The genus Alcides Hlibner, [ 1 822 1 (Lepidoptera Uraniidae). The Family Uraniidae contains some of the most beautiful moths known to

lepidoptcrists. It includes four subfamilies (Auzcinac, Epipleminae, Microniinae and Uraniinac), some ninety genera and about seven hundred species.

This is a tropical family which is found in Africa, the Americas, and the Indo-Australian region. Diurnal and nocturnal species are found within this

family and the day-flying ones are usually vibrantly colored and covered with iridescent scales and tails which have led many to mistake them for

butterflies. Several species are known to be migratory, but very little is known about the details of these migrations. One of these species was once

considered by the French lepidopterist Jean Baptiste Antoine Dechauffor de Boisduval (1799-1879) as the most beautiful of the creation.” The

subfamily Uraniinae contains seven genera. Three of them include species of moderate size and with resemblance to those in the Microniinae, while the

other four include large and strongly tailed species. Their hosts include plants belonging to the Euphorbiaceae and it appears that some of these moths

are models for certain species of some butterfly families. Among the Uraniinae we find the swallow tail moths of the genus Alcides Hubner, which are

brightly colored and probably distasteful to birds and other potential predators. Alcides is distributed in the Indo-Australian region and contains ten

species. Most fly by day but some have also been reported with crepuscular activity. Their larvae feed on leaves of plants in the genera Endospermum

and Omphalea which carry certain poisons that appear to protect the caterpillars, and eventually the adults, from predation. The caterpillars pupate in a

sparse cocoon in crevices or between dead leaves. The imagoes are characterized by front wings of triangular shape with a large, curved band that starts

in the Costal margin, crosses the discal area, and ends in the anal angle. A second, parallel, thinner band, is also observed in the apical angle of the wing

and starts also in the costal margin (without touching it), and goes to the anal angle, but vanishes before reaching it. The colors in both bands are always

iridescent green or blue. The hind wings have a wide band in the discal area, and when the wings are spread it looks like a continuation of the large band

in the forewing. This band also presents iridescent colors. All species in the genus present short tails in the hind wings. The current systematic

knowledge and taxonomy of the genus is incomplete and the status of some taxa remains uncertain. Even though ten species have been described so far,

the systematics of the group is still somehow uncertain. They are knowm from localities in Indonesia, Papua New Guinea, and the Moluccas, New'

Hebrides and Salomon Islands. However, the only exception to this distribution, the species Alcides met a unis (Hopffer) [=A. zodiaca (Butler)], is

endemic of Queensland, Northern Australia. The illustrated specimen is the Holotype of Alcides privitera Vincigucrra, This is a beautiful species

somehow similar to A. metaums but has been found in Sarong, Irian Jaya, Indonesia.

Jorge M. Gonzalez, California State University, Fresno, Department of Plant Sciences, Fresno, CA 93740-8033 (Research Associate, McGuire Center

for Lepidoptera & Biodiversity), USA; e-mail: gonzalez.jorge.m@gmail.com

Biodiversity Journal, 2013, 4 (2): 263-268

Description of a new species of Amphidromus Albers, 1 850

from Sumba, Indonesia (Gastropoda Pulmonata Camaenidae)

David P. Cilia

29, Triq il-Palazz 1-Ahmar, Santa Venera, Malta; e-mail: (dpcilia@gmail.com)

ABSTRACT The camaenid Amphidromus ( Syndromus ) iunior n. sp. from an isolated forest in the east

of Sumba island in the Indonesian archipelago is described. Its closest named relative is

Amphidromus (, Syndromus ) abbasi Chan et Tan, 2010, and some conchological features are

common for both species. However, the new species is smaller, with consistent differences

in shell thickness, pattern and pigmentation.

KEY WORDS Camaenidae; Amphidromus iunior n. sp.; Sumba; Indonesia.

Received 16.01.2013; accepted 14.04.2013; printed 30.06.2013

INTRODUCTION

The polyphyletic family Camaenidae Pilsbry,

1895 (Scott, 1996) is distributed in three distinct

clades (Wade et al., 2007), geographically separated

mainly by the Pacific Ocean. Synapomorphies

amongst the members of the different clades are ab-

sent, and the family is characterized by the absence of

vaginal accessory organs found in other members of

the otherwise monophyletic Helicoidea Rafinesque,

1815 (Wade at al., 2006). According to molecular

techniques employed by Wade et al. (2007), the Aus-

tralasian fraction of this family was found to be syn-

onymous with Bradybaenidae Pilsbry, 1934.

Amongst the most speciose genera in this Aus-

tralasian clade is the arboreal Amphidromus Albers,

1850. Extending over a wide geographical area

from India to Australia (Laidlaw & Solem, 1961;

Solem, 1983), it is divided into two or three sub-

genera historically differentiated mostly through

conchological characteristics. Of these, the sub-

genus Syndromus Pilsbry, 1900 (comprising the

newly described species discussed herein) consists

of animals with small shells (20-40 mm high, 10-

25 mm wide) and a short epiphallic caecum, which

according to Lok & Tan (2008) includes one dex-

tral, one enantiomorphic, and 44 sinistral taxa. New

species have recently been assigned to this subgenus

by Sevems (2006), Dharma (2007) and Chan & Tan

(2010), though Scucharit & Panha (2006) shed doubt

on the taxon’s monophyly. A new species of Syndro-

mus collected on east Sumba is described herein.

ABBREVIATIONS. Depositories: BP = collec-

tion of Bama Pall-Gergely, Mosonmagyarovar, Hun-

gary; DC = collection of David P. Cilia, Santa

Venera, Malta; FMNH = Field Museum of Natural

History, Chicago, Illinois; HUJ = Hebrew University

of Jerusalem, Israel; JA = collection of John Abbas,

Jakarta, Indonesia; MNHN = Museum National

d'histoire Naturelle, Paris, France; NHMUK = Natu-

ral History Museum, Fondon, United Kingdom;

NMNH = National Museum of Natural History,

Mdina, Malta; TAU = Zoological Department of Tel

Aviv University, Israel. Morphology and anatomy:

D = diameter; H = height; H/D = height to diameter

ratio; min. = minimum value; med. = median value;

max. = maximum value.

MATERIALS AND METHODS

The mean value of two readings for height (in-

264

David P. Cilia

eluding the reflected peristome), width (including the

reflected peristome) and oblique apertural height for

a random selection of 2 1 adult specimens was taken

using a dial caliper of a resolution of 50 jam. Results

were rounded off to the nearest 0. 1mm. Whorls were

counted, including the nucleus. Statistical data was

formulated and used together with peculiar morpho-

logical characteristics to differentiate the new species

from similar ones, most significant of which is the

geographically and phylogenetically close A abbasi

Chan et Tan, 2010. Topotypical adult shells of the lat-

ter were similarly measured and morphometrically

compared to those of the new species. Systematics

in the present paper follow Bouchet & Rocroi (2005).

SYSTEMATICS

Superfamily Helicoidea Rafmesque, 1815

Family Camaenidae Pilsbry, 1895

Subfamily Camaeninae Pilsbry, 1895

Genus Amphidromus Albers, 1850

Subgenus Syndromus Pilsbry, 1900

Type species Helix contraria Muller, 1774

Amphidromus ( Syndromus ) iunior n. sp.

Examined material. Holotype: small forest

close to Mangili village, east part of Sumba Island,

East Nusa Tenggara, Indonesia (-10°05'32"N,

120°42'08"E), at an altitude of about 560 m above

sea level, leg. JA: MNHN 23265. Paratypes (20

specimens): same data as holotype: DC RG217 (3);

HUJ 53490 (2); JA unreg. (3); MNHN 23266 (2);

FMNH 328102 (2); TAU 75175 (1); NHMUK

20120044 (3); NMNH unreg. (2); BP unreg. (2).

Description of holotype (Figs. 1-6). Sinistral,

conical, thin, smooth, glossy and translucent shell,

14 mm wide and 28 mm tall. Dull yellow base

colour intensifying towards the ultimate whorl, with

brown markings in the form of two well-defined

subperipheral spiral striae, faint subordinate ones,

and blurred spaced vertical columns featuring a re-

duction in their colour intensity and frequency to-

wards the final whorl. These columns are

interrupted halfway through by a perpendicularly

crossing spiral band lacking in pigment, another of

which is present just beneath the suture. Nucleus

dark purple-brown, its colour descending and fad-

ing out along the apical whorl just above the suture,

appearing in top view as a dark spiral over the pale

base. The number of the moderately convex whorls

is between six and seven, the final of which is

smoothly rounded. Aperture is vertically elongated

and teardrop-shaped, its height constituting about

40% of the shell’s total. Peristome fragile and ex-

panded, only thickened and reflected at its columel-

lar side. The reflected segment is not completely

fused to the base of the ultimate whorl, in most

cases leaving an obliquely oriented pinhole um-

bilicus. Internal aspect glossy, with the brown mark-

ings showing through.

Variability. The width varies from 1 3 to 1 6 mm

and the height from 23 to 34 mm (Figs. 8-22; Table

1). Whereas the subperipheral spiral striae are al-

ways present, very brief interruptions occasionally

occur. Additional spiral striae may unusually take

the form of parallel brown dashes that also coincide

in a perpendicular direction, forming columns.

Etymology. The species is named for the

youngest son of John Abbas, who encountered this

hitherto unrecorded species during one of his

expeditions. Another species of Amphidromus

named for John, A. (S.) abbasi, is closely related to

this species, but is significantly larger, amongst

other differences. The meaning of iunior therefore

takes on another meaning, referring to the relative

size of the snail (iunior is the comparative form of

iuvenis, meaning young in Latin).

Distribution. Only known from the type locality.

Comparative notes. Two Syndromus (sensu

lato) lineages are represented on the volcanic is-

land of Sumba (Laidlaw & Solem, 1961; Chan &

Tan, 2008, 2010). One lineage is represented by

the A (S.) latestrigatus Schepman, 1892 complex,

including A. (S.) latestrigatus sumbaensis Fulton,

1896 and A. (S.) floresianus Fulton, 1897. The

other lineage has only recently been discovered,

with the first species described being A. (S.) ab-

basi (see Chan & Tan, 2008: p. 9, figs. 1.1 -1.6;

2010: p. 247, figs. 1A-C). The new species, a sec-

ond addition to this latter lineage, is currently only

known from a patch of forest close to Mangili in

eastern Sumba which lies on a Pleistocene lime-

stone platform, and as such it is reproductively

separated from A abbasi, which is located at Lan-

ggaliru, about 122km in a northwesterly direction,

in another isolated forest patch, here on a late Cre-

taceous formation of sedimentary and volcanic

origin (Lytwyn et al., 2001).

Description of a new species of Amphidromus from Sumba, Indonesia (Gastropoda Pulmonata Camaenidae) 265

Figures 1-6. Shells of Amphidromus iunior n. sp., small forest close to Mangili village, east part of Sumba island, East Nusa

Tenggara, Indonesia - holotype. Figure 7. Map of Sumba showing the type localities of A. iunior n. sp. (Mangili, marked

with a white square) and of A. abbasi (Langgaliru, marked with a white circle). Inset map shows the Indonesian archipelago.

266

David P. Cilia

Figures 8-22. Shells of Amphidromus iunior n. sp., small forest close to Mangili village, east part of Sumba island, East Nusa

Tenggara, Indonesia. Figs. 8-12: paratype (FMNH 328102). Figs. 13-17: paratype (NHMUK 20120044). Figs. 18-22: paratype

(DCRG217).

Description of a new species of Amphidromus from Sumba, Indonesia (Gastropoda Pulmonata Camaenidae) 267

Figures 23-32. Shells of Amphidromus abbasi, forest in Langgalira, southwest part of Sumba island, East Nusa Tenggara, Indo-

nesia (-09°45'44"N, 119°38'33"E) (DC RGA698).

H (mm)

D (mm)

H/D

mean

min.

med.

max.

mean

min.

med.

max.

A. iunior n. sp. holotype

27.65

14.25

1.94

A. iunior n. sp. type series

27.70

23.30

27.65

33.80

14.31

12.65

14.25

15.75

1.93

A. abbasi topotype

37.78

35.60

38.00

39.50

18.86

18.00

18.95

19.65

2.01

Table 1. Condensed data for shell measurements of the type series of A. iuniorn. sp. (21 specimens) and a topotypical sample

of A. abbasi (6 specimens - DC R.GA698) for comparison. All measurements are in mm.

