kobelt W., 1881. Catalog der im europaischen Faunen-
gebiet lebenden Binnenconchylien. 2. Auflage.
Fischer, Cassel, 294 pp.
Kiister H. C., 1847-1862. Die Schliessschnecken und die
verwandten Gattungen {ClciUSilicl, Baled, CylitldrclltL
490
Fabio Liberto et alii
Megaspira) . In Abbild ungen nach der Natur mit
B e sc h reib u n g e n . S y s te m atis c h e s C o n c h y lie n -C ab in e t
von Martini und Chemnitz, 14: 1-355, Pis. 1-38.
Manganelli G., Bodon M., Favilli L. & Giusti F., 1995.
Gastropoda Pulmonata. In: M in elli A. , Ruffo S.& La
Posta S. (Eds.), Checklist delle specie della fauna
italiana, 16. Calderini, Bologna, 60 pp.
M Ollendorff von O., 1875. Studien zur Systematik der
Clausilien. Nachrichtsblatt der deutschen Malako-
zoologischen G esellschaft, 7: 17-24.
Monterosato T., 1892. Conchiglie terrestri delle Isole
adiacenti la Sicilia. A tti della Reale Accademia di
Scienze Lettere e Belle A rti di Palermo, 2: 1-34.
Nordsieck H., 1979. Zur Anatomie und Systematik der
Clausilien, XXI. Das System der Clausilien II: Die
rezenten europischen Clausilien. Archiv fur M o 1 -
luskenkunde, 109: 249-275.
Nordsieck H ., 2002. Contributions to the knowledge of
the Delimini (Gastropoda: Stylommatophora:
Clausiliidae), Mitteil ungen der Dtschen Malako-
zoologischen G esellschaft, 67: 27-39.
Nordsieck H ., 2007. Worldwide Door Snails (Clausil-
iidae), recent and fossil. ConchBooks, Hackenheim,
2 14 pp .
Nordsieck N . , 2013. Revisory remarks on the species of
Siciliaria Vest from N. W. Sicily. Available at.:
w w w .hnords.de/printable/5356429d6b 1 1 adcOb/5356
42aldcl365e05/index.html - last access: 10. III. 2015.
PfeifferL., 1 856. Berichtiiberweitere Mittheilungen des
Herrn Zelebor. M ala k o z o o lo g is c h e Blatter, 3: 175-
186, PI. 1,2.
Pfeiffer L., 1 859. Monographia Heliceorum viventium.
Sistens descriptiones system aticas etcriticas omnium
huius familiae generum et specierum hodie cog-
nitarum. Lipsiae, 5: x i i + 565 pp.
Pfeiffer L., 1 8 77. Monographia heliceorum viventium.
Sistens descriptiones system aticas etcriticas omnium
huius familiae generum et specierum hodie cog-
nitarum. Lipsiae, 8: 2 + 729 pp.
Reitano A., Liberto F. & Sparacio I., 2007. Nuovi dati su
M o 11 u sc h i terrestri e dulciacquicoli di Sicilia. 1° Con-
tribute (Gastropoda Pro sobranchia N e o ta e n io g lo s s a ;
Gastropoda Pulmonata Basommatophora, Stylomma-
tophora). II N aturalista siciliano, 3 1: 3 1 1-3 30.
Sacchi C.F., 1955. Contributo alia conoscenza faunistica
della Campania. Ricerche malacologiche nella
regione sorrentina. II. Appunti b io g e o g rafic i . Annu-
ario dell’Istituto e Museo di Zoologia Universita di
Napoli, [ 1 954] 6: 14 pp.
Sacchi C ,F., 1 956. I Molluschi terrestri nelle relazioni
b io g e o g ra fic h e tra Italia ed Africa. Estratto da:
Archivio Botanico e Biogeografico Italiano, 32.
Valbonesi, Forli, 31 pp.
Sacchi C.F., 1957. Componenti storiche e fattori ambien-
ali nelle fisionomie zoologiche della Sicilia. Bollet-
tino di Zoologia, 24: 633-683.
Schmidt A., 1868. System der europaischen Clausilien
und ihrer nachsten Verwandten. Th. Fischer, Cassel,
[1-2], 176 pp.
Vest W.V., 1 867. Uber den S c h lie s s a p p ara t der Clausi-
lien. Verhandlungen und Mittheilungen des sieben-
biirgischen Vereins flir Naturwissenschaften zu
H erm annstadt, 18: 161-174.
Wagner A.J., 1913. Iconographie der Land- & Siiss-
w asser-M ollusken mit vorzuglicher B e rti c k s ic h ti-
gung der europaischen noch nicht abgebildeten Arten
von E.A. Rossmassler, fortgesetzt von Dr. W. Kobelt.
Die familie der Clausiliidae. Neue Folge. 21: 1-20,
PL 5 7 1 -580.
Wagner A. J., 1924. System atisches Verzeichnis der mir
heute bekannten Arten und Forme n der Clausiliidae,
III. Annales Zoologici Musei Polonici Historiae
N aturalis, 3: 99-1 26.
WagnerA., 1925. Studien uberdie Systematik, Stammes-
geschichte und g e o g rap h is c h e Verbreitung des Genus
Delima (Hartmann) A.J. Wagner. Annales Zoologici
Musei Polonici Historiae N aturalis, 4: 1-73, Pis. 1 -
17 .
W e lte r- S c h u lte s F., 2012. European non-marine
molluscs, a guide for species identification. Planet
Poster Editions, Gottingen, 760 pp.
Westerlund C .A ., 1 8 7 8. Monografi ofver p alao ark tis k a
regionens Clausilier. Berling, Lund, 1-27, 1-1 84,
1-12 p p .
Westerlund C.A., 1884. Fauna der in der Palaarktischen
Region (Europa, Caukasien, Sibirien, Turan, Persien,
Kurdistan, Arm enien, M e s o p o ta m ie n , Kleinasien,
Syrien,Arabien, Egypten, Trip o lis, Tunesien.Agerien
und Marocco) lebenden Binnen-conchylien.
Karlskrona, Vol. 4, Gen. BttleCl Prid. & ClciUSiUci
Drap., vii + 212 + 18 pp.
Westerlund C.A., 1892. Spicilegium malacologicum.
Neue Binnen-Conchylien in der palaarktischen
Region. III. Nachrichtsblatt der Deutschen Malako-
zoologischen Gesellschaft, 24: 185-201.
Westerlund C.A., 1901. Synopsis molluscorum in regione
palaearctica viventium ex typo ClciUSiUci Drap. -
Zapiski Imperatorskoj Akademii Nauk po fiziko-
matematicheskomu Otdeleniyu - Memoires de 1 1
Academie Imperiale des Sciences de St. Petersbourg,
Classephysico-mathematique 11: [1,2], [1-37], 1-203.