The family of Mutillidae (Hymenoptera Aculeata), Velvet ants (Mutillidae) arc a family of insects belonging to the order of Hymenoptera and within

that to the Vespoidea superfamily of the Aculeata suborder, in which the female ovipositor serves as a sling, Mulillids are known already from Late

Cretaceous and Eocene. The family Mutillidae, with about 4200 described species, is among the largest solitary wasp families parasitoids on enclosed

hosts at preimaginal stage of other holometabolous insects, buth the estimated number of species is much higher The mutillid faunas of the Palaearctic,

Oriental and Afrotropical regions are the most similar on the tribal and generic levels and represent the nucleus of the world fauna. Velvet ants occur in all

zoogeograph i cal regions, but the vast majority of them are found in tropical and subtropical regions of the world being abundant in arid and semiarid

areas. There are more than 500 species of mutillids in 54 genera known in the Palaearctic region. About 1 70 species were found in Europe. A peculiar

feature for the family is the extreme sexual dimorphism: the males are winged (rarely brachypterous or apterous) and possess a normal apocritan

mesosoma, The females are wingless, and appear to be rather setae, colourful ants. They can easily be differentiated from ants by the lack of petiole

nodes, which are present on all ant species. In addition, the mesosomatic (thoracic) segments are completely fused and have at most two segments. The

metasoma (abdomen) contains six visible terga (dorsal surface of any body segment) and a "felt line" of dense, closely appressed hairs is located

laterally on the second metasomatic tergum. Females are generally densely pubescent and predominantly black, brown, or reddish, often marked with

bright spots or bands that are usually white, yellow, or red. Because of their extreme sexual dimorphism, sex associations cannot be made on

morphological grounds alone; most species and even many genera are known from one sex only. In most cases the males are so different from the

females that only males and females caught in the act of copulation can be claimed to belong to the same species with certainty. It may be revealed in

many cases that certain male and female specimens thought to be distinct species may belong to the same species, therefore the number of existing

species may be lower than the actual number. Knowledge of the natural history' of mutillid wasps is usually restricted to the habitat in which the adults

are collected and/or the flowers where are established to feed. Recently it has been estimated that the hosts of only about 2-3 % of the described mutillid

species are known, and for some subfamilies there arc no records at all. All the species are parasites in the nests of solitary and social bees, wasps, ants or

beetles. Females of mutillids arc apterous and spend most of the daytime in burrows in ground or under grass turf's. Problems encountered by any female

mutillid mainly involve finding suitable host individuals and penetrating their enclosures for egg laying. Solitary' hosts arc generally scattered and often

concealed. Mutillids apparently use odour signals (kairomomes) while actively running in suitable locations for finding such hosts, and must spend

much time searching, with little prospect of finding numerous hosts. Females search for the ground nests of hosts and deposit their eggs in the brood

cells. They usually search for mature larv ae (or prepupae) and paralyze the larva with the sting and after that they lay an egg into the brood cell. The

mutillid larvae are always ectoparasite ids of host stages which are enclosed in some sort of package (cell, cocoon, puparium, ootheca) and which are not

actively feeding. The quantity ofaccessible food determines the resulting size of the eventual imago. Males are winged and live only a few' weeks. They

search for virgin females to mate them usually flying not high above the ground. Some mostly tropical species use special dances when males fly and

carry females connected on the abdomen within the mating process, otherwise in species living in sand dunes and deserts females carry apterous males

on their backs. European mutillids usually have one generation per year (univoltine): both sexes emerge in summer, males die in the beginning of

autumn, females overwinter and attack nests of hosts in early or late spring,

Marcello Romano, Piazza A. Cataldo, 10- 90040 Capaci (Palermo), Italy; e-mail: marcellr@libero.it Cover: photos by M. Romano

Biodiversity Journal, 2013, 4 (4): 443-450

Preliminary contribution to the knowledge of Coleoptera Bu-

prestidae from Atlantic rainforest regions of Rio Doce and Ita-

colomi State Parks of Minas Gerais, Brazil.The genera Agrilus

Curtis, 1825, Autarcontes Waterhouse, 1 887, and Geralius Ha-

rold, 1 869

Gianfranco Curletti 1 *, Servio P. Ribeiro 2 & Letizia Migliore 2

'Museo Civico di Storia Naturale, via S. Francesco di Sales, 188 - 10022 Carmagnola, Italy; e-mail: gianfranco.curletti@yahoo.it

2 Laboratorio de Ecologia Evolutiva de Insetos de Dossel, Departamento de Biodiversidade, Evolusao e Meio Ambiente, ICEB,

Universidade Federal de Ouro Preto, Campus Morro do Cruzeiro, Ouro Preto, MG, Brasil.

‘Corresponding author:

ABSTRACT A checklist of species belonging to Agrilus Curtis, 1 825, Autarcontes Waterhouse, 1 887, and

Geralius Harold, 1869 genera found in Rio Doce and Itacolomi Estadual Parks is given. 19

species are listed, 5 of which are new for the science and here described: Agrilus pirilampo

n. sp., A. disorientatus n. sp., A. coal n. sp., A. rarestriatus n. sp., A. taediosus n. sp. Most

species were found in one sampling season and reflect a quite favourable rainforest for xy-

lophagous species. The high dominance of Leguminosae large tree species could be related

to this pattern.

KEY WORDS Coleoptera; Agrilus', Autarcontes', Geralius', Brasil; Rio Doce; Itacolomi.

Received 19.09.2013; accepted 29.10.2013; printed 30.12.2013

INTRODUCTION

The present fauna has been collected in two

State parks covered predominantly with Atlantic

rainforest, at upper and mid-basin of the Doce river,

in the Central region of Minas Gerais State, Brazil.

The forests of these parks have been monitored for

insect herbivores and ants for many years. For the

Rio Doce State Park, a lowland forest, 1 3 years of

research were accomplished on canopy insects from

three distinct geomorphologies, as part of a long

term study of natural (lake-forest) and human made

(pasture-forest) ecotones, based on insect interac-

tions with the main dominant tree species, Mabea

fistulifera Mart. (Euphorbiaceae) and Byrsonima

sericea DC (Malpiguiaceae), respectively. In the

montane Park of Itacolomi, another long term re-

search area, comparative entomological studies

were set in three permanent plots in distinct succes-

sional forests, running since 2006.

Any of those entomological researches showed

that, in general, the number of species per sample

unit (tree crowns) were below expected for a tropi-

cal forests (Campos et al., 2006a; Ribeiro et al.,

2008; Fagundes et al., 2012; Neves et al., 2013),

and different causes may influence these findings.

In Rio Doce, the rainforest may be extremely re-

cent, coming from a xeric habitat that suffered a cli-

matic change no longer than 9 thousand years ago

(Overloop, 1981; Werneck et al., 2011). In Ita-

colomi, the locally cold and unpredictable winter

may be related to a severe lack of large tropical in-

vertebrate species (Espirito Santo et al., 2012),

along with the loss of connections with other moun-

444

G. ClIRLETTI, S.P. RiBEIRO & L. MlGLIORE

tain ecosystems due to hundreds of years since first

human occupation. Nevertheless, both forests may

favour xylophagous insects due to an intense gap

dynamic found in both.

The present study shows preliminary results from

a first survey on xylophagous, namely Buprestidae

species, in both regions.

ACRONYMS AND ABBREVIATIONS. The

Natural History Museum, London, England

(BMNH); Museo Civico di Storia Naturale, Carma-

gnola, Italy (MCCI); Musee National d’Histoire

Naturelle, Paris, France (MNHN) ; Museu de Zo-

ologia Uni vers itade Sao Paulo, Brazil (MZUSP);

Narodni Muzeum, Prague, Czech Republic

(NMPC); Parque Estadual Itacolomi, Brazil (PEIT);

Parque Estadual Rio Doce, Brazil (PERD).

Study area

Rio Doce State Park - PERD. This is the largest

continuous preserved Atlantic rainforest fragment

(35,974 ha) in Minas Gerais State, Brazil

(19°48 , 18”-19°29 , 24”S, 42°38 , 30 ,, -42°28 , 18 ,, W).

Altitude varies from 230 to 515 m above sea level;

climate type is Aw (tropical hot semi-humid), with

wet seasons from October to March and dry seasons

from April to September (Gilhuis, 1986). The pre-

dominant vegetation is semi-deciduous seasonal for-

est, with 20% to 50% deciduous trees (Veloso et al.,

1991). Also, this is the largest natural lake system of

South America, and 10% of the Park is covered with

lakes, result of changes in the Doce river positioning

and blocking of old drainages, along with increasing

rainfall (see below).

Itacolomi State Park - PEIT. The Itacolomi

State Park is a 7,543 ha conservation area, at south

of Serra do Espinhago (20°22'30"S-43°32'30 M W).

The area is located in a transition between Atlantic

Forest and Cerrado, with physiognomies of grass-

land and montane evergreen forest types, with alti-

tudes between 700 and 1700 m above sea level. The

climate is typically seasonal tropical, understanding

the types Cwa and Cwb of Koppen, in lowest and

highest areas, respectively (SEMAD/IEF/PRO-

MATA, 2007). The studied areas are reported as

human disturbed since the colonial times. More re-

cently, it belonged to a farm of tea, Camellia sinen-

sis (L.) Kuntze. After the abandonment of the

culture about 40 years ago, the species Eremanthus

erythropappus DC (Asteraceae) colonized the most

former plantations, resulting in a monodominant

forests, which have been gradually substituted by a

more diverse wet forest along time. In deep valleys,

a more wet and diverse forest always existed and

was preserved due to difficult access and water pro-

tection for the farms. Flooded small valleys are

dominated by Myrcia species and resemble sub-

tropical flood forests.

General ecology and biogeographic evolution

For reasons still under study, these forest canopies

tend to have a particularly poorer leaf-herbivore in-

sect fauna than observed in canopies of wet and

close equatorial forests (Campos et al., 2006b,

Ribeiro et al., 2008). However, the present findings

for Buprestidae are astonishingly diverse, especially

for Rio Doce State Park. Most species were found

in one sampling season and reflect a quite favourable

rainforest for xylophagous species. The dominance

of large deciduous and semi-deciduous Legumi-

nosae tree species could be related to this pattern.

Although this is a wet continuous forests, Rio Doce

Park is quite heterogeneous, and the area where

most of species were found was probably a drier

vegetation type no longer than four thousand years

ago (Overloop, 1981; Wemeck et al., 2011). The cli-

mate shift during the Holocene contributed to the

origin of the lakes in the Doce river mid-basin, that

define enormously this landscape (Pflug, 1969;

Meis & Monteiro, 1979; Meis & Tundisi, 1986;

Peronico & Castro, 2008). Hence, adaptations to a

climate shifted toward a wetter seasonal ecosystem

may be on course, and severe and frequent tree

death is normally observed even within natural and

well preserved forest spots. Campos et al. (2006a,

b) and Ribeiro et al. (2008) suggest that high gap

formation influences canopy insect fauna. In addi-

tion, Castro et al. (2012) found a highly diverse lit-

ter ant fauna, and clearly an intense debris

production ought to influence such finding.

MATERIALS AND METHODS

The material was taken by several methods dur-

ing expeditions in November 2010, March 2012,

and September 2012. The priority was the use of

entomological umbrella, which was positioned

Coleoptera Buprestidae from Rio Doce and Itacolomi State Parks, Brazil. The genera Agrilus, Autarcontes, and Geralius 445

below branches of trees in the upper canopy or in

the foliage in the borders of forest as mentioned

above. Systematic samplings by beating on the

branches have been developed along both projects

on several marked trees. In addition, qualitative and

semi-quantitative methods, such as spider net,

sticky traps, breeding larvae and direct sampling

were added specifically for Buprestidae sampling.

The use of sticky traps, contrary to African and

Palaearctic faunae, was ineffective, probably be-

cause the Neotropical Buprestidae species are not

attracted by the colours, surely not by the yellow

colour (G. Curletti, pers. obs.).

The specimens collected are preserved dry,

after extraction of genitalia. The pictures are made

with microscope and assembled with Combine Z4

program.

RESULTS

List of species found in PERD

Geralius furciventris (Chevrolat, 1838)

Examined material. 1 specimen female: Brazil,

Minas Gerais, Marlieria, PERD, SD - Mab. PL7B -

6. II. 2001, Ribeiro S.P. leg.

Remarks. Species (Chevrolat, 1838: 88 sub

Stenogaster ) known for Argentina, Bolivia, Brazil,

Paraguay, Pem. Biology unknown.

Autarcontes mucoreus (Klug, 1825)

Examined material. 1 specimen female: Brazil,

Minas Gerais, Marlieria, PERD, SD - Mab. PL7B -

6. II. 2001, Ribeiro S.P. leg.

Remarks. Species described of Minas Gerais,

Uberaba (Klug, 1825: 428, sub Buprestis). Known

for Brazil only.

Agrilus {Agrilus) gracchus Obenberger, 1935

Examined material. Three specimens females:

Brazil, P. E. Rio Doce, 20.XI.20 10, G. Curletti and

L. Migliore leg. (MCCI).

Remarks. Species described from Minas Gerais,

Mar de Hespanha (Obenberger, 1935: 135); type

NMPC. Endemic of Brazil. Biology unknown.

Agrilus {Agrilus) zikani Obenberger, 1935

Examined material. 1 female specimen: Brazil,

P. E. Rio Doce, 20. XI. 20 10, G. Curletti and L.

Migliore leg.

Remarks. Species described from Minas Gerais

(Obenberger, 1935: 126); type NMPC. Endemic of

Brazil. Biology unknown.

Agrilus {Agrilus) pirilampo n. sp.

Examined material. Holotype female (Fig. 1):

Brazil, P. E. Rio Doce, Perd Vinhatico, 08.IX.2000,

S. Ribeiro leg. (MZUSP).

Description of holotypus. Length 14 mm,

elongate. Dorsal color black with green reflections

on elytra and two orange spots at the apex. Vertex

strongly depressed, as wide as half anterior edge of

pronotum. Frons concave with orange round spot at

the base. Clypeus small, without carina. Pronotum

wider anteriorly, with lateral edges subrounded but

sinuate before the base forming posterior angles

acute. Anterior edge incise in middle. Disc gibbous,

with two round depressions in the medium-anterior

part. Sculpture composed by thin and thickened

striae. Premarginal carinula absent. Lateral carinae

very open ahead, separate at the base. Prosternal

gular lobe small, widely sinuate. Scutellum con-

cave, small, with transversal carina nearly visible.

Elytra with both apices rounded and microdenticu-

late. Ventral side black, glabrous, with a lateral

orange spot at the sides of the antepenultimate ven-

trite. Same spot at the corresponding laterotergum.

Legs black, with all claws mucronate. Metatarsus

shorter than metatibia. First metatarsomere longer

than the sum of the following three (1 >2+3+4).

Etymology. The two yellow spots on elytra are

very evident and for this reason A. pirilampo n. sp.

remember the lights produced by the Elateridae

species belonging to the Pyrophorini tribe, named

pirilampos by the Brazilian people.

Remarks. For the dimensions, color and elytral

spots, A. pirilampo n. sp. appears unique on the

South American fauna.

Agrilus {Agrilus) sedyi Obenberger, 1933

Examined material. 1 male: Brazil, P. E. Rio

Doce, 19.XI.20 10, G. Curletti and L. Migliore leg.

446

G. ClIRLETTI, S.P. RlBEIRO & L. MlGLIORE

Remarks. Species described from Sao Paulo

(Obenberger, 1933 a: 12), endemic of Brazil.

Agrilus {Agrilus) luederwaldti Obenberger, 1933

Examined material. 1 male: Brazil, P. E. Rio

Doce, 19.XI.20 10, G. Curletti and L. Migliore leg.

Remarks. Species described from Santa Catha-

rina (Obenberger, 1933a: 11), endemic of Brazil.

Agrilus {Agrilus) arnus Gory, 1841

Examined material. 1 female, Brazil, P. E. Rio

Doce, 19.XI.20 10, G. Curletti and L. Migliore leg.

Remarks. Species widespread (Gory, 1841:

232), quoted from Brazil, Argentina, Colombia.

Type in MNHN.

Agrilus {Agrilus) disorientatus n. sp.

Examined material. Holotype female (Fig. 2):

Brazil, P. E. Rio Doce, 16.III.2012, G. Curletti and

L. Migliore leg. (MZUSP).

Description of holotypus. Length 6.8 mm.

Elytra brown with 8 (4+4) spots of yellow pube-

scence along the suture. Pronotum as wide as 1/3 of

anterior edge of pronotum, darker than the vertex,

reddish. Frons black, with red reflections at the base,

glabrous. Clypeus separate by frons by a transversal

small carina. Antennae serrate from IV antennomere.

Pronotum wider anteriorly, posterior angles right.

Yellow pubescence along the lateral edges; disc with

regular, transverse, thin sculpture. Premarginal cari-

nae entire, joined to lateral edge before the half

length. Marginal carinae subparallel, separate from

base. Prostemal gular lobe rounded; prostemal plate

with lateral edges sinuate in middle. Scutellum trans-

versely carinate. Elytra with apex acute, ending by a

tip. The pubescent spot are placed respectively in the

humeral callus, at 1/3, at 2/3 and before the apex: the

basal (humeral) couple rounded, the second elongate,

the third oval, the fourth smaller, less visible, reduced

to a short line along the suture. Ventral side bronze,

with a line of white pubescence in median part of the

basal stemite and a round spot at the margins of all

remaining. Legs brown like the ventral side;

metatarsus shorter than metatibia, with a first article

longer than the sum of the following two (l>2+3).

All claws bifid, but with the teeth separate.

Etymology. Disorientatus = out of bearings.

The specimen was found in a dead branch of Platy-

menia foliosa Benth incised and killed by Oncideres

sp. (Cerambycidae). All specimens had already

come out from the wood; only this specimen, still

alive, was in the pupal cell, destined to a sure

death because it was turned to the inside of the

branch.

Remarks. Several South-American species have

four spots on elytra, but A. disorientatus is unique

for the shape of the apical acute apex.

Agrilus {Agrilus) consentaneus Kerremans, 1897

Examined material. 3 specimens, 2 males

and 1 female, found in PERD the 07-15.11.2011,

L. Migliore leg.

Remarks. To this species (Kerremans, 1897: 89)

we attribute the specimens collected.

Agrilus {Agrilus) gileti Obenberger, 1933

Examined material. 1 male: Brazil, Minas

Gerais, PERD, 19. XI. 20 10 G. Curletti and L.

Migliore leg. (MZUSP).

Remarks. Species described from Sao Paulo

(Obenberger, 1933a: 19); type inNMPC.

Agrilus {Agrilus) coal n. sp.

Examined material. Holotype male (Figs. 3,

4): Brazil, P.N. Rio Doce, 19 0 45’48”S-

42°37’54 ,, W, 18.XI.2010, 288 m, G. Curletti and

L. Migliore leg. (MZUSP).

Description of holotypus. Length 5.6 mm.

Pronotum and elytra black. Vertex as wide as 1/3 of

anterior edge of pronotum with punctiform sculp-

ture. Frons golden, flat, glabrous. Clypeus without

transversal carina. Antennae serrate from article IV,

black at the base, gold from antennomere V. White

pubescence on the cheek. Pronotum wider in the

middle of length, with lateral margins rounded but

sinuate before the base forming acute angles. Disc

with two elongate small depressions in middle,

after the vertex and before the scutellum. Sculp-

ture formed by transversal striae alternate to inter-

striae composed by other more superficial and thin

striae in number of 3-5 every interstria. White pu-

bescence little visible along the lateral edges. Pre-

Coleoptera Buprestidae from Rio Doce and Itacolomi State Parks, Brazil. The genera Agrilus, Autarcontes, and Geralius 447

marginal carina entire, joined to the edges before

the half length. Marginal carinae subparallel, sepa-

rate from the base. Prostemal gular lobe entire but

not rounded. Scutellum transversely carinate. Elytra

glabrous; apex hardly denticulate with median tooth

bigger and stumpy. Ventral side bronze with latero-

marginal spot of white pubescence on the ventrites.

Legs with all claws bifid, but with internal median

and posterior teeth shorter and squat. Metatarsus

shorter of metatibia; first metatarsomere longer than

the sum of the following two (l>2+3). Aedeagus

fusiform, median lobe acute (Fig. 4).

Etymology. After the black colour.

Remarks. For the shape, color, abdominal pu-

bescence, A. coal n. sp. is similar to A. lestageanus

Obenberger, 1935 from Brazil (type in NMPC).

This last differs especially for having different

pronotum sculpture, apical apex, frons green.

Agrilus {Agrilus) rarestriatus n. sp.

Examined material. Holotype female (Figs. 5,

6): Brazil, Minas Gerais, PERD, 19.XI.2010 G.

Curletti and L. Migliore leg. (MZUSP).

Description of holotypus. Length 5.8 mm.

Pronotum dark green like the vertex, but reddish

along the sides, elytra black. Vertex as wide as the

half of anterior edge of pronotum. Frons glabrous,

red wine brilliant. Clypeus without transversal ca-

rina. Antennae brief, serrate from antennomere IV.

Pronotum wider anteriorly, sinuate before the pos-

terior angles that are acute. Anterior edge protrud-

ing in middle, between the eyes. Disc with

superficial striae, with unusual structure remembering

the afrotropical A. buani Curletti et Vayssieres,

2007, but more rarefied (Fig. 6). Premarginal ca-

rina absent; marginal carinae joined at the base.

Prostemal gular lobe cut in middle of anterior edge.

Scutellum transversely carinate. Elytrae with each

apex rounded and denticulate. Disc with white pu-

bescence veiy short but well visible, regularly dis-

posed. Lateroterga whit longer white pminose

pubescence. Abdomen dark bronze, with pubes-

cence like elytra. Same colour in the legs: all claws

like A. disorientatus n. sp. Metatarsus shorter than

metatibia; first metatarsomere as long as the sum

of the following two (1=2+3).

Etymology. After the peculiar structure of the

pronotum striae.

Remarks. For the pronotum sculpture A. rares-

triatus n. sp. is unique in the Brazilian agrilofauna.

The specimen was collected on herbaceous small

bush, still unidentified, that is probably the host

plant.

Agrilus {Agrilus) taediosus n. sp.

Examined material. Holotype male (Figs. 7,

8): Brazil, Minas Gerais, PERD, 17. XI. 20 10 G.

Curletti and L. Migliore leg. (MZUSP).

Description of holotypus. Length 6.5 mm.

Dorsal colour dark bronze. Vertex as wide as the

half of the anterior edge of pronotum. Eyes small,

little visible from the back. Frons green-bronze,

glabrous, widely furrowed, without transverse ca-

rina before the clypeus. Antennae bronze, brief, ser-

rate from IV antennomere. Pronotum gibbous,

depressed at the sides, wider anteriorly, with curved

lateral edges and sinuate before the posterior angles

that are acute. A wide superficial, longitudinal fur-

row in middle. Sculpture composed by numerous

transversal striae alternate to small points. Pre-

marginal carina entire, superficial, little visible.

Marginal carinae joined at the base. Prostemal gular

lobe cut in middle of anterior edge. Scutellum trans-

versely carinate. Elytrae with each apex rounded

and denticulate. Elytra stumpy, glabrous, with apices

rounded and microdenticulate. White pubescence

on the lateroterga only, with glabrous abdomen. All

claws mucronate. Metatarsus shorter than metatibia.

First metatarsomere as long as the sum of the fol-

lowing two (1=2+3). Aedeagus small and thin with

median lobe acute (Fig. 8).

Etymology. From the Latin taedium = bore-

dom, for the want of remarkable characters.

Remarks. On the whole of cited characters, A.

taediosus n. sp. is close to A. needhami Obenberger,

1933b described from Sao Paulo (type in NMPC).

The two species are principally distinguishable for

having the very different shape of aedeagus.

Agrilus {Agrilus) vanini Curletti et Migliore, in press

Examined material. 1 female: Brazil, Minas

Gerais, PERD, (19 0 45’48”S - 42 0 37’54”W),

20.XI.20 10, G. Curletti and L. Migliore leg. (type

in MZUSP).

448

Figure 1 .Agrilus ( Agrilus ) pirilampo n. sp., holotype. Figure 2. A. (A.) disorientatus n. sp., holotype. Figure 3. A. (A. ) coal

n. sp., holotype. Figure 4. Idem, aedeagus in dorsal view, 1.2 mm. Figure 5. A. (A.) rarestriatus n. sp., holotype. Figure 6.

Idem, pronotum. Figure 7. A. (A. ) taediosus n. sp. holotype. Figure 8. Idem, aedeagus in dorsal view, 1.5 mm. Scale bar =

1 mm.

G. CURLETTI, S.P. RlBEIRO & L. MlGLIORE

Coleoptera Buprestidae from Rio Doce and Itacolomi State Parks, Brazil. The genera Agrilus, Autarcontes, and Geralius 449

List of species found in PEIT

Agrilus (Agrilus) sp.

Examined material. 1 female specimen: Brazil,

Minas Gerais, PEIT, 11.XI.2007, G. Curletti andL.

Migliore leg.

Remarks. This specimen close to A. gileti

Obenberger, 1933, belongs probably to a new

species, but the morphological characters are no

sufficient for the description.

Agrilus (Agrilus) clazon Obenberger, 1933

Examined material. 1 female specimen: Brazil,

Minas Gerais, PEIT, 11.XI.2007, G. Curletti andL.

Migliore leg.

Remarks. Species described from Sao Paulo

(Obenberger, 1933b: 80); type in NMPC. The speci-

men from Itacolomi is more bronze, probably for

the recent capture.

Agrilus (Agrilus) badius Kerremans, 1897

Examined material. 1 male specimen: Brazil,

Minas Gerais, PEIT, 11.XI.2007, G. Curletti and L.

Migliore leg.

Remarks. Species described from Minas Gerais

(Kerremans, 1897: 79, Caraga); type in BMNH.

Agrilus (Agrilus) octavius Obenberger, 1935

Examined material. 1 female specimen, Brazil,

Minas Gerais, PEIT, 15-30.III.2012, G. Curletti and

L. Migliore leg.

Remarks. Species described from Sao Paulo

(Obenberger, 1935: 124); type in NMPC.

ACKNOWLEDGMENTS

We wish to thanks the staff members for the

help in the field, especially Geraldo dos Santos

Adriano. The IEF, the State Institute of Forestry

and the administration of Rio Doce and Itacolomi

State Park, for providing permits and logistical fa-

cilities for the work. We thank FAPEMIG and

PELD/CNPq for financial support. SPR is a re-

searcher funded by CNPq.

REFERENCES

Campos R.I., Vasconcelos H.L., Ribeiro S.P., Neves F.S.,

Soares J.P., 2006a. Relationship between tree size and

insect assemblages associated with Anadenanthera

macrocarpa. Ecography, 29: 442-450.

Campos R.I., Soares J.P., Martins R.P., Ribeiro S.P.,

2006b. Effect of habitat structure on ant assemblages

associated to two pioneer tree species (Hymenoptera:

Formicidae). Sociobiology, 47: 722-723.

Castro F.S., Gontijo A.B., Castro P.T.A., Ribeiro S.P.,

2012. Annual and seasonal changes in the structure

of litter-dwelling ant assemblages (Hymenoptera:

Formicidae) in Atlantic semideciduous forests.

Psyche, 95971: 1-12.

Chevrolat L.A.A., 1838. Centurie de Buprestides. Revue

Entomologique, 5 (1837): 41-110.

Curletti G. & Migliore L., in press. O genero Agrilus

Curtis, 1829 nas colegoes do Museu de Zoologia da

Universidade de Sao Paulo (Coleoptera Buprestidae).

Papeis Avulsos de Zoologia.

Curtis J., 1825. British Entomology; being illustrations

and descriptions of the genera of insects found in

Great Britain and Ireland: containing coloured figures

from nature of the most rare and beautiful species,

and in many instances of the plants upon which they

are found. London, Printed for the author, Volume 2,

plates 51-98 with text, not paginated.

Espirito Santo N.B., Ribeiro S.P. & Lopes J.F.S., 2012.

Evidence of Competition Between Two Canopy Ant

Species: Is Aggressive Behavior Innate or Shaped by

a Competitive Environment?. Vol. 2012, Article ID

783945, 8 pages doi: 10. 1155/2012/783945

Gilhuis J.P., 1986. Vegetation Survey of the Parque

Florestal Estadual do Rio Doce, MG, Brasil. Master

Tesis, Universidade Federal de Vigosa & Wageningen

Agricultural University, Vigosa, Brasil.

Gory H.L., 1841. Histoire naturelle et iconographie des

insectes Coleopteres. Supplement aux Buprestides. P.

Dumenil, Paris. Volume 4, livraisons 43-52, genera:

Melanophila (pp. 73-77), Buprestis (pp. 107-

111,126), pp. 127-356, Stigmodera, Colobogaster,

Chrysobothris, Belionota , Castalia, Poecilonota,

Zemina, Stenogaster, Agrilus, Amorphosoma, Eu-

merus, Coraebus, Anthaxia, Evagora, Sphenoptera,

Sponsor, Brachys, Trachys, Aphanisticus.

Harold E. Von., 1869. Berichtigungen und Zusatze zum

Catalogus Coleopterorum synonymicus et systemat-

icus. Coleopterologische Hefte, 5: 105-125.

Kerremans C., 1897. Contribution a l'etude de la faune

intertropicale Americaine. Voyages de M. E. Gounelle

au Bresil. Buprestides. Memoires de la Societe ento-

mologique de Belgique, 6: 1-146.

Klug J.C.F., 1825. Entomologiae Brasilianae specimen

alteram, sistens insectoram coleopterorum nondum

450

G. ClIRLETTI, S.P. RiBEIRO & L. MlGLIORE

descriptorum centuriam. Nova Acta Physico-Medica

Academiae Caesareae-Leopoldino-Carolinae. Natu-

rae Curiosoram, 12: 419-476.

Meis M.R.M. & Monteiro A.M.F., 1979. Upper Quater-

nary “rampas”: Doce river valley, Southeastern Bra-

zilian plateau. Zeitschrift f fir Geo-morphologie, 23:

132-151.

Meis M.R.M. & Tundisi J.G., 1986. Geomorphology and

limnological processes as a basis for lake typology:

the middle Rio Doce Valley Lake System. Anais da

Academia Brasileira de Ciencias, 58: 103-120.

Neves F.S., Sperber C.F., Campos R.I., Soares J.P. &

Ribeiro S.P., 2013. Contrasting effects of sampling

scale on insect herbivores distribution in response to

canopy structure. Revista de Biologia Tropical, 61:

125-137.

Obenberger J., 1933a. Hg-n/ws-Studien II. Wiener Ento

mologische Zeitung, 50: 6-24.

Obenberger J., 1933b. Nove druhy rodu Agrilus Steph.

(Col. Bupr.). De Agrili generis speciebus novis (Col.

Bupr.). Acta Entomologica Musaei Nationalis Pragae,

11: 15-81.

Obenberger J., 1935. De novis Agrili generis regionis

Neotropicae speciebus (Col Bupr.). Nove druhy

krascg z rodu Agrilus z Neotropicke oblasti. Acta En-

tomologica Musaei Nationalis Pragae, 13: 108-140.

Obenberger J., 1935. Synonymia Agrilorum. Acta So-

cietatis Entomologicae Cechosloveniae, 32: 43.

Overloop E., 1981. Post-Glacial to Holocene transi-

tion in a peatlayer of lakes jacare (Rio Doce

Basin, Brazil). Bulletin des Societe Beige de Geolo-

gie, 90: 107-119.

Peronico C. & Castro P.T., 2008. Analise faciologica e

interpretagao palinologica de testemunho recuperado

de ambiente lacustre assoreado no Parque Esta-

dual do Rio Doce (MG). Revista Brasileira de

Geociencias, 38: 654-660.

Pflug R., 1969. Quaternary lakes of eastern brazil.

Photogrammetria, 24: 29-35.

Ribeiro S.P., Soares J.P., Campos R.I. & Martins R.P.,

2008. Insect herbivores species associated to pioneer

tree species: contrasting within forest and ecotone

canopy habitats. Re vista Brasileira de Zoociencias,

10: 141-152.

SEMAD/ IEF/ PROMATA. 2007. Plano de Manejo do

Parque Estadual do Itacolomi. Relatorio tecnico. 236

pp.

Souza Fagundes de R., Del Claro, Ribeiro S.P., 2012. Ef-

fects of the Trophobiont Herbivore Calloconophora

pugionata (Hemiptera) on Ant Fauna Associated with

Myrcia obovata (Myrtaceae) in a Montane Tropical

Forest. Psyche (Cambridge, 1874). , v.2012, p. 1—8.

Veloso H.P., Rangel-Filho A.L.R. & Lima J.C., 1991.

Classificagao da Vegetagao Brasileira, Adaptada a um

Sistema Universal. IBGE, Brasil.

Waterhouse C.O., 1887. New genera and species of

Buprestidae. The Transactions of the Entomological

Society of London, 1887: 177-184.

Wemeck F.P., Costa G.C., Colli G.R., Prado D.E. & Sites

J.W., 2011. Revisiting the historical distribution of

seasonally dry tropical forests: new insights based on

palaeodistribution modelling and palynological evi-

dence. Global Ecological Biogeography, 20: 272-

288.

Biodiversity Journal, 2013, 4 (4): 451-466

The genus Lichenophanes Lesne, 1 899 in Italy (Coleoptera

Bostrichidae) and short considerations on the saproxy-

lophagous beetle-fauna of Nebrodi Mountains (Sicily)

Calogero Muscarella 1 , Ignazio Sparacio 2 , Andrea Liberto 3 & Gianluca Nardi 4 *

'Cooperativa Silene, Via D’Ondes Reggio, 8A Scala G - 90127 Palermo, Italy; e-mail: calogero@silenecoop.org

2 Via E. Notarbartolo, 54 int. 13 - 90145 Palermo, Italy; e-mail: isparacio@inwind.it

’Via Camillo Pilotto, 85 F/15 - 00139 Roma, Italy; e-mail: andrea.liberto@ansa.it

4 MiPAAF, Corpo Forestale dello Stato, Centro Nazionale per lo Studio e la Conservazione della Biodiversita Forestale “Bosco

Fontana” di Verona, Sede di Bosco Fontana, Strada Mantova 29 - 46045 Marmirolo (MN), Italy; e-mail: l_nardi@hotmail.com

^Corresponding author

ABSTRACT The Italian distribution and ecology of Lichenophanes numida Lesne, 1899 and L. varius

(Illiger, 1801) are summarized; both species are recorded for the first time from Sicily, and

L. varius also from Veneto and Molise Regions. L. varius is a protected species, at different

levels, in most of European countries in which it occurs, so its main threatening factors are

discussed. Finally, the importance of the dead wood for the conservation of saproxylic beetle-

fauna of Sicilian forests, is underlined.

KEY WORDS distribution; saproxylic fauna; global warming.

Received 21.09.2013; accepted 11.11.2013; printed 30.12.2013

INTRODUCTION

The genus Lichenophanes Lesne, 1899

(Coleoptera Bostrichidae Bostrichinae) occurs

with 41 species in Europe, Asia, Africa and Ameri-

cas, especially in more mesic areas (Ivie, 2002;

Borowski & WQgrzynowicz, 2007).

As stated by Lesne (1899) in the original de-

scription of this genus, its etymology refers to the

mimetic lichen-like appearance of the colouration

of many of its species. Lichenophanes is distin-

guished from other genera of Bostrichinae (cf.

Lesne, 1899, 1901; Ivie, 2002; Bahillo de la Puebla

et al., 2007) by the shape of the apophysis of the

first stemite dilated ventrally, and by the presence,

on each side of the articles of the antennal club, of

a pairs of sensory rounded dimples (Lesne, 1899,

figs. 57-59; De Marzo & Porcelli 1989, fig. 5b) with

well-defined margins and covered by a dense

golden pubescence (Fig. 1).

From a biological point of view, this genus in-

cludes secondary saproxylophagous beetles that, so,

during the larval stages feed on wood yet partially

degraded by other organisms.

In the Palaearctic Regions are present only

three species (Borowski & W^grzynowicz, 2007),

two of them occur in Europe: L. numida Lesne,

1899 and L. varius (Illiger, 1801), both live also in

Italy (Audisio et al., 1995; Nardi, 2004; Borowski,

2007; Borowski & W^grzynowicz, 2007). New

records of these species give an opportunity to re-

vise their distribution in Italy and to summarize

(Table 1; Figs. 2-7) the morphological features for

their identification.

452

C. Muscarella, I. Sparacio, A. Liberto & G. Nardi

Lichenophanes varius

Lichenophanes numida

Colour brown

Colour dark brown

Length 5.5-13 mm

Length 9-14 mm

Clypeus without reddish, dense and long hairs

Clypeus with reddish, dense and long hairs

Pronotum slightly longer than wide (Figs. 2, 3) (Lesne,

1899: figs. 66-67)

Pronotum slightly wider than long (Fig. 4) (Lesne,

1899: fig. 68)

Apex of elytra (visible ventrally) not enlarged (Fig. 6)

with crenulate margin

Apex of elytra (visible ventrally) enlarged (Fig. 7)

with smooth margin

Intercoxae process of the abdomen rounded or truncate

at the apex (Fig. 5)

Intercoxae process of the abdomen pointed at the

apex

Ventral surface of the abdomen with dense and regular

punctures

Ventral surface of the abdomen with sparse punctures

on the midline, denser on the sides

Apical declivity of the elytra generally with numerous

pubescent areas (Fig. 10)

Apical declivity of the elytra generally with only two

pubescent areas in the upper part

Lobes of the parameres of the aedeagus wider than long

with parallel sides to the basal half and the apical half

abruptly truncated and occupied by a sensory and

pubescent area (Bahillo de La Puebla et al., 2007: fig. 9c)

Lobes of the parameres of the aedeagus as long as wide,

regularly decreasing in width from the base to the apex,

with outer margin regularly rounded (Bahilo de La Pue-

bla, 2007: figs. 9a-b)

Table 1. Morphological characters that differentiate the two species: Lichenophanes varius and L. numida (cf. Lesne,

1899, 1901; Porta, 1929; Bahillo de la Puebla et ah, 2007).

MATERIALS AND METHODS

In this paper, all the available Italian data are crit-

ically revised and listed together with new records.

For each record, the following information, when

available, is provided: region, province, commune,

locality, biotope, metres a.s.l., geographic coordi-

nates, date, collector, additional information on the

finding, number of specimen/s (es.), collection, pos-

sible published data source (bibliographic reference

or website, in parenthesis); the symbol “(!)” refers

to examined records. The labels of the examined

specimens are generally written in Italian; hereun-

der, the regions and the collecting methods were

translated in English; the same is valid for literature

records. The mainland Italian regions are listed

from north to south, and from west to east, all to-

ponyms are listed alphabetically. When deemed

useful for the discussion of both species, material

examined from other countries (“Other material

examined”) is also provided. The material was iden-

tified according to Lesne (1899, 1901) and Bahillo

de la Puebla (2007). Comments and interpretations

are given in square brackets.

ACRONYMS. Specimen depositories: CFA =

F. Angelini, Francavilla Fontana (Brindisi), Italy

(F. Angelini, pers. com., 2013); CDS = D. Sechi,

Cagliari, Italy; CLE = P. Leo, Cagliari, Italy; CLI =

A. Liberto, Rome, Italy; CMO = L. Mola, Castel

Mella (Brescia), Italy; CMU = C. Muscarella,

Palermo, Italy; CNA= G. Nardi, Cisterna di Latina

(Latina), Italy; CNBFVR = Centro Nazionale per

lo Studio e la Conservazione della Biodiversita

Forestale “Bosco Fontana” di Verona, Sede di

Bosco Fontana, Mannirolo (Mantua), Italy; CPA

= G. Pace, Rome, Italy (G. Pace, pers. com., 2013);

CSP = I. Sparacio, Palermo, Italy; CVO = V. Vomero,

Rome, Italy (V. Vomero, pers. com., 1995); MCGB

= G. Binaghi c/o Museo Civico di Storia Naturale

“Giacomo Doria”, Genoa, Italy; MCGD = A. Dodero

c/o Museo Civico di Storia Naturale “Giacomo

Doria”, Genoa, Italy (R. Poggi, in litteris, 1996);

The genus Lichenophanes in Italy and considerations on the saproxylophagous beetle-fauna of Nebrodi Mountains (Sicily) 453

Figure 1. Antenna of the genus Lichenophanes. Figures 2-7. Morphological characters of Lichenophanes varius and

L. numida, see Table 1 (cf. Lesne, 1899, 1901; Bahillo de la Puebla et al., 2007).

MCGM = C. Mancini c/o Museo Civico di Storia

Naturale “Giacomo Doria”, Genoa, Italy; MCSV =

Museo Civico di Storia Naturale, Verona, Italy;

MCSVB = M. Burlini c/o Museo Civico di Storia

Naturale, Verona, Italy; MCSVS = A. Sette c/o

Museo Civico di Storia Naturale, Verona, Italy;

MCZRE = C. Emery c/o Museo Civico di Zoologia,

Rome, Italy; MCZRG = G. Ganev c/o Museo Civico

di Zoologia, Rome, Italy; MCZRL = P. Luigioni c/o

Museo Civico di Zoologia, Rome, Italy; MNHN =

Museum National d’Histoire Naturelle, Paris,

France; MSNF = Museo di Storia Naturale dell’

Universita degli Studi di Firenze, Sezione di Zo-

ologia "Fa Specola", Florence, Italy (F. Cianferoni,

pers. com., 2013).

Main collectors: AF = A. Fiberto; AM = A.

Molinu; AS = A. Sette; AV= A. Verdugo; CE = C.

Esposito; CM = C. Muscarella; DB = D. Birtele; DS =

D. Sechi; DW = D. Whitmore; EC = E. Colonnelli;

FA= F. Angelini; FI = F. Izzillo; GA= G. Altadonna;

GM = G. Magnani; GN = G. Nardi; GP = G. Pace;

GS = G. Sanaa; IG = I. Ganev; IS = I. Sparacio; FM =

F. Mola; MBr = M. Bracalini; MBu = [M.] Burlini;

MG = M. Gigli; ME = M. Fopresti; PC = P Cerretti;

PF = P. Feo; RD = R. De Togni.

Other abbreviations and recurrent terms used in

Records: Bosco = Wood; ca = circa = about; coll. =

collection; dint, di = dintomi di = environs of; env.

= environs; es. = specimen/s; ex = emerged from

wood of; FEI = “Forum Entomologi Italiani”

(http://www.entomologiitaliani.net) (accessed 13

August 2013); Fago = Fake; loc. = locality; M. =

Monte = Mount; prov. = province; presso = near;

sdb = same data but; Via = Road; wdc = without

date of collection; wfd = without further data; ! =

material examined.

RESULTS

Revised quotations and new records of the

species in Italy with details are listed below.

454

C. Muscarella, I. Sparacio, A. Liberto & G. Nardi

Lichenophanes numida Leslie, 1899

Lichenophanes numida Lesne, 1899: Lesne, 1901:

92; Audisio et al., 1995: 7; Urbano Granero,

2002: 30; Nardi 2004; Bahillo de la Puebla et

al., 2007: 163; Borowski & Wqgrzynowicz,

2007: 97; Lopez-Colon & Bahillo, 2011: 3;

Baena & Zuzarte, 2013: 33.

Lichenophanes numida Lesne, [18]99: Winkler,

1927: 796.

Lichenophanes numida Lesne: Luigioni, 1929: 640;

Porta, 1929: 415.

Lichenophanes numida Lesne, 1898 [sic!]: Vrydagh,

1960: 10.

Lichenophanes numidica [sic!] Lesne, 1899: An-

gelini, 1998: 20.

Lichenophanes numida (Lesne, 1899) [sic!]: Fer-

nandez-Carillo et al., 2001: 42; Murria Beltran,

2002: 198.

Italian records (Fig. 8). Basilicata: Matera

prov.: Lago di San Giuliano, loc. Ponte Cagnolino,

VIII. 1992, FA, light trap (Angelini, 1998); sdb, 27-

31. VII. 1993, 1 ex (CFA). Sardinia: no exact local-

ity and date (Winkler, 1927; Luigioni, 1929; Porta,

1929; Vrydagh, 1960; Audisio et al., 1995; Feman-

dez-Carillo et al., 2001; Lopez-Colon et al., 2001;

Murria Beltran, 2002; Urbano Granero, 2002;

Nardi, 2004; Bahillo de la Puebla et al., 2007;

Borowski & Wqgrzynowicz, 2007; Lopez-Colon

& Bahillo, 2011; Baena & Zuzarte, 2013); Cagliari

prov.: “Gonnos, en juin (Baudi [leg.] in Coll.

Oberthiir)” (Lesne, 1901); “Gonnos, VI.[18]91, A.

varia ”, 1 es. (MCGD); Gesturi, Giara VIII.2004,

ex Quercus suber, DS, 3 es. (CDS); idem,

VIII. 2005, 1 es. (CDS); Sinnai, M. Sette Fratelli,

VIII. 2005, ex Quercus suber , DS, 3 es. (CDS); Vil-

laputzu, Rio Ollastu, m. 80, 17.VI.2007, collecting

at light, DS, 1 es. (CDS); San Nicolo Gerrei

19.VI.2013, collecting at light, DS, 1 es. (CDS);

Nuoro prov.: Orune, wfd, 1 es. (MCGD); ibidem,

wfd, 2 es. (MCGM) (!); ibidem, wdc, coll. De-

marchi, 2 es. (MCGB) (!); Oristano prov.: Mago-

madas, Nigolosu, VII.1994, PL, 1 es. (CLE) (!);

Sassari prov.: Budduso, loc. Sos Canales,

14.VI.1997, AM, 2 es. (CLI) (!); Sassari, Rio Bun-

nari, 8. VI. 1997, DS, collecting at light, 1 es. (CDS)

(!). Sicily: Palermo prov.: Bosco della Ficuzza,

Torretta Torre, 940 m, Plot Conecofor SIC1, UTM

33 S 357671 4194110, 5.V.2004, under the bark of

a Quercus sp. tree, GN, 3 es. remains (CNBFVR)

(!); [Bosco della] Ficuzza, Torretta Torre, 986 m,

UTM 33 S 359993 4196856, 28.VI.2005, DB, PC,

ML & DW, collecting at light, 1 es. (CNA) (!);

[Monreale], Lago Scanzano, 8.VII.2008, IS, direct

collection, 1 es. (CSP) (!).

Other material examined. Algeria: Tizi Ouzou,

Parc Nat. [= National] d’Akfadou, 800-1000 m, 17-

18.VI.1982, GS, 4 es. (MCSV); Ft. [= Foret]

d’Akfadou, 1000 m, 4-7.VI.1980, GS & GM, 2 es.

(MCSV). Spagna: Cadiz, Los Barrios, Arroyo

Valdeinfierno, P.N. [= National Park] Los Al-

comocales, 155 m, 10.VI.2008 , AV, 4 es. (CMU).

Chorotype. W-Mediterranean: Morocco, Alge-

ria, Tunisia, Portugal, Spain and Italy (Borowski,

2007; Borowski & Wogrzynowicz, 2007). Borowski

(2007) recorded this species also from “AFR” [=

Afrotropical Region], very probably on the basis of

unpublished data, since no other literature record

from this Region (see also Borowski &

Wqgrzynowicz, 2007) is known.

Ecology. The adults of this species, as those of

L. varius, are nocturnal (Espanol, 1956), and are

often collected at light (Lesne, 1905; Espanol,

1956, 1974, as L. numida Lesne, 1898 [sic!]; An-

gelini, 1998; Murria Beltran, 2002; Urbano Gra-

nero, 2002; Llinares & Navarro, 2003; !); they

remain hidden in their tunnels, under bark or in the

cracks of trunks during the day (Espanol, 1956).

The adults are collected chiefly from May to Au-

gust (Bahillo de la Puebla et al., 2007). The main

host plant is Quercus suber L. (cf. Bahillo de la

Puebla et al., 2007), but this beetle develops also

on Q. canariensis Willd. (FEI, 2012), Q.pyrenaica

Willd. (Baena & Zuzarte, 2013), Eucalyptus spp.

(Lesne, 1901, 1905; Pic, 1905) and Populus nigra

L. (Lopez-Colon & Bahillo de la Puebla, 2011). On

High Atlas (Morocco), this species reaches an alti-

tude of 2000 m a.s.l. (Kocher, 1956).

Remarks. L. numida is quite spread in the south-

ern and central parts of the Iberian Peninsula

(Bahillo de la Puebla et al., 2007), but according to

Llinares & Navarro (2003) its presence in this area,

as for other saproxylic beetles, is due to passive in-

troductions with woods from Maghreb.

About the occurrence of this species in Sardinia,

it must be underlined that the old above records from

“Gonnos” refer probably to one of the following to-

ponyms of the Cagliari province: Gonnoscodina,

Gonnosfanadiga, Gonnosno, Gonnostramatza. The

species after the record of Lesne (1901), was cited

The genus Lichenophanes in Italy and considerations on the saproxylophagous beetle-fauna of Nebrodi Mountains (Sicily) 455

generically from Sardinia by several authors (Win-

kler, 1927; Luigioni, 1929; Porta, 1929; Vrydagh,

1960; Audisio et al., 1995; Lopez-Colon, 2000; Fer-

nandez-Carillo et al., 2001; Lopez-Colon et al.,

2001; Murria Beltran, 2002; Urbano Granero, 2002;

Nardi, 2004; Bahillo de la Puebla et al., 2007;

Borowski & Wqgrzynowicz, 2007; Lopez-Colon &

Bahillo, 2011; Baena & Zuzarte, 2013) but the same

Lesne (1938) has omitted (or has overlooked?) this

record. The new records confirm definitively the oc-

currence of this species in Sardinia. While, the

generic record for “S” (= Peninsular Italy) of Audi-

sio et al. (1995) is an error (Liberto & Nardi, unpub-

lished data), but the occurrence of the species in this

area was later reported by Angelini (1998). The

species is here firstly recorded from Sicily. The

above first Sicilian specimens come from an old

mixed oak-forest belonging to the Quercetum gus-

sonnei vegetational association (cf. Mason et al.,

2006), while the latest specimen comes from a dense

planting of Eucalyptus trees situated among stands

of the same above-mentioned oak-forest; so at least

two of the host plants of this beetle occur in these

Figure 8. Distribution in Italy of Lichenophanes numida.

Dot = references; square = new places; R = regional reports;

? = dubious locations.

Sicilian sites (cf. Gianguzzi & La Mantia, 2004).

Various kinds of traps (window flight, Malaise, pit-

fall) were used in this forest (cf. Mason et al., 2006),

but the species was not intercepted by these meth-

ods. The same situation was obseived in various Sar-

dinian sites (cf. Cerretti et al., 2009; Nardi et al.,

2011). In Italy this species is rare and localized; be-

fore this paper, the sole recent record was from

Basilicata Region (Angelini, 1998).

Lichenophanes varius (Illiger, 1801)

Apate varia Illiger: Comolli, 1837: 37.

Apate varia 111.: Villa & Villa, 1844: 62; Bertolini,

1875: 143.

Apate varia Illig.: Bargagli, 1873: 40.

Apate varians [sic!] 111. ( Dufouri Latr. [= dufourii

(Latreille, 1805)]): Costa, 1884: 28.

Lichenophanes varius Illiger, 1801: Lesne, 1898:

470; Lesne, 1901:470.

Lichenophanes varius Illig.: Bertolini, 1904: 76;

Luigioni, 1920: 207; Porta, 1929: 415; Grandi,

1956: 417; Tassi, 1963: 26; De Marzo & Por-

celli, 1989: 88; Gobbi, 2002: 46.

Lichenophanes varius 111.: Luigioni, 1929: 640;

Faggioli, 1955: 174; Frediani, 1961: 3.

Lichenophanes varius : Borchert, 1938: 78; Gobbi,

1984: 54,58.

Lichenophanes varius (Illiger, 1801): Audisio et al.,

1995: 7; Nardi, 1997: 177-178; Gobbi, 2000:

195, 221; Nardi, 2004; Nardi & Zaharadnik,

2004: 127; FEI, 2011, 2012; Pezzi, 2013: 110.

Italian records (Fig. 9). Lombardy: no exact

locality and date (Bertolini, 1875, 1904; Luigioni,

1929; Porta, 1929; Borchert, 1938); “p” [= plain]

(Villa & Villa, 1844); Brescia prov.: Capriano del

Colle, Bosco delle Colombere, 20.VI.20 12, LM,

1 es. (CMO) (FEI, 2012); Como prov.: “campagne

di Rovello Porro” (Comolli, 1837); Mantua

prov.: Marmirolo, Bosco della Fontana, Stand

12, [3 1 . V-] 14. VI. 2000, FM, QrlB3 [= trunk win-

dow trap on an untreated standing Quercus robur

tree], 1 es. (CNBFVR) (!) (Nardi & Zaharadnik,

2004). Veneto: Verona prov.: Illasi, Parco Perez-

Pompei, 18.V-12. VII. 2010, AS, trappola [= trap

= wine trap (cf. Allemand & Aberlenc, 1991)], 1

es. and 1 es. remains (MCSVS) (!); ibidem,

20.V.2011, RD & AS, ex Carpinus betidus , 1 es.

(MCSVS) (!); ibidem, 12.VI.2011, RD & AS, 1

456

C. Muscarella, I. Sparacio, A. Liberto & G. Nardi

es. (MCSVS) (!); ibidem, 21.VI.2011, RD & AS,

ex Carpinus betulus, 1 es. (MCSVS) (!); sdb

2. VII. 2011, 3 es. (MCSVS) (!); sdb 7.VIII.2011, 1

es. (MCSVS) (!); Musella, 5.VIII.1981, AS, ex

Quercus sp., 5 es. (MCSVS) (!); sdb 5.IX.1982, ex

Carpino [( Carpinus betulus )], 1 es. (MCSVS) (!);

Negrar, [Fraz.] Montecchio, 500 m, 11.VII.2011,

RD & AS, ex Carpinus betulus, 1 es. (MCSVS) (!);

[Sommacampagna] (Verona), [Fraz.] Custoza, Val

dei Mulini, 25.11.2010, AS, 2 es. remains (MCSVS)

(!). Emilia-Romagna: Ravenna prov.: Mezzano,

Zona di Protezione Speciale (ZPS) “Bacini ex zuc-

cherificio di Mezzano”, 2006-2013 (Pezzi, 2013).

Tuscany: Grosseto prov.: Grosseto, 20.VII.2011,

MBr, ex Quercus pubescens, 4 es. (FEI, 2011).

Lazio: no exact locality and date (Luigioni, 1929;

Porta, 1929; Borchert, 1938); Frosinone prov.:

[Monti Lepini,] Supino, loc. Pian della Croce, 1100

m, 9.VII.2006, GP, under the bark of an unidentified

tree (maybe a Beech ( Fagus sylvatica), 27 es.

(CPA); Latina prov.: Monti Lepini, Norma, Monte

Arrestino vers. SW, 750 m ca, 25.VII.2002, GN &

CE, about 16 hours, a death specimen carried by a

Camponotus ( Camponotus ) vagus (Scopoli, 1763)

(Hymenoptera, Formicidae) (M. Mei det., 2013) on

a trunk of a fallen Quercus pubescens tree, under

an old tree of the same species covered with Poly-

poraceae, 1 es. (CGN) (!); Parco Nazionale del Circeo

[= Circeo National Park], Sabaudia, 1. VI. 1996, GP,

under the bark of a Quercus sp. tree, 2 es. (CLI) (!);

Rome prov.: [Manziana,] Bosco di Manziana,

12.III.2012, under the bark of a death standing

Quercus sp. tree, MG, 1 es. remains (FEI, 2012);

Roma [= Rome], VI. 1922, Delbue leg., on walls, 1

es. (MCZRL) (!) (Tassi, 1963; Nardi, 1997); dint,

di Roma [= Rome], presso Via Appia Antica,

2.VII.1963, F. Tassi leg., about 20 hours while flut-

tering around the canopy of a large oak tree, 1 es.

(Tassi, 1963); Roma, Nuovo Salario, 1995, intro-

duced with woods (CVO) (Nardi, 1997); Sasso Fur-

bara, 250 m, 2.VI.1996, FI, ex Quercus suber, 1 es.

(CLI) (!); sdb 22.V.1996, 1 es. (CLI) (!); Viterbo

prov.: Bassano [di Sutri = Bassano Romano], VII,

under the bark of a dead Beech [(Fagus sylvatica )],

1 es. (Luigioni, 1920); Bassano [di] Sutri,

18.VII.1909, Luigioni leg., 1 es. (MCZRL) (!); Tus-

cania, 13-24.IV. 1972, ex Quercus suber (Gobbi,

1984: 54); dint. Tuscania, 5.V.1972, ex dry branks

of Quercus suber (Gobbi, 1984: 58). Molise: Cam-

pobasso prov.: Campomarino, Fraz. Campomarino

Lido, 17.VI.1992, AL, on Populus sp., 1 es. (CLI)

(!). Campania: no exact locality and date (Luigioni,

1929; Porta, 1929); Naples prov.: no exact locality

and date (Luigioni, 1929; Porta, 1929); Naples

prov.: “Naples” (MNHN) (Lesne, 1898, 1901);

“Nap” [= Neapel = Naples] (Borchert, 1938);

Napoli [= Neaples], wfd [very probably collected

during the XIX century], 1 es. (MSNF); “Nap.” [=

Napoletano = Naples area] (Bertolini, 1875); “n” [=

ditto] (Bertolini 1904); [Naples,] Capodimonte,

27.VI.1867, 2 es. (MCZRE) (!); sdb 29.VI.1867, 1

es. (MCZRE) (!); dint, di Napoli [= Naples], Capodi-

monte, VI. 19 11, Anguis. [= Anguissola] leg., 1 es.

(MCZRL) (!). Apulia: Foggia prov.: Gargano,

Foresta Umbra (Faggioli, 1956); ibidem, [4. VII-

4.VIII.1955], direct collection on dry woods

(Grandi, 1956); Villaggio Amendola, 21.V.1999, P.

Crovato leg., on Ulmus sp., 1 es. (CLI). Basilicata:

Matera prov.: Policoro, 25.V.1986, 1 female (De

Marzo & Porcelli, 1989); Potenza prov.: Foresta

[demaniale] Gallipoli-Cognato, 500 m, 14.VI.1994,

G. Gobbi leg., taken from a little cell in a death

brank of an Acer monspessulanum tree, 2 es.

(Gobbi, 2002); Foresta demaniale Gallipoli-Cog-

nato, 500 m, 15.VI.1996, AL, under the bark of a

fallen Quercus cerris tree, 1 es. (CLI) (!). Sicily:

Messina prov.: [Monti Nebrodi,] Biviere di M. Sori,

VII. 1938, MBu, 1 es. (MCSVB) (!); [Monti Ne-

brodi,] Monte Soro, Piano Vescovo, 1400 m,

1. VII. 1989, IS, 1 es. (CSP) (!); [Monti Nebrodi,]

Messina, Monte Trefmate, 20.VII.2013, GA & CM,

[on a stump of a Quercus cerris tree], 1 es. (CMU)

(!); Monti Nebrodi, tra [= between] Caronia e [=

and] Capizzi, 1300 m, 37°56 , 01”N/14°30 , 67”E,

6. VII. 2005, AL, on a Quercus cerris tree, 1 es.

(CLI) (!). Sardinia: no exact locality and date

(Bertolini, 1875, 1904; Porta, 1929; Audisio et al.,

1995; Nardi, 2004); no exact locality and date,

“Baudi, Ghiliani [leg.]” (Bargagli, 1873); Nuoro

prov.: “un individuo sotto le cortecce delle Elci

sul Monte Chiesa di Aritzo: luglio” [= a specimen

under the barks of Holms [( Quercus ilex)] on

Mount Chiesa of Aritzo: July] (Costa, 1884).

Other material examined. [Bulgaria:] Svile-

uprad, 10.VII.1981, IG, 1 es. (MCZRG) (!). Czech

Republic: Moravia, Breclav, 2.VI.2007, M. Brabech

leg., ex [unspecified tree], 1 es. (CNA) (!); sdb

2. VII.2008, 1 es. (CNA) (!). France: Corse du sud,

env. Porto Vecchio, Rau Scopa Piana, 100 m,

7. VI. 1999, EC, 2 es. (CLI) (!). Greece: Thessaly

The genus Lichenophanes in Italy and considerations on the saproxylophagous beetle-fauna of Nebrodi Mountains (Sicily) 457

(Trikala), Oros Antihassia, Vlahava env., 600 m,

29.VI.1999, AL, on Quercus sp., 1 es. (CLI) (!).

[Romania:] Banat, Mehadia, wfd, coll. Burlini, 2

es. (MCSV) (!).

Chorot ype . Turano -Europ eo -Mediterranean

(cf. Nardi & Zaharadnik, 2004; Borowski, 2007;

Borowski & Wqgrzynowicz, 2007).

Ecology. The larvae are saproxylophagous and

develop in the wood (branks and rotting trunks) of

many broadleaved tree genera (e.g. Alnus, Carpi-

nus, Castanea, Fagus, Populus, Quercus and 777/a)

(cf. Lesne, 1901; Sahlberg, 1913; Iablokoff, 1945;

Espanol, 1955; Horion, 1961; Damoiseau, 1966;

Koch, 1989, as Lichenophanus [sic!] varius; Za-

haradnik, 1996; Nardi & Zaharadnik, 2004; Ricarte

et al., 2009) and roots of Glycyrrhiza glabra L.

(Frediani, 1961); in Italy this species was found in

Acer monspessulanum L., Carpinus betulus L.,

Quercus sp., Q. pubescens Willd., and Q. suber

(Gobbi, 1984; FEI, 2011; Gobbi, 2002; !); more-

over, adults were collected on the following trees:

Quercus sp., Q. cerris L., Q. ilex L., Q. robur L.,

Fagus sylvatica L., and Populus sp. (Costa, 1884;

Tassi, 1963; Nardi & Zaharadnik, 2004; !).

The females seem to prefer for oviposition the

trunk and smaller branches of large decaying logs,

but with wood still compact, positioned on the

sunny clearings of forests exposed to the East (Klin-

gelhoffer, 1843; Perris, 1850 as Apate dufourii).

L. varius , similarly to other Bostrichidae, is able

also at the adult stage of digging tunnels in the wood.

These galleries have a circular entrance that pene-

trates into the wood horizontally for about 1-2 cm

and then fold down with a straight stretch of about

2-3 cm. After the oviposition, the female reaches the

entrance of the cavity where shortly after dies occlud-

ing the entiy; the small larvae, after hatching, go to-

wards the bottom, filling the gallery of sawdust, to

build a small niche where to end the development

(Klingelhoffer, 1843; Reitter, 1911). The larval stage

lasts for more than five years (Recalde Irazun & San

Martin Moreno, 2012) and during this long period,

the larvae may be preyed upon by birds such as wood-

peckers and owls, other beetles such as Cleridae and,

occasionally, even mammals such as dormice (Glis

glis Linnaeus, 1766) (Flopmann, 2010). The pupal

stage begins in late April and lasts about two weeks.

The flickering occurs in early summer and the speci-

mens remain active until August (Klingelhoffer,

1843; Lesne, 1901; Espanol, 1955).

Figure 9. Distribution in Italy of Lichenophanes varius. Dot

= references; square = new places; asterisk = general reports;

R = regional reports; ? = dubious locations.

The adult is typically crepuscolar and nocturn

and is often attracted by artificial lights (Dajoz,

1961; Flechtner, 2000; Bahillo de la Puebla et al.,

2007); according to Iablokoff (1945) it is particu-

larly active in humid and rainy nights. L. varius dur-

ing the day remains hidden under the bark, in the

cracks of trunks or in its tunnels (Lesne, 1901;

Iablokoff, 1943; Espanol, 1955; JTorion, 1961),

from where it is easy to get it out by blowing out

the entrance with tobacco smoke (Perris, 1850).

According to some authors (cf. Iablokoffl, 1943,

1945; Espanol, 1955), L. varius might attack only

wood invaded by the mycelia of Biscogniauxia num-

mularia (Bull.) Kuntze (1891) (Pyrenomycetes, Xy-

lariaceae). This peculiar ecology, is probably, as

observed for other saproxylophagous beetles (cf.

Rejzek & Vlasak, 2000), one of the causes of the rar-

ity of this species. The above Sicilian records seem

to confirm this requirement since a large decay of

the beechs caused by Biscogniauxia nummularia is

known from one (Monte Soro) of the above men-

tioned collection sites (Torta et al., 2009), while the

beetle from Monte Trefmaite was collected on a

458

C. Muscarella, I. Sparacio, A. Liberto & G. Nardi

stump of an oak ( Quercus sp.) (Fig. 10) tree infested

by the black ascocarpi of another species of the same

genus: B. mediterranea (De Not.) Kuntze (1891) (A.

La Rosa, pers. com., 2013). This fungus lives on

oaks ( Quercus spp.) while B. nummularia lives on

beech (cf. Franceschini et al., 2009; Torta et al.,

2009). B. mediterranea is recorded also from Bosco

della Ficuzza (cf. Torta et al., 2009), where L. nu-

mida was collected (!). In recent years there has been

a resurgence of attacks from these phytopathogenic

fungi in Mediterranean forest stands. This is in re-

sponse to climate change that essentially led to an

increase in average temperatures and an altered pat-

tern of annual rainfalls, which caused water stress

to trees, limiting their tolerance to adverse factors

(Franceschini et al., 2009). This resurgence probably

can favour the populations of Lichenophanes spp.

Remarks. The alleged rarity of L. varius in

much of its range, including Italy (Tassi, 1963;

Horion, 1969; Flechtner, 1999; Gobbi, 2000; Zaha-

radnik & Nardi, 2004; Moulin, 2007; Lakatos &

Molnar, 2009; Recalde Irurzun & San Martin

Moreno, 2012), and its fragmented distribution is

due not only to lack of research, but above all to the

peculiar ecological requirements.

In fact, several authors (Cymorek, 1969; Geiser,

1994, 1998; Flechtner, 1999, 2000; Schillhammer,

2003) consider L. varius as "primary forest species",

a species associated with primary forests that since

the last European glaciation have not undergone en-

vironmental disturbances. Horion (1969) considers

L. varius as a pre-glacial forest relict ("praglaziales

urwaldrelikt"), that as a result of climate change re-

mained confined to relict forest islands ("urwaldin-

seln") thus forming disjunct populations spread over

a fragmented distribution area. To confirm this dis-

tribution model Horion (1969) claims to have iden-

tified the area of discontinuity, located along the

valley of the Rhine River, between the French and

North-German populations of L. varius and other

beetles; the existence of this area confirms the exis-

tence of two glacial forest islands that have worked

to refuge areas.