268

David P. Cilia

A number of morphological characteristics are

valuable in differentiating between the two species.

A. abbasi is larger, with its smallest specimens

exceeding by about 2mm in width and in height the

largest A. innior n. sp. specimens encountered (Table

1). The degree of conchological intraspecific vari-

ability in A. abbasi significantly exceeds that of the

A. iunior n. sp. population. It is also thicker-shelled,

reaching a degree of opacity lacking in A. iunior n.

sp. (Figs. 23-32). The two subperipheral brown spiral

bands may be extremely faint or absent in A. abbasi ,

and considerable variation in base colour occurs in

the topotypic population, with a spectrum ranging

from pink to yellow-brown, whereas this is never the

case for A. iunior n. sp. The columnar markings in the

former are frequently resolved to a series of dashes,

in the latter they are generally fused and continuous,

apart from the one main perpendicular interruption

through which the yellowish base colour appears. The

peristome of A. abbasi is more strongly reinforced

and its degree of fusion at the columellar side is rarely

incomplete; in A. iunior n. sp., the umbilical zone is

always narrowly perforated (closed in A. abbasi ). Pig-

mentation of the nucleus, a feature of significant di-

agnostic value amongst Nusa Tenggara Syndromus,

according to Goldberg & Sevems (1997), is always

restricted to a spot in A. abbasi, varying between

pinkish and brown, while in A. iunior n. sp. it is

always dark, descending and diminishing gradually

along the first whorl in proximity of the suture.

ACKNOWLEDGEMENTS

The author would like to thank John Abbas

(Jakarta, Indonesia), for making the specimens

available for study, Fred Naggs and Jonathan Ablett

(NHMUK), Jochen Gerber (FMNH), Philippe Bou-

chet and Virginie Heros (MNHN), John J. Borg

(NMHM), and Henk Mienis (HUJ) for technical

assistance, Fabio Liberto and Salvatore Giglio

(Cefalu, Italy) for photography and plate layout,

and anonymous reviewers for their suggestions.

REFERENCES

Bouchet P. & Rocroi J.-P., 2005. Classification and nomen-

clator of gastropod families. Malacologia, 47: 1-397.

Chan S.Y. & Tan S.K., 2008. On a new species of Am-

phidromus {Syndromus) (Gastropoda: Pulmonata:

Camaenidae) from Sumba Island, Indonesia. Occa-

sional Molluscan Papers, 1: 6-10.

Chan S.Y. & Tan S.K., 2010. On two new species of Am-

phidromus (Gastropoda: Camaenidae) from the Les-

ser Sunda Islands, Indonesia. The Raffles Bullettin of

Zoology, 58: 245-249.

Dharma B., 2007. Report on fossil Amphidromus and de-

scription of new species and subspecies of recent and

fossil Amphidromus from Indonesia (Gastropoda,

Pulmonata: Camaenidae). Schriften zur Malakozoo-

logie, 23: 45-78 + pis. 9-14.

Goldberg R.L. & Severns M., 1997. Isolation and evolu-

tion of the Amphidromus in Nusa Tenggara. Ameri-

can Conchologist, 25: 3-7.

Laidlaw F.F. & Solem A., 1961. The land snail genus Am-

phidromus-. a synoptic catalogue. Fieldiana (Zo-

ology), 41: 507-677.

Lok A.F.S.L. & Tan S.K., 2008. A review of the Singa-

pore status of the green tree snail, Amphidromus atri-

callosus perakensis Fulton, 1901 and its biology.

Nature in Singapore, 1: 225-230.

Lytwyn J., Rutherford E., Burke K. & Xia C. 2001. The

geochemistry of volcanic, plutonic and turbiditic

rocks from Sumba, Indonesia. Journal of Asian Earth

Sciences, 19: 481-500.

Scott B., 1996. Phylogenetic relationships of the Camae-

nidae (Pulmonata: Stylommatophora: Helicoidea).

Journal of Molluscan Studies, 62: 65-74.

Severns M., 2006. A new species and a new subspecies

of Amphidromus from Atauro Island, East Timor

(Gastropoda, Pulmonata, Camaenidae). Basteria, 70:

23-28.

Solem A., 1983. First record of Amphidromus from Aus-

tralia, with anatomical notes on several species (Mol-

lusca: Pulmonata: Camaenidae). Records of the

Australian Museum, 35: 153-166.

Sutcharit C. & Panha S., 2006. Taxonomic review of the

tree snail Amphidromus Albers, 1850 (Pulmonata:

Camaenidae) in Thailand and adjacent areas: Sub-

genus Amphidromus. Journal of Molluscan Studies,

72: 1-30.

Wade C.M., Hudelot C., Davison A., Naggs F. & Mordan

P.B., 2007. Molecular phylogeny of the helicoid land

snails (Pulmonata: Stylommatophora: Helicoidea),

with special emphasis on the Camaenidae. Journal of

Molluscan Studies, 73: 411-415.

Wade C.M., Mordan PB. & Naggs F., 2006. Evolutionary

relationships among the Pulmonate land snails and

slugs (Pulmonata, Stylommatophora). Biological

Journal of the Linnean Society, 87: 593-610.

Biodiversity Journal, 2013, 4 (2): 269-274

Reproductive cycle of the European anchovy Engraulis encra-

sicolus (Linnaeus, 1 758) (Clupeiformes Engraulidae) in the gulf

of Skikda (Algerian East coasts)

L/amine Mezedjri 1 *, Ahmed Kerfouf 2 &AliTahar 3

'Departement SNV, Faculte des sciences, Universite du 20 aout 55, Skikda 21000, Algerie; email: mezedjri.lyamine@gmail.com

2 Laboratoire d’ecodeveloppement des espaces, departement des sciences de l’environnement, faculte des sciences, universite Djillali

Liabes, Sidi Bel-Abbes 22000, Algerie; email: kerfoufalnned@yahoo.fr

3 Laboratoire de biologie vegetale et environnement, departement de biologie, Faculte des sciences, Universite Badji Mokhtar, An-

naba 23000, Algerie; email: prjahar _ali@hotmail.com

’Corresponding author

ABSTRACT A study on the sexual cycle of the European anchovy, the pelagic fish Engraulis encrasicolus

(Linnaeus, 1758) (Clupeiformes Engraulidae), was carried out in Algerian East coasts over

a year (July 2008-June 2009). Annual sex-ratio (SR) showed that females dominate with an

average male sex-ratio of 39.35%. The European anchovy spawning period ranges from April

to October with peaks in the warmest months, strictly dependent on temperature. The go-

nado-somatic index was updated monthly which allowed us to divide the entire cycle into

four successive phases: (i) a phase of slow maturation which occurs between the end of winter

and the beginning of spring; (ii) a phase of significant sexual activity; (iii) a phase of emission

of the sexual products which corresponds to the warmest months; and, finally, (iv) a phase of

sexual rest occurring in the coldest months. Monthly variation of either hepato-somatic index

or mesenteric fat reserves informed us about the origin of the energetic deposits, particularly

important for the development of the gonads. In both sexes, the first sexual maturity is reached

at a fish total length (Lt) of 12.5 cm.

KEY WORDS Algerian East-coasts; Engraulis encrasicolus', reproduction; sex-ratio; first sexual maturity.

Received 16.02.2013; accepted 14.05.2013; printed 30.06.2013

INTRODUCTION

Reproduction and sexual behaviour of the Euro-

pean anchovy Engraulis encrasicolus (Linnaeus,

1758) (Clupeiformes Engraulidae) were the subject

of many studies focusing on the connection between

these items and food behaviour, plumpness of fishes

and ecological factors in natural populations of this

species (Fage, 1911, 1920; Dorel, 1986; Hemida,

1987; Whitehead et al., 1988; Djabali & Hamida,

1989; Pertierra, 1992; Prouzet & Metuzals-Sebedio,

1994; E.R.H., 1996; Kara, 2001; Mezedjri & Tahar,

2005, 2006; Kada et al., 2009). This work brings new

information on the reproduction of E. encrasicolus

in the Gulf of Stora, Skikda (Algerian East coasts).

Sex-ratio, gonado-somatic and hepato-somatic in-

dexes, adiposity and size at first sexual maturity were

studied. Obtained results were compared with those

reported in literature for several Mediterranean areas.

MATERIALS AND METHODS

The study was carried out on fishes caught by

means of sliding seines in the Gulf of Stora (Alge-

270

L. Mezedjri.A. Kerfouf & A.Tahar

ria) during one year, from July 2008 to June 2009.

Just with the unloading, a sample of 1 kg at mini-

mum was taken at the port of Stora, monthly. On

each fish a series of measures were made (Table

1) including length, by using an ichthyio-meter

with meadows millimetre; total and eviscerated

weights, by means of a balance with an accuracy

of 0.01 g; and gonadic and hepatic weights to the

nearest 0.0001 g.

Determination of maturity stages was carried

out according to the method recommended by the

ERJJ team (E.R.H., 1996), in particular, by taking

into account the degree of fattening of each spec-

imen according to an empirical four degree-scale

which is a derivative of the scale of Nikolsky

(1963). In literature there is a divergence on the for-

mula for the sex-ratio. In our work we used the for-

mula which gives sex-ratio as a percentage of males

by the following relation: SR= (Males Number/Total

Number)xl00. Then, the chi-squared (x 2 ) test was

used to evaluate the variation of the observed values

of the sex-ratio compared to the theoretical propor-

tion of 50% (Dagnelie, 2006). We supposed as hy-

pothesis H 0 : sex-ratio = 50% and tested this

hypothesis by calculating the value of x 2 obs by the

formula:

X 2 = (nr / F) + (f~ / F ) - n

where, m: number of males; f: number of fe-

males; n = m + f; F = n/2. When x 2 0 b s > T\-a at ^

degree of freedom we rejected the hypothesis H 0 at

the level a = 0.05. Please note that this test is valid

only for number of males or females higher than 5

(Dagnelie, 2006). The gonado-somatic index (GSI)

N°

CODE

DESCRIPTION

Total length

Total Weight

Eviscerated Weight

Wgon

Gonadic Weight

Whep

Hepatic Weight

Adip

Adiposity

Sex

Identification of sex and stage of ma-

turity

Table 1 . Measures made on fishes for the reproduction survey.

is an index expressed in percentage. It is calculated

starting from the relationship between the weight

of the gonads (Wgon) and the eviscerated weight of

fish (We). We preferred to use the weight of emp-

tied specimens in the place of the total weight to

eliminate the variations due to the repletion state of

the stomach. The GSI is calculated for each indi-

vidual according to the following formula (Kara,

1997): GSI=(Wgon/We) x 100. GSI values reflect

changes in gonads weight during the reproductive

cycle, thus making us possible to monitor their

maturation (Barnabe, 1976). The hepato-somatic

index (HSI) is expressed in percentage as well. This

relationship links the hepatic weight (Whep) and

the eviscerated weight (We) according to the follow-

ing formula (Kara, 1997): HSI= (Whep/We) x 100.

Its changes are connected to the variations of he-

patic weight during the reproductive cycle. Monthly

variations of adiposity (Adip), i.e. the mesenteric

fat, were followed during the entire period (July

2008-June 2009). All the specimens caught during

the reproduction period (from April to October)

were distributed, according to their size, in classes

of sizes with an interval of 0.5 cm. Then, we took

in consideration the frequency of the mature indi-

viduals in relation to the total number of specimens

belonging to each class. Size at first sexual maturity

is determined as the size of 50% of the mature in-

dividuals (Barnabe, 1976; Kara, 1997). This para-

meter is calculated separately for each sex, and then

the two sexes are compared. All calculations have

been executed with the software MINITAB for

analysis and statistical treatment of the data, version

16 for Windows (2010).

RESULTS AND DISCUSSION

Sex-ratio

On a whole of 803 examined individuals, we ob-

served 316 males and 487 females with an average

male sex-ratio of 39.35% (x 2 = 36.41; P<0.001).

Monthly variations of sex-ratio are shown in Table

2. Broadly speaking, the sex-ratio was always in

favour of females (with a male SR ranging from

18.42% to 35.62%, 0<P<0.05) except during Sep-

tember, when we noted a value of sex-ratio in favour

of males (SR=7 1 .96%, P<0.001). SR values assessed

in October 2008 and March 2009 (56.30% and 40%,

Reproductive cycle of the European anchovy Engraulis encrasicolus in the gulf of Skikda (Algerian East coasts) 271

respectively) were not statistically significant. Fe-

males remained dominant for 7 months, particularly

during the warmest ones (from April to August),

when the anchovies come closer to the Algerian co-

asts, where they are captured easily. This leads us to

say that the Gulf of Stora may be considered as a

zone of spawning of the European anchovy.

Month

No. of

males

No. of

females

Sex-ratio

X2 obs

Jul-08

18.4211

30.31579 ***

Aug

100

33.3333

16.66667 ***

Sep

71.9626

20.64486 ***

Oct

56.3025

1.89076 ns

Nov

Dec

Jan-09

31.5789

5.15789 *

Feb

22.5225

33.52252 ***

Mar

3.2 ns

Apr

35.6164

6.04109 *

May

23.0769

11.30769 ***

Jun

0.4 ns

Total

316

487

39.3524

36.41469 ***

Table 2. Monthly variations of male sex-ratio for Engraulis

encrasicolus from the gulf of Stora between July 2008 and

June 2009. p > a = 0,05: (ns) not significant, p < a = 0,05:

(*) significant, p < a = 0,001 : (***) very highly significant.

Gonado-somatic index

Graphically, monthly fluctuations of gonado-so-

matic index (GSI) values of E. encrasicolus are

shown in figure 1. For females, GSI showed the high-

est values in summer with a maximum in August

(GSI=4.31%), decreased at the beginning of au-

tumn, the lowest values being recorded in winter

with a minimum in February (0.65%), and then

gradually and notably increased in spring. Males

ex-hibited more or less a similar trend, with the ma-

ximum value recorded in September (3.39%) and

the least in January (0.86%). Noteworthy, the in-

crease in GSI values through the spring period

(March- June) was less marked than in females.

Monthly variations of the gonado-somatic index re-

vealed that the reproduction period of the anchovy

in our area spreads between April and October. This

is in line with Whitehead et al. (1988) who affirm

that the peak of spawning generally coincides with

the warmest months, the limits of the spawning sea-

son being dependent on the temperature. Plotting

the evolution of GSI values showed that the repro-

ductive cycle of E. encrasicolus is characterized by

four successive phases: (i) a phase of slow matura-

tion which starts in March and continues until June;

(ii) a phase of significant and intense sexual activity

which occurs during the warm-est months (from

June to September); (iii) a phase of decline starting

in September during which the testicles and the

ovaries gradually loose their mass; and finally (iv)

a phase of rest, coinciding with the coldest months

(from December to Feb-ruary), when fishes become

sexually inactive.

These findings once again support the hypothe-

sis that temperature exerts a strong influence on the

sexual cycle of the European anchovy, with high

temperatures leading to spawning and low temper-

atures preventing sexual activity (Whitehead et al.,

1988; Djabali & Hamida, 1989).

Hepato-somatic index

For the hepato-somatic index (Fig. 2) we ob-

served, in females, a maximum in July

(HSI=1 .58%) and, then, a clear reduction up to Oc-

tober (0.85%), with the lowest values in January

(0.53%) when it dramatically increased to reach a

maximum in February (1.17%). After that, HSI

gradually decreased up to June (0.64%). In males

we observed the same trend, although absolute val-

ues were slightly lower than in females. HSI, com-

pared with GSI, was less marked. During the

reproduction period HSI values decreased probably

due to the consumption of fat deposits, reaching a

peak in winter (at the end of February) which sug-

gests a possible storage of reserves in the liver dur-

ing the sexual rest, followed by the reduction of HSI

from March on, when sexual maturity begins. It is

interesting to note that, especially for females, HSI

and GSI values did not always show complemen-

tary trends, thus suggesting that energy reserves

used to allow the development of the gonads may

not be accumulated exclusively in the liver.

272

L. Mezedjri.A. Kerfouf & A.Tahar

Djabali & Hamida (1989) during a study on the

European anchovy, carried out between 1983 and

1984, concluded that the liver didn’t have any role

in the maturation process of the sexual products;

this implies that the two parameters (GSI and HSI)

should be studied together.

Adiposity

Changes in mesenteric deposits in E. encrasi-

Figure 1. Monthly variations of GSI in Engraulis encrasi-

colus from the gulf of Stora.

Figure 2. Monthly variations of the HSI for E. encrasicolus

from the gulf of Stora.

Figure 3. Monthly variations of adiposity for E. encrasico-

lus from the gulf of Stora.

Figure 4. Size at first sexual maturity in females of E. en-

crasicolus from the gulf of Stora.

Figure 5. Size at first sexual maturity in males off. encra-

sicolus from the gulf of Stora.

colus are shown in figure 3. The maximum values

were recorded in September (males) and January

(females) while, in both sexes, the lowest ones

were in April.

Taking into account that the highest values were

observed in winter (during the sexual rest) and the

lowest ones in the warmest months, these findings

confirm the hypothesis, maintained by Djabali &

Hamida (1989), that gonadic reserves may origin

mainly from mesenteric fat deposits.

Year 2008 2009

Calss of size

Reproductive cycle of the European anchovy Engraulis encrasicolus in the gulf of Skikda (Algerian East coasts) 273

Size at first sexual maturity

The size at the first sexual maturity (Lt) is roughly

the same for both sexes and corresponds to 12.5

cm (Figs. 4, 5). As can be seen, our data are in line

with most of the Lt values available for E. encra-

sicolus populations in the Mediterranean (Table 3),

suggesting that, in the areas analysed, there are not

big differences in environmental factors, particu-

larly water temperatures, population density and

food availability, while Lt = 6.5 cm reported for

the lagoon of Nador (Morocco) might be possibily

due to disturbing issues including high fishing

pressures.

In conclusion, the present study not only gave

information on any biological aspects of Engraulis

encrasicolus from the guld of Skikda (Algerian

East-coasts), which up to now were deficient, but

also contributed to provide a preliminary base for

monitoring the changes in biological indexes con-

nected to the reproduction cycle of this species in

order to ensure its sustainable management in the

Mediterranean area.

Lt (cm)

Sex

Locality

Author

13.0

Males

Golf of Biscay

Dorel, 1986

13.0

Females

Golf of Biscay

Dorel, 1986

11.2

Males

Bay of Algiers

Djabali &

Hamida, 1989

11.6

Females

Bay of Algiers

Djabali &

Hamida, 1989

11.5

Indefinite

Coast of

Catalonia

Pertierra, 1992

12.5

Both sexes

Golf of Stora

(Skikda)

Mezedjri &

Tahar, 2005; 2006

12.5

Both sexes

Golf of Stora

(Skikda)

present paper

06.5

Both sexes

the lagoon of

Nador (Morocco)

Kada et al., 2009

Table 3. Size at first sexual maturity of Engraulis encrasi-

colus in some regions of the Mediterranean Sea.

ACKNOWLEDGEMENTS

Authors wish to thank two anonymous referees

for the their suggestions which significantly im-

proved the manuscript.

REFERENCES

Bamabe G., 1976. Contribution a la connaissance de la bio-

logie du loup Dicentrarchus labrax (L.) (Poisson Se-

raranide) de la region de Sete. These d’etat, Universite

des Sciences et Techniques du, Montpellier, 426 pp.

Dagnelie P., 2006. Statistique theorique et appliquee.

Tome 2: Inferences statistique a une et a deux dimen-

sions. De Boeck, Bruxelles, 659 pp.

Djabali F. & Hamida F., 1989. Reproduction de l'anchois

( Engraulis encrasicolus, Linne 1758) de la region d' Al-

ger. Bulletin de Pinstitut des sciences de la mer et de

l’amenagement du littoral ISMAL. Pelagos, 7: 11-14.

Dorel D., 1986. Poissons de l'Atlantique nord-est rela-

tions taille-poids. Institut Franqais de Recherche pour

l'Exploitation de la Mer. IFREMER, Nantes, 165 pp.

E.R.H., 1996. Evaluation de la ressource halieutique. Mi-

nistere des peches et des ressources halieutiques.

Rapp. Campagne, 40 pp.

Fage L., 1911. Recherches sur la biologie de l’anchois

(Engraulis encrasicolus Linne): races-age-migra-

tions. Annales de l'lnstitut oceanographique Monaco,

2: 1-40.

Fage L., 1920. Engraulidae, Clupeidae. Report on the

Danish Oceanographical Expeditions 1908-1919 to

the Mediterranean and adjacent seas, Vol. 2, 140 pp.

Hemida F., 1987. Contribution a l’etude de l’anchois En-

graulis encrasicolus (Linne, 1758) dans la region

d’ Alger. Biologie et exploitation. These de magistere,

USTHB.

Kada O., Abdellaoui S., Ramdani M. & Nachit D., 2009.

Contribution a 1’ identification et a la caracterisation

biologique et dynamique de l’anchois de la lagune de

Nador (Maroc). Bulletin de l’lnstitut Scientifique,

Rabat, section Sciences de la Vie, 31: 91-98.

Kara M.H., 1997. Cycle sexuel et fecondite du loup Di-

centrarchus labrax (poisson Moronide) du golfe

d’Annaba. Cahiers de Biologie Marine, 38: 161-168.

Kara M.H., 2001. Biologie, dynamique et evaluation des

ressources halieutiques coheres du littoral Est-alge-

rien. Projet de recherche: Ministere de l’Enseigne-

ment Superieur et de la Recherche scientifique.

Mezedjri L. & Tahar A., 2005. Reproduction de l’an-

chois Europeen dans le golfe de Skikda, Algerie :

Actes du Sixieme Congres Maghrebin des Sciences

de la Mer Monastir, Tunisie (18-22 decembre 2005).

Abstract book.

Mezedjri L. & Tahar A., 2006. Reproduction de l’anchois

Europeen Engraulis encrasicolus dans le golfe de

Stora, Skikda. 8eme conference internationale des

limnologues d’expression franqaise « CILEF 2006 ».

Hammamet, Tunisie. Abstract book.

Minitab 16 Statistical Software (2010). [Computer soft-

ware]. State College, PA: Minitab, Inc. (www.mini-

tab.com).

274

L. Mezedjri.A. Kerfouf & A.Tahar

Nikolsky G.V., 1963. The ecology of fishes. Academic

Press. London, 352 pp.

Pertierra J.P., 1992. Fisheries biology of the anchovy (En-

graulis encrasicolus ) in the Catalan coast. Ph.D. The-

sis, Universitat Politeenica de Catalunya, Spain.

Prouzet P. & Metuzals-Sebedio K.I., 1994. Population

structure and reproductive biology. Stock discrimi-

nation studies using morphometric and genetic data.

In: Cendrero O. (Ed.), Improvement of Stock Asses-

sment by direct Methods, Its Application to the An-

chovy ( Engraulis encrasicolus ) in the Bay of Biscay.

Final Report. Projet DG XIV EU, ref MA. 2. 495.

Whitehead P.J.P., Nelson G.J. & Wongratana T., 1988.

FAO species catalogue Vol. 7. Clupeoid fishes of the

world. An annotated and illustrated catalogue of the

herrings, sardines, pilchards, sprats, shads, anchovies

and wolf-herrings. Part 2. Engraulididae. FAO Fishe-

ries Synopsis, 7: 305-579.