However, as noted by Nagel (1971), in central

Europe this species is mainly distributed in areas that

during the ice ages were devoid of forest islands, or

in countries (e.g. Romania, Hungary and former Yu-

goslavia), which, although forested, were still cov-

ered by permafrost for most of the year, a condition

that would inhibit the survival of L. varius.

Ponel (2007), however, argues that the fragmented

distribution of L. varius is attributable to the human

impact that from the Neolithic has gradually reduced

the extent of primary forest grind into portions, rele-

gating in these forestry islands the relict saproxy-

lophagous species with high ecological requirements

and determining elsewhere the extinction.

For all these reasons, L. varius is included in the

Red Lists, both regional and national, in several Eu-

ropean countries (Mason et al., 2013), in addition, it

is considered rare in environmental conditions not

compromised (Bulgarini et al., 2004) and its habitat

of choice is considered in strong regression through-

out Europe (Franc, 2004; Fischer et al., 2012).

L. varius is referred to as CR (Critically Endan-

gered) in Poland (Glowacinski, 1992; Pawlowski et

al., 2002) and in Germany (Geiser, 1998); “stark

gefahrdet” (highly endangered) in the lander of

Baden- Wuerttemberg and Brandenburg, “vom

aussterben bedrohf ’ (endangered) in the land of

Bavaria (see Bussler, 2013; Biiche & Moller, 2005),

“Rare” in the Upper Silesia (Kubisz et al., 1998),

“stark gefahrdet” in Austria ( Geiser, 1994), “VU”

(Vulnerable) in Slovakia (Holecova & Franc, 2001).

It is not among the species mentioned by a proposal

for an Italian Red List of invertebrates (Cerfolli et

al., 2002), probably due to lack of available data. It

is also classified as “NT” (Near Threatened) in the

European Red List of Saproxylic Beetles developed

by Nieto & Alexander (2010 ) on behalf of the

IUCN (International Union for Conservation of Na-

ture, 2013). Finally, L. varius was included by

Brustel (2001) in a list of saproxylophagous beetles

that are bioindicators of high-quality mature French

woodlands. It is not surprising, therefore, that this

species in Sicily has been found just on Nebrodi

Mountains, that are the Sicilian mountain range that

includes forest environments of greater naturalness

(cf. Sabella & Sparacio, 2004).

The old Italian regional records were listed also

by most of subsequent authors (see above), never-

theless Luigioni (1929) has not listed the species

from Sardinia; the sole detailed records from this

island (Bargagli, 1873; Costa, 1884) are prior to the

description of L. numida. In this framework, it is

possible that the two species have been confused,

nevertheless L. varius occurs also in the nearby

Corsica Island (Lesne, 1901; Bertolini, 1904; Lui-

gioni, 1929; Porta, 1929; Sainte-Claire Deville,

1937; Borchert, 1938; Vrydagh, 1960; Horion,

The genus Lichenophanes in Italy and considerations on the saproxylophagous beetle-fauna of Nebrodi Mountains (Sicily) 459

Figure 10. Lichenophanes varius from Monte Trefmaite

(Nebrodi Mountains, Sicily). Photo by C. Muscarella.

1961; !), which has also a similar biogeographical

history (cf. De Jong, 1998), so only the study of the

old Sardinian specimens (Bargagli, 1873; Costa,

1884) can confirm their identifications, but unfor-

tunately their re-examination has so far been impos-

sible. The above-listed new records from Veneto,

Molise and Sicily, suggest that L. varius probably

occurs in most of Italian regions. The new records

show an unexpected diffusion of the species in the

Verona province (Veneto), but in this province it

was not intercepted by various kinds of traps (win-

dow flight, Malaise, pitfall) in the relatively undis-

turbed broadleaved woods of Mount Baldo (Spada,

2008; Nardi & Spada, 2008). In this province the

species develops also on Carpinus betulus (!) that

in Italy is a new host plant, while it was already

listed by Damoiseau (1966) and Ricarte et al.

(2009). At Bosco della Fontana (Lombardy, Mantua

prov.), L. varius is very rare; this nature reserve

(236.11 ha) includes one of the last remaining

oak-hornbeam forest ( Querco-Carpinetum boreoi-

talicum ) in the Po plain. This reserve is a forest-habi-

tat island, completely surronded by cultivated fields

(cfr. Mason et al., 2002; Mason, 2004). In this lo-

cality a sole specimen was trapped during 2000

(Nardi & Zaharadnik, 2004), although this forest,

since 1988, was object of monitoring of the ento-

mofauna (Mason et al., 2002; Cerretti et al., 2004a;

D’Amen et al., 2013a; Comacchia & Nardi, unpub-

lished data), also at canopy-level (Cerretti et al.,

2004b; Stireman et al., 2011; Birtele & Hardersen,

2012). In Lombardy, this species was not trapped

(Nardi, unpublished data) also in forest sites situ-

ated along the Ticino Valley Regional Park (cf.

Della Rocca et al., 2013), in spite of being recorded

from the bordering Swiss Ticino Region (Luigioni,

1929; Borchert, 1938; Fontana, 1947). The above

site from Molise Region is in an area that in the past

was covered by large woodland formations, and it

is near two remaining forests. The former forest is

Bosco Fantine (Campomarino, Campobasso prov.)

and is 2-3 km in a beeline; it is an hygrophil wood

survived to a reclamation (“Bonifica Ramitelli”)

made during the first half of the twentieth century

(Taffetani, 2011), the second forest is a few km

south of Bosco Fantine, a little beyod the regional

border Molise-Apulia, in the commune of Marina

di Chieuti (Foggia prov.), and is a well preserved

but unprotecteded small hygrophilic wood. The

record from Villaggio Amendola (Apulia, Foggia

prov.) comes from a row of elms and is veiy surpis-

ing, since the environment in this locality is very

degraded: it is constituted by cultivated fields or by

fallow fields with Ferula sp. (Apiaceae) along the

State Road Garganica near a militar airport, more-

over in the neighbouring area no forest is present.

At the time, there was a row of elms not very old,

all cut during the previous year. On these elms, be-

fore their cutting, were present also some Bupresti-

dae (Coleoptera) typical of this tree: e.g. Anthaxia

( Anthaxia ) senicula (Schrank, 1789), = A. (A.) deau-

rata (Gmelin, 1790, and A. (A.) manca (Linnaeus,

1767) (Liberto, unpublished data). This locality is

in the historic Appenine region of Capitanata (south-

ern Molise and northern Apulia); in all this historic

region there is a grid of these rows of elms with func-

tion of windbreak, nowadays gradually replaced

with Eucalyptus sp. The dispersal flight capacity of

this species is probably low (Liberto & Nardi, un-

published data), so probably the occurence in this

site, may reflect a past habitat situation when the

forest density was probably higher than that mea-

sured actually (cf. Pratesi & Tassi, 1979; Ranius,

2006; Taffetani, 2011).

Finally, it must be underlined that the type speci-

men of this specis is probably lost (Vrydagh, 1962).

SHORT CONSIDERATIONS ON THE SA-

PROXYLOPHAGOUS BEETLE-FAUNA OF

SICILY

The discovery of L. varius in various Italian re-

gions and its ecological value, as documented above,

460

C. Muscarella, I. Sparacio, A. Liberto & G. Nardi

further underline the importance of conservation of

forest environments which often act as refuge-area

for many saproxylophagous species; they, represent

a fauna consortium highly specialized in the pro-

cesses of degradation of wood and creation of eco-

logical niches of vital importance for the development

of numerous other organisms which, according to

some estimates, are up to about 30% of the overall

biodiversity of a forest (Pignatti et al., 2009).

Moreover, in recent years, the “dead wood” has

become a key indicator to assess the well-being of

forest environments and the implementation of their

management (Mason et al., 2003; MCPFE, 2003;

European Environment Agency, 2007); the role of

the “dead wood” in the global ecology of a forest,

in terms of biodiversity, has been highlighted and

shown in numerous works (Wermelinger & Duels,

2002; Mason et al., 2003; Schlaghamersky, 2003;

Tagliapietra, 2003; Speight & Good, 2003; Hahn &

Christensen, 2004; Humphrey et al., 2004; Ranius

& Fahrig, 2006; Travaglini et al., 2007; Bishop et

ah, 2009; Pignatti et al., 2009; La Mantia et al.,

2010; D' Amen et al., 2013b, 2013c) .

In particular, on Nebrodi Mountains (Sicily),

where L. varius was recently collected, different

saproxylophagous beetles that are included in the

Habitat Directive (cf. Trizzino et al., 2013) can be

found as well: Rosalia alpina Linnaeus, 1758, Ce-

rambyx cerdo Linnaeus, 1758 (Cerambycidae) and

Osmoderma cristinae Sparacio, 1994 (Cetoniidae).

Moreover, in this area are present many taxa en-

demic of Sicily, in rarefaction and with an extremely

limited range: Ropalopus siculus (Stierlin, 1864),

Clytus clavicornis Reiche, 1860, Grammoptera

viridipennis Pic, 1893 (Cerambycidae), Gnorimus

decempunctatus Heifer, 1833 (Cetoniidae), and Lu-

canus tetraodon sicilianus Thunberg, 1806 (Lu-

canidae) (Sparacio, unpublished data). The Nebrodi

Mountains host at least 70 species (Sabella & Spara-

cio, 2004; Sparacio, unpublished data) included in

the “European Red List of Saproxylic Beetles”

(Nieto & Alexander, 2010).

The paper of Nieto & Alexander (2010), al-

though worthy of further updates, shows that in

Sicily there are at least 112 of the 436 mentioned

species (Muscarella & Sparacio, unpublished data).

Then it occurs that Sicily (0.24 % of European ter-

ritory) hosts approximately 25% of the saproxy-

lophagous beetles considered threatened or

endangered in Europe.

This faunal community according to the "2nd

National Report on the implementation of the Habi-

tats Directive and the conservation status of habitats

and species in Italy" is particularly threatened (La

Posta et al., 2008). For this reason the EU directives

on the protection of the saproxylic fauna have been

implemented by the Italian State which has enacted

specific laws, both regional and national, and pro-

posed guidelines for the management of dead wood

and fauna associated with it (Campanaro et al.,

2011; Trizzino et al., 2013).

The main factor of disturbance of such an im-

portant wildlife is the short-sighted management of

forests carried out by some administrations that do

not consider the dead wood as fundamental to the

ecological balance of the forest but rather as a threat

to health, a vector of pests and insects "harmful "

or, at best, as a simple source of firewood. As a re-

sult, the indiscriminate falling of dead stumps not

yet fallen, and removing decayed and senescent

trees have a negative impact on the populations of

saproxylophagous beetles with inevitable repercus-

sions on the entire food chain in forest ecosystems

causing biodiversity loss (see Biscaccianti & Lo-

renzetti, 2012).

ACKNOWLEDGMENTS

Thanks are due to Giovanni Altadonna (Messina),

Fernando Angelini (Francavilla Fontana, Brindisi),

Daniele Baiocchi (Rome), Daniele Birtele (Centro

Nazionale per lo Studio e la Conservazione della

Biodiversita Forestale “Bosco Fontana” di Verona,

Marmirolo, Mantua), Pierfilippo Cerretti (Centro

Nazionale per lo Studio e la Conservazione della

Biodiversita Forestale “Bosco Fontana” di Verona,

Verona), Fabio Cianferoni (Museo di Storia Natu-

rale dell’Universita degli Studi di Firenze, Sezione

di Zoologia “La Specola”, Florence, Italy), Enzo

Colonnelli (Rome), Paolo Comacchia (Porto Man-

tovano, Mantua), Paolo Crovato (Naples), Carmine

Esposito (Velletri, Rome), Enzo Gatti (Belluno),

Maurizio Gigli (Rome), Fabio Gorian (Centro

Nazionale per lo Studio e la Conservazione della

Biodiversita Forestale “Bosco Fontana” di Verona,

Verona), Francesco Izzillo (Naples), Alfonso La

Rosa (Palermo), Piero Leo (Cagliari), Agostino

Letardi (ENEA, S. Maria di Galeria, Rome), Mas-

simo Lopresti (Centro Nazionale per lo Studio e la

The genus Lichenophanes in Italy and considerations on the saproxylophagous beetle-fauna ofNebrodi Mountains (Sicily) 461

Conservazione della Biodiversita Forestale “Bosco

Fontana” di Verona, Verona), Franco Mason (Cen-

tro Nazionale per lo Studio e la Conservazione della

Biodiversita Forestale “Bosco Fontana” di Verona),

Maurizio Mei (Museo di Zoologia, Sapienza Uni-

versita di Roma, Rome, Italy), Alessandro Molinu

(Sassari), Giuseppe Pace (Rome), Emanuele Piat-

tella (Museo di Zoologia, Sapienza Universita di

Roma, Rome, Italy), Roberto Poggi (Museo Civico

di Storia Naturale “Giacomo Doria”, Genoa), Maria

Concetta Sala (Palermo), Daniele Sechi (Cagliari),

Laura Spada (Centro Nazionale per lo Studio e la

Conservazione della Biodiversita Forestale “Bosco

Fontana” di Verona, Marmirolo, Mantua), Antonio

Verdugo (Cadiz), Augusto Vigna Taglianti (Museo

di Zoologia, Sapienza Universita di Roma, Rome,

Italy), Vincenzo Vomero (Rome), Daniel Whi-

thmore (Natural History Museum, London), and

Alberto Zilli (Museo Civico di Zoologia, Rome),

for help in the field or in the lab, and/or for enabling

us to study the collections in their care.

REFERENCES

Allemand R. & Aberlenc H.P., 1991. Une methode effi-

cace d’echantillonnage de Pentomofaune des

frondaisons: le piege attractife aerien. Mitteilungen

der Schweizerischen Entomologischen Gesellschaft/

Bulletin de la Societe entomologique Suisse, 64:

293-305.

Angelini F., 1986. Coleottero fauna del Massiccio del

Pollino (Basilicata-Calabria) (Coleoptera). Entomo-

logica, 21: 37-125.

Angelini F., 1998. Coleotterofauna reperita mediante

trappola luminosa in due stazioni umide della Basili-

cata (Italia meridionale). Quaderni della stazione di

ecologia del civico museo di storia naturale di Fer-

rara, 11: 7-37.

Audisio P., Gobbi G., Liberti G. & Nardi G., 1995.

Coleoptera Polyphaga VIII (Bostrichoidea, Cleroidea,

Lymexyloidea). In: Minelli A., Ruffo S. & La Posta

E. (Eds). Checklist delle specie della fauna italiana,

54. Calderni, Bologna, 27 pp.

Baena M. & Zuzarte A.J., 2013. Contribution to the study

of Bostrichidae from Portugal and update of the

catalogue of the Iberian fauna (Coleoptera, Bostrichi-

dae). Zoologica baetica, 24: 25-51.

Bahillo de la Puebla R, Lopez-Colon J.I. & Baena M.,

2007. Los Bostrichidae Latreille, 1802 de la fauna

ibero-balear (Coleoptera). Heteropterus, Re vista de

Entomologia, 7: 147-227.

Bargagli P., 1873. Materiali per la fauna entomologica

dell’isola di Sardegna. Coleotteri. Bullettino della So-

cieta Entomologica Italiana, 5: 34-49.

Bertolini S., 1875. Catalogo sinonimico e topografico dei

Coleotteri d’ Italia. Supplemento del Bullettino della

Societa entomologica italiana, 6 (IV) (1873): 141—

156.

Bertolini S., 1904. Catalogo dei Coleotteri d’ltalia. Ed.

Rivista italiana di Scienze naturali, Siena (1 899—

1904), 144 pp.

Birtele D. & Hardersen S., 2012. Analysis of vertical strat-

ification of Syrphidae (Diptera) in an oak-hornbeam

forest in northern Italy. Ecological Research, 27:

755-763. doi: 10. 1007/sl 1284-012-0948-2

Bishop D., Majka C., Bondrup-Nielsen S. & Peck S.,

2009. Deadwood and saproxylic beetle diversity in

naturally disturbed and managed spruce forests in

Nova Scotia. In: Majka C. & Klimaszewski J. (Eds.).

Biodiversity, Biosystematics, and Ecology of Cana-

dian Coleoptera II. ZooKeys, 22: 309-340.

Biscaccianti A. & Lorenzetti E., 2012. Entomofauna ed

effetti delle utilizzazioni forestali - Giornata di studio

sul monitoraggio vegetazionale, faunistico e di fun-

zionalita ecosistemica e inventariazione forestale.

Centro Ricerche Floristiche dell’Appennino, Aquila.

Available at: http://www.gransassolagapark.it/pag-

ina.php?id=282

Borchert W., 1938. Die Verbreitung der Kafer Deutsch-

lands, Tatsachen und Folgerungen. W. Borchert,

Schonebeck (Elbe), [5] + 137 + 9 + 47 (doublemaps)

pp.

Borges A. & Meriguet B., 2005. Espace naturel sensible:

la carriere de Vigny. Inventaire des coleopteres saprox-

yliques. OPIE - Office pour les insecte et leur envi-

ronnement, 27 pp.

Borowski J. & Wqgrzynowicz P., 2007. World catalogue

of Bostrichidae (Coleoptera). Mantis Publishing,

Olsztyn, 248 pp.

Borowski J., 2007. Family Bostrichidae Latreille,

1802. In: Lobl I. & Smetana A. (Eds). Catalogue of

Palaearctic Coleoptera, Elateroidea, Derodontoidea,

Bostrichoidea, Lymexyloidea, Cleroidea, Cucujoidea,

Volume 4. Apollo Books. Stenstrap, pp. 320-328.

Bruneau de Mire P., 2008. Role des reserves de

Fontainebleau dans le maintien en Ile-de-France

d’une faune saproxylique. Especes indicatrices de la

naturalite d’un massif forestier. Annales scientifiques

de la Reserve de Biosphere Transfrontaliere Vosges

du Nord-Pfalzerwald, 14: 79-89.

Brustel H., 2001. Coleopteres saproxyliques et valeur

biologique des foret frangaises, perspectives pour la

conservation du patrimoine naturel. These de l’lnsti-

tut National Polytechnique de Toulouse, 327 pp.

Biiche B. & Moller G., 2005. Rote Liste und Gesam-

tartenliste der holzbewohnenden Kafer (Coleoptera)

462

C. Muscarella, I. Sparacio, A. Liberto & G. Nardi

von Berlin mit Angaben zu weiteren Arten. In: Der

Landesbeauftragte fair Naturschutz und Landschaft-

spflege / Senatsverwaltung fur Stadtentwicklung

(Hrsg.): Rote Listen der gefahrdeten Pflanzen und

Tiere von Berlin, CD-ROM, 69 pp.

Bulgarini F., Teofili C. & Bologna G., 2004. GLOBAL

200 ERC - Ecoregional Conservation - II processo di

conservazione ecoregionale e la sua applicazione in

Italia. Servizi Editoriali WWF Italia, Roma, 50 pp.

Bussler H. (Ed.), 2013. Rote liste gefahrdeter Heteromera

(Coleoptera: Tenebrionoidea) und Teredilia (Coleoptera:

Bostrichoidea) Bayems. BayLfU/166: 140-145.

Campanaro A., Bardiani M., Spada L., Carnevali L.,

Montalto F., Antonini G., Mason F. & Audisio R

(Eds), 2011. Linee guida per il monitoraggio e la con-

servazione dell’entomofauna saproxilica. Quaderni

Conservazione Habitat, 6: 1-8 pp. + CD-ROM.

Cerfoli F., Petrassi F. & Petretti F. (Eds), 2002. Libro

Rosso degli Animali d’ Italia - Invertebrati. WWF

Italia - ONLUS - Roma, 83 pp.

Cerretti, P, Hardersen S., Mason F., Nardi G., Tisato M.

& Zapparoli M. (Eds), 2004a. Invertebrati di una

foresta della Pianura Padana, Bosco della Fontana,

Secondo contributo. Conservazione Habitat Inverte-

brati, 3: 1-304.

Cerretti, P, Whitmore D., Mason F., & Vigna Taglianti

A., 2004b. Survey on the spatio-temporal distribution

of tachinid flies - using Malaise traps (Diptera,

Tachinidae). In: Cerretti, P, Hardersen S., Mason F.,

Nardi G., Tisato M. & Zapparoli M. (Eds), Inverte-

brati di una foresta della Pianura Padana, Bosco della

Fontana, Secondo contributo. Conservazione Habitat

Invertebrati, 3: 229-256.

Cerretti P, Mason F., Minelli A., Nardi G. & Whitmore

D. (Eds), 2009 Research on the Terrestrial Arthro-

pods of Sardinia. Zootaxa, 2318: 1-602.

Comolli A., 1837. De Coleopteris novis ac rarioribus

minusve cognitis provinciae Novocomi. Fusi et Socii,

Ticini, I- VI + 7-54 pp.

Costa A., 1884. Notizie ed osservazioni sulla Geo-Fauna

Sarda. Memoria Terza. Risultamento di ricerche fatte

in Sardegna nella estate del 1883 (presentata nelF

adunanza del 10 maggio 1884). Atti della reale

Accademia delle Scienze fisiche e matematiche di

Napoli, 1, serie 2a, fasc. 9 (1884): 1-64.

Cymorek S., 1969. 67. Familie: Bostrychidae (Bohrka-

fer). In: Freude H., Harde K.W. & Lohse G.A. (Eds),

Die Kafer Mitteleuropas Band 8. Goecke & Evers,

Krefeld, pp. 13-25.

Dajoz R., 1961. Notes sur la faune entomologique de la

foret de la Massane (P.O.). V. Premiers resultats

obtenus a l’aide d’un piege lumineux a ultra- violets.

Bulletin mensuel de la Societe linneenne de Lyon, 30:

147-149.

D’Amen M., Birtele D., Zapponi L. & Hardersen S.,

2013a. Patterns in diurnal co-occurrence in an assem-

blage of hoverflies (Diptera: Syrphidae). European

Journal of Entomology, 110 (4): 649-656.

D’Amen M., Bombi P., Campanaro A., Zapponi L.,

Bologna M. & Mason F., 2013b. Protected areas and

insect conservation: questioning the effectiveness of

Natura 2000 network for saproxylic beetles in Italy.

Animal Conservation, 16: 370-378.

D’Amen M., Bombi P., Campanaro A., Zapponi L.,

Bologna M. & Mason F., 2013c. Possible directions

in the protection of the neglected invertebrate biodi-

versity. Animal Conservation, 16: 383-385.

Damoiseau R., 1966. Les Bostrychidae du Musee de

Moravie a Brno (Coleoptera - Cucujoidea). Casopis

Moravskeho Musea (Acta Musei Moraviae), 51:

291-300.

De Jong H., 1998. In search of historical biogeographic

patterns in the western Mediterranean terrestrial

fauna. Biological Journal of the Linnean Society, 65:

99-164.

Della Rocca F., Stefanelli S., Campanaro A. & Bogliani

G., 2013. Dead wood threshold for saproxylic beetles

in Po Plain woodlands, Italy. 7th Symposium and

Workshop on Conservation of Saproxylic Beetles,

12-14 May 2012, Granada - Spain, p. 31.

De Marzo L. & F. Porcelli, 1989. Struttura delle “aree sen-

soriali” antennali in alcuni Bostrichidi (Coleoptera).

Entomologica, 22 (1987): 87-95.

Espanol F., 1955. Los bostriquidos de Cataluna y Baleares

(Col. Cucujoidea). Publicaciones del Institute de Bio-

logia Aplicada, 21:1 07- 135.

Espanol F., 1956. Lictidos y bostriquidos de la zona

mediterranea de Marruecos. Publicaciones del Insti-

tute de Biologia Aplicada, 24: 73-75.

Espanol F., 1974. Nuevos dates sobre los Bostrychidae

de la fauna espanola (Col. Cucujoidea). Graellsia, 28:

37-44.

European Environment Agency, 2007. Halting the loss

of biodiversity by 2010: proposal for a first set of

indicators to monitor progress in Europe. EEA

Technical report, 11, 182 pp.

Faggioli D., 1956. Campagna di ricerche dell’Istituto di

Entomologia dell’Universita di Bologna nella

“Foresta Umbra” (Gargano). II. Elenco delle specie

raccolte. (Primo lotto). Bollettino dell’Istituto di En-

tomologia dell’Universita degli Studi di Bologna, 21 :

167-177.

Fernandez-Carillo E., Fernandez-Carillo J.L. & Lopez

Colon I.J., 2001. Los bostrichidae de la provincia de

Ciudad Real (Coleoptera). Boletin de la Sociedad En-

tomologica Aragonesa, 29: 42.

Fischer R., Waldner R, Camicer J., Coll M., Dobbertin

M., Ferretti M., Hansen K., Kindermann G., Lasch-

Born P., Lorenz M., Marchetto A., Meining S., Niem-

inen T., Penuelas J., Rautio P., Reyer C., Roskams P.

& Sanchez G., 2012. The Condition of Forests in

The genus Lichenophanes in Italy and considerations on the saproxylophagous beetle-fauna of Nebrodi Mountains (Sicily) 463

Europe. 2012 Executive Report. ICP Forests, Ham-

burg, 24 pp. Available at http://www.icp-

forests.org/RepEx.htm [accessed 24 December

2013],

Flechtner G., 1999. “Urwaldrelikt” nach uber 100 Jahren

in Stadtwald wiederentdeckt (Coleoptera: Bostrychi-

dae). Nachrichten des Entomologischen Vereins

Apollo, N.F. 20(1): 25-26.

Flechtner G., 2000. Weitere “Urwaldrelikte” im Frank-

furter Stadtwald wiederentdeckt (Coleoptera). Nach-

richten des Entomologischen Vereins Apollo, N.F. 20

(3/4): 321-326.

Flo(3mann S., 2010. Lichenophanes varius (Illiger,

1801), ein interessanter Bohrkafer fur Thiiringen

(Coleoptera: Bostrychidae). Thuringer Faunistische

Abhandlungen, 15: 163-165.

Fontana R, 1947. Contribuzione alia fauna coleotterolog-

ica Ticinese. Seconda edizione ampliata e aggiornata.

Bollettino della Societa Ticinese di Scienze Naturali,

42: 16-94.

Franc V., 2004. Beetles (Coleoptera) of the Strazovske

vrchy Mts with special reference to bioindicatively

significant species. In: Franc V. (Ed.), Strazovske

vrchy Mts - research and conservation of nature. Pro-

ceedings of the conference, Belusske Slatiny, October

1 &2,2004,pp. 103-115.

Franceschini A., Fongo S. & Moricca S., 2009. Avversita

biotiche e mutamenti climatici in ambienti forestali. In:

Ciancio O. (ed.), Atti del Terzo Congresso Nazionale

di Selvicoltura per il miglioramento e la conser-

vazione dei boschi italiani. 16-19 ottobre 2008,

Taormina (Messina), volume secondo. Accademia

italiana di Scienze Forestali, Firenze, pp. 605-610.

Frediani D., 1961. Ricerche morfo-biologiche sul Sinoxy-

lon perforans Schrk. (Coleoptera Bostrychidae).

Bollettino del Faboratorio di Entomologia agraria

“Filippo Silvestri”, 19: 1-52.

Geiser R.H., 1994. Bostrychidae, Bohrkafer: 145-146.

In: Jach M.A. (red.), Rote Fiste der gefahrdeter Kafer

Osterreichs (Coleoptera). In: Gepp J. (Ed.), Rote Fis-

ten gefahrdeter Tiere Osterreichs, Band 2. Bun-

desministerium fiir Umwelt, Junged und Familie,

Graz, pp. 107-200.

Geiser R., 1998. Rote Fiste der Kafer (Coleoptera). In:

Bundesamt fiir Naturschutz (Hrsg.): Rote Fisten der

gefahrdeten Tiere in Deutschland. Schr.- Schriften-

reihe fiir Fandschaftspflege und Naturschutz, 55:

168-230.

Gianguzzi F. & Fa Mantia A., 2004. Fe serie di vege-

tazione della Riserva “Bosco Ficuzza, Rocca Busam-

bra, Bosco del Cappelliere e Gorgo del Drago”

(provincia di Palermo) con allegata Carta della Veg-

etazione (scala 1:20.000). II Naturalista siciliano, 28:

265-326.

Gobbi G., 1984. Appunti su Coleotteri Cleroidei preda-

tori di xilofagi (Coleoptera: Trogositidae, Cleridae,

Melyridae). Bollettino delTAssociazione Romana di

Entomologica, 38 (1983): 51-62.

Gobbi G., 2000. Gli artropodi terrestri e la tutela degli

ecosistemi in Italia. II Naturalista siciliano, 24: 1 89—

223.

Gobbi G., 2002. Nuovi reperti di Coleotteri nel biotopo

di Policoro (Matera) ed in altre aree protette della

Fucania (Coleoptera). Annali del Museo Civico di

Storia Naturale di Ferrara, 5: 41-50.

Grandi G., 1956. Campagna di ricerche delTIstituto di

Entomologia delTUniversita di Bologna alia “Foresta

Umbra” (Gargano). I: Ricapitolazione dei risultati

ottenuti. Annali. Accademia Italiana di Scienze Fore-

stali, 4: 405-418.

Hahn K. & Christensen M., 2004. Dead wood in Euro-

pean forest reserves - a reference for forest manage-

ment. In: Marchetti M. (ed.), “Monitoring and

indicators of forest biodiversity in Europe - from

ideas to operationally” . Firenze (Italy) 12-15 No-

vember 2003. EFI Proceedings, 51: 181-191.

Holecova M. & Franc V., 2001. Cerveny (ekosozolog-

icky) zoznam chrobakov (Coleoptera) Slovenska.

Ochrana prirody (SOP SR Banska Bystrica). In:

Balaz D., Marhold K. & Urban P. (Eds.), Cerveny zoz-

nam rastlin a zivoci-chov Slovenska. Ochrana prirody

(SOP SR Banska Bystrica), Suppl. 20: 111-128.

Horion A., 1961. Faunistik der Mitteleuropaischen Kafer.

Band VIII: Clavicornia. 2. Teil (Thorictidae bis

Cisidae), Teredilia, Coccinellidae. Uberlingen - Bo-

densee, Kommissionsvergal Buchdriickerei Aug.

Feyel, I-XV + 375 pp.

Horion A., 1 969. Neunter Nachtrag zum Verzeichnis der

mitteleuropaischen Kafer. Entomologische Blatter,

65: 1-47.

Humphrey J.W., Sippola A.F., Femperiere G., Dodelin

B., Alexander K.N.A. & Butler J.E., 2004. Deadwood

as an indicator of biodiversity in European forests:

from theory to operational guidance. In: Marchetti M.

(Ed.), “Monitoring and indicators of forest biodiver-

sity in Europe - from ideas to operationally” Firenze

(Italy) 12-15 November 2003. EFI Proceedings, 51,

pp. 193-206.

Iablokoff A.K., 1943. Ethologie de quelques Elaterides

du Massif de Fontainebleau. Memories Museum Na-

tional d’Histoire Naturelle, n.s., 18: 81-160 + 9 pis.

Iablokoff A.K., 1945. Notes sur le Lichenophanes varius

111. F’Entomologiste, 1: 70-72.

IUCN, 2012. Red Fist Categories and Criteria: Version

3.1. Second edition. Gland, Switzerland and Cam-

bridge, UK: IUCN. IV+ 32 pp.

IUCN, 2013. The IUCN Red Fist of Treatened Species,

http ://www.iucnredlist.org/ details/ 1577 60/ 1

464

C. Muscarella, I. Sparacio, A. Liberto & G. Nardi

Ivie M.A., 2002. Bostrichidae Latreille 1802. In: Arnett

R.H.Jr (t), Thomas M.C., Skelley P.E. & Frank J.H.

(Eds), American Beetles. Volume 2. Polyphaga:

Scarabaeoidea through Curculionoidea. CRC Press,

Boca Raton, London, New York, Washington: 233-244.

Klingelhoffer H.O., 1843. Mittheilungen aus dem tage-

buche des Hrn. Klingelhoffer Oberlieut in Darmstadt.

Stettiner Entomologische Zeitung, 4: 86-87.

Koch K., 1989. Die Kafer Mitteleuropas. Okologie Band

2. Goecke & Evers, Krefeld, 382 pp.

Kocher L., 1956. Catalogue commente des Coleopteres

du Maroc, fascicule IV. Clavicomia et groupes voisins.

Travaux de Tlnstitut Scientifique Cherifien. Serie

zoologie, 11: 1-136.

Kubisz D., Kuska A., Pawlowski J., 1998. Czerwona

Lista Chrzqszczy (Coleoptera) Gornego Slqska [=

Red List of Upper Silesian Beetles (Coleoptera)].

Centrum Dziedzictwa Przyrody Gornego Slqska,

Raporty Opinie, 3: 8-68.

La Mantia T., Bellavista M., Giardina G. & Sparacio I.,

2010. Longhorn beetles of the Ficuzza woods

( W Sicily, Italy) and their relationship with plant diver-

sity (Coleoptera Cerambycidae). Biodiversity Jour-

nal, 1: 15-44.

La Posta A., Dupre E. & Bianchi E. (Eds), 2008. At-

tuazione della Direttiva Habitat e stato di conser-

vazione di habitat e specie in Italia [2° rapporto

nazionale]. Ministero dell’Ambiente e della Tutela

del Territorio. Roma, 52 pp.

Lakatos F. & Molnar M., 2009. Mass Mortality of Beech

( Fagus sylvatica L.) in South-West Hungary. Acta

Silvatica & Ligaria Hungarica, 5: 75-82.

Lesne P., 1899. Revision des Coleopteres de la famille

des Bostrychides. 3e Memoire Bostrychinae. Annales

de la Societe entomologique de France, 67 (1898):

438-621.

Lesne P., 1901. Synopsis des Bostrychides paleartiques.

L’Abeille, 30: 73-136.

Lesne P., 1905. Notes additionnelles et rectificatives sur

les Bostrychides paleartiques. L’Abeille, 30: 249-

254.

Lesne P., 1938. Bostrychidae. In: Junk W. (Ed.), Coleoptero-

rum Catalogus vol. 10 pars 161. S. Schenkling, Ber-

lin, 84 pp.

Llinares A. & Navarro J., 2003. Primer registro para Cor-

doba (Andalucia, Espana) de Lichenophanes numida

Lesne, 1899 (Coleoptera, Bostrichidae). Bolletin de

la Sociedad Andaluza de Entomologia, 8: 7-9.

Lopez-Colon J.I.,. 2000. Los Bostrichidae Latreille, 1802

de la fauna de Marruecos (Coleoptera). Biocosme

Mesogeen, 16(1999): 171-221.

Lopez-Colon J.I., Melic Bias A., Gonzalez Pena C.F.,

Beltran Valen J.R. & Blaso-Zumeta J., 2001. Familia:

Bostrichidae Latreille, 1802 (Insecta: Coleoptera).

Catalogus de la entomofauna aragonesa (Sociedad

entomologica Aragonesa), 25: 15-28.

Lopez-Colon J.I. & Bahillo de la Puebla, 201 1. El chopo,

nuevo arbol-huesped para Lichenophanes numida

Lesne, 1899 (Coleoptera, Bostrichoidea, Bostrichi-

dae). Boletin de la Asociacion Entomologica As-

turiana, 3: 6-7.

Luigioni P., 1920. Contributo alio studio della fauna

coleotterologica del Lazio. Atti delTAccademia Pon-

tificia de’ Nuovi Lincei, 73: 186-214.

Luigioni P., 1929. I Coleotteri d’ltalia. Catalogo sinon-

imico-topografico-bibliografico. Memorie della Pon-

tificia Accademia delle Scienze, I Nuovi Lincei (2)

13: [4 +] 1-1159 [+ 1],

Mason F., 2004. Dinamica di una foresta della Pianura

Padana. Bosco della Fontana. Seconda edizione con

linee di gestione forestale. Rapporti scientifici 1, Arte

Grafiche Chiribella, Bozzolo, Mantova, 224 pp.

Mason F., Cerretti P., Tagliapietra A., Speight M.C.D. &

Zapparoli M. (Eds), 2002. Invertebrati di una foresta

della Pianura Padana, Bosco della Fontana, Primo

contributo. Conservazione Habitat Invertebrati 1.

Gianluigi Arcari Editore, Mantova, 176 pp.

Mason F., Nardi G. & Tisato M. (Eds), 2003. Proceed-

ings of the International Symposium “Dead wood: a

key to biodiversity”, Mantova, May 29th-31st 2003.

Sherwood 95, Suppl. 2, 100 pp.

Mason F., Cerretti P., Nardi G., Whitmore D., Birtele D.,

Hardersen S. & Gatti E., 2006. Aspects of biological

diversity in the CONECOFOR plots. IV. The Inver-

tebrate Biodiv pilot project. In: Ferretti M., Petric-

cione B., Bussotti F. & Fabbio G. (Eds), Aspects of

biodiversity in selected forest ecosystems in Italy:

status and changes over the period 1996-2003. Third

report of the Task Force on Integrated and Combined

(I&C) evaluation of the CONECOFOR programme.

Annali dell’Istituto Sperimentale per la Selvicoltura,

30, Supplemento 2: 51-70.

Mason F., Campanaro A., Horak J., Istrate P., Munteanu

N., Buche B., Tezcan S., Mendez M., Dodelin B.,

Alexander K. & Schlaghamersky, J. 2010. Licheno-

phanes varius. In: IUCN 2013. IUCN Red List of

Threatened Species. Version 2013.2. Available at:

http://www.iucnredlist.Org/details/157760/l.

MCPFE, 2003. State of Europe’s Forests 2003. The

MCPFE Reports on Sustainable Forest Management

in Europe. Vienna, Austria 2003, 1 15 pp.

Moulin N., Jolivet S., Meriguet B. & Zagatti P., 2007.

Methodologie de suivis scientifiques des especes

patrimoniales (faune) sur le territoire du Parc naturel

regional du Vexin frangais - Entomofaune. OPIE -

PNR Vexin frangais, 66 pp.

Murria Beltran F., 2002. Primer registro de Lichenopha-

nes numida (Lesne, 1 899) para Aragon (Coleoptera ,

Bostrichidae). Boletin de la Sociedad entomologica

Aragonesa (S.E.A.), 30: 198.

The genus Lichenophanes in Italy and considerations on the saproxylophagous beetle-fauna of Nebrodi Mountains (Sicily) 465

Nagel P., 1971. Lichenophanes varius Illiger 1801 (Co-

leoptera, Bostrychidae) - Bemerkungen zu seiner

Chorologie und kausalen Zoogeographie. Faunis-

tisch-floristische Notizen aus dem Saarland, 4: 23-

27.

Nardi G., 1997. Coleoptera Dermestidae, Lyctidae,

Bostrichidae, Ptinidae. In: Zapparoli M. (Ed.), Gli In-

setti di Roma.Comune di Roma, Dipartimento X

Area Risorsa Suolo e Tutela Ambientale, Quaderni

dell’Ambiente, 6: 177-179.

Nardi G., 2004. Fauna Europaea: Bostrichidae. In:

Audisio P. (Ed.), Fauna Europaea: Coleoptera 2, Bee-

tles. Fauna Europaea version 1.0, available at

http://www.faunaeur.org [accessed 16 December

2013 as version 2.6.2 of 29 August 2013].

Nardi G. & Spada L., 2008. Biodiversita degli Artropodi

nella Riserva Naturale Integrale “Lastoni - Selva

Pezzi”, Monte Baldo (Verona). Report (non pubblicato)

per Regione Veneto - Veneto Agricoltura. MiPAAF,

Corpo Forestale dello Stato, Centro Nazionale per lo

Studio e la Conservazione della Biodiversita Forestale

“Bosco Fontana” di Verona, Verona, 59 pp.

Nardi G., Whitmore D., Bardiani M., Birtele D., Mason

F., Spada L. & Cerretti P. (Eds), 2011. Biodiversity

of Marganai and Montimannu (Sardinia). Research

in the framework of the ICP Forests network. Con-

servazione Habitat Invertebrati, 5. Cierre Edizioni,

Sommacampagna, Verona, 896 pp. + 1 map.

Nardi G. & Zahradnik P., 2004. Bostrichidae and Anobi-

idae (Coleoptera). In: Cerretti P., Hardersen S.,

Mason F., Nardi G., Tisato G. & Zapparoli M. (Eds),

Invertebrati di una foresta della Pianura Padana,

Bosco della Fontana, Secondo contributo. Conserva-

zione Habitat Invertebrati, 3: 125-139.

Nieto A. & Alexander K.N.A., 2010. European Red List

of Saproxylic Beetles. Publications Office of the Eu-

ropean Union, Luxembourg, I— VIII + 45 pp.

Pawlowski J., Kubisz D. & Mazur M., 2002: Coleoptera-

chrzqszcze. In: Glowacinski Z. (Eds), Red List of

Threatened Animals in Poland. Polish Academy of

Sciences, Institute of Nature Conservation, Cracow,

150 pp.

Perris E., 1850. Moeurs et metamorphoses de YApate

capicina, Fabr., de YApate sexdentata, Oliv., de

YApate sinuata. Fab. et de YApate dufourii, Latr.

Annales de la Societe entomologique de France, Series

2, 8: 563-565.

Pezzi G., 2013. L’entomofauna della Zona di Protezione

Speciale (ZPS) “Bacini ex zuccherificio di Mezzano”

(Ravenna). 5° contributo. Coleotteri Adefagi (escl.

Carabidae s.l.) e Polifagi (escl. Scarabaeiformia) (In-

secta Coleoptera). Quademo di Studi e Notizie di Sto-

ria Naturale della Romagna, 37: 105-122.

Pic N., 1905. Notes sur divers Coleopteres. Bulletin de

la Societe entomologique de France, 1905: 181-182.

Pawlowski J., Kubisz D. & Mazur M., 2002. Coleoptera

Chrzqszcze. In: Glowacinski Z. (Ed.), Czerwona

Lista Zwierzqt Ginqcych i Zagrozonych w Polsce.

IOP PAN, Krakow, pp. 88-1 10.

Pignatti G., De Natale F., Gasparini P. & Paletto A., 2009.

II legno morto nei boschi italiani secondo Flnventario

Forestale Nazionale. Forest@, 6: 365-375 [online:

2009-11-23] - doi: 10.3832/efor0598-006

Ponel R, 2007. Beetle Records - Postglacial Europe.

Encyclopedia of Quaternary Science, pp. 267-275.

Porta A., 1929. Fauna Coleopterorum Italica. III. Diver-

sicornia. Stabilimento Tipografico Piacentino, Pia-

cenza, 465 pp.

Pratesi F. & Tassi F., 1979. Guida alia natura della Puglia,

Basilicata e Calabria. A. Mondadori, Milano, 253 pp.

Ranius T., 2006. Measuring the dispersal of saproxylic

insects: a key characteristic for their conservation.

Population Ecology, 48 (3): 177-188. doi:

10.1 007/s 1 0 1 44-006-0262-3

Ranius T. & Fahrig L., 2006. Targets for maintenance of

dead wood for biodiversity conservation based on ex-

tinction thresholds. Scandinavian Journal of Forest

Research, 21: 201-208.

Recalde Iruzun J.I. & San Martin Moreno A.F., 2012.

Registro de Lichenophanes varius (Illiger, 1801) en

Navarra (Norte de Espana) y otros Bostrichoidea de

la misma masa forestal (Coleoptera: Bostrichidae y

Anobiidae). Arquivos Entomoloxicos, 6: 63-64.

Reitter E., 1911. Fauna germanica: die Kafer des

deutschen Reiches - III. Band: nach der analytischen

Methode bearbeitet. K.G. Lutz, Stuttgart: 436 pp. +

48 Tafn.

Rejzek M. & Vlasak J., 2000. Larval nutrition and female

oviposition preferences of Necydalis ulmi Chevrolat,

1838 (Coleoptera : Cerambycidae). Biocosme Me-

sogeen, 16 (1999): 55-66.

Ricarte A., Jover T., Marcos-Garcia M.A., Mico E. &

Brustel H., 2009. Saproxylic Beetles (Coleoptera)

and hoverflies (Diptera: Syrphidae) from a Mediter-

ranean forest: towards a better understanding of their

biology for species conservation. Journal of Natural

History, 43 (9-12): 583-607.

Sabella G. & Sparacio I., 2004. II ruolo dei parchi si-

ciliani nella conservazione di taxa di insetti di parti-

colare interesse naturalistico (Insecta Coleoptera et

Lepidoptera Rhopalocera). II Naturalista siciliano,

28: 477-508.

Sahlberg J., 1913. Coleoptera mediterranea orientalia,

quae in Aegypto, Palaestina, Syria, Caramania atque

in Anatolia occidentali anno 1904 collegerunt John

Sahlberg et Unio Saalas. Finska Vetenskaps Soc.,

Forhandlingar, 55 (1912-1913), Afd. A, N.19: 1-282.

Sainte-Claire Deville J., 1937 (f). Catalogue raisonne des

Coleopteres de France. (Complete et publie par A.

Mequignon). L’Abeille, 36: 265-372.

466

C. Muscarella, I. Sparacio, A. Liberto & G. Nardi

Schillhammer H., 1998. Bemerkenswerte Kaferfunde aus

Osterreich (VII) (Coleoptera). Koleopterologische

Rundschau, 68: 265-267.

Schlaghamersky J., 2003. Saproxylic invertebrates of

floodplains, a particularly endangered component of

biodiversity. In: Mason F., Nardi G. & Tisato M.

(Eds), Proceedings of the International Symposium

“Dead wood: a key to biodiversity”, Mantova, May

29th-3 1st 2003. Sherwood 95, Suppl. 2: 15-18.

Spada L., 2008. Artropodi geofili della Riserva Naturale

Integrale “Lastoni - Selva Pezzi” Monte Baldo. Tesi

di Laurea (non pubblicata) in Scienze Naturali (rela-

tore Prof. A. Minelli), Anno Accademico 2006-2007.

Universita degli Studi di Padova, 150 pp.

Speight M.C.D. & Good J.A., 2003. Development of

eco-friendly forestry practices in Europe and the

maintenance of saproxylic biodiversity. In: Mason F.,

Nardi G. & Tisato M. (Eds), Proceedings of the In-

ternational Symposium “Dead wood: a key to biodi-

versity”, Mantova, May 29th-31st 2003. Sherwood

95, Suppl. 2: 73-78.

Speight M.C.D. , 1989. Les invertebres saproxyliques et

leur protection. Collection Sauvegarde de la Nature

n°42. Conseil de l’Europe, Strasbourg, 77 pp.

Stireman J.O. Ill, Cerretti P., Whitmore D., Hardersen S.

& Gianelle D., 2011. Composition and stratification

of a tachinid (Diptera: Tachinidae) parasitoid com-

munity in a European temperate plain forest. Insect

Conservation and Diversity 5: 346-357. doi:

10.1 111/j. 1752^1598. 20 11.00168.x

Taffetani F. (Ed.), 2011. II Bosco Fantine: un’areaumida

retrodunale di elevato valore naturalistico ed ambien-

tale nel Comune di Campomarino (CB). I quaderni

della Selva, 4: 1-131.

Tagliapietra A., 2003. Importanza biologica del legno

morto. In: Cavalli E. & Mason F. (Eds), Tecniche di

ripristino del legno morto per la conservazione della

faune saproxiliche II progetto LIFE Natura

NAT/IT/99/6245 di “Bosco della Fontana” (Mantova,

Italia). Rapporti Scientifici, 2. Gianluigi Arcari

Editore, Mantova: 23-29.

Tassi F., 1963. Reperti 2. - Lichenophanes varius Illig.

(Col. Bostrychidae). Bollettino dell’Associazione

Romana di Entomologia, 18: 26.

Torta L., Burruano S., Sidoti A. & Granata G., 2009. Lat-

ifoglie in Sicilia: un laboratorio di casi fitopatologici.

In: Ciancio O. (Ed.), Atti del Terzo Congresso Nazio-

nale di Selvicoltura per il miglioramento e la conser-

vazione dei boschi italiani. 16-19 ottobre 2008,

Taormina (Messina), volume secondo. Accademia

italiana di Scienze forestali, Firenze, pp. 691-696.

Travaglini D., Barbati A., Chirici G., Lombardi F., Marchetti

M. & Corona P., 2007. ForestBIOTA data on dead-

wood monitoring in Europe. Plant Biosystems, 141:

222-230. doi: 10.1080/11263500701401778

Trizzino M., Audisio P., Bisi F., Bottacci A., Campanaro

A., Carpaneto G.M., Chiari S., Hardersen S., Mason

F., Nardi G., Preatoni D.G., Vigna Taglianti A., Zauli

A., Zilli A. & Cerretti P. (Eds), 2013. Gli artropodi

italiani in Direttiva Habitat: biologia, ecologia, ri-

conoscimento e monitoraggio. Quaderni Conserva-

zione Habitat, 7. CFS-CNBFVR, Centro Nazionale

Biodiversita Forestale. Cierre Grafica, Sommacam-

pagna, Verona, 256 pp.

Urbano Granero J.M., 2002. Primera cita de Licheno-

phanes numida Lesne, 1899 (Coleoptera: Bostrichi-

dae) para Huelva (Andalucia). Boletin de la SAE, 4:

30-32.

Villa A. & Villa G.B., 1844. Catalogo dei Coleopteri della

Lombardia. In: Cattaneo C. (Ed.) Notizie naturali e

civili sulla Lombardia, vol. 1 . Coi tipi di Giuseppe

Berdardoni di Giovanni, Milano, 77 pp. [reprint]. [In

reality, this title is not present in the book; this reprint

assemble two paragraphs of the chapter VIII of the

book: “V. Insetti della Lombardia”, pp. 406-415 and

“Catalogo degli Insetti Coleopteri della Lombardia”,

pp. 416-477].

Vrydagh J.M., 1960. Contribution a F etude des Bostrychi-

dae. 23. Collection de la Section Zoologique du

Musee National Hongrois a Budapest. Bulletin Insti-

tut royal des Sciences naturelles de Belgique, 36: 1-

32.

Vrydagh J.M., 1962 (f). Contribution a F etude des

Bostrychidae (Coleoptera). 31. Troisime collection

du Musee Zoologique de FUniversite Humboldt a

Berlin. Bulletin Institut royal des Sciences naturelles

de Belgique, 38: 1-47.

Wermelinger B. & Duelli R, 2002. Die Insekten im Okosys-

tem Wald. Bedeutung, Anspriiche, Schutz. Werden-

berger Jahrbuch 2003 (16. Jg). Buchs, Buchs Medien:

104-112.

Winkler A., 1927. Tededilia. In: Winkler A. (Ed.), 1924-

1932. Catalogus Coleopterorum Regionis Palaearcti-

cae. Albert Winkler Verlag, Wien: 795-816.

Zaharadnik P., 1996. Coleoptera: Bostrichoidea. In:

Rozkosny R. & Vanhara J. (Eds), Terrestrial Inver-

tebrates of the Palava Biosphere Reserve of UNE-

SCO, III. Folia Facultatis Scientiarum Naturalium

Universitatis Masarykianae Brunnensis, Biologia, 94:

465-470.

Biodiversity Journal, 2013, 4 (4): 467-470

First record of Pipistrellus pygmaeus (Leach, 1 825) (Mam-

malia Chiroptera) in Sicily (Southern Italy)

Gaetano Fichera 1 , Mauro Mucedda 2 *, Paolo Catalano 1 & Ermanno Pidinchedda 2

'Centro Speleologico Etneo, Via Valdisavoia, 3 - 95123 Catania, Italy

2 Centro Pipistrelli Sardegna, Via G. Leopardi, 1 - 07100 Sassari, Italy; e-mail: m.mucedda@tiscali.it

"■Corresponding author

ABSTRACT We report the capture through mist-nets of two adult females Pipistrellus pygmaeus (Leach,

1825) (Mammalia Chiroptera), along the course of Simeto river, in Adrano (Catania, Sicily,

Italy). Their presence has been reported for the first time in Sicily, increasing the number of

bat species present in the region to twenty three.

KEY WORDS Chiroptera; Pipistrellus pygmaeus', Sicily; Simeto river.

Received 09.10.2013; accepted 22.11.2013; printed 30.12.2013

INTRODUCTION

In Sicily we know only two species of Pipistrel-

lus Kaup, 1829 genus (Mammalia Chiroptera): P.

pipistrellus (Schreber, 1774) and P kuhlii (Kuhl,

1817), both widely distributed all over the Italian

territory.

Recently, the P. pygmaeus (Leach, 1825) was

genetically separated from P. pipistrellus (Barratt

et al., 1997), and they are considered two cryptic

species (Russo & Jones, 2000), therefore its dis-

tribution in Italy, particularly, is little known and

incomplete. Up to now the presence of P. pygmaeus

was reported in Valle d’ Aosta (Debernardi & Pa-

triarca, 2008), Lombardy (Martinoli & Spada,

2008), Tuscany (Agnelli et al., 2005), Umbria

(Spilinga et al., 2013), Lazio and Campania

(Russo & Jones, 2000), Abruzzo (Russo et al.,

2011) and Sardinia (Veith et al., 2011).

Thus, until now Sicily was one of the Italian re-

gions where the presence of P pygmaeus had not

been verified. In this publication we report the pres-

ence of P. pygmeus in Sicily, that was captured

twice with mist-nets along the Simeto river.

MATERIALS AND METHODS

The area of Sicily where the research was con-

ducted is situated at 216 m a.s.l., along the Simeto

river, the second most long Sicilian river, under

the Acquedotto Biscari bridge (Adrano, Catania)

(Fig. 1). This location is characterized by clay

outcrops and a tamarisk vegetation along the

river. Nearby there are citrus crops, vegetables

and uncultivated areas.

The captures were made using two 6-meter long

mist nets (19 mm mesh and 5 pockets each) one on

top of the other for a total height of 4 meters. They

were positioned on the river, a few inches above the

water surface supported by telescopic poles. The

nets were positioned half an hour before sunset and

removed 3 hours after it. The biometric measure-

ments of animals were performed with a calliper

(0.1 mm accuracy) and with a pesola (0.5 g accu-

racy). The recordings of bats sounds were made

with a Pettersson D 240 Bat-detector, in Time ex-

pansion mode, assisted by a Zoom H2 recorder. The

computer processing were carried out with the Pet-

tersson Batsound software.

468

G. Fichera, M. Mucedda, P. Catalano & E. Pidinchedda

All specimens were promptly released at the end

of operations. All activities were carried out with

the authorization of the Sicilian Regional Assesso-

rate for Agricultural and Food Resources (1742-

01/06/2012) and of the Ministry for the

Environment and Territorial and Sea Protection

(0009358-07/06/2012).

Figure 1. The pointer indicates the location of capture of

Pipistrellus pygmaeus in Sicily.

RESULTS

During the research conducted in the areas sur-

rounding Mount Etna on the 1 0th of July and the

8th of September 2013, two capture sessions were

carried out using mist nets along the River Simeto,

and two P. pygmaeus specimens were caught. They

were both adult females, one of them was a lactat-

ing, whose biometric data are reported in Table 1 .

Date

Sex

Forearm

length

Weigth

July 10 2013

Female

lactating

29.0 mm

5.0 g

September

8 2013

Female

30.9 mm

4.9 g

Table 1 . Measures of Pipistrellus pygmaeus.

The animals were identified on the basis of the

following morphological characteristics: the pres-

ence of a ridge between the nostrils, the cells be-

tween the veinings of the wings typical of the

species, as indicated by Dietz & Elelversen (2004).

The color of the fur is of a lighter brown compared

to the P. pipistrellus , almost blond as already ob-

served for specimens of Sardinia (personal obser-

vation) and as shown in the images of Dietz et al.

(2007) (Fig. 2). The animals were released im-

mediately after examination. The validity of the

species was also confirmed by the recording of the

sounds emitted by the two bats released from the

hand in open space. Only the final part of the

recorded sequences were analysed, to avoid in-

fluence by the hand release. Three calls from each

bat were measured from power spectra, given the

following values of Frequency of Maximum Energy

(FMaxE): 56.5 ± 0.2 kHz for the first bat and 56.6

±0.7 kHz for the second bat.

Sounds of other specimens of P. pygmaeus with

typical feeding buzz were also recorded, showing

foraging activity in this locality (Fig. 3).

DISCUSSION AND CONCLUSIONS

Figure 2. The specimen of Pipistrellus pygmaeus captured

on July 10 2013. Note the light-colored fur.

In Sicily, Agnelli et al. (2008) report a list of

20 species of bats, some of which still have to be

First record of Pipistrel I us pygmaeus (Leach, 1825) (Mammalia Chiroptera) in Sicily (Southern Italy)

469

Figure 3. Spectrogram of a capture sequence of a Pipistrellus pygmaeus with final feeding buzz.

confirmed. More recently Hypsugo cfr. darwinii

(Tomes, 1859) (Veith et al., 2011) and Myotis

bechsteinii (Kuhl, 1817) (Di Salvo et al., 2012)

were added and the presence of Barbastella bar-

bastellus (Schreber, 1774) was also confirmed (Mu-

cedda et al., 2012).

Di Salvo et al. (2009) only hypothesized the pre-

sence of P. pygmaeus in Sicily, by bioacoustic re-

cordings that didn't allow to distinguish the species

from Miniopterus schreibersii with certainty.

The capture of P pygmaeus allows us to add a

new species to the bat fauna of Sicily and brings the

number of confirmed species of bats living in the

island to 23: Rhinolophus ferrumequinum (Schre-

ber, 1774), R. hipposideros (Bechstein, 1800), R.

mehelyi Matschie, 1901, R. euryale Blasius, 1853,

Myotis bechsteinii (Kuhl, 1817), M. blythii (Tomes,

1857), M. capaccinii (Bonaparte, 1837), M. dauben-

tonii (Kuhl, 1817), M. emarginatus (Geoffroy,

1806), M. myotis (Borkhausen, 1797), M. mystaci-

nus (Kuhl, 1817), M. nattereri (Kuhl, 1817), Pip-

istrellus pipistrellus (Schreber, 1774), P. kuhlii

(Kuhl, 1817),/! pygmaeus (Leach, 1825 ), Hypsugo

savii (Bonaparte, 1837), H. cfr. darwinii (Tomes,

1859), Eptesicus serotinus (Schreber, 1774), Bar-

bastella barbastellus (Schreber, 1774), Nyctalus la-

siopterus (Kuhl, 1817), Plecotus austriacus

(Fischer, 1829), Miniopterus schreibersii (Kuhl,

1817), Tadarida teniotis (Rafmesque, 1814).

The finding also allows us to ensure its repro-

duction on the island and to expand the range of

the species in Italy, of which it is the southern

limit. Future research should aim at identifying

roosts, to define the distribution of P. pygmaeus in

Sicily and establish the conservation measures that

have to be taken.

ACKNOWLEDGEMENTS

We thank Mariacristina Borrello for the help

during the field work, Marianna Catalano and

Emily Halliday for the translation.

REFERENCES

Agnelli P., Di Salvo I., Russo D. & Sara M., 2008. Chi-

rotterofauna della Sicilia (Mammalia Chiroptera). In:

AA.VV., 2008. Atlante della Biodiversita della Si-

cilia: Vertebrati terrestri. Collana Studi e Richerche

dell’ ARPA [Agenzia Regionale per la Protezione

dell’Ambiente - Sicilia], n.6. ARPA Sicilia, Palermo,

25-40.

Agnelli P., Guaita C. & Vergari S., 2005. Un contributo

per la pianificazione degli interventi nella Tenuta di

San Rossore (Pisa): lo studio della chirottero fauna.

In: Prigioni C., Meriggi A., Merli E., 2005. V Con-

470

G. Fichera, M. Mucedda, P. Catalano & E. Pidinchedda

gresso Italiano di Teriologia. Arezzo, 10-12 Novem-

bre 2005, 80.

Barratt E.M., Deaville R., Burland T.M., Bruford M.W.,

Jones G., Racey RA. & Wayne R.K., 1997. DNA an-

swers the call of pipistrelle bat species. Nature, 387:

138-139.

Debernardi P. & Patriarca E., 2008. Prima segnalazione

di Myotis bechsteinii, Myotis daubentonii, Myotis

nattereri, Nyctalus leisleri, Pipistrellus pygmaeus,

Plecotus macrobullaris e Tadarida teniotis in Valle

d’Aosta. Aggiomamento deirinventario dei chirotteri

noti per la regione. Revue Valdotaine d'Histoire

Naturelle, 62: 5-27

Dietz C. & von Helversen O., 2004. Illustrated

identification key to the bats of Europe. Tuebingen

and Erlangen (Germany), Electronic publication, Ver-

sion 1, 72 pp.

Dietz C., von Helversen O. & Nill D. 2007. Handbuch

der fledermause Europas und Nordwestafrikas. Bi-

ologie. Kennzeichen. Gefardung, fmack kosmos Ver-

lag, Stuttgart, 399 pp.

Di Salvo I., Russo D. & Sara M., 2009. Habitat prefer-

ences of bats in a rural area of Sicily determined by

acoustic surveys. Hystrix, the Italian Journal of

Mammalogy, 20: 137-146.

Di Salvo I., Fulco A., Sara M. & Russo D., 2012. Oc-

curence of Bechstein’s bat Myotis bechsteinii (Chi-

roptera: Vespertilionidae) in Sicily. Natura Rerum, 2:

75-80.

Martinoli A., & Spada M., 2008. Pipistrello pigmeo. In:

Vigorita V., Cuce L. (a cura di). La fauna selvatica in

Lombardia. Rapporto 2008 su distribuzione, abbon-

danza e stato di conservazione di uccelli e mam-

miferi. Regione Lombardia, p. 282.

Mucedda M., Fichera G. & Pidinchedda E., 2012. Record

of Barbastella Barbastellus (Schreber, 1774) (Chi-

roptera, Vespertilionidae) in Sicily after 56 years.

Natura Rerum, 1: 79-81.

Russo D., Cistrone L., Garonna A.P. & Jones G., 2011.

The early bat catches the fly: daylight foraging in so-

prano pipistrelle bats. Mammalian Biology, 76: 87-

89.

Russo D. & Jones G., 2000. The two cryptic species of

Pipistrellus pipistrellus (Chiroptera: Vespertilionidae)

occur in Italy: evidence from echolocation and social

calls. Mammalia, 64: 187-197.

Spilinga C., Russo D., Carletti S., Jimenez Grijalva M.P.,

Sergiacomi U. & Ragni B., 2013. Chirotteri dell’Um-

bria. Distribuzione geografica ed ecologica. Regione

Umbria. Universita degli Studi di Perugia, 187 pp.

Veith M., Mucedda M., Kiefer A. & Pidinchedda E.,

201 1. On the presence of pipistrelle bats {Pipistrellus

and Ifypsugo; Chiroptera: Vespertilionidae) in Sardi-

nia. Acta Chiropterologica, 13: 89-99.

Biodiversity Journal, 2013, 4 (4): 471-482

A review of the impacts of invasive aquatic weeds on the bio-

diversity of some tropical water bodies with special reference

to Lake Victoria (Kenya)

Peninah Aloo 1 *, William Ojwang 2 , Reuben Omondi 2 , James Murithi Njiru 3 & Dalmas Oyugi

'Department of Aquaculture and Fisheries Management, School of Natural Resources and Environmental Studies, Karatina Uni-

versity, RO. Box 1957, Karatina, Kenya

2 Kenya Marine and Fisheries Research Institute, RO Box 1881, Kisumu, Kenya

’Department of Fisheries and Aquatic Sciences, Moi University, P.0 Box 3900, Eldoret, Kenya

4 Kenya Wildlife Service Training Institute, Kenyatta Avenue, P.O. Box 842, Naivasha, Kenia

*Corresponding author: e-mail: aloopenina@yahoo.com

ABSTRACT Aquatic weeds may be defined as troublesome or unsightly plants growing in abundance in

aquatic situations where they are not wanted. These plants are either adapted to continuous

supplies of water or are at least tolerant to waterlogged soil conditions for substantial periods

of time. The wide range of adaptation to varying amounts of water, and the impossibility of

sharply distinguishing between aquatic and terrestrial environments, makes it difficult to

precisely define an aquatic plant. The menace of water weeds is reaching alarming propor-

tions in many parts of the world, especially in tropical water bodies where they have led to

serious ecological and economic losses. Lake Victoria, Kenya, which is the largest freshwater

body in the tropics, has undergone serious ecological changes including over-exploitation

of its fishery resources, degradation of the catchment area, introduction of exotic fish species

and invasion by the water hyacinth, Eichhornia crassipes (Mart.) Solms (Pontederiaceae),

among others. The presence of the weed in the lake has led to many problems including

blockage of water pumps, reduced fishing activities and increase in water borne diseases

such as schistosomiasis. Positively, aquatic weeds constitute a free crop of great potential

value; they are a highly productive crop that requires no tillage, fertilizer, seed, or cultivation.

Moreover, these plants have the potential for exploitation as animal feed, human food, source

of food to some aquatic organisms, soil additives, fuel production, wastewater treatment,

source of raw materials and habitat to many organisms. This paper reviews the effects of

aquatic weeds in aquatic systems with examples from some selected waterbodies and special

reference to Lake Victoria.

KEY WORDS Invasive; Lake Victoria; weeds; tropical; waterbodies.

Received 14.10.2013; accepted 22.11.2013; printed 30.12.2013

INTRODUCTION

Aquatic weeds are higher plants that grow in

water or in wet soils but some weeds may endure

periods of desiccation. They usually occur along the

shores of water bodies like dams, lakes and along

rivers and river mouths. The distribution, perma-

nency and quality of water available for their occu-

pation govern their distribution and ecology. The

most variable environmental factors of basic eco-

logical importance for aquatic plants are the length

of the period during which water is present; whether

472

P. Aloo, W. Ojwang, R. Omondi.J.M. Njiru & D. Oyugi

the water is still or moving; the availability of plant

nutrients; and the quality and quantity of light pene-

tration into the water. Factors that influence the

establishment and distribution of aquatic plants in-

clude: depth, topography, types of substrate, expo-

sure to currents and/or wind and water turbidity.

The distribution of macrophytes is often related to

their method of attachment (Sculthorpe, 1976).