Biodiversity Journal, 2013, 4 (2): 275-280

Amauta hodeei (Oberthur, 1881) and its subspecies (Lepi-

doptera Castniidae), with comments on the life and times of

Brother Apolinar Maria

Jorge M. Gonzalez 1 *, Roberto Vinciguerra 2 & Sergio D. Rios 3

'California State University, Fresno, Department of Plant Sciences, Fresno, CA 93740-8033 (Research Associate, McGuire Center

for Lepidoptera & Biodiversity), USA; e-mail: gonzalez.jorge.m@gmail.com

2 Via XX Settembre, 64, 1-90141 Palermo, Italy

’Museo Nacional de Historia Natural de Paraguay, Sucursal 1 Campus, Central XI, San Lorenzo, Departamento de Biologia, Fa-

cultad de Ciencias Exactas y Naturales, Universidad Nacional de Asuncion, Paraguay. E-mail: sergiord40@gmail.com

*Corresponding author

ABSTRACT In 1948, a fired destroyed the La Salle Museum in Bogota, Colombia, which was built with

a great effort by the La Salle religious teaching congregation, but with the particular and de-

cisive input of Brother Apolinar Maria. He became a champion of the study of the natural

history of Colombia and through the museum, he established numerous connections with

scientists and naturalists worldwide. Some rare Castniidae were among the numerous speci-

mens of the fauna he traded with museums around the world. General information about

Brother Apolinar Maria, the La Salle Museum, and the subspecies of Amauta hodeei (Cast-

niidae) are provided here in an attempt to improve and stimulate the interest in such a re-

markable naturalist and some rare and almost unknown species of the South American Fauna.

One of these ssp. (A. hodeei kruegeri ) is reported from Ecuador, thus increasing knowledge

about its geographical distribution.

KEY WORDS Museo La Salle Bogota; Castniidae; Amauta hodeei hodeei', Amauta hodeei kruegeri.

Received 22.02.2013; accepted 08.04.2013; printed 30.06.2013

INTRODUCTION

Massive riots followed the assassination of po-

litical leader and presidential candidate Jorge Elie-

cer Gaitan (1903-1948) in Bogota, Colombia, on

April 9, 1948. The 10 hour riot resulted in a fire that

left thousands of dead and injured, and much of

downtown Bogota was destroyed by the effects of

a widespread fire (Alape, 1994). The Museum of

Natural History La Salle was among those buildings

that disappeared because of the fire (Figs. 2, 3). It

originally contained samples of over 73,000 speci-

mens of several groups, and by 1930, some 37,706

of those specimens were insects (Salazar, 1999).

The success of this museum was mainly the product

of the efforts of Nicolas Seiller (1867-1949), better

known as Brother Apolinar Maria (Figs. 1, 4)

(Velez & Salazar, 1991; Salazar, 1999; Diaz Meza

et al., 2006). The museum and its annex library

were consumed by the fire and, as a result, Apolinar

Maria entered a deep state of depression and died

in December, 1949 (Salazar, 1999).

Seiller was born in Alsace, France, in 1867.

Some time later, he went to Reims, where he

eventually joined the religious Brotherhood of La

Salle.

276

J.M. GonzAlez, R.Vinciguerra & S.D. Rios

As customary in the brotherhood, he changed

his name to Apolinar Maria and was then sent to

Colombia, where he arrived in 1904 (Lamas, 1979).

Thanks to his interest in the natural sciences, he

founded the Natural History Society of La Salle In-

stitute in Bogota, in 1912 (Lamas, 1979). With the

help of other brothers and local amateur naturalists,

he also built the museum that would eventually con-

tain the largest collection of Lepidoptera in Colom-

bia (Lamas, 1979; Salazar, 1999). Most importantly,

he emphasized the need of knowing and sharing in-

formation with scientists around the world, and with

this vision, he sent countless Colombian specimens

to several institutions and private American and

European collections in an effort to trade, but also

obtain the appropriate identification of his adopted

country’s fauna and flora (Lamas, 1979; Rodriguez,

2002; Freile & Cordova, 2008). Together, with col-

leagues and pupils, he published numerous papers

on Lepidoptera, described many new taxa, and

made observations on Colombian natural history

(Lamas, 1979, 2013).

Figure 1. Brother Apolinar Maria examining specimens inside the Natural History Museum, La Salle, Bogota (Picture borro-

wed from Diaz Meza et al. 2006). Figure 2. La Salle Museum being devoured by the flames, 1948. Figure 3. La Salle Museum

after the flames were controlled, 1948. Figure 4. Nicollas Seiller, best known as Brother Apolinar Maria (1867-1949).

Amauta hodeei and its subspecies (Lepidoptera Castniidae), with comments on the life and times of Brother Apolinar Maria 277

Through the Museum he established communi-

cations with scientists and naturalists around the

world, including renowned entomologists such as

Charles Oberthiir (1845-1924), Paul Dognin (1856-

1931), Eugene Louis Bouvier (1856-1944), Harri-

son Gray Dyar jr. (1866-1929), Anton H. Fassl

(1876-1922), William Schaus (1858-1942), Johan-

nes Karl Rober (1861-1942), Arnold Schiiltze

(1875-1948), and Romualdo Ferreira D’ Almeida

(1891-1969) (Lamas 1979; Salazar, 1999; Velez &

Salazar, 1991; Julian Salazar, pers. comm.).

Apolinar Maria is considered one of the most re-

levant and recognized Colombian naturalists and

the precursor of the studies of Lepidoptera in the

country (Salazar 1999; Rodriguez, 2002). Aside

from the collecting he and other brothers from his

congregation did while travelling to different Co-

lombian locations, he hired people to collect fauna

and flora specimens, mainly from Muzo, Boyaca,

along the forests that border Rio Magdalena and

other effluents, like Rio Carare, as well as Villavi-

cencio, Meta (Apolinar Maria, 1915; Julian Salazar,

pers. comm., 2012).

He published some of his findings, including the

descriptions of several new species, in various jour-

nals and newsletters (Lamas, 1979; 2013). In Lepi-

doptera, he mainly published about butterflies,

however he also wrote several papers or notes with

information about moths (Salazar, 1999; Lamas,

2013). Among those works, he wrote about some

Castniidae found in Colombia (Apolinar Maria,

1915, 1945). Brother Apolinar Maria recognized

and made brief comments about ten species and se-

veral subspecies in the Castniidae collected in Co-

lombia and deposited at Bogota’s La Salle Museum

(Apolinar Maria, 1915, 1945; Salazar, 1999).

Among the over 3 1 taxa belonging to 2 1 species

of Castniidae found in Colombia (Hernandez-Baz

et al., 2012), some belong to the genus Amauta

Houlbert, 1918. According to Lamas (1995) it con-

tains four species: A. ambatensis (Houlbert, 1918),

A. cacica (Herrich- Schaffer, [1854]), A. hodeei

(Oberthiir, 1881), and A. papilionaris (Walker,

[1865]). They are all species of medium to large

dimensions, with triangular forewings, commonly

dark brown to black, with some white/cream and

orange markings in both wings and they are se-

xually dimorphic (Miller, 1986). The genus is dis-

tributed from Guatemala, through Central America

and down to South America, where specimens have

been found in Colombia, Peru, Ecuador, Bolivia

and Brazil (Miller, 1986, 1995; Lamas, 1995). Un-

fortunately, the known information on the several

species in the genus is still fragmentary and scarce,

and data found in several insect Museums is far

from adequate (Miller, 1986; Miller & Sourakov,

2009).

Even though we have been able to study Castnii-

dae specimens from many museums worldwide, we

could locate only few of Amauta hodeei. Among the

known specimens of both subspecies (A. hodeei ho-

deei and A. hodeei kruegeri), including those men-

tioned in the literature, a few were collected by

Brother Apolinar Maria (or his group of collectors).

Both subspecies are known to occur in Colombia;

however, a specimen of A. hodeei kruegeri collec-

ted in Ecuador is reported herein, thus expanding

its known distribution.

Specimens examined are in the following col-

lections: AMNH = American Museum of Natural

History, New York, USA; MGCLB = McGuire

Center for Lepidoptera & Biodiversity, (Allyn Mu-

seum Collection), Gainesville, Florida, USA;

MHN-UC = Natural History Museum, University

of Caldas, Manizales, Colombia; MffNP = Natural

History Museum, Paris, France; NMNH-SI = Na-

tional Museum of Natural Histoiy, Smithsonian In-

stitution, Washington D.C., USA; GPC = G. Penati

Collection, Milan, Italy; RVC = Roberto Vinci-

guerra Collection, Palermo, Italy.

COMMENTS ON THE TWO KNOWN SSP.

OF AMAUTA HODEEI (OBERTHUR, 1881)

Amauta hodeei hodeei (Oberthiir, 1881)

This subspecies was originally described (as

Castnia hodeei ) by Oberthiir (1881) from material

collected in the Colombian region along Santa Rosa

and the Carare river. Years later, Schaus (1896) de-

scribed Castnia corrupta from Colombia, while

Oberthiir (1925) illustrates a species he names C.

apollinaris [nomem nudum (Lamas, 1995)] based

on a male specimen collected in Colombia, and

compares and discusses similarities and differences

with the species C. corrupta. That same year, Knop

(1925) describes C. amazona from Bogota, Colom-

bia. Those species have been established as syno-

nyms of hodeei hodeei (Lamas, 1995).

278

J.M. GonzAlez, R.Vinciguerra& S.D. Rios

Unfortunately, this is a rare taxon and almost no-

thing is known about its biology and ecology. In

fact, very few specimens are known from insect col-

lections worldwide. Joicey & Talbot (1925) men-

tioned that they received three specimens (two

males and a female) sent by Brother Apolinar Maria

which were originally collected in Muzo and Villa-

vicencio. Gonzalez & Salazar (2003) also reported

a male and a female from Guamoco and Cauca Val-

ley (Caucaval).

Examined material (with notes). COLOMBIA:

1 male, 1 female, Bogota, 1931, Coll. Fre. Apoli-

naire Marie (MHNP) [even though the labels (Fig.

10) clearly state that this material was collected in

Bogota, they were most certainly collected in the

Muzo region where Brother Apolinar Maria had

peasants hired to collect specimens. Since the La

Salle Museum was located in Bogota, most Institu-

tions that received material from Apolinar Maria,

commonly assumed that “Bogota” was the collec-

Figure 5. Amauta hodeei hodeei, male, Bogota, Colombia (MHNP). Figure 6, Idem, male, Guamoco Colombia (AMNH).

Figure 7. Idem, female, Cauca Valley, Colombia (AMNH). Figure 8. A. hodeei kruegeri, Esmeraldas, Ecuador (RVC). Figure

9. Idem, Rio Calima, Colombia (MGCLB). Figure 10. Labels of two A. hodeei hodeei specimens collected or sent by

Brother Apolinar Maria (MHNP). Scale (for figures 5-9): 10 mm.

Amauta hodeei and its subspecies (Lepidoptera Castniidae), with comments on the life and times of Brother Apolinar Maria 279

ting locality]; 1 male?, Muzo, Boyaca, [the occiden-

tal sector with influence of the middle area of the

Magdalena River], Coll. Apolinar Maria [reported

in Apolinar Maria (1945). This specimen was surely

lost in the Museum’s fire]; 1 male, Campo Santo,

Bogota, 1922, Coll. L. Pfeiffer (NMNH-SI) [this is

the type specimen of Castnia corrupta described by

Schaus (1896) and now considered a synonym of A.

hodeei hodeei (Lamas, 1995)]; 1 female, Colombia,

Boyaca, Coll. ? (NMNH-SI); 1 male, Guamoco,

Coll. F. Johnson (AMNH); 1 female, Caucaval

[Cauca Valley], Coll. F. Johnson (AMNH); 1 male,

Santa Rosa-Carare, Coll. Oberthiir [mentioned in

Oberthiir (1881) and Houlbert (1918)].

Amauta hodeei kruegeri (Niepelt, 1927)

Only males are known of this rare subspecies,

and they differ from the nominate subspecies be-

cause of its darker, blackish, background color, and

also because of the very distinctive white and

orange markings in the hind wings (Figs. 5-9).

Niepelt (1927) described it (as Castnia kriigeri )

based on a male collected in “West Colombia.” We

have been able to examine four specimens, all

males, from four different collections. The fore-

wings are devoid of any markings except for a small

light amber band in the sub-apical region. They are

larger than the sub-apical creamy spot observed in

males of the nominate subspecies (Figs. 5, 6, 8, 9).

A large, creamy-white spot can be observed in the

tormal region of the hind wing. The spot turns

orange at the edge of the wing. Such markings are

clearly noticeable, better defined, and larger in the

four studied specimens than in the similar (but

smaller and faddier) markings found in males of the

nominate species (Figs. 5, 6, 8, 9).

Examined material. COLOMBIA: 1 male, Rio

Dagua, W. Colombia, ex coll. H. Gerstner (GPC);

1 male, Rio Calima, VII, [19] 84, Allyn Museum

Ace. 1991-13, Allyn Museum Photo No. 850827-

15/16, Slide No. M-7132, male append. Jacqueline

Y. Miller (MGCLB); 1 male, Valle, Queremal -

Km 55, 9-IV-85, Leg. J.A Salazar (MJJN-UC).

ECUADOR: 2 males, Esmeraldas, (RVC).

All specimens known to us (and listed above) of

A. hodeei hodeei were collected east of the Western

Cordillera while those of A. hodeei kruegeri were

collected on the western slope of the Western Cor-

dillera in Colombia, and closer to the coast and west

01 "O'CTW 00'fftrW 79XrtrW 78 WW 7T‘UCrW 76WW 75 , 0 , 0"W 74'0 r (rW 73*0 - 0"W

— I 1 i i I I I I I

8 1WW &0 a cnrw 79*ff(TW 70WW 77 e CrO“W 76 WW 75 WW 74 1 <WW 73WW

Figure 1 1 . Map showing localities and ecological regions

where specimens of Amauta hodeei ssp. have been collected

in Colombia and Ecuador. A. hodeei hodeei: A1 : Magdalena,

Guamoco; A2: Magdalena, Santa Rosa and Carare; A3:

Magdalena, Muzo; D4: Paramo, Bogota; A. hodeei kruegeri :

B6: Choco, Rio Dagua; C5: Cauca, Rio Calima; E7: East

Ecuador, Durango; F: Napo.

of the Ecuadorian Western Cordillera (Fig. 11). The

Western Cordillera of Colombia continues along the

Western Cordillera of Ecuador and they seem to be

a geographic barrier that clearly separates the po-

pulations of both subspecies. The populations of

both ssp. appear to be highly dispersed along their

geographic range, but the low number of specimens

known and the lack of knowledge about their bio-

logy and ecology prevent us from providing any in-

formation about their conservation status.

ACKNOWLEDGEMENTS

We are greatly indebted to curators and persons

in charge of the revised insect collections mentio-

ned herein. We appreciate the kindness and help of

Ronald Erwin Rengifo and the personnel in charge

of the photographic register of the La Salle Mu-

seum, Bogota, Colombia, who provided us with

pictures of Brother Apolinar Maria (Figs. 1, 4). Also

thanks to Julian Salazar who provided us with pic-

tures of the La Salle Museum while burning and

after the fire was extinguished (Figs. 2, 3). Thanks

280

J.M. GonzAlez, R.Vinciguerra& S.D. Rios

to Joan Sharma and Martin Valencia (California

State University, Fresno, USA) for their help to

assemble the plates and to Andrea C. Gonzalez and

William Hale for proof reading the manuscript.

Our friend and colleague Roberto Vinciguerra

passed away shortly after this manuscript was

submitted to BJ for consideration. We (JMG &

SDR) will like to dedicate this work to his memory.

REFERENCES

Alape A., 1994. El Bogotazo: memorias del Olvido. Abril

9 de 1948. Planeta, Colombia, 653 pp.

Apolinar Maria, 1915. Miscelanea. Capturas interesantes.

Boletin de la Sociedad de Ciencias Naturales del Ins-

titute La Salle, 3: 141-144.

Apolinar Maria, 1945. Miscelanea entomologica. I-Algo

sobre castnidos colombianos. Revista de la Academia

Colombiana de Ciencias Experimentales, Fisicas &

Naturales, 6: 322.

Diaz Meza C.J., Jimenez J.R. & Turriago D., 2006. His-

toricidad, Saber y Pedagogia, Una Mirada al Modelo

Pedagogico Lasallista en Colombia 1915-1935. Uni-

versidad de La Salle, Bogota, Colombia. 170 pp.

Freile J.F. & Cordoba S., 2008. Historia de la omitologia

en la region andina: el ejemplo de Colombia y Ecua-

dor. Omitologia Neotropical, 19 (suppl.): 381-389.

Gonzalez J.M. & Salazar J.A., 2003. Adicion a la lista de

castnidos (Lepidoptera: Castniidae: Castniinae) co-

nocidos de Colombia. Boletin Cientifico del Museo

de Historia Natural, Universidad de Caldas, Maniza-

les, 7: 47-56.

Houlbert C., 1918. Revision monographique de la sous-

famille des Castniinae. Etudes de Lepidopterologie

Comparee,15: 1-730.

Joicey J.J. & Talbot G., 1925. Notes on some Lepidop-

tera, with descriptions of new forms. Annals and Ma-

gazine of Natural History, (9)16 (96): 633-653.

Knop T., 1925. Castnia amazona nov. spec, und Papilio

androgeus (Cr.) nov. var .fassli. Entomologische Zeit-

schrift, 39: 42-43.

Lamas G., 1979. Brother Apolinar Maria (1867-1949)

and his contributions to Colombian lepidopterology.

Journal of the Lepidopterists Society, 33: 65-67.

Lamas G., 1995. A critical review of J. Y. Miller’s chec-

klist of the Neotropical Castniidae (Lepidoptera). Re-

vista Pemana de Entomologia, 37: 73-87.

Lamas G., 2013. Bibliography of Butterflies. An Anno-

tated Bibliography of the Neotropical Butterflies and

Skippers (Lepidoptera: Papilionoidea and Hesperioi-

dea). Revised Electronic edition, 672 pp.

Miller J.Y., 1986. The Taxonomy, Phylogeny, and Zoogeo-

graphy of the Neotropical Castniinae (Lepidoptera:

Castnioidea: Castniidae). Ph.D. Thesis. University of

Florida, Gainesville, 571 pp.

Miller J.Y., 1995. Castniidae. In: Heppner J.B. (Ed.).

Checklist: Part 2. Hyblaeoidea-Pyraloidea-Tortricoi-

dea. Atlas of Neotropical Lepidoptera, 3: 133-137,

176-177. Association for Tropical Lepidoptera/Scien-

tific Publishers, Gainesville.

Miller J.Y. & Sourakov A., 2009. Some observations on

Amauta cacica procera (Boisduval) (Castniidae:

Castniinae) in Costa Rica. Tropical Lepidoptera Re-

search, 19: 113-114.

Niepelt W., 1927. Neue Falter von Columbien. Interna-

tionale Entomologische Zeitchrift, 21: 239-241.

Oberthur C., 1881. II. Lepidopteres d’Amerique. Etudes

d’Entomologie, 6:xxv - xxvi, 27-38.

Oberthur C., 1925. Lepidopteres de l’Amerique du Sud.

Eudes de Lepidopterologie. Comparee, 9: 63-64.

Rodriguez G., 2002. El naturalista Antoine Rouhaire

(Hermano Niceforo Maria) y el desarrollo de la Zoo-

logia sistematica en Colombia. Revista de la Acade-

mia Colombiana de Ciencias, 26: 229-237.

Salazar J.A., 1999. Notas biograficas a una historia de la

Lepidopterologia en Colombia durante el siglo XX.

Boletin Cientifico del Museo de Historia Natural,

Universidad de Caldas, Manizales, 3: 71-102.

Schaus W., 1896. New species of Heterocera from tropi-

cal America. Journal of the New York Entomological

Society, 4: 147-154.

Velez J.H. & Salazar J.A., 1991. Mariposas de Colombia.

Villegas Eds., Bogota, Colombia, 167 pp.

Biodiversity Journal, 2013, 4 (2): 281-284

Further records of two uncommon Crustaceans in Italian

seas: Maja goltziana D'Oliveira, 1 888 (Decapoda Brachyura

Majidae) and Xaiva biguttata (Risso, 1816) (Decapoda Bra-

chyura Portunidae)

Joachim Langeneck 1 * & Davide Di Franco 2

'Via Derna 13B, 56126 Pisa, Italy; email: jo.langeneck@gmail.com

2 Via S. Maria dei Monti 38/D, 95030, Gravina, Catania, Italy; email: davide_di_franco@hotmail.it

* Corresponding author

ABSTRACT The presence of Maja goltziana D'Oliveira, 1888 (Decapoda Brachyura Majidae) is confirmed

in the central Tyrrhenian Sea and in the Tuscan Archipelago. Xaiva biguttata (Risso, 1816)

(Decapoda Brachyura Portunidae) is first recorded from the Western Sardinian coast and the

Eastern Sicilian coast. The presence and spread of M. goltziana are considered a consequence

of climatic changes. The presence of A. biguttata in Italian seas was probably underestimated,

and its supposed rarity should be reassessed.

KEY WORDS Maja goltziana ; Xaiva biguttata', Mediterranean Sea; new record; climatic changes.

Received 3.04.2013; accepted 9.06.2013; printed 30.06.2013

INTRODUCTION

Maja goltziana D'Oliveira, 1888 (Decapoda

Brachyura Majidae) and Xaiva biguttata (Risso,

1816) (Decapoda Brachyura Portunidae) are two

Crustaceans generally considered rare in the Italian

seas. In this paper are reported some further records

of these species, and their current status in the Ital-

ian seas is discussed.

ABBREVIATIONS. CC = F. Crocetta collec-

tion, Napoli, Italy; CD = D. Di Franco collection,

Gravina, Catania, Italy; CF = J. Fangeneck collec-

tion, Pisa, Italy.

Maja goltziana D’Oliveira, 1888

Examined material. One male specimen (Figs.

1 , 2) (carapace length 49 mm; carapace width 42

mm, including rostral and longest lateral spines)

found in a gillnet set near the coast of Elba Island

(Figurian Sea) at a depth of 40-50 m, May 2008

(CF). One male specimen (carapace length 98 mm;

carapace width 76 mm, including rostral and lon-

gest lateral spines) trawled on deep circalittoral bot-

toms at Fa Maddalena Island, October 2010 (CC).

Remarks. The first specimen is a subadult

male; the similar M. squinado (Herbst, 1788) is

fairly common in the collecting area. While adult

specimens of each species can be easily distin-

guished, the difference between subadult specimens

is thinner. A visible difference between the speci-

men of M. goltziana and a subadult of M. squinado

is the central series of five large spines, which inM

squinado are much smaller and shaped as tubercles.

However, a remarkable affinity with M. squinado

consists in the absence of the strong antero-dorsal

spine on the meri of the pereiopods, as it is de-

282

Joachim Langeneck & Davide Di Franco

scribed by Holthuis & Gottlieb (1958). A compari-

son between the second right pereiopod of both

species shows that the legs of M. squinado (Fig. 3)

and of sub-adult specimen of M. goltziana (Fig. 4)

are, in fact, more similar than the legs of this last

one compared to the adult of M. goltziana (Fig. 5).

The absence of the spine on the meri in young spec-

imens has been confirmed by further observations

on eastern Mediterranean specimens (Galil, pers.

comm.), but this characteristic was not highlighted

in literature so far; it is remarkable that the larger

specimen, caught at La Maddalena Island, shows

well developed spines on the pereiopods.

Xaiva biguttata (Risso, 1816)

Examined material. One male specimen (Fig.

6) (carapace length 8 mm; carapace width 7 mm)

found stranded on the Is Arenas beach (Cuglieri,

Western Sardinia) (CL) (August 2008). One male

specimen (Fig. 7) (carapace length 9 mm; carapace

width 9 mm) collected alive on shell grit at a depth

of 1.7 m (Cannizzaro, Catania, July 2008, Eastern

Sicily) (CD).

RESULTS AND DISCUSSION

M. goltziana is a widespread species, and it is

considered quite common in the Eastern Atlantic

Ocean (Manning & Holthuis, 1981; D'Udelcem

DAcoz, 1999; Henriksen, 2009) and in the Eastern

Mediterranean Sea (Holthuis & Gottlieb, 1958;

Artiiz, 2006; Lelli et al., 2008; Ate§ et al., 2010).

The first record of the species in the Mediterranean

Sea dates back to the late 50's off the Israel coast

(Holthuis & Gottlieb, 1958); in the Italian seas the

species has been recorded first in the Ionian Sea

(Pastore, 1983) and in the Sicilian Straight (Pipitone

& Arculeo, 2003), while just recently it was recor-

ded in the Adriatic Sea (Pallaoro & Dulcic, 2004),

in the Tyrrhenian Sea (Soppelsa et al., 2005; Cro-

cetta, 2007) and in the Ligurian Sea (Vignoli et al.,

2004). Therefore, the new records do not widen the

known distribution of the species; however, they

confirm the presence of M. goltziana in the Western

Tyrrhenian Sea and in the Ligurian Sea (Fig. 8).