Aquatic plants, like most water organisms, are

more widely distributed throughout the world than

terrestrial plants. This is because factors or condi-

tions required by aquatic plants are uniform in

general than those to which land plants must adapt

to. The aesthetically pleasing appearance and

unique growth form of floating aquatic weeds

have been responsible for their spread to various

tropical and sub-tropical countries by humans dur-

ing the 1800s and 1900s. In Kenya, aquatic weeds

were used to grace aquaria and ornamental ponds

from where they escaped into natural or artificial

water bodies causing serious problems (Njuguna,

1992).

Macrophytes are among the most productive

plant communities in the world (Sculthorpe, 1976)

and are known to provide nutrition for humans and

herbivorous animals. In general, water plants have

both positive and negative importance to man,

either directly or indirectly (Mitchell, 1974).

Study area

Lake Victoria (Fig. 1) is the largest tropical lake

in the world. The lake is shared by Kenya (6% by

area), Uganda (43%) and Tanzania (51%). It has a

mean depth of 40 m, maximum depth of 84 m,

shoreline of 3450 km, a water retention time of 140

years and a catchment area of 193 000 km 2 , which

extends into Rwanda and Burundi. It is situated

1134 m above sea level and has a surface area of

68,800 squared kilometers. The lake is roughly in

square shape; its greatest length and width being

about 400 and 320 Ion, respectively. Much of the

lake is less than 40 m deep and the deepest part, 60-

90 m, is in the northeast. The bottom is mainly cov-

ered by a thick layer of organic mud, but with

patches of hard substrate, sand, shingle or rock

(Scholtz et al., 1990). The coastline is intended with

bays and gulfs. The Kagera and Nzoia rivers are the

principal influents while the only outflow occurs to

the River Nile via Lake Kyoga.

The lake receives nearly 80% of its water inputs

from rainfall and a similar amount is lost via eva-

poration (Bootsma & Flecky, 1993) (Table 1).

Parameter

Characteristic

Lake area (km 2 )

68,800

Maximum Depth (m)

(92)

Mean Depth (m)

Volume (km 3 )

2,760

Catchment (km 2 )

194,000

Outflow (0)(km 3 /y)

Inflow (I)(km 3 /y)

Precipitation (P)(km 3 /y)

100

Evaporation (km 3 /y)

100

Flushing Time (V/0)(years)

140

Residence time (V/(P+I)(years)

Table 1 : Catchment and basin parameters of Lake Victoria.

Source: Bootsma & Hecky, 1993; IDEAL 1990; Welcomme,

1972; Sigel & Coulter, 1996.

Lake Victoria is an important source of afford-

able protein food in the form of fish. It provides

employment, income, and export earnings to the ri-

parian communities. It is a source of water, mainly

taken without treatment, and is also used for trans-

port. As well as their food value, the fish species of

the lake are of important evolutionary significance

and have been extensively studied.

Lake Victoria Environmental Management

Programme (LVEMP) (2001) reported that the

quality of Lake Victoria waters has been deterio-

rating over the last 30 years due to increased in-

flow of nutrients into the lake. Population increase

within the catchment area also raises the need for

more food hence, there is widespread use of fer-

tilizers to boost food production, adding to the

agricultural load in the lake. Concentrations of

phosphorus have increased substantially within

deep waters while nitrogen levels have increased

closer to the shores.

Impacts of invasive aquatic weeds on the biodiversity of some tropical water bodies with special reference to Lake Victoria 473

Figure 1. Map of Lake Victoria.

As a consequence of the increase in nutrients

levels, vast algal blooms have become a common

feature of Lake Victoria, with blue green algae

dominating, causing de-oxygenation of the water,

increased sickness for populations using the lake

water and high water treatment costs together

with loss of deep water fish species. Fish inhabit-

ing shallow waters are also at risk from periodic

up welling of hypoxic waters. In addition, reports

also indicate increased invasion of the lake by

aquatic weeds including the abnoxious water

weed, Eichhornia crassipes (Mart.) Solms (Pont-

ederiaceae).

Types and characteristics of aquatic weeds

Aquatic macrophytes can be sub-divided into

four groups on the basis of their water requirements,

life forms and habitats. The categories include:

submerged, emergent, floating leafed and free-

floating macrophytes.

Submerged macrophytes are plants that are

usually rooted in the substrate but are completely

under the water surface. These plants cannot be

established at a depth more than 10m and are ad-

versely affected by large fluctuation in the water

level. Submerged plants have thin finely dissected

leaves adapted for rapid exchange of nutrients

with water. Examples include Potamogeton pecti-

natus L. (Potamogetonaceae), Najas horrid A.

Braun ex Magnus (Hydrocharitaceae) and Val-

lisneria spiralis L. (Alismataceae). The growth

rate of submerged plants has been recorded to

range from 10 mg dry matter per day for Cerato-

phylum demersum L. (Ceratophyllaceae) to over

80 mg in Hydrilla verticillata (L.f. Royle) (Hy-

drocharitaceae) (Denny, 1972).

Emergent macrophytes include plants which

are rooted with their principal photosynthetic sur-

faces above the water. They are markedly exclu-

sive since they always occur in dense

monospecific stands reminiscent of monocultures.

Boyd (1969) suggested that this comes about

because of their success in taking up and storing

nutrients, which are later made available to young

growing organs by translocation. Emergent plants

such as Typha sp. (Typhaceae) and Cyperus pa-

pyrus L. (Cyperaceae) are generally rigid and are

not dependant on water for support. They have

both high growth rate and high biomass (Symoens

et al., 1981). The emmergents such as Typha can

only occur in shallow water or damp soils along

the shoreline and are unlikely to survive large

changes in lake level (Mitchell, 1974).

Floating leafed macrophytes are plants that are

rooted but have floating leaves. Examples of such

macrophytes are Nymphaea lotus L. (Nymphaeaceae)

and Trapanatans L. (Trapaceae). These are usually

restricted in the shallower areas of water bodies.

Free-floating macrophytes are aquatic plants

that float on the water surface. Typical examples

of free floating macrophytes include Salvinia

molesta D. Mitch. (Salviniaceae) and E. cras-

sipes. These plants are independent of depth and

substratum requirements as well as largely unaf-

fected by changes in lake levels; hence they are

the greatest threat to any water body (Adeniji,

1979). A review of literature makes it clear that in

the tropics, the greatest threat is posed by this

group of macrophytes.

DISCUSSION

Problems of aquatic weeds

Aquatic plants are considered nuisance when

excessive growth interferes with desired water uses

474

P. Aloo, W. Ojwang, R. Omondi.J.M. Njiru & D. Oyugi

in a number of ways (Adeniji, 1979). In line with

increased industrialisation, travel and communica-

tion, agricultural productivity, increased human

population and changes in consumption, problems

associated with aquatic weeds have increased in the

last century (Davis & Hirji, 2003).

The menace of some aquatic plants may reach

alarming rates especially in the tropical regions

where warm water fosters plant growth. The

problem is aggravated by the increasing en-

richment of water bodies by poor land use practices

and other effluents from the human and industrial

wastes. Less than 20 of the 700 species of macro-

phytes are considered weeds (Triest, 1993).

Although originally perceived as a practical prob-

lem for fishing and navigation, recent research in-

dicates that aquatic weeds are now also considered

a threat to biological diversity affecting fish fauna,

plant diversity and other freshwater life and food

chains (Garry et al., 1997). Excessive population

of macrophytes, especially floating and emergent

weeds, results in exclusion of light from the lower

layers of water. This can cause reduction of pri-

mary production and may also affect the various

physical and chemical characteristics of water.

They can also lead to blockage of hydroelec-

tric installations and water intakes and irrigation

outlets. Submerged and free floating plants have

also been reported to interfere with fishing and

navigation activities. The invasion by aquatic

weeds of any environment has always led to

increase in diseases such as malaria and bilharzia,

whose vectors use the weeds as breeding grounds

as in Lake Victoria (Johnson et al., 1990; Twongo,

1996). The weeds also provide hiding grounds for

dangerous organ-isms such as snakes.

It is evident that aquatic weeds are a total nui-

sance from the point of view of the human use of

the water where they occur. It is important to eval-

uate the effects of the weeds on the entire ecosystem

for example, on the invertebrates, benthic fauna,

fish and large animals.

Effects of aquatic weeds on aquatic organisms

Mitchell (1974) stated that the appearance of

macrophytes in any aquatic ecosystem leads to an

increase in the density of other plants and animal

species. Therefore, two major effects of these weeds

on the ecosystem are habitat diversification and

food source for the organisms.

a) Habitat diversification

Macrophytes physically change aquatic ecosys-

tems in several ways which are important in habi-

tat diversification. They slow down the movement

of sediments in fast stretches of rivers (Butcher,

1933); consequently, the number of species in-

creases in previously unproductive regions. The

mesh formed by finely divided leaves of some

submerged species ( Potamogeton sp.), and the

networks of filaments formed by some algae pro-

vide excellent refuge for juvenile fish (Mitchell,

1974). The structurally complex habitats formed

by macrophytes coupled with periodic low oxy-

gen levels is also known to protect several fish

species from both excessive human exploitation

and predation (Chapman et al., 1996; 2002). Sev-

eral insect larvae and fish species such as the tilapi-

ines use submerged macrophytes materials in the

construction of nests. The older rhizomes of reed

swamp emergents such as papyrus or any soft

plant material present in shallow water provide a

burrowing substrate for insects such as mayflies.

These invertebrate fauna form a crucial link in the

foodweb of natural and man-made tropical lakes

(Petr, 1970).

Investigations by Bowmalcer (1968, 1973) and

Mitchell (1976) in Lake Kariba demonstrated that

Salvinia mats play an important role as a habitat

for young fish and their prey and, that mobile mats

swept over the lake by wind assist in the distribu-

tion of fish fry. Reports from Lake Naivasha in

Kenya indicated that the abundance of some inver-

tebrates such as the water boatman Micronecta

scutellaris (Stal, 1858), Hemiptera Corixidae, was

closely related to S. molesta mats (Aloo, 1988).

Perhaps the most important contribution of macro-

phytes to the ecology of Nyumbaya Mungu reser-

voir in Tanzania was their close association with

the fish and the aufwuchs communities. In this

reservoir, the major shoals of commercial tilapiine

cichlids fishes such as Sarotherodon pangani

(Lowe, 1955) and S. jipe (Lowe, 1955) (= genus

Oreochromis) were reported to be closely associ-

ated with aquatic plants such as the Cyperus arti-

culates L. (Mitchell, 1974).

Impacts of invasive aquatic weeds on the biodiversity of some tropical water bodies with special reference to Lake Victoria 475

Emergent plants such as C. papyrus also pro-

vide special conditions needed by swamp inhabi-

tants including the swamp worm Almaemini

(Clark et al., 1987). This worm is adapted to life

under anaerobic conditions and can even live for

several days without oxygen. In Lake Volta, Petr

(1968) demonstrated the importance of the inver-

tebrates associated with weed beds and sub-

merged plant surfaces while in Lake Kariba,

Bowmaker (1968) noted their abundance among

the floating mats of Salvinia. McLachan (1969)

reported that the appearance of submerged macro-

phytes such as Potamogeton and emergents such

as Ludwigia (Onagraceae) in Lake Kariba re-

sulted in marked changes in the benthic fauna.

Total mud species numbers increased from 28 to

46 in the bottom close to the base of these plants.

In Lake Victoria, the introduction of E. crassipes

led to an increase in the population of leeches,

dragon fly nymphs and several mollusc species

which were reported to occur under hyacinth

mats (Twongo, 1996).

Lurthermore, the macrophytes coverage, par-

ticularly the swampy areas are known to subject

fish populations in particular to extreme physico-

chemical regimes that in the long-term acts as

natural selection forces responsible for speciation.

The fish assemblage in Lake Victoria which was

previously composed of the haplochromine ci-

chlids was thought to have evolved through this

speciation mechanism. According to postulation

by Greenwood (1974) and Kaufman (1992), these

unique habitats together with small water bodies

around Lake Victoria acted literally as boiler

plates for speciation and the multiplication of ci-

chlids.

b) Food source

Several studies using stable isotopes have reaf-

firmed plants, particularly aquatic plants (both C3

and C4 plants) as the main source of energy in

most water bodies (Ojwang et al., 2004; 2007).

Other studies using traditional gut contents analy-

sis have shown that herbivorous tilapiine cichlids

fishes such as Tilapia rendalli (Boulenger, 1897)

and T. zillii (Gervais, 1948) feed on aquatic plants

as their main diet. Though there are some indica-

tions of preference for submerged plants ( Elodea ,

Ceratophyllum and Najas ), floating plants (duck-

weeds, Wolffia spp., Araceae) and emergent plants

( Typha ) are also eaten (Sculthorpe, 1976; Black-

burn et al., 1971). The development of submerged

vegetation in the lower Volta was reported to have

profound effects on the whole ecosystem. Lish

found food and shelter in the beds of hydrophytes.

Populations of Tilapia spp. and Neritina sp.

(Gastropoda Neritidae), a gastropod which feeds

on Vallisneria (Hydrocharitaceae) increased re-

markably, a phenomenon which was attributed to

increase in productivity in the ecosystem by both

Potamogeton and Vallisneria plant species (Hall

& Pople, 1968).

Other fish species such as Disticho dusschenga

Peters, 1852, D. mossambicus Peters, 1852 (Disti-

chodontidae) and T. rendalli also fed on Pota-

mogeton and Vallisneria. T. rendalli was reported

to be the only fish specialized in grazing on macro-

phytic plants in Nyumbaya Mungu, although the

scavengers such as Clarias mossambicus (Peters,

1852) (Clariidae) and Synodontis punctulata

(Gunther, 1889) (Mochokidae) occasionally in-

gested plant fragments (Bailey et al., 1978). In the

Kenyan Lake Naivasha, Oluoch (1983) reported

an abundance of macrophytes in the stomach of

the crayfish Procambarus clarkii Girard, 1852

(Crustacea Cambaridae).

It is not only fish species that depend on macro-

phytes as source of energy. In Lake Cabora Bassa,

Bond & Roberts (1978) observed hippos, baboons,

monkeys, warthog and kudus feeding on E. cras-

sipes particularly during the dry season. Water-

fowls such as the Egyptian geese fed to a limited

extent on stranded plants (Davies et al., 1975).

Lake Naivasha supports 31,000 ducks which for-

age on a wide range of aquatic plants (6 spp.) and

43,000 coots Fulica cristata Gmelin, 1789 (Aves

Rallidae) that depend largely on the submerged

plant Naja spectinata (Pari.) Magn., Bivalvia

Najadaceae (Watson & Parker, 1970; Clark et al.,

1987).

The lake also supports a large population of

Myocaster coypus (Molina, 1782), Rodentia My-

ocastoridae, which is reported to enjoy the emergent

vegetation around the lake shore. Recent studies in-

dicate that the decline in aquatic macrophytes in

Lake Naivasha has led to the low numbers of the

crayfish, P. clarkii (Harper, 2010).

476

P. Aloo, W. Ojwang, R. Omondi.J.M. Njiru & D. Oyugi

c) Other benefits of aquatic plants

A few species of aquatic plants are directly im-

portant to man for food and for materials used in

building/constructions. The Indian rice, water chest-

nut and delta potatoes are sources of food. Some

weeds (Nile cabbage, water spinach) are used as

vegetable among many African communities. In

some parts of the world, the bulrush is used for

building boats, floor mats and wall partitions. The

papyrus, C. papyrus, has been utilized for weaving

baskets, making mats and thatching huts especially

in rural communities (Prescott, 1969).

Limnologically, aquatic plants and shoreline

vegetation play important roles including beach

building, the filling in of lake margins with the

accompanying aging and eutrophication and pre-

vention of shore erosion. A few aquatic plants bring

about the deposition of lime, thus, after a long pe-

riod of time, produce useful marl deposits. Besides

these, there are many interactions between aquatic

plants, water chemistry and the nature of bottom de-

posits (Prescott, 1969).

Macrophytes strands can act as a filter for ex-

cessive nutrients, which would otherwise lead to

eutrophication of adjacent water bodies. Although

the absorption may not in itself ensure the re-

moval as the plants might re-release them on de-

composition, the wet low oxygen soils favour

denitrification by bacteria leading to loss of nu-

trients. The plants also remove heavy metals, bio-

cides and other toxins from the water temporarily

into their tissues. In theory, this could harm or-

ganisms higher up the food chain, however, the

plants have various biological, chemical, bio-

chemical and physical processes which immobi-

lize, transform and fix the contaminants (Malthy,

1986).

Recent changes in the distribution and di-

versity of macrophytes in Lake Victoria

Lake Victoria has undergone drastic ecological

changes since the introduction of alien fish species

in the 50s and 60s. Prior to these introductions, the

lake’s fisheries were dominated by haplochromines

which were alleged to have hindered the establish-

ment of macrophytes in the inshore areas by con-

stantly disturbing the substrate (Witte et al., 1992).

The large populations of haplochromines were later

decimated by the piscivorous Nile perch, Lates

niloticus Linnaeus, 1758 (Latidae).

Aquatic plants along the lake’s shoreline were

dominated by papyrus, C. papyrus. Floating is-

lands which were common during the rainy sea-

sons consisted of populations of the emergent

macrophytes which were sloughed by currents and

winds. The other plants on floating islands were

few mainly comprising climbers such as Ipomoea

aquatic Forssk., Convolvulaceae. Before the inva-

sion of the lake by water hyacinth, succession of

aquatic plants were not as drastic, a phenomenon

earlier reported by Russell (1942) and Penfound

& Earle (1948).

Macrophytes in Lake Victoria mostly occur in

the vicinity of river mouths and sheltered bays.

River mouths favour the establishment of macro-

phytes because of sedimentation from rivers while

the incoming nutrients stimulate the growth of

various plant species. More than 30 families and

60 species of macrophytes have been recorded

from Lake Victoria. These are dominated by Cyper-

aceae, of which many species have not been

identified.

E. eras sipes (Fig. 2) was first recorded on the

Ugandan side of Lake Victoria in 1988 (Twongo,

1991). The weed reached Lake Victoria from River

Kagera which originates in Burundi through

Rwanda and Tanzania and enters the lake in

Uganda. While the introduction of E. crassipes may

have been accidental, the weed was originally

transported from its native home in South America

to be used as an-ornamental plant. By 1992 the

weed was firmly established in most suitable

littoral environments along the entire shoreline of

Uganda (Twongo, 1996). Water hyacinth had

already been reported in Tanzania (AAPS, 1990) as

well as in the sheltered bays in Kenya.

The weed was confirmed to be present on the

Kenyan side of Lake Victoria in 1991, although

unconfirmed reports of its presence at the River

Yala swamp had been received as early as 1990

(Mailu, 1999). The spread of the weed across Lake

Victoria was greatly aided by strong winds which

swept large masses of the weed from the bays along

the north-western shores across the lake, to the

northern and possibly north-eastern shores.

Currently, E. crassipes is widespread on the

Kenyan waters of Lake Victoria occurring abun-

dantly in Winam Gulf, Karungu Bay, Muhuru

Impacts of invasive aquatic weeds on the biodiversity of some tropical water bodies with special reference to Lake Victoria All

Bay, Luanda, Usenge, Asembo Bay, Osieko

Beach and along Yala Swamp. Small mats of the

weed were reported to occur at Sio Port, Mbita

Point and Nzoia River (Aloo, 2003). The weed is

estimated to occupy a total of 5,000 hectares on

the Kenyan side of Lake Victoria. The geo-climatic

requirements of water hyacinth favour its growth

and reproduction in the lake. Factors that lead to

its increased growth and production include

nutrients especially nitrogen and phosphorus.

Lake Victoria is nutritionally enriched by drainage

from agricultural, discharge from factories, urban

waste and inadequately treated sewage effluents.

The population of water hyacinth has, therefore,

spread rapidly under the nutritionally favourable

conditions and ambient temperatures in Lake

Victoria.

Macrophytes succession in Lake Victoria

after water hyacinth invasion

On the establishment of water hyacinth in Lake

Victoria, the floating mat provided substrate for

the establishment of other plants including terres-

trial plant species. Succession of the aquatic plants

has also been accelerated due to the shading effect

on other plants and increase of organic matter

upon the death and eventual sinking of the plant

biomass. Pistia stratiotes L. (Araceae), Azolla pin-

nata R. Br. and A. nilotica Decne. ex Mett. (Azol-

laceae) were the most common floating

macrophytes in Lake Victoria before the invasion

of water hyacinth. The grass, Vossia cuspidata

(Roxb.) Griff. (Poaceae) formed a thin layer

behind P. stratiotes followed by C. papyrus ,

Aeschynomene elaphroxylon Taub. (Fabaceae),

Phragmites australis (Cav.) Trin. ex Steud. (Poaceae)

and Typha domingensis Pers. (Typhaceae). On

establishment in mid-1990s, the large populations

of water hyacinth both smothered and pushed,

landwards, P. stratiotes and Azolla spp. This com-

pact mass, the resulting debris and proximity to

land led to the increase of V cuspidata and

Echinochloa pyramidalis P. Beauv. (Poaceae)

communities by acting as a substrate.

Other emergents which were reported to be closely

associated with water hyacinth mats were Hibiscus

diversifolius Jacq. (Malvaceae) and T. domigensis.

Figure 2. Water hyacinth, Eichhornia crassipes.

These emergents in turn shaded and competed for

nutrients resulting in the death of the latter. After

the invasion of water hyacinth, floating islands

consisted mainly of the hyacinth and V cuspidata.

Aquatic communities that were decimated by water

hyacinth included the floating leafed N. lotus and

T. natans and the submerged plants, C. demersum

and N. horrida.

Effects of Water Hyacinth in Lake Victoria

The problems created and damage caused by

E. crassipes are greater than those due to any other

floating aquatic weed in Lake Victoria. It is no

wonder that the weed has been described as truly

the world 4 s worst aquatic weed (Harley, 1990).

The weed forms dense, impenetrable mats, which

impede or prevent boat traffic in natural water

courses, artificial or natural lakes, irrigation and

flood mitigation canals. Detailed assessment of the

impact of water hyacinth infestation on a given

ecosystem and its resources might justify the huge

economic tag that is required to control the vast

expanses of the weed around the shores of Lake

Victoria (Twongo, 1996). The rapid spread of E.

crassipes in Lake Victoria led to serious economic,

social, health and environmental impacts in the

riparian states.

The weed has affected the fisheries, water

supply and transport, environmental health and

power generation, interfering with the lives of

over 30 million people that depend on the lake

(Aloo, 2003). Fish production in the lake has

declined since the weed has blocked many fish

landing beaches; it has infested sheltered bays

478

P. Aloo, W. Ojwang, R. Omondi.J.M. Njiru & D. Oyugi

which are breeding grounds of fish while in open

waters the mats of the weed has swept away and

entangled fleets of nets. The mats also pose

obstruction to fisheries exploitation which has led

to higher operation costs hence increase in fish

prices. Many landing beaches have been aban-

doned and income-generation from the sale of fish

has been negatively affected.

Water hyacinth has caused clogging of water

pipes, leading to frequent water shortages in

lakeshore urban centres. The infestation has occa-

sionally resulted in disruptions, delays and rising

operation costs in the transport sector yet water

transport is the cheapest means of transport for the

lakeside communities. The weed has grossly inter-

fered with cross border trade between the three East

African states. Decomposition of the weed requires

large amounts of oxygen which leads to a drop in

levels of oxygen in areas infested by the weed,

reducing survival of aquatic organisms especially

fish. The decomposed material also leads to pollu-

tion of the water making it unfit for any use. The

weed has also been reported to impact negatively

on the health of lakeside communities, since it pro-

vides optimal habitat for mosquitoes and snails

which are vectors for malaria and bilharzia respec-

tively (Twongo, 1996).

While some studies have been carried out in

Kenya to investigate the association of water

hyacinth with various vector borne diseases, large

number of snails are usually found among water

hyacinth mats in Lake Victoria. Dazo et al. (1966)

reported a positive association between bilharzia

vectors Bulinus truncatus Audouin, 1837 (Gas-

tropoda Planorbidae) and Biomphalaria alexan-

drina Ehrenberg, 1831 (Gastropoda Planorbidae)

and various aquatic macrophytes including water

hyacinth in Egypt. There is an urgent need to

monitor the impact of water hyacinth infestation on

incidence of water associated vector-borne diseases.

The weed also provides a hiding place for dan-

gerous organisms such as snakes. Water hyacinth is

now a widespread weed occurring in dams, rivers

and home aquaria all over Kenya. The weed has

formed a green carpet on the Nairobi dam, situated

in Motoine River Valley South of Kibera slums in

Nairobi, covering the entire water surface. Today,

the dam has completely dried out thus affecting the

recreational activities that existed in the dam before

invasion by water hyacinth (Aloo, pers. obs.).

Currently, the weed has been reported to be a

threat to the Athi River, which is Kenya’s second

largest river. In Masinga dam, a few patches of

the weed have been observed floating on the

water.

Positive attributes of water hyacinth in Lake

Victoria

Aquatic weeds have some positive effects in the

water bodies where they occur. In Lake Victoria,

reports have indicated that there has been a resur-

gence of the endemic fish species especially some

Cichlidae of the genus Haplochromis , Clarias

gariepinus (Burchell, 1 822) (Clariidae), Protopte-

rus aethiopicus Heckel, 1851 (Protopteridae) and

Elephant-snout fish Mormyrus kannume Forsskal,

1775 (Mormyridae) as a result of the invasion of

the lake by the weed (Bugenyi & Van de Knaap,

1998; Njiru et al., 2000; Ogari, 2000). This is be-

cause mats of the weed provide hiding habitat for

many fish species (Ogari, 2000). The swampy

areas, particularly the remaining wetlands along

the shores of Lake Victoria, are recognized as refu-

gia to some of the native fish populations (Denny,

2000). Most of the fish species inhabiting the

swampy areas are tolerant of extreme environmen-

tal conditions (Chapman et al., 2002). For example,

P. aethiopicus and C. gariepinus are physiologi-

cally hardy and have structures that enable them

access to free atmospheric oxygen.

The structural complexities of the swampy

habitats coupled with periodic low oxygen levels

protect a number of indigenous fish species from

both excessive human exploitation and predation

from Nile perch. Species richness has been

observed to be specifically higher in the swampy

areas along the shores of the lake than in the inte-

rior areas (Chapman et al., 1996; 2002; Balirwa et

al., 2003). These are areas that physiologically limit

the foraging abilities of Nile perch, which frequent

waters of higher oxygen content. On the other

hand, the unique ability of macrophytes to improve

water clarity favours both mate recognition

amongst the haplochromine cichlids and at the

same time promoting the coexistence of the species

with the Nile perch, since the predatory techniques

of Nile perch are best suited to waters of interme-

diate turbidity.

Impacts of invasive aquatic weeds on the biodiversity of some tropical water bodies with special reference to Lake Victoria 479

E. crassipes has been inhabited by many aquatic

insects which are an important diet to the juveniles

of many fish species. In some parts of the lake,

there has been increase in the abundance of

aquatic birds which depend on water hyacinth as

their food and resting stations in the course of their

foraging activities. Water hyacinth has also been

utilized for various purposes such as fodder, raw

materials and for water purification purposes

(Twongo, 1996). The weed’s role as water purifier

is of great importance in Lake Victoria where

many lake side municipalities’ wastewater treat-

ment plants are presently broken or are ill main-

tained, thus the amount of raw sewage flowing

directly into Lake Victoria is enormous. This en-

hances the proliferation of the water hyacinth and

other aquatic weeds.

Re-appearance of water hyacinth in Lake

Victoria

Currently, water hyacinth has abundantly reap-

peared in the lake (pers. obs.). Biological control

method through the introduction of the natural

enemies of the plant, Neochetina eichhorniae

Warner, 1970 and N. bruchi Hustace, 1926

(Coleoptera Curculionidae) was widely used in the

early years of invasion. This method was believed

to have reduced the total area covered by hyacinth

from 6000 ha in April 1998 to 400 ha in 2001

(Wawire & Ochiel, 2001).

Today, there have been serious resurgences of

the weed especially around Homa Bay and Kisumu

areas. Unpublished reports also indicate that there

is an ecological association between water hyacinth

and the hippo grass which is also found abundantly

in the lake. Thus the case of succession of the

weeds in Lake Victoria is a subject of interest to

ecologists and require further studies.

CONCLUSIONS

The impacts of aquatic weeds are diverse and

their significance in ecosystems, especially in

large water bodies cannot be ignored. The weeds

need to be managed prudently to allow sustain-

able exploitation and exploration of mitigation

measures, to minimize potential negative impacts

often associated with their massive proliferation.

The spread of aquatic weeds in the tropics is a

matter of great concern and concerted mitigation

efforts are required to allow for continued use of

enormous aquatic resource potentials in the trop-

ics. This is critical as the majority of livelihoods

in the tropics depend on the water bodies and their

resources. In Lake Victoria, plants succession due

to introduction of invasive species, E. crassipes

has led to several ecological changes in the lake

with far-reaching socio-economic consequences

(Aloo, 2003).

Although aquatic weeds have caused serious

problems in many waterbodies, their positive ef-

fects cannot be ignored especially on the biodi-

versity of such systems. The weeds not only

provide shelter to many aquatic organisms but

they are also a source of food to some fish species

and other vertebrates and provide raw materials

for use in several livelihood activities (Dean,

1969; Bond & Roberts, 1978; Mitchell, 1974;

Oluoch, 1983). In Lake Victoria, for example, the

appearance of Eichhornia has led to an increase in

the catches of C. gariepinus, Haplochromis spp.,

P. aethiopicus, M. kannume and many hapl-

ochromine species (Njiru et al., 2000). Many of

these fish species were a favourite dish to local

communities in the pre-Nile perch Lake Victoria.

Given the magnitude of aquatic weeds infestation

in Lake Victoria, this paper recommends the fol-

lowing:

- a survey of the extent and nature of the aquatic

weeds infestation should be undertaken to establish

the magnitude of the problem in Kenya’s aquatic

systems including small water bodies;

- multidisciplinary research should be carried

out on aquatic weeds to understand their effects on

the aquatic systems and their potential benefits to

both humans and other organisms;

- there is need to investigate the relationship bet-

ween water hyacinth and the hippo grass in Lake

Victoria;

- control strategies should take into account the

potential effects of the method on the flora and

fauna found in the water body.

ACKNOWLEDGEMENTS

The authors wish to acknowledge Karatina Uni-

versity, Kenya Marine and Fisheries Research In-

480

P. Aloo, W. Ojwang, R. Omondi.J.M. Njiru & D. Oyugi

stitute, Moi University and Kenya Wildlife Service

Training Institute.

REFERENCES

Adeniji H.A., 1979. Framework document on special

problem of man-made lakes. S.I.L. Workshop on

African Limnology, UNEP Head-Quarters, Nairobi,

Kenya.

All African Press Service (AAPS), 1990. Harmful

hyacinth resists elimination: Science and Technology

Feature of the All Africa Press Service, January 1990,

Nairobi, Kenya.

Aloo P.A, 1988., Ecological studies of the Blackbass,

Microp terns salnioides (Lacepede) in Lake Naivasha,

Kenya. MSc. Thesis, University of Nairobi, Kenya.

Aloo P.A, 2003. Biological diversity of the Yala Swamp

lakes, with special emphasis on fish species compo-

sition in relation to changes in the Lake Victoria

Basin (Kenya): threats and conservation measures.

Biodiversity and Conservation, 12: 905-920.

Balirwa J.S., Chapman C.A., Chapman L.J., Cowx I.G.,

Geheb K., Kaufman L., Lowe-McConnell R.H.,

Seehausen O., Wanink J.H., Welcomme R.L. & Witte

F., 2003. Biodiversity and fishery sustainability in the

Lake Victoria Basin: An unexpected marriage?

Bioscience, 53: 703-715.

Bailey R.G., Churchfield S. & Pimm R., 1978. Obser-

vation on the zooplankton and littoral macroinverte-

brates of Nyumba Ya Mungu reservoir, Tanzania.

The Biological Journal of Linnean Society, 10: 93-107.

Blackburn R.D., Sutton, D.L. & Taylor, T., 1971. Bio-

logical control of aquatic weeds. Journal of Irrigation

Drainage, Division of American Society Civil En-

gineers, 97: 421-32.

Bond W.J. & Roberts M.G., 1978. The colonization of

Cabora Bassa, Mozambique, a new man-made lake,

by floating macrophytes. Hydrobiologia, 60: 243-

259.

Bootsma H.A. & Hecky R.E., 1993. Conservation of

the African Great Lakes: A Limnological Perspective.

Conservation Biology, 7: 644-656.

Bowmaker A.P., 1968. Preliminary observations on some

aspects of the biology of the Sinamwenda estuary,

Lake Kariba. Proceeding Trans. Rhodesia Science.

Association, 53: 3-8.

Bowmaker A.P., 1973. A hydrobiological study of the

Mwenda River and its mouth, Lake Kariba. PhD.

Thesis, University of Witwaters rand, 576 pp.

Boyd C.E., 1969. Production, mineral nutrient absorption

and biochemical assimilation by Justicia americana

and Alternanthera philoxeroides. Archiv fur Hydro-

biolologie, 66: 139-60.

Bugenyi F.W.B. & Van De Knaap M., 1998. Lake

Victoria: an example of ACP-EU Fisheries Research.

European Commission Fisheries Cooperative Bul-

letin, 10: 20-21.

Butcher R.W., 1933. Studies on the ecology of rivers: I.

On the distribution of macrophytic vegetation in

the rivers of Britain. Journal of Ecology, 21: 59-91.

Clark F., Beeby A. & Kirby P, 1987. A study of the

Macro-invertebrates of Lake Naivasha, Oloidien and

Sonachi, Kenya.In: Harper D.M. (Ed.) Studies on the

Lake Naivasha ecosystem 1982-1984. Final Report

to Kenya Government.

Chapman L.J., Chapman C.A., Ogutu-Ohwayo R., Chan-

dler M., Kaufman L. & Keiter A.E., 1996. Refugia

for endangered fishes from an introduced predator in

Lake Nabugabo, Uganda. Conservation Biology, 10:

554-561.

Chapman L.J., Chapman C.A., Nordlie F.G. & Rosen-

berger A.E., 2002. Physiological refugia: swamps,

hypoxia tolerance, and maintenance of fish diversity

in the Lake Victoria region. Comparative Biochem-

istry and Physiology, 133: 421-437.

Davies B.R., Hall A & Jackson P.B.N., 1975. Some

ecological aspects of the Cabora Bassa dam. Bio-

logical Conservation, 8: 189-201.

Davis R. & Hirji R., 2003. Water Resources and Envi-

ronment, technical note C.2. Environmental Flows:

Case Studies. Management of aquatic plants. Techni-

cal note G4. The World Bank Publication, Washing-

ton, 30 pp.

Dazo B.C., Hairstone N.G. & Dadwood I.K., 1966. The

ecology of Bulinus truncatus and Biomphlaria alexan-

drina and its implication for the control of bilharzia-

sis in the Egypt 49 project Area: Bulletin. Wld Hlth

Org., 35: 339-359.

Dean J.L., 1969. Biology of the Cray fish Orconectes

causeyi and its use for control of aquatic weeds in

trout lakes. Tech, paper, 24. U.S. Fish and Wildlife

Service, Washington, 15 pp.

Denny P, 1972. Lakes of South Western Uganda. I. Phys-

ical and chemical studies on Lake Bunyonyi. Fresh

Water Biology, 2: 143-158.

Denny P., 2000. Eichhornia crassipes (Mart) Solms -

Laub - ecological implications and impacts on the

fish of Lake Victoria: possible research strategies.

Paper presented during Lake Victoria Fisheries Work-

shop. Entebe, Uganda 1st -4th June, 2000.

Garry H., Waage J. & Phiri G., 1997. The Water hyacinth

problem in Tropical Africa. Report prepared for the

first meeting of an International Water Hyacinth

Consortium held at the World Bank, Washington,

18th- 19th, March, 1997.

Greenwood P.H., 1974. The Cichlid fishes of Lake Vic-

toria, East Africa: the biology and evolution of a

Impacts of invasive aquatic weeds on the biodiversity of some tropical water bodies with special reference to Lake Victoria 481

species flock. Bulletin of the British Museum Natural

History (Zoology) Supplement, 6: 1-134.

Hall J. & Pople W., 1968. Recent vegetational changes

in the lower Volta River. Ghana Journal of Science,

8: 24-29.

Harley K.L.S., 1990. The role of biological control in

the management of water hyacinth ( Eichhornia cras-

sipes ). Biocontrol News and Information, 11: 11-12.

Harper D.M., 2010. Lake Naivasha: Can anybody man-

age nature’s uncertainty? Paper presented at the ARK

II International conference held at the Kenya Wildlife

Training Institute, Naivasha, Kenya. 16th to 19th No-

vember, 2010.

Johnson T.C., Cohen A. J., Talbolt M.R., Kelts K., Lehman

J.T., Lingstin D.A., Odada E.O., Tambalan A.F.,

McGill T. & Tiecellin J., 1990. Paleoclimatology of

the African Rift Valley Lakes. Gordon and Breach,

Amsterdam, 640 pp.

Kaufman L., 1992. Catastrophic change in species-rich

freshwater ecosystems: the lessons of Lake Victoria.

Bioscience, 42: 846-858.

MailuA.M., 1999. Water hyacinth: distribution, impact

and control Measures. KARI Technical Note Series

No. 3, Nairobi, Kenya, 7 pp.

Maltby E., 1986. Waterlogged Wealth. International In-

stitute for Environment and Development, London,

200 pp.

McLachlan A.J., 1969. The effect of aquatic macrophytes

on the variety and abundance of benthic fauna in a

newly created lake in the tropics (Lake Kariba).

Archiv fur Hydrobiologie, 66: 222-231.

Mitchell D.S., 1974. Aquatic vegetation and its use and

control. Unesco, Paris. Benton Jnr, pp. 99-106.

Mitchell D.S., 1976. The marginal fish fauna of Lake

Kariba. Kariba studies, 8: 109-162.

Njiru M., Othina A., Tweddle D. & Cowx I.G., 2000.

Lake Victoria invasion by water hyacinth: a blessing

for Lake Victoria Fisheries. In : Book of Abstracts,

Lake Victoria 2000: A new Beginning (International

Conference). LVFO, Jinja, Uganda, pp. 80-81.

Njuguna S.G., 1992. Floating aquatic weeds in Kenya.

In: Crafter S.A, Njuguna S.G. & Howard G.W.

(Eds). Wetlands of Kenya. Proceedings of the

KWWG Seminar on Wetlands of Kenya, National

Museums of Kenya, Nairobi, Kenya, 3-5 July 1991.

183 pp.

Ogari J., 2000. Impact of exotic fish species and invasive

water weeds such as the water hyacinth on Lake

Victoria fisheries. Paper presented during “Lake

Victoria 2000” conference, Jinja, Uganda. 16th -19th

May, 2000.

Ojwang W.O., Kaufman L., Asila A.A., Agembe S. &

Michener B., 2004. Isotopic Evidence of Functional

Overlap amongst the Resilient Pelagic Fishes of

Lake Victoria, Kenya. Hydrobiologia, 529: 27-35.

Ojwang W.O., Kaufman L., Soule E. & Asila A. A.,

2007. Evidence of stenotopy and anthropogenic in-

fluence on carbon source for tow major riverine

fishes of Lake Victoria watershed. Journal of Fish

Biology, 70: 1430-1446.

Oluoch A., 1983. A study of some aspects of the distri-

bution, biology, ecology and fishery of Crayfish Pro-

cambarus clarkii (Girard) population in Lake

Naivasha. M. Sc. thesis, University of Nairobi,

Kenya.

Penfound W.T. & Earle T.T., 1948. The biology of the

water hyacinth. Ecological Monographs, 18: 447-

472.

Petr T., 1968. Population changes in aquatic invertebrates

living on two water plants in a tropical man-made

lake. Hydrobiologia, 32: 449-85.

Petr T., 1970. Macroinvertebrates of flooded trees in the

man-made Volta Lake (Ghana) with special reference

to the mayfly Povilla adusta Navas. Hydrobiologia,

36: 373-398.

Prescott G.W., 1969. How to know aquatic plants. C.

Brown Publishers, Dubuque, 171 pp.

Scholz C.A., Rosendahl B.R., Versfelt J.W. & Rach N.,

1990. Results of high-resolution echo-sounding of

Lake Victoria. Journal of African Earth Sciences, 1 1 :

25-32.

Sculthorpe C.D., 1976. The Biology of Aquatic Vascular

Plants. Edward Arnold, London, 610 pp.

Russell R. J., 1942. Flotant. Geographical Review, 32:

74-98.

Symmoens J.J., Burgis M. & Gaudet J.J., 1981. The ecol-

ogy and utilization of African Inland waters. SIL

Workshop. UNEP Proceedings. Girgiri, Kenya,

191 pp.

Triest L., 1993. Problems created by excessive macro-

phytic growth in tropical freshwater ecosystems and

strategies for control. In: Tilzer M.M. & Khandker M.

(Eds). Hypertrophic and polluted Freshwater ecosys-

tems: Ecological Bases for Water Resource Manage-

ment Proc. Int. Symp. Limnol., 25-28 November,

1991. Department of Botany, University of Dhaka,

Bangladesh, pp. 137-153.

Twongo T., 1991. Water hyacinth on Lakes Kyoga and

Kwania. Special report No. 3 November 1991. In:

Thompson K. (Ed). The water hyacinth in Uganda.

Ecology, Distribution, Problems and strategies for

control. Work-shop: 22-23 October 1991. FAO

TCP/UGA/9 153/A.

Twongo T., 1996. Growing impact of water hyacinth on

near shore environments on Lakes Victoria and

Kyoga (East Africa). In: Johnson T.C. & Odada E.O.

(Eds). The Limnology, Climatology and Paleclima-

tology of the East African Lakes. Gordon and Breach,

Amsterdam, pp. 633-642.

482

P. Aloo, W. Ojwang, R. Omondi.J.M. Njiru & D. Oyugi

Watson E. & Parker I.S.C., 1970. The ecology of Lake

Naivasha, Kenya. The identification and description

of some important components for a model. African

Scientists, 2.

Wawire N.W. & Ochiel R.S., 2001. Manuscript presented

at the 1st LVEMP National Scientific Conference on

Lake Victoria. Tom Mboya Labour College, 24th-

28th September 2001, Kisumu, Kenya.

Welcomme R.L., 1972. A brief on the flood plain fishery

of Africa. African Journal of Tropical Hydrobiology

and fisheries, 1: 67-76.

Witte E, Goldschmidt T., Goudswaard PC., Ligtvoet W.,

van Oijen M.J.P. & Wannink J.H., 1992. Species

extinction and concomitant ecological changes in

Lake Victoria. Netherland Journal of Zoology, 42:

214-232.

Biodiversity Journal, 2013, 4 (4): 483-500

New faunistic data on Trochoidea ( Trochoidea ) caroni (De

shayes, 1832) (Gastropoda Pulmonata Hygromiidae)

Nicola Maio 1 *, Agnese Petraccioli 1 , Paolo Crovato 2 , Nabil Amor 3 & Gaetano Odierna 1

'Dipartimento di Biologia, Complesso Universitario di Monte S. Angelo, Universita degli Studi di Napoli Federico II, Edificio 7,

Via Cinthia, 21 - 80126 Naples, Italy

2 Via S. Liborio 1, - 80134 Naples, Italy

3 Unite de Recherche: Genetique, Biodiversite et Valorisation des Bioressources, UR/09-30. Institut Superieur de Biotechnologie

de Monastir, Monastir 5000, Tunisie

’Corresponding author: nicomaio@unina.it

ABSTRACT Trochoidea ( Trochoidea ) caroni (Deshayes, 1832), with T. (T.) elegans (Gmelin, 1791) and

T. (T.) trochlea (Pfeiffer, 1846), belong to a group of species (“ elegans group”), which,

although considered valid by most authors, show a not completely defined systematic position.

In this work, we report four new records (Terracina and Spigno Satumia for Latium, Sorrento

for Campania and Castagneto Carducci for Tuscany) and confirm two historical records

(Ischia Island for Campania and Manfredonia for Apulia) of T. caroni from Italy, and all

published and new faunistic data about the presence of this species in the Sicilian Islands,

Capri Island and Maltese Islands. Moreover, we report some historical data from the Western

Mediterranean Sea: two bibliographical records from the Balearic Islands, not verified, and

three records from North Africa (Tunis in Tunisia, Annaba in Algeria and an unmarked locality

in Morocco). We also report the up to now historical faunistic data of T. trochlea, a not well

known species with an undefined status, and of some populations of T. elegans with raised

whorl. It is not yet possible to draw a certain conclusion on the particular biogeography of

these taxa, but the new data presented in this work, enhance the current knowledge on the

distribution of T. caroni which would be larger than previously known.

KEY WORDS Trochoidea', land gastropods; Italy; Mediterranean; faunistics.

Received 19.09.2013; accepted 13.11.2013; printed 30.12.2013

INTRODUCTION

Trochoidea ( Trochoidea ) caroni (Deshayes,

1832) (Gastropoda Pulmonata Hygromiidae) is

morphologically characterised by a trochiform,

pyramidal, dextrorse shell with 8-10 well raised

whorls having a sharp marginal keel, finely ribbed

external surface and small umbilicus. The height-

to-diameter ratio (h/d) of the spire ranges from 1.00

to 1.30, in some cases as low as 0.88: see Sacchi’s

form (Sacchi, 1955a).

Anatomically, T. (72) caroni is characterised

(Giusti et al., 1995) by penial papilla elliptical or reni-

form in transverse section with long C-shaped

groove on one side; four pleats defining two lozenges,

each surrounding the opening of an appendicula into

the vagina; atrial appendix containing crest-like

structure; long penial flagellum. T. (T.) caroni lives

in garigue and grassy meadow environments, under

stones and detritus, sometimes also on calcareous

rocks or at their base. It is currently reported in Italy

as a living species in Central Western Sicily and on

484

N. Maio, A. Petraccioli, P. Crovato, N. Amor & G. Odierna

Capri (Alzona, 1971; Cossignani & Cossignani,

1995; Manganelli et al., 1995). The aim of the

present paper is to provide new data on the presence

of T. caroni in certain regions of Italy and to collect

all data in the literature and available from museum

collections on its present and past distribution.

T. caroni has close anatomical affinity (“ elegans

group”) with T. (I) elegans (Gmelin, 1791), T. (T.)

spratti (Pfeiffer, 1 846), a species endemic to the is-

land of Malta, and T. (T.) cumiae (Calcara, 1847),

endemic to the island of Lampedusa (Giusti et al.,

1995). In particular, the systematic and biogeographi-

cal positions of T. caroni , T. elegans and T. trochlea

(Pfeiffer, 1846), the latter a little-known North-

African species, are still unclear.

T. trochlea was described as a new species by

Pfeiffer (1846) but the original types are not known.

According to the original description (Pfeiffer,

1 846), this species is morphologically characterised

by a conical shell with eight keeled, crenulated

whorls, having diameter about the same as height,

or slightly greater (h/d = 1.04 in Pfeiffer (1846),

1.05 in Bourguignat (1864) with a variety B minor

reaching 1.14). In the continuation of Rossmassler’s

“Iconographie”, Kobelt (1877) also reports similar

measurements for Helix trochlea Pfeiffer [“Diam.

et alt. 7-8 Mm”, h/d= 1.14]. Bourguignat (1864)

adds other distinctive characters with respect to T.

elegans , such as more accentuated striations, small

umbilicus, crenulated keel and more imbricated

whorls that overlap each other. Its distribution is

thought to include Algeria and Northern Tunisia

(Pfeiffer, 1853; Morelet, 1853; Bourguignat, 1864,

1868; Kobelt, 1877; De Saint-Simon, 1882; Le-

toumeux & Bourguignat, 1887; Monterosato, 1893;

Sacchi, 1955a).

Morphologically, T. elegans is characterised by

a conical shell with 6-7 whorls of variable height

and diameter greater than height. It is distinguished

from T. caroni by a different height-to-diameter

ratio (h/d) of the spire, in the range 0.82-0.62

(Sacchi, 1954) or less (“ scitula ” form); external sur-

face with fine regular growth ridges; umbilicus

small. Its distribution is European- Western Mediter-

ranean (Southern France, Spain, Greece, Tunisia,

Algeria and perhaps Morocco; (Pallary, 1904)) with

isolated populations in Belgium, Northern France,

Sardinia and Corsica; introduced into Great Britain

from the early 1800s (Ferussac, 1821), into South

Carolina (USA) from before 1875 (Mazyclc, 1876)

and perhaps the Balearic Islands (Gargominy,

2011). In Italy it is present in Figuria, Tuscany,

Umbria (Misuri, 1907), Fatium and doubtful in

Sicily. It is a xerophilous species with wide ecolog-

ical valence colonising coastal, ruderal and culti-

vated calcareous environments. The biometric

character hitherto used therefore does not discrimi-

nate the three species with absolute certainty and

various anatomical similarities also exist. Since

there are currently no other diagnostic criteria, it

was decided to list the data available in the literature

and from museum collections, exactly as reported

by the original authors or collectors, also for T.

trochlea and T. elegans.

MATERIALS AND METHODS

In order to obtain an exhaustive picture of the

distribution of T. caroni in Italy, field surveys were

conducted and their results were compared with his-

torical data obtained from other sources. This re-

search was carried out in addition to the

data-gathering in the literature and by studying

malacological collections in different Italian and

foreign museums and private collections. The field

studies were conducted between September 2002

and September 2006 on Capri and in Sicily and in

the period 2006-2009 in Fatium. The species iden-

tification was based on Giusti et al. (1995); the tax-

onomical and nomenclatural arrangement follows

Manganelli et al. (1995) and Bank (2011).

RESULTS

Trochoidea ( Trochoidea ) caroni (Deshayes, 1832)

Original description. Deshayes, 1832, species

n. 142, pag. 262, locus typicus “Sicile”: “ Helice de

Caron. Helix Caroni. Nob. H. testa conica, elon-

gato-pyramidata, trochiformi, longitudinaliter

striata, striis confertis, sublamellosis ; anfractibus

numerosis, basi carinatis; carina squamosa; aper-

tura depressa, quadrangulari, simplici; labro tenui.

Diametre de la base 9 millim., longueur 11 millim”.

[h/d = 1.22

Ferussac (1821) reports the species for the first

time as: “ Helix elata Faure Biguet [J. P. Faure-Biguet]

espece n. 304, p. 46, “Hab. F’ile de Capree”, without

New faunistic data on Trochoidea (Trochoidea) caroni (Deshayes, 1832) (Gastropoda Pulmonata Hygromiidae) 485

cU.-fW

VlEUX

PftiEROe , * st

det. : towi . CftFSPN

\tbn . tocAci-

£.2, : 2.#2-

Museum Nil HSsL W*t- Pan*

!NVERT£BR£S MARINS

ET MALACOLOGY

Figure 1 . Syntype n° Moll 23356 from Palermo collected by “M. Caron” (collection Mollusques, Museum National d'Histoire

Naturelle, Paris), apertural view. Figure 2. Idem, umbilical view. Figure 3. Idem, apical view. Figure 4. Idem, original labels.

Photos by P. Maestrati, Museum National d'Histoire Naturelle, Paris.

any description. Thus the name is now considered

a nomen nudum.

Deshayes (1832) [according to Sherborn (1924:

p. 1110) the correct year of publication is 1832]

published the first valid description of the species,

assigning the name “ Helix Caroni ”. The original

type series is still kept in the National Museum of

Natural History in Paris: a syntype from Palermo

collected by “M. Caron”, to whom the species is

dedicated (syntype: “1 lot de 1 specimen, Typothe-

que Leg. Caron, n° Moll 23356”) and other 8 syn-

types from “Sicilia” (syntypes: “1 lot de 8

specimens, Typotheque, n° Moll 23357”) (Maio,

pers. obs.; Figs. 1-4).

Rossmassler (1837: pp. 23, 24, fig. 344), who

gives Faure-Biguefs description of Helix elata,

considers the species a synonym of Helix caroni.

Examined material. Latium. The first re-

port of T. caroni in Latium is by Statuti (1882) who

mentions finding a single shell on the beach be-

tween Porto di Badino and Torre Olevola, a few km

from Terracina in Latina Province. The author sus-

tains that the specimen reached the beach with flot-

sam from nearby rivers of the Pontine Plain. The

studies conducted in this area produced 12 shells

(three juveniles) from the Municipality of Spigno

Saturnia (Latina) in spring and summer 2006 (Figs.

5-7). The specimens have an h/d ratio greater than

1.40. Their good state of conservation suggests that

they come from a population still present in the

area. However, this could not be confirmed as no

live specimens were found (Petraccioli et al., 2011).

A live population of T. caroni was identified in

1986 (Hallgass, 2007; com. pers.; Figs. 8-10,

25) at San Silviano in the Ausoni Mountains, near

Terracina (Latina), at an altitude of 200 m. Speci-

mens have an h/d ratio between 1.10 and 1.30. The

species is indicated generically in Latium by

Welter-Schultes (2012).

Campania. The first historical report of T. caroni

in Campania dates back to Ferussac (1821) who re-

ports data of Faure-Biguet for Capri. The species is

referred to as “ Helix elata Faure Biguet”. Ross-

massler (1837) also reports it from Capri as Faure-

Biguefs Helix elata. Achille Costa (1840, sub

Corocolla (sic!) elata) claims to have found it under

stones on the road to S. Giacomo (Capri, Naples)

and lists it among rare species. Philippi (1836,

1844, sub Helix elata Faure Biguet) confirms the

presence of the species on Capri. Monterosato

(1893) reports Xerophila caroni Desh. ( =turrita

Ph.) from “isola di Capri (Philippi)”.

486

N. Maio, A. Petraccioli, P. Crovato, N. Amor & G. Odierna

In his writings, Bellini (1899, 1900, 1901, 1910,

1915, 1920, 1922, 1924, 1926) reports it with var-

ious names: Helix (Xerophila) turrita Phil., Helix

(Tropidocochlis) turrita Phil., Tropidocochlis elatus

Faure Biguet, Tropidocochlis turritus Phil., H. ( Tropi-

docochlis ) data Faure Biguet, Tropidocochlis Ca-

roni Deshayes, H. ( Xerophila ) data Fer. var.

caprensis and claims it to be a rare species on Capri,

though locally abundant in litter at Castiglione, Arco

Naturale, Via Krapp and Pizzolungo. Bellini (1901)

describes a hypsometric zone of Capri between

Valletta (120 m) and Mt. S. Michele (245 m), which

he calls “zona di H. data e H. surrentina , \ He dis-

tinguishes an eastern subzone where H. data pre-

dominates. Bellini (1899; 1900) also describes a

variety caprensis (Bellini, 1899: pp. 4-5), “raranella

parte occidentale dell’isola, nelle zolle incolte al di

sotto del Castello Barbarossa” [rare in the western

part of the island, in uncultivated turf and below

Castello Barbarossa]. Original specimens collected

by Bellini are kept in the malacological collection

of the Zoological Museum of Naples University

Federico II and in the museum of the Centro

Caprense “Ignazio Cerio” on Capri (Maio, pers.

obs.). Sacchi (1953) reports its presence on Capri,

on Mt. Tuoro and Mt. Solaro, in sparse garigues with

rosemary, as: “endemita del gruppo Helicella ( Tro -

choidea) elegans Gm., a spira elevata e a forma tur-

ricolata, che corrisponde alle forme descritte dal

Bellini (1915) come H. caroni Desh. e H. data

Faure”. Subsequently (Sacchi, 1955a, 1955b, 1956a),

he states that “ Helicella ( Trochoidea ) caroni Desh.

and H. data Faure” are indistinguishable on Capri.

Three specimens found in Via Tragara in August

1979 are kept in the S. Duraccio collection (ex coll.

Pirozzi no. 69). During our study on Capri, we

found T. caroni in 10 stations ranging in altitude

from about 25 m to 200 m, but only in two of these

stations did we find live specimens (Table 1, Figs.

11-13, 27). They were found at the base of the

trunks or near superficial roots of mainly legumi-

nous plants typical of Mediterranean maquis,

growing on calcareous rock in the littoral zone (Pe-

traccioli et al., 2005a, 2005b, 2006). The h/d ratio

was about 1.05.

There is an interesting report of fossils of this

species from Capri. Segre (1959) reports it as Helix

caronii (sic!) Desh. (F. Settepassi det.) found in cal-

carenite of Villa Jovis (Monte di Lauro, Capri), a

well-cemented, medium to coarse grained calcare-

ous sandstone from the Lower Pleistocene (Barat-

tolo et al., 1992; Petraccioli et al., 2007).

Sacchi (1953) states that it is possible to find

“una piccola forma, (...) mostrante affinita con He-

licella elegans e forme simili” at the base of coastal

cliffs between Amalfi and Positano (Salerno).

Specimens

Sampling stations

Date

Habitats

Alt. m

UTM

Living

Trackfrom via Mesola to Fortino

Mesola (Anacapri)

17.04.2004

Dry stone walls,

mediterranean maquis

37-150

3289

Living

Fortino Mesola (Anacapri)

17.04.2004

Mediterranean maquis

3289

Shells

“Belvedere o Piazzetta delle Noci”

(Capri)

15.04.2004

Pinewood, dry stone walls.

In humus

194

3789

Shells

Scala Fenicia, quarto tratto, Capri

06.04.2003

On calcareous rock

67-100

3489

Shells

Via Krupp (Capri)

18.07.2003,

01.11.2003

On calcareous rock

25-125

3588

Shells

Via Arco Naturale (Capri)

18.07.2003

In litter

3788

Shells

Via Pizzolungo (Capri)

16.04.2004

In litter

50-100

3688

Shells

Via Cesina (Capri)

16.11.2003

Dry stone walls. In debris

150-200

3689

Shells

Giardini Augusto (Capri)

24.01.2004

Gardens. In debris

75-96

3688

Shells

Monte Tamborio (Capri)

07.11.2003

On calcareous rock. In debris

200

3689

Table 1. Faunistic data of Trochoidea ( Trochoidea ) caroni from Capri Island.

New faunistic data on Trochoidea (Trochoidea) caroni (Deshayes, 1832) (Gastropoda Pulmonata Hygromiidae) 487

Finally, two shells of juvenile specimens of T. ca-

roni were found in humus at Sorrento (Naples)

(March 1984, G. Fasulo legit) (Figs. 14-16). The

site unfortunately no longer exists, due to road-

works that destroyed the original habitat. Their h/d

ratio was about 0.95.

Delle Chiaje (1841) reports Helix elegans Drap.

from Ischia Island. The plate was previously pub-

lished by the author in the atlas “Memorie sulla Sto-

ria e Notomia degli Animali senza vertebre del

Regno di Napoli” dating between 1822 and 1830

(Delle Chiaje, 1830) (Fig. 23). The figure clearly

indicates that the species in question is actually T.

caroni, which at the time had not yet been de-

scribed. The synonymy is also confirmed by Benoit

(1862, 1875) and Statuti (1882).

Apulia. A specimen of T. caroni found at Man-

fredonia (Foggia Province), collected by W. Kobelt

in 1878, is kept in the L. Pfeiffer collection at the

Senckenberg Forschungsinstitut und Naturmuseum

in Frankfurt am Main, Germany (R. Janssen, pers.

com.; see Table 3). Its h/d ratio is 7. 0/6. 6 = 1.06.

Tuscany. A specimen found of T. caroni at

Castagneto Carducci (Livorno Province) in 1983 is

kept in the S. Duraccio collection (ex coll. Pirozzi

no. 69F, G. D’Anna legit). Its h/d ratio is 8. 5/7.0 =

1 . 21 .

Sicily. The first historical report of T. caroni

from Sicily dates back to Femssac (1821), who cites

W.E. Leach’s report of “Helix elata Faure Biguet”.

Subsequently, Deshayes (1832) described Helix Ca-

roni on a specimen from Sicily (Figs. 1-4).

Philippi (1836: pp. 137-138, pi. 8, fig. 17) de-

scribes it in his first volume as a new species with

the name of Carocolla turrita on specimens from

Palermo (Fig. 24). He also reports Carocolla elata

Faure-Biguet (pi. 8, fig. 16) from Palermo, Sciacca

(Agrigento Province) and from Pachino (Siracusa

Province). In his second volume (Philippi, 1844),

he reports Helix turrita Ph. as a synonym of H. ca-

roni Desh. from Palermo. Rossmassler (1837) also

cites the same works.

Pirajno (1840, sub Helix elata), reports it

“presso Scillato [Palermo] alia Portella di mare”.

Describing the Trapani area, Power (1842, as

Caracolla elata ) reports T. caroni “sul monte S.

Giuliano sotto erbe e pietre” and near the walls of

the Gastello di Erice (Sparacio, 2012). Aradas &

Maggiore (1840) reported T. caroni as H. elata Fer.

(Elice alta) in their "Catalogo delle Conchiglie

viventi e fossili", as being “frequente in Palermo e

Sciacca, rarissima nei nostri dintorni. Trovasi pure

alle Madonie presso Scillato alia portella di Mare”,

and as H. turrita Phil. (Elice torriciuolata) as being

“comune in Palermo, e alle falde di Monte Pelle-

grino, abita in terreni aridi”.

Calcara (1845) reported T. caroni sub Helix

elata in the Palermo area, on the banks of the River

Oreto and as Helix caroni from "Malaspina, alle

Croci, falde di Monte Pellegrino, sponde dell’ Oreto,

a Sferracavallo e Carini".

Benoit (1862, p. 206, pi. 5, fig. 18) describes a

local form of Helix cfr. caroni with a less accentu-

ated conical form and rounded apex, as Helix Se-

quentiana. He found it in the Eastern Madonie

Mountains and near Cefalu (Palermo). He also de-

scribes a variety dilatata of H. elata Faure-Biguet

that he found near Palermo and on Mt. S. Giuliano

(or Mt. Erice) near Trapani. He describes H. caroni

Deshayes on Mt. Pellegrino, on the banks of the

River Oreto, “alle Croci”, at Carini, near Palermo,

as well as at Calatafimi, near the Temple of Segesta,

Trapani (Benoit, 1875, 1881). Some specimens of

the Mina Palumbo collection bear Fiume Oreto and

Palermo as collection sites (Liberto et al., 2010).

De Saint-Simon (1882) reports Helix caroni

generically from Sicily. Westerlund (1889) reports

it from Palermo as Helix caroni, from the Madonie

Mountains as Helix sequentiana, and generically

from Sicily as Helix turrita.

Monterosato (1893) reports Xerophila caroni

Desh. (= H. Caroni, Ben. - 111. Sist. 1857, t. V. f.

14) from Favignana and the Egadi islands (Benoit).

He also states that the species of this group are: “a)

X. Caroni, Desh. (= turrita , Ph.) - Prov. di Palermo,

Nicosia, Siracusa; b) H. exacta, Monts, mss. =H.

Caroni, Ben. - 111. Sist. T. V, f. 15 (Calatafimi); c)

H. elata, Faure-Biguet et var. dilatata, Ben. - Prov.

di Trapani; d) =H. Seguenziana, Ben. H. Ipyramis,

Ph. - Calatafimi, Sciacca ed altre localita Siciliane”.

Monterosato (1894) later cites H. (Xerocochlea)

Caroni Desh. “comune e caratteristica delle parti

basse del Monte Pellegrino e dei suoi dintorni sino

a S. Polo, in tutta la parte che lo circonda”.

De Gregorio (1895, sub Helix elata Faure Biguet)

reports T. caroni from “Fiume Anapo in Siracusa

e (...) Carini”; in the same paper (1895, as Helix

elata Var. dilatata Ben.) also from Monte San

Giuliano.

488

N. Maio, A. Petraccioli, P. Crovato, N. Amor & G. Odierna

Specimens

Localities (Municipality)

Province

Date

Collection

Collector

Shell

Monte Pellegrino

Palermo

1949

“V.Wiese” Haus der Natur,

Cismar (Germany)

Brandt

17 es.

via Sampolo (Palermo)

Palermo

1910-60

“C. Alzona” MCSNM

Monterosato

3 es.

San Vito (Palermo)

Palermo

1910-60

“C. Alzona” MCSNM

Monterosato

4 es.

Monreale

Palermo

1910-60

“C. Alzona” MCSNM

Monterosato

6 es.

Salapamta

Trapani

1910-60

“C. Alzona” MCSNM

2 es.

Erice

Trapani

Ott. 1951

“C. Alzona” MCSNM

C. Sacchi

2 es.

Agrigento

1910-60

“C. Alzona” MCSNM

11 es.

Porto Empedocle

Agrigento

Ott. 1951

“C. Alzona” MCSNM

C. Sacchi

5 es.

Siracusa

1910-60

“C. Alzona” MCSNM

Shells

Sferracavallo (Palermo)

Palermo

15.09.1978

Coll. I. Sparacio

I. Sparacio

Living

Bosco della Ficuzza (Corleone)

600 m s.l.m.

Palermo

21.11.1979

Coll. I. Sparacio

I. Sparacio

Living

Capaci (Capaci)

Palermo

17.02.1980

Coll. I. Sparacio

I. Sparacio

Living

Foci Fiume Pollina (Pollina)

Palermo

24.05.1981

Coll. I. Sparacio

I. Sparacio

Living

Bolognetta

Palermo

19.06.1987

Coll. I. Sparacio

I. Sparacio

Living

Isola delle Femmine

Palermo

15.08.1990

Coll. I. Sparacio

I. Sparacio

Living

Golfo di Cofano, Castelluzzo

(San Vito Lo Capo)

Trapani

14.10.1984

Coll. I. Sparacio

I. Sparacio

Living

Golfo di Cofano, fascia costiera

(S. Vito Lo Capo)

Trapani

06.10.1987

Coll. M. Cuomo

M. Cuomo

Living

Monte Cofano, Castelluzzo

(San Vito Lo Capo)

Trapani

04.1991

Coll. I. Sparacio

I. Sparacio

Living

Calatafimini (Calatafimi Segesta)

Trapani

05.12.1984

Coll. I. Sparacio

I. Sparacio

Shells

Favignana, Cala Rossa

Trapani

25.08.1990

Coll. I. Sparacio

I. Sparacio

Living

localita Lido Cornino

(Custonaci)

Trapani

Cossignani &

Cossignani, 1995

Living

Monte Erice (Erice)

Trapani

Giusti et al., 1995

Living

Favignana (Is. Egadi)

Trapani

G. Manganelli

Giusti et al., 1995

Living

San Vito Lo Capo

Trapani

20.08.2006

Coll. I. Sparacio

I. Sparacio

Living

Favignana (Is. Egadi)

Trapani

Reitano, 2006b

Living

Zone aride costiere per Cala

Rossa (Terrasini)

Palermo

2006

Reitano, 2006a

Living

Monti delle Madonie

Palermo

2006

Reitano, 2006b

Living

Dintomi di Grotta Conza,

Sferracavallo

Palermo

2012

Reitano et al.,

2012

Shells

North side of the city

Agrigento

10.09.2008

Coll. W. Renda

W. Renda

Living

Erice

Trapani

26.08.2008

Coll. W. Renda

W. Renda

Living

Isola delle Femmine

Palermo

16.10.2008

Coll. W. Renda

W. Renda

Living

Capaci

24.09.2010

I. Sparacio

Table 2. Faunistic data of Trochoidea ( Trochoidea ) caroni from Sicily.