M. goltziana is an eurybathic species with sub-

tropical affinity, and has been first recorded in the

warmest Mediterranean sectors; its spread in central

and Northern Mediterranean Sea is considered as a

consequence of climatic changes (Vignoli et al.,

2004; Soppelsa et al., 2005). The species seems to

have spread either geographically (in the Adriatic

and Tyrrhenian Sea) and numerically (Lelli et

al., 2007, recorded more than 50 specimens in

Lebanese waters, whereas every previous record

concerned at most two or three specimens). There-

fore, the hypothesis of the climatic changes conse-

quence appears to be persuasive.

X biguttata is known in the Eastern Atlantic

Ocean (Manning & Holthuis, 1981; D'Udelcem

D'Acoz, 1999; Vieira & Morato, 2001) and in the

Mediterranean Sea, where it was recorded along

the Spanish coast (Garcia Raso & Jimenez Millan,

1981), in the Eastern Mediterranean Sea (Lewin-

sohn & Holthuis, 1986; Ate§ et al., 2010) and

along the Italian coasts (Pastore, 1977; Pipitone

& Arculeo, 2003; Bedini, 2004). The Italian

records refer to the southern Tyrrhenian Sea, (Pi-

pitone & Arculeo, 2003) and the Ionian Sea (Pa-

store, 1977); recently Bedini recorded one

subadult female (carapace length 7 mm; carapace

width 6 mm) on the Tuscan coast on Posidonia

meadow (Bedini, 2004 and pers. comm.). The

species has never been officially reported from

the western Sardinian coast before. The record

from Cannizzaro (Catania) confirms the presence

of the species in the Ionian Sea (Fig. 9). X bigut-

tata has been recorded throughout the Mediter-

ranean Sea (Lewinsohn & Holthuis, 1986), but

with quite a few specimens for each record; there-

fore, the species is generally considered rare in

the Mediterranean Sea (Garcia Raso & Jimenez

Millan, 1981; Bedini, 2004), while it is consid-

ered more common in the Atlantic Ocean (Vieira

& Morato, 2001). In the Mediterranean Sea the

abundance of X biguttata is probably higher than

what is commonly considered; however, it is hard

to find this species as fishing by-catch because of

its littoral habitat and small size.

ACKNOWLEDGEMENTS

We wish to thank Prof. R. Bedini, Dr. V. Vi-

gnoli for providing data, and especially Dr. F. Cro-

cetta for data and suggestions.

Further records of two uncommon Crustaceans in Italian seas: Maja goltziana and Xaiva biguttata

283

Figure 1. Maja goltziana, Rio Marina, Elba Island (Ligurian Sea). Figure 2. Idem, particular of the dorsal spines. Figure 3.

Second right pereiopod of subadult M. squinado. Figure 4. Second right pereiopod of subadult M. goltziana. Figure 5. Second

right pereiopod of M. goltziana (photo by courtesy of Dr. V. Vignoli). Figure 6. Xaiva biguttata, ecdysis from Western Sardinia,

in frontal and ventral view. Figure 7. X biguttata, specimen from Cannizzaro (Eastern Sicily), in frontal and ventral view.

284

Joachim Langeneck & Davide Di Franco

!z X

r\ C LT" ' • v S

r ' C ,j\ x

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Figure 8. Map showing the

known records of Maja

goltziana in the Italian seas.

The current record are repre-

sented by stars. 2: two speci-

mens (Vignoli et al., 2003). 3:

three specimens (Pallaoro &

Dulcic, 2004).

Figure 9. Map showing the

known records of Xaiva bi-

guttata in Italian seas. The

current records are represen-

ted by stars.

REFERENCES

Boccone Artuz M.L., 2006. First record of Maja

goltziana Oliveira, 1888 in Saros Bay, north Ageis.

Zoonatantia, 33-34.

Ate§ A.S., Kocata§ A. & Ozcan T., 2010. An updated list

of decapod crustaceans on the Turkish coast with a

new record of the Mediterranean shrimp, Processa

acutirostris Nouvel and Holthuis 1957 (Caridea,

Processidae). North-Western Journal of Zoology, 6:

209-217.

Bedini R., 2004. Gli animali delle praterie a Posidonia

oceanica : dai macroinvertebrati ai pesci. Bandecchi

& Vivaldi, Pontedera, 543 pp.

Crocetta F., 2007. First record of Maja goltziana D’Oli-

veira, 1888 (Decapoda, Brachyura, Majidae) in the

western Tyrrhenian Sea. Crustaceana, 80: 631-633.

D’Udekem D’Acoz C., 1999. Inventaire et distribution

des crustaces decapodes de TAtlantique nord-orien-

tale, de la Mediterranee et des eaux continentales

adjacentes au nord de 25°N. Museum National d’Hi-

stoire Naturelle, Collection Patrimoine Naturels 40,

Paris, 383 pp.

Garcia Raso J.E. & Jimenez Millan F., 1981. Algunas

species de decapodos (Crust.) del litoral malagueno.

Boletin de la Asociacion Espanola de Entomologia,

4: 19-24.

Henriksen C.S., 2009. Investigation of crustacean from

shelf areas in the Gulf of Guinea, with special em-

phasis on Brachyura. Master thesis, University of

Bergen, Norway, 71 pp.

Holthuis L.B. & Gottlieb E., 1958. An annotated list of

the decapod Crustacea of the Mediterranean coast of

Israel, with an appendix listing the Decapoda of the

eastern Mediterranean. Bulletin of Research Council

of Israel, 7B: 1-126.

Lewinsohn Ch. & Holthuis L.B., 1986. The Crustacea

Decapoda of Cyprus. Zoologische Verhandelingen,

230: 1-64.

Lelli S., Carpentieri P,, Colloca F. & Ardizzone G.D., 2008.

The spiny spider crab Maja goltziana (Crustacea:

Majidae) in south Lebanese waters. Marine Biodiver-

sity Records, 1, e83 doi: 10. 10 17/S 1755267207008676.

Manning R.B. & Holthuis L.B., 1981. West African bra-

chyuran crabs. (Crustacea: Decapoda). Smithson

Contributions to Zoology, 306: 1-379.

Pallaoro A. & Dulcic J., 2004. On the occurrence of the

spider crab Maja goltziana (Crustacea: Majidae) an

alien species in the Adriatic Sea. Journal of the Ma-

rine Biological Association of the United Kingdom,

84: 1007-1008.

Pastore M., 1977. Presenza di Thia scutellata (Fabricius)

e Xaiva biguttata (Risso) nel Golfo di Taranto (Mar

Jonio). Thalassia Salentina, 7: 83-90.

Pastore M., 1983. An Oxyrhynch crab new to the Ionian

Sea: Maja goltziana D’Oliveira, 1888 (Decapoda:

Brachyura). Crustaceana, 45: 232-237.

Pipitone C. & Arculeo M., 2003. The marine Crustacea

Decapoda of Sicily (central Mediterranean Sea): a

checklist with remarks on their distribution. Italian

Journal of Zoology, 70: 69-78.

Soppelsa O., Crocetta F. & Pipitone C., 2005. Maja gol-

tziana D’Oliveira, 1888 (Decapoda: Brachyura,

Majidae) in the Southern Tyrrhenian Sea. Crusta-

ceana, 78: 121-124.

Vieira C.M. & Morato T., 2001. First record of the crabs

Pirimela denticulata (Montagu 1808) and Xaiva

biguttata (Risso, 1816) (Crustacea: Decapoda) from

the Azores. Life and Marine Sciences, 18 A: 89-91.

Vignoli V., Caruso T. & Falciai L., 2004. Decapoda

Brachyura from Monte Argentario (Mediterranean

Sea, central Tyrrhenian). Crustaceana, 77: 177-186.

Biodiversity Journal, 2013, 4 (2): 285-300

No end in sight? Further new records of amphibians and rep-

tiles from Phong Nha - Ke Bang National Park, Quang Binh

Province, Vietnam

Vinh Quang Luu 1,5,6 , Truong Quang Nguyen 2,6 , CuongThe Pham 2 , Kien Ngoc Dang 3 , Thanh NgocVu 4 , Sladjana

Miskovic 5 , Michael Bonkowski 6 & Thomas Ziegler 5,6 *

'Department of Wildlife, Faculty of Natural Resource and Environmental Management, Vietnam Forestry University, Xuan Mai,

Chuong My, Fla Noi, Vietnam; e-mail: qvinhfuv@yahoo.com.au

institute of Ecology and Biological Resources, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet, Hanoi, Viet-

nam; e-mail: nqt2@yahoo.com

Thong Nha - Ke Bang National Park, Bo Trach District, Quang Binh Province, Vietnam

4 Faculty of Biology, Vietnam National University, Hanoi, University of Science, Department of Vertebrate Zoology, Zoological

Museum, 334 Nguyen Trai Str., Thanh Xuan, Hanoi, Vietnam; e-mail: thanhdouc@gmail.com

5 AG Zoologischer Garten Koln, Riehler Strasse 173, D-50735 Cologne, Germany; e-mail: ziegler@koelnerzoo.de

"Zoological Institute, University of Cologne, Ziilpicher Strasse 47b, D-50674 Cologne, Germany

’Corresponding author

ABSTRACT We report 1 1 new records of amphibian and reptile species and subspecies on the basis of

newly collected specimens from the UNESCO World Heritage Site Phong Nha - Ke Bang

National Park, Quang Binh Province, Vietnam: Ingerophrynus macrotis, Limnonectes

gyldenstolpei, Babina chapaensis, Theloderma corticate, T. stellatum, Scincella rufocaudata,

Oligodon cinereus pallidocinctus, Parahelicops annamensis, Rhynchophis boulengeri,

Sinomicrurus macclellandii and Protobothrops mucrosquamatus. The record of T. corticate

from Phong Nha - Ke Bang National Park is the southernmost record generally known, that

of T. stellatum the northernmost record within Vietnam, and, most remarkably, the finding

of Limnonectes gyldenstolpei represents the first country record for Vietnam. In addition,

we report the second known specimen and the first adult male of Sphenomorphus tetradacty-

lus, a species recently described based on a single female only. At time, 151 species of am-

phibians and reptiles are known from Phong Nha - Ke Bang National Park, including 50

species of amphibians, 12 species of turtles, 31 species of lizards, and 58 species of snakes.

In addition, an updated list of the local herpetofauna is provided, including recent taxonomic

or nomenclatural changes.

KEY WORDS herpetofauna; taxonomy; distribution; limestone habitat; Truong Son Mountain Range.

Received 7.04.2013; accepted 24.05.2013; printed 30.06.2013

INTRODUCTION

Phong Nha - Ke Bang National Park in the

Truong Son Mountain Range in central Vietnam is

known as one of the country’s most famous pro-

tected areas in terms of herpetofaunal diversity. Zie-

gler & Herrmann (2000) published a first,

preliminary list of the karst forest area's herpetodi-

versity, based on own fieldwork, collection-based

research and first Vietnamese reports, comprising

96 amphibian and reptile species. Four years later,

the total number of amphibian and reptilian species

known from the area as a result of further field work

was increased to 128 species (Ziegler et al., 2004).

286

Vinh Quang Luu etalii

In a third updated herpetofaunal list for the area,

Ziegler et al. (2006) reported of 140 amphibian and

reptilian species, demonstrating a still ongoing in-

crease in new species records. Ziegler et al. (2007)

published a comprehensive review of the snake di-

versity of Phong Nha - Ke Bang including nine for-

merly not yet recorded species, thus increasing the

total number of snakes known from Phong Nha -

Ke Bang to 59 species. Hendrix et al. (2008) pro-

vided an updated anuran list comprising 47 taxa

recorded for the Phong Nha - Ke Bang National

Park, in which five species were recorded for the

first time from that karst forest area. Recently, Zie-

gler & Vu (2009) published an updated checklist of

the amphibians and reptiles from Phong Nha - Ke

Bang National Park with a total of 138 species, in-

cluding 45 species of amphibians and 93 species of

reptiles. The total number of recorded amphibians

and reptiles has decreased in this overview, because

several doubtful or unconfirmed records, e.g., men-

tioned in Ziegler et al. (2007) and Hendrix et al.

(2008), were removed from the list provided by Zie-

gler & Vu (2009), viz. Bombina maxima (Boulen-

ger, 1905), Eutropis chapaensis (Bourret, 1937),

Scincella rupicola (Smith, 1916), Sphenomorphus

buenloicus (Darevslcy et Nguyen, 1983), and Ma-

layemys subtrijuga (Schlegel et Muller, 1844). In

the years 2010 and 2011, additional herpetological

field surveys were conducted in Phong Nha - Ke

Bang National Park and extension area which re-

vealed the existence of a number of so far not re-

ported amphibian and reptile species. In addition to

the eleven new herpetofaunal records for Phong

Nha - Ke Bang National Park we provide an up-

dated list of amphibians and reptiles occurring in

this area.

MATERIALS AND METHODS

Field surveys were conducted in the Phong Nha

- Ke Bang National Park area by Thomas Ziegler,

Thanh Ngoc Vu, KienNgoc Dang, and Sladjana Mi-

skovic (TZ and others) from June to July 2010, as

well as from 12th July to 2nd August, and from 12th

September to 1st October 2011 by Truong Quang

Nguyen, Cuong The Pham, Dai Van Nguyen, Hang

Thi An, and Kien Ngoc Dang (TQN and others)

(Figs. 1, 2). After taking photographs, specimens

were anaesthetized, fixed in 40-70% ethanol (am-

Figure 1. Map showing the location of Phong Nha - Ke

Bang National Park, Quang Binh Province, Vietnam.

phibians) or 80-85% ethanol (reptiles) and subse-

quently stored in 70% ethanol. Measurements were

taken with a digital calliper to the nearest 0.1 mm.

ABBREVIATIONS, a.s.l. = above sea level; SVL

(snout- vent length) = from tip of snout to anterior

margin of cloacal; TaL = tail length, from posterior

margin of cloacal to tip of tail; tenninology of mor-

phological characters followed Nguyen et al. (2012)

for amphibians and anuran webbing formula followed

Glaw & Vences (2007) (except for forearm length,

FAL = from axilla to elbow and hand length, HAL =

from base of outer palmar tubercle to tip of finger III),

Phung & Ziegler (2011) for lizards, and David et al.

(2012) for snakes. Specimens were deposited in the

collections of the Institute of Ecology and Biological

Resources (IEBR), Vietnam Academy of Science and

Technology Hanoi, Vietnam, Phong Nha - Ke Bang

National Park (PNKB), Quang Binh Province, Viet-

nam, and the Zoologisches Forschungsmuseum

Alexander Koenig (ZFMK), Bonn, Germany.

Further new records of amphibians and reptiles from Phong Nha - Ke Bang National Park, Quang Binh Province, Vietnam 287

RESULTS

BUFONIDAE

Ingerophrynus macrotis (Boulenger, 1887)

Big-eyed Toad / Coe tai to (Fig. 3)

Examined material. One juvenile ZFMK

94263 collected by TZ and others during night time

in June 2010 in the leaf litter of karst forest near

Tam Co Cave, Tan Trach Commune, Bo Trach Dis-

trict and one adult female IEBR A.2013.7 (SVL:

58.7 mm) collected on 15 July 2011 by TQN and

others in Da Lat forest, Thuong Hoa Commune,

MinhHoa District (17°40.124’N, 105°55.031’E, at

an elevation of 3 12 m a.s.l.).

Morphological characters. SVL of adult fe-

male 58.7 mm; head wider than long (HW 20.3 mm,

HL 18.7 mm); snout truncate, protruding, longer

than horizontal diameter of eye (SL 6.6 mm, ED 5.9

mm); canthus rostralis rounded, loreal region con-

cave; interorbital distance wider than intemarial dis-

tance (IOD 4.7 mm, IND 4.1 mm); nostril closer to

the tip of snout than to the eye (SN 1.7 mm; EN 3.6

mm); pupil rounded; tympanum distinct (TD 3.2

mm); parotoid gland 11.1 mm; vomerine teeth ab-

sent; tongue rounded posteriorly. Forelimbs: FAL

15.4 mm, HAL 30.1 mm; fingers free of webbing,

relative length of fingers: II<I V <I<III ; tips of fingers

rounded, without discs; dermal fringe along outer

finger absent; palmar tubercles distinct. Hindlimbs:

femur longer than tibia and foot length (FML 26.0

mm, TBL 24.4 mm, FTL 23.2 mm); toes long and

thin, relative length of toes: I<II<V<III<IV; tips of

toes rounded; webbing basal; tarsal fold absent, der-

mal fringe along outer toe absent; subarticular tu-

bercles present; inner metatarsal tubercle present

(IMT 2.4 mm), outer metatarsal tubercle small

(OMT 1.8 mm). Dorsal skin of body covered with

tubercles or warts in different sizes, those on head

smallest, few enlarged tubercles around vent.

Coloration in preservative: dorsal skin greyish

brown with some symmetrical darker markings on

snout and interorbital region; a V-shaped conversely

mark in front of shoulder and dark spots near mid-

dle of back; dorsal tubercles on body, limbs grey to

yellow; tympanum dark grey; upper lip with yellow

and grey flecks. Ventral skin and concealed parts of

limbs dirty greyish-yellow (determination after

Inger et al., 1999).

Distribution. In Vietnam, /. macrotis has been

recorded from Thanh Hoa, Ha Tinh, Thua Thien-

Hue, Da Nang, Quang Nam, Dak Lak, Lam Dong,

and Dong Nai provinces. Our finding represents the

first record for Phong Nha - Ke Bang National Park

and for Quang Binh Province. Elsewhere, this

species is known from northeastern India, Myan-

mar, Laos, Thailand, Cambodia, and Malaysia

(Nguyen et al., 2009; Frost, 2013).

DICROGLOSSIDAE

Limnonectes gyldenstolpei (Andersson, 1916)

Gyldenstolpe’s Frog / Ech gin-den-x-ton-pi (Fig. 4)

Examined material. One adult male IEBR

A. 20 13. 8 collected on 14 September 2011 in the

evergreen forest, Hoa Son Commune, Minh Hoa

District (17°42.166’N, 105°47.957’E, at an eleva-

tion of 449 m a.s.l.)

Morphological characters. SVL 69.6 mm;

head longer than wide (HL 35.2 mm, HW 34.4

mm); snout longer than horizontal diameter of eye

(SL 12.7 mm; ED 7.2 mm); canthus rostralis

rounded; nostril directed laterally; loreal region

concave; interorbital distance broader than intemar-

ial distance (IOD 8.5 mm, IND 7.3 mm); nostril

closer to the tip of snout than to the eye (EN 7.1

mm; SN 4.7 mm); tympanum rounded, longer than

tympanum-eye distance (TD 7.4 mm, TEY 6.4

mm); vomerine teeth present; tongue notched pos-

teriorly; vocal sac indistinct in males. Forelimbs:

FAL 13 mm, HAL 32 mm; fingers free of webbing,

relative length of fingers: II<IV<I<III; tips of fin-

gers rounded, without discs; dermal fringe along

outer finger absent; palmar tubercles present; nup-

tial pad present in males. Hindlimbs: femur shorter

than tibia and foot length (FML 32.7 mm, TBL 36.7

mm, FTL 34.7 mm); toes long and thin, relative

length of toes: I<II<V<III<IV; tips of toes rounded;

webbing formula: Io(0)-(l)iIIo(0)-(l)iIIIo(l/2)-

(l)iIVo(2)-(0) iV; dermal fringe along outer toe ab-

sent; subarticular tubercles present; inner metatarsal

tubercle present (IMT 4.3 mm), outer metatarsal tu-

bercle indistinct. Dorsal skin of head smooth with

a swollen flap (10.6 mm length and 10.2 mm

width); dorsal surface of forelimbs, thigh and tarsus

smooth; supratympanic fold distinct, from eye to

shoulder; ventral surface smooth.

288

Vinh Quang Luu etalii

Coloration in preservative: head grey with a

cream broad stripe between eyes; dorsum brown;

dark dorsolateral spots; dark bars present on upper

surface of forelimbs, tibia and thigh; venter cream

with dark spots on throat (determination after Tay-

lor, 1962; Ohler & Dubois, 1999).

Distribution. The specimen from Phong Nha -

Ke Bang National Park represents the first country

record of Limnonectes gyldenstolpei for Vietnam.

Elsewhere, this species is known from Laos, Cam-

bodia, and Thailand (Frost, 2013).

Remarks. The male specimen differs from the

juvenile female in the description of Ohler & Du-

bois (1999) by having the tibia longer than femur

(TBL 36.7 mm, FML 32.7 mm versus TBL 11.6

mm, FML 11.7 mm).

RANIDAE

Babina chapaensis (Bourret, 1937)

Chapa Frog / Chang sa pa (Fig. 5)

Examined material. Two adult males IEBR

A.2013.9 and ZFMK 94258, one adult female

IEBR A.2013.10 collected on 14 September 2011

and one adult male ZFMK 94259 collected by

TQN and others on 16 September 2011 in the forest

near Cha Lo Village, Hoa Son Commune, Minh

Hoa District (17°42.213’N, 105°47.748’E, at an

elevation of 570 m a.s.l.).

Morphological characters. SVL 43.0-46.0

mm in males (mean ± SE 44.2 ± 1.6, N = 3), 56.9

mm in the female; head longer than wide (HL 17.3-

20.4 mm, HW 14.9-18.0 mm); snout longer than

horizontal diameter of eye (SL 6. 9-7. 8 mm; ED 4.7-

5.6 mm); canthus rostralis rounded; nostril directed

laterally; loreal region concave; interorbital distance

narrower than internarial distance (IOD 3. 3-4. 5

mm, IND 5. 1-6.3 mm); nostril closer to the eye than

to the tip of snout (EN 2. 9-4.0 mm; SN 3. 5-4. 2

mm); tympanum rounded, longer than tympanum-

eye distance (TD 3. 9-4. 2 mm, TEY 0.3- 1.4 mm);

vomerine teeth present; tongue notched posteriorly;

vocal sac present in males. Forelimbs: FAL 8.8-10.3

mm, HAL 20.2-23.7 mm; fingers free of webbing,

relative length of fingers: II<I=IV<III; tips of fin-

gers rounded, without discs; dermal fringe along

outer finger absent; palmar tubercles distinct; nup-

tial pad present in males. Hindlimbs: femur shorter

than tibia and foot length (FML 21.4-25.2 mm, TBL

24.5-29.5 mm, FTL 24.1-29.2 mm); toes long and

thin, relative length of toes: I<H<V<III<IV; tips of toes

rounded; webbing formula: Io(l)-(l)iIIo(l/2)-

(2)iIIIo(l)-(2)iIVo(2)-(l/2)iV; dermal fringe along

outer toe absent; subarticular tubercles present;

inner metatarsal tubercle present (IMT 2. 2-2. 5 mm),

outer metatarsal tubercle indistinct.

Dorsal surface of head and dorsum smooth; dor-

solateral fold distinct; lateral sides smooth; a small

fold present along arm; ventral surface smooth. Col-

oration in preservative: head and dorsum light

brown with a cream vertebral stripe, edged in dark

brown, running from behind the eye to vent; poste-

rior part of dorsum with some dark spots; upper jaw

with a cream stripe, from below the nostril to axilla;

dorsolateral fold yellowish brown, edged in black

laterally; upper surface of tibia and thigh with some

dark bars; venter cream (determination after Bourret,

1942; Chuaynkem et al., 2010).

Distribution. In Vietnam, B. chapaensis has been

recorded from Lao Cai, Bac Giang, Ha Tinh, Kon

Turn, Gia Lai, and Dak Lak provinces (Nguyen et al.,

2009). This is the first record of this species from

Phong Nha - Ke Bang National Park as well as from

Quang Binh Province. Elsewhere, this species is

known from Laos and Thailand (Nguyen et al., 2009).