New faunistic data on Trochoidea (Trochoidea) caroni (Deshayes, 1832) (Gastropoda Pulmonata Hygromiidae) 489

Many specimens (more than 20) of T. caroni

from Sicily are kept in the malacological collection

of the Zoological Museum of Naples. These speci-

mens, reported with various names such as: Helix

caroni , H. ( Turricula ) turrita, H. (T.) caroni. Helix

(T.) exact a, date to the second half of the nineteenth

century and were acquired under the direction of

Achille Costa; most come from localities in Palermo

Province or from near the city (exact toponyms are

not indicated); some were donated by the eminent

Sicilian malacologist Tommaso De Maria Allery di

Monterosato (Maio, pers. obs.).

Bellini (1915) reports the species as Tropido-

cochlis Caroni Deshayes “nei dintomi di Palermo,

Nicosia [Enna] e Siracusa”.

In October 1953, Sacchi (1955b, 1956a, 1956b)

reported it as Helicella ( Trochoidea ) caroni-elata

from Sicily, island of Mozia (Isola dello Stagnone,

Marsala, Trapani) and Capo Gallo (Palermo), and

in November 1953 as Helicella ( Trochoidea ) caroni

on silicicolous substrates at Mistretta (Messina

Province) and on Isola delle Feminine (Palermo),

stating that the latter form differed from the former

“per la conchiglia molto alta e turricolata”. Sacchi

(1955b, 1956b) also reports finding fossil speci-

mens of H. elata on Levanzo, Favignana and Ma-

rettimo (Egadi Islands, Trapani) in pink sandstone.

Some specimens from Porto Empedocle (Agri-

gento) are kept in the Fobbecke Museum in Diis-

seldorf (Maio, pers. obs.). They were collected by

J. Boscheinen in 1974 and determined as Tro-

choidea elata (Ferussac, 1819) by T. Meijer in

1975. Specimens of this species are reported from

different sites in Trapani Province, Palermo and Agri-

gento between 1949 and 2010 (Tab. 2). Fiorentino

et al. (2004) reports it from the Egadi Islands.

Finally, the present authors found various colonies

of T. caroni at Cala Rossa (Terrasini, Palermo)

(Figs. 17-19, 26, 28) in November 2004, at Gi-

acalone (Palermo) in October 1995 and at S. Vito Fo

Capo and Macari (Trapani Province) in June 2006.

The h/d ratio ranged from 1.14 to more than 1.20.

Malta. The species was first documented as a

fossil from the Quaternary of Malta by Trechmann

(1938, sub Xeroclivia sp.). Four specimens from

Marfa Point in the North of the island, determined by

Trechmann, are kept in the Department of Paleon-

tology of the British Natural Histoiy Museum of Fon-

don (Sacchi, 1955b). Giusti et al. (1995) found many

juvenile fossil specimens and shell fragments similar

to Sicilian populations of T. caroni in deposits at

Cirkewwa dating to the Fower and Upper Quaternary

(Hunt, 1997; Kolouch, 2003). Their h/d ratio ranged

from 1.24 to more than 1.29 (Giusti et al., 1995).

Recent specimens of T. caroni were reported

from the Maltese Islands in the nineteenth century:

Gulia (1874) and Feilden (1879) reported speci-

mens that Giusti et al. (1995) assume to have been

stranded transported by the sea. A specimen from

Malta, found in the first half of the nineteenth cen-

tury, is kept in the Philipp P. Carpenter collection

of the Field Museum of Natural History in Chicago

(Maio, pers. obs.). Originally labelled Iberus trochlea

Pfr., it was later determined by K. H. Beckmann as

T. caroni. Four specimens (two recent without date

and two Quaternary) from Malta, determined as

Trochoidea cfr. caroni, are kept in the Paul Sammut

collection in Rabat, donated to Heritage Malta,

the cultural agency of the Government of Malta

(Various authors, 2007).

North Africa. Monterosato (1893) reports Xe-

rophila caroni Desh. ( =turrita Ph.) from “Tunisi

(auct. [Monterosato?])” and H. elata, Faure-Biguet

et var. dilatata, Ben. from “Bona (Kob.)”.

Bellini (1915) reports the species as Tropidoco-

chlis Caroni Deshayes from Tunis.

Four specimens from Annaba (= Bona, Algeria)

attributed to T. caroni are kept in the Fobbecke Mu-

seum in Diisseldorf (Germany). They were collected

by Carl Emil Fischke in 1868 and determined by

him as Trochoidea elata (Ferussac, 1819) (Figs. 20-

22). Their h/d ratio ranges from 0.97 to 1.12. Two

specimens of the species from Morocco, dating to

the first half of the twentieth century, are kept in the

Walter F. Webb collection of the Field Museum of

Natural History in Chicago (USA). Originally la-

belled “ Helicella sequentiana (Benoit)”, they were

determined as T. caroni by K. H. Beckmann.

All these specimens were examined directly by

one of the authors (Maio, pers. obs.) and despite the

shell variations also present in North African popu-

lations of this group (see Bourguignat, 1864: vol. 1,

T. 32, f. 4-22), seem to belong to T. caroni.

Balearic Islands (Spain). Bullen (1910) re-

ports fossils of “Trochula elata (Faure-Biguet)” in

two different Holocene deposits on the island of

Majorca (Balearic Islands), namely at Porto Pi and

Alcudia. Sacchi (1954, 1955a) mentions an old re-

port of Helicella ( Trochoidea ) elata from Majorca,

considered doubtful by Jaekel (1952).

490

N. Maio, A. Petraccioli, P. Crovato, N. Amor & G. Odierna

Figs. 5-22. Shells of T. caroni. Figs. 5-7. Spigno Satumia, height 8.8 mm, diameter 6.1 mm: apertural view (Fig. 5), umbilical

view (Fig. 6), apical view (Fig. 7). Figs. 8-10. Terracina, height 9.6 mm, diameter 7.1 mm: apertural view (Fig. 8), umbilical

view (Fig. 9), apical view (Fig. 10). Figs. 1 1-13. Capri, height 6.7 mm, diameter 6.6 mm: apertural view (Fig. 1 1), umbilical

view (Fig. 12), apical view (Fig. 13). Figs. 14-16. Sorrento, juvenile shell, height 3.3 mm, diameter 3.5 mm: apertural view

(Fig. 14), apical view (Fig. 15), umbilical view (Fig. 16). Figs. 17-19. Terrasini, height 11.1 mm, diameter 9.6 mm: apertural

view (Fig. 1 7), umbilical view (Fig. 1 8), apical view (Fig. 1 9). Figs. 20-22. Annaba (Algeria) Lobbeclce Museum, Diisseldorf,

Germany, height 9.6 mm, diameter 8.6 mm: apertural view (Fig. 20), umbilical view (Fig. 21), apical view (Fig. 22).

diameter 8.6 mm: apertural view (Fig. 20), umbilical view (Fig. 21), apical view (Fig. 22).

New faunistic data on Trochoidea (Trochoidea) caroni (Deshayes, 1832) (Gastropoda Pulmonata Hygromiidae) 491

Fig. 23. Carocolla elata Faure-Biguet and C. turrita Philippi, 1836 illustrated in Philippi (1836), pi. 8, fig. 16 and 17, re-

spectively. Fig. 24. Helix elegans from Ischia illustrated in Delle Chiaje (1841), pi. 72, figs. 3, 4. Fig. 25. Live specimen

of Trochoidea caroni from Terracina, height 7.8 mm, diameter 7.0 mm. Fig. 26. Live specimen of T. caroni from Terrasini,

height 12.5 mm, diameter 10.4 mm. Fig. 27. Live specimen of T. caroni from Capri, height 6.8 mm, diameter 6.6 mm. Fig.

28. T. caroni from Terrasini, detail of two eggs (diameter 1.5 e 1.7 mm).

492

N. Maio, A. Petraccioli, P. Crovato, N. Amor & G. Odierna

Trochoidea ( Trochoidea ) trochlea (Pfeiffer, 1846)

Original description. Pfeiffer, 1846, species

no. 302, p. 69, locus typicus: “Patria ?”: “ Helix

trochlea Pfr. - T. perforata, conico-turrita, trochlearis,

costulato-striata, albida, interdum fusco-maculata

vel basi fusco -fas data; anfr. 8 acute crenulato-

carinati, late exserti, utriuque planulati, ultimus

basi subtilius et confertius striatus, convexius cuius,

antice non descendens; apertura securiformis; perist.

acutum, margine basali arcuato, intus leviter labi-

ato, columellari subdilatato, ad perforationem sub-

reflexo. Diam. 7 2/3, alt. 8 mm [h/d=1.04] - Patria?

( Coll. Cuming. y\

Examined material. The first faunistic re-

ports of T. trochlea date back to Morelet (1853,

sub Helix trochlea Pfr.) from the promontory of

Ippona, Capo di Guardia near Bona [present town

of Annaba, author’s note], Algeria, and to Bour-

guignat (1864) who reports four varieties of He-

licella trochlea Pfeiffer (var. minor, depressa,

fusco-maculata and hypozona ) from Bugia [present

town of Bejai'a, author’s note], Algiers and at

Capo di Guardia, near Annaba (Algeria). Bourguig-

nat (1868) also reports it from Tunisia at Kamart

(Tunis), in the ruins of Carthage and on the east-

ern slope of Djebel Ahmar (Tunisia). Specimens

with this determination from Annaba are kept in

the Lobbecke Museum in Diisseldorf (1868, legit

Lischke) and have h/d ratios in the range 0.68-

1.04 (mean: 0.86) (Maio, pers. obs.; Figs. 29-31).

Kobelt (1877) writes: “This very attractive species

represents [=substitutes?] the group of the “ Hel .

Caroni ” in Algeria, where it is found in the Prov-

inces of Algiers and Constantine in various

points” (Fig. 32). De Saint-Simon (1882) reports

specimens of Helix trochlea collected at Capo di

Guardia, near Bona, of which he examined the

radula. Fetoumeux & Bourguignat (1887) also re-

port it from Tunisia, as well as from Bona, Bugia

and Algiers (Algeria). Monterosato (1893) reports

H. trochlea Pfeiff. from “Algeria a Bona e nel de-

serto del Sahara (Mouss.)” and H. speciosa,

Monts, as “nuova forma vicina alia trochlea ma

pur distinta” from Salaparuta (Trapani Province)

in Sicily (Prof. A. Palumbo).

Wenz (1923) reports this species as a “Pliocen.

S. - Diluv.” fossil at "Cap de Garde bei Bone” and

Joleaud (1936) also reports it as a fossil from Capo

di Guardia in the “panchina quatemario antica del

posto”. Sacchi (1955a) reports finding specimens

of Helicella ( Trochoidea ) trochlea Drap. in Eastern

Algeria at Capo Carbon, near Bugia and at Capo di

Guardia.

In the F. Pfeiffer collection of the Senckenberg

Forschungsinstitut und Naturmuseum in Frankfurt

am Main (Germany), there is a specimen reported

as “Helix trochlea Pf., coll. O. Reinhardt ex Par-

reyss” from "Sicilia" (h/d=0.81) and nine specimens

reported as “ Trochula trochlea Pf., coll. C. Bout ex

H. Rolle” from "Sierra Nevada, Hispania"

(h/d=0.72). In the same collection there are other

specimens with labels indicating provenance from

sites in Algeria and Tunisia (R. Janssen, pers. com.;

see Table 3).

Trochoidea ( Trochoidea ) elegans (Gmelin, 1791)

Original description. Gmelin, 1791, species

no. 229, p. 3642, locus typicus “Barbaria et australi

Europa”: “ Helix elegans. H. testa pyramidali alba

umbilicata: anfractibus sex acutis planiusculis

marginatis. Habitat terrestris in Barbaria et australi

Europa. [Barbaria or Berberia: the Mediterranean

coastal area of Barbary and the Barbary States:

Tunisia, Algeria and Morocco, author’s note]”.

Examined material. During the present

study, some populations with particularly elevated

spire were found in the literature. Morphologically

very similar to T. caroni, they were of uncertain tax-

onomic attribution. The reports are included to

complete the information on this particular group

of land snails.

Sacchi (1955a) reports finding specimens of He-

licella elegans with elevated spire and morphologi-

cal characteristics very similar to H. elata on Ibiza.

In the same paper he reports “molto turricolate”

forms of H. elegans from Tunis and specimens of

Helicella ( Trochoidea ) elegans stated to belong to

“forme assai alte di spira” from Southern France.

Sacchi (1954, 1955a) does not exclude the hypothe-

sis of the presence of turriculate forms in the Balea-

ric Islands, though he considers them the result of

geographical isolation of populations of H. elegans

(reported, moreover, by Bofill I Poch & D’Aguillar-

Amat (1924) as Helix ( Trochula ) elegans).

Two historical reports of T. elegans by Ross-

massler (1837, Tafel XXVI, Fig. 345, sub H. elegans

Gm.) and by Aradas & Maggiore (1840: specie

FXVI, Elice elegante H. elegans Gm) from Sicily

New faunistic data on Trochoidea (Trochoidea) caroni (Deshayes, 1832) (Gastropoda Pulmonata Hygromiidae) 493

Figures 29-31. Shell of Trochoidea trochlea from Annaba (Algeria), Lobbecke Museum, Dusseldorf, Germany, height 8.1

mm, diameter 10.2 mm: apertural view (Fig. 29), umbilical view (Fig. 30), apical view (Fig. 31). Figure 32. Figure n. 1470

of Helix trochlea from plate 146 in Kobelt (1877).

are also noteworthy. In particular, the latter authors

write: “Rara: gli esemplari che possediamo ci sonper-

venuti da Palermo ad eccezione di un solo ritrovato

nella plaia di Catania. Diam. lin. [Linea siciliana (o

linia) = 1.79234583 mm] 5 X A [= 9.86 mm], altezza

lin. 2 y 2 [=4.48 mm] (h/d: 0.46). Coll. (A.G.) [=

Abate Emiliano Guttadauro?]". Philippi (1836) also

reports it with the name Carocolla elegans Lamk.

from Palermo and Pantelleria Island. The same

citation is repeated by Monterosato (1893) with the

comment “Non e stato piu ritrovata, ne credo si

trovi”.

DISCUSSION

On the basis of the data collected, the current dis-

tribution of T. caroni, limited to Capri and Sicily

(Manganelli et al., 1995), proves to be much wider

(Fig. 33). In peninsular Italy, new sites of collection

are in fact reported: one at Terracina with live speci-

mens, one at Spigno Saturnia with many fresh

empty shells, one in Sorrento with two empty shells

of juveniles, one at Castagneto Carducci with a

shell of an adult specimen, one at Manfredonia with

a shell in a museum collection, dating to 1878, and

a bibliographic specimen from Ischia, dating to

1822-1830. It is also important to mention the re-

ports by Statuti (1882: Porto Badino and Torre

Olevola near Terracina) and the fossil reports from

Capri by Segre (1959) attributed by him to the

Lower Pleistocene, when Capri was joined to the

Sorrentina peninsula (Barattolo et al., 1992; Ca-

passo Barbato & Gliozzi, 1995). This report there-

fore demonstrates the ancient presence of the

species on the island of Capri.

Of similar interest is the finding of fossil speci-

mens on Malta, which may demonstrate colonisa-

tion from Sicily during the Quaternary (Giusti et al.,

1995; Hunt, 1997; Kolouch, 2003). T. caroni may

have been present on the island until last century at

least, as historical reports from the mid and late

nineteenth century suggest (Gulia, 1874; Feilden,

1879).

Reports from North Africa are also interesting

but require further confirmation, since local forms

of T. trochlea or other taxa cannot be excluded.

Reports from the Balearic Islands cannot be

verified since the original specimens cited in the

papers could not be traced.

In the light of this new data, it is possible to say

that the distribution of T. caroni is certainly wider

than hitherto known, although it is not yet possible

to draw conclusions on the origin and spread of this

population. Likewise for T. trochlea and T. elegans

with elevated spires, it is impossible to determine

the correct systematic, phylogenetic and biogeo-

graphic position of the taxa and their effective rela-

494

N. Maio, A. Petraccioli, P. Crovato, N. Amor & G. Odierna

tionships with T. caroni. The morpho-biometric

criterion used to date does not enable clear discrim-

ination between the various populations. T. trochlea

should have an h/d between 1.14 and 0.70 or lower,

T. caroni between 1.30 or higher and 0.88 and T.

elegans between 0.82 and 0.62 or lower. Thus the

h/d ratios of the three forms overlap and the same

is true of their distributions.

Studies of anatomical characters do not solve

the problem. Interestingly, in their times, von

Ihering (1892) and Hesse (1934) already observed

that the anatomy of specimens of T. caroni from

Capri was indistinguishable from that of Sicilian

specimens. Moreover, Caziot (1908) and Sacchi

(1956a) considered that the anatomy of H. elegans

from Tunisia was identical to that of Sicilian H. ca-

roni-elata. Giusti et al. (1995) observed an evident

similarity of genital systems between T. caroni from

Sicily and T. elegans from Tuscany and Algeria,

sustaining that anatomically, all species of the “ ele-

gans group” ( T. elegans , T. caroni, T. spratti and T.

cumiae) were indistinguishable.

Sacchi (1955a; 1956b) considers Helix caroni-

elata and H. trochlea valid species, distinct from H.

elegans by virtue of “their particular geographic dis-

tribution” and the “remarkable morphological char-

acterisation” of their shells, even if he views all

these species as a single species complex (“ciclo di

forme”). Giusti et al. (1995), too, suggest that these

species probably all belong to a hypothetical “su-

perspecies” or “species group”, the validity of

which has yet to be confirmed.

According to Liggia (2013), T. trochlea Pfeiffer

1846 is synonymous with T. elegans (Gmelin

1791), but the site does not provide sources or other

references for this conclusion.

Conservation status

In the Red List of the International Union for the

Conservation of Nature (IUCN), version 2013.1.

(Falkner et al., 2011), the status of T. caroni is clas-

sified as “LC” (Least Concern) or “lowest risk, not

endangered or low risk”. In the Red List of the 27

countries of the European Union it is classified as

“LC” and is considered a “European endemic

species” (Cuttelod et al., 2011). T. elegans is also

classified as “LC” by the IUCN (Gargominy, 2011)

and in the European Red List it is considered a “Eu-

ropean endemic species” (Cuttelod et al., 2011). In

view of the extreme localisation of currently known

populations of these species, we think they are near

threatened and worthy of protection at regional and

national level.

Figure 33. Sites of finding of Trochoidea caroni in the Western Mediterranean.

New faunistic data on Trochoidea (Trochoidea) caroni (Deshayes, 1832) (Gastropoda Pulmonata Hygromiidae) 495

Species

Localities

Collection

Collector, date,

name reported

T. elegans

Spagna: Sierra Nevada

Slg. C. Bout ex H. Rolle

As T. trochlea

T. trochlea

Algeria

Slg. W. Kobelt

T. trochlea

Algeria: Maskara

Slg. W. Kobelt

T. trochlea

Algeria: Oran

Slg. Rossmassler

ex Terver 1944

T. trochlea

"Sicilia"

Slg. 0. Reinhardt

ex Parreyss

T. trochlea

Algeria: Cape de Garde,

Constantine

Slg. Jetschin

ex G. Schneider 1881

T. trochlea

Algeria: Bona

Slg. Jetschin

ex 0. Stoll 1899 ex R. Mousson

T. trochlea

Algeria: Bone [Cap de Garde]

Slg. Jetschin

ex Kobelt 1887

T. trochlea

Algeria: Bone

Slg. Jetschin

ex M. Braun 1887

T. trochlea

Tunisia: “Umgebung von Tunis”

Slg. 0. Reinhardt

ex? 5.1891

T. trochlea

Algeria: Bone, Cap de Garde

Slg. W. Kobelt

T. trochlea

Tunisia: “SW Biserta bei la

PecherieamSee von Biserta”

Slg. H. Kaltenbach 15.3.1956

T. trochlea

Tunisia: “PaBhohe s. Biserta

von Tunisaus”

Slg. H. Kaltenbach 14.3.1956

T. caroni

Is. Capri

Slg. C. R. Boettger 1906

As T. elata

T. caroni

Is. Capri

Slg. K. L. Pfeiffer 1928

As T. elata

T. caroni

Is. Capri

Slg. J. Blum 3.1891

As T. elata

T. caroni

Is. Capri

Slg. W. Kobelt 1878

As T. elata

T. caroni

Is. Capri: Mte. Solaro

Slg. K. L. Pfeiffer 1928

As T. elata

T. caroni

Is. Capri

ex Fitz-Gerald 1878 as T. elata

T. elata

Napoli

Slg. C. Bosch ex H. Rolle

As T. elata

T. caroni

S -Italia: Manfredonia

Slg. W. Kobelt 1878

As T. elata

T. caroni

Is. Capri

Slg. S. H. Jaeckel

As T. elata

T. caroni

Is. Capri

Slg. H. Kaltenbach ex 0. Schmidt

As T. elata

T. caroni

Is. Capri

Slg. Jetschin

ex Rousseau 1901 as T. elata

T. caroni

Is. Capri: M.te Solaro

Slg. Schlickum 19.8.1956

As T. elata

T. caroni

Is. Capri

Slg. W. Kobelt

ex Heynemann as T. elata

T. caroni

Is. Capri

Slg. Jetschin

ex Kobelt 1880 as T. elata

T. caroni

Is. Capri

Slg. Ehrmann

4.1894 as T. elata

T. caroni

Is. Capri

Slg. L. Henrich

As T. elata

T. caroni

Is. Capri

Slg. H. Kaltenbach ex 0. Schmidt

ex A. Kohler 1906 as T. elata

T. caroni

Sicilia

Slg. Rossmassler

ex Philippi as T. elata

T. caroni

Sicilia

Slg. 0. Reinhardt

ex Parreyss as T. elata

T. caroni

Sicilia

Slg. 0. v. Moellendorff

As T. elata

T. caroni

Sicilia

Slg. Rossmassler

ex Hueb du Pavilion as T. elata

T. caroni

Sicilia: Trapani

Slg. S. H. Jaeckel

As T. elata

T. caroni

Sicilia: San Vito

Slg. K. L. Pfeiffer 1930

As T. elata

Table 3. Faunistic data of Trochoidea sp. della “Collezione L. Pfeifer” of the Senckenberg Forschungsinstitut und

Naturmuseum (R. Janssen, pers. com.) of Frankfurt am Main, Germany.

496

N. Maio, A. Petraccioli, P. Crovato, N. Amor & G. Odierna

Species

Localities

Collection

Collector, date,

name reported

T. caroni

Sicilia

Slg. Jetschin

ex Linnaea 1885 as T. data

T. caroni

Sicilia

ex Parreyss as T. data

T. caroni

Sicilia: Trapani

Slg. K.L.Pfeiffer ex K.

Henschel 1890

As T. data

T. caroni

Sicilia: Palermo

Slg. C. Bosch ex H. Rolle

As T. data dilatata

T. caroni

Sicilia: M.te San Giuliano

Slg. K. L. Pfeiffer 1930

As T. data

T. caroni

Sicilia: Trapani-Marsala

Slg. K. L. Pfeiffer 1930

As T. data

T. caroni

Sicilia: Trapani

Slg. Jetschin

ex Caruana Gatto as T. data

T. caroni

Sicilia: Trapani

Slg. Jetschin

ex Kobelt 1880 as T. data

T. caroni

Sicilia: Sferracavallo near

Palermo

Slg. W. Kobelt 1878

As T. data

T. caroni

Sicilia

Slg. W. Kobelt

ex Benoit as T. data

T. caroni

Sicilia

Slg. Rossmassler

ex Ziegler as T. data

T. caroni

Sicilia: Calatafimi

Slg. W. Kobelt

ex Monterosato as T. data

T. caroni

Sicilia: Trapani

Slg. W. Kobelt 1878

As T. data

T. data

Sicilia: Palermo

Slg. Rossmassler

ex Philippi as T. data caroni

T. data

Sicilia

Slg. W. Kobelt

ex Westerlund as T. data caroni

T. data

Sicilia

Slg. 0. Reinhardt

ex Parreyss as T. data caroni

T. data

[Sicilia]

Slg. Knotte

As T. data caroni

T. data

Sicilia

Slg. 0. Boettger

ex C. Koch as T. data caroni

T. data

Sicilia

Slg. K. Harhagen

As T. data caroni

T. data

Sicilia

Slg. 0. v. Moellendorff

As T. data caroni

T. data

Sicilia

Slg. A. Gyssea

As T. data caroni

T. data

Sicilia

Slg. W. Kobelt

ex Benoit as T. data caroni

T. data

Sicilia: Mte. Pellegrino

near Palermo

ex ? 4.1889 as T. data caroni

T. data

Sicilia

Slg. S. H. Jaeckel

ex Scholtz as T. data caroni

T. data

Sicilia: Palermo

Slg. G. Nagele ex

Monterosato 1897

As T. data caroni

T. data

Sicilia: Palermo

Slg. G. Nagele

As T. data caroni

T. data

Sicilia: Palermo

ex ? As T. data caroni

T. data

Sicilia: Palermo

ex Linnaea 1883 as

T. data caroni

T. data

Sicilia: Sferracavallo near

Palermo

Slg. S. H. Jaeckel 4.1935

As T. data caroni

T. data

Sicilia: Palermo

Slg. Jetschin

ex Wohlberedt 1903 as

T. data caroni

T. data

Sicilia: Palermo

Slg. Th. Kriiper

As T. data caroni

T. data

Sicilia: Palermo

Slg. H. Kaltenbach

ex A. Kohler as T. data caroni

T. data

Sicilia

Slg. Jetschin

ex Zellweger as T. data caroni

T. data

Sicilia: near Palermo

Slg. Jetschin

ex Kobelt 1880 as T. data

caroni

Table 3. Faunistic data of Trochoidea sp. della “Collezione L. Pfeiffer” of the Senckenberg Forschungsinstitut und

Naturmuseum (R. Janssen, pers. com.) of Frankfurt am Main, Germany.

New faunistic data on Trochoidea (Trochoidea) caroni (Deshayes, 1832) (Gastropoda Pulmonata Hygromiidae) 497

Species

Localities

Collection

Collector, date,

name reported

T. elata

Sicilia: Palermo

Slg. W. Kobelt

ex Monterosato as T. elata

caroni

T. data

Sicilia: Palermo, San Polo

Slg. Jetschin

ex A. Bofill 1900 as

T. elata caroni

T. elata

Sicilia: Sferracavallo near

Palermo

Slg. H. Kaltenbach ex A.

Kohler 1913

As T. elata caroni

T. elata

Sicilia: Palermo

Slg. C. Bosch ex H. Rolle

As T. elata caroni

T. elata

Sicilia: Palermo

Slg. W. Kobelt

As T. elata caroni

T. elata

Sicilia

ex ? as T. elata caroni

T. elata

Sicilia: Sferro Cavallo

Slg. C. Bosch ex H. Rolle

As T. elata caroni

Table 3. Faunistic data of Trochoidea sp. della “Collezione L. Pfeiffer” of the Senckenberg Forschungsinstitut und

Naturmuseum (R. Janssen, pers. com.) of Frankfurt am Main, Germany.

ACKNOWLEDGMENTS

We thank: Joseph Boscheinen (Malakozoologie

Curator, Stadt Diisseldorf Aquazoo, Lobbecke-

Museum, Germany) for loan of specimens of T. ca-

roni e T. trochlea preserved in the Lobbecke

Museum; Ronald Janssen (Sektionsleiter Malakolo-

gie, Senckenberg Forschungsinstitut und Naturmu-

seum, Frankfurt am Main, Germany) for

informations and photos of the specimens of “L.

Pfeiffer collection”. Folco Giusti (University of

Siena, Italy), Giuseppe Manganelli (University of

Siena, Italy), Simone Cianfanelli (University of

Florence, Italy) and Rafael La Pema (University of

Bari) for precious advices given; Rudiger Bieler

(Curator and Director, Invertebrates Department of

Zoology, Field Museum of Natural History di

Chicago, USA), Jochen Gerber (Collections Mana-

ger, Invertebrates Department of Zoology, Field

Museum of Natural History, Chicago, USA), Vir-

ginie Heros and Philippe Maestrati (rispectively

Chargee de conservation, collection Mollusques,

Departement Systematique et Evolution, and Pho-

tographer, Museum National d'Histoire Naturelle,

Paris, France), Monica Leonardi (Museo Civico di

Storia Naturale, Milano, Italy), Giacomo Doria and

Maria Tavano (Museo Civico di Storia Naturale

"Giacomo Doria", Genova, Italy), Aldo Giacomo

Segre (Rome, Italy), Fabio Parenti (Rome, Italy),

Ignazio Sparacio (Palermo, Italy), Walter Renda

(Amantea, Italy), Agatino Reitano (S. G. Galemo,

Catania, Italy), Fabio Liberto (Cefalu, Italy),

Giuseppe Fasulo (Napoli, Italy), Sergio Duraccio

(Naples, Italy) and Mario Cuomo (Naples, Italy) per

le informazioni inedite fomite; the Centro Caprense

“I. Cerio”, Capri (Naples, Italy) and, particolarly,

the President Claudio Giordano for collaboration.

A particular thank goes to Alessandro Hallgass

(Rome, Italy) for specimens from Terracina.

REFERENCES

AA.VV., 2007. Annual Report 2006-2007. Heritage

Malta, Valletta, 196 pp.

Alzona C., 1971. Malacofauna Italica. Catalogo e bibli-

ografia dei molluschi viventi, terrestri e d’acqua

dolce. Atti della Societa Italiana di Scienze Naturali

e del Museo Civico di Storia Naturale di Milano,

111: 1-433.

Aradas A. & Maggiore G., 1840 [71841]. Catalogo ra-

gionato delle conchiglie viventi e fossili di Sicilia es-

istenti nelle collezioni del Dottor Andrea Aradas e

dell'estinto Abate D. Emiliano Guttadauro Casinese.

Memoria IV. Seguito dei Gasteropodi. Letta nella se-

duta ordinaria del di 23 agosto 1840. Atti Accademia

Gioenia di Scienze Naturali di Catania, 17: 53-106.

Bagni L., 2013. Species: Trochoidea elegans Gmelin,

1791 - ID: 4531250015. Available at: http://www.ba-

gniliggia.it/WMSD/HtmSpecies/453 12500 1 5 .htm.

Worldwide Mollusc Species Database. Last update:

September 30th, 2013.

Bank R.A., 2011. Fauna Europaea: Mollusca Gastropoda.

Fauna Europaea version 2.4. Checklist of the land

and freshwater Gastropoda of Italy. Fauna Europaea

Project: 1-49. Last update: July 24th, 2011.

http://www.nmbe.unibe.ch/sites/default/files/uplo-

ads/pubinv/fauna_europaea_gastropoda_o f_italy.pdf

Barattolo F., Cinque A., D’Alessandro E., Guida M.,

Romano P. & Russo Ermolli E., 1992. Geomorfolo-

498

N. Maio, A. Petraccioli, P. Crovato, N. Amor & G. Odierna

gia ed evoluzione tettonica quaternaria dell’Isola

di Capri. Studi Geologici Camerti, Vol. Spec. 1:

221-229.

Bellini R., 1899. Ulteriori osservazioni sui molluschi

terrestri dei dintorni di Napoli. Rivista Italiana di

Scienze Naturali, 19: 100-118.

Bellini R., 1900. I molluschi extramarini dell’isola di

Capri. Bollettino della Societa Zoologica Italiana, 19:

29-55.

Bellini R., 1901. Alcune osservazioni sulla distribuzione

ipsometrica dei molluschi terrestri nell’isola di Capri.

(Sunto). Monitore Zoologico Italiano, 12: 180-182.

Bellini R., 1910. Osservazioni geomorfologiche sulk

Isola di Capri. Atti della Societa Italiana di Scienze

Naturali, 49: 244-260.

Bellini R., 1915. I molluschi extramarini dei dintorni di

Napoli. Bollettino della Societa dei Naturalisti in

Napoli, 27: 149-194.

Bellini R., 1920. Osservazioni nell’Isola di Capri a

conferma dell’antico stato della regione tirrena.

Bollettino della Societa dei Naturalisti in Napoli, 32:

125-140.

Bellini R., 1922. Nel campo della fauna dell’Isola di

Capri. Le pagine dell’Isola di Capri (agosto 1922):

3-4.

Bellini R., 1924. La distribuzione geografica dei Mol-

luschi terrestri e l’ipotesi della Tirrenide. Rivista di

Biologia, 6: 218-221.

Bellini R., 1926. Osservazioni a favore della «Tirrenide».

Bollettino della Societa Geologica Italiana, 45: 1 84—

192.

Benoit L., 1862. Illustrazione sistematica critica icono-

grafica de' testacei estramarini della Sicilia Ulteriore

e delle isole circostanti. Quaderno 4. Stabilimento

tipografico G. Nobile, Napoli, pp. 181-248, PI. 7-8.

Benoit L., 1875. Catalogo delle conchiglie terrestri e flu-

viatili della Sicilia e delle isole circostanti. Bullettino

della Societa Malacologica Italiana, 1: 129-163.

Benoit L., 1881. 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. Atti della Reale Accademia Peloritana, 3:

43-216.

Bofill I., Poch A. & De Aguillar-Amat J.B., 1924. Mala-

cologia de les illes Pitiuses. Trabajos del Museo de

Ciencias Naturales de Barcelona, 10: 1-64.

Bourguignat J.R., 1864. Malacologie de l’Algerie ou

histoire naturelle des animaux mollusques terrestres

et fluviatiles recueillis jusqu'a ce jour dans nos pos-

sessions du nord de l'Afrique. Bouchard-Huzard,

Paris, 1: i-xii + 294 pp., 32 Pis.

Bourguignat J.R., 1868. Souvenirs d'une exploration

scientifique dans le nord de l'Afrique. Vol. III.

Histoire malacologique de la Regence de Tunis.

Challamel Aine, Paris, 36 pp.

Bullen R., 1910. Notes on (I) Pleistocene, (II) Holocene,

(III) Recent non-marine shells from Mallorca; (IV)

marine shells associated with the Holocene deposits;

(V) marine shells from Alcudia, Mallorca; (VI) non-

marine shells from Manresa, Cataluna. Proceedings

of the malacological society of London, 9: 118-122.

Calcara P., 1845. Esposizione dei molluschi terrestri

e fluviatili dei dintorni di Palermo. Atti della Ac-

cademia di Scienze e Lettere di Palermo, 1: 1-46.

Capasso-Barbato L. & Gliozzi E., 1995. Biochronolog-

ical and paleogeographical implication of a well-bal-

anced late Middle Pleistocene fauna from Quisisana -

Certosa (Capri, Southern Italy). Bollettino della So-

cieta Paleontologica Italiana, 34: 235-261.

Caziot E., 1908. Etude sur les Helix elegans Drapamaud,

et seitula Cristofori & Jan. Bulletin de la Societe

zoologique de France, 33: 183-186.

Cossignani T. & Cossignani V., 1995. Atlante delle

conchiglie terrestri e dulciacquicole italiane. L'lnfor-

matore Piceno, Ancona, 208 pp.

Costa A., 1840. Molluschi. In: Statistica fisica ed econom-

ica dell’isola di Capri. Esercitazioni accademiche

degli aspiranti naturalisti ovvero raccolta di memorie

dell’ accademia di tal nome, Napoli, 2, pp. 64-72.

Cuttelod A., Seddon M. & Neubert E., 2011. European

Red List of Non-marine Molluscs. Publications Of-

fice of the European Union, Luxembourg, 98 pp.

De Gregorio A., 1895. Appunti su talune conchiglie

estramarine di Sicilia viventi e fossili con la spie-

gazione delle tavole dell’ opera di Benoit. II Natura-

lista siciliano, 14: 183-212.

De Saint-Simon A., 1882. Etude des Helix du groupe de

V elegans. Bulletin de la Societe d'histoire naturelle

de Toulouse, 16: 191-227.

Delle Chiaie S., 1830 [1822]. Memorie sulla storia e

notomia degli animali senza vertebre del Regno di

Napoli. Figure. Napoli: taw. I-IV e I-CIX.

Delle Chiaie S., 1841. Descrizione e notomia degli ani-

mali invertebrati della Sicilia citeriore osservati vivi

negli anni 1822-1830. Tomo II. Molluschi Gastero-

pedi. Stabilimento tipografico di C. Batelli e comp.,

Napoli, 146 pp.

Deshayes G.P., 1832 [1830]. Histoire naturelle des Vers.

Par Bruguiere et De Lamarck, continuee par Mr. P.

G. Deshayes. Tome second, part 2. In: Encyclopedic

Methodique (Dictionaire Encyclopedique Methodique),

ou par ordre de matieres. Agasse, Paris, pp. j-vij[=l-

7], pp. 185-271.

Falkner G., Falkner M. & von Proschwitz T., 2011. Tro-

choidea caroni. In: IUCN 2013. IUCN Red List of

Threatened Species. Version 2013.1. Downloaded on:

www.iucnredlist.org. Last view: 07 October 2013.

Feilden H.W., 1879. The land and fresh-water mollusca

of Maltese Group. The Zoologist, 3: 193-199.

Ferussac A.E.J.P.J.F. d’Audebard de, 1821. Tableaux

systematiques des animaux mollusques classes en

New faunistic data on Trochoidea (Trochoidea) caroni (Deshayes, 1832) (Gastropoda Pulmonata Hygromiidae) 499

families naturelles, dans lesquels on a etabli la con-

cordance de tous les systemes; suivis d'un prodrome

general pour tous les mollusques terrestres ou fluvi-

atiles, vivants ou fossiles. Bertrand, Paris, Sowerby,

Londres, 110 pp.

Fiorentino V., Cianfanelli S., Manganelli G. & Giusti F.,

2004. I molluschi non marini delle isole Egadi

(Canale di Sicilia): biodiversita e conservazione. So-

ciety italiana di Ecologia, XIV Congresso Nazionale,

4-6 ottobre 2004, Siena: 89 [poster presentation],

Gargominy O., 2011. Trochoidea elegans. In: IUCN

2013. IUCN Red List of Threatened Species. Version

2013.1. Disponibile su: www.iucnredlist.org. Last up-

date: October 7th, 2013.

Giusti F., Manganelli G. & Schembri P.J., 1995. The non-

marine molluscs of the Maltese Islands. Museo Re-

gional di Scienze Naturali, Monografie, 15: 1-607.

Gulia G., 1874. [Untitled]. II Barth (Malta), 1: 467.

Gmelin J.F., 1791. Caroli a Linne, systema naturae. Tom.

I. Pars VI. Beer, Lipsiae, pp. 3021-3910.

Halgass A., 2007. Trochoidea ( Trochoidea ) caroni.

http ://www. naturamediterraneo .com/forum/topic .asp

?T0PIC_ID=22967. Last view: 07 October 2013.

Hesse P., 1934. Zur Anatomie und systematik palearktis-

cher Stylommatophoren. Zweiter Teil. Zoologica,

Stuttgart, 85: 1-59.

Hunt C.O., 1997. Quaternary deposits in the Maltese

Islands: a microcosm of environmental change in

Mediterranean lands. Geojoumal, 41: 101-109.

Jaeckel S., 1952. Die Mollusken der Spanischen Mittle-

meer-Inseln. Mitteilungen aus dem Zoologischen

Museum in Berlin, 28: 53-143.

Joleaud L., 1936. Etude geologique de la region de Bone

e de la Calle. Bulletin Service Carte geologique

Algerie, 12: 1-199.

Kobelt W., 1877. Iconographie der Land- & Siisswasser-

Mollusken mit vorziiglicher Beriicksichtigung der eu-

ropaischen noch nicht abgebildeten Arten von E. A.

Rossmassler. V. Band. Mit dreissig Tafeln. -Kreidel,

Wiesbaden, pp. [1-3], 1-129, Taf. 121-150.

Kolouch L.R., 2003. Suchozemsti, sladkovodni a brakieti

mikkysi ostrovu Malty Terrestrial, freshwater, and

brakish mollusca of Malta islands. Malacologica

Bohemoslovaca (Ceskoslovenska slimac), 2: 43-50.

Letoumeux A. & Bourguignat J.R., 1887. Prodrome de

la malacologie terrestre et fluviatile de la Tunisine.

Exploration scientifique de la Tunisine. Imprimerie

National, Paris, 166 pp.

Liberto F., Giglio S., Reitano S., Colomba M.S. & Spara-

cio I., 2010. Molluschi terrestri e dulciacquicoli di

Sicilia della collezione F. Mina Palumbo di Castel-

buono. Monografie naturalistiche, 2. Edizioni

Danaus, Palermo, 136 pp.

Manganelli G., Bodon M., Favilli L. & Giusti F., 1995.

Gastropoda Pulmonata. In: Minelli A., Ruffo S. & La

Posta S. (Eds). Checklist delle specie della fauna ital-

iana, 16. Calderini, Bologna, 60 pp.

Mazyck W.G., 1876. On the Occurrence of Helix Ter-

restris , Chemnitz, in North America. Proceedings of

the Academy of Natural Sciences of Philadelphia, 28:

127-128.

Misuri A., 1907. Sul genere Helix nella provincia delF

Umbria. Bollettino della Societa Zoologica Italiana,

Roma, Serie II, Vol. 8: 54 pp. Pp. 54 (172 - 192; 274

- 306) con 3 tav.

Monterosato T. [Di Maria Tommaso Marchese Allery di

Monterosato], 1893. Conchiglie terrestri delle Isole

adiacenti la Sicilia. Atti della Reale Accademia di

Scienze Lettere e Belle Arti di Palermo, (III serie)

2/1892: 1-34.

Monterosato T. [Di Maria Tommaso Marchese Allery di

Monterosato], 1894. Conchiglie terrestri viventi e

fossili di Monte Pellegrino. II Naturalista siciliano,

13: 165-173.

Morelet A., 1853. Catalogue des Mollusques terrestres et

fluviatiles de FAlgerie, Paris, 4: 280-505.

Pallary P., 1904. Quatrieme contribution a F etude de la

faune malacologique du NO de FAfrique. Memoires

de la Societe geologique de France. Paleontologie,

52: 1-59.

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. Bollettino della sezione Campania

ANISN, 16: 19-48.

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.

IV International Congress of the European Malaco-

logical 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., 2006. 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: 77-86.

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.

Petraccioli A., Marrone N., Giannetti F., Crovato P.,

Guarino F.M. & Maio N., 2011. Checklist prelim-

inare dei Gasteropodi continentali del Rio Santa

Croce (Latina, Lazio). Notiziario S.I.M., 29: 21-23.

Pfeiffer L., 1846. Symbolae ad hystoriam Heliceorum.

Sectio tertia. Th. Fischer, Cassellis, 100 pp.

500

N. Maio, A. Petraccioli, P. Crovato, N. Amor & G. Odierna

Pfeiffer L., 1848. Monographia heliceorum viventium.

Sistens descriptiones systematicas et criticas omnium

hums familiae generum et specierum hodie cogni-

tarum. Vol. 1. F. A. Brockhaus, Lipsiae. pp. I-XXXII

[= 1-32], 484 pp.

Pfeiffer L., 1853. Monographia heliceorum viventium.

Sistens descriptiones systematicas et criticas omnium

huius familiae generum et specierum hodie cogni-

tarum. Vol. 3. F. A. Brockhaus, Lipsiae. pp. I- VIII [=

1-8], 711pp.

Philippi R.A., 1836. Enumeratio Molluscorum Siciliae

cum viventium turn in tellure tertiaria fossilium, quae

in itinere suo observavit. Vol. 1, Schropp, Berolini

(Berlino), xiv + 267 pp.

Philippi R.A., 1844. Enumeratio molluscorum Siciliae

cum viventium turn intellure terziaria fossilium, quae

in itinere suo observavit. Vol. II. E. Anton, Halis

Saxonum, 303 pp.

Pirajno E., 1840. Catalogo dei molluschi terrestri e fluvi-

atili delle Madonie e luoghi adiacenti. Cefalu, 41 pp.

Power J., 1842. Guida per la Sicilia. Stabilimento

Poligrafico di F. Cirelli, Napoli, 382 pp.

Reitano A., 2006a. Trochoidea caroni (Deshayes, 1830).

http://www.sim-online.it/risorse/moll_vivi/gas-

tropoda%20terrestri/_gastropoda%20terrestri.htm.

Last update: April 28th, 2007.

Reitano A., 2006b. Hygromidae. http://freeweb.su-

pereva.com/siciliaria/Hygromidae.htm. Last update:

June 28th, 2007.

Reitano A., Liberto F., Giglio S., Grasso R. & Spena

M.T., 2012. Terrestrial Molluscs from R.N.I. “Grotta

Conza” (Palermo, North-Western Sicily) (Gastropoda

Architaenioglossa Pulmonata). Biodiversity Journal,

3: 555-570.

Rossmaessler E.A., 1837. Iconographie der Land- und

Siisswasser-Molluscken, mit vorzuglicher Beriick-

sichtigung der europaischen noch nicht abgebildeten

Arten. Heft 5-6. Arnold, Dresden e Leipzig, 70 pp.

Sacchi C.F., 1953. Un’escursione malacologia sulla

penisola sorrentina ed a Capri. Atti della Societa

Italiana di Scienze Naturali e del Museo Civico di

Storia Naturale, 92: 109-116.

Sacchi C.F., 1954. Contributo alia conoscenza dei popo-

lamenti delle piccole isole mediterranee. II. Cenni

biogeografici sulla malacofauna di Iviza (Pitiuse).

Bollettino di Zoologia, 21: 1-40.

Sacchi C.F., 1955a. Contributo alia conoscenza faunistica

della Campania. Ricerche malacologiche nella re-

gione sorrentina. I. Note sistematiche di alcune eli-

celline. Annuario dell’Istituto e Museo di Zoologia

Universita di Napoli, 6/1954: 1-14.

Sacchi C.F., 1955b. Contributo alia conoscenza faunistica

della Campania. Ricerche malacologiche nella re-

gione sorrentina. II. Appunti biogeografici. Annuario

dell’Istituto e Museo di Zoologia Universita di

Napoli, 6/1954: 1-14.

Sacchi C.F., 1956a. Sulla sistematica di alcune elicelline

(Stylomm. Helicidae) della fauna italiana. Annuario

dell’Istituto e Museo di Zoologia Universita di

Napoli, 7/1955 (6): 1-16.

Sacchi C.F., 1956b. II contributo dei molluschi terrestri

alle ipotesi del “Ponte Siciliano”. Elementi tirrenici

ed orientali nella malacofauna del Magreb. Archivio

Zoologico Italiano, 40/1955: 49-181.

Segre A., 1959. Isola di Capri, scala 1: 25000. Carta Geo-

logica d’ltalia, foglio n. 196 (Penisola Sorrentina -

Isola di Capri). Servizio Geologico Italiano. Roma.

Sherborn C.D., 1924. Index Animalium sive index

nominum quae ab A. D. MDCCLVIII generibus et

speciebus animalium imposita sunt. Sectio secunda a

kalendis ianuariis, MDCCCI usque ad finem decem-

bris, MDCCCL. Part V. (C-Ceyl). - London, by order

of the Trustees, pp. 945-1196.

Sparacio I., 2012. Le conoscenze sulla malacofauna

terrestre e dulciacquicola nella Sicilia dei primi de-

cenni del 1800 attraverso gli scritti di Jeannette

Power. II Naturalista siciliano, 36: 375-386.

Statuti A., 1882. Catalogo sistematico e sinonimico dei

Molluschi terrestri e fluviatili viventi nella Provincia

Romana. Atti delFAccademia Pontificia de’ nuovi

Lincei, 34/1881: 1-89.

Trechmann C.T., 1938. Quaternary conditions in Malta.

Geology Magazine London, 75: 1-26.

Von Ihering H., 1892. Morphologie und Systematik des

Genitalapparates von Helix. Zeitschrift Wis-

senschaftliche Zoologie, 54: 425-520.

Welter- Schultes F.W., 2012. European non-marine mol-

luscs, a guide for species identification. Planet Poster

Editions, Gottingen, pp. 679 pp.

Wenz W., 1923 - Pars 18. Gastropoda extramarina ter-

tiaria. II. In: C. Diener (Ed.), Fossilium catalogus. I:

Animalia. W. Junk, Berlin, pp. 353-736.

Westerlund C.A., 1889. Fauna der in der palaarctischen

Region (Europa, Kaukasien, Sibirien, Turan, Persien,

Kurdistan, Armenien, Mesopotamien, Kleinasien,

Syrien, Arabien, Egypten, Tripolis, Tunesien, Algerien

und Marocco) lebenden Binnenconchylien. II. Gen.

Helix. Friedlander, Berlin, pp. 1-473, 1-31, 1-8.

Biodiversity Journal, 2013, 4 (4): 501-506

Glycaspis brimblecombei Moore, 1 964 (Hemiptera Psyllidae)

invasion and new records in the Mediterranean area

Kheddar Reguia 1 & Francisco Javier Peris-Felipo 2 *

'Department of Agricultural sciences, Faculty agro-veterinary, Universityof Saad. Dahlab BP 270,09000 Blida, Algeria

laboratory of Entomology and Pest Control. Institute Cavanilles of Biodiversity and Evolutionary Biology. University of

Valencia. c/. Catedratico Jose Beltran, 2 - 46980 Paterna, Valencia, Spain

■"Corresponding author: Francisco.peris@uv.es

ABSTRACT Glycaspis brimblecombei Moore, 1964 (Hemiptera Psyllidae) is a sap-sucking insect known

for being a severe pest in several Eucalyptus spp. This paper provides new information about

the first record of G. brimblecombei in Algeria and Greece, and new information about its

biology, analyzing the evolution and reasons of its expansion. It is also the first record of the

parasitic wasp Psyllaephagus bliteus Riek in Algeria, frequently used as biological control

of this psyllid. Samplings were carried out in European and North African Mediterranean

countries from 2011 to 2013.

KEY WORDS Glycaspis brimblecombei', Psyllidae; Psyllae phagusbliteus; Mediterranean areas; invasion.

Received 22.10.2013; accepted 29.11.2013; printed 30.12.2013

INTRODUCTION

The red gum lerp psyllid, Glycaspis brimble-

combei Moore, 1964 (Hemiptera Psyllidae), is a

sap-sucking insect of Australian origin (Moore,

1964), which currently shows a widespread distri-

bution outside its native range due to frequent in-

troductions. It was detected in California, U.S.A. in

1998 (Brennan et al., 1999), in Mexico in 2000

(Castillo, 2003), in the Hawaiian Islands in 2001

(Nagamine & Heu, 2001), in Chile in 2002 (San-

doval & Rothmann, 2003), in Brazil (Santana et al.,

2003) and Mauritius (Sookar et al., 2003) in 2003,

in Madagascar in 2004 (Hollis, 2004), in Argentina

in 2005 (Bouvet et al., 2005), in Ecuador in 2007

(Onore & Gara, 2007), in Venezuela (Rosales et al.,

2008), Peru (Burckhardt et al., 2008) and Iberian

Peninsula (Hurtado & Reina, 2008; Valente &

Hoodkinson, 2009; Prieto-Lillo et al., 2009) in

2008, in Morocco in 2009 (Maatouf & Lumaret,

2012), in Italy in 2010 (Laudonia & Garonna, 2010)

and recently in France (Cocquempot et al., 2012).

This psyllid is considered a serious pest that

causes severe defoliation and some tree mortality on

several Eucalyptus spp., being included in the EPPO

list of quarantine species since 2002. It mainly feeds

on Eucalyptus camaldulensis Dehnh., although it

can also be found on other species including E. rudis

Endl., E. globulus Labill., E. diversicolor F. Muell.,

E. sideroxylon A. Cunn. ex Woolls, E. nicholii

Maiden et Blakely, E. lehmannii (Schauer) Benth.,

in California and E. blakelyi Maiden, E. nitens H.

Deane et Maiden, E. tereticornis Sm., E. dealbata

A. Cunn. ex Schauer, E. bridgesiana R. T. Baker, E.

brassiana S.T. Blake, E. mannifera Mudie, in Aus-

tralia (Moore, 1970; Brennan et al., 1999; 2001).

However, in the Mediterranean area is primarily as-

sociated with red eucalyptus (E. camaldulensis), fre-

quently used in urban and rural forestiy programmes

(Peris-Felipo et al., 2010).

502

Kheddar Reguia & Francisco Javier Peris-Felipo

G. brimblecombei can be distinguished from

other species due to the length of its cephalic pro-

jections, located below the eyes and known as genal

processes (Laudonia & Garonna, 2010). Adults

show sexual dimorphism mainly on the body size,

with females slightly larger than males (size varying

between 2.5 and 3.1 mm). The body is light green,

sometimes with yellow spots. During oviposition,

females lay yellow ovoid eggs, individually or in

groups and without any protection. Nymphs are yel-

lowish orange with grey wing rudiments. Nymphs

secrete honeydew, which builds a white cover for

protection until the adult stage is reached. This

cover, also called a shield, is conic in shape and is

built by several layers linked to each other. The

nymph and the protective shield, which reaches a

maximum size of 3.0 x 2.0 mm (Ide et al., 2006),

grow at the same rate. Oviposition takes place on

leaves and, in the case of adverse weather; the eggs

enter into a quiescence period until conditions be-

come favourable. After hatching, nymphs rapidly

develop into pupal instars, producing the protective

shield and readying the insect for final development

into adulthood (Laudonia & Garonna, 2010). Once

the adult stage is reached, reproductive activity

swiftly takes place, fertilized females oviposit and

nymphs hatch a few days later, starting another

cycle. G. brimblecombei may undergo a multivol-

tine cycle but variations of the life cycle have been

observed in different geographical contexts. For in-

stance, in Australia and California there are two to

four generations per year whilst in Chile the life

cycle is postponed for about one month during the

spring-summer period (Hidalgo, 2005). In fact, bio-

logical control programs have been applied in Aus-

tralia and California using the parasitoid

Psyllaephagus bliteus Riek, as it is known for its

efficiency in parasiting several psyllid species, in-

cluding G. brimblecombei (Riek, 1962).

The present paper provides biological and distri-

bution data of G. brimblecombei around the Mediter-

ranean countries and information about its

progressive expansion from the first record.

MATERIALS AND METHODS

Samples were carried out from 2011 to 2013

around the Mediterranean area excluding the eastern

countries. In order to determine the degree of ex-

pansion of Glycaspis several Eucalyptus spp. (E.

camaldulensis , E. globulus , E. bosistoana F. Muell.

and E. gomphocephala A. Cunn. ex DC.) were

checked. Eucalyptus- trees showing positive pres-

ence were sampled, and leaves and insects collected.

The samples were either hand-picked or collected

with entomological nets. Leaves were taken to the

laboratory for observation under light stereomicro-

scope. Samplings around different areas of Valencia

province (Spain) and North-West of Algiers (Alge-

ria) were also taken to deteimine the presence of the

parasitoid wasp Psyllaephagus blitteus.

RESULTS

The new records of G. brimblecombei increase

our knowledge of its distribution in countries where

it has not been discovered until this research such

as Algeria and Greece.

Samples confirm that this psyllid is well estab-

lished in South of Europe and North of Africa, hav-

ing expanded very fast since the first time that was

cited in Europe in 2008 (Spain) (Hurtado & Reina,

2008). During four years G. brimblecombei has

spread throughout Eastern and Southern Spain

(Peris-Felipo et al., 2010), Central and South of Italy

(Laudonia & Garonna, 2010; Peris-Felipo et al.,

2010) and France (Cocquempot et al., 2012), being

now reported also in North-West of Algiers (Alge-

ria) and the Peloponnese region (Greece) (Fig. 1).

There were differences between areas relating

to the species of Eucalyptus infested by Glycapsis :

E. camaldulensis was the only species affected in

Europe while E. gomphocephala and to a lesser ex-

tent E. globules, were also infested in Algeria.

There were no psyllids reported in E. bosistoana.

In addition, in Algeria during 2012, samples to

know the global population and activity of G. brim-

blecombei were done. In order to represent on the

same graph the abundance, temperature (T) and

precipitation (P), the Walter ratio (P = 3T) was used.

The study shows that the population of Glycaspis

was significantly higher from late May to early Au-

gust in all regions, coinciding with low precipita-

tions (less than 50 mm) and temperate temperatures

(15 to 25°C) (Fig. 2).

Despite reported damage caused by the species

on Eucalyptus around the world, our observations

have not revealed any negative effects so far,

Glycaspis brimblecombei Moore, / 964 (Hemiptera Psyllidae) invasion and new records in Mediterranean area 503

Figure 1 . Distribution of Glycaspis brimblecombei.

6000

5000

4000

3000

2000

1000

Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

* Glycaspis

8 .

Figure 2. Evolution of the population of Glycaspis brimblecombei with temperature and precipitation in Algeria.

though it should be noted that further research is re-

quired. Although our research has not identified any

significant effect on the trees’ health-status, the at-

tack of this sucking insect can indeed leave large

quantities of honeydew on leaves, facilitating sub-

sequent attacks by fungi resulting in fiimagina syn-

dromes. Moreover, when trees support high popu-

lation levels, some discoloration of leaves or, at

least, the adoption of a yellowish green colour has

been detected. In fact, as time progresses, these

fungi attacks produce leaf discoloration, falling,

stunted growth and general abatement of plant

504

Kheddar Reguia & Francisco Javier Peris-Felipo

vigour. This general deterioration could facilitate

further attacks by other insect pests or the death of

branches or even whole trees (Bouvert et al., 2005;

Ide et al., 2006; Hurtado & Reina, 2008).

As a result, biological control programs using the

natural wasp enemy Psyllaephagus bliteus has been

used in Australia and California (Riek, 1 962) and

thus knowing the distribution of this encyrtid be-

comes relevant in the case of a pest. In Europe, this

parasitoid wasp was discovered for the first time in

different areas from Sicily (Italy) (Caleca et al.,

2011) and Huelva (Spain) (Perez-Otero et al., 201 1)

and this paper provides the first record in Algeria.

Once known Glycaspis biology, distribution and

its expansion, next step was trying to answer the hy-

pothesis about the means used for its dispersion.

Countries connections were analysed finding that

there were motorways and waterways linking all

countries were Glycaspis was found (Figs. 3, 4).

On one hand, waterways data analyses demon-

strate that Sea connection joins all Mediterranean

countries were Glycaspis is present (Fig. 3). Check-

ing carefully sea lanes it is possible to observe that

there are numerous good routes from the Portuguese

and Spanish area to other Spanish regions (i.e. Ba-

learic Islands), areas (i.e. Sardinia) or countries (i.e.

France, Greece, Italy or Algeria). On the other hand,

motorways networks (Fig. 4) allow a good connec-

tion between Portugal, Spain, France and Italy

thanks to the Mediterranean motorway. However,

there is no connection with Greece, indicating that

Glycaspis might have reached this country by the

Mediterranean Sea. A similar situation could be ap-

plied to Italy, as the evolution of G. brimblecombei

expansion shows it arrived to this area after being

found in the Iberian Peninsula. However, in the cen-

tral areas of Spain and Italy this possibility disap-

pears and therefore a land-based expansion would

be more feasible.

Despite that both motor and waterways are

good candidates for explaining Glycapsis distribu-

tion, there is a lack of knowledge about the species

biology and thus it is not possible to ignore ways

of dispersion used by other arthropods such as the

Aphids, which rely on the wind to colonize other

trees. These data suggest that Glycaspis expansion

was done via sea and land: the maritime route

would have dominated in coastal areas while the

land route would have encouraged its dispersion

to offshore areas.

The rapid colonization of Mediterranean coun-

tries by G. brimblecombei demands the conduction

of organic studies aimed at obtaining a better

knowledge of its distribution, population character-

istics, possible impact and potential natural ene-

mies. Once these aspects are clarified, appropriated

Figures 3, 4. Waterways (Fig. 3) and motorways (Fig. 4) connections in Mediterranean countries with Glycaspis.

Glycaspis brimblecombei Moore, / 964 (Hemiptera Psyllidae) invasion and new records in Mediterranean area 505

control measures should be adopted to prevent sig-

nificant damage on trees and economic loss.

REFERENCES

Bouvet J.P.R., Harrand L. & Burckhardt D., 2005.

Primera cita de Blastopsylla occidentalis y Gly-

caspis brimblecombei (Hemiptera: Psyllidae) para la

Republica Argentina. Revista de la Sociedad Ento-

mologica Argentina, 64: 99-102.

Brennan E.B., Gill R.J., Hrusa G.F. & Weinbaum S.A.,

1999. First record of Glycaspis brimblecombei

(Moore) (Homoptera: Psyllidae) in North America:

initial observations and predator associations of a po-

tentially serious new pest of Eucalyptus in California.

Pan-Pacific Entomologist, 75: 55-57.

Brennan E.B., Hrusa G.F., Weinbaum S.A. & Levison Jr

W., 2001. Resistance of Eucalyptus species to Gly-

caspis brimblecombei (Homoptera: Psyllidae) in the

San Francisco Bay area. Pan-Pacific Entomologist,

77: 249-253.

Burckhardt D., Eozada PW. & Diaz B.W., 2008. First

record of the red gum lerp psyllid Glycaspis brim-

blecombei (Hemiptera: Psylloidea) from Peru.

Mitteilungen der Schweizerischen Entomologische

Gesellschaft, 81: 83-85.

Caleca V., Fo Verde G. & Maltese M., 2011. First record

in Italy of Psyllaephagus bliteus Riek (Hymenoptera

Encyrtidae) parasitoid of Glycaspis brimblecombei

(Hemiptera Psyllidae). II Naturalista siciliano, 35:

435-444.

Castillo J.V., 2003. Condicion de insectos forestales en

Mexico. In: Proceedings of the 1st Joint Meeting of

the 12th National Symposium on Forest Parasites and

the 54th Eastern Forest Insect Work Conference,

Guadalajara, Mexico, p. 61

Cocquempot C., Malausa J.C., Thaon M. & Brancaccio

F., 2012. The Red Gum Ferp Psyllid ( Glycaspis

brimblecombei Moore) introduced don French eu-

caliptus trees (Hemiptera, Psyllidae). Bulletin de la

Societe Entomologique de France, 117: 363-370.

Hidalgo F.O., 2005. Evaluacion de la preferencia de Gly-

caspis brimblecombei Moore por diversos hospeda-

dores del genero Eucalyptus F’Herit. en la region

metropolitana, Chile. Universidad de Chile, Chile,

40 pp.

Hollis D., 2004. Australian Psylloidea: Jumping plant lice

and lerps insects. Australia Biological Resources

Study, Canberra, XVI + 216 pp.

Hurtado A. & Reina I., 2008. Primera cita para Europa

de Glycaspis brimblecombei Moore (Hemiptera:

Psyllidae), una nueva plaga del eucalipto. Boletin

Sociedad Entomologica Aragonesa, 43: 447-449.

Ide M.S., Munoz C.A., Beeche M.C., Moncada J.E.,

Jacques F.R., Gonzalez P.E. & Goycoolea C.P.,

2006. Deteccion y control biologico de Glycaspis

brimblecombei Moore (Hemiptera: Psyllidae). Sub-

departamento de Vigilancia y Control de Plagas Fore-

stales y Exoticas Invasoras. Gobierno de Chile.

Ministerio de Agricultura. S.A.G., Chile, 32 pp.

Faudonia S. & Garonna A.P, 20 10. The red gum lerp

psyllid, Glycaspis brimblecombei , a new exotic pest

of Eucalyptus camaldulensis in Italy. Bulletin of In-

sectology, 63: 233-236.

Maatouf N. & Fumaret J.P., 2012. Eco-ethology of new

invasive pest species on eucaliptus plantation of Mo-

rocco. Annales de la Societe Entomologique de

France, 48: 289-297.

Moore K.M.,1964. Observations on some Australian for-

est insects. 18. Four new species of Glycaspis (Ho-

moptera: Psyllidae) from Queensland. Proceedings

of the Finnean Society of New South Wales, 89:

163-166.

Moore K.M., 1970. Observations on some Australian for-

est insects. 23. A revision of the genus Glycaspis (Ho-

moptera: Psyllidae) with descriptions of seventy-three

new species. Australian Zoologist, 15: 248-341.

Nagamine W.T & Heu R.A., 2001. Red gum lerp psyllid

Glycaspis brimblecombei Moore (Homoptera:

Psyllidae). New Pest Advisory, 01-02. (http:// ha-

waii.gov/hdoa/pi/ppc/npa-l/npa01-02_rpsyllid.

pdf)

Onore G. & Gara R.F., 2007. First record of Glycaspis

brimblecombei (Hemiptera: Psyllidae) in Ecuador,

biological notes and associated fauna. Extended Ab-

stracts of the 4th European Hemiptera Congress

Ivrea, Turin, Italy, 10-14 September 2007, 41-42.

Perez-Otero, R., Borrajo, P, Mansilla J.P. & Ruiz, F.,

2011. Primera cita en Espana de Psyllaephagus bli-

teus Riek (Hymenoptera, Encyrtidae), parasitoide de

Glycaspis brimblecombei Moore (Hemiptera, Psylli-

dae). Boletin de Sanidad Vegetal Plagas, 37: 37-44.

Peris-Felipo F.J., Bernues-Baneres A., Perez-Faorga

Arias E. & Jimenez-Peydro R., 2010. Nuevos datos

sobre la distribucion en Espana de Glycaspis brim-

blecombei Moore, 1964 (Hemiptera: Psyllidae), plaga

de Eucalyptus camaldulensis. Boletin Asociacion

espaiiola Entomologia, 33: 517-526.

Prieto-Fillo E., Rueda J., Hernandez R. & Selfa J., 2009.

Primer registro del psilido rojo del eucalipto, Gly-

caspis brimblecombei (Homoptera: Psyllidae), en la

Comunidad Valenciana. Boletin Sanidad Vegetal

Plagas, 35:277-281.

Riek E.F., 1962. A new encyrtid (Hymenoptera, Chalci-

doidea) genus of parasites of lerp-forming psyllids

on Eucalyptus. Proceedings of the Finnean Society

of New South Wales, 87: 283-285.

506

Kheddar Reguia & Francisco Javier Peris-Felipo

Rosales C.J., Lobosque O., Carvalho R, Bermudez L. &

Acosta C., 2008. Glycaspis brimblecombei Moore

(Hemiptera: Psyllidae). “Red Gum Lerp”: nueva

plaga forestal en Venezuela. Entomo tropica, 23:

103-104.

Sandoval A. & Rothmann S., 2003. Deteccion del psilido

de los eucaliptos rojos, Glycaspis brimblecombei

Moore (Hemiptera: Psyllidae) en Chile. In: 24 Con-

greso Nacional de Entomologia, Resumenes.

(http://www.udec.cl/insectos/res.menes.html)

Santana D.L.Q., Mendezes A.O., Silva H.D., Bellote

A.F.J. & Favaro R.M., 2003. O Psilideo de concha

(' Glycaspis brimblecombei ) em eucalipto. Comunica-

gao Tecnica, 105.

Sookar P., Seewooruthun S.I. & Ramkhelawon D., 2003.

The red gump lerp psyllid, Glycaspis brimblecombei ,

a new pest of Eucalyptus sp. in Mauritius. AMAS,

Food and Agricultural Research Council, R.duit,

Mauritius.

Valente C. & Hodkinson I. 2009. First record of the red

gum lerp psyllid

, Glycaspis brimblecombei Moore (Hem: Psyllidae), in

Europe. Journal of Applied Entomology, 133: 315-

317.

Biodiversity Journal, 2013, 4 (4): 507-552

Monograph

First report on the herpetofauna of TayYenTu Nature Re-

serve, northeastern Vietnam

Vera L. Hecht 1 , CuongT. Pham 2 ,TaoT. Nguyen 3 , Truong Q. Nguyen 2,4 , Michael Bonkowski 4 & Thomas

Ziegler 1,4 *

*AG Zoologischer Garten Koln, Riehler Strasse 173, - 50735 Cologne, Germany; e-mail: ziegler@koelnerzoo.de

institute of Ecology and Biological Resources, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet Road, Hanoi,

Vietnam; e-mail: nqt2@yahoo.com

3 Vietnam National Museum of Nature, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet Road, Hanoi, Vietnam;

e-mail: nguyenthientao@gmail.com

4 Zoological Institute, University of Cologne, Ziilpicher Strasse 47b, D-50674 Cologne, Germany; email: m.bonkowski@uni-koeln.de

’Corresponding author

ABSTRACT A total number of 76 species of amphibians and reptiles were recorded during recent field

surveys from the Tay Yen Tu Nature Reserve in Bac Giang Province, northeastern Vietnam,

comprising one caecilian species, one newt species, 34 species of anurans, 18 species of

lizards, and 22 species of snakes. Thirty species are reported for the first time from Yen

Tu Nature Reserve as well as for Bac Giang Province. Among the recorded species, five

are currently known only from Vietnam. A high level of species diversity and endemism

of the herpetofauna underlines the importance of biodiversity conservation in this nature

reserve, which covers a major part of the remaining lowland evergreen forest in northea-

stern Vietnam.

KEY WORDS Amphibians; Bac Giang Province; diversity; new records; reptiles.

Received 24.10.2013; accepted 30.11.2013; printed 30.12.2013

INTRODUCTION

The Tay Yen Tu Nature Reserve is located in

Bac Giang Province, about 100 kin northeast of

Hanoi (Tordoff et al., 2004; Ha et al., 2010). This

nature reserve is situated in the western side of the

Yen Tu massif, which is known as the largest gran-

ite formation in northeastern Vietnam (Ha et al.,

2010; Fig. 1). The main habitat type of the Yen Tu

massif is evergreen broad-leaved tropical forest

(Averyanov et al., 2003; Tordoff et al., 2004; Fig.

2). The total area of the Tay Yen Tu Nature Reserve

comprises 16,466 ha and includes two non-con-

tiguous sectors: the Thanh Son-Luc Son sector and

the Khe Ro sector. The Tha nh Son-Luc Son sector

is centered on the 1,068 m high Mount Yen Tu,

whereas the Khe Ro sector is allocated around the

886 m high Mount Da Bac (Tordoff et al., 2004).

A comprehensive study on the biodiversity has

not been conducted in Tay Yen Tu Nature Reserve

to date, however, the results of preliminary field re-

search have indicated that this area supports a num-

ber of species of national or global conservation

concern (Ha et al., 2010). Four new species have

been discovered from Yen Tu Mountain in the last

decade: Sphenomorphus cryptotis Darevsky, Orlov

et Cue, 2004, Scincella devorator Darevsky, Orlov

et Cue, 2004 (Darevsky et al., 2004), Tylototriton

508

Vera L. Hecht et alii

vietnamensis Bohme, Schottler, Nguyen et Kohler,

2005 (Bohme et al., 2005), and Odorrana yentuensis

Tran, Orlov et Nguyen, 2008 (Tran et al., 2008). In

addition, three new country records were recently

reported from this nature reserve: Shinisaurus

crocodilurus Ahl, 1930, Amphiesmoides ornaticeps

(Werner, 1924), and Rhacophorus maximus

Gunther, 1858 (Le & Ziegler, 2003; Nguyen et al.,

2008; Nguyen et al., 2010a). The discoveries of new

reptiles and amphibians from Yen Tu Mountains un-

derscore the unrealized biodiversity of northeastern

Vietnam. Based on the results of our recent field sur-

veys in the period between 2008 and 2010, we

herein provide the first list of amphibians and repti-

les recorded from Tay Yen Tu Nature Reserve.

MATERIALS AND METHODS

Field surveys in the Tay Yen Tu Nature Reserve

took place in April 2008, in May and October 2009,

and from April to July 2010. Survey transects were

set up along streams, ponds, and forest paths within

both sectors of the nature reserve. The habitat sur-

rounding the ponds consisted of bamboo forest or

mixed forest of bamboo and secondary forest; the

streams were located in disturbed primary forest or

secondary low-land forest (Figs. 3-5). In the Thanh

Son - Luc Son Sector, the following streams were

surveyed: Suoi Tuyen I (= stream 2); Suoi Tuyen II

(= stream 1); Khe cam 1 (= stream 3); stream 4;

stream 5; Ba Bep Stream (= stream 6). In addition,

the surroundings of the following ponds were sur-

veyed: Ao Cua; Khe Cam 1; Khe Cam 3; Deo Gio;

Ba Bep; May Klioan; Da Lua 1; Da Lua 2; Lai Am

1 and Lai Am 2. In the Khe Ro Sector, night excur-

sions were conducted in the forest near the Vung

Tron Ranger Station; further surveys took place

nearby Dong Ri and Dong Thong Ranger Stations.

Surveyed sites are situated at elevations between 75

and 600 m above sea level.

Specimens collected during field surveys were

deposited in the museum collections. Taxonomic

identifications were made following Boulenger

(1912), Pope (1935), Smith (1935, 1943), Bourret

(1936, 1942, 2009), Manthey & Grossmann

(1997), Ziegler (2002), Bain & Nguyen (2004a, b),

Darevsky et al. (2004), Tran et al. (2008), Vogel et

al. (2009), Nguyen et al. (2011a, b), and Nguyen

et al. (2012). Ventral scales of snakes were counted

according to Dowling (1951). Systematics and

nomenclature generally followed Nguyen et al.

(2009) and Zaher et al. (2009). The gender was

identified based on external sexual characters, and

if required, from the inspection of the gonads after

dissection.

ABBREVIATIONS. Abbreviations of collec-

tors. NQT: T.Q. Nguyen; NTT: T.T. Nguyen; NVS:

S. V. Nguyen; PTC: C.T. Pham; TTT: T.T. Tran; TZ:

T. Ziegler; VH: V.L. Hecht.

Collections. Institute of Ecology and Biological

Resources (IEBR), Hanoi, Vietnam; Vietnam Na-

tional Museum of Nature ( VNMN), Hanoi, Viet-

nam; Vinh Phuc College of Education, Vietnam

(TYT numbers); and Zoologisches Forschungsmu-

seum Alexander Koenig (ZFMK), Bonn, Germany.

Others abbreviations. NR: nature reserve; asl:

above sea level; ED: horizontal eye diameter (eye

length); EN: distance between anterior corner of

eye and nostril; ET: distance between posterior cor-

ner of eye and tympanum; HL: head length; HW:

head width; IN: internarial distance; IntOrb: in-

terorbital distance; NtoS: distance between nostril

and tip of snout; SL: snout length; SVL: snout-vent

length (from tip of snout to cloaca); TaL: tail length

(from posterior margin of cloaca to tip of tail); TD:

maximum tympanum diameter; TiL: tibia length;

TiW: tibia width; uEL: width of upper eyelid.

RESULTS

SYSTEMATICS

AMPHIBIA

GYMNOPHIONA

ICHTHY OPHIID AE

Ichthyophis bannanicus Yang, 1984

Examined material. One adult male VNMN

1359 (SVL 233 mm), collected by NTT, 10 July

2010, 450 m asl (Fig. 6).

Morphological characters. Tail length 2.42

mm, tail width 2.98 mm, tail tip pointed; number of

annuli: total annuli 365, annuli interrupted by vent

4, post- vent annuli 5. Colouration in life: dorsal sur-

face of head, body and tail lilac; lateral stripe bright

yellow, broad, interrupted posteriorly, extending

from behind tentacle on upper jaw to posterior end

First report on the herpetofauna of TayYen Tu Nature Reserve, northeastern Vietnam

509

CHINA

LAOS

1 8 °N

4 5

Figure 1. Map showing the Tay Yen Tu Nature Reserve in Bac Giang Province, Northern Vietnam. Figure 2. Vegetation

type of the Tay Yen Tu Nature Reserve. Figures 3-5. Habitat types in Tay Yen Tu Nature Reserve. Photos by T. Ziegler

and C.T. Pham.

510

Vera L. Hecht et alii

of vent; ventral surface bright lilac (determination

after Nishikawa et al., 2012).

Distribution. This is a widespread species in

northern Vietnam, from Cao Bang and Tuyen

Quang provinces southwards to Hoa Binh Prov-

ince. I. bannanicus was previously recorded from

the eastern side of the Yen Tu Mountain in Hai

Duong Province (Nguyen et al., 2009). Our find-

ing represents the first record of this species from

Tay Yen Tu NR and from Bac Giang Province.

Elsewhere, the species has been recorded from

China and Laos (Nguyen et al., 2009; Nishikawa

et al., 2012).

Remarks. The specimen was found at night on

the forest path near Ba Bep Stream in a bamboo for-

est near Mau Village.

CAUDATA

SALAMANDRIDAE

Tylototriton vietnamensis Bohme, Schottler,

Nguyen et Kohler, 2005

Examined material. Two adult males IEBR

A. 2013. 57 (SVL 53 mm, TaL 55 mm), IEBR

A.2013.58 (SVL 62 mm, TaL 65 mm) collected by

PTC, June 2010 and July 2012, 200-500 m asl

(Figs. 7, 8).

Morphological characters. Habitus moder-

ately stout; head broader than body, slightly sloping

in profile; snout short, truncate in dorsal view,

rounded in profile and protruding beyond lower

jaw; nostrils close to snout tip; upper lips thick,

fleshy and overlapping lower lip under eye region;

vomeropalatine teeth in two rows; tongue with poorly

developed tongue pad, lacking a free posterior mar-

gin; a low vertebral tubercular ridge, extending

from top of head to base of tail; two lateral rows of

larger warts, extending from insertion of fore arms

to base of tail; glands and warts relatively small, cov-

ering most of dorsal surfaces; venter almost smooth;

parotoids greatly enlarged, slightly projecting back-

wards; gular fold absent; fingers without webbing,

toes with basal webbing; tail laterally compressed;

dorsal and ventral tail fin moderately developed; tail

tip acuminate in profile. Colouration in life: dorsal

and ventral surfaces brownish tan; finger and toe

tips cream coloured; tail brownish tan with ventral

tail fin creamy yellow; cloacal region bordered with

cream-yellow (determination after Bohme et al.,

2005; Nishikawa et al., 2013).

Distribution. This species is currently known

only from Lang Son, Bac Giang, and Phu Tho prov-

inces in northern Vietnam (Nguyen et al., 2009;

Nishikawa et al., 2013).

Remarks. Eggs, larvae, subadults and adults

were found in and around diverse ponds during

day and night in secondary forests (Bernardes et

al., 2013).

ANURA

BUFONIDAE

Duttaphrynus melanosticus (Schneider, 1799)

Examined material. Identification was based

on direct observation in the field and photographs

(Fig. 9): cranial crests conspicuous, black and

more distinct on supraorbital region; parietal crest

absent; parotoid gland prominent, elongated;

tympanum distinct, rounded; dorsum and upper

surface of limbs with conical, spiny warts; warts

more rounded and smaller on flanks; ventral sur-

face granular; fingers free of webbing, toes with

webbing at base. Colouration in life: upper head

and dorsum yellowish grey to dark brown with

black spines; ventral cream (determination after

Bourret, 1942; Ziegler, 2002).

Distribution. This is a common species in

Vietnam as well as in Asia (Nguyen et al., 2009).

Therefore, we did not collect any voucher speci-

mens.

Remarks. Several individuals were found during

the day and in the evening in the grassland near the

Dong Thong Ranger Station.

Ingerophrynus galeatus (Gunther, 1864)

Examined material. One subadult female

ZMFK 92837 (SVL 39.04 mm), collected by TZ

and NTT, October 2009, ca. 400 m asl (Fig. 10).

Morphological characters. Snout short, ob-

tuse, protruding in profile; pupil horizontal; vomer-

ine teeth absent; canthus rostralis with well

developed bone-crest, continuing on supraorbital

and parietal region, parietal crest absent; interorbital

First report on the herpetofauna of TayYen Tu Nature Reserve, northeastern Vietnam

511

distance narrower than upper eyelid width (IntOrb

3.23 mm, uEL 3.67 mm); a well-developed bone

crest beyond eye above tympanum, reaching oval

parotoid gland; tympanum distinct, smaller than eye

length (TD 1.85 mm, ED 4.04 mm); dorsal skin

with warts, ventral skin granular; finger I longer

than II; toes 1/3- 1/2 webbed; two metatarsal tuber-

cles present; tarsal fold absent; tibio-tarsal articula-

tion reaching eye. Colouration in life: dorsum grey

with reddish warts and brown to dark grey mar-

bling; lips with some bars; limbs with dark trans-

verse bands, ventral surface pale yellowish grey

with some dark markings (determination after

Bourret, 1942; Inger et al., 1999; Ziegler, 2002;

Tran et al., 2010).

Distribution. In Vietnam, this species has

been recorded from Lao Cai in the North to Dak

Lak, Lam Dong, and Dong Nai provinces in the

South. This is the first record of I. galeatus from

Tay Yen Tu NR as well as from Bac Giang Prov-

ince. Elsewhere, the species has been reported

from China, Laos, and Cambodia (Nguyen et al.,

2009).

Remarks. A single specimen was found in the

late afternoon on the forest ground near Suoi Tuyen

I. Another juvenile was encountered on the forest

ground near Ba Bep Pond in the afternoon on 3

June 2010.

MEGOPHRYIDAE

Leptobrachium cf. chapaense (Bourret, 1937)

Examined material. One adult female ZFMK

92838 (S VL 76.55 mm), collected by TZ and NQT,

27 May 2009, ca. 350 m asl (Fig. 11).

Morphological characters. Head large,

flattened; snout round; pupil vertical; upper part

of iris white; vomerine teeth absent; supratympa-

nic fold distinct; tympanum indistinct (TD 4.69

mm); fore limbs long and slender, fingers free of

webbing; webbing of toes rudimental; inner

metatarsal tubercle present, outer metatarsal tu-

bercle absent; tibio-tarsal articulation reaching

tympanum; tibia length 26.72 mm; dorsal and

ventral skin smooth. Colouration in life: dorsum

reddish to greenish brown with some irregular

black spots and marbling, flanks dark brown with

white spots, upper surface of limbs lighter with

dark bars; throat light reddish-brown, finely speck-

led with white, venter blackish grey with white

spots (determination after Bourret, 1942; Ziegler,

2002; Bain & Nguyen, 2004a).

Distribution. This species has been recorded

from northern and central Vietnam. This is the first

record of L. chapaense from Tay Yen Tu NR as well

as from Bac Giang Province. Elsewhere, the species

is known from China, Myanmar, Laos, and Thai-

land (Nguyen et al., 2009).

Remarks. ZFMK 92838 was found on the

path in the bamboo forest near Vung Tron Ranger

Station. In October 2009, a tadpole of L. cf. cha-

paense was collected in the shallow part of a

small stream.

Leptolalax nyx Ohler, Wollenberg, Grosjean,

Hendrix, Vences, Ziegler et Dubois, 2011

Examined material. One adult female IEBR

3659 (SVL 33.94 mm), collected by NQT, 10 April

2008, ca. 350 m asl (Fig. 12).

Morphological characters. Head longer than

wide (HL 11.27 mm, HW 10.71 mm); vomerine

teeth present; snout longer than eye (SL 5.75 mm,

ED 4.61 mm); interorbital distance slightly wider

than upper eyelid width (IntOrb 4.09 mm, uEL 3.91

mm); nostrils oval, closer to tip of snout than to eye

(NtoS 2.49 mm, EN 3.26 mm); tympanum distinct

round, greater than distance between tympanum

and eye (TD 3.32 mm, ET 2.44 mm); supratym-

panic fold distinct; small webbing on feet and nar-

row fringes on toes; dermal ridges under toes poorly

developed; finger tips slightly enlarged; dorsum

with glandular warts and short elongate ridges.

Colouration in life: dorsal pattern distinct including

dark outlines on warts and folding, a light W-shaped

brown marking in the shoulder region; dark spots

on flanks present; ventral surface whitish (determi-

nation after Ohler et al., 2011).

Distribution. This species was previously

known only Rom the type locality in Ha Giang Prov-

ince, Vietnam (Ohler et al., 2011). This is the first

record of the species from Tay Yen Tu NR as well

as from Bac Giang Province.

Remarks. Specimen was collected at night on

a tussock grass in a rocky stream near Dong Ri

Ranger Station.

512

Vera L. Hecht et alii

Ophryophryne microstoma Boulenger, 1903

Examined material. Two adult males ZFMK

92839 and ZFMK 93906, collected by TZ and

NQT, 26 May 2009, elevation 250 m asl; one adult

female ZFMK 92840, collected by TZ and NTT,

October 2009, 250-370 m asl; two adult males

VNMN 1320 (SVF 34.72 mm) and VNMN 1340

(SVF 33.99 mm), collected by TZ and NTT, June

2010, 400-600 m asl (Fig. 13).

Morphological characters. Relatively large

Ophryophryne , SVF 33.99-46.3 mm; head small;

snout shorter than eye (SF 2.90-3.64 mm, ED

3.69-4.49 mm), obliquely obtuse, strongly pro-

nounced; loreal region vertical, concave; nostril

closer to tip of snout than to eye (EN 1.79-2.15

mm, NtoS 1.16-1.6 mm); interorbital distance as

broad as upper eyelid; conical dermal appendix

on upper eyelid present; eye larger than tympa-

num (ED 3.69-4.49 mm, TD 2.04-2.37 mm);

tympanum distinct; arms and legs slender; palmar

tubercle indistinct; subarticular tubercles indis-

tinct; finger I shorter than II; tibio-tarsal articula-

tion reaching eye or at least armpit; tibia about

four times longer than wide (TiF 15.77-22.05

mm, TiW 3.48-4.81 mm); toe webbing rudimen-

tal; dorsal skin with small warts, with symmetric

glandular ridges; males without externally visible

vocal sacs. Colouration in life: dorsum greyish or

light brown, upper surface of limbs with trans-

verse bars; a light bar present below the eye;

flanks with some small black spots; ventral sur-

face cream with dark marbling (determination

after Bourret, 1942; Ohler, 2003).

Distribution. In Vietnam, this species has

been recorded from Fao Cai and Ha Giang in

the North southwards to Dak Fak and Earn Dong

provinces. This is the first record of O. micros-

toma from Tay Yen Tu NR as well as from Bac

Giang Province. Elsewhere, the species is known

from China, Faos, Thailand, and Cambodia

(Nguyen et al., 2009).

Remarks. All five specimens were found at

night on branches, ferns or stones near rocky

streams. Ohler (2003) stated that vocal sacs are vi-

sible in the males of O. microstoma , however, they

are indistinct in our specimens.

Xenophrys major (Boulenger, 1908)

Examined material. One adult male VNMN

1345 (SVF 77.23 mm), collected by TZ and NTT,

24 June 2010, 300 m asl.

Morphological characters. Head flat, slightly

wider than long (HE 27.81 mm, HW 28.75 mm);

snout obliquely obtuse, strongly pronounced;

pupil vertical; tympanum distinct, approximately

half of eye diameter (TD 3.52 mm, ED 8.23 mm);

supratympanic fold distinct; nostrils closer to eye

than to tip of snout (EN 4.61 mm, NtoS 5.10 mm);

interorbital distance a bit narrower than upper

eyelid width (IntOrb 6.93 mm, uEL 7.17 mm);

tips of fingers and toes swollen; finger I longer

than II; toes 1/4 webbed; tibio-tarsal articulation

reaching the position between eye and tympanum;

tibia length (TiL 39.27 mm), half of snout-vent

length; dorsal skin nearly smooth, partly granular,

glandular ridges on back and upper surface of

limbs present; supratympanic fold present. Coloura-

tion in life: dorsal head and body brown, a dark

brown triangular pattern present between the

eyes; lateral sides of head dark brown; upper lip

with a white stripe, running from nostril backward

to shoulder; supratympanic fold edged in white;

dorsal surface of limbs with dark bars; ventral sur-

face white (determination after Bourret, 1942;

Tran et al., 2010).

Distribution. This is a widespread species in

Vietnam, from Lao Cai and Ha Giang provinces

southwards to Dak Lak and Dong Nai provinces.

Elsewhere, the species has been reported from

India, Bangladesh, China, Myanmar, Laos, and

Thailand (Nguyen et al., 2009).

Remarks. The single specimen was found at

night on a dry and loamy forest path.

MICROHYLIDAE

Kalophrynus interlineatus (Blyth, 1854)

Examined material. One adult female ZFMK

92841 (SVL 46.61 mm), collected by TZ and NTT,

October 2009, 160-180 m asl (Fig. 14).

Morphological characters. Snout pointed,

slightly pronounced, somewhat longer than eye (SL

5.30 mm, ED 4.11 mm); vomerine teeth absent;

First report on the herpetofauna of TayYen Tu Nature Reserve, northeastern Vietnam

513

Figure 6. Ichthyophis bannanicus. Figures 7, 8. Larvae and adult of Tylototriton vietnamensis . Figure 9. Duttaphrynus

melanostictus. Figure 10. Ingerophrynus galeatus. Figure 11. Leptobrachium cf. chapaense. Figure 12. Leptolalax nyx.

Figure 13. Ophryophryne microstoma. Photos by T. Ziegler and T.Q. Nguyen.

514

Vera L. Hecht et alii

pupil horizontal; interorbital distance broader than

upper eyelid (IntOrb 4.42 mm, uEL 2.99 mm); tym-

panum distinct, about 2/3 of eye length (TD 2.59

mm, ED 4.11 mm); fingers free of webbing; toes

rudimentally webbed; subarticular tubercles distinct;

inner and outer metatarsal tubercles present; tibio-

tarsal articulation reaching to the middle position

between axilla and groin; dorsal skin with tubercles,

larger in size on belly and on upper surface of thighs;

supratympanic fold indistinct; dorsolateral fold nar-

row and granular. Colouration in life: dorsal head

and body light brown, one black spot present on

each side of the hips; flanks dark brown; ventral sur-

face whitish (determination after Bourret, 1942).

Distribution. In Vietnam, this species has been

reported from Lang Son, Phu Tho, Bac Giang, Ninh

Binh, Quang Binh, Dong Nai, and Kien Giang

provinces. Elsewhere, the species is known from

India, China, Myanmar, Laos, Thailand, Cambodia,

and Indonesia (Nguyen et al., 2009).

Remarks. The female specimen was found in the

late afternoon, on the ground near pond Khe Cam 1

in a bamboo forest. Our specimen differs from Bour-

ret’s (1942) description by its longer snout (versus

a snout as long as eye) and the absence of black mar-

king in the loreal region, under the dorsolateral fold,

in the cloacal region and at the back of thigh.

Microhyla butleri Boulenger, 1900

Examined material. One adult male VNMN

1326 (SVL 22.39 mm), collected by TZ and NTT,

July 2010, ca. 400 m asl; one adult female IEBR

A. 2013. 59 (SVL 25.55 mm), collected by VH, 3

July 2010, 450 m asl (Fig. 15).

Morphological characters. Snout rounded,

pronounced, longer than eye (SL 3.14-3.55 mm,

ED 1.68-2.02 mm); vomerine teeth absent; pupil

horizontal; interorbital distance 1.9-2. 6 times

broader than upper eyelid (IntOrb 2.48-2.85 mm,

uEL 0.96-1.47 mm); tympanum invisible; fingers

free of webbing, with slightly developed discs; toes

with small discs, webbed at base; subarticular tu-

bercle prominent; inner and outer metatarsal tuber-

cles present; tibio-tarsal articulation reaching

between eye and tip of snout; dorsal skin smooth;

supratympanic fold present; ventral skin smooth,

cloacal region granular; males with visible vocal

sacs. Colouration in life: dorsal head and body grey

with brownish and reddish markings; one whitish

stripe from eye to beginning of arm; limbs with

dark transverse bars; ventral surface whitish, throat

and chest mottled with dark brown (determination

after Bourret, 1942; Manthey & Grossmann, 1997;

Ziegler, 2002; Bain & Nguyen, 2004b).

Distribution. This is a widespread species in

Vietnam. Elsewhere, the species has been reported

from China, Myanmar, Laos, Thailand, Cambodia,

Malaysia, and Singapore (Nguyen et al., 2009).

Rjemarks. VNMN 1326 was found on a stone

covered by moss in Suoi Tuyen I at night while

IEBR A.2013.59 was collected on the ground of the

bamboo forest near Lai Am Pond in the evening.

Microhyla heymonsiV ogt, 1911

Examined material. One adult male VNMN

1327 (SVL 20.23 mm) collected by TZ and NTT, 4

June 2010, 440 m asl; one adult male IEBR

A. 2013. 60 (SVL 21.21 mm) collected by VH, 7

June 2010, ca. 400 m asl (Fig. 16).

Morphological characters. Snout obtusely

pointed, pronounced, somewhat longer than eye

(SL 2.84-2.90 mm, ED 2.08-2.30 mm); vomerine

teeth absent; interorbital distance 1.2-1. 5 times

broader than upper eyelid (IntOrb 2.06-2.30 mm,

uEL 1.54-1.72 mm); tympanum hidden; fingers

free of webbing, with small discs; finger I shorter

than II; palmar tubercle present; toes webbed rudi-

mentally; subarticular tubercles well developed;

inner and outer metatarsal tubercles small; tibio-tarsal

articulation reaching between eye and tip of snout;

dorsal and ventral skin smooth; supratympanic fold

indistinct. Colouration in life: dorsal head and body

grey or light brown, with a white stripe from tip of

snout to cloaca, and a small dark spot in the center

of the back; lateral sides of head and flanks dark

brown to black; anterior part of thighs, cloacal re-

gion and lower parts of feet black; limbs with thin

transverse bars; ventral surface white to grey; males

with black vocal sacs (determination after Bourret,

1942; Manthey & Grossmann, 1997).

Distribution. This is a widespread species in

Vietnam. However, this is the first record of M. hey-

monsi from Tay Yen Tu NR as well as from Bac

Giang Province. Elsewhere, the species has been

recorded from India, China, Laos, Thailand, Cambo-

dia, Malaysia, and Indonesia (Nguyen et al., 2009).

First report on the herpetofauna of TayYen Tu Nature Reserve, northeastern Vietnam

515

Remarks. Specimens were found during the day

time or at night on the ground near small ponds in

the bamboo forest.

Microhyla pulchra (Hallowell, 1861)

Examined material. One subadult ZFMK

93905 (SVL 17.58 mm), collected by VH, 8 June

2010, 75 m asl (Fig. 17).

Morphological characters. Snout obtusely

pointed, slightly pronounced, longer than eye (SF

2.52 mm, ED 2.04 mm); vomerine teeth absent;

pupil round; interorbital distance broader than

upper eyelid (IntOrb 1.84 mm, uEF 1.41 mm);

tympanum indistinct; tips of fingers not enlarged;

fingers free of webbing, finger I shorter than II;

toes almost 1/2 webbed; subarticular tubercles

well developed; metatarsal tubercle large; tibio-

tarsal articulation reaching between eye and tip of

snout; dorsal skin smooth; a distinct fold present

between posterior edges of the eyes; ventral skin

smooth; cloacal region granular. Colouration in

life: dorsum light brown, with a dark brown A-

shaped pattern on the back, containing several

dark and light lines, outer part bordered by several

light lines; canthus rostralis and flanks dark

brown; limbs with transverse bars; groin and an-

terior part of thigh yellow; ventral surface whitish

yellow, chin and throat with black marbling (de-

termination after Bourret, 1942; Ziegler, 2002;

Bain & Nguyen, 2004b).

Distribution. This is a common species in Viet-

nam. Elsewhere, the species is known from India,

China, Faos, Thailand, and Cambodia (Nguyen et

al., 2009).

Remarks. A single specimen was found in the

morning on grassland near the Dong Thong Ranger

Station. Our specimen well matched the descrip-

tions of Bourret (1942) and Ziegler (2002), but it

differs in having only one metatarsal tubercle in-

stead of two small ones.

DICROGFOSSIDAE

Fejervarya limnocharis (Gravenhorst, 1829)

Examined material. One adult male ZFMK

93908 (SVL 46.05 mm), collected by TZ and NQT,

27 May 2009; one subadult ZFMK 93907 (SVL

24.73 mm), collected by Marta Bernardes, 7 June

2010, 75-90 m asl (Fig. 18).

Morphological characters. Head long, nar-

row; snout pointed, longer than eye (SL 4.44-7.40

mm, ED 3.34-5.53 mm); tongue bifid; vomerine

teeth present; canthus rostralis obtuse; loreal region

oblique; internostril and upper eyelid broader than

interorbital distance (IN 2.52-2.93 mm, uEL 2.27-

3.77 mm, IntOrb 1.66-2.29 mm); tympanum dis-

tinct (TD 1.72-2.48 mm); supratympanic fold

distinct; tips of fingers and toes pointed; finger I

longer than II; toes half webbed; outer metatarsal

tubercle very small; tibio-tarsal articulation reach-

ing eye; tibia 3-4 times longer than broad (TiL

13.08-23.99 mm, TiW 3.17-7.44 mm); dorsal skin

granular, with several irregular dermal folds; nup-

tial pad present in males. Colouration in life: dorsal

head and body greyish green with camouflage-

pattern; light vertebral stripe present or absent; lips

with dark vertical bars; limbs with transverse bars

or spots; ventral surface white; throat marbled with

black in males (determination after Bourret, 1942;

Manthey & Grossmann, 1997; Ziegler, 2002;

Goodall & Faithfull, 2010).

Distribution. This is a common species not

only in Vietnam but also in Asia, from Afghanistan,

Pakistan, hidia eastwards to China and Japan, south-

wards to Malaysia, Indonesia, and the Philippines

(Nguyen et al., 2009).

Remarks. The adult male was found on a stone

in a stream and the subadult was found on grassland

about 30 m away from a small stream at night.

Limnonectes bannaensis Fitzinger, 1843

Examined material. One adult female IEBR

A. 2013. 61 (SVL 47.43 mm), collected by PTC,

June 2010, ca. 350 m asl (Fig. 19).

Morphological characters. Head large, flat-

tened, wider than long (HL 19.70 mm, HW 20.9

mm); snout rounded, longer than eye diameter (SL

7.64 mm, ED 6.96 mm); canthus rostralis absent;

loreal region oblique and slightly concave; vomer-

ine teeth present; tongue bifurcated posteriorly;

lower jaw with two tooth-like, bony structures; tym-

panum hidden; supratympanic fold present; arms

short, fingers without webbing, tips of fingers ob-

tuse or slightly swollen, finger I as long as finger

516

Vera L. Hecht et alii

II, finger II and III with lateral skin-ridges; subar-

ticular tubercles on fingers and toes large; hind

limbs short; tibio-tarsal articulation reaches tympa-

num or eye; toes short, nearly fiill-webbed tips of

toes dilated into small discs; subarticular tubercles

fairly large; inner metatarsal tubercle large, outer

metatarsal tubercle absent; specimen with smooth

skin above. Colouration in life: dorsal head and

back greenish brown with small dark blotches;

limbs with dark brown bars; a short, dark transverse

line present between eyes; upper and lower lip with

vertical dark stripes; ventral surface whitish or yel-

lowish and with numerous light grey or brownish

blotches on throat, chest and limbs (determination

after Ye et al., 2007; McLeod, 2010).

Distribution. In Vietnam, this species has been

recorded from Lao Cai Province in the North to

Dong Nai and Kien Giang provinces in the South.

Elsewhere, it is known from China, presumably to

be found in Myanmar, and Laos (Nguyen et al.,

2009; McLeod, 2010).

Remarks. The specimen was found on stone in

the stream Suoi Tuyen II in the evening. McLeod

(2010) stated that Limnonectes kuhlii (Tschudi,

1838) is a complex of cryptic species. The “true”

L. kuhlii is known only from the type locality in

Java, Indonesia, and specimens identified as L.

kuhlii from China and Vietnam should be assigned

to L. bannaensis.

Quasipaa acanthophora Dubois et Ohler, 2009

Examined material. One male IEBR

A. 2013. 62 (SVL 102.23 mm), two females IEBR

A. 2013. 63 (SVL 87.26 mm), IEBR A.2013.64

(SVL 89.59 mm) collected by PTC, June 2010,

300-500 m asl; one subadult ZFMK 92843 (SVL

57.46 mm) collected by TZ and NTT, October

2009, ca. 350 m asl (Fig. 20).

Morphological characters. Head wider than

long (HW 32.16-39.93 mm, HL 29.77-36.5 mm);

snout rounded, slightly protruding, longer than eye

diameter (SL 12.28-15.10 mm, ED 8.44-11.46

mm); vomerine teeth present; canthus rostralis in-

distinct; loreal region concave; nostrils oval, with

flap of skin laterally; nostrils closer to eyes than to

tip of snout (EN 5.08-5.93 mm, NtoS 4.50-4.76

mm); interorbital distance narrower than intemarial

distance and upper eyelid (IN 7. 14-8.54 mm, IntOrb

5.08-5.93 mm, uEL 7.32-9.08 mm); tympanum in-

distinct; supratympanic fold prominent; pineal ocel-

lus present, between anterior borders of eyes; arm

short; fingers free of webbing; finger I longer than

II; tips of fingers rounded, slightly enlarged, without

discs; toes short and thin; tips of toes rounded,

distinctly enlarged, without discs; toes fully webbed;

dorsal head, body and limbs shagreened with regu-

larly disposed glandular warts; upper part of flanks

shagreened with elongated glandular warts, lower

part with foldings; dorsolateral folds absent; thigh

shagreened with thin foldings; legs shagreened with

thin foldings and homy spinules; tarsus smooth;

ventral surface smooth; belly with transversal fold-

ings. Colouration in life: dorsal surface light brown,

with grey spots; dorsal surface of limbs with trans-

verse bars; supratympanic fold darker; lips with ver-

tical bars ventral surface yellowish white, gular

region with black marbling (determination after

Dubois & Ohler, 2009). The males are larger in size,

black nuptial spines present on prepollex and finger

I (two separated pads).

Distribution. This species was previously

known only from the type locality in Lang Son Prov-

ince, Vietnam (Dubois & Ohler, 2009). This is the

first record of this species from Tay Yen Tu NR as

well as from Bac Giang Province.

Remarks. The specimens were found on stones

in the stream Suoi Tuyen I and Suoi Tuyen II in the

evening.

Occidozyga martensii (Peters, 1867)

Examined material. One adult female ZFMK

92844 (SVL 24.95 mm), collected by TZ, 29 May

2009, 200 m asl (Fig. 21).

Morphological characters. Head longer than

wide (HL 9.37 mm, HW 7.88 mm); vomerine teeth

absent; tongue rounded posteriorly; eyes directing

laterally; tympanum hidden; supratympanic fold

weakly developed; fingers and toes with small

discs; finger I longer than II; tibio-tarsal articulation

reaching eye; toes fully webbed, except toes IV and

V; intercalary cartilage tubercles absent; inner

metatarsal tubercle present; dorsal skin smooth an-

teriorly with some tubercles on the posterior part of

back; ventral skin smooth. Colouration in life: dor-

sum beige grey, with a darker stripe between eyes;

one light vertical bar from eye to snout; limbs with

First report on the herpetofauna of TayYen Tu Nature Reserve, northeastern Vietnam

517

Figure 14. Kalophrynus interlineatus. Figure 15. Microhyla butleri. Figure 16. M. heymonsi. Figure 17. M.pulchra. Figure

18. Fejervarya limnocharis. Figure 19. Limnonectes bannaensis. Figure 20. Quasipaa acanthophora. Figure 21. Occido-

zyga martensii. Photos by T. Ziegler and C.T. Pham.

518

Vera L. Hecht et alii

transverse bars; ventral surface white, gular region

marbled with black (determination after Bourret,

1942; Manthey & Grossmann, 1997; Ziegler, 2002).

Distribution. This is a widespread species in

Vietnam, from Lao Cai Province in the North to

Dong Nai and Ba Ria-Vung Tau provinces in the

South. However, this is the first record of O.

martensii from Tay Yen Tu NR and from Bac

Giang Province. Elsewhere, the species is known

from China, Laos, Thailand, and Cambodia

(Nguyen et al., 2009).

Remarks. The specimen was found in the

stream. Some other individuals were sighted in the

water or the riverine of ponds during the day time.

RANIDAE

Amolops ricketti (Boulenger, 1899)

Examined material. One subadult female

ZFMK 92845 (SVL 33.75 mm), collected by TZ

and NQT, 25-29 May 2009, 300-400 m asl (Figs.

22, 23).

Morphological characters. Head somewhat

longer than broad (HL 12.58 mm, HW 11.91 mm);

snout round, pronounced, longer than eye length

(SL 5.28 mm, ED 4.07 mm); vomerine teeth present;

canthus rostralis distinct; loreal region concave;

nostril at the midway between eye and tip of snout

(EN 2.89 mm, NtoS 2.62 mm); interorbital distance

narrower than internarial distance, but broader than

upper eyelid (IN 4.41 mm, IntOrb 3.21 mm, uEL

2.91 mm); tympanum indistinct, about 1/3 of eye

length (TD 1.17 mm); finger short, with lateral der-

mal fringe, with enlarged discs, discs with circum-

marginal groove as wide as tympanum diameter;

finger I shorter than II; subarticular tubercles small;

tibio-tarsal articulation reaching to tip of snout;

heels overlapping; tibia four times longer than wide,

about 2/3 of SVL (TiL 20.44 mm, TiW 5.00 mm);

toes fully webbed, without tarsal fold; intercalary

cartilage tubercles absent; toe discs with circum-

marginal groove, smaller than finger discs; inner

metatarsal tubercle oval; outer metatarsal tubercle

absent; dorsal skin with flattened granules; supratym-

panic fold distinct; ventral surface granular. Coloura-

tion in life: dorsum yellowish grey with larger

reddish spots; upper surface of limbs with darker

transverse bars; ventral surface yellowish white,

gular region marbled (determination after Bour-

ret, 1942).

Distribution. In Vietnam, this species has been

recorded from Lao Cai and Ha Giang provinces

southwards to Hoa Binh Province (Nguyen et al.,

2009). Records of this species in northwestern and

central Vietnam must be confirmed, as it looks simi-

lar to A. cremnobcitus Inger et Kottelat, 1998 and A.

spinapectoralis Inger, Orlov et Darevsky, 1999. This

is the first record of A. ricketti from Tay Yen Tu NR

and from Bac Giang Province. Elsewhere, this

species is known from China (Nguyen et al., 2009).

Remarks. The specimen was found near a rocky

stream near Vung Tron Ranger Station. Further

individuals were observed on branches and stones

inside and nearby streams and cascades. The speci-

men matched the description of Bourret (1942)

well, but its snout is longer than the eye length in-

stead of being shorter than the eye length.

Babina chapaensis (Bourret, 1937)

Examined material. Four adult males ZFMK

92846, VNMN 1318, ZFMK 92847 and VNMN

1321, collected by TZ, 25 July 2010, 450-500 asl

(Fig. 24).

Morphological characters. SVL 36.78-

42.58 mm; head flattened, as long as or longer than

wide (HL 13.22-16.16 mm, HW 13.14-15.50 mm);

snout obtuse, round, longer than eye length (SL

6.28-6.89 mm, ED 4.18-4.70 mm); vomerine teeth

present; canthus rostralis distinct; loreal region con-

cave; nostrils at the midway between eye and tip

of snout (EN 2.71-3.10 mm, NtoS 2.78-3.27 mm);

internarial distance as broad as or broader than in-

terorbital distance and wider than upper eyelid (IN

4.78-5.31 mm, IntOrb 3.58-4.55 mm, uEL 2.99-

3.18 mm); tympanum distinct, smaller than eye (TD

3.14-3.86 mm, TD/ED 0.73-0.90); tip of fingers

with small discs; tibiotarsal articulation reaching tip

of snout; heels overlapping; tibia 3.84^4.30 times

longer than wide (TiL 21.56-24.83 mm, TiW 5.40-

6.26 mm); toe tips with small discs, with circum-

marginal groove; toes half webbed; inner metatarsal

tubercle present; outer metatarsal tubercle small;

dorsal skin smooth, with fine granular pattern on

back, thigh and cloacal region; dorsolateral fold dis-

tinct; upper lip with a fold from below the nostril to

anterior axilla, ending in a small wart. Colouration

First report on the herpetofauna of TayYen Tu Nature Reserve, northeastern Vietnam

519

in life: dorsum brown, with a light vertebral stripe

from interorbital region to cloaca; a narrow black

line from nostrils to groin, bordering dorsolateral

fold; loreal region and tympanum dark brown;

flanks yellowish green or grey, with some black

spots; upper surface of limbs reddish brown with

transverse dark bars; back of thigh with brown mar-

bling; dermal fold on upper lip white; ventral sur-

face white, yellow in posterior part; lower surface

of arms with grey or black spots (determination

after Bourret, 1942).

Distribution. In Vietnam, this species has been

recorded from Lao Cai, Bac Giang, Ha Tinh, Kon

Turn, Gia Lai, and Dak Lak provinces. Elsewhere,

the species is known from Laos and Thailand

(Nguyen et al., 2009).

Remarks. Specimens were found at night on the

ground in the bamboo forest near Da Lua Pond.

Hylarana guentheri (Boulenger, 1882)

Examined material. Two adult males IEBR

A.2013.65 (SVL 71.31 mm), IEBRA.2013.66 (SVL

72.85 mm) collected by PTC, June 2010; one adult

female IEBR A.3013.67 (SVL 73.10 mm) collected

by PTC, October 2011, 100-300 m asl (Fig. 25).

Morphological characters. Head wider than

long (HL 19.71-20.11 mm, HW 25.59-26.75 mm);

vomerine teeth present; intemarial distance broader

than upper eyelid (IntOrb 6.32-6.63 mm, uEL 4.50-

5.30 mm); tip of snout pointed, strongly projecting;

canthus rostralis distinct; loreal region moderately

oblique, concave; nostril closer to tip of snout than

to eye (EN 6.10-7.95 mm, NtoS 3.10-5.91 mm);

tympanum distinct (TD 6.14-6.50 mm); supratym-

panic fold prominent; fingers slender and rather

long, free of weebing, finger tips swollen, finger I

longer than finger II, subarticular tubercles large;

toes 3/4 webbed, tips of toes dilated into small discs,

with a median groove, subarticular tubercles rather

small, tarsal fold absent; inner metatarsal tubercle

present, outer metatarsal tubercle very small; skin

above and below smooth; dorsolateral fold distinct.

Colouration in life: dorsal surface of head and body

grey, light brown, golden to reddish brown, uniform

or with dark brown spots; lateral head and flanks

with a dark line or band, bordering the dorsolateral

fold; tympanum dark brown or reddish; limbs with

brown crossbars, back of thighs yellow with black

mottling; ventral surface white or yellow, throat or

chest speckled with brown (determination after

Bourret, 1942; Ziegler, 2002).

Distribution. This is a common species in low-

land areas of Vietnam. Elsewhere, the species has

been reported from China, Taiwan, Myanmar, and

Laos (Nguyen et al., 2009).

Remarks. The specimens were found in a small

stream near Mau village.

Hylarana maosonensis Bourret, 1937

Examined material. One adult male ZFMK

92850 (SVL 40.43 mm), collected by TZ and NTT,

October 2009, ca. 500 m asl (Fig. 26).

Morphological characters. Head longer than

wide (HL 15.40 mm, HW 14.24 mm); vomerine

teeth present; snout round, slightly pronounced,

shorter than eye length (SL 6.05 mm, ED 6. 14 mm);

loreal region oblique, concave; nostril closer to tip

of snout than to eye (EN 3.50 mm, NtoS 2.60 mm);

interorbital distance narrower than internarial dis-

tance and upper eyelid (IN 4.17 mm, IntOrb 3.33

mm, uEL 3.90 mm); tongue notched posteriorly;

pupil rounded; tympanum distinct, half of eye length

(TD 3.04 mm); fingers free of webbing, with small

discs; subarticular tubercle round, distinct; tibio-

tarsal articulation reaching between eye and tip of

snout; discs of toes larger than discs of fingers; toes

3/4 webbed; intercalary cartilage tubercles absent;

tibia 4 times longer than wide (TiL 22.65 mm, TiW

5.23 mm); subarticular tubercles small, round; inner

metatarsal tubercle small, elongated; outer metatarsal

tubercle small; tarsal fold absent; dorsolateral fold

distinct, interrupted posteriorly; tubercles present on

dorsal surface of body and thighs and upper part of

flanks; supratympanic fold prominent; a large tuber-

cle in posterior corner of the mouth; flanks with

elongated warts below dorsolateral fold; two rows

of elongated warts on each thigh; ventral surface

smooth. Colouration in life: dorsum yellowish

brown, flanks lighter with black spots; limbs with

dark transverse bars; tympanum brown; lips white

with small black spots; ventral surface whitish, chin

mottled with dark brown (determination after Bour-

ret, 1942; Inger et al., 1999).

Distribution. This species has been recored in

northern and central Vietnam, from Lao Cai and Ha

Giang provinces southwards to Quang Tri and Thua

520

Vera L. Hecht et alii

Thien-Hue provinces. Elsewhere, the species is

known from Laos (Nguyen et al., 2009).

Rem arks. The specimen was found at night in

Suoi Tuyen I.

Hylarana cf. nigrovitatta (Blyth, 1856)

Examined material. One adult female IEBR

A.2013.68 (SVL 46.43 mm), collected by TZ and

NQT, 28 May 2009, 200 m asl (Fig. 27).

Morphological characters. Elead longer than

wide (HL 18.26 mm, HW 15.91 mm); vomerine

teeth present; snout round, longer than eye (SL 6.70

mm, ED 5.93 mm); loreal region oblique, concave;

nostril closer to tip of snout than to eye (EN 3.57

mm, NtoS 3.13 mm); intemarial distance broader

than interorbital distance and upper eyelid (IN 4.60

mm, IntOrb 4.17 mm, uEL 3.66 mm); tongue

notched posteriorly; pupil rounded; tympanum dis-

tinct (TD 3.70 mm); fingers free of webbing, tips of

fingers swollen; toes 3/4 webbed; subarticular tuber-

cles small; inner metatarsal tubercle small, elongated;

outer metatarsal tubercle small; dorsolateral fold dis-

tinct; skin above and below smooth. Colouration in

life: dorsal head and body greyish brown, a dark

stripe present on each side of head and upper part of

flanks, bordering the dorsolateral fold; tympanum

brown; limbs with dark crossbars, back of thighs yel-

low with black mottling; ventral surface whitish (de-

termination after Bourret, 1942; Inger et al., 1999).

Distribution. In Vietnam, this species complex

has been recorded from Cao Bang and Lang Son

provinces in the North to Dong Nai and Kien Giang

provinces in the South. This is the first record of H.

cf. nigrovittata from Tay Yen Tu NR and from Bac

Giang Province. Elsewhere, the species is known

from India, Nepal, China, Myanmar, Laos, Thailand,

Cambodia, and Malaysia (Nguyen et al., 2009).

Remarks. The specimen was found at night in

a rocky stream near Vung Tron Ranger Station. H.

nigrovittata represents a so far only partly under-

stood complex of cryptic species (e.g., Gawor et al.,

2009).

Hylarana taipehensis (Van Denburgh, 1909)

Examined material. Three adult males IEBR

A. 2013. 69 (SVL 27.27 mm), IEBR A.2013.70

(SVL 28.56 mm), IEBR A.2013.71 (SVL 25.14

mm); one adult female IEBR A. 2013.72 (SVL

37.71 mm) collected by PTC, October 2011, 200-

300 m asl (Fig. 28).

Morphological characters. Head longer than

wide (HL 10.19-14.42 mm, HW 7.80-10.15 mm);

vomerine teeth present; nostril closer to tip of snout

than to eye (EN 2.76-4.20 mm, NtoS 1.96-2.32

mm); interorbital distance wider than upper eyelid

(IntOrb 3.45-5.17 mm, uEL 2.32-3.56 mm); tym-

panum distinct, about 2/3 of eye diameter (TD

2.13-2.80 mm, ED 2.83-3.47 mm); fingers small,

free of webbing, tips of fingers swollen; toes 3/4

webbed; dorsolateral fold distinct; skin above and

below smooth. Colouration in life: dorsum green,

with two stripes from behind the eyes to groins;

ventral surface white or yellow (determination after

Bourret, 1942; Inger et al., 1999).

Distribution. This is a widespread species in

Vietnam, from Lao Cai and Ha Giang provinces

southwards to Dong Nai and Kien Giang provinces.

Elsewhere, the species has been reported from India,

Nepal, Bangladesh, China, Taiwan, Myanmar, Laos,

Thailand, and Cambodia (Nguyen et al., 2009).

Remarks. The specimens were found at night in

Ba Bep.

Odorrana bacboensis Bain, Lathrop, Murphy,

Orlov et Ho, 2003

Examined material. Two adult females IEBR

A. 2013. 73 (SVL 106.19 mm) and ZFMK 92848

(SVL 108.44 mm), collected by TZ and NQT, 26-

27 May 2009, 300-500 m asl (Fig. 29).

Morphological characters. Head longer than

wide (HL 37.27-39.46 mm, HW 35.65-36.00 mm);

vomerine teeth present; tongue notched posteriorly;

snout round, slightly prominent, longer than eye (SL

15.58-16.72 mm, ED 10.14—11.80 mm); canthus ros-

tralis distinct; interorbital distance narrower than in-

ternarial distance but broader than upper eyelid

(IntOrb 9.00-10.24 mm, IN 10.37-10.92 mm, uEL

7.55-8.90 mm); pupil round; tympanum distinct,

half of eye diameter (TD 4.83-5.08 mm, ED 10.14-

11.80 mm); supratympanic fold distinct; fingers free

of webbing; finger discs larger than those of toes,

with circummarginal groove; tibiotarsal articulation

reaching tip of snout; tibia 4 times longer than wide

(TiL 60.90-61.45 mm, TiW 15.55-15.60 mm); toes

fully webbed; outer metatarsal tubercle absent; inner

First report on the herpetofauna of TayYen Tu Nature Reserve, northeastern Vietnam

521

Figures 22, 23. Amo/ops ricketti. Figure 24. Babina chapaensis. Figure 25. Hylarana guentheri.

Figure 26. H. maosonensis. Figure 27. H. cf. nigrovitatta. Photos by T. Ziegler and C.T. Pham.

522

Vera L. Hecht et alii

metatarsal tubercle large, flat; tarsal fold absent; dor-

sal skin smooth; dorsolateral fold distinct. Coloura-

tion in life: dorsum brown to dark brown, with tiny,

irregular, black blotches; flanks brown, with some

large black spots; throat and chest region white (de-

termination after Bain et al., 2003).

Distribution. In Vietnam, this species has been

recorded in Lao Cai, Tuyen Quang, Bac Kan, and

Nghe An provinces (Nguyen et al., 2009). This is

the first record of O. bacboensis from Tay Yen Tu

NR and from Bac Giang Province.

Remarks. The specimens were found on the

rock in a small cascade stream (Suoi Tuyen I).

O dorr ana graminea (Boulenger, 1900)

Examined material. One adult female VNMN

1346 (106.68 mm), collected by TZ and NTT, June

2010; one adult female ZFMK 92849 (SVL 80.37

mm), collected by TZ and NTT, October 2009,

300-500 m asl (Fig. 30).

Morphological characters. Head longer

than wide (HF 30.47-35.14 mm, HW 26.89-32.93

mm); vomerine teeth present; tongue notched pos-

teriorly; snout round, longer than eye (SF 12.72-

16.40 mm, ED 7.40-9.67 mm); canthus rostralis

distinct; interorbital distance wider than intemarial

distance and upper eyelid (IntOrb 8.08-1 1 .45 mm,

IN 7.44-10.11 mm, uEF 6.84-7.14 mm); pupil

round; tympanum distinct, approximately half of

eye diameter (TD 4.17-4.22 mm, ED 7.40-9.67

mm); supratympanic fold distinct; fingers free of

webbing; finger discs larger than discs of toes,

with circummarginal groove; tibiotarsal articula-

tion reaching beyond tip of snout; tibia about

4.64-5.91 times longer than wide (TiF 52.27-

65.1 1 mm, TiW 8.85-14.02 mm); toes fully webbed;

outer metatarsal tubercle absent; inner metatarsal

tubercle elongate; dorsal skin smooth, flank with

tubercles; dorsolateral fold absent. Colouration in

life: dorsum green with black spots; sides of head

and flanks brownish grey; lips white; hind limbs

with distinct dark bars; webbing dark grey; ventral

surface whitish (determination after Bourret, 1942;

Bain et al., 2003).

Distribution. In Vietnam, this species has been

recorded from Bac Kan, Fang Son, and Earn Dong

provinces. This is the first record of O. graminea

from Tay Yen Tu NR and from Bac Giang Province.

Elsewhere, the species is known from China

(Nguyen et al., 2009).

Remarks. The specimens were collected at

night time in a small cascade stream.

O dorr ana nasica (Boulenger, 1903)

Examined material. One adult male IEBR

A.2013.74 (SVL 73.45 mm), collected by TZ and

NTT, October 2009, 300-500 m asl (Fig. 31).

Morphological characters. Head longer than

wide (HL 27.04 mm, HW 23.77 mm); vomerine

teeth present; snout obtusely pointed, prominent,

longer than eye (SL 12.75 mm, ED 8.22 mm); can-

thus rostralis distinct; internarial distance broader

than interorbital distance and upper eyelid (IN 8.41

mm, IntOrb 6.14 mm, uEL 5.17 mm); tympanum

distinct, half of eye diameter (TD 4.52 mm, ED

8.22 mm); supratympanic fold absent; fingers free

of webbing; finger I longer than II; finger discs larger

than those of toes, with circummarginal groove;

tibiotarsal articulation reaching beyond tip of snout;

tibia 4.48 times longer than wide (TiL 44.48 mm,

TiW 9.92 mm); toes fully webbed; outer metatarsal

tubercle absent; inner metatarsal tubercle elongate;

dorsal skin smooth; dorsolateral fold present; exter-

nal gular sacs absent; nuptial pads present. Coloura-

tion in life: dorsum dark green; dorsolateral fold and

canthus rostralis white; sides of head and flanks

brownish grey; lips whitish; tympanum dark brown;

limbs with dark bars; ventral surface white (deter-

mination after Bourret, 1942; Bain et al., 2003).

Distribution. In Vietnam, this species has been

recorded from Lao Cai and Cao Bang provinces

southwards to Ha Tinh and Thua Thien-Hue prov-

inces. This is the first record of O. nasica from Tay

Yen Tu NR and from Bac Giang Province. Else-

where, this species is known from China, Laos, and

Thailand (Nguyen et al., 2009).

Remarks. IEBR A.2013.74 was found at night

in a broad, deep part of a rocky stream near the

Dong Thong Ranger Station.

Odorrana yentuensis Tran, Orlov et Nguyen, 2008

Examined material. One adult male ZFMK

92851 (SVL 43.14 mm) and two adult females

ZFMK 92852 (SVL 63.01 mm) and IEBR

First report on the herpetofauna of TayYen Tu Nature Reserve, northeastern Vietnam

523

A.20 13.76 (SVL 59.29 mm) collected by TZ and

NTT, October 2009, 400-600 m asl; two adult

males VNMN 1320 (SVL 46.00 mm) and VNMN

1344 (SVL 45.72 mm) collected by TZ and NTT,

June 2010, 350-600 m asl (Fig. 32).

Morphological characters. Males smaller

than females (SVL males 43.14-46.00 mm, females

59.29-63.1 mm); head longer than wide (HL 16.21 —

22.83 mm, HW 13.43-20.45 mm); snout long,

rounded anteriorly; vomerine teeth present; eye

larger than tympanum (TD/ED 0.47-0.72); lingers

free of webbing, with small discs, with circum-

marginal groove; finger I longer than II; subarticu-

lar tubercles round, prominent; toes almost fully

webbed; subarticular tubercles distinct, rounded;

dorsal skin rough, posterior part of back with small

tubercles; ventral skin smooth; dorsolateral fold dis-

tinct; supratympanic fold distinct; males with nup-

tial pad and external vocal sacs. Colouration in life:

dorsum brownish grey, with several irregular brown

and/or green spots; lips white; dorsolateral fold bor-

dered by a black line, discontinuous posteriorly;

limbs with transverse bars; webbing grey; belly and

ventral surface of limbs immaculate yellowish

white (detennination after Tran et al., 2008).

Distribution. This species is currently known

only from Bac Giang Province, Vietnam (Tran et

al., 2008; Nguyen et al., 2009).

Remarks. Four specimens were found on rocks

at night in Suoi Tuyen I and II or at the shore of

Suoi Tuyen I. The specimens matched the descrip-

tion of Tran et al. (2008) well but differ in a lower

ratio of tympanum diameter to eye diameter (0.47-

0.72 instead of 0.81-0.84).

Rana johnsi Smith, 1921

Examined material. One adult male ZFMK

92853 (SVL 45.76 mm), collected by TZ and NTT,

October 2009, 250-300 m asl (Fig. 33).

Morphological characters. Head as broad as

long (HW 15.48 mm, HL 15.70 mm), flattened;

snout obtusely pointed, pronounced, longer than

eye (SL 6.97 mm, ED 5.61 mm); tympanum dis-

tinct, about 3/4 of ED (TD 4.09 mm); interorbital

distance broader than internostril distance and

upper eyelid (IntOrb 4.00 mm, IN 3.60 mm, uEL

3.04 mm); pupil horizontal; vomerine teeth present;

fingers free of webbing without discs; subarticular

tubercles prominent; tibio-tarsal articulation reach-

ing beyond tip of snout; heels overlapping; tibia five

times longer than wide (TiL 30.22 mm, TiW 5.63

mm); toes with small discs, almost fully webbed;

tarsal fold absent; subarticular tubercles distinct;

inner metatarsal tubercle oval, prominent; outer

metatarsal tubercle absent; dorsal skin smooth, with

some small tubercles; some short, oblique dermal

folds on limbs; a A-shaped fold between shoulders;

supratympanic fold present; dorsolateral fold dis-

tinct anteriorly, interrupted on hip. Colouration in

life: dorsum light brown, upper surface of limbs

with greyish transverse bars; flanks whitish brown;

a small black stripe from nostril to eye; tympanum

covered by a black lozenge; sides of limbs with dark

pattern; ventral surface white or cream; gular region

marbled with grey; thigh yellow (determination

after Bourret, 1942).

Distribution. This is a widespread species in

Vietnam, from Lao Cai and Ha Giang provinces

southwards to Lam Dong and Dong Nai provinces.

Elsewhere, this species is known from China,

Taiwan, Laos, Thailand, and Cambodia (Nguyen et

al., 2009).

Remarks. The specimen was found on the

ground of broad leaved forest. Another, most likely

a female, was observed at Ba Bep Pond during the

day time.

RHACOPHORIDAE

Chiromantis vittatus (Boulenger, 1887)

Examined material. Two adult males VNMN

1341-1342, collected by TZ and NTT, June 2010,

350-400 m asl; one adult male IEBR A.2013.77,

collected by VH, 7 June 2010, 400 m asl (Fig. 34).

Morphological characters. SVL 19.52-25.91

mm; vomerine teeth absent; tongue bifid; pupil hor-

izontal; tympanum indistinct, 0.34-0.60 times of ED

(TD 1.05-2.21 mm, ED 3.06-3.69 mm); snout

pointed, as long as diameter (SL 3.38-3.94 mm);

canthus rostralis obtuse; loreal region slightly

oblique; nostrils closer to tip of snout than to eye

(EN 1.85-2.41 mm, NtoS 1.58-1.91 mm); interor-

bital distance as broad as or broader than upper eye-

lid (IntOrb 2.67-3.25 mm, uEL 1.90-2.85 mm);

fingers free of webbing, tips of fingers with enlarged

discs; finger I shorter than II; tibio-tarsal articulation

524

Vera L. Hecht et alii

Figure 28. Hylarana taipehensis. Figure 29. Odorrana bacboensis. Figure 30. Odorrana graminea. Figure 31. Odorrana

nasica. Figure 32. Odorrana yentuensis. Figure 33. Rana johnsi. Photos by C.T. Pham, T. Ziegler, and T.Q. Nguyen.

First report on the herpetofauna of TayYen Tu Nature Reserve, northeastern Vietnam

525

reaching eye or beyond but not to tip of snout; limbs

without fringes; toes 3/4 webbed, discs well devel-

oped; cloacal fold absent; inner metatarsal tubercle

tiny; skin smooth; gular region, posterior part of ven-

tral surface and posterior parts of femur granular;

nuptial pads absent. Colouration in life: dorsum yel-

low or brown with light yellow stripes from nostril over

eye to groin; flanks yellowish; ventral surface whitish

(determination after Bourret, 1942; Ziegler, 2002).

Distribution. This is a widespread species in

Vietnam, from Lang Son Province southwards to

Dong Nai and Ba Ria-Vung Tau provinces. Else-

where, this species is known from India, China,

Myanmar, Laos, Thailand, and Cambodia (Nguyen

et al., 2009).

Remarks. All three specimens were found at

night on plants about 1-2 m above the ground near

the ponds.

Kurixalus bisacculus (Taylor, 1962)

Examined material. Two adult males IEBR

A. 2013. 78 (SVL 29.13 mm) IEBR A. 2013.79

(SVL 29.75 mm) collected by PTC, June 2010,

300-500 m asl (Figs. 35, 36).

Morphological characters. Head longer than

wide (HL 10.66-12.80 mm, HW 9.62-11.66 mm);

snout pointed anteriorly; tympanum distinct, smaller

than eye (ED 4.21-4.49 mm, TD 2.41-2.75 mm);

intemarial distance narrower than interorbital dis-

tance (IN 3.30-3.39 mm, IntOrb 4.07^4.41 mm);

vomerine teeth in two low ridges, arising near inner

edges of choanae; fingers free of webbing, tips of fin-

gers with enlarged discs; toes 3/4 webbed, discs well

developed; dermal fringes along outer edges of fore

arm and tarsus present; some scattered flat tubercle

present on head, eyelids, and occiput, sparse on dor-

sum, more dense and larger on flanks, finely granuler

on mmp; chin granular, chest nearly smooth; venter

and lower part of sides granular; vocal sacs present

in males. Colouration in life: dorsal head and body

light or reddish brown with green marking, occiput

with a dark green marking in triangular shape; tym-

panum brownish; chin cream with dark spots; throat,

chest, venter, and underside of limbs cream (deter-

mination after Taylor, 1962; Yu et al., 2010).

Distribution. This species has been reported

from Lao Cai and Ha Giang provinces in the North

to Gia Lai Province in the South. However, the ex-

tent of this species in Central Vietnam needs to be

confirmed, as it looks similar to K. banaensis

(Bourret, 1939). Elsewhere, the species is known

from China, Myanmar, Laos, Thailand, and Cam-

bodia (Nguyen et al., 2009).

Remarks. Specimens were found at night on the

tree branches and shrubs near ponds or streams. Yu et

al. (2010) regarded the previous records of K. verru-

cosus (Boulenger, 1893) in Vietnam as K. bissaculus.

Polypedates megacephalus Hallowell, 1861

Examined material. One adult female VNMN

1322 (SVL 70.15 mm), collected by TZ and NTT,

26 June 2010, ca. 350 m asl (Fig. 39).

Morphological characters. Head slightly

longer than wide (HL 26.30 mm, HW 25.12 mm);

vomerine teeth present; snout obtusely rounded,

longer than eye (SL 11.93 mm, ED 6.61 mm); can-

thus rostralis distinct; loreal region vertical; nostrils

closer to tip of snout than to eye (EN 7.45 mm, NtoS

3.66 mm); interorbital distance wider than intemarial

distance and upper eyelid (IntOrb 9.57 mm, IN 5.27

mm, uEL 4.93 mm); tympanum distinct, as large as

eye (TD 5.04 mm); fingers free of webbing; disc of

finger III half of TD (disc of finger III 2.00 mm); toes

almost fully webbed; lateral dermal fringes present;

subarticular tubercles present; outer metatarsal tuber-

cle present; tibio-tarsal articulation reaching tip of

snout; skin smooth; supratympanic fold distinct;

vocal sacs absent. Colouration in life: dorsum orange

brown, with some black blotches and sometimes

with a X-shaped pattern on neck; upper surface of

limbs orange brown with transverse bars; flanks and

posterior side of thigh with distinct dark reticulation;

throat and chest mottled in brown; ventral surface

white (determination after Bourret, 1942; Manthey

& Grossmann, 1997; Kuraishi et al., 2012).

Distribution. In Vietnam, this species has been

recorded from Cao Bang, Vinh Phuc, and Hai

Duong provinces. Kuraishi et al. (2012) suggested

that the populations of the P. leucomystax complex

in southern China and northern Vietnam contain at

least two species, P. megacephalus and P. mutus.

Elsewhere, the species is known from India, China,

Myanmar, Taiwan, Laos, Thailand, and Japan

(Nguyen et al., 2009).

Remarks. A single specimen was found at night

on a tree branch in Suoi Tuyen II.

526

Vera L. Hecht et alii

Polyp edates mutus (Smith, 1940)

Examined material. One female VNMN

A. 2013. 14 (SVL 89.44 mm), two males IEBR

A. 2013. 80 (SVL 62.92 mm), ZFMK 92854 (SVL

65.36 mm), collected by TZ and NQT, 27 May

2009, 200-500 m asl (Figs. 37, 38).

Morphological characters. SVL 62.92-

89.44 mm; head longer than wide (HL 21.51-

30.89 mm, HW 19.36-29.13 mm); vomerine teeth

present; snout pointed, longer than eye (SL 10.10-

14.03 mm, ED 6.31-10.29 mm); canthus rostralis

distinct, slightly concave; loreal region vertical;

nostrils closer to tip of snout than to eye (EN 6.46-

9.42 mm, NtoS 2.75-3.90 mm); interorbital dis-

tance wider than internarial distance and upper

eyelid (IntOrb 5.52-8.14 mm, IN 3.90-5.73 mm,

uEL 5.51-7.73 mm); tympanum distinct, approxi-

mately half of eye diameter (TD 3.44-5.36 mm);

fingers free of webbing; disc of finger III as large

as tympanum (disc of finger III 2. 6-5. 2 mm); toes

almost fully webbed; toe discs smaller than finger

discs; lateral dermal fringe present; subarticular

tubercles present; outer metatarsal tubercle present;

tibio-tarsal articulation reaching tip of snout; dor-

sal skin smooth; upper surface of limb granular;

ventral surface smooth; supratympanic fold present;

vocal sacs absent. Colouration in life: dorsum

beige yellow or brown, with or without dark

blotches; a X-shaped pattern present on neck;

limbs with dark transverse bars; dark brown stripe

bordering supratympanic fold from eye beyond

arm; flanks sometimes with few dark brown spots;

back of thigh with large white spots; belly yellow-

ish white; throat finely mottled or spottet (deter-

mination after Ziegler, 2002; Ziegler et al., 2006;

Kuraishi et al., 2012).

Distribution. This is a widespread species in

Vietnam. Elsewhere, this species is known from

China, Myanmar, Laos, and Thailand (Nguyen et

al., 2009).

Remarks. All three specimens were found at

night on tree branches in about E5-2 m above a

stream. A couple of P. mutus in amplexus was pho-

tographed on the forest path in June 2010. Further

individuals were sighted on tree trunks or leaves

between 0.5 and 3 m above the streams or around

ponds during the day and at night time.

Rhacophorus maximus Inger, 1966

Examined material. Two adult males IEBR

3653, IEBR 3680, collected by NQT, 10-13 May

2008; one adult male VNMN 1535 and one adult

female VNMN 1538, collected by NTT, June 2009,

250-500 m asl (Fig. 40).

Morphological characters. SVL 74.34-

13 L0 mm; head broader than long (HW 28. 1-36.74

mm, HL 26.4-34.48 mm); vomerine teeth present;

nostril oval, closer to tip of snout than to eye; tym-

panum round, smaller than eye diameter (TD 4.4-

5.64 mm, ED 7.16-9.41 mm); interorbital distance

wider than internarial distance (IntOrb 9.5-1 1.97

mm, IN 7.9-10.72 mm); supratympanic fold distinct;

fingers and toes completely webbed; metacarpal

tubercle well developed in males; males with vocal

sacs. Colouration in life: dorsal surface of head

and body uniformly green; a narrow white stripe

present along the flanks; ventral surface cream

(determination after Anders & Rai, 2002; Nguyen

et al., 2008).

Distribution. In Vietnam, this species has been

recorded from Bac Giang and Nghe An provinces.

Elsewhere, the species is known from India, Nepal,

China, and Thailand (Nguyen et al., 2009).

Remarks. Specimens were found on trees and

shrubs (ca. 1-3 m above the ground) near Ao Cua

pond or streams in secondary forests near Dong Ri

Ranger Station.

Rhacophorus rhodopus (LiuetHu, 1960)

Examined material. Two adult males ZFMK

92855, IEBR A.2013.81 (VH28) collected by TZ

and NQT, 27 May 2009, 400 m asl; one adult male

VNMN A. 2013. 15, collected by VH, 7 June 2010,

ca. 430 m asl (Fig. 41).

Morphological characters. SVL 39.89-

40.17 mm; head as long as or longer than wide

(HL 14.41-14.81 mm, HW 13.18-14.63 m);

vomerine teeth present; tongue notched poste-

riorly; snout pointed, longer than eye diameter (SL

6.44-6.69 mm, ED 4.53-5.57 mm); pupil rounded;

nostril closer to eye than to tip of snout (EN 2.65-

3.27 mm, NtoS 3.00-3.11 mm); interorbital dis-

tance broader than internarial distance and upper

eyelid (IntOrb 6.44-6.69 m, IN 3.87-4.17 mm,

uEL 3.60-4.46 mm); tympanum distinct, about

First report on the herpetofauna of TayYen Tu Nature Reserve, northeastern Vietnam

527

half of eye diamter (TD 1.79-2.45 mm); fingers

almost fully webbed; finger I shorter than II; sub-

articular tubercles distinct; toes fully webbed;

tarsal fold present; inner metatarsal tubercle present,

outer metatarsal tubercle absent; subarticular tu-

bercles small, distinct; tibio-tarsal articulation

reaching between eye and tip of snout; tibia about

five times longer than wide (TiL 19.09-19.63 mm,

TiW 3.52-3.74 mm); cloacal dermal fringe present;

skin smooth. Colouration in life: dorsum reddish

brown with or without small black spots; large

black blotches on axilla and flanks present or ab-

sent; transverse bands on hind limbs absent or in-

distinct; ventral yellowish white; webbing of

fingers and toes reddish orange (determination

after Bourret, 1942; Lui & Hu, 1960).

Distribution. In Vietnam, this species has been

recorded from Lao Cai Province southwards to Lam

Dong and Dong Nai provinces. The first record of

R. rhodopus from Tay Yen Tu NR was reported by

Nguyen et al. (2008). Elsewhere, the species is

known from India, China, Myanmar, Laos, Thai-

land, and Cambodia (Nguyen et al., 2009).

Remarks. All specimens were found at night on

trees and bushes, ca. 0.5-1. 5 m above the ground,

around ponds in secondary forest. The female is

bigger than the male. The female has yellowish

orange flanks with some black blotches. Our speci-

mens match the descriptions of Bourret (1942) and

Lui & Hu (1960), but show some differences: toes

not fully webbed, transverse bands on hind limbs

indistinct and black blotches on axilla and flanks

absent in some individuals.

Theloderma asperum (Boulenger, 1886)

Examined material. Two adult males IEBR

A.2013.82 (SVL 31.92 mm, IEBR A.2013.83 (SVL

32.32 mm), collected by PTC, 8 July 2012, eleva-

tions 300-500 m asl (Fig. 42).

Morphological characters. Head broader

than long (HW 11.73-12.01 mm, HL 10.59-11.81

mm); snout rounded, as long as the eye (SL 3.66-

4.20 mm, ED 3.63-3.79 mm); canthus rostralis in-

distinct; loreal region slightly concave; nostril

closer to tip of snout than to eye (NtoS 1.05-1.48

mm, EN 2.61-2.71 mm); interorbital distance wider

than upper eyelid (IntOrb 4.69-4.81 mm, uEL 2.3-

2.4 mm); tympanum distinct (TD 2.45-3.20 mm);

vomerine teeth absent; tongue notched behind; fin-

gers free of webbing, tips of fingers with enlarged

discs; toes 3/4 webbed, tips of toes with large discs;

inner metatarsal tubercle present, small; tibiotarsal

articulation reaching tip of snout; skin on dorsum

and flanks with granular tubercles; throat smooth;

venter granular. Colouration in life: dorsal surface

blackish or greyish brown with large white blotches

covering head, loreal regions, anterior parts of dor-

sum, upper part of flanks and hip; dark transverse

bars on hind limbs present; head with short trans-

verse brown line between eyes; ventrally body and

limbs blackish with whitish marbling; iris pinkish

brown (determination after Bourret, 1942; Taylor,

1962; Neang & Holden, 2008).

Distribution. This is a widespread species in

Vietnam, from Lai Chau, Lao Cai, and Ha Giang

provinces in the North to Lam Dong and Dong Nai

provinces in the South. T. asperum was previously

known from the eastern side of the Yen Tu Moun-

tain in Hai Duong Province (Nguyen et al., 2009).

Our finding represents the first record of this

species from Tay Yen Tu NR and from Bac Giang

Province. Elsewhere, this species is known from

India, China, Myanmar, Laos, Thailand, Cambodia,

and Malaysia (Nguyen et al., 2009).

Remarks. Specimens were collected in a tree

hole in bamboo forest near Mau village.

Theloderma corticate (Boulenger, 1903)

Examined material. Two adult males IEBR

A. 2013. 84 (SVL 67.68 mm), IEBR A.2013.85

(SVL 57.48 mm) collected by PTC, 10 July 2012,

elevations 300-500 m asl (Fig. 43).

Morphological characters. Head wider than

long (HL 25.07-26.72 mm, HW 27.5-28.82 mm);

snout longer than eye diameter (SL 10.43-12.19

mm; ED 6.37-7. 14 mm); canthus rostralis rounded;

loreal region concave; interorbital distance wider

than internarial distance (IntOrb 7.78-8.87 mm, IN

4.19-4.67 mm); nostril closer to tip of snout than

to eye (NtoS 2.21-2.85 mm; EN 9.22-9.34 mm);

tympanum oval, greater than tympanum-eye dis-

tance (TD 5.29-5.30 mm, ET 3.69-4.32 mm);

vomerine teeth present; tongue notched posteriorly;

tips of fingers and toes enlarged into round discs;

webbing present at base of fingers III and IV; der-

mal fringe along outer sides of arm and tarsus

528

Vera L. Hecht et alii

present; palmar tubercles distinct; toes almost fully

webbed; subarticular tubercles present; inner

metatarsal tubercle present; outer metatarsal tuber-

cle absent; dorsal surface of head, body and limbs

covered with tubercles or warts of different sizes;

ventral skin with small tubercles; nuptial pad

present; external vocal sac absent. Colouration in

life: dorsal surface green marbled with reddish

brown spots; dark brown bars present on upper sur-

face of fore and hind limbs; ventral surface yellow

with green marbling (determination after Inger et

al., 1999; Orlov et ah, 2006).

Distribution. This species has been recorded

only from Northern Vietnam: Ha Giang, Tuyen

Quang, Cao Bang, Lang Son, Vinli Phuc, and Son La

provinces (Nguyen et al., 2009). This is a new record

for Tay Yen Tu NR and for Bac Giang Province.

Remarks. Specimens were discovered in a tree

hole in secondary forest near Dong Thong Ranger

Station.

Theloderma lateriticum Bain, Nguyen et Doan,

2009

Examined material. Two adult males VNMN

1215 (SVL 21.8 mm) and VNMN 1216 (SVL 23.1

mm), collected by PTC and NTT, 10 April 2010,

elevation ca. 300 m asl (Fig. 44).

Morphological characters. Head longer than

wide (HW 7. 5-8.4 mm, HL 8. 3-9.0 mm); vomerine

teeth absent; nostril nearer to tip of snout than to

eye; interorbital distance greater than width of

upper eyelid; tympanum distinct, with distinct tym-

panic annulus (TYD 2. 0-2. 6 mm); fingers free of

webbing; toes 1/4 webbed; outer metatarsal tubercle

present; dorsal skin granular, bearing keratinized

spicules, raised on small, isolated bumps; dermal

fringes on the post axial portions of the limbs ab-

sent; nuptial pad present on finger I; vocal sacs ab-

sent. Colouration in life: dorsum rusty brown with

a mid-dorsal black spot; lip with small white spots;

upper portion of flanks with black blotches; throat,

chest and belly grey-brown with cream spots (iden-

tification after Bain et al., 2009).

Distribution. This species is known only from

the type locality in Lao Cai Province, Vietnam

(Bain et al., 2009). This is the first record of T.

lateriticum from Tay Yen Tu NR as well as from

Bac Giang Province.

Remarks. Two specimens were found at night

time on the tree near a rocky stream, surrounded by

the mixed secondary forest of small hardwoods and

shrubs near Dong Ri Ranger Station.

SQUAMATA

SAURIA

AGAMIDAE

Acanthosaura lepidogaster (Cuvier, 1829)

Examined material. One subadult female

IEBR 3660 collected by NQT on 10 April 2008, ca.

350 m asl; one juvenile ZFMK 9283 1 , collected by

TZ and NQT, 28 May 2009, 200 m asl; and one

juvenile VNMN 1348, collected by TZ and NTT,

May 2010, ca. 400 m asl (Fig. 45).

Morphological characters. SVL subadult fe-

male 69.31 mm, juveniles 29.14-34.31 mm (n = 2),

TaL subadult female 114.00 mm, juveniles 43.41-

46.99 mm (n = 2); head large; postorbital spine

small, shorter than half the diameter of orbit; two

spines present above tympanum; nuchal crest spines

8; dorsal crest present, low; supralabials 9-14; in-

fralabials 9-13; tympanum visible; mental small,

pentagonal; body compressed, not flattened; dorsal

scales heterogeneous, keeled; ventral scales strongly

keeled, midbody scales in 114-123 rows; femoral

pores absent; lamellae 17-19 under finger IV, 20-22

under toe IV; hind limbs long, reaching up to or over

tip of snout. Colouration in life: dorsum green to dark

brown, a bright rhombic pattern usually present on

the neck; back and tail with dark cross bars (deter-

mination after Ziegler, 2002; Bourret, 2009).

Distribution. This is a common species in

Vietnam, from Lao Cai and Ha Giang provinces

southwards to Binh Phuoc and Dong Nai prov-

inces. Elsewhere, the species is known from China,

Myanmar, Laos, Thailand, and Cambodia (Nguyen

et al., 2009).

Remarks. Specimens were collected during the

day or at night on trees in mixed secondary forests

of hardwood, bamboo and shrub.

Draco maculatus (Gray, 1845)

Examined material. One adult male IEBR

3671, collected by NQT, 12 April 2008, 150 m asl;

First report on the herpetofauna of TayYen Tu Nature Reserve, northeastern Vietnam

529

Figure 34. Chiromantis vittatus. Figures 35, 36. Kurixalus bisacculus. Figures 37, 38. Polypedates mutus.

Photos by T. Ziegler and C.T. Pham.

530

Vera L. Hecht et alii

Figure 39. Polypedates megacephalus. Figure 40. Rhacophorus maximus. Figure 41. R. rhodopus. Figure 42.

Theloderma asperum. Figure 43. T. corticate. Figure 44. T. lateriticum. Photos by T. Ziegler and C.T. Pham.

First report on the herpetofauna of TayYen Tu Nature Reserve, northeastern Vietnam

531

one adult female IEBR A.0946, collected by TZ and

NQT, 27 May 2009, 150 m asl; and one adult fe-

male VNMN 1347, collected by TZ and NTT, May

2010, ca. 350 m asl (Fig. 46).

Morphological characters. SVL male

71.93 mm, females 75.75-78.97 mm (n = 2), TaL

male 116.29 mm, females 113.18-131.51 mm (n

= 2); head small; upper head scales heteroge-

neous, strongly keeled; one spiny scale present on

the back of the eyebrow arch; tympanum covered

by small scales: two incisors on the upper jaw;

nostrils directing laterally outwards; supralabials

7-9, smooth; infralabials 8-9; gular pouch trian-

gular, covered by small scales, very long in males

(often twice longer than head length), shorter in

females; patagium supported by five ribs; dorsal

scales heterogeneous, mostly smooth; lateral

scales enlarged; scales on the back edges of thighs

and tail base fringe-like; ventral scales smooth or

feebly keeled, as large as or slightly smaller than

dorsal scales; forelimbs reaching over the tip of

the snout, hind limbs reaching to elbow or armpit.

Colouration in life: dorsal head and body grey-

ish-brown; patagium with variable markings, vary-

ing in form and colour (determination after

Boulenger, 1912; Manthey & Grossmann, 1997;

Bourret, 2009).

Distribution. This is a widespread species in

Vietnam, from Bac Kan Province in the North to

Lam Dong and Ba Ria-Vung Tau provinces in the

South. This is the first record of D. maculatus from

Tay Yen Tu NR and also from Bac Giang Province.

Elsewhere, the species is known from India, China,

Myanmar, Laos, Thailand, Cambodia, and Malaysia

(Nguyen et al., 2009).

Remarks. Specimens were collected during the

day in the secondary forest.

Physignathus cocincinus Cuvier, 1829

Adult males, females as well as juveniles were

observed during our field work in Yen Tu NR (Fig.

47). They can be easily identified due to their char-

acteristic colour pattern, the large sizes, and the

long tail, gular pouch and supraorbital spines ab-

sent; dorsal crest well developed in males. Coloura-

tion in life: dorsal surface green with a touch of

rost-brown and several very narrow transverse

white stripes on the body and anterior part of the

tail of which the first ones on the body directing

backwards; wide black rings on tail; throat whitish;

belly yellowish green (determination after Ziegler,

2002 ).

Remarks. Several individuals of P. cocincinus

were seen on trees, about 1-2 m above the water in

Suoi Tuyen I and other large streams in Ba Bep area.

EUBLEPHARIDAE

Goniurosaurus lichtenfelderi (Mocquard, 1897)

Examined material. Three adult males IEBR

3695, ZFMK 89229, IEBR A.0827 and two adult

females IEBR 3696, ZFMK 89228, collected by

NQT between 12 and 15 April 2008, 250-300 m asl

(Fig. 48).

Morphological characters. Body shape ro-

bust, SVL 77.1-104.3 mm; external nares bor-

dered by 5-6 nasal scales; supraorbital region with

a row of slightly enlarged tubercles; outer surface

of upper eyelid composed of granular scales, about

one-half the size of those on top of head and with-

out enlarged tubercles; internasals 1-2; supral-

abials 8-10; preorbital scales 15-19; eyelid fringe

scales 51-58; postmentals 2-5; gular region below

lower jaws without enlarged tubercles; paraverte-

bral tubercles 21-27; scale rows around midbody

117-130, granular scales surrounding tubercles

11-13; axillary pockets shallow; subdigital lamel-

lae under toe IV 18-22; precloacal pores in males

30-32, in females 17-21. Colouration in life: iris

reddish-brown; dorsal ground colour of head, body

and limbs brown, without small dark brown

blotches; nuchal loop thin, posteriorly rounded (in

U-shape); dorsal body bands between limb inser-

tions 2, thin, immaculate yellow; gular region with-

out dark spots (Smith, 1935; Ziegler et al., 2008c;

Nguyen, 2011).

Distribution. This species has been recorded

only from northeastern Vietnam (Nguyen et al.,

2009; Nguyen, 2011).

Remarks. Specimens of G. lichtenfelderi were

collected on the forest floor, under decayed trees or

under rocks near streams. This species is nocturnal

and it inhabits the mixed forest of wooden trees,

bamboo, and shrubs.

532

Vera L. Hecht et alii

GEKKONIDAE

Hemidactylus frenatus Schlegel, 1836

Examined material. Two adult females IEBR

A. 2013. 86 and ZFMK 92832, collected by TZ and

NQT, 26 May 2009, 150 m asl (Fig. 49).

Morphological characters. SVL 58.43-

59.08 mm (n = 2), tail regenerated; head covered

with small scales; snout obtuse, longer than distance

between eye and tympanum (SL 7.68-8.30 mm, ET

4.23-4.57 mm); eye covered by transparent mem-

brane, without moveable eyelid; tympanum small,

rounded, longer than half of ED (TD 2.25 mm, ED

3.44-3.77 mm); supralabials 10-11 + 5-7; infral-

abials 8-10 + 3-7; dorsal scales small, grainy; ven-

tral scales imbricate; finger more or less

paddle-like, broadened at base, with divided trans-

verse or oblique lamellae under fingers; 5-6 divided

lamellae under thumb; 9-10 divided lamellae under

finger IV; terminal segments of finger I to V free,

with claw; 12-13 broadened lamellae under toe IV;

midbody scales in 119-121 rows; 10-12 broadened

femoral scales on each side; tail strongly dorso-

ventrally compressed, with lateral denticulation (de-

termination after Bourret, 2009).

Distribution. This is a common species in Viet-

nam. Elsewhere, the species is known from India,

Nepal, Sri Lanka, Maldives, China, Taiwan, Myan-

mar, Thailand, Malaysia, Indonesia, Philippines,

New Guinea, Australia, Japan, Polynesia, Microne-

sia, Melanesia, Solomon Islands, Somalia, Madagas-

car, Mauritius, Reunion, Rodrique, Comoro Island,

Samoa, and New Caledonia (Nguyen et al., 2009).

Remarks. Both specimens were found at night

on the walls of Vung Tron Ranger Station.

Gekko palmatus Boulenger, 1907

Examined material. One adult female IEBR

3638 (SVL 73.53 mm) collected by NQT and PTC

on 8 April 2008 and one adult male IEBR

A.2013.75 (SVL 64.32 mm) collected by TZ, NTT

and PTC in October 2009, ca. 300 m asl (Fig. 50).

Morphological characters. Moderate-sized

gecko (SVL < 80 mm); nares in contact with rostral;

intemasal single, smaller than supranasal; postmen-

tals enlarged; interorbital scales between anterior

comers of the eyes 30 and 36; dorsal tubercle rows

9 and 1 1 ; ventral scales between mental and cloacal

slit 163 and 183; midbody scale rows 123 and 142;

ventral scale rows 40 and 44; subdigital lamellae

under toe I 11 or 12, under toe IV 12 or 13; finger

and toe webbing present at base; tubercles absent

on upper surface of fore and hind limbs; precloacal

pores 24 in the male, absent in the female; postcloa-

cal tubercle single; tubercles present on dorsal sur-

face of tail base; subcaudals enlarged. Colouration

in life: dorsal surface of head and body grey with a

small light blotch on neck and four larger blotches

between shoulder and sacrum; flanks with some

small light spots between limb insertions; limbs

with light spots and bars; dorsal tail with 8 or 9 light

bands; throat, venter, and precloacal region yellow-

ish cream with dark dots (identification after

Nguyen et al., 2013).

Distribution. In Vietnam, this species has been

recorded from Yen Bai, Lang Son, Vinh Phuc,

Quang Ninh, and Quang Binh provinces. The

record of G. palmatus from Bac Giang Province

was previously confused with G. chinensis Gray,

1842 (Nguyen et al., 2009, 2013). Elsewhere, the

species is known from China (Nguyen et al., 2009).

Remarks. Both specimens were found at night

on rocky walls along a stream near Dong Ri Ranger

Station.

LACERTIDAE

Takydromus kuehneiVm Denbourgh, 1909

Examined material. One adult male VNMN

1330, collected by TZ and NTT, May 2010, ca. 200

m asl (Fig. 51).

Morphological characters. SVL 52.95 mm,

TaL 167.32 mm; supralabials 7; infralabials 6-7; chin

shields in 4 pairs; dorsal scales in 6 rows at midbody,

without a non-contiguous vertebral row of smaller

scales; ventral scales widened, in 6 rows at midbody;

lateral scales in 12 rows at midbody on each sides,

smaller in size than dorsal and ventral scales; femoral

pores 4 on each side; subdigital lamellae broadened.

Colouration in life: upper surface of head and body

brown, ventral surface whitish-cream, dorsal surface

of limbs and tail base with black spots; upper part of

flanks black with numerous yellowish spots, upper

part of flanks light brown; head dark brown with

First report on the herpetofauna of TayYen Tu Nature Reserve, northeastern Vietnam

533

some black spots (determination after Ziegler &

Bischoff, 1999; Ziegler, 2002).

Distribution. In Vietnam, this species has been

recorded from Quang Ninh, Son La, Hoa Binh,

Ninh Binh, Thanh Hoa, Nghe An, and Ha Tinh

provinces. This is the first record of T. kuehnei from

Tay Yen Tu NR and from Bac Giang Province. Else-

where, the species has been reported from China

(Nguyen et al., 2009).

Remarks. The specimen was collected in the

evening, on a ferny leaf in the secondary forest.

SHINISAURIDAE

Shinisaurus crocodilurus Ahl, 1930

During our night excursions in summer 2010,

several individuals of crocodile lizards have been

found and susequently released after taking mea-

surements and photographs. Maximum SVL was

167.97 mm, maximum TaL was 208.91 mm. The

morphological characters accorded with the de-

scriptions provided by Le & Ziegler (2003) and

Ziegler et al. (2008a) (Fig. 52).

Distribution. This species is only known from

southern China and northeastern Vietnam (Nguyen

et al., 2009).

Remarks. Most specimens of S. crocodilurus

were found on branches or palm tree leaves above

rocky streams in the evergreen forest at elevations

between 364-450 m asl, only one was seen hiding

under a rock. Further ecological data of this species

from Tay Yen Tu NR will be published elsewhere.

SCINCIDAE

Ateuchosaurus chinensis Gray, 1845

Examined material. One juvenile female IEBR

A.0947, collected by TZ and NTT, October 2009,

150 m asl (Fig. 53).

Morphological characters. SVL 43. 14 mm,

tail lost; supranasals absent; prefrontals small, sep-

arated, touching both loreals laterally; frontal an-

teriorly truncated, approximately three times

longer than wide, twice longer than frontoparietal

and interparietal together, twice longer than its dis-

tance to tip of snout; frontoparietals separated

from each other, as long as interparietal; a small

transparent spot present on interparietal; parietals

small, in contact; nuchals absent; nostril in a single

nasal; supraoculars 4; supralabials 6; infralabials

7; postmental undivided; chin shields in 2 pairs;

lower eyelid scaly; tympanum deeply sunken; ven-

tral scales in 30 rows at midbody; paravertebral

scales 53; limbs with short lamellae, 17-18 under

toe IV. Colouration in life: dorsum brown, each

scale with a darker spot in the center; flanks mot-

tled with black and white spots; ventral surface

cream (determination after Nguyen et al., 2008;

Bourret, 2009).

Distribution. In Vietnam, this species has been

recorded from Ha Giang, Lang Son, Bac Giang, and

Nghe An provinces. Elsewhere, the species is

known from China (Nguyen et al., 2009).

Remarks. The juvenile female was found at

night (22:00), under leaf litter in the bamboo forest

near Pond Khe Cam 1 . The number of paravertebral

scales is somewhat higher than in Nguyen et al.

(2008) (53 vs. 48-51).

Eutropis longicaudatus (Hallowell, 1856)

Examined material. One adult male IEBR

A.2010.13, collected by NQT, August 2001, 200-

300 m asl.

Morphological characters. SVL 79.56

mm; supranasals in contact; frontonasal broader

than long; postnasal single; prefrontals touching

each other; lower eyelid scaly; supraoculars 4;

supraciliaries 6; supralabials 7; infralabials 5; pari-

etals separated; nuchals in 1 pair; tympanum deeply

sunken; dorsal scales bicarinate; midbody scales

in 27 rows; limbs overlapping when adpressed

along the body; lamellae under toe IV 26-27; pre-

cloacal shields 4, enlarged. Colouration in alco-

hol: dorsal head and body brown; lateral sides

dark brown to black; ventral surface cream (de-

termination after Manthey & Grossmann, 1997;

Ziegler, 2002; Bourret, 2009; Nguyen et al.,

2011b).

Distribution. This is one of the most common

species in Vietnam. Elsewhere, the species has been

reported from China, Taiwan, Laos, Thailand, Cam-

bodia, and Malaysia (Nguyen et al., 2009).

534

Vera L. Hecht et alii

Figure 45. Acanthosaura lepidogaster. Figure 46. Draco maculatus. Figure 47. Physignathus cocincinus. Figure 48. Go-

niurosaurus lichtenfelderi. Figure 49. Hemidactylus frenatus . Figure 50. Gekko palmatus. Figure 51. Takydromus kuehnei.

Figure 52. Shinisaurus crocodilurus . Photos by T.Q. Nguyen, T. Ziegler and C.T. Pham.

First report on the herpetofauna of TayYen Tu Nature Reserve, northeastern Vietnam

535

Remarks. The adult male was found in the mor-

ning in a plantation forest near Dong Thong Village.

Eutropis multifasciatus (Kuhl, 1820)

Morphological characters. The identification

was based on direct observations and photographs

(Fig. 54): head covered with large shields, pre-

frontals in broad contact; tympanum deeply sunken;

supranasals present, separated from each other;

lower eyelid scaly; one single postnasal; limbs well

developed, dorsal scales tri-carinate; midbody scales

in 30 rows; dark lateral band present (determination

after Smith, 1935; Ziegler, 2002; Bourret, 2009).

Distribution. This is a common species in Viet-

nam (Nguyen et al., 2009). Elsewhere, the species

has been reported from India, throughout China, In-

dochina southwards to the Philippines and New

Guinea (Nguyen et al., 2009).

Remarks. One subadult specimen was observed

in the evening of 8 June 2010 near the Dong Thong

Ranger Station.

Plestiodon tamdaoensis (Bourret, 1937)

Examined material. One adult VNMN 1351,

collected by TZ and NTT, June 2010; one adult fe-

male IEBR A.2010.06, collected by NQT, August

2001, ca. 300 m asl (Fig. 55).

Morphological characters. SVL male 120.2

mm, female 87.6 mm, tail of the male regenerated

(TaL 108.7 mm), of the female lost; supranasals large,

in contact with each other; postnasal single; postmen-

tals 2; prefrontals in contact with each other; loreals

3; lower eyelid scaly; supraoculars 4; supraciliaries

8-9; frontoparietals in contact with each other; inter-

parietal larger than frontoparietals; parietals sepa-

rated; nuchals in 2 pairs; supralabials 8-9; infralabials

7; tympanum deeply sunken, with 3 small lobules on

the anterior edge; dorsal scales smooth; midbody

scales in 24 rows; paravertebral scales 42; lamellae

under toe IV 19; precloacal shields 2, enlarged; limbs

overlapping when adpressed along body. Colouration

in alcohol: dorsal head and body brown; lateral band

black-brown; ventral surface cream (determination

after Hikida et al., 2001; Bourret, 2009).

Distribution. In Vietnam, this species has been

recorded from Ha Giang, Cao Bang, Bac Kan, Vinh

Phuc, Bac Giang, Hai Duong, Hoa Binh, and Nghe

An provinces. Elsewhere, the species is known

from China (Nguyen et al., 2009).

Remarks. Both specimens were found at night

in the mixed secondary forest.

Sphenomorphus cryptotis Darevsky, Orlov et

Ho, 2004

Examined material. Two adult males ZFMK

92833 and VNMN A.2013.16, collected by TZ and

NTT, October 2009, ca. 500 m asl (Fig. 56).

Morphological characters. SVL 70.50-

78.06 mm, TaL 127.67 mm; supranasals absent;

rostral touching frontonasal; prefrontals in contact

with each other; parietals in contact posteriorly;

supraoculars 4; supralabials 7-8; infralabials 7;

postmental undivided; primary temporal single;

external ear present, without lobules, tympanum su-

perficial; dorsals larger than laterals and ventrals;

midbody scales in 34-36 rows; paravertebral scales

75; ventrals 80; limbs well developed, overlapping

when adpressed along body; lamellae under toe IV

18-19. Colouration in life: dorsum and tail base

bronze brown with a vertebral row of large black

blotches; upper lateral zone with a dark grey stripe,

in width of 4-5 scales, from behind eye to tail base,

paler on distal tail; two rows of light spots on upper

and lower margins of the dark stripe; lower lateral

zone light grey (determination after Darevsky et al.,

2004; Nguyen, 2011; Nguyen et al., 2011a).

Distribution. This species is known only from

Vietnam, in Lao Cai, Quang Ninh, Bac Giang, and

Nghe An provinces (Nguyen et al., 2009; Nguyen,

2011 ).

Remarks. Both specimens were found at night

on leaves, ca. 0. 2-0.5 m above the stream and on

the rock in Suoi Tuyen I.

Sphenomorphus incognitus (Thompson, 1912)

Examined material. Two subadults TYT 62,

150, three adult females TYT 644, 675, 676, col-

lected by TTT in May 2006; one adult male IEBR

A. 0823, two sudadults IEBR A.0825-A.0826

collected by NQT and NVS in August 2001; one

subadult female IEBR 3637, two adult females

IEBR 3687-3688, collected by NQT between 9 and

536

Vera L. Hecht et alii

13 April 2008, 200^100 m asl; one subadult ZFMK

92834, collected by TZ and NTT, October 2009, ca.

350 m asl (Fig. 57).

Morphological characters. Size (SVL 79.6-

103.9 mm, n = 6); prefrontals separated from each

other; supralabials 7; primary temporals 2; external

ear present, without lobules, tympanum deeply

sunken; midbody scales in 36-44 rows; dorsal

scales smooth, paravertebral scales 67-80, not

widened; limbs well developed; subdigital lamellae

under toe IV 19-24. Colouration in life: dorsum and

tail base greyish brown or brown with irregular

black dots; dorsolateral area with white spots; upper

lateral zone with a black stripe, not clearly defined,

in width of 3-4 scales, from behind eye to tail base,

interrupted by light spots; lower lateral zone light

grey with dark marbling or spots (determination

after Nguyen et al., 2012).

Distribution. S. incognitus was recorded for

the first time from Vietnam by Nguyen et al. (2012).

Elsewhere, the species is known from China and

Taiwan (Nguyen et al., 2012).

Remarks. S. incognitus inhabits the mixed for-

est of small wooden trees, bamboo, and shrub, at

elevation between 200-400 m. Specimens were

found both on the forest floor and in streams.

Sphenomorphus indicus (Gray, 1853)

Examined material. One adult VNMN 1329,

collected by TZ and NTT, 26 June 2010, ca. 300 m

asl (Fig. 58).

Morphological characters. SVL 75.91 mm,

tail lost; supranasals absent; prefrontals in contact

with each other; parietals slightly in contact;

nuchals absent; lower eyelid scaly; supraoculars 4;

supraciliaries 8; supralabials 7; infralabials 7; pri-

mary temporals 1/2; external ear present, with 1-3

very small lobules or without lobules, tympanum

deeply sunk; postmental undivided; midbody scales

in 36 rows; limbs overlapping when adpressed

along body; lamellae under toe IV 17-18. Coloura-

tion in life: dorsum and tail base bronze brown with

irregular black dots; light dorsolateral stripe present

on neck and shoulder; upper lateral zone with a dark

stripe, in width of 2-3 scales, from behind eye to

tail base, paler on distal tail, light bars sometimes

present on lower lateral zone; ventral surface

whitish (determination after Smith, 1935; Manthey

& Grossmann, 1997; Ziegler, 2002; Nguyen, 2011;

Nguyen et al., 2011a).

Distribution. S. indicus is a common species

in Vietnam. However, this is the first record of the

species from Tay Yen Tu NR and from Bac Giang

Province. Elsewhere, the species is known from

India, Bhutan, China, Taiwan, Myanmar, Laos,

Thailand, Cambodia, Malaysia, and Indonesia

(Nguyen et al., 2009).

Remarks. A single specimen was found in the

evening on the forest ground near Suoi Tuyen I.

Sphenomorphus tonkinensis Nguyen, Schmitz,

Nguyen, Orlov, Bohme et Ziegler, 2011

Examined material. One adult male VNMN

1331, collected by TZ and NTT, 26 June 2010, ca.

400 m asl (Fig. 59).

Morphological characters. SVL 42.82 mm,

TaL 5 1 .43 mm; head longer than wide; supranasals

absent; prefrontals in contact with each other;

supralabials 7; infralabials 6; nuchals absent; pri-

mary temporals 2; external ear present, without

lobules, tympanum slightly sunken; limbs overlap-

ping when adpressed along body; midbody scales

in 34 rows; dorsal scales smooth; paravertebrals

74; lamellae under toe IV 15. Colouration in life:

dorsum and tail base bronze brown with black,

discontinuous vertebral line reaching to first third

of tail; upper part of flanks with black bars, inter-

rupted by small, light spots in the neck; tail orange

brown; ventral surface cream (determination after

Nguyen et al., 2011a).

Distribution. In Vietnam, this species has been

recorded from Vinh Phuc, Quang Ninh, and Hai

Phong provinces. This is the first record of S. tonk-

inensis from Tay Yen Tu NR as well as from Bac

Giang Province. Elsewhere, the species is known

from China (Nguyen et al., 2011).

Remarks. The specimen was found at night in

forest near Suoi Tuyen II.

Tropidophorus hainanus Smith, 1923

Examined material. One adult male ZFMK

92835, collected by TZ and NTT, October 2009, ca.

400 m asl; one adult male VNMN 1343, collected

by TZ and NTT, June 2010, 440 m asl (Fig. 60).

First report on the herpetofauna of TayYen Tu Nature Reserve, northeastern Vietnam

537

Morphological characters. SVL 37.77^42.31

mm (n = 2), TaL: 46.86 mm; head longer than wide;

head shields striated; supranasals absent; frontonasal

undivided; prefrontals separated from each other;

tympanum superficial; supralabials 6-7; infralabials

5; postmental undivided; nuchals absent; limbs over-

lapping when adpressed along body; dorsals and lat-

erals keeled; midbody scales in 30-32 rows; ventrals

43-44, cycloid; paravertebral scales 44; lamellae

under toe IV 16-18; precloacal shields 2, enlarged.

Colouration in life: dorsal head and body brown,

with bright, narrow, transverse bars bordered with

black, the first two V-shaped; flanks cream, upper

part of flanks with large whitish dark bordered spots;

ventral surface white; throat with dark longitudinal

lines (determination after Bourret, 2009 and Nguyen

et al., 2010b).

Distribution. In Vietnam, this species has been

recorded from Lai Chau, Lao Cai, and Ha Giang

provinces southwards to Dak Lak and Dak Nong

provinces. Elsewhere, the species is known from

China (Nguyen et al., 2009).

Remarks. One specimen (ZFMK 92835) was

found at night on a rock in Suoi Tuyen I and another

specimen (VNMN 1343) was collected on the for-

est ground near Ba Bep Pond in the late morning

(11:40).

Tropidophorus sinicus Boettger, 1886

Examined material. One adult female VNMN

A.2013.17 and one adult male ZFMK 92836, col-

lected by TZ and NTT, October 2009, 400-600 m

asl (Fig. 61).

Morphological characters. SVFmale 53.12

mm, female 60.74 mm, TaF male 66.69 mm, female

76.24 mm; head longer than wide; head shields

striated; supranasals absent; frontonasal divided;

prefrontals in contact; loreal single; tympanum dis-

tinct, superficial; supralabials 5; infralabials 5; post-

mental divided; midbody scales in 32 rows; dorsal

scales keeled; paravertebral scales 43-45; lamellae

under toe IV 17-21 . Colouration in life: dorsal head

and body dark brown, with large, transverse pale

yellow bars; flanks with smaller light spots; ventral

surface white (determination after Bourret, 2009;

Nguyen et al., 2010b; Nguyen, 2011).

Distribution. In Vietnam, this species has been

recorded from Cao Bang, Bac Kan, Lang Son, Bac

Giang, Hai Duong, and Quang Ninh provinces.

Elsewhere, the species is known from China

(Nguyen et al., 2009).

Remarks. Both specimens were found on rocks

in Suoi Tuyen I.

SERPENTES

CALAMARIIDAE

Calamaria pavimentata Dumeril etBibron, 1854

Examined material. One juvenile VNMN 1349

(SVL 134.78 mm, TaL 10.33 mm), collected by TZ

and NTT, June 2010, ca. 250 m asl (Fig. 62).

Morphological characters. Head not distinct

from neck; intemasals absent; rostral as wide as

high, clearly visible from above; frontal longer than

wide but shorter than parietals, approximately two

times wider than supraocular; pupil round; loreal ab-

sent; preocular single; postocular single; temporals

absent; supralabials 4, second and third touching the

eye, fourth largest; infralabials 5, first touching men-

tal; mental groove present; chin shields in 2 pairs, in

contact medially; dorsal scales smooth, in 13:13:13

rows; ventrals 160; cloacal shield undivided; sub-

caudals 24, divided; tail with pointed tip. Coloura-

tion in life: dorsum brown, dark collar in nuchal

region, outermost dorsal scale rows with dark line;

neck with orange transverse band; ventral surface

yellowish; tail with 2 yellow spots at base and 2

other ones close to the tip, ventral surface with a thin

dark median line (determination after Smith, 1943;

Ziegler et al., 2008b; Nguyen et al., 2009).

Distribution. In Vietnam, this species has been

recorded from Lao Cai and Cao Bang provinces

southwards to Quang Nam and Lam Dong prov-

inces. This is the first record of C. pavimentata from

Tay Yen Tu NR and from Bac Giang Province. Else-

where, the species is known from China, Myanmar,

Laos, Cambodia, Thailand, Malaysia, Indonesia,

and Japan (Nguyen et al., 2009).

Remarks. The specimen was found at night in

the mixed forest of bamboo and small hardwood.

Calamaria septentrionalis Boulenger, 1890

Examined material. One adult male IEBR

3713 (SVL 287.36 mm, TaL 28.45 mm), collected

538

Vera L. Hecht et alii

Figure 53. Ateuchosaurus chinensis. Figure 54. Eutropis multifasciatus . Figure 55. Plestiodon tamdaoensis. Figure 56.

Sphenomorphus cryptotis. Figure 57. S. incognitus. Figure 58. S. indicus. Figure 59. S. tonkinensis. Figure 60. Tropidophorus

hainanus. Figure 6F T. sinicus. Photos by T. Ziegler, T.Q. Nguyen, and C.T. Pham.

First report on the herpetofauna of TayYen Tu Nature Reserve, northeastern Vietnam

539

by NQT, NTT and PTC, 18 June 2009, ca. 350 m

asl (Fig. 63).

Morphological characters. Head not distinct

from neck; intemasals absent; rostral as wide as

high, barely visible from above; frontal longer than

wide but shorter than parietals, 2.5 times wider than

supraocular; pupil round; loreal absent; preocular

single; postocular single; temporals absent; supral-

abials 4, second and third touching the eye, fourth

largest; infralabials 5, first touching mental; mental

groove present; chin shields in 2 pairs, in contact

medially; dorsal scales smooth, in 13:13:13 rows;

ventrals 157; cloacal shield undivided; subcaudals

17, divided; tail tapering with rounded end. Coloura-

tion in life: dorsum dark brown; outermost dorsal

scale rows yellow, edged in black below; neck with

two large orange blotches; ventral surface yellow-

ish; ventral scales with dark outermost comers; tail

with 2 yellow spots at base and 2 other ones close

to the tip, ventral surface with a thin dark median

stripe (determination after Smith, 1943; Ziegler et

al., 2008b; Nguyen et al., 2009).

Distribution. In Vietnam, this species has been

recorded from Lao Cai and Ha Giang southwards to

Ha Tinh and Quang Binh provinces. Elsewhere, the

species is known from China (Nguyen et al., 2009).

Remarks. The specimen was found at night in

the bamboo forest near Mau Village.

COLUBRIDAE

Ahaetulla prasina (Boie, 1827)

This is a widespread species in Vietnam (Fig.

64) . Elsewhere, the species is known from India

throughout China and Indochina, southwards to In-

donesia and the Philippines (Nguyen et al., 2009).

One specimen was seen during the day time in

June 2010 in the bamboo forest near Mau Village.

Species identification was based on photographs:

Head very long, with a very long, pointed snout;

canthus rostralis very sharp; pupil horizontal; body

green, with interstitial white and black stripes (de-

termination after Smith, 1943; Ziegler, 2002).

Cyclophiops multicinctus (Roux, 1907)

This is a widespread species in Vietnam (Fig.

65) . However, this is the first record of C. multi-

cinctus from Tay Yen Tu NR and from Bac Giang

Province. Elsewhere, the species is known from

China and Laos (Nguyen et al., 2009).

One individual was seen at night in October

2009 in branches above a forest stream near Dong

Ri Ranger Station. Identification was based on pho-

tographs: head and anterior part of body green,

posterior part of body and tail brown with some

pale black and white spots forming interrupted bars

on body; ventral surface yellow (determination after

Smith, 1943; Ziegler et al., 2007).

Lycodon futsingensis (Pope, 1928)

Examined material. One adult male IEBR A

2013.87, collected byTZ and NTT, October 2009,

ca. 350 m asl; one adult male VNMN 1350, col-

lected by TZ and NTT, June 2010, ca. 450 m asl

(Fig. 66).

Morphological characters. SVL 512.52-

527.51 mm (n = 2), TaL 145.02-147.76 mm (n

2): head distinct from neck; snout long, prominent;

pupil vertically elliptic; rostral large; internasals

wider than long, not touching loreal and preocular;

frontal hexagonal; parietals longer than wide; nasal

divided; loreal single, not entering orbit; preocular

single; postoculars 2; temporals 2+3; supralabials

8, third to fifth or fourth and fifth entering orbit; in-

fralabials 8-9; mental groove present; dorsal scales

smooth, in 17:17:15 rows; vertebrals not enlarged;

ventrals 3-4 + 198-200, laterally distinctly angu-

lated; cloacal shield undivided; subcaudals 76-81,

divided. Colouration in life: back brownish grey to

dark brown, with 23-24 light brown bands on body,

12 bands on tail; first band starting at ventrals 15-

20; some bands in Y-shape, more distinct in poste-

rior part of body; head grey with a lighter band,

from eye to neck; belly cream, posterior part mot-

tled; lower surface of tail dark brown (determina-

tion after Bourret, 1936; Vogel et al., 2009).

Distribution. In Vietnam, this species has been

recorded from Lao Cai and Cao Bang provinces

southwards to Quang Binh and Da Nang provinces.

Elsewhere, the species is known from China

(Nguyen et al., 2009).

Remarks. Both specimens were found at night

in Suoi Tuyen I Stream. One was found on a root

near the water and the other specimen was collected

on a tree branch, about 2 m above the ground.

540

Vera L. Hecht et alii

Ly codon meridionalis Bourret, 1936

Examined material. One adult female VNMN

1333, collected by TZ and NTT, 24 June 2010, ca.

360 m asl (Fig. 67).

Morphological characters. SVL 632.13 mm,

TaL 164.23 mm; head distinct from neck; rostral

twice as wide as high; intemasals wider than long;

prefrontals wider than long; frontal small, narrow-

ing posteriorly, shorter than parietals; posterior

nasal longer than anterior one; pupil vertically el-

liptic; loreal single, entering orbit at a point, not

touching intemasals; preocular single; postoculars

2; temporals 2+2/3; supralabials 8/9, third to fifth

or fourth and fifth touching the eye; infralabials

9/10; chin shields in 2 pairs; mental groove present;

dorsal scale rows 17:17: 15; outer dorsal scale rows

not keeled; vertebral scales not enlarged; ventrals

2+231ventrals; cloacal shield undivided; subcau-

dals 98, divided. Colouration in life: dorsum black,

with 93 yellow transverse bands on body and 24 on

tail; upper surface of head black with yellow su-

tures; flanks yellow with irregular black spots; ven-

tral surface yellow; subcaudals dark brown to black

with light sutures (determination after Bourret,

1936; Smith, 1943).

Distribution. In Vietnam, this species has been

recorded from Lao Cai and Ha Giang provinces

southwards to Ninh Binh Province. Elsewhere, the

species is known from China and Laos (Nguyen et

al., 2009).

Remarks. The specimen was found in the

evening (ca. 19:00) on a branch, about 1.6 m

above the water in Suoi Tuyen I Stream. Some

other individuals were seen on branches about 1-

2 m above the ground.

Oligodon chinensis (Gunther, 1888)

Examined material. One adult male IEBR

3708, collected by NQT, NTT and PTC, 17 June

2009, ca. 300 m asl; one adult female VNMN 1352,

collected by TZ and NTT, June 2010, ca. 250 m asl

(Fig. 68).

Morphological characters. SVL male 460.2

mm, female 574.06 mm, TaL male 125.85 mm, fe-

male 103.87 mm; head indistinct from neck; inter-

nasals 2; prefrontals 2; frontal longer than its

distance to tip of snout, as long as parietals; nasal

divided or undivided; loreal single; preocular sin-

gle; postoculars 2; suboculars absent; temporals

1+2; supralabials 8, fourth and fifth touching the

eye; infralabials 8-9; mental groove present; dorsal

scale rows 17:17:15; ventrals 1-2 + 173-182,

slightly laterally angulated; cloacal shield undi-

vided; subcaudals 52-63, divided. Colouration in

life: dorsal surface brown to reddish brown with

11-13 distinct dark spots on body and 4 on tail;

narrow, interrupted, black bands present between

those spots; an inverse V-marking present behind

head, distinct; dark spot on temporal region present

or absent; belly white, with rectangular blotches

(determination after Bourret, 1936; Smith, 1943;

David et al., 2008).

Distribution. In Vietnam, this species has been

recorded from Lao Cai and Lang Son southwards

to Gia Lai provinces. This is the first record of O.

chinensis from Tay Yen Tu NR and from Bac Giang

Province. Elsewhere, the species is known from

China (Nguyen et al., 2009).

Remarks. Both specimens were found in the

mixed secondary forest of small hardwood and

bamboo. Another specimen was observed in the

evening (20:30) near the stream close to Khe Cam

Pond on 4 July 2010. Our specimens agreed with

the descriptions of Bourret, (1936), Smith, (1943)

and David et al. (2008), but showed some minor

differences: dorsal dark spots cover only two in-

stead of three vertebral scales and nasal of IEBR

3708 divided on one side and undivided on the

other side.

Rhadinophis prasinus (Blyth, 1854)

Examined material. One adult male VNMN

1335, collected by TZ and NTT, 26 June 2010, ca.

400 m asl (Fig. 69).

Morphological characters. SVL 596.98 mm,

TaL 230.71 mm; head distinct from neck; pupil

round; intemasals in contact with each other, wider

than long but shorter than prefrontals; rostral wider

than high; prefrontals 2; frontal longer than wide,

as long as its distance to tip of snout, shorter than

parietals, not touching preocular; loreal single;

preocular single; postoculars 2; temporals 2+2/3;

supralabials 9, fourth to sixth touching the eye; in-

fralabials 8; mental groove present; dorsal scale

rows 19:19:15; ventrals 6+196; cloacal shield di-

First report on the herpetofauna of TayYen Tu Nature Reserve, northeastern Vietnam

541

Figure 62. Calamaria pavimentata. Figure 63. C. septentrionalis. Figure 64. Ahaetulla prasina. Figure 65. Cyclophiops

multi cinctus . Figure 66. Lycodon futsingensis . Figure 67. L. meridionalis . Figure 68. Oligodon chinensis. Photos by

T.Q. Nguyen and T. Ziegler.

542

Vera L. Hecht et alii

vided; subcaudals 111, divided. Colouration in life:

dorsal surface entirely green; flanks yellowish

green; interstitial skin black and white, more dis-

tinct in the anterior part of body; belly whitish-

green (determination after Bourret, 1936; Smith,

1943; Manthey & Grossmann, 1997).

Distribution. In Vietnam, this species has been

known from Lao Cai, Bac Kan, Thai Nguyen, Vinh

Phuc, Nghe An, Quang Binh, and Gia Lai prov-

inces. This is the first record of R. prasinus from

Tay Yen Tu NR as well as from Bac Giang Pro-

vince. Elsewhere, the species is reported from India,

China, Myanmar, Laos, Thailand, and Malaysia

(Nguyen et al., 2009).

Remarks. A single specimen was found in the

evening (19:00) on a tree branch, approximately

1 .60 m above the ground in Suoi Tuyen I. Our speci-

men differs from the descriptions of Bourret (1936)

and Manthey & Grossmann (1997) in having three

instead of two posttemporals on the left side.

Rhynchophis boulengeri (Mocquard, 1897)

Examined material. One adult male IEBR

A.20 13.88, collected by TZ and NTT, October

2009, ca. 350 m asl (Fig. 70).

Morphological characters. SVL 645.15

mm, TaL 252.28 mm; head long, distinct from

neck; conical appendix present, covered by small

scales on upper jaw, directing upwards, somewhat

longer than its distance to eye; pupil round; nostril

surrounded by 2 nasals; internasals half size of pre-

frontals; frontal broad anteriorly, shorter than pari-

etals; loreal single, longer than high; preocular

single, touching frontal; postoculars 2; temporals

2+2+3; supralabials 9, fourth to sixth touching eye;

infralabials 10; dorsal scales smooth, in 19:19:15

rows; mental groove present; ventrals 221; cloacal

shield divided; subcaudals 137, divided. Coloura-

tion in life: upper surface green; a black line run-

ning from nostril to the eye and continuing to neck;

ventral surface light green; laterally angulated re-

gion of ventrals white (determination after Bourret,

1936; Pope, 1935; Smith, 1943).

Distribution. In Vietnam, this species has been

recorded from Son La, Thai Nguyen, Vinh Phuc,

Hanoi, Quang Ninh, Hai Phong, and Ha Tinli prov-

inces. This is the first record of R. boulengeri from

Tay Yen Tu NR as well as from Bac Giang Prov-

ince. Elsewhere, the species is known from China

(Nguyen et al., 2009).

Remarks. The adult male was found in the

evening (19:00) on a tree branch, about 1.60 m

above the ground at Suoi Tuyen I. A small bird was

found in its stomach. The scale counts of IEBR

A2013.88 are somewhat higher than in the descrip-

tion of Smith (1943) (221 ventrals instead of 207-

216 and 137 subcaudals instead of 123-132).

Sibynophis chinensis (Gunther, 1899)

Examined material. One adult male VNMN

1353, collected by TZ and NTT, May 2010, ca. 300

m asl (Fig. 71).

Morphological characters. SVL 367.25 mm,

TaL 153.31 mm, tail tip broken; rostral wider than

high; intemasals 2, in contact with each other; pre-

frontals 2; parietals 2, touching upper postocular;

pupil round; loreal single; preocular single, twice

as high as loreal; postoculars 2; temporals 2+2;

supralabials 9, fourth to sixth touching the eye; in-

fralabials 9; mental groove present; chin shields in

2 pairs; dorsal scales smooth, in 17:17:17 rows;

ventrals 164; cloacal shield divided; subcaudals 75,

divided. Colouration in alcohol: dorsal head olive

grey with two narrow black crossbars behind the

eyes and another large one on neck; a dark stripe

running from nostril to the eye and continuing to

neck; ground colour of body and tail light reddish

grey, dorsum with a vertebrally interrupted line of

dark spots and two interrupted lines of light spots

laterally; ventral surface yellowish- white, darker on

ventral tail; each ventral with one dark spot on lat-

eral edge and two dark spots in the center, these

spots on subcaudals forming dark lines on ventral

tail (determination after Pope, 1935; Bourret, 1936;

Smith, 1943).

Distribution. In Vietnam, this species has been

recorded from Lao Cai, Cao Bang, Bac Kan, Vinh

Phuc, Hoa Binh, Ninh Binh, Quang Tri, and Gia Lai

provinces. This is the first record of S. chinensis

from Tay Yen Tu Nature Reserve and from Bac

Giang Province. Elsewhere, the species is known

from China and Taiwan (Nguyen et al., 2009).

Remarks. The single specimen was found on

the forest floor. The specimen has fewer ventral

scales than reported by Smith (1943) (164 instead

of 168-183).

First report on the herpetofauna of TayYen Tu Nature Reserve, northeastern Vietnam

543

LAMPROPHIIDAE

Psammodynastes pulv erulentus (Boie,1827)

Only one individual was seen at night in Octo-

ber 2009 on branches next to a forest trail about 1 .5

m above the ground (Fig. 72).

Morphological characters. Identification

was based on photographs: head elongated, in

triangular shape; dorsal surface of head and neck

brown with some dark stripes; dorsal body and

tail brown with some light spots, edged in black

(determination after Smith, 1943; Ziegler et al.,

2002 ).

Distribution. Although P. pulv erulentus is

known almost from entire Southeastern Asia and

from Vietnam (Nguyen et al., 2009), this is the first

record of the species from Tay Yen Tu NR as well

as from Bac Giang Province.

NATRICIDAE

Amphiesmoides ornaticeps (Werner, 1924)

Examined material. One subadult male

VNMN 1355, collected by TZ and NTT, June 2010,

ca. 200 m asl (Fig. 73).

Morphological characters. SVL 284.29 mm,

TaL 139.01 mm; head distinct from neck; rostral

hexagonal, wider than high; intemasals 2, in con-

tact, as long as prefrontals, narrowing anteriorly;

prefrontals 2, touching loreal; frontal hexagonal,

longer than wide; nostrils laterally; pupil round; lo-

real single, in contact with nasal; preocular single,

postoculars 3; temporals 2+2/3; supralabials 9,

fourth to sixth entering orbit; infralabials 8; mental

groove present; dorsals strongly keeled, midbody

scale rows 19; ventrals 2+160; cloacal shield di-

vided; subcaudals 122, divided. Colouration in al-

cohol: upper surface of body and tail greyish brown,

anterior part of body with white squarrish net;

flanks cream, with some brown marbling; head

light brown; eye bordered by two vertical white

streaks, edged in black, posterior one large; ventral

surface cream (determination after Pope, 1935;

Nguyen et al., 2010a).

Distribution. In Vietnam, this species has been

recorded from Bac Giang, Hoa Binh, and Nghe An

provinces. Elsewhere, the species is known from

China (Nguyen et al., 2009).

Remarks. A single specimen was found on the

forest floor near Mau Village. Another specimen

was collected in a small stream in bamboo forest

near Mau Village in April 2008.

Opisthotropis lateralis Boulenger, 1903

Examined material. One adult male ZFMK

93904, collected by TZ and NQT, 28 May 2009,

200 m asl (Fig. 74).

Morphological characters. SVL 407.06

mm, TaL 64.61 mm; rostral broader than high; in-

ternasals 2, as long as broad, in contact with each

other; prefrontal single; frontal longer than wide,

shorter than parietals, twice as broad as supraocu-

lar; nostrils directing upwards, in the upper part of

single nasal; pupil round; loreal single, longer than

high, not touching intemasals; preoculars 2; pos-

toculars 2; temporals 1+2; supralabials 10, sixth

touching the eye; infralabials 9; mental groove

present; dorsal scales smooth, in 17:17:17 rows;

ventrals 2+184; cloacal shield divided; subcaudals

49, divided. Colouration in life: dorsal surface dark

greyish brown, with 7 longitudinal lines on back;

one black line on third dorsal scale row; 3 outer

rows of dorsal scales orange; ventral surface yel-

lowish white (determination after Pope, 1935;

Bourret, 1936; David et al., 2011).

Distribution. In Vietnam, this species has been

recorded from Cao Bang, Lang Son, Vinh Phuc,

Quang Ninh, Bac Giang, Hai Duong, and Hoa Binh

provinces. Elsewhere, the species is known from

China (Nguyen et al., 2009).

Remarks. The specimen was found at night

(2 1 :00) in a small water pool in a stream near Vung

Tron Ranger Station. The specimen matched the de-

scriptions of Pope (1935), Bourret (1936) and

David et al. (2011) but showed some differences:

more ventral scales (2+184 instead of maximum

173), prefrontal entire instead of semi-divided, a

longer total length (471.67 mm instead of maxi-

mum 437 mm), and a lower ratio of tail to total

length (0.137 instead of 0.25).

544

Vera L. Hecht et alii

Figure 69. Rhadinophis prasinus. Figure 70. Rhynchophis boulengeri. Figure 71. Sibynophis chinensis. Figure 72.

Psammodynastes pulverulentus. Figure 73. Amphiesmoides ornaticeps. Figure 74. Opisthotropis lateralis. Photos by T.

Ziegler and T.Q. Nguyen.

First report on the herpetofauna of TayYen Tu Nature Reserve, northeastern Vietnam

545

Rhabdophis subminiatus (Schlegel, 1837)

Identification was based on photographs (Fig.

75): head distinct from neck; eye large, black;

upper head bluish; neck with a black blotch, fol-

lowed by a U-shaped yellow band; a black verti-

cal stripe below the eye, directing backwards;

anterior part of body reddish brown, posterior part

yellowish brown, with numerous black or dark

grey scales forming a regular pattern; tail brown;

ventral surface cream (determination after Zie-

gler, 2002).

Distribution. This is a common species in Viet-

nam. Elsewhere, the species is known from India

throughout China and Indochina southwards to In-

donesia (Nguyen et al., 2009).

Remarks. Several individuals were seen during

our survey in summer 2010. One was crawling on

the ground in the bamboo forest and some were div-

ing in ponds at the forest edge near Mau Village

during day time.

Sinonatrix aequifasciata (Barbour, 1908)

Examined material. One adult female VNMN

1332, collected by TZ and NTT, 24 June 2010, ca.

400 m asl.

Morphological characters. SVL 360.48 mm,

TaL 109.68 mm; head distinct from neck; rostral

broader than high; intemasals 2, in contact, twice as

long as wide; prefrontals 2; frontal longer than wide,

shorter than parietals; nostrils lateral; pupil round; lo-

real single, as long as high; preocular single; postoc-

ulars 2; subocular single; temporals 2+3; supralabials

9, fifth touching the eye; infralabials 9; mental

groove present; dorsal scales keeled, in 19:19:17

rows; ventrals 5+141, no lateral edges; cloacal shield

divided; subcaudals 75, divided. Colouration in life:

head brownish above; back olive grey, with 1 8 black

double-bands on body, 10 on tail; flanks with dark

markings, in X-shape; ventral surface yellow, with

some traces of the black bands (determination after

Bourret, 1936; Vogel et al., 2004).

Distribution. In Vietnam, this species has been

recorded from Lao Cai, Ha Giang, Cao Bang, Lang

Son, Vinh Phuc, Bac Giang, Nghe An, and Ha Tinh

provinces. Elsewhere, the species is known from

China (Nguyen et al., 2009).

Remarks. The specimen was found in the

evening (22:30), on a tree branch in Suoi Tuyen I.

The specimen matched the descriptions of Bourret

(1936) and Vogel et al. (2004) well, but showed two

minor differences: only one subocular on the left

side instead of two and 18 bands on body and 10 on

tail instead of 20 and 12 bands, respectively.

Sinonatrix percarinata (Boulenger, 1899)

Examined material. One juvenile IEBR 3707

(SVL1 88.09 mm, TaL 62.05 mm), collected by

NQT, NTT, PTC, June 2009, elevation ca. 400 m

asl; one adult female VNMN 1357, collected by TZ

and NTT, 3 June 2010, ca. 400 m asl (Fig. 76).

Morphological characters. SVL female

423.28 mm, TaL female 154.31 mm; head distinct

from neck; rostral twice as wide as high; internasals

2, in contact, longer than wide; prefrontals 2; nos-

trils lateral; pupil round; nasal divided; loreal sin-

gle; preocular single; postoculars 2-3; suboculars

1-2; temporals 2+3; supralabials 9, fourth and fifth

touching the eye; infralabials 9-10; mental groove

present; dorsal scales keeled, except for outermost

rows, in 17/19:19:17 rows; ventrals 2-3 + 136/140;

cloacal shield divided; subcaudals 72 or 75, di-

vided. Colouration in life: dorsal head and back

olive greyish; brown body with 28-3 1 transverse

dark lozenge-shaped bands on back, dark bands

continuing to belly, forming rings; tail with 13-20

black bands, similar to those on body; ventral sur-

face yellow (determination after Pope, 1935; Bour-

ret, 1936; Smith, 1943; Ziegler, 2002).

Distribution. In Vietnam, this species has been

recorded from Lao Cai and Ha Giang southwards

to Gia Lai and Dong Nai provinces. Elsewhere, the

species is known from India, China, Taiwan, Myan-

mar, Laos, and Thailand (Nguyen et al., 2009).

Remarks. A juvenile was found in the evening

and the female was found during the day time in Lai

Am Pond.

PAREATIDAE

P areas hamptoni (Boulenger, 1905)

Examined material. One adult male VNMN

1356, collected by TZ and NTT, 10 July 2010, ca.

400 m asl (Fig. 77).

546

Vera L. Hecht et alii

Morphological characters. SVL 335.38 mm,

TaL 113.05 mm; head distinct from neck; body

strongly compressed; snout short; rostral not visible

from above; internasals in contact, half of pre-

frontals; nostrils lateral; pupil vertically elliptic;

prefrontals touching the eye; frontal shorter than pari-

etals; loreal single, not touching the eye; preocular

single, in triangle shape; eyes diameter greater than

the distance from eye to tip of snout; postocular sin-

gle; suboculars 2/3; temporals 2+3; supralabials 7,

fourth and fifth touching the eye; infralabials 6/7;

mental very small, mental groove absent; chin

shields in 3 pairs; dorsals smooth, midbody scale

rows 15, vertebral scales slightly enlarged; ventrals

186; cloacal shield undivided; subcaudals 84, di-

vided. Colouration in life: dorsal surface reddish

brown, with black bands on the interstitial skin;

ventral surface yellow; upper head dark, with two

parallel longitudinal stripes on neck, bordered in

black (determination after Bourret, 1936; Smith,

1943; Cox, 1991).

Distribution. In Vietnam, this species has been

recorded from Lao Cai and Ha Giang southwards

to Lam Dong and Dong Nai provinces. This is the

first record of P. hamptoni from Tay Yen Tu NR and

from Bac Giang Province. Elsewhere, the species

is known from China, Myanmar, Laos, and Cam-

bodia (Nguyen et al., 2009).

Remarks. Another adult male was seen in the

evening (20:00) on a tree branch, 1.5 m above the

ground near Ba Bep Stream.

P areas margaritophorus (Jan, 1866)

Examined material. One adult female VNMN

1354, collected by TZ and NTT, 10 July 2010, ca.

400 m asl (Fig. 78).

Morphological characters. SVL 437.00 mm,

TaL 83.95 mm; head distinct from neck; body not

strongly compressed; eye moderate; intemasals in

contact, shorter than prefrontals; prefrontals in con-

tact, entering orbit; frontal as long as wide, longer

than its distance to tip of snout, shorter than pari-

etals; loreal single; preocular absent; postocular sin-

gle; supralabials 7, fourth and fifth entering orbit;

infralabials 7; temporalia 2+2; mental groove ab-

sent; mental very small; chin shields in 3 pairs; dor-

sal scales smooth, midbody scale rows 15;

paravertebral scales not enlarged; ventrals 150,

without lateral edges; cloacal shield undivided; sub-

caudals 39, divided. Colouration in life: dorsum

dark grey with transverse rows of spots, spot edged

in white anteriorly and in black posteriorly; ventral

surface white, with numerous brown spots (deter-

mination after Bourret, 1936; Smith, 1943; Man-

they & Grossmann, 1997).

Distribution. In Vietnam, this species has been

recorded from Vinh Phuc and Hai Duong provinces

southwards to Kien Giang Province. This is the first

record of P. margarithophorus from Tay Yen Tu NR

and from Bac Giang Province. Elsewhere, the species

is known from China, Myanmar, Laos, Thailand,

Cambodia, Malaysia (Nguyen et al., 2009).

Remarks. The female specimen was found at

night (22:00) on the forest path near Ba Bep Stream.

TYPHLOPIDAE

Ramphotyphlops braminus (Daudin, 1803)

Examined material. One subadult VNMN

1336 (SVL 119.92 mm, TaL 2.76 mm), collected by

TZ and NTT, June 2010, elevation ca. 300 m asl.

Morphological characters. Small sized,

worm-like snake; nostril sourrounded by 2 nasals;

preocular single, touching second and third supral-

abials; eye rudimental; mental groove present; ven-

trals as large as lateral scales; dorsal scales smooth,

cycloid, in 20 rows; tail cylindrical. Colouration in

alcohol: dorsal surface dark brown, ventral surface

grey; lower side of head and cloacal region greyish

white; throat with 3 white scales forming a line; tip

of tail whitish (determination after Pope, 1935;

Bourret, 1936; Smith, 1943; Manthey & Gross-

mann, 1997; Ziegler, 2002).

Distribution. This is a widespread species in

Vietnam. Elsewhere, the species is known from San-

sibar, Tansania, Mosambique, Somalia, Camerun,

Benin, Togo, Ivory Coast, Senegal, Gabun, Mada-

gascar, Comores, Nosy Be, Mauritius, Iran, Pak-

istan, India, Sri Lanka, Nepal, Bangladesh, Bhutan,

China, Myanmar, Laos, Thailand, Cambodia,

Malaysia, Singapore, Indonesia, New Guinea, Philip-

pines, Japan, Melanesia, Micronesia, Australia,

New Caledonia, Solomon Islands, Vanuatu, Saudi

Arabia, Guatemala, Mexico, USA, and Hawaii

(Nguyen et al., 2009).

First report on the herpetofauna of TayYen Tu Nature Reserve, northeastern Vietnam

547

Remarks. A single specimen was found on the

forest floor between leaf litter.

XENODERMATIDAE

Achalinus rufescens Boulenger, 1888

Examined material. One adult female VNMN

1334, collected by TZ and NTT, 24 June 2010, 320-

400 m asl (Fig. 79).

Morphological characters. SVL 424.37 mm,

TaL 95.12 mm; head slender; suture between inter-

nasals twice as long as suture between prefrontals;

frontal wider than long, half as long as parietals, 4

times wider than supraocular; parietals 2; loreal

single, entering orbit; preocular and postocular ab-

sent; temporals 2+2, anterior upper one touching

the eye; supralabials 6, fourth and fifth touching the

eye, first very small, sixth very long; infralabials 5;

mental wider than long; chin shields in 3 pairs;

mental groove present; dorsal scales strongly keeled,

tri-carinated, midbody scale rows 23; ventrals 63;

cloacal shield undivided; subcaudals 5+152, undi-

vided. Colouration in life: dorsal head and body

light brown, iridescent; vertebral zone darker grey-

ish brown; lips yellowish; ventral surface whitish-

yellow (determination after Bourret, 1936; Pope,

1935; Smith, 1943; Ziegler, 2002).

Distribution. In Vietnam, this species has been

recorded from Lao Cai, Cao Bang, Bac Kan, Vinh

Phuc, Hai Duong, and Ha Tinh provinces. This is

the first record of A. rufescens from Tay Yen Tu NR

and from Bac Giang Province. Elsewhere, the

species is known from China (Nguyen et al., 2009).

Remarks. The specimen was found at night (ca.

23:00) on a forest path near Suoi Tuyen I.

ELAPIDAE

Bungarus fas ciatus (Schneider, 1801)

Identification was based on photographs (Fig.

80): triangular body with a prominent vertebral

ridge along the back; tail tip blunt; body and tail al-

ternately banded in black and yellow; a large black

mark on nape, continued in a point on head to be-

tween the eyes, bordered on each side by yellow

(determination after Smith, 1943).

Distribution. This is a widespread species in

Vietnam and in Asia (Nguyen et al., 2009).

Remarks. One individual was seen crawling on

the ground of the bamboo forest close to a pond dur-

ing the night of 26 June 2010.

VIPERIDAE

Trimeresurus stejnegeri (Schmidt, 1925)

Identification was based on photographs (Fig.

81): head triangle-shaped; loreal pitches present;

body green, with longitudinal parallel bicoloured

(white and red) stripe on the outer edge of the ven-

trals (determination after Smith, 1943; Bain &

Nguyen, 2004a).

Distribution. This is a widespread species in

northern and central Vietnam, from Lao Cai and Ha

Giang provinces southwards to Quang Binh Prov-

ince and Da Nang City. Elsewhere, the species is

known from China, Taiwan, and Myanmar (Nguyen

et al., 2009).

Natural history notes: Three individuals were

sighted at night in the bushes and trees besides a for-

est stream and close to Khe Cam 1 Pond, in October

2009 and in May 2010.

DISCUSSION

Based on our field work during the period from

2008 to 2010, a total of 76 species were recorded

from the Tay Yen Tu NR, comprising 36 species of

amphibians and 40 species of reptiles. The families

with the most diverse species richness are Ranidae

(11 species), Rhacophoridae (9 species), Scincidae

(10 species), and Colubridae (8 species) (Fig. 82).

Because of the lack of convincing diagnostic char-

acters, the identification of several amphibian species

is still not clearly solved yet, for example Lepto-

brachium cf. chapaense, and Hylarana cf. nigrovit-

tata, but also Ichthyophis bannanicus and

Limnonectes bannaensis (see Gawor et al., 2009;

Nguyen et al., 2009; McLeod, 2010; Nishikawa et

al., 2012). Further studies based on additional voucher

specimens and molecular comparisons are required

to confirm the taxonomic placement of aforemen-

tioned species (or species complexes).

548

Vera L. Hecht et alii

Figure 75. Rhabdophis subminiatus. Figure 76. Sinonatrix percarinata. Figure 77. Pareas hamptoni. Figure 78. P. marga-

ritophorus. Figure 79. Achalinus rufescens. Figure 80. Bungarus fasciatus. Figure 81. Trimeresurus stejnegeri. Photos by

C.T. Pham, T. Ziegler, and T.Q. Nguyen.

First report on the herpetofauna of TayYen Tu Nature Reserve, northeastern Vietnam

549

Figure 82. Species richness of amphibian and reptile families from Tay Yen Tu Nature Reserve

(y-axis represents species numbers).

Thirty of the recorded species (or 39.5% of the

total species number) are new records for Tay Yen

Tu Nature Reserve as well as for Bac Giang Prov-

ince. Remarkably, five of the recorded species are

currently known only from Vietnam, namely Tylo-

totriton vietnamensis, Quasipaa acanthophora ,

Odorrana bacboensis, O. yentuensis, and Goni-

urosaurus lichtenfelderi. The high level of species

diversity and endemism of the herpetofauna under-

scores the biodiversity conservation potential of the

Tay Yen Tu NR. Although this nature reserve har-

bors a considerable number of endemic and rare

species, its biodiversity is heavily threatened by

habitat degradation and over harvesting associated

with wildlife trade (Nguyen, 2011). To preserve the

unique biodiversity of Yen Tu Mountain (see also

Ha et al., 2010), we must not only continue with

research, in particular the ecological requirements

of the local flora and fauna, but also the protection

of the habitat must be improved. Only by doing so,

we can guarantee the long-term survival of the

unique lowland forest system and its in part en-

demic diversity.

ACKNOWLEDGEMENTS

We thank the directorate of the Tay Yen Tu Nature

Reserve, Forest Protection Department of Bac Giang

and Quang Ninh provinces for support of our field

work and issuing relevant permits. V.L. Hecht and

C. T. Pham are grateful to M. Bemardes and D. Karbe

(Cologne), T.T. Tran (Vinh Phuc) for their assistance

in the field. For the loan of specimens, we are grate-

ful to C.X. Le and T.H. Tran (Hanoi), W. Bohme and

D. Rodder (Bonn). We thank E. Sterling (New York)

and K. Koy (Berkeley) for providing the map. Field

surveys in Yen Tu Mountains were partially funded

by the European Association of Zoos and Aquaria

550

Vera L. Hecht et alii

(EAZA), World Association of Zoos and Aquariums

(WAZA), Nagao Natural Environment Foundation,

Japan (Small-scale research grant programme for

T.T. Nguyen), Vietnam Academy of Science and

Technology (Small grant for young researchers for

C. T. Pham), and Cologne Zoo. Internship of C.T.

Pham in Germany was funded by the Deutsche

Foschungsgemeinschaft (DFG) - International Col-

laboration Grant (BO 1907/ 11-1). Research of T.Q.

Nguyen in Germany is funded by the Alexander von

Humboldt Foundation (VIE 1143441).

REFERENCES

Anders C. & Rai K. R., 2002. Rhacophorus maximus

Gunther, 1859 “1858”. In: Schleich H. H. & Kastle

W. (Eds). Amphibians and reptiles of Nepal. Bi-

ology, Systematics, Field Guide, A. R. G. Ganter

Verlag K. G., pp. 337-340.

Averyanov L.V., Phan L.K., Nguyen H.T. & Harder D.K.,

2003. Phytogeographic review of Vietnam and adja-

cent areas of eastern Indochina. Komarovia, 3: 1—83.

Bain R.H. & Nguyen T.Q., 2004a. Herpetofaunal diver-

sity of Ha Giang Province in northeastern Vietnam,

with description of two new species. American Mu-

seum Novitates, 3453: 1—42.

Bain R.H. & Nguyen T.Q., 2004b. Three new species of

narrow-mouth frog (Genus: Micro hy l a) from In-

dochina, with comments on Microhyla annamensis

and Microhyla palmipes. Copeia, 3: 507-524.

Bain R.H., Lathrop A., Murphy R.W., Orlov N. & Ho

C.T., 2003. Cryptic species of a cascade frog from

Southeast Asia: Taxonomic revisions and descrip-

tions of six new species. American Museum Novi-

tates, 3417: 1-60.

Bain R.H., Nguyen T.Q. & Doan K.V., 2009. A new

species of the genus Theloderma Tschudi, 1838

(Anura: Rhacophoridae) from northwestern Vietnam.

Zootaxa, 219: 58—68.

Bernardes M., Rodder D., Nguyen T.T., Pham T.C.,

Nguyen T.Q. & Ziegler T., 2013. Habitat characteri-

zation and potential distribution of Tylototriton viet-

namensis in northern Vietnam. Journal of Natural

History, 47: 1161-1175.

Bohme W., Schottler T., Nguyen T.Q. & Kohler J., 2005.

A new species of salamander, genus Tylototriton

(Urodela: Salamandridae), from northern Vietnam.

Salamandra, 41: 215—220.

Boulenger G.A., 1912. A vertebrate fauna of the Malay

Peninsula from the Isthmus of Kra to Singapore in-

cluding the adjacent islands - Reptilia and Batrachia.

Taylor and Francis, London, 294 pp.

Bourret R., 1936. Les Serpentes de Flndochine. I. Etudes

sur la Faune. Henri Basuyau & Cie, Vols. 1 & 2,

Toulouse, 141+ 505 pp.

Bourret R., 1942. Les Batraciens de Flndochine. Institute

Oceanographique de Flndochine, Hanoi, x + 547 pp.,

4 pis.

Bourret R., 2009. Les Lizards de Flndochine. Edition

Chimaira, Frankfurt am Main, 624 pp.

Cox M.J., 1991. The snakes of Thailand and their hus-

bandry. Krieger Publishing Company, Malabar, Flor-

ida, 526 pp.

Darevsky I.S., Orlov N.L. & Ho C.T., 2004. Two new ly-

gosomine skinks of the genus Sphenomorphus

Fitzinger, 1843 (Sauria, Scincidae) from northern

Vietnam. Russian Journal of Herpetology, 11: 1 1 1 —

120 .

David P., Pham C.T., Nguyen T.Q. & Ziegler T., 2011. A

new species of the genus Opisthotrophis Gunther,

1872 (Squamata: Natricidae) from the highlands of

Kon Turn Province, Vietnam. Zootaxa, 2758: 43-56.

David P., Vogel G. & Pauwels O.S.G., 2008. A new

species of the genus Oligodon Fitzinger, 1826

(Squamata: Colubridae) from Southern Vietnam and

Cambodia. Zootaxa, 1939: 19—37.

Dowling H.G., 1951. A proposed standard system of

counting ventrals in snakes. British Journal of Her-

petology, 1: 97—99.

Dubois A. & Ohler A., 2009. A new species of the genus

Quasipaa (Anura, Ranidae, Dicroglossinae) from

northern Vietnam. Alytes, 27: 49—61.

Gawor A., Hendrix R., Vences M., Bohme W. & Ziegler

T., 2009. Larval morphology in four species of

Hylarana from Vietnam and Thailand with comments

on the taxonomy of H. nigrovittata sensu latu (Anura:

Ranidae). Zootaxa, 2051: 1—25.

Goodall D. & Faithfull S., 2010. U Minh Thuong National

Park - Kien Giang Province, Vietnam Amphibian and

Reptile Survey 7-21 September 2009. Wildlife At

Risk, Vietnam.

Ha Q.M., La C.M., Tran S.N. & Do Q.V., 2010. Tay Yen

Tu Nature Reserve: Biodiversity conservation value

and development potential. Publishing House for

Science and Technology, Hanoi, 35 pp.

Hikida T., Lau M.W.-N. & Ota H., 2001. A new record

of the Vietnamese Five-Lined Skink, Eumeces

tamdaoensis (Reptilia: Scincidae), from Hong Kong,

China, with special references to its sexual dimor-

phism. The Natural History Journal of Chulalongkom

University, 1: 9—13.

Inger R.F., Orlov N. & Darevsky I., 1999. Frogs of Viet-

nam: A report on new collection. Fieldiana Zoology,

92: 1 — 46

Kuraishi N., Matsui M., Hamidy A., Belabut D.M.,

Ahmad N., Panha S., Sudin A., Yong H.-S., Jiang J.-

P, Ota H., Ho T.T. & Nishkawa K., 2012. Phyloge-

First report on the herpetofauna of TayYen Tu Nature Reserve, northeastern Vietnam

551

netic and taxonomic relationships of the Polypedates

leucomystax complex (Amphibia). Zoologica Scripta,

42: 54-70.

Le Q.K. & Ziegler T., 2003. Fist record of Chinese Croc-

odile Lizard outside of China: Report on a population

of Shinisaurus crocodilurus Ahl, 1930 from North-

eastern Vietnam. Hamadryad, 27: 193—199.

Liu C.C. & Hu S.Q., 1960. Preliminary report of Am-

phibia from southern Yunnan. Acta Zoologica Sinica,

11: 509-533 [in Chinese with English abstract],

Manthey U. & Grossmann W., 1997. Amphibien und

Reptilien Sudostasiens. Natur & Tier Verlag, Mun-

ster, 512 pp.

Mcleod D.S., 2010. Of least concern? Systematics of a

cryptic species complex: Limnonectes kuhlii (Am-

phibia: Anura: Dicroglossidae). Molecuar Phyloge-

netics and Evolution, 56: 991—1000.

Neang T. & Holden J., 2008. A field guide to the amphib-

ians of Cambodia. Fauna and Flora Internationnal.

Phompenh, 127 pp.

Nguyen S.V., Ho C.T. & Nguyen T.Q., 2009. Herpeto-

fauna of Vietnam. Edition Chimaira, Frankfurt am

Main, 768 pp.

Nguyen T.Q., 2011. Systematics, ecology, and conserva-

tion of the lizard fauna in northeastern Vietnam, with

special focus on Pseudocalotes (Agamidae), Goni-

urosaurus (Eublepharidae), Sphenomorphus and

Tropidophorus (Scincidae) from this country. Disser-

tation, University of Bonn, April, 2011, 229 pp.

Nguyen T.Q., David P., Tran T.T., Luu V.Q., Le Q.K. &

Ziegler T., 2010a. Amphiesmoides ornaticeps

(Werner, 1924), an addition to the snake fauna of

Vietnam, with a redescription and comments on the

genus Amphiesmoides Malnate, 1961 (Squamata: Na-

tricidae). Revue suisse de Zoologie, 117: 45—56.

Nguyen T.Q., Koch A. & Ziegler T., 2009. A new species

of reed snake Calamaria Boi, 1827 (Squamata:

Colubridae), from central Vietnam. Hamadryad, 34:

1 - 8 .

Nguyen T.Q., Nguyen S.V., Orlov N., Hoang T.N.,

Bohme W. & Ziegler T., 2010b. A review of the genus

Tropidophorus (Squamata, Scincidae) from Vietnam

with new species records and additional data on

natural history. Zoosystematics and Evolution, 86:

5-19.

Nguyen T.Q., Schmitz A., Nguyen T.T., Orlov N.L.,

Bohme W. & Ziegler T., 2011a. Review of the genus

Sphenomorphus Fitzinger, 1843 (Squamata: Sauria:

Scincidae) in Vietnam, with description of a new

species from northern Vietnam and southern China

and the first record of Sphenomorphus mimicus

Taylor, 1962 from Vietnam. Journal of Herpetology,

45: 145-154.

Nguyen T.Q., Stenke R., Nguyen H.X. & Ziegler T.,

2011b. The terrestrial reptile fauna of the Biosphere

Reserve Cat Ba Archipelago, Hai Phong, Vietnam.

In: Schuchmann K.-L. (Ed.), Tropical Vertebrates in

a Changing World. Bonner zoologische Monogra-

phien, 57, pp. 99—115.

Nguyen T.Q., Tran T.T., Hoang N.V., Bohme W. &

Ziegler T., 2008. Rediscovery and redescription of

Ateuchosaurus chinensis Grey, 1845 (Squamata:

Sauria: Scincidae) from northeastern Vietnam. Her-

petology Notes, 1: 17—21.

Nguyen T.Q., Tran T.T., Nguyen T.T., Ziegler T. &

Bohme W., 2012. First record of Sphenomorphus

incognitus (Thompson, 1912) (Squamata: Scincidae)

from Vietnam with some notes on natural history.

Asian Herpetological Research, 3: 147—150.

Nguyen T.Q., Wang Y-Y., Yang Y-H., Lehmann T., Le

M.D., Ziegler T. & Bonkowski M., 2013. A new

species of the Gekko japonicus group (Squamata:

Sauria: Gekkonidae) from the border region between

China and Vietnam. Zootaxa, 3652: 501—518.

Nguyen T.T., Tran T.T., Nguyen T.Q. & Pham C.T., 2008.

Distribution: Rhacophorus maximus (Nepal Flying

Frog). Herpetological Review, 39: 364.

Nishikawa K., Matsui M. & Nguyen T.T. 2013. A new

species of Tylototriton from northern Vietnam (Am-

phibia: Urodela: Salamandridae). Current Herpetol-

ogy, 32: 34—49.

Nishikawa K., Matsui M. & Orlov N.L., 2012. A new

striped Ichthyophis (Amphibia: Gymnophiona:

Ichthyophiidae) from Kon Turn Plateau, Vietnam.

Current Herpetology, 31: 27—38.

Ohler A., 2003. Revision of the genus Ophryophryne

Boulenger, 1903 (Megophryidae) with description of

two new species. Alytes, 21: 23—44.

Ohler A., Wollenberg K.C., Grosjean S., Hendrix R.,

Vences M., Ziegler T. & Dubois A., 2011. Sorting out

Lalos: description of new species and additional

taxonomic data on megophryid frogs from northern

Indochina (genus Leptolalax, Megophryidae, Anura).

Zootaxa, 3147: 1—83.

Orlov N., Dutta S., Ghate H. & Kent Y., 2006. New spe-

cies of Theloderma from Kon Turn province (Viet-

nam) and Nagaland state (India) (Anura:

Rhacophoridae). Russian Journal of Herpetology, 13:

135-154.

Pope C.H., 1935. The Reptiles of China - Turtles, Croc-

odiles, Snakes, Lizards. Natural History of central

Asia. Vol X. American Museum of Natural History,

New York, 604 pp.

Smith M.A., 1935. The fauna of British India including

Ceylon and Burma. Reptilia and Amphibia. Vol. II.

Sauria. Taylor and Francis, London, 440 pp.

Smith M.A., 1943. The fauna of British India, Ceylon

and Burma. Reptilia and Amphibia. Vol. III. Ser-

pentes. Taylor and Francis, London, 525 pp.

552

Vera L. Hecht et alii

Taylor E.H., 1962. The amphibian fauna of Thailand.

University of Kansas Science Bulletin, 63: 265-599.

Tordoff A.W., Tran B.Q., Nguyen T.D. & Le H.M. (Eds),

2004. Sourcebook of existing and proposed protected

areas in Vietnam. BirdLife International in Indochina

and Ministry of Agriculture and Rural Development,

second edition, Hanoi.

Tran D.A.T., Le Q.K., Le K.V., Thanh N.V., Nguyen T.Q.,

Bdhme W. & Ziegler T., 2010. First and preliminary

frog records (Amphibia: Anura) from Quang Ngai

Province, Vietnam. Herpetology Notes, 3: 111—119.

Tran T.T., Orlov N.L. & Nguyen T.T., 2008. A new

species of cascade frog of Odorrana Fei, Yi et Huang,

1990 genus (Amphibia: Anura: Ranidae) from Bac

Giang Province (Yen Tu Mountain Range, North-

eastern Vietnam). Russian Journal of Herpetology,

15: 212-224.

Vogel G., David P, Pauwels O.S.G. & Brachtel N., 2004.

On the occurance of the Watersnake Sinonatrix ae-

quifasciata (BARBOUR, 1908) (Serpentes, Colubri-

dae, Natricinae) in Vietnam. Hamadryad, 29:

110-114.

Vogel G., David P, Pauwels O.S.G., Sumontha M., Nor-

val G., Hendrix R., Vu T.N. & Ziegler T., 2009. A re-

vision of Ly codon ruhstrati (Fischer 1886) auctorum

(Squamata Colubridae), with the description of a new

species from Thailand and a new subspecies from the

Asian mainland. Tropical Zoology, 22: 131—182.

Ye C.Y., Fei L., Xie F. & Jiang J.P, 2007. Anew Ranaidae

species from China - Limnonectes bannanensis (Ra-

nidae: Anura). Zoological Research, 28: 545—550.

Yu G., Zhang M. & Yang J., 2010. A species boundary

within the Chinese Kurixalus odontotarsus species

group (Anura: Rhacophoridae): new insights from

molecular evidence. Molecular Phylogenetics and

Evolution, 56: 942—950.

Zaher H., Grazziotin F.G., Cadle J.E., Murphy R.W.,

Moura-Leite J.C. & Bonatto S.L., 2009. Molecular phy-

logeny of advanced snakes (Serpentes, Caenophidia)

with an emphasis on South America xenodontines: a

revised classification and descriptions of new taxa.

Papeis Avulsos de Zoologia, 49: 115—153.

Ziegler T., 2002. Die Amphibien und Reptilien eines

Tieflandfeuchtwald-Schutzgebiets in Vietnam. Natur

& Tier Verlag, Munster, 342 pp.

Ziegler T. & Bischoff W., 1999. Takydromus ( Platyplaco -

pus) kuelmei vietnamensis ssp. n., eine neue Schnel-

llaufereidechsen-Unterart aus Vietnam (Reptilia:

Squamata: Lacertidae). Salamandra, 35: 209—226.

Ziegler T., Hendrix R., Vu T.N., Vogt M., Forster B. &

Dang K.N., 2007. The diversity of a snake commu-

nity in a karst forest ecosystem in the central Truong

Son, Vietnam, with an identification key. Zootaxa,

1493: 1-40.

Ziegler T., Le Q.K., Vu T.N., Hendrix R. & Bohme W.,

2008a. A comparative study of crocodile lizards

( Shinisaurus crocodilurus Ahl, 1930) from Vietnam

and China. Raffles Bulletin of Zoology, 56: 181—187.

Ziegler T., Nguyen S. V. & Nguyen T. Q., 2008b. Anew

reed snake of genus Calamaria Boie (Squamata:

Colubridae) from Vietnam. Current Herpetology,

27: 71-80.

Ziegler T., Nguyen T. Q., Schmitz A., Stenke R. & Rosier

H., 2008c. Anew species of Goniurosaurus from Cat

Ba Island, Hai Phong, northern Vietnam (Squamata:

Eubleppharidae). Zootaxa, 1771: 16—30.

Ziegler, T., Ohler, A., Vu Ngoc Thanh, Le Khac

Quyet, Nguyen Xuan Thuan, Dinh Huy Tri & Bui

Ngoc Thanh (2006): Review of the amphibian and

reptile diversity of Phong Nha - Ke Bang National

Park and adjacent areas, central Truong Son, Viet-

nam. In: Vences, M., Kohler, J., Ziegler, T. & W.

Bohme (Hrsg.): Herpetologia Bonnensis II: 247-

262. Proceedings of the 13th Ordinary General

Meeting of the Societas Europaea Herpetologica,

Bonn: 262 pp.

Biodiversity Journal, 2013, 4 (4): 553-556

First record of Hierophis gemonensis (Laurenti, 1 768) (Reptilia

Serpentes Colubridae) in the Aegean island of Tsougria,

Northern Sporades, Greece

Mauro Grano 1 *, Cristina Cattaneo 2 & Augusto Cattaneo 3

1 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 The presence of Hierophis gemonensis (Laurenti, 1768) (Reptilia Serpentes Colubridae) in

Tsougria, a small island of the Northern Sporades, Greece, is here recorded for the first time.

KEY WORDS Aegean islands; Balkan whip snake; Hierophis gemonensis ; Northern Sporades; Tsougria.

Received 05.11.2013; accepted 02.12.2013; printed 30.12.2013

INTRODUCTION

The Balkan whip snake, Hierophis gemonensis

(Laurenti, 1768) (Reptilia Serpentes Colubridae), is

widespread along the coastal areas of Slovenia,

Croatia, Bosnia-Erzegovina, Montenegro, Albania

and Greece (Vanni et al., 2011). The basic colour is

silver gray to dark green with some spots only on

one third of the body, tending to regular stripes on

the tail. Melanistic specimens are also known

(Dimitropoulos, 1986; Schimmenti & Fabris,

2000). The total length is usually less than 130 cm,

with males larger than females (Vanni et al., 2011).

Regarding the Aegean islands, this species has

an irregular distribution, therefore its presence in this

area can be defined discontinuous (Clark, 1989). H.

gemonensis has been found on these islands:

Northern Sporades

Asproniso (= Aspro): Buchholz & Schultze-

Westrum, 1964; Kock, 1979.

Euboea: Werner, 1933; Kock, 1979.

Argo- Saronic islands

Aegina: Werner, 1937; Wettstein, 1953; Kock,

1979; Clark, 1989.

Psili: Clark, 1973, 1989; Kock, 1979.

Tolon: Clark, 1973, 1989; Kock, 1979.

Stavronissos, Dhokos, Trikkeri (archipelago of

Hydra): Clark, 1989.

Kythera: Boulenger, 1893; Kock, 1979.

Crete: Boettger, 1888; Sowig, 1985.

Cretan islets

Gramvoussa: Wettstein, 1953; Kock, 1979.

Gavdos: Wettstein, 1953; Kock, 1979.

Gianyssada: Wettstein, 1953; Kock, 1979.

Dia: Raulin, 1869; Kock, 1979.

Theodori: Wettstein, 1953.

Karpathos: Broggi, 1994.

The population of the island of Gyaros (Cy-

clades), previously assigned to H. ( Coluber ) gemo-

nensis (Mertens, 1968), and successively

considered an endemic species, H. gyarosensis

(Mertens, 1968), by Schatti (1988) and Bohme

(1993), is actually referred to H. viridiflavus car-

bonarius (Bonaparte, 1833) (Utiger & Schatti,

2004), on the bases of molecular and, in part, mor-

phological data (the structure of the hemipenis is

different). The western whip snake on Gyaros was

introduced casually on the island in historical times.

554

M. Grano, C. Cattaneo & A. Cattaneo

In fact, the values of the ventral lepidosis (194-205),

colour pattern (dark and without spots ventrally) and

the geographical location of the island surrounded

by other islands inhabited by DoJichophis caspius

(Gmelin, 1789), as well, would led to assign the

ophidic population of Gyaros to D. caspius. More-

over, despite as stated by Schatti (1988), the melan-

oticism is present in the micro insular Aegean

populations of D. caspius (see Alonissos and, per-

haps, Patmos) (Cattaneo, 1998, 2008; Broggi, 2010).

Study area

Tsougria is a small island off the SE coast of

Skiathos (longitude: 23°29 , 58.63”E; latitude

39°07’20.17”N). (Fig. 1) The area of 1.14 Ion 2 and

the altitude 90 m a.s.l. This small island, just like

Skiathos and other nearby islets (except for

Asproniso, that consists of limestone), are com-

posed of igneous and metamorphic rocks (Ferenti-

nos, 1972). Two small pounds occur on the island

behind the dunes of brackish water surrounded by

a belt of Juncus acutus L.; during our visit to the is-

land only one still contained water. Tsougria is cur-

rently uninhabited, but there was human presence

in the past. This is testified by the presence of small

valleys in the western part of the island exploited

for the cultivation of olive trees. These old olive

trees are now mixed with a shrub vegetation con-

sisting of Erica manipuliflora Salisb., Cistus creti-

cus L. and Dittrichia viscosa (L.) Greuter. The rest

of the island is covered by woodlands of Pinus

halepensis Mill, (with undergrowth of Erica ar-

borea L. and Pistacia lentiscus L.) alternated with

Figure 1. Study area.

stretches of dense and intricate maquis where pre-

dominant elements are Pistacia lentiscus , Quercus

coccifera L., Phillyrea latifolia L., Olea europaea

subsp. oleaster Hoffmg. et Link. Remains of two

rural buildings and information obtained from the

inhabitants of Skiathos, indicate that the anthropic

use of the island occurred until the 1970s of the last

century. As sign of the activities carried out in the

island, remain two groups of buildings actually

crumbling. The first was used for processing oil, the

other one was a farm with lodge on the upper floor

and stalls on the lower one. The area in front of the

farm provided a well for fresh water and some tanks

for watering the animals.

RESULTS AND CONCLUSIONS

Six records of H. gemonensis were found in the

island of Tsougria in August 2013: four pieces of

exuvia, one carcass and a live specimen. Two of the

four pieces of exuvia were rather dark. The live

specimen was a young adult of 60 cm total length

(tail 19 cm). It was found in front of the farm, under

an iron sheets set in the shade, near a dry stone wall

(Fig. 2) The soil covered by the sheet was wet de-

spite the diy spell. Colour and pattern were typical

of the species (Figs. 3, 4)

At Tsougria were also found Hemidactylus turci-

cus Linnaeus, 1758 and Lacerta trilineata Bedriaga,

1878. H. turcicus has been frequently observed under

wooden tables and iron sheet near the two groups of

buildings and around the little Church of Aghios

Floros. L. trilineata instead, has been observed

within the rushes. In this island, as indeed in Skiathos,

the Northern Sporades lizard, Podarcis erhardii

ruthveni (Wemer, 1930), is absent (Wettstein, 1953;

Gruber & Schultze-Westmm, 1971; Cattaneo, 1997).

Only Bergman (1995) observed specimen of P.

erhardii in Skiathos, nearby the main town where it

was probably introduced by boats that connect

Skiathos to nearby islands. But this record has not

been confirmed. On the other hand is documented its

occurrence in the other surrounding islets, such as

Repi, Asproniso, Tsougriaki, Arkos, Maragos. In

both Skiathos and Tsougria P erhardii is lacking, but

L. trilineata occurs (Gmber & Schultze-Westrum,

1971). This has raised many questions among her-

petologists leading to more or less plausible hy-

potheses. It was assumed that P erhardii forming part

First record of Hierophis gemonensis (Laurenti, 1 768) in the Aegean island of Tsougria, Northern Sporades, Greece 555

Figure 2. Tsougria island: the place of discovery of the Hierophis gemonensis.

Figures 3, 4. Hierophis gemonensis from Tsougria island.

in the pabulum of L. trilineata (Gruber & Schultze-

Westrum, 1971), would have suffered a selective

pressure by the latter species with the result of an

extinction on both above mentioned islands and

possibility of existence only in the surrounding islets,

where L. trilineata is absent. However it seems not

confirmed by the case of Arkonissi, a small island

which is also part of Skiathos archipelago, where the

two species are sympatric (however in the absence

of ophidic saurophagous species).

The same could regard H. gemonensis. The

plausible competitive interaction with Malpolon

monspessulanus Hermann, 1804 also present at

Skiathos (Buchholz & Schultze-Westrum, 1964;

Cattaneo, 1997) could be a relevant factor in the

distribution of H. gemonensis in the Northern Spo-

rades (as well as in the islands of the Argo-Saronic

archipelago: Clark, 1989). In fact, the exclusive

presence of H. gemonensis in Asproniso and Tsougria,

could be a consequence of the selective pressure

operated by M. monspessulanus , that would allow

the Balkan whip snake to survive only in these two

small islands surrounding Skiathos. Considering

the ophiophagy and the aggressive behavior of this

large snake, it could have played an important role

to obstacle the coexistence between the two species.

Also in the Argo-Saronic islands usually these two

snakes are not sympatric (Clark, 1989). The first

report of H. gemonensis for the Northern Sporades

is related to one specimen found in the island of

Asproniso, very close to Skiathos (Buchholz &

Schultze-Westrum, 1964). In this paper is given the

first record for the island of Tsougria and the sec-

ond for the Northern Sporades archipelago and

556

M. Grano, C. Cattaneo & A. Cattaneo

confirmed the occurrence on some of the small is-

lands around Skiathos of this species, whose relict

presence on the main island could possibly be

detected in the future (in 1 994 the third author fre-

quently sighted in Skiathos snakes perhaps attribu-

table to this species).

REFERENCES

Bergman J., 1995. Neues zur Herpetofauna der Insel

Skiathos, Nordliche Sporaden, Griechenland.

Herpetofauna, 17: 26-28.

Boettger O., 1888. Verzeichnis der von Hem E. von Oer-

tzen aus Griechenland und aus Kleinasien mitge-

brachten BatrachierundReptilien. Sitzungsberichte der

koniglich preussischen Akademie der Wissenschaften

zu Berlin, 5: 139-186.

Bohme W., 1993. Coluber gyarosensis Mertens, 1968 -

Gyaros-Pfeilnatter. - In: Bohme W. (Ed.), Handbuch

der Reptilien und Amphibien Europas, 3/1, Schlangen

(Serpentes). Aula, Wiesbaden, 111-114.

Boulenger G.A., 1893. Catalogue of the snakes in the

British Museum (Natural History). Containig the

families Typhlopidae, Glauconiidae, Boidae, Ilysiidae,

Uropeltidae, Xenopeltidae, and Colubridae Aglyphae,

part. Trustees, London, 440 pp.

Broggi M.F., 1994. Feldherpetologischen Beobachtungen

und Bemerkungen zu schiitzensweten Biotopen auf

griechischen Insel (Amphibia; Reptilia; Griechen-

land). Herpetozoa, 7: 29-34.

Broggi M.F., 2010. The herpetofauna of Alonissos

(Northern Sporades, Greece) (Amphibia, Reptilia).

Herpetozoa, 23: 71-78.

Buchholz K.F. & Schultze-Westrum T., 1964. Zur

Kenntnis der Schlangenfauna der Nordlichen Spo-

raden. Zoologischer Anzeiger, 173: 127-136.

Cattaneo A., 1997. L’erpetofauna delTisola greca di

Skiathos (Sporadi settentrionali). Atti della Societa

Italiana di Scienze Naturali e del Museo Civico di

Storia Naturale di Milano, 136: 145-156.

Cattaneo A., 1998. Gli Anfibi e i Rettili delle isole greche

di Skyros, Skopelos e Alonissos (Sporadi setten-

trionali). Atti della Societa Italiana di Scienze Natu-

rali e del Museo Civico di Storia Naturale di Milano,

139: 127-149.

Cattaneo A., 2008. Osservazioni sull'ofidiofauna delle

isole egee di Leros e Patmos (Dodecaneso) (Reptilia

Serpentes). II Naturalista siciliano, 32: 201-219.

Clark R.J., 1973. New locality records for Greek reptiles.

British Journal of Herpetology, 4:31 1-312.

Clark R.J., 1989. A checklist of the herpetofauna of the

Argo-Saronic gulf District, Greece. British Herpeto-

logical Society Bulletin, 28: 8-24.

Dimitropoulos A., 1986. Some notes on the colour and

pattern variation of the Greek snake fauna in relation

to geographic distribution. Biologia Gallo-hellenica,

12: 463—47 1 .

Ferentinos G.C., 1972. The geology-petrology of the is-

land of Skiathos. PhD. thesis, University of Patras.

Gmber U. & Schultze-Westmm T., 1971. Zur Taxonomie

und Okologie der Cycladen Eidechse ( Lacerta

erhardii ) von den Nordlichen Sporaden. Bonner

Zoologische Beitrage, 22: 101-130.

Kock D., 1979. Zwei Shlangen neu fiir Kephallinia,

Ionische Inseln, Griechenland (Reptilia: Serpentes,

Colubridae). Senckenbergiana biologica, 60: 7-11.

Mertens R., 1968. Eine schwarze Zornnatter von den

Cycladen: Coluber gemonensis gyarosensis n. subsp.

Senckenbergiana biologica, 49: 181-189.

Raulin V., 1869. Description physiologique de Tile de

Crete. Zoologie. Actes de la Societe Linneenne de

Bordeaux, 24: 691-692.

Schatti B., 1988. Systematik und Evolution der

Schlangengattung Hierophis Fitzinger, 1843. Zurich,

Inaugural-Dissertation, 50 pp.

Schimmenti G. & Fabris V., 2000. Note sull’erpetofauna

dell’isola di Krk (Croazia nordoccidentale). Atti del

I Congresso Nazionale della Societas Herpetologica

Italica. Museo Regionale Scienze naturali di Torino:

643-652.

Sowig P., 1985. Beitrage zur Kenntnis der Verbreitung

und Okologie der Amphibien und Reptilien Kretas.

Salamandra, 21: 252-262.

Utiger U. & Schatti B., 2004. Morphology and phyloge-

netic relationships of the Cyprus racer, Hierophis

cypriensis, and the systematic status of Coluber gemo-

nensis gyarosensis Mertens (Reptilia: Squamata:

Colubridae). Revue Suisse de Zoologie, 111: 225-238.

Vanni S., Nistri A., Lanza B. & Bressi N., 2011. Hierophis

gemonensis (Laurenti, 1768). In: Corti C., CapulaM.,

Luiselli L., Razzetti E., Sindaco R. (Eds), Fauna

d’ltalia: Reptilia, Vol. XLV. Edizioni Calderini de II

Sole 24 ORE, Editoria Specializzata S.r.l., Bologna:

505-509.

Werner F., 1933. Ergebnisse einer zoologischen Studien

und Sammelreise nach den Inseln des Agaischen

Meeres. I. Reptilien und Amphibien. Sitzungs-

berichte/Akademie der Wissenschaften in Wien,

Mathematisch-Naturwissenschaftliche Klasse

Abteilung, 142: 103-133.

Werner F. 1937. Beitrage zur Kenntnis der Tierwelt des

Peloponnes, der Inseln Kythira und Euboa sowie der

kleinen Inseln im Saronischen Golf. Sitzungs-

berichte/Akademie der Wissenschaften in Wien, Mathe-

matisch-Naturwissenschaftliche Klasse Abteilung,

146: 135-153.

Wettstein O. 1953. Herpetologia aegaea. Sitzungs-

berichte/Osterreichische Akademie der Wissenschaften,

Mathematisch-Naturwissenschaftliche Klasse

Abteilung, 162: 651-833.

Biodiversity Journal, 2013, 4 (4): 557-564

Trend of a population of Wild Rabbit Oryctolagus cuniculus

(Linnaeus, 1 758) in relation to Domestic Sheep Ovis aries dries

(Linnaeus, 1 758) grazing within a small insular protected area

Agatino Maurizio Siracusa & Ettore Petralia

Department of Biological, Geological and Environmental Sciences - Section of Animal Biology, University of Catania, via Androne

81 - 95124 Catania, Italy

^Corresponding author: amsira@unict.it

ABSTRACT The wild rabbit Oryctolagus cuniculus (Linnaeus, 1758) (Mammalia Leporidae) if present

at high density can cause significant damages to crops and natural vegetation. The aim of

this study, carried out from 2002 to 2008 in the Natural Reserve “Complesso Immacolatelle

e Micio Conti”, in the foothills of Mount Etna, was to investigate the relationship between

the wild rabbit and the presence of grazing domestic sheep Ovis aries aries (Linnaeus, 1758)

(Mammalia Bovidae), considering also the possible synergistic effect of these two herbivores

on the natural vegetation in a small protected area. Precipitation during the month of Sep-

tember correlated statistically significantly with the density of rabbits. A comparison between

the density values obtained during the period under study did not show statistically signifi-

cant differences with the exception of 2006 and 2007 when, within the area, was present a

flock of sheep (200-250 individuals), free to graze in the reserve. The abundance of rabbits

in 2002-2005 and 2008 is equal to 6.97% more than the expected value; in the absence of

the grazing sheep flock (2006-2007) instead the population assumed density values signifi-

cantly higher than expected. Since grazing of rabbits, especially in summer, is critical for

the subsequent composition of herbaceous vegetation, for the purposes of conservation and

protection of natural or semi natural environments, especially with a small extension, it is

desirable to properly assess and manage the presence of the wild rabbit, especially in areas

where it is not original, for the effects it may have on plant communities and populations of

arthropods, and to control sheep grazing as well to contain the caused damages.

KEY WORDS Oryctolagus cuniculus', Ovis aries aries', over grazing; small protected area.

Received 28.11.2013; accepted 22.12.2013; printed 30.12.2013

INTRODUCTION

The wild rabbit Oryctolagus cuniculus (Linna-

eus, 1758) (Mammalia Leporidae) is considered to

be endemic of the Iberian Peninsula (Dobson M.,

1998; Alves et al., 2008; Amori et al., 2008) and

since the Neolithic is widespread in the rest of the

Mediterranean (Hardy et al., 1994; Ferrand &

Branco, 2007; Alves et al., 2008; Amori et al., 2008).

It has demonstrated to be a successful colonizer,

since is currently present in at least 150 islands

around the Mediterranean Basin (Masseti & De Mari-

nis, 2008). It is a nocturnal and strongly gregarious

animal, which can live in colonies the size of which

is proportional to the availability of food. In Sicily

is naturalized and widely distributed, although non-

uniformly and with very variable density (Amori et

al., 1996; AA.VV., 2008). Individuals present in this

558

Agatino Maurizio Siracusa & Ettore Petralia

area show intermediate size within the range of vari-

ability known for populations of Spain (Andalusia)

and France (Camargue) (Siracusa et al., 2007; Lo

Valvo et al., 2008); it is supposed that the current

biometric characteristics of the species are the result

of immissions for the purpose of restocking with

strains of different origins (Lo Valvo et al., 2008).

In Southern Spain ecosystems it is a key species;

it participates in determining the community struc-

ture of predators and scavengers, as well in support-

ing their populations density (Delibes & Hiraldo,

1981; Ferrer & Negro, 2004; Delibes-Mateos et al.,

2007). Besides it plays a strong influence on the habi-

tat of other species (“ecosystem engineers”) by con-

suming vegetation, dispersing the seeds, spreading

latrines and digging burrows (Delibes-Mateos et al.,

2008; Delibes-Mateos et al., 2009; Galvez-Bravo et

al. 2009). It has a significant role even outside the

areas of origin; being prey for many vertebrates, it

maintains, at low density of grazing, a greater plant

diversity and sustains a better quality of habitat for

some species of invertebrates (Norbuiy, 1996;

Wray, 2006). In contrast if present at high density

can cause significant damage to crops and natural

vegetation. In the Iberian Peninsula the wild rabbit

is in decline from early ‘50s (Ward., 2005; Williams

et al., 2007), and currently the state of conservation

in this region, based on the IUCN criteria, is as-

sessed as “Vulnerable” (Virgos et al., 2007). Ac-

cording to Smith & Boyer (2008; IUCN, 2013), the

species is globally evaluated as near threatened

(NT). For these reasons, it is necessary to manage

the wild rabbit through scientific criteria to maintain

auto-sustainable populations and prevent (or inter-

vene in the case of) significant damage to the

economy or to the natural heritage.

The aim of the present study was to investigate

the relationship between wild rabbit and domestic

sheep Ovis dries dries (Linnaeus, 1758) (Mammalia

Bovidae), considering also the possible synergistic

effect of these two grazing herbivores on the annual

development of natural vegetation in a small

protected area.

MATERIALS AND METHODS

Study area

The study area is the Natural Reserve “Com-

plesso Immacolatelle e Micio Conti”, an area in the

foothills of Mount Etna. It is included within the

SIC ITA070008 (“Complesso Grotte Immaco-

latelle, Micio Conti e boschi limitrofi”) and covers

about 70 hectares; it is divided into a Zone A of ab-

solute restriction and a Zone B where the fruition

is permitted (Fig. 1).

In the area habitats are affected by agro-pastoral

activities, which determined the degradation of the

original vegetation. The tree cover consists of

Quercus virgiliana (Ten.), Quercus amplifolia Gus.,

which are accompanied by Celt is australis L.; there

are also several individuals of Olea europea L., re-

mains of an ancient cultivation. The herbaceous

vegetation that colonizes the most degraded areas

has several species typical of arid and uncultivated

lands as Hyparrhenia hirta (L.), Carlina corymbosa

L., Lobularia marittima (L.), Dittrichia graveolens

(L.) Greuter, Micrometria graeca (L.), Mandragora

autumnalis Bertol and the endemic Helitropium

bocconei Guss. Finally, the crags are covered with

plant species characteristic of the Mediterranean

maquis including Euphorbia dendroides L. asso-

ciated with various other sclerophyllous such as

Olea oleaster (Hoffmg. et Lk.), Pistacia therebintus

L., Rhamnus alaternus L. and liana species as Smilax

aspera L., Asparagus acutifolius L. and A. albus L.

Monitoring survey design

For counting rabbits was applied the method

proposed by Taylor & Williams (1956), used for

similar studies in the Donana National Park (Spain)

(Moreno & Villafuerte, 1992) and in several pro-

tected areas of Sicily (Caruso & Siracusa, 2001;

Siracusa et al., 2004; Siracusa et al., 2005; Siracusa

et al., 2007). This methodology consists in counting

the number of excrement in fixed detection stations

scattered randomly in the study area. The data may

provide information on population size and numer-

ical trends over time (Meriggi, 1991). This method

has been confirmed successful for the good corre-

lation between the number of excrements and the

density of animals (Wood, 1988; Palomares, 2001).

To obtain an estimation of the density of animals

(D) in an area of identified surface, knowing the

number of excrements deposited daily by the ani-

mal (r) and the period of time in which these have

been deposited (t), we applied the formula of

Eberhardt & Van Etten (1956):

D = d/rt

where d is the mean density of excrement for sam-

Trend of a population of Wild Rabbit in relation to Domestic Sheep in a small insular protected area

559

Figure 1. View from above of the study area; red line represents the boundaries of the Reserve.

pling station. Due to the lack of experimental data

for Sicily, r was considered equal to 350, the value

reported by Moreno & Villafuerte (1992) for the

Mediterranean environments, value however not far

from those found in the UK and Australia (Lockley

in Pages, 1980; Wood, 1988). The parameter (t) is

instead the number of days between the cleaning of

stations and counting.

The number of stations (n = 30), with circular

shape and large 1.54 m 2 , was chosen in function

of the abundance of excrement (> 30 escr./l .54 m 2 )

found at the site subjected to investigation. This

value was defined based on asymptotic functions

of the ratio number of plots-mean value of excre-

ment found. The number of pellets per station also

showed an aggregate distribution (variance con-

siderably higher than the average); extreme values

found tend to alter the average because the distri-

bution of data appeared irregular (Fowler &

Cohen, 1993).

Between 2002 and 2008 we calculated the an-

nual density of the rabbit population in the Reserve

through monthly surveys of the population (see also

Siracusa et al., 2005). At the same time have been

measured some meteorological parameters (mean

annual temperature, mean annual precipitation, total

precipitation and annual total precipitation for each

month of the year) in order to verify a possible in-

fluence of these on the annual density of wild rab-

bits. The correlations were made both considering

the values of the same year and those of the pre-

vious year in order to identify a possible delayed

influence of these parameters.

In the course of the study was estimated as

well (by direct counting) the number of grazing

sheep present, with the purpose to verify for any

interference with the population of wild rabbit,

also to assess the amount of grazing pressure of

both sheep and rabbits. Was also calculated an ex-

pected value of density (spec. /ha) of wild rabbit,

combining the number of species of Poacae present

and the altitude above sea level using 9 different

sample areas by the technique of Multiple Regres-

sion. For the comparison between the abundance

of rabbits in different years under investigation

was used instead the x 2 test. For the correlation

560

Agatino Maurizio Siracusa & Ettore Petralia

between variables (meteorological parameters and

density) was used the Spearman rank correlation

coefficient (rs).

Statistical analyses were performed using the

software STATISTIC A 5.0.

RESULTS

The annual average density of wild rabbits was

found to be 44.79± 27.34 (s.d.) ind./ha, with a

minimum value of 25.6 ind./ha in 2005 and a

maximum value of 94.6 ind./ha in 2007 (Fig. 2).

The value of density can be predicted by the

number of species of Poacae and the altitude above

sea level (R 2 = 0.822; F 2 6 = 13.881; P < 0.006), where

both variables resulted as predictive (p <0.003)

(Fig. 3).

The expected value of density is equal to 36.3

(± 95.0 % 51.1-21.5) ind./ha. None of the climatic

variables correlated statistically significantly with

the density of the species, with the exception of

the significant negative correlation between the

density of rabbits and the annual amount of precip-

itation during the month of September (r s =- 0.929;

n=7; p=0.002).

The comparison between the density values ob-

tained does not show statistically significant differ-

ences during the years 2002, 2003, 2004, 2005 and

2008 (% 2 =2.74, df=4, p < 0.60). However, there

are differences with the abundance values observed

in 2006 and 2007 (x 2 =19.51, df=2, P=0.000).

Noteworthy, during the all period, with the ex-

ception of 2006 and 2007, within the area was pre-

sent a flock of sheep (200-250 individuals) free to

graze in the reserve.

DISCUSSION

In Sicily, the main factors that seem to affect the

density of this species are the extension of aban-

doned ex-cultivated areas (and / or uncultivated

areas) (Caruso & Siracusa, 2001; Siracusa et al.,

2007), the altitude above sea level and the number

of species of Poacae (Caruso & Siracusa, 2001;

Siracusa et al., 2007).

It’s rather less probable that the wild rabbit is

generally subject to limitation by the community

of predators (Trout & Tittensor, 1989; Moreno &

Villafuerte, 1992; Trout et al., 2000; Caruso &

Siracusa, 2001; Siracusa et al., 2004; Siracusa et

al., 2007). Habitat fragmentation is also consid-

ered one of the causes of the decline of local pop-

ulations of wild rabbit (Virgos et al., 2003); the

site is not isolated from the surrounding areas and

the degree of fragmentation of the territory, for the

presence of the wild rabbit, can be considered of

small entity with the opportunity for individuals

to disperse and maintain normal phenomenon of

immigration/ emigration.

In another area with Mediterranean climate, pre-

dominantly semiarid, in the NE of Spain, Calvete

et al. (2004) found a positive correlation between

Figure 1. Annual trend of wild rabbit abundance observed

during the years 2002-2008 within the Reserve “Immaco-

latelle-Micio Conti"

EipefWnsUM

Figure 2. Expected and observed values of wild rabbit den-

sity at nine different sites in the Etna area (the arrow indi-

cates the Reserve “Immacolatelle-Micio Conti").

Trend of a population of Wild Rabbit in relation to Domestic Sheep in a small insular protected area

561

the annual mean temperature, the average of rainfall

in the months of February and May, and abundance

of rabbits in different sites, while the average rain-

fall for the months of September and November

correlated negatively.

In this protected area of Mount Etna the total

rainfall for the month of September is negatively

correlated with the density per year. It is possible

that heavy rainfall in early autumn can determine

an increase of winter mortality due to various fac-

tors such as greater difficulty in thermoregulation,

excessive soil moisture with limited plant growth

resulting in food shortages and delays in beginning

of the breeding season (Calvete et al., 2004). Me-

teorological events, concentrated in a few days can

also have negative effects on rabbit populations,

especially in open areas (Palomares, 2003).

The abundance of rabbits in 2002-2005 and

2008 (29.30 ± 4.48 ind./ha) is equal to 6.97% more

than the expected value. In the absence of the graz-

ing sheep flock (2006-2007) instead the population

assumed density values (83.1 ind./ha) significantly

higher than expected (23.8 ind./ha); under these con-

ditions the grazing of rabbits, especially in late sum-

mer, resulted in high proportions. Generally

speaking, the remarkable grazing of rabbits in the

summer is critical as fundamental for the subsequent

composition of herbaceous vegetation in the follow-

ing spring (Myers & Poole, 1963). To this it should

be also added the grazing activity of sheep.

As an indication, if one considers 390.9 gr the

food ingested daily by a rabbit (Meakin et al.,

2002) for its metabolic needs, the annual demand

per hectare of the population is on average 17.5 kg

of vegetables. It should also be estimated that a

sheep consumes the equivalent of 10-16 rabbits

(Myers et al., 1994).

In addition, the density values occurred in 2006

and 2007 are high when compared with other areas

of Mount Etna (Siracusa et al., 2004). An excessive

presence of rabbits, because of overgrazing, may

adversely affect the normal evolution in the succes-

sion of the vegetation (Zeevallcing & Fresco in

Begon et al., 1986) determining a reduction of plants

diversity and sometimes even causing the extinction

of some species. The most important factors that

cause these changes are the selection by the rabbit

of some species and parts of plants, especially dur-

ing the critical period of vegetative growth, and the

mechanical destmction of the soil surface; both of

these factors facilitate, in competition, some species

than others (Myers & Pool, 1963).

Elerbivores influence the species richness of

plants in many terrestrial ecosystems by affecting

their structure, composition and function; the num-

ber of plant species is also closely linked to the size

of the area (AA.VV. in Albon et al., 2007; Olofsson

et al., 2008). Grazing and deposition of excrements

of rabbits creates spatial heterogeneity in vegetation

and soil resources.

An overexploitation of vegetation, due to an

excessive number of rabbits, tends so to reduce the

heterogeneity and structural complexity of plant

communities with indirect effects on populations

of arthropods (both phytophagous and predators -

'intermediate disturbance hypothesis'), in particu-

lar on the abundance of many species with conse-

quent negative effects on functionality of the

ecosystem considered (compare for example Moss

& Hassal, 2006).

It should be also taken into consideration the

high impact of sheep, due to their behaviour that

tends to be aggregative and with limited displace-

ments, with consequent action of disturbance in

limited areas. Among the effects of the removal of

sheep there are significant changes in the compo-

sition and structure of vegetation, with clear reper-

cussions on populations of wild herbivores (Hope

et al., 1996). Are also documented the effects of

sheep grazing on the arthropod communities

through changes in the composition and structure

of vegetation heterogeneity (Dennis, 2003).

Wild rabbits and sheep, as well as other domes-

tic ruminants are herbivorous grazers that often feed

on the same plant species; for this reason among

them there is a strong competition for food

(Dawson & Ellis, 1979; Soriguer, 1983).

CONCLUSIONS

The relationship wild rabbit-vegetation-habitat

has been studied thoroughly both in the area where

the species is native and where it has been intro-

duced (see for example Copson & Whinam, 1998;

Moller et al., 1997; Norbury & Norbury, 1996;

Zeevalking & Fresco, 1977; Denyer et al., 2007

and their references). A moderate grazing is con-

sidered positive for vegetation, contributing to in-

crease diversity; the disappearance of the rabbit in

562

Agatino Maurizio Siracusa & Ettore Petralia

some areas has in fact led to the development of

dominant plant species with consequent effects on

plant communities (such as decrease or disappear-

ance of the most sensitive species and reduction

of biological diversity).

For the purposes of conservation and protection

of natural or semi natural environments (especially

with small extension) it is desirable to properly

assess and manage the presence of the wild rabbit,

especially in areas where it is not original, and to

control as well sheep grazing in order to contain the

caused damages.

ACKNOWLEDGEMENTS

We wish to thank the staff of the Natural Re-

serve “Complesso Immacolatelle e Micio Conti”,

especially Dr. Salvo Cartarrasa and Dr. Silvia

Giuffrida who made possible the realization of this

study.

REFERENCES

AA. VV, 2008. Atlante della Biodiversita della Sicilia:

Vertebrati terrestri. Studi e Ricerche, 6, Arpa Sicilia,

Palermo, 536 pp.

Albon S.D., Brewer M.J., O’ Brien S., Nolan A.J. & Cope

D., 2007. Quantifing the grazing impacts associated

with different herbivores on rangelands. Journal of

Applied Ecology, 44: 1176-1187.

Amori G., Angelici F.M., Prigiont C. & Vigna Taglianti

A., 1996. The Mammal Fauna of Italy: A Review.

Hystrix, the Italian Journal of Mammalogy, 8: 3-7.

Amori G., Contoli L. & Nappi A., 2008. Mammalia II:

Erinaceomorpha, Soricomorpha, Lagomorpha,

Rodentia. Fauna d'Italia,V. XLIV, ed. Calderini,

Bologna, 786 pp.

Begon M.J., Harper J.L. & Townsend C.R., 1986. Ecol-

ogy: individuals, populations, and communities. Sin-

auer Associates, Sunderland, Massachusetts, 1068

pp. + xii.

Camso S. & Siracusa A.M., 2001. Factors affecting abun-

dance of Wild rabbits ( Oryctolagus cuniculus) in

agro-ecosistem of the Mount Etna Park. Hystrix, the

Italian Journal of Mammalogy, 12: 45-49.

Copson G. & Whinam J., 1998. Response of Vegetation

on Subantarctic Macquarie Island to Reduced Rabbit

Grazing. Australian Journal of Botany, 46: 15-24.

Dawson T.J. & Ellis B.A., 1979. Comparison of the diets

of yellow-footed rock-wallabies and sympatric

herbivores in western New South Wales. Australian

Wildlife Research, 6: 245-254.

Delibes M. & Hiraldo F., 1981. The rabbit as prey in the

Iberian Mediterranean ecosystem. In: Meyers K.,

Machines C.D. (Eds). Proceedings of the World

Lagomorph Conference. University of Guelph, On-

tario, pp. 614-622.

Delibes-Mateos M., Delibes M., Ferreras P. & Villafuerte

R. , 2008. Key role of European rabbits in the conser-

vation of the Western Mediterranean Basin hotspot.

Conservation Biology, 22: 1 106-1 117.

Delibes-Mateos M., Ferreras P. & Villafuerte R., 2009.

European rabbit population trends and associated fac-

tors: a review of the situation in the Iberian Penisula.

Mammal Review, 39: 124-140.

Delibes-Mateos M., Redpath S.M., Angulo E., Ferreras

P. & Villafuerte R., 2007. Rabbits as a keystone

species in southern Europe. Biological Conservation,

137: 149-156.

Dennis P., 2003. Sensitivity of upland arthropod diversity

to livestock grazing, vegetation stmcture and land-

form. Agriculture and Environment, 1: 301-307.

Dobson M., 1998. Mammal distributions in the western

Mediterranean: the role of human intervention. Mam-

mal Review, 28: 77-88.

Eberhardt L. & Van Etten R.C., 1956. Evaluation of the

Pellet Group Count as a Deer Census Method.

Journal of Wildlife Management, 20: 70-74.

Fa J.E., Sharpies C.M. & Bell D.J., 1999. Habitat corre-

lates of European rabbit ( Oryctolagus cun iculus ) dis-

tribution after the spread of RVHD in Cadiz Province,

Spain. Journal of Zoology, 249: 83-96.

Ferrand N. & Branco M., 2007. The evolutionary history

of the European rabbit ( Oryctolagus cuniculus ): major

patterns of population differentiation and geographic

expansion inferred from protein polymorphism. Weiss

S. & Ferrand N. (Eds). Phylogeography of Southern

European Refugia. Springer: Netherlands, pp. 207-

235.

Ferrer M. & Negro J.J., 2004. The near extinction of two

large European predators: super specialist pay a price.

Conservation Biology, 18: 344-349.

Galvez-Bravo L., Belliure J. & Rebollo S., 2009. Euro-

pean rabbits as ecosystem engineers: warrens in-

crease lizard density and diversity. Biodiversity and

Conservation, 18: 869-885.

Hardy C., Vigne J.D., Casane D., Dennebouy N.,

Mounolou J.C. & Monnerot M., 1994. Origin of Eu-

ropean rabbit ( Oryctolagus cuniculus ) in a Mediter-

ranean island: Zooarchaeology and ancient DNA

examination. Journal of Evolutionary Biology, 7:

217-226.

Hope D., Picozzi N., Catt D.C. & Moss R., 1996. Effects

of reducing sheep grazing in the Scottish Highland.

Journal of Range Management, 49: 301-310.

Trend of a population of Wild Rabbit in relation to Domestic Sheep Ovis aries aries in a small insular protected area 563

Denyer J.L., Hartley S.E., John E.A., 2007. Small mam-

malian herbivore determines vegetation response to

patchy nutrient inputs. Oikos, 116: 1186-1192.

Lo Valvo M., La Scala A., Giacalone G. & Ticali S.,

2008. Conteggi di Coniglio selvatico Oryctolagus

cuniculus (Linnaeus, 1758), in aree campione della

provincia di Palermo. In: Prigioni C., Meriggi A. &

Merli E. (Eds) 2008. VI Congr. It. Teriologia, Hys-

trix, the Italian Journal of Mammalogy (N.S.) Suppl.

2008: 1-124.

Massed M. & De Marinis A.M., 2008. Prehistoric and

historic artificial dispersal of Lagomorphs on the

Mediterranean islands. In: Alves P.C., Ferrand N. &

Hacklander K. (Eds). Lagomorph biology. Evolution,

ecology, and conservation. Springer, Berlin; Heidel-

berg; New York: pp. 13-25.

Meriggi A., 1991. L’uso dei dati di popolazione per la

gestione delle specie oggetto di prelievo. In: Fasola

M. (red.) Atti II Seminario Italiano Censimenti Fau-

nistici dei Vertebrati. Suppl. Ric. Biol. Selvaggina,

XVI: 1-820.

Meakin I., Adamson P. & Hoy S., 2002. Methods for esti-

mating daily food intake of wild birds and mammals.

Central Scienze Laboratory DEFRA. Final report, pp.

1 - 22 .

Myers K. & Pool W.E., 1963. A study of the biology of

the Wild Rabbits, Oryctolagus cuniculus (L.) in con-

fined population IV. The effect of rabbit grazing on

sown pastures. Journal of Ecology, 51: 435-451.

Myers K., Parer I., Wood D. & Cooke B.D., 1994. The

history and biology of a successful colonizer: the rab-

bit in Australia. In: Thompson H.V. & King C.M.

(Eds), Oxford, Oxford Science Publications.

Moller H., Clapperton B.K. & Fletcher D.J., 1997. Den-

sity of rabbits ( Oryctolagus cuniculus L.) in the

Mackenzie Basin, South Island, New Zeland. New

Zeland Journal of Ecology, 21: 161-167.

Moreno S. & Villafuerte R., 1992. Seguimiento de las

poblaciones de Conejo en le Parque Nactional de

Donana. Estaction Biologica de Donana (C.S.I.C.),

77 pp.

Moss, A. & M. Hassall., 2006. Effects of disturbance on

the biodiversity and abundance of isopods in tem-

perate grasslands. European Journal of Soil Biology,

42: 254-268.

Norbury D., 1996. The effects of rabbit on conservation

values. Science of Conservation: 34: 1-32.

Norbury D.C. & Norbury G.L., 1996. Short-term effects

of rabbit grazing on a degraded short-tussock grass-

land in Central Otago. New Zeland Journal of Eco-

logy, 20: 285-288.

Olofsson J., De Mazancourt C. & Crawley M.J., 2008.

Spatial heterogeneity and plant species richness at

different spatial scales under rabbit grazing. Oecolo-

gia, 156:825-834.

Pages M.V., 1980. Methodes d'etude de la repartition

spatiale du lapin de garenne dans deux milieux

differents du Languedoc. Bulletin Men. O.N.C., n°

special, Dec. 1980: 111-126.

Palomares F., 2001. Comparison of 3 methods to estimate

rabbit abundance in a Mediterranean environment.

Wildlife Society Bulletin, 29: 578-585.

Palomares F., 2003. The negative impact of heavy rains

on the abundance of a Mediterranean population of

European rabbits. Mammalian Biology, 68: 224-234.

Siracusa A.M., Caruso S. & Leonardi G., 2005. Abundance

and seasonal fluctuations of the rabbit ( Oryctolagus

cuniculus) in agrocoenoses of Mount Etna, Sicily. Atti

della Societa italiana di scienze naturali e del Museo

civico di storia naturale di Milano, 146: 117-126.

Siracusa A.M., Giuffrida S. & Messina A., 2004. Densita’

e fluttuazioni numeriche di una popolazione di Coniglio

selvatico ( Oryctolagus cuniculus) (Mammalia Lago-

morpha) in un’area protetta della Sicilia. II Natural-

ista siciliano, 28: 1195-1203.

Siracusa A.M., Leonardi G., Grasso R, Di Maio M.C.,

Caruso S. & Petralia A., 2007. Indagini sulle popo-

lazioni di coniglio selvatico ( Oryctolagus cuniculus)

in due aree protette della Provincia di Ragusa. Atti e

Memorie dell’Ente Fauna siciliana, 9 (2002-03-04):

49-86.

Smith A. T. & Boyer A.F., 2008. Oryctolagus cuniculus.

In: IUCN 2013. IUCN Red List of Threatened

Species. Version 2013.2. <www.iucnredlist.org>.

Downloaded on 25 November 2013.

Soriguer R.C., 1983. Consideraciones sobre el efecto de

los conejos y los grandes herbivoros en los pastizales

de la Vera de Donana. Donana. Acta Vertebrata,

10:155-168.

Taylor R.H. & Williams R.M., 1956. The use of pellets

counts for estimating the density of populations of

wild rabbit, Oryctolagus cuniculus. The New Zealand

Journal of Science and Technology, 38: 236-256.

Trout R.C. & Tittensor A.M., 1989. Can predators regu-

late wild Rabbits Oryctolagus cuniculus population

density in England and Walles?. Mammal review, 19:

153-173.

Trout R.C., Langton S., Smith G.C. & Haines-Young R.

H., 2000. Factors affecting the abundance of rabbits

{Oryctolagus cuniculus) in England and Wales. Jour-

nal of Zoology, 252: 227-238.

Virgos E., Cabezas-Diaz S., Malo A., Lozano J. & Lopez-

Huertas D., 2003. Factors shaping European rabbit

abundance in continuous and fragmented populations

of central Spain. Acta Theriologica, 48: 113-122.

Virgos E., Cabezas-Diaz S. & Lozano J., 2007. Is the

wild rabbit {Oryctolagus cuniculus) a threatened

species in Spain? Sociological constraints in the con-

servation of the species. Biodiversity and Conserva-

tion, 16: 3489-3504.

564

Agatino Maurizio Siracusa & Ettore Petralia

Zeevalking H.J. & Fresco L.F.M., 1977. Rabbit grazing

and species diversity in a dune area. Vegetatio, 35:

193-196.

Ward D., 2005. Reversing Rabbit Decline: One of the

biggest challenges for nature conservation in Spain

and Portugal. Cat Specialist Group, Lagomorph Spe-

cialist Group, 54 pp.

Williams D., Acevedo P., Gortazar C., Escudero M.A.,

Labarta J.L., Marco J. & Villafuerte R., 2007. Hunting

for answers: rabbit ( Oryctolagus cuniculus ) popula-

tion trends in northeastern Spain. European Journal

of Wildlife Research, 53: 19-28.

Wood D. H., 1988. Estimating rabbit density by counting

dung pellets. Australia Wildlife Research, 15: 665-

671.

Wray S., 2006. A guide of rabbit management. CIRIA,

London, 12 pp.

Biodiversity Journal, 2013, 4 (4): 565-570

Remarks on the genus Sphecodes Latreille, 1 804, pinguiculus

Perez, 1 903 species-group (Hymenoptera Apidae) from Italy:

a reply to the article of Schwarz & Gusenleitner (20 1 2)

Vittorio Nobile 1 & Giuseppe FabrizioTurrisi 2

'Via Psaumida, 17, lotto 25 - 97100, Ragusa, Italy; e-mail: nobilevittorio@tin.it

2 via Cristoforo Colombo 8 - 95030, Pedara, Catania, Italy; e-mail: tuiTisifabrizio@yahoo.it

’Corresponding author

ABSTRACT The present paper is a reply to the article by Schwarz & Gusenleitner (2012) and deals with

the taxonomy of the genus Sphecodes Latreille, \&04, pinguiculus Perez, 1903 species-group.

Particularly, this study, after a critical revision of the morphological characters used to the

identification of the species of the genus Sphecodes from Italy, confirms the results of Nobile

& Turrisi (2004) and allows the revalidation of all species which they described; therefore

are considered valid species the following taxa: S. campadellii Nobile et Turrisi, 2004, S.

combai Nobile et Turrisi, 2004, S. banaszaki Nobile et Turrisi, 2004, S. marcellinoi Nobile et

Turrisi, 2004, S. walteri Nobile et Turrisi, 2004, S. iosephi Nobile et Turrisi, 2004, S. tomar-

chioi Nobile et Turrisi, 2004.

KEY WORDS Sphecodes', pinguiculus species-group; taxonomic discussion; species revalidated.

Received 15.12.2013; accepted 27.12.2013; printed 30.12.2013

INTRODUCTION

The identification of the species of the cleptopar-

asitic genus Sphecodes Latreille, 1804 is somewhat

problematic (Meyer, 1919; Bogusch& Straka, 2012)

and, to facilitate the identification, several Authors

have been recognized the usefulness of the morphol-

ogy of the antennomeres. Whereas the females have

the antennomeres normally shaped, instead the males

have mostly antennomeres ventrally gibbous. In

more detail, the gibbous process is always placed dis-

tally on antennomere, whereas proximally, beginning

from the base, it is present a variously shaped and

more or less deep notch (anular, half-moon like, etc.)

covered by short, bristly and white setae, which form

sensillar plates. At low magnification, these sensillar

plates resemble white spots, defined differently by

several Authors: “reticulatio subtilis grisea”

(Thomson, 1872); “graaktigt rotfalt” (Aurivillius,

1903); “svobepletter” (Jorgensen, 1921); “pilose

facets” (Mitchell, 1960); “flagellar sensilla” (Agren

& Svensson, 1982); “Haarfleck” (Hagens, 1882;

Meyer, 1919; Bliithgen, 1923; Wamcke, 1992);

“white setae” (Nobile & Turrisi, 2004); “felt-like

pubescence” (Bogusch & Straka, 2012). It is note-

worthy that on each antennomere, it exists a comple-

mentarities, e.g. if the notched area is relatively

small, conversely, the gibbous area is wider, and vice

versa. These peculiar morphological features of

antennomeres are constant within a species but show

distinct patterns among different Sphecodes species,

thus providing good characters for identification, as

confirmed by careful investigation of antennae of

eleven Swedish Sphecodes species carried out by

Agren & Svensson (1982), who pointed out as fol-

lowing indicated:

566

Vittorio Nobile & Giuseppe FabrizioTurrisi

a) on antenna of both males and females there

are about ten diverse types of sensillar structures;

b) some sensilla are erect, hair-like, short and

white (e.g., sensilla trichodea, etc.), whereas, some

other are flattened, not evidently protruding above

cuticle (e.g., sensilla placodea, etc.);

c) in the males, on the ventral surface of flagel-

lomeres, the erected sensilla (Haarfleck, etc.) are

placed basally, covering the notched area, whereas,

the flattened sensilla cover the distal, gibbous area

of the same flagellomeres;

d) in the females, the differently shaped sensilla

are mixed up, covering uniformly the flagellomeres.

Based solely on the morphology of the male

antenna of some Sphecodes- species, Agren &

Svensson (1982) provided a key for their identi-

fication. Utilizing these antennal characters, as

well as other ones, Warncke (1992) provided a

key to the male of the West Palaearctic Sphecodes-

species. However, in the species belonging to the

S. pinguiculus species-group, the antennal flagel-

lum of males, at least beginning from the third flagel-

lomere, is uniformly covered by sensilla thus,

without the characteristic sequence of notch-gibbous

areas. Nobile & Turrisi (2004) have been recorded

S. pinguiculus and additionally have described

seven new Italian species of Sphecodes , having a

similar antennal morphology of the former species.

Based on antennal morphology and other com-

monly shared features, Nobile & Turrisi (2004)

proposed the institution of the S. pinguiculus

species-group, providing a comprehensive key to

species. Recently, these newly described species

have been synonymised by Schwarz & Gusenleit-

ner (2012), on the basis of the examination of type

material.

In the present paper, the distinctive characters

of the seven species of Sphecodes described by

Nobile & Turrisi (2004) are confirmed and pointed

out, demonstrating the inconsistency of the syn-

onymies by Schwarz & Gusenleitner (2012) and

thus confirming the validity of these species.

MATERIALS AND METHODS

The present paper is based on primary type ma-

terial of the seven species described by Nobile &

Turrisi (2004), preserved in the collection of Zool-

ogische Staatssammlung Miinchen (Germany).

RESULTS

In the following, we point out the results of the

reexamination of each species of the S. pinguiculus

species-group described by Nobile & Turrisi

(2004), which confirm their validity.

Sphecodes banaszaki Nobile et Turrisi, 2004

Sphecodes banaszaki , Nobile & Turrisi, 2004. En-

tomofauna, 25 (8): 120 (Italia).

Sphecodes marginatus, Schwarz & Gusenleitner,

2012. Entomofauna, 33 (8): 74.

From the comparison between S. marginatus Ha-

gens, 1882 and S. banaszaki Nobile et Turrisi, 2004

(the latter retained synonym of S. marginatus by

Schwarz & Gusenleitner, 2012), and the descriptions

provided by Hagens (1882) and Meyer (1919), as

well as the identification keys provided by Wamcke

(1992) and Bogusch & Straka (2012), it is possible to

point out many remarkable differences between the

two considered species, the most important being:

• S. banaszaki has the flagellomeres, excluding

the basal two, uniformly covered by sensilla on the

ventral surface (Haarfleck, sensilla), without the

characteristic sequence of alternate notch and gib-

bous areas, as also pointed out by the same Schwarz

& Gusenleitner (2012: fig. 1); conversely, in S.

marginatus , each flagellomere, excluding the basal

three, bears a basal white spot of sensilla alternate

with an evident and well protruded distal gibbous

area, as also clearly showed by Bogusch & Straka

(2012: fig. 122);

• the male genital capsule of S. banaszaki has

the membranous area of the gonostylus wide and

quadrangular shaped (Nobile & Turrisi, 2004: fig.

3; Schwarz & Gusenleitner, 2012: figs. 7a-d), thus,

quite different from S. marginatus , which has a less

wide and triangular shaped membranous area (Ha-

gens, 1882: fig. 18; Wamcke, 1992: fig. 32; Bo-

gusch & Straka, 2012: figs. 165, 166). Moreover,

the apex of the sclerifed part of the gonostylus of S.

banaszaki is more robust and more developed than

S. marginatus.

These strong differences between S. banaszaki

and S. marginatus , affecting important features of

head and metasoma, and clearly not running within

intraspecific variation, have been overlooked by

Schwarz & Gusenleitner (2012); moreover, while S.

Remarks on the genus Sphecodes pinguiculus species-group (Hymenoptera Apidae) from Italy

567

banaszaki belongs to the S. pinguiculus species-

group, S. marginatus belongs to the S. miniatus Ha-

gens, 1882 species-group (Bogusch & Straka, 2012).

S. banaszaki is deeply different from S. margi-

natus , and taking also into account the differences

with the other species of the same group, S. bana-

szaki Nobile et Turrisi, 2004 is valid species.

Sphecodes campadellii Nobile et Turrisi, 2004

Sphecodes campadellii , Nobile & Turrisi, 2004. En-

tomofauna, 25 (8): 118 (Italia).

Sphecodes geoffrellus , Schwarz & Gusenleitner,

2012. Entomo fauna, 33 (8): 74.

From the comparison between S. geoffrellus

(Kirby, 1802) and S', campadellii (the latter retained

synonym of S. geoffrellus by Schwarz & Gusenleit-

ner, 2012) and the descriptions provided by Hagens

(1882) and Meyer (1919), as well as the identifica-

tion keys provided by Wamcke (1992) and Bogusch

& Straka (2012), it is possible to point out many re-

markable differences between the two considered

species, the most important being:

• S. campadellii has flagellomeres, excluding

the basal two, uniformly covered by sensilla on the

ventral surface (Haarfleck, etc.), without the

characteristic sequence of alternate notch and gib-

bous areas (similarly to that presented by Schwarz

& Gusenleitner, 2012: fig. 1); instead S. geoffrellus

has each of the median flagellomeres covered for at

most 3/4 of the surface by a white spot of sensilla

alternate with evident gibbous areas.

Moreover, S. campadellii belongs to the S. pin-

guiculus species-group, whereas S. geoffrellus be-

longs to a different group.

S. campadellii is deeply different from S. geof-

frellus, and taking also into account the differences

with the other species of the same group, S. cam-

padellii Nobile et Turrisi, 2004 is valid species.

Sphecodes combai Nobile et Turrisi, 2004

Sphecodes combai, Nobile & Turrisi, 2004. Ento-

mofauna, 25 (8): 119 (Italia).

Sphecodes marginatus, Schwarz & Gusenleitner,

2012. Entomofauna, 33 (8): 75.

From the comparison between S. marginatus

and S. combai (the latter retained synonym of S.

marginatus by Schwarz & Gusenleitner 2012) and

the descriptions by Hagens (1882) and Meyer

(1919), as well as the identification keys provided

by Wamcke (1992) and Bogusch & Straka (2012),

it is possible to point out many remarkable dif-

ferences between the two considered species, the

most important being:

• S. combai has flagellomeres, excluding the

basal two, uniformly covered by sensilla on the ven-

tral surface (Haarfleck, etc.), without the character-

istic sequence of alternate notch and gibbous areas

(similarly to that presented by Schwarz & Gusen-

leitner, 2012: fig. 1); instead, ins', marginatus, each

flagellomere, excluding the basal three, bears a

basal white spot of sensilla alternate with an evident

and well protmded distal gibbous area, as also clearly

showed by Bogusch & Straka (2012: fig. 122);

• S. combai has the first metasomal tergite

polished and shiny, due to the absence of microsculp-

ture, with fine, superficial and scattered punctures

(distance between punctures 1.0-3. Ox puncture

diameter); instead S. banaszaki (= S. marginatus ?)

has the first metasomal tergite dull, due to the pre-

sence of microsculpture, with coarse, deep and

dense punctures (distance between punctures 1.0-

1 .5x puncture diameter) (Nobile & Turrisi, 2004;

Schwarz & Gusenleitner, 2012: fig. 4).

These strong differences between S. combai and

S. marginatus, affecting important features of head

and metasoma and clearly not mnning within in-

traspecific variation, have been overlooked by

Schwarz & Gusenleitner (2012); moreover, while

S. combai belongs to the S. pinguiculus species-

group, S. marginatus belongs to the S. miniatus

species-group (Bogusch & Straka, 2012).

S. combai is deeply different from S. margina-

tus, and taking also into account the differences

with the other species of the same group, S. combai

Nobile et Turrisi, 2004 is valid species.

Sphecodes marcellinoi Nobile et Turrisi, 2004

Sphecodes marcellinoi, Nobile & Turrisi, 2004. En-

tomofauna, 25 (8): 121 (Italia).

Sphecodes marginatus, Schwarz & Gusenleitner,

2012. Entomofauna, 33 (8): 75.

From the comparison between S. marginatus

and S. marcellinoi (the latter retained synonym of

S. marginatus by Schwarz & Gusenleitner, 2012)

568

Vittorio Nobile & Giuseppe FabrizioTurrisi

and the descriptions provided by Hagens (1882) and

Meyer (1919), as well as the identification keys pro-

vided by Wamcke (1992) and Bogusch & Straka

(2012), it is possible to point out many remarkable

differences between the two considered species, the

most important being:

• S. marcellinoi has flagellomeres, excluding the

basal two, uniformly covered by sensilla on the ven-

tral surface (Haarfleck, etc.), without the character-

istic sequence of alternate notch and gibbous areas

(similarly to that presented by Schwarz & Gusen-

leitner, 2012: fig. 1); instead, in S. marginatus, each

flagellomere, excluding the basal three, bears a basal

white spot of sensilla alternate with an evident and

well protruded distal gibbous area, as also clearly

showed by Bogusch & Straka (2012: fig. 122);

• S. marcellinoi has a slightly arcuate clypeus,

nearly straight; conversely, S. walteri (= S. margina-

tus ?) has a strongly arcuate clypeus;

• S. marcellinoi has fine, superficial and scat-

tered punctuation on frons (distance between punc-

tures about 2. Ox puncture diameter); instead S.

banaszaki (= S. marginatus ?) has coarse, deep and

dense punctuation on frons (distance between punc-

tures less than puncture diameter) (Nobile & Tur-

risi, 2004; Schwarz & Gusenleitner, 2012: fig. 2).

These strong differences between S. marcellinoi

and S. marginatus , affecting important features of

head and metasoma, clearly not running within

intraspecific variation, have been overlooked by

Schwarz & Gusenleitner (2012); moreover, while

S. marcellinoi belongs to the S. pinguiculus species-

group, S. marginatus belongs to the S. miniatus

species-group (Bogusch & Straka, 2012).

S. marcellinoi is deeply different from S. margina-

tus , and taking also into account the differences

with the other species of the same group, S. mar-

cellinoi Nobile et Turrisi, 2004 is valid species.

Sphecodes walteri Nobile et Turrisi, 2004

Sphecodes walteri, Nobile & Turrisi, 2004. Ento-

mofauna25, (8): 122 (Italia).

Sphecodes marginatus, Schwarz & Gusenleitner,

2012. Entomofauna, 33 (8): 75.

From the comparison between S. marginatus

Hagens, 1882 and S. walteri (the latter retained syn-

onym of S. marginatus by Schwarz & Gusenleitner,

2012) and the descriptions provided by Hagens

(1882) and Meyer (1919), as well as the identifica-

tion keys provided by Wamcke (1992) and Bogusch

& Straka (2012), it is possible to point out many re-

markable differences between the two considered

species, the most important being:

• S. walteri, has flagellomeres, excluding the

basal two, uniformly covered by sensilla on the ven-

tral surface (Haarfleck, etc.), without the character-

istic sequence of alternate notch and gibbous areas

(similarly to that presented by Schwarz & Gusen-

leitner, 2012: fig. 1); instead in S. marginatus, each

flagellomere, excluding the basal three, bears a

basal white spot of sensilla alternate with an evident

and well protruded distal gibbous area, as also clearly

showed by Bogusch & Straka (2012: fig. 122);

• S. walteri has the clypeus strongly arcuate;

instead, S. marcellinoi (= S. marginatus ?) has the

clypeus slightly arcuate, nearly straight;

• S. walteri has irregular, coarse, deep and mod-

erately dense punctuation on frons (distance be-

tween punctures 1.0-2. Ox puncture diameter);

instead S. marcellinoi (= S. marginatus ?) has nearly

regular, fine, superficial and scattered punctuation

on frons (distance between punctures about 2. Ox

puncture diameter).

These strong differences between S. walteri and

S. marginatus, affecting important features of the

head and clearly not mn nin g within intraspecific vari-

ation, have been overlooked by Schwarz & Gusen-

leitner (2012); moreover, while S. walteri belongs

to the S. pinguiculus species-group, S. marginatus

belongs to the S. miniatus species-group (Bogusch

& Straka, 2012).

S. walteri is deeply different from S. marginatus,

and taking also into account the differences with the

other species of the same group, S. walteri Nobile

et Turrisi, 2004 is valid species.

Sphecodes iosephi Nobile et Turrisi, 2004

Sphecodes iosephi, Nobile & Turrisi, 2004. Ento-

mofauna, 25 (8): 123 (Italia).

Sphecodes marginatus, Schwarz & Gusenleitner,

2012. Entomofauna, 33 (8): 75.

From the comparison between S. marginatus

and S. iosephi (the latter retained synonym of S.

marginatus by Schwarz & Gusenleitner, 2012) and

the descriptions provided by Hagens (1882) and

Meyer (1919), as well as the identification keys pro-

Remarks on the genus Sphecodes pinguiculus species-group (Hymenoptera Apidae) from Italy

569

vided by Wamcke (1992) and Bogusch & Straka

(2012), it is possible to point out many remarkable

differences between the two considered species, the

most important being:

• S. iosephi has flagellomeres, excluding the

basal two, uniformly covered by sensilla on the ven-

tral surface (Haarfleck, etc.), without the character-

istic sequence of alternate notch and gibbous areas

(similarly to that presented by Schwarz & Gusen-

leitner, 2012: fig. 1); instead, ins', marginatus, each

flagellomere, excluding the basal three, bears a

basal white spot of sensilla alternate with an evident

and well protruded distal gibbous area, as also clearly

showed by Bogusch & Straka (2012: fig. 122);

• S. iosephi has irregular, coarse, deep and very

dense punctuation on frons (distance between punc-

tures about equal to puncture diameter); moreover,

the vertex is punctate-carinulate, with punctuation

coarse, deep and dense; instead, in S. marcellinoi

(= S. marginatus ?) the frons and vertex have punc-

tuation, with nearly regular, fine, superficial and

scattered punctuation (distance between punctures

about 2. Ox puncture diameter);

• in S. iosephi the first metasomal tergite bears

fine, superficial and scattered punctuation (distance

between punctures 1.0- 3. Ox puncture diameter); in-

stead S. banaszaki (= S. marginatus ?), has the first

metasomal tergite bearing coarse, deep and dense

punctures (distance between punctures 1.0-1 .5x

puncture diameter) (Nobile & Turrisi, 2004;

Schwarz & Gusenleitner, 2012: fig. 4);

• S. iosephi has a stout genital capsule and the

sclerified part of the gonostylus extends veiy slightly

beyond the membranous part (Nobile & Turrisi,

2004: fig. 6; Schwarz & Gusenleitner, 2012: figs.

10a- lOd); instead S. banaszaki (= S. marginatus ?)

has the sclerified part of the gonostylus well ex-

tended beyond the membranous part (Nobile & Tur-

risi, 2004: fig. 3; Schwarz & Gusenleitner, 2012:

figs. 7a-7d).

These strong differences between S. iosephi and

S. marginatus , affecting important features of head

and metasoma, clearly not running within intraspe-

cific variation, have been overlooked by Schwarz

& Gusenleitner (2012); moreover, while S. iosephi

belongs to the S. pinguiculus species-group, S.

marginatus belongs to the S. miniatus species-group

(Bogusch & Straka, 2012).

S. iosephi is deeply different from S. margina-

tus , and taking also into account the differences

with the other species of the same group, S. iosephi

Nobile et Turrisi, 2004 is valid species.

Sphecodes tomarchioi Nobile et Turrisi 2004

Sphecodes tomarchioi , Nobile & Turrisi, 2004. En-

tomofauna, 25 (8): 124 (Italia).

Sphecodes marginatus , Schwarz & Gusenleitner,

2012. Entomofauna, 33 (8): 76.

From the comparison between S. marginatus

and S. tomarchioi (the latter retained synonym of S.

marginatus by Schwarz & Gusenleitner, 2012) and

the descriptions provided by Hagens (1882) and

Meyer (1919), as well as the identification keys pro-

vided by Wamcke (1992) and Bogusch & Straka

(2012), it is possible to point out many remarkable

differences between the two considered species, the

most important being:

• S. tomarchioi has flagellomeres, excluding the

basal two, uniformly covered by sensilla on the ven-

tral surface (Haarfleck, etc.), without the character-

istic sequence of alternate notch and gibbous areas

(similarly to that presented by Schwarz & Gusen-

leitner, 2012: fig. 1); instead, ins', marginatus , each

flagellomere, excluding the basal three, bears a basal

white spot of sensilla alternate with an evident and

well protmded distal gibbous area, as also clearly

showed by Bogusch & Straka (2012: fig. 122);

• S. tomarchioi has irregular, coarse, deep and

very dense punctuation on frons and vertex (dis-

tance between punctures less than puncture diame-

ter); instead S. marcellinoi (= S. marginatus ?) has

frons and vertex bearing nearly regular, fine, super-

ficial and scattered punctuation (distance between

punctures about 2. Ox puncture diameter);

• S. tomarchioi has the first metasomal tergite

bearing fine, superficial and scattered punctuation

(distance between punctures 2. 0-3. Ox puncture di-

ameter); instead S. banaszaki (= S. marginatus ?)

has the first metasomal tergite with coarse, deep and

dense punctures (distance between punctures 1.0-

1.5x puncture diameter) (Nobile & Turrisi 2004;

Schwarz & Gusenleitner, 2012: fig. 4);

• the genital capsule of S. tomarchioi (Nobile

& Turrisi, 2004: fig. 7; Schwarz & Gusenleitner,

2012: figs. 1 la-1 Id) is significantly differentiated

from that of S. marginatus, due to the presence of a

long, slender and strongly curved toward inner distal

process of the sclerified part of gonostylus; instead,

570

Vittorio Nobile & Giuseppe FabrizioTurrisi

in S. marginatus this process is absent, with the di-

stal sclerified part of gonostylus stouter and straight

toward apex (Hagens, 1882: fig. 18; Wamcke, 1992:

fig. 32; Bogusch & Straka, 2012: figs. 165, 166).

These strong differences between S. tomarchioi

and S. marginatus , affecting important features of

head and metasoma, clearly not running within in-

traspecific variation, have been overlooked by

Schwarz & Gusenleitner (2012); moreover, while S.

tomarchioi belongs to the S. pinguiculus species-

group, S. marginatus belongs to the S. miniatus

species-group (Bogusch & Straka, 2012).

S. tomarchioi is deeply different from S. margina-

tus , and taking also into account the differences

with the other species of the same group, S. tomar-

chioi Nobile et Turrisi, 2004 is valid species.

CONCLUSIONS

Schwarz & Gusenleitner (2012) studied and

compared the type specimens of all the seven

species described by Nobile & Turrisi (2004), taking

into account the dimensions of the body, the features

of genital capsule and, in some cases, the distal part

of the antenna. Schwarz & Gusenleitner (2012) crit-

icized the bad preparation of the genital capsules, as

well as the drawings provided by Nobile & Turrisi

(2004); moreover, they retained some specimens,

namely those belonging to S. walteri and S. iosephi ,

too early collected, thus with cuticle of some parts,

useful for identification, too soft (the last flagellom-

eres and some detail of the genital capsule). With re-

gard to the drawings provided by Nobile & Turrisi

(2004), we reject the statement by Schwarz &

Gusenleitner (2012), because if a strict comparison

is made between these drawings and the photo-

graphs provided by Schwarz & Gusenleitner (2012)

and Bogusch & Straka (2012), it is possible to ascer-

tain not only the substantial adherence of the fea-

tures showed in the drawings with the photographs,

but also the usefulness of these features for identifi-

cation of the species, and clearly appears the para-

dox of having a species, namely S. marginatus

(sensu Schwarz & Gusenleitner, 2012), including

the additional six taxa described by Nobile & Turrisi

(2004), too variable and definitely without clear

diagnostic features, thus introducing an inacceptable

taxonomic treatment of the species concept within

S. pinguiculus species-group.

ACKNOWLEDGMENTS

We would to thank Schwarz M. and Gusen-

leitner F. for giving us the opportunity to point out

the validity of the species described in Nobile &

Turrisi (2004) especially through the usefulness of

the antenna morphology. Thanks also to Schmidt S.

(Zoologische Staatssammlung Miinchen, Germany)

for allowing the examination of the type material.

REFERENCES

Agren L. & Svensson B.G., 1982. Flagellar sensilla of

Sphecodes bees (Hymenoptera, Halictidae). Zoo-

logica Scripta, 11: 45-54.

Aurivillius C., 1903. Steklar. Hymenoptera. 1. Gaddsteklar.

Aculeata. Svensk Insektfauna, 13: 1-90.

Bliithgen P., 1923. Beitrage zur Systematik der Bienen-

gattung Sphecodes Latr. Deutsche Entomologische

Zeitschrift, 1923: 441-513.

Bogusch R & Straka J., 2012. Review and identification

of the cuckoo bees of central Europe (Hymenoptera:

Halictidae: Sphecodes). Zootaxa, 3311: 1-41.

Hagens D. von, 1882. Uber die mannlichen Genitalien

der Bienen-Gattung Sphecodes. Deutsche Entomolo-

gische Zeitschrift, 26: 209-229.

Jorgensen L., 1921. Bier. Danmark Fauna, 25: 1-11.

Meyer R., 1919. Apidae-Sphecodinae. Archiv fur

Naturgeschichte, 58: 79-242.

Mitchell T.B., 1960. Bees of the eastern United States. I.

Technical Bulletin of North Carolina Agriculture

Experimental Station, 141: 1-538.

Nobile V. & Turrisi G.F., 2004. Contribution to the

knowledge of Italian cleptoparasitic Bees. X. The

genus Sphecodes Latreille, “ pinguiculus Perez”

group, with description of new species (Hymenoptera,

Apoidea, Halictidae). Entomofauna, 25: 117-132.

Schwarz M. & Gusenleitner F., 2012. Zur Kenntnis

der von Nobile V. & G.F. Turrisi (2004) aus Italien

beschriebenen Sphecodes- Arten (Hymenoptera, Api-

dae). Entomofauna, 33: 73-80.

Thomson C.G., 1872. Hymenoptera Scandinaviae. Skan-

dinaviens Hymenoptera. Vol. 2, Berling, Lund, 286 pp.

Warncke K., 1992. Die westpalaearktischen Arten der

Bienengattung Sphecodes Latr. (Hymenoptera,

Apidae, Halictinae). Bericht dder Naturforschenden

Gesellschaft Augsburg, 52: 9-64.

Biodiversity Journal, 2013, 4 (4): 571-580

On the presence of Dreissena polymorpha Pallas, 1 77 1 and

Sinanodonta woodiana woodiana (Lea, 1 834) in Sicily (Bivalvia)

Maria Stella Colomba 1 *, Fabio Liberto 2 , Agatino Reitano 3 , Rosario Grasso 4 , Davide Di Franco 5 & Ignazio

Sparacio 6

'Universita di Urbino “Carlo Bo”, Dipartimento di Scienze Biomolecolari, via Maggetti 22 (loc. Sasso) - 61029 Urbino (PU), Italy

2 Strada Provinciale Cefalu - Gibilmanna, 93 - 90015 Cefalu, Palermo, Italy; e-mail: fabioliberto@alice.it

3 Via Gravina, 77 - 95030 Tremestieri Etneo, Catania, Italy; e-mail: tinohawk@yahoo.it

4 Dipartimento di Scienze Biologiche, Geologiche ed Ambientali, Universita degli Studi di Catania, Via Androne, 81 - 95124 Catania,

Italy; e-mail: rosagra@unict.it

5 Via S. Maria dei Monti 38/D - 95030, Gravina, Catania, Italy; email: davide_di_franco@hotmail.it

6 Via E. Notarbartolo 54 int. 13 - 90145 Palermo, Italy; e-mail: isparacio@inwind.it

* Corresponding author: mariastella.colomba@uniurb.it

ABSTRACT In this paper we report on the presence of two non-native and invasive species of bivalve

mollusks in Sicily: Sinanodonta woodiana woodiana (Lea, 1834) (Bivalvia Unionidae) na-

tive to East Asia and Dreissena polymorpha (Pallas, 1771) (Bivalvia Dreissenidae) native

to the Ponto-Caspian area, with a particular focus on how the presence of these alien species

can lead to imbalances in the delicate equilibrium of Sicilian freshwater invertebrates and,

above all, does threaten the existence of Unio elongatus gargottae Philippi, 1836 an island's

native bivalve suffering from strong rarefaction.

KEY WORDS invasive species; bivalve mollusks; aquatic biotopes; Sicily.

Received 28.11.2013; accepted 20.12.2013; printed 30.12.2013

INTRODUCTION

The problem of non-native and invasive species,

and particularly those who have settled in the Euro-

Mediterranean area, has become in recent years ex-

tremely interesting in relation to the large number

of species that have been reported.

All habitats and all groups of animals and

plants are affected by this phenomenon and the

impact on native ecosystems is almost always dev-

astating (Bogan, 1993; Lydeard et al., 2004).

The Sicilian aquatic biotopes are not an ex-

ception, with an increasing colonization by al-

lochthonous organisms observed both in historical

times that during the last few decades, for the fre-

shwater mollusks:

- Haitia acuta (Drapamaud, 1805): Cianfanelli et

al., 2007)

- Radix auricularia (Linnaeus, 1758): Liberto et al.,

2010

- Corbicula fluminea (Muller, 1774): Mienis, 1991

- Anodonta sp.: Zettler & Richard, 2003

- Potamopyrgus antipodarum (J.E. Gray, 1843):

Zettler & Richard, 2003; Bodon et al. 2005;

Cianfanelli et al., 2007

- Melanoides tuberculatus (O.L. Muller, 1774): Rei-

tano et al., 2007

- Helisoma duryi (Wetherby, 1879): Manganelli et

al. 1995; Reitano et al., 2007

- Ferrissia fragilis (Tryon, 1863): Marrone et al.,

2011.

572

Maria Stella Colomba et alii

Recently, some observations have indicated to

us the presence in Sicily of Sinanodonta woodiana

woodiana (Lea, 1834) (Bivalvia Unionidae) (per-

sonal observations; Distefano, 2013; Firrito, 2013)

one of the most invasive aquatic macroinvertebrate

species. Subsequent research carried out in the field

allowed to highlight the presence of another al-

lochthonous: Dreissena polymorpha (Pallas, 1771)

(Bivalvia Dreissenidae).

ACRONYMS. The materials used for this study

are deposited in the following private collections:

R. Grasso collection, Dipartimento di Scienze

Biologiche, Geologiche ed Ambientali, Universita

degli Studi di Catania, Italy (CGR); S. Giglio col-

lection, Cefalu, Italy (CGS); F. Liberto collection,

Cefalu, Italy (CL); A. Reitano collection, Tremestieri

Etneo, Italy (CR); I. Sparacio collection, Palermo,

Italy (CS).

MATERIALS AND METHODS

All specimens were collected by sight directly

in the field. Photographs were taken with a digital

camera. Voucher specimens were stored in collec-

tions indicated below. Toponyms (place-names)

are reported following the Portale Cartografico

Nazionale (PCN, http://www.pcn.minambiente.it

/PCN/), Map IGM 1:25000. Taxonomical refer-

ences are based on the checklist of "Fauna Eu-

ropaea", version 2.0 (Araujo 2009a, 2009b)

available at: http://www.faunaeur.org.

RESULTS

BIVALVIA

DREISSENIDAE

Dreissena polymorpha Pallas, 1771

Zebra mussel

Examined material. Italy, Sicily, Ragusa, Lago

di Santa Rosalia, 385 m, 36°59T9"N 14°46'47"E,

29.IX.20 12, 3 specimens with soft tissues (CR)

(Figs. 1, 2).

Description. Mytiliform bivalve (25-35 mm in

size) with relatively flattened ventral margins,

roundend dorsal margins and pointed umbo.

Brownish-yellowish in color with dark and light

coloured zigzag banding. Viewed from the inside,

the shell has a large septum to which the anterior

adductor and retractor muscles are attached (Killeen

et al., 2004) (Figs. 3-7).

Figure 1. Current geographic distribution of Dreissena polymorpha and Sinanodonta woodiana in Sicily. Dot: Ragusa,

Lago di Santa Rosalia.

On the presence of Dreissena polymopha and Sinanodonta woodiana woodiana in Sicily (Bivalvia)

573

Figure 2. Sicily, Ragusa: Lago di Santa Rosalia. Figures 3-7. Dreissena polymorpha from Sicily, Lago di Santa Rosalia.

574

Maria Stella Colomba et alii

Distribution and Biology. D. polymorpha is

native to the Ponto-Caspian Region of Eastern Eu-

rope. Its invasion history dates back to the late 18th

century in Russia and during the 19th century the

species spread out in most of inner water systems

of central and western Europe (Kinzelbach, 1992;

Strayer & Smith, 1993; Minchin et al., 2002); it has

been reported also in Spain, in the southern part of

the Ebro river (Araujo & Halcon, 2001; Araujo et

al., 2004) and in Northern Ireland (Millane et al.,

2008). In 1988, it first appeared in the Great Lakes

of North America and rapidly spread along the

Mississippi River (Strayer, 1991; Strayer et al.,

1994; Strayer & Smith, 1996; Strayer et al., 1998;

May et al., 2006). It was found for the first time in

Italy in 1969 in Lake Garda (Giusti & Oppini,

1973; Franchini, 1976). Subsequently, D. polymor-

pha has been reported for several lakes of Northern

Italy (Bianchi et al., 1974; Camusso et al., 2001;

Morpurgo & Thaler, 2002; Dalfreddo & Maiolini,

2003, Roncaglio & Borsani, 2005) and central

Italy (Spilinga et al., 2000; Bodon et al., 2005; Lori

& Cianfanelli, 2006).

D. polymorpha is characterized by a high fecun-

dity, ranging from about 30,000 to 40,000 eggs/

female per year. Veliger larvae lead a planktonic

life for, usually, 8-10 days, then settle and attach to

the bottom by byssal filaments. The average life

span is 3-6 years. Adults anchored to the substra-

tum can reach a high density, in the order of thou-

sands per square meter. Hence, filtrating capacity

of zebra mussels usually cause severe damages at

both local and system-levels, including changes in

species composition and density of native benthic

invertebrates; decreases in phytoplankton and zoo-

plankton densities, in chlorophyll concentration

and suspended matter; increase in water trans-

parency with a consequent growth of macrophytes

(Mackie et al., 1991; Therriault et al., 2013).

Remarks. In September 2013 a population of

D. polymorpha was found in the artificial Lake

Santa Rosalia (Ragusa, Sicily, Italy), which is the

first record of D. polymorpha in Sicily. Water dis-

charge due to introduction of fish was probably

the main vector that made it possible the spread

of D. polymorpha in the lake. As noted, the species

seems to be successfully established in the lake

basin with large aggregations (dmses) and it is likely

that a future further increase in its spreading rate

and diffusion will occur leading these organisms

to colonize the basin of Irminio river and other

basins. Indeed, Lake Santa Rosalia is connected

to Scicli by a 16 km aqueduct and two other pipe-

lines will be built to connect the Lake to Marina

di Ragusa and Santa Croce Camerina (Ragusa),

which might behave as highways for the diffusion

of the species.

According to the Global Invasive Species Data-

base of IUCN (2006) D. polymorpha is one of the

100 World's Worst Invasive Alien Species, and its

negative impacts on the ecosystem and especially

its economical damage are well known.

UNIONIDAE

Sinanodonta woodiana woodiana (Lea, 1834)

Chinese pond mussel

Examined material. Italy, Sicily, Ragusa, Lago

di Santa Rosalia, 36°59T3"N 14°46'50"E, 385 m,

21. VII. 20 12, 1 specimen (CGR); Italy, Sicily,

Ragusa, Lago di Santa Rosalia, 36°59T9"N

14°46'47"E, 385 m, 29.IX.2012, 4 specimens (CR);

idem, legit Reitano A., 3 specimens (CL); idem,

legit Reitano A., 3 specimens (CGS); idem, legit

Reitano A., 3 specimens (CS) (Figs. 8-10).

Description. From a morphological standpoint,

S. woodiana collected in Sicily show typical char-

acters of the species: wide shell (maximum length

ca. 300 mm), with deeply rounded ventral margin,

surface irregularly rippled to corrugated, umbonal

rugae prominent widely spaced, subconcentric or

slightly transverse ripples without prominent nod-

ules, posterior pedal retractor scar very narrow;

hinge without teeth. The size of some specimens

exceed 190 mm in length.

Distribution and Biology. S. woodiana is a

species of East Asia (Baba, 2000) that has recently

been reported as being invasive worldwide. The

expansion of S. woodiana’s range began in the

second half of the twentieth century. To date, S.

woodiana can be found in the Indonesian islands

(Djajasasmita, 1982), Central America (Watters,

1997), Europe (Hungary: Petro, 1984; Romania:

Sarkany-Kiss, 1986; France: Girardi & Ledoux,

1989; Slovakia: Kosel, 1995; Czech Republic:

Beran, 1997; Austria: Reischutz, 1998; Poland:

Bohme, 1998; Ukraine: Yurishinets & Komiushin,

2001; Germany: Gloer & Zettler, 2005; Serbia:

On the presence of Dreissena polymopha and Sinanodonta woodiana woodiana in Sicily (Bivalvia)

575

Figures 8-10. Sinanodonta woodiana woodiana from Sicily, Lago di Santa Rosalia.

576

Maria Stella Colomba et alii

Paunovic et al., 2006; Montenegro: Tomovic et

al., 2013; see also Sarkany-Kiss et al., 2000;

Kraszewski, 2007; Douda et al., 2012), and North

America (Benson, 2011).

It was first reported in Italy in 1996 (Manganelli

et al., 1998), and in about 15 years it formed colonies

in many Italian regions: Emilia-Romagna and

Latium (Manganelli et al., 1998; Fabbri & Landi,

1999; Lodde et al., 2005a; 2005b; Albano, 2006),

Tuscany and Veneto (Niero, 2003; Lori & Cian-

fanelli, 2006), Marche, Piedmont, Umbria and

Lumbardy (Solustri & Nardi, 2006), and Campania

(De Vico et al., 2007); in Sicily, Anodonta sp. was

recorded by Zettler & Richard (2003) and, recently,

the presence of S. woodiana was reported in some

web pages from Ragusa, Lago di Santa Rosalia

(Distefano, 2013; Firrito, 2013).

S. woodiana larvae, like all Unionidae (Casta-

gnolo et al., 1980; Aldridge & Mclvor, 2003), go

throught an obligatory parasitic stage (known as

glochidium); glochidia larvae parasitize, by cling-

ing with a kind of hook, the fins or gills of fish for

several days to weeks; afterwards they detach from

the host and fall to the bottom where mature and

start to conduct a free life. The great success of S.

woodiana has been attributed to the worldwide in-

troduction for commercial purposes of its sympatric

fish hosts (mainly carp species from East Asia)

(Watters, 1997). S. woodiana is a broad generalist,

and it may parasitize even on novel native hosts

(Kiss, 1995; Watters, 1997; Sarkany-Kiss et al.,

2000; Douda et al., 2012), in addition both juve-

niles and adults of S. woodiana can successfully

cope with a wide range of environmental conditions.

Thermal conditions, water flow and substrate

characteristics mostly determine the distribution

and density of S. woodiana (Kraszewski &

Zdanowski, 2007). According to Demayo et al.

(2012), this species prefers habitats with higher

temperatures (the optimal thermal conditions vary

within 10 and 35°C).

Remarks. The presence of S. woodiana in Lake

Santa Rosalia, an artificial basin for irrigation built

in 1980 on Irminio river (Ragusa, South-eastern

Sicily), may have been caused by the introduction

of several species of fish from farms to promote

sport fishing. The discovery in 2012 of specimens

larger than 190 mm, aged more than 8 years, allows

us to hypothesize that the entrance of S. woodiana

in the lake took place at least 10 years ago.

According to the Global Invasive Species Data-

base of IUCN (2006), S. woodiana is one of the 100

World's Worst Invasive Alien Species.

DISCUSSION AND CONCLUSIONS

Generally, the introduction of alien species in a

given territory produces nearly always negative

effects on the presence and population dynamics of

native species. Particularly, the filtrating capacity

of D. polymorpha and S. woodiana may affect or-

ganic matter pathways within the sediment and the

water column with serious consequences for the

surrounding ecosystem (Vaughn & Hakenkamp,

2001; Kraszewski & Zdanowski, 2007).

In Sicily, the increase in population density of

these species and the diffusion to other lakes and

rivers can be expected to produce a negative impact

on native sedentary benthic invertebrates. For

example, native unionid mussels may be threatened

by competition for food, space and hosts, as has al-

ready been observed in other Italian regions (Fabbri

& Landi, 1999; Niero, 2003).

In Sicily, the presence of the genus Unio Philipson,

1788, the only major bivalve native to the island is

particularly critical. According to Bodon et al.

(2005) and Prie et al. (2012) two species of Unio

are present in Italy, Unio pictorum in the Isonzo

Basin and Unio elongatus (Pfeiffer, 1825) in the rest

of the Italian territory, including Sicily (the latter

previously reported also as U. mancus Lamarck,

1819). Unio populations from Sicily are considered

by Zettler & Richard (2003) and Araujo (2009a) as

belonging to the endemic subspecies U. elongatus

gargottae Philippi, 1836.

In the second half of the nineteenth century,

many taxa were described or reported for the genus

Unio in major rivers and lakes of Sicily (see for

example: Philippi, 1836- 1844; Benoit, 1875, 1882;

Monterosato, 1896; Castagnolo et al., 2002).

During the twentieth century the records of Unio

in Sicily were rare (Naselli-Flores et al., 1996; per-

sonal observations), thus indicating a dramatic de-

cline in the distribution and abundance of Unio

populations mainly caused by pollution and im-

proper wetland management (drainage, channel al-

terations, water intake, cementification of

riverbanks, or total destruction of habitats). How-

ever, some observations in the field have allowed

On the presence of Dreissena polymopha and Sinanodonta woodiana woodiana in Sicily (Bivalvia)

577

to testify the presence of U. elongatus gargottae in

a few basins of western Sicily with small but stable

populations monitored successfully since 2003 to

2007. Now, this survival strategy of the native Unio

could be threatened by the presence and strong in-

vasiveness of S. woodiana.

REFERENCES

Albano P.G., 2006. Ritrovamenti presso la spiaggia di

Palo Laziale (Roma). Notiziario S.I.M., 24: 21-22.

Aldridge D.C. & Mclvor A.L., 2003. Gill evacuation and

release of glochidia by Unio pictorum and Unio

tumidus (Bivalvia: Unionidae) under thermal and hy-

poxic stress. Journal of Molluscan Studies, 69: 55-

59.

Araujo R., 2009a. Fauna Europaea: Bivalvia, Unionidae,

Sinanodonta. Fauna Europaea version 2.0 Available

at: http://www.faunaeur.org (Accessed 30 Sept.

2013).

Araujo R., 2009b. Fauna Europaea: Bivalvia, Dreis-

senidae, Dreissena. Fauna Europaea version 2.0

Available at: http://www.faunaeur.org (Accessed 30

Sept. 2013).

Araujo R., Valladolid M. & Gomez I., 2004. Life cycle

and density of a newcomer population of zebra mus-

sels in the Ebro River, Spain. In: van Der Velde G.,

Rajagopal S. & Bij de Vaate A. (Eds) The Zebra Mus-

sel in Europe, Backhuys Publishers, Leiden, pp. 1 83—

189.

Araujo R. & Halcon R.M.A., 2001 . El mejillon cebra en el

Ebro: un grave caso de riesgo ambiental en Aragon.

Naturaleza Aragonesa, 8: 39-46.

Baba K., 2000. An area-analytical zoogeographical clas-

sification of Paleartic Unionaceae species. Bollettino

Malacologico, 36: 133-140.

Benoit L., 1875. Catalogo delle conchiglie terrestri e flu-

viatili della Sicilia e delle Isole circostanti. Bullettino

della Societa Malacologica italiana, 1: 129-163.

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.

Benson A. J., 2011. Sinanodonta woodiana. USGS nonin-

digenous aquatic species database, http://nas.er.

usgs.gov/queries/FactSheetaspx?speciesID=2824

(Accessed 30 Sept. 2013).

Beran L., 1997. First record of Sinanodonta woodiana

(Mollusca, Bivalvia) in the Czech Republic. Acta

Societatis Zoologicae Bohemicae, Praha, 61: 1-2.

Bianchi I., Girod A. & Mriani M., 1974. Densita, strut-

ture di popolazione e distribuzione di Dreissena po-

lymorpha nel bacino idrografico del Benaco. Archiv

fur Molluskenkunde, 104: 97-105.

Bohme M., 1998. Ein neuer Fundort der Chinesischen

Teichmuschel ( Sinanodonta woodiana ) in Mitteleu-

ropa. Heldia, 2: 166.

Bodon M, Cianfanelli S., Manganelli G, Pezzoli E. &

Giusti F., 2005. Molluschi Bivalvi. In: Ruffo S. &

Stoch F. (Eds), Checklist e distribuzione della fauna

italiana. Memorie del Museo Civico di Storia Natu-

rale di Verona, 2. serie, Sezione Scienze della Vita

16: 83-84 + CD.

Bogan A.E., 1993. Freshwater bivalve extinctions (Mol-

lusca: Unionoida): a search for causes. American

Zoologist, 33: 599-609.

Camusso M., Balestrini R. & Binelli A., 2001. Use of

zebra mussel ( Dreissena polymorpha) to assess trace

metal contamination in the largest Italian subalpine

lakes. Chemosphere, 44: 263-270.

Castagnolo L., Franchini D. & Giusti F., 1980. Bivalvi.

Guide per il riconoscimento delle specie animali delle

acque interne italiane. C.N.R., Verona, 64 pp.

Castagnolo L., Nagel K.O. & Cencetti E., 2002. Gliunion-

idi italiani della “Collezione Paulucci” conservati nel

Museo Zoologico “La Specola” di Firenze. Atti

della Societa toscana di Scienze naturali, Memorie,

Serie B, 109: 29-79.

Cianfanelli S., Lori E. & Bodon M., 2007. Non-indige-

nous freshwater molluscs and their distribution in

Italy. In: Gherardi F. (Ed.), Biological invader in in-

land waters: profiles, distribution, and threats. Sprin-

ger, Dordrecht. Chapter 5, pp. 103-121.

Dalfreddo C. & Maiolini B., 2003. II popolamento

malacologico di alcuni laghi trentini a confronto 70

anni dopo. Studi Trentini di Scienze Naturali, Acta

Biologica, 80: 175-177.

Demayo C.G., Cabacaba K.M.C. & Torres M.A.J., 2012.

Shell shapes of the Chinese pond mussel Sinan-

odonta woodiana (Lea, 1834) from Lawis Stream in

Iligan City and Lake Lanao in Mindanao, Philip-

pines. Advances in Environmental Biology, 6: 1468-

1473.

De Vico G., Maio N. & Castagnolo L., 2007. Prima seg-

nalazione di Anodonta ( Sinanodonta ) woodiana (Lea,

1834) (Mollusca: Bivalvia: Unionidae) per il sud

Italia. Notiziario S.I.M., 25: 23-25.

Distefano S., 2013. Le conchiglie alia diga di Santa Ros-

alia. ragusanes.com [4.X.2013]: http://www.ragusa-

news.com/articolo/34800/le-conchiglie-alla-diga-di-

santa-rosalia (Accessed 20 Sept. 2013).

Djajasasmita M., 1982. The occurrence of Anodonta

woodiana Lea, 1837 in Indonesia (Pelecypoda, Union-

idae). The Veliger, 25: 175.

Douda K., Vrtilek M., Slavik O. & Reichard M., 2012.

The role of host specificity in explaining the invasion

578

Maria Stella Colomba et alii

success of the freshwater mussel Anodonta woodiana

in Europe. Biological Invasions, 14: 127-137.

Fabbri R. & Landi D., 1999. Nuove segnalazioni di

molluschi, crostacei e pesci esotici in Emilia Roma-

gna e prima segnalazionedi Corbicula fulminea (O.F.

Muller, 1774) in Italia (Mollusca Bivalvia, Crustacea

Decapoda, Osteichthyes Cypriniformes). Quaderno

di Studi e Notizie di Storia Naturale della Romagna,

12: 9-20.

Firrito A., 2013. www.facebook.com/groups/faunasicil-

iana/permalink/466947273420625/(Accessed 30

Oct. 2013).

Franchini D.A., 1976. Sostituzione degli insediamenti

malacologici autoctoni ad opera della Dreissena

polymorpha (Pallas) in una stazione del Benaco

(Torri del Benaco, VR). Favori del Simposio sui Mol-

luschi terrestri e dulcicoli delFItalia Settentrionale,

Mantova, 10-11/5/1975. Favori della Societa Mala-

cologica Italiana, 13: 89-95.

Girardi H. & J.C. Fedoux, 1989. Presence d’ Anodonta

woodiana (Fea) en France (Mollusques, Famelli-

branches, Unionidae). Bulletin Mensuel de la Societe

Finneenne de Fyon, 58: 286-290.

Giusti F. & Oppi E., 1973. Dreissena polymorpha

(Pallas) nuovamente in Italia. (Bivalvia, Dreis-

senidae). Memorie del Museo Civico di Storia Natu-

rale di Verona, 20: 45-49.

Gloer P. & Zettler M.F., 2005. Kommentierte Artenliste

der SiiBwassermollusken Deutschlands. Malakolo-

gische Abhandlungen, 23: 3-23.

Killeen I., Aldridge D. & Oliver. G., 2004. Freshwater

Bivalves of Britain and Ireland. Preston, Montford.

Field studies council Publication, 1 19 pp.

Kinzelbach R., 1992. The main features of the phylogeny

and dispersal of the Zebra Mussel Dreissena Poly-

morpha. In: Neuman D. & Jenner HA (Eds), The

zebra mussel Dreissena polymorpha, Ecology, Bio-

logical Monitoring and First Applications in Water

Quality management. Fimnologie Aktuell, 4: 5-17.

Kiss A., 1995. The propagation, growth and biomass of

the Chinese huge mussel ( Anodonta woodiana wood-

iana Fea, 1834) in Hungary. Dissertation, University

of Agricultural Sciences of Godollo, 38 pp.

Kosel V., 1995. The first record of Anodonta woodiana

(Mollusca, Bivalvia) in Slovakia. Acta Zoologica

Universitatis Comenianae, Bratislava, 39: 3-7.

Kraszewski A., 2007. The continuing expansion of Sinan-

odonta woodiana (Fea, 1834) (Bivalvia: Unionidae)

in Poland and Europe. Folia Malacologica, 15: 65-

69.

Kraszewski A. & Zdanowski B., 2007. Sinanodonta

woodiana (Fea, 1834) (Mollusca) a new mussel

species in Poland: occurrence and habitat preferences

in a heated lake system. Polish Journal of Ecology,

55: 337-356.

IUCN, 2006. Global Invasive Species Database.

www.issg.org (Accessed 30 Sept. 2013).

Fiberto 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, 136 pp.

Fodde A., Palmerini E. & Castagnolo F., 2005a. Repro-

ductive strategy of an Anodonta woodiana (Fea,

1834) (Mollusca, Bivalvia, Unionidae) population

from a Modena canal, a non-indigenous species in

Italy. INWAT: Biological invasionsin inland waters.

May 5-7, 2005, Florence: 46-47.

Fodde A., Palmerini E., Sala F. & Castagnolo F., 2005b.

Anodonta woodiana (Fea, 1834) (Mollusca: Bivalvia:

Unionidae), a non-indigenous species wide-spread in

Italy: comparison of the biological cycle in native

countries (Far East) and in Italy (Modena canals). IV

International Congress of the European Malacologi-

cal Societies. October 10-14 2005, Naples (Italy).

Notiziario S.I.M., 23: 70.

Fori E. & Cianfanelli S., 2006. New records of Dreissena

polymorpha (Pallas, 1771) (Mollusca: Bivalvia:

Dreissenidae) from Central Italy. Aquatic Invasions,

1:281-283.

Fydeard C., Cowie R.H., Ponder W.F., Bogan A.E.,

Bouchet P., Clark S.A., Cummings K.S., Frest T.J.,

Gargominy O., Herbert D.G., Hershler R., Perez

K.E., Roth B., Seddon M., Strong E.E. & Thompson

F.G., 2004. The global decline of nonmarine mol-

lusks. Bioscience, 54: 321-330.

Mackie G.F., Gibbons W.N., Muncaster B.W. & Gray

I.M., 1991. The zebra mussel Dreissena polymor-

pha a synthesis of European experiences and a pre-

view for north America. Queen’s Printer for Ontario,

135 pp.

May G.E., Gelembiuk G.W., Panov V.E., Orlova M.I. &

Eunmi Fee C.E., 2006. Molecular ecology of zebra

mussel invasions. Molecular Ecology, 15: 1 021—

1031.

Manganelli G., Bodon M., Favilli F., Giusti F., 1995.

Gastropoda Pulmonata. In: Minelli A., Ruffo S. Fa

Posta S. (Eds.), Checklist delle specie della fauna

italiana, 16. Calderini, Bologna, 60 pp.

Manganelli G., Bodon M., Favilli F., Castagnolo F. &

Giusti F., 1998. Checklist delle specie della fauna

d’ Italia, molluschi terrestri e d’acqua dolce. Errata

ed addenda, 1. Bollettino Malacologico, 33: 151—

156.

Marrone F., To Brutto S. & Arculeo M., 2011. Cryptic

invasion in Southern Europe: The case of Ferrissia

fragilis (Pulmonata: Ancylidae) Mediterranean popu-

lations.Versita Biologia, 66: 484-490.

Mienis H.K., 1991. Some remarks concerning asiatic

clams invading Europe with a note on sample of Cor-

On the presence of Dreissena polymopha and Sinanodonta woodiana woodiana in Sicily (Bivalvia)

579

bicula fluminea (Muller, 1774) from Trapani, Sicily.

Notiziario S.I.M., 9: 137-139.

Millane M., Kelly-Quinn M. & Champ T., 2008. Impact

of the zebra mussel invasion on the ecological in-

tegrity of Lough Sheelin, Ireland: distribution popu-

lation characteristics and water quality changes in the

lake. Aquatic Invasions, 3: 271-281.

Minchin D., Lucy F. & Sullivan M., 2002. Zebra mussel:

impacts and spread. In: Leppakoski et al. (Eds), In-

vasive aquatic species of Europe. Springer

Science+Business Media Dordrecht, 135-146.

Monterosato T. Di Maria di, 1896. Note intorno alle

Najadi siciliane. II Naturalista Siciliano, 15: 6-20.

Morpurgo M. & Thaler B., 2002. Ritrovamento di Dreis-

sena polymorpha (Pallas) (Mollusca, Bivalvia) nel

Lago Grande di Monticolo (Alto Adige, Italia).

Gredleriana, 2: 219-222.

Naselli-Flores L., Zava B. & Chemello R. 1996. Mol-

luschi delle Acque interne Siciliane. Segnalazione di

Unio elongatus C. Pfeiffer, 1825 (Bivalvia: Union-

idae). Bollettino Malacologico, 31: 163-168.

Niero I., 2003. Sulla presenza in Veneto e centra Italia di

Anodonta woodiana woodiana (Lea, 1834) (Mol-

lusca, Bivalvia). Bollettino del Museo civico di Storia

naturale di Venezia, 54: 29-33.

Paunovic M., Csany B., Simic V., Stojanovic B. & Cakic

P., 2006. Distribution of Anodonta ( Sinanodonta )

woodiana (Lea, 1834) in inland waters of Serbia.

Aquatic Invasions, 1: 154-160.

Petra E., 1984. Occurrence of Anodonta woodiana (Lea,

1834) (Bivalvia: Unionacea) in Hungary. Allattanu

Kozlemenyek, 71: 181-191.

Philippi R.A., 1836-1844. Enumeratio molluscoram sicilia

e cum viventium in tellure tertiaria fossilium, quae in

itinere suo observavit. Berolini, XIV+267 pp., 12 tav.

(1836); Halis, Saxonum, IV+303 pp., 16 tav. (1844).

Prie V., Puillandre N. & Bouchet P., 2012. Bad taxonomy

can kill: molecular reevaluation of Unio mancus

Lamarck, 1819 (Bivalvia: Unionidae) and its accepted

subspecies. Knowledge and Management of Aquatic

Ecosystems, 405: 1-18.

Reischutz P.L., 1998. Vorschlag fur deutsche Namen der

in Osterreich nachgewiesenen Schnecken- und

Muschelarten. Nachrichtenblatt der Ersten Vorarlber-

ger Malakologischen Gesellschaft, 6: 3 1—44.

Reitano A., Liberto F. & Sparacio I., 2007. Nuovi dati su

Molluschi terrestri e dulciacquicoli di Sicilia. 1° Con-

tribute (Gastropoda Prosobranchia Neotaenioglossa;

Gastropoda Pulmonata Basommatophora, Stylom-

matophora). II Naturalista siciliano, 31:31 1-330.

Roncaglio P. & Borsani G., 2005. Analisi della strattura

di popolazione del mollusco bivalve Dreissena poly-

morpha (PALLAS, 1771) nel Sebino (Lombardia,

Italia settentrionale). Natura Bresciana, Annali del

Museo civico di Scienze Naturali di Brescia, 34:

49-53.

Sarkany-Kiss A., 198 6 . Anodonta woodiana (Lea, 1834)

a new species in Romania (Bivalvia: Unionacea).

Travaux du Museum d’Histoire Naturelle “Grigore

Antipa”, 28: 15-17.

Sarkany-Kiss A., Sirbu I. & Hulea O., 2000. Expansion

of the adventive species Anodonta woodiana (Lea,

1834) (Mollusca, Bivalvia, Unionoidea) in central

and eastern Europe. Acta Oecologica Carpatica, 7:

49-57.

Solustri C. & Nardi G., 2006. Una nuova stazione di

Anodonta woodiana woodiana (Lea, 1834) nell’

Italia centrale) (Mollusca, Bivalvia, Unionidae).

Quademo di Studi e Notizie di Storia Naturale della

Romagna, 23: 1-8.

Spilinga C., Chiappafreddo U. & Pirisinu Q., 2000.

Dreissena polymorpha (Pallas) al Lago Trasimeno.

Rivista di idrobiologia, 39: 145-152.

Strayer D.L., 1991. Projected distribution of the zebra

mussel, Dreissena polymorpha, in North America.

Canadian Journal of Fisheries and Aquatic Sciences,

48: 1389-1395.

Strayer D.L. & Smith L.C., 1993. Distribution of the

zebra mussel ( Dreissena polymorpha ) in estuaries

and brackish water. In: Nalepa T. F. & Schloester D.

W. (Eds), Zebra Mussels, Biology, Impacts, and Con-

trol. Lewis Publishers, Boca Raton, Florida, pp.715-

727.

Strayer D.L. & Smith L.C., 1996. Relationships between

zebra mussels ( Dreissena polymorpha ) and unionid

clams during the early stage of the zebra mussel

invasion of the Hudson River. Freshwater Biology,

36: 771-779.

Strayer D.L., Hunter D.C., Smith L.C. & Borg C.K.,

1994. Distribution, abundance, and roles of fresh-

water clams (Bivalvia, Unionidae) in the freshwater

tidal Hudson River. Freshwater Biology, 31: 239-

248.

Strayer D.L., Smith L.C. & Hunter D.C., 1998. Effects

of the zebra mussel ( Dreissena polymorpha ) invasion

on the macrobenthos of the freshwater tidal Hudson

River. Canadian Journal of Zoology, 76: 419-425.

Therriault T.W., Weise A.M., Higgins S.N., Guo Y. &

Duhaime J., 2013. Risk assessment for three dreisse-

nid mussels ( Dreissena polymorpha, Dreissena ros-

trifonnis bugensis, and Mytilopsis leucophaeata) in

Canadian Freshwater Ecosystems. DFO Canadian

Science Advisory Secretariat Research Document

2012/174. V+88 pp.

Tomovic J., Zoric K., Simic V., Kostic M., Kljajic Z.,

Lajtner J. & Paunovic M., 2013. The first record of

the Chinese pond mussel Sinanodonta woodiana

(Lea, 1834) in Montenegro. Archives of Biological

Sciences, 65: 1525-1531.

580

Maria Stella Colomba et alii

Yurishinets V.I. & Korniushin A.V., 2001. The new

species in the fauna of Ukraine Sinanodonta wood-

iana (Bivalvia, Unionidae), its diagnostics and pos-

sible ways of introduction. Vestnik Zoologii, 35:

79-84.

Vaughn C.C. & Hakenkamp C.C., 2001. The functional

role of burrowing bivalves in freshwater ecosystems.

Freshwater Biology, 46: 1431-1446.

Watters G.T., 1997. A synthesis and review of the ex-

panding range of the Asian freshwater mussel An-

odonta woodiana (Lea, 1834) (Bivalvia: Unionidae).

The Veliger, 40: 152-156.

Zettler M.L. & Richard D., 2003. Bemerkungen iiber

SuBwassermollusken Siziliens unter besonderer

Berucksichtigung von Theodoxiis meridionalis (Philippi,

1836). Malakologische Abhandlungen, 21: 29-38.

581

NEW TAXA PUBLISHED IN THE BIODIVERSITY JOURNAL 4, 2013

Elamena soonthronkitti n. sp. (Decapoda Hymenosomatidae) lisd

Kulabtong, Kunlapapuk et Sottiyothin, 2013, 4: 130

lisd:zoobank.org:act:0A58AF90-F7CF-4DEE-9174-0B7FCE5094A6

Aporrhais etrusca n. sp. (Gastropoda Aporrhaidae)

Brunetti et Forli, 2013, 4: 195

lisd:zoobank.org:act:E3D7705C-49 12-408 1-A1F5-9EC9C3226762

Carabus (Carabus) szeli changbaicus n. ssp. (Coleoptera Carabidae Carabinae)

Rapuzzi et Li, 2013, 4: 217

lisd:zoobank.org/2312226E-98A3-44AC-A0C3-CB5BCB7E9891

Amphidromus (Syndromus) iunior n. sp. (Gastropoda Pulmonata Camaenidae)

Cilia, 2013 , 4: 264

lisd-um:lsid:zoobank.org:act:7BE6FD46-B4F9-458B-B341-03C14204248E

Madrasostes bartolozzii n. sp. (Coleoptera Scarabaeoidea Hybosoridae Ceratocanthinae)

Ballerio, 2013, 4: 380

lisd:zoobank.org:act:B9069EFD-2873-40BE-B6A9-0B67CFC74F92

Agrilus (Nigritius) massaronei n. sp. (Coleoptera Buprestidae)

Curletti, 2013, 4: 407

lisd:zoobank.org:act:01B9C526-FF3D-41B9-8B27-6E50876697BC

Agrilus (Nigritius) ivorianus n. sp. (Coleoptera Buprestidae)

Curletti, 2013, 4: 408

lisd:zoobank.org:act:6B72EE2E-297B-4355-A8D5-F5B4F46211C4

Agrilus (Nigritius) yangambii n. sp. (Coleoptera Buprestidae)

Curletti, 2013, 4: 408

lisd:zoobank.org:act:7983DlF0-A734-4E4D-ABAD-387010F6EDDC

Agrilus (Nigritius) bancoi n. sp. (Coleoptera Buprestidae)

Curletti, 2013, 4: 410

lisd:zoobank.org:act:BlB3D637-4BE6-4AF6-92C4-4B4A8E9650BA

Agrilus (Agrilus) pirilampo n. sp. (Coleoptera Buprestidae)

Curletti, Ribeiro et Migliore, 2013, 4: 445

lisd:zoobank.org:act:AE9470AB-8175-487C-B12D-6276B93D9F02

Agrilus (Agrilus) disorientatus n. sp. (Coleoptera Buprestidae)

Curletti, Ribeiro et Migliore, 2013, 4: 446

lisd:zoobank.org:act:76CD7400-D839-4124-A6D4-81D8ECFCE924

Agrilus (Agrilus) coal n. sp. (Coleoptera Buprestidae)

Curletti, Ribeiro et Migliore, 2013, 4: 446

lisd:zoobank.org:act:2F76AFE3-A745-4318-B96C-78845E234B14