RHACOPHORIDAE

Theloderma corticale (Boulenger, 1903)

Tonkin Bug-eyed Frog / Ech cay san bac bo (Fig. 6)

Examined material. Three adult males IEBR

A.2013.11, ZFMK 94262 collected on 2 1 July 2011

by TQN and others in Da Lat forest, Thuong Hoa

Commune, Minh Hoa District (17°39.032’N,

105°54.774’E, at an elevation of 516 m a.s.l.) and

PN-KB 2011.204 collected by TQN and others on

28 July 2011 in Cha Noi forest, Xuan Trach Com-

mune, Bo Trach District (17°37.758’N,

106°05.893’E, at an elevation of 470 m a.s.l.).

Morphological characters. SVL of males

61.3-69.7 mm; head wider than long (HL 24.1-27.5

mm, HW 26.3-28.6 mm); snout longer than hori-

zontal diameter of eye (SL 9.0-10.0 mm; ED 6.2

mm); canthus rostralis rounded; loreal region con-

cave; interorbital distance wider than internarial dis-

Further new records of amphibians and reptiles from Phong Nha - Ke Bang National Park, Quang Binh Province, Vietnam 289

Figure 2. Karst forest in Thung Hoa Commune, extension area of Phong Nha - Ke Bang National Park. Figure 3. Big-eyed

Toad, lngerophynus macrotis. Figure 4. Gyldenstolpe’s Frog, Limnonectes gyldenstolpei. Figure 5. Chapa Frog Babina

chapaensis. Figure 6. Tonkin Bug-eyed Frog, Theloderma corticale. Figure 7. Taylor’s Bug-eyed Frog, Theloderma stel-

latum. Photos: T. Q. Nguyen & T. Ziegler.

290

Vinh Quang Luu etalii

tance (IOD 6. 1-6.5 mm, IND 3. 7-4. 4 mm); nostril

closer to tip of snout than to eye (SN 2.5 mm; EN

7. 1-8.3 mm); vocal sac absent; tympanum oval,

greater than tympanum-eye distance (TD 4. 0-4. 9

mm, TEY 3. 3-3. 9 mm); vomerine teeth present;

tongue notched posteriorly. Forelimbs: FAL 12.2-

14.8 mm, HAL 33.2-34.7 mm; relative length of fin-

gers: I<II<IV<III; tips of fingers and toes enlarged

into round discs; webbing present at base of fingers

III and IV; dermal fringe along outer finger present;

palmar tubercles distinct; nuptial pad present.

Hindlimbs: tibia longer than femur and foot length

(TBL 31.8-34.0 mm, FML 25.5-32.2 mm, FTL

27.8-30.5 mm); relative length of toes:

I<II<III<V<I V; webbing formula: Io(0)-(0)iIIo(0)-

(0)iIIIo(0)-(l)iIVo(l)-(0)iV; dermal fringe along

outer toe absent; subarticular tubercles present;

inner metatarsal tubercle present (IMT 3. 1-4.9

mm); outer metatarsal tubercle absent.

Dorsal surface of body covered with tubercles

or warts of different sizes, those on head and back

biggest; ventral skin with small tubercles. Col-

oration in preservative: dorsal colour olive-green

marbled with red-brown spots; dark brown bars

present on upper surface of fore and hind limbs;

ventral surface greyish yellow (determination after

Inger et al., 1999; Orlov et al., 2006).

Distribution. I corticale is currently known

only from northern Vietnam: Ha Giang, Tuyen

Quang, Cao Bang, Lang Son, Vinh Phuc, and Son

La provinces (Nguyen et al., 2009; Frost, 2013).

This is a new record for Phong Nha - Ke Bang Na-

tional Park and for Quang Binh Province as well as

the southernmost known record of the species.

Theloderma stellatum Taylor, 1962

Taylor’s Bug-eyed Frog / Ech cay san tay-lo (Fig. 7)

Examined material. One adult female ZFMK

94261 collected by TZ and others during night time

on a tree trunk nearby a forest stream and one adult

male IEBR A.20 1 3 . 1 2 collected by TQN and others

on 28 July 2011 in Cha Noi forest, Xuan Trach

Commune, Bo Trach District (17°37.649’N,

106°05.806’E, at an elevation of 517 m a.s.l.).

Morphological characters. SVL 32.7-35.0

mm; head as long as wide (HL 13.5-13.9 mm, HW

13.6-14.0 mm); snout longer than horizontal diame-

ter of eye (SL 5. 3-5. 4 mm; ED 4. 5-4. 7 mm); can-

thus rostralis rounded; loreal region concave;

interorbital distance wider than intemarial distance

(IOD 3. 6-3. 9 mm, IND 2.0-2. 1 mm); nostril closer

to tip of snout than to eye (SN 1.6- 1.7 mm; EN 3.2-

4.0 mm); tympanum rounded, greater than tympa-

num-eye distance (TD 2. 2-2. 9 mm, TEY 1.0 mm);

vomerine teeth present; tongue notched posteriorly.

Forelimbs: FAL 7. 9-8.4 mm, HAL 17.5-17.9 mm;

relative length of fingers: I<II<IV<III; tip of fingers

and toes enlarged into large discs; webbing basal;

dermal fringe along outer finger absent; palmar tu-

bercles indistinct; nuptial pad present. Hindlimbs:

tibia longer than femur and foot length (TBL 17.6-

18.3 mm, FML 16.6-16.9 mm, FTL 15.6-16.0 mm);

relative length of toes: I<II<III=V<IV; webbing for-

mula: Io( 1 )-( 1 )iIIo( 1 /2)-(2)iIIIo( 1 )-(2)iI Vo(2)-( 1/2)

iV; dermal fringe along outer toe absent; subarticu-

lar tubercles present; inner metatarsal tubercle

present (IMT 1.3- 1.5 mm).

Dorsal skin of head and body, upper surface of

fore-arm, tibia, and tarsus, with tubercles covered

in whitish granular asperities; ventral s kin smooth.

Coloration in preservative: dorsal head and body

brownish with cream speckles; black spots present

on snout and black marking present between shoul-

ders; upper surface of thigh, tibia, tarsus, and foot

with transverse dark bars; discs pinkish; chin and

venter dark brown with light flecks (determination

after Taylor, 1962; Inger et al., 1999; Orlov et al.,

2006; Nguyen & Nguyen, 2008).

Distribution. In Vietnam, T. stellatum has been

recorded from Kon Turn, Gia Lai, Dak Lak, Dong

Nai, and Kien Giang provinces (Nguyen et al.,

2009). This is the first record of the species from

Phong Nha - Ke Bang National Park as well as from

Quang Binh Province, which at the same time is

the northernmost country record. Elsewhere, this

species is known from Thailand, southern Laos and

Cambodia (Nguyen et al., 2009).

SCINCIDAE

Scincella rufocaudata (Darevsky et Nguyen, 1983)

Red-tailed ground skink / Than lan phe-no duoi do

(Fig. 8)

Examined material. One adult male IEBR

A.20 13. 13 collected on 21 July 2011 by TQN and

others in Da Lat forest, Thuong Hoa Commune,

MinhHoa District (17°39.665’N, 105°55.800’E, at

Further new records of amphibians and reptiles from Phong Nha - Ke Bang National Park, Quang Binh Province, Vietnam 291

an elevation of 448 m a.s.l.), 2 adult females IEBR

A. 20 13. 14 collected on 29 July 2011 by TQN and

others in Khe Ma forest, Thuong Hoa Commune,

MinhHoa District (17°3 9. 41 0’N, 106°03.592’E, at

an elevation of 300 m a.s.l.), and ZFMK 94256 col-

lected on 20 July 2011 by TQN and others in Hoa

Son Commune, Minh Hoa District (17°39.120’N,

105°59.678’E, at an elevation of 250 m a.s.l.) and

one juvenile ZFMK 94257 collected on 18 July

20 1 1 by TQN and others in Thuong Hoa Commune,

Minh Hoa District (17°40.057’N, 105°56.049’E, at

an elevation of 513 m a.s.l.).

Morphological characters. SVL 45.5 mm in

the male, 45.9-47.2 mm in females; TaL 51.0 mm

in one female, regenerated or lost in others; snout

obtuse; rostral wider than long, nostril in the nasal;

frontonasal large; prefrontals separated from each

other; frontal longer than frontoparietal; parietals

large, in contact posteriorly; nuchals absent; supraoc-

ulars 4; supraciliaries 8; loreals 2; supralabials 7;

temporals 2 + 2, upper overlapped by lower one; in-

fralabials 6; ear-opening oval; lower eyelid with a

transparent window; midbody scales in 30-34 rows,

smooth; paravertebral scales 67-69; ventral scales

61-66; enlarged precloacals 2; limbs pentadactyl;

subdigital lamellae under fourth toe 15-20.

Coloration in preservative: dorsal head and body

brown with a row of dark vertebral spots, a black

stripe present on upper part of the side; upper sur-

face of tail reddish; venter cream (determination

after Stuart & Emmett, 2006; Nguyen et al., 2011a).

Distribution. In Vietnam, S. rufocaudata is

known from Thua Thien - Hue, Quang Nam, Kon

Turn, Gia Lai, Dak Lak, and Ba Ria - Vung Tau prov-

inces (Nguyen et al., 2009). Records of this species

in northern Vietnam (Bac Kan, Thai Nguyen and

Vinh Phuc provinces) were reidentified as S. tonki-

nensis by Nguyen et al. (2011a). Elsewhere, this

species is recorded from Laos and Cambodia

(Nguyen et al., 2009).

Remarks. The red-tailed ground skink was trans-

ferred from the genus Sphenomorphus to Scincella

by Nguyen et al. (2011a).

Sphenomorphus tetradactylus (Darevsky et

Orlov, 2005)

Four-fingered skink / Than lan phe-no bon ngon

(Fig. 9)

Examined material. One adult male IEBR

A.2013.15 collected on 17 July 2011 by TQN and

others in Thuong Hoa Commune, Minh Hoa Dis-

trict (17°40.057’N, 105°56.049’E, at an elevation

of 513 m a.s.l.).

Morphological characters. Size small, SVL

36 mm, TaL 49.9 mm, tail tip lost; rostral wider than

high, in contact with frontonasal; frontonasal broa-

der than long; prefrontals small, separated from

frontal; supraoculars 4/4, first two in contact with

frontal on each side; loreal single; lower eyelid

scaly; external ear openings hidden with slightly re-

cessed auricular depression; supraciliaries 7/7, first

in contact with frontal; supralabials 6/6, third to fifth

Figure 8. Red-tailed Ground Skink Scincella rufocaudata. Figure 9. Four-fingered Skink Sphenomorphus tetradactylus.

Photos: K. N. Dang & T. Q. Nguyen.

292

Vinh Quang Luu etalii

below the eye; infralabials 7/7; nuchals 2/3; mid-

body scale rows 20; paravertebral scales 48; ventral

scales 51; enlarged precloacals 2; limbs very short,

forelimb tetradactyl, first shortest; hind limbs pen-

tadactyl; subdigital lamellae under fourth toe 9/10.

Coloration in preservative: dorsal head and body

brown with longitudinal dark shapes along dorsum;

venter cream with brown spots; legs dark above with

indistinct black marks; free margins of upper and

lower eyelids not edged in white (determination after

Darevsky & Orlov, 2005; Nguyen et al., 2011a).

Distribution. S. tetradactylus is currently

known only from Phong Nha - Ke Bang National

Park, Quang Binh Province, Vietnam (Nguyen et

al., 2011a).

Remarks. S. tetradactylus was originally de-

scribed by Darevsky & Orlov (2005) as a member

of the genus Leptoseps based on the holotype

collected from Phong Nha - Ke Bang National Park.

However, Nguyen et al. (2011a) removed this spe-

cies from the genus Leptoseps to the genus Sphe-

nomorphus. This is the second known specimen and

the first reported adult male of the species.

COFUBRIDAE

Oligodon cinereus pallidocinctus (Bourret, 1934)

Guenther’s Kukri Snake / Ran khiem xam (Fig. 10)

Examined material. One male, PNKB S.0154,

collected by K.D. Ngoc in 2009.

Morphological characters. SVL 495 mm;

TaL 90 mm; maxillary teeth 16/16; loreal 1/1;

supralabials 8/8, fourth and fifth entering orbit;

infralabials 8/8; preoculars 2; postoculars 2; temporals

1/2+2; dorsal scale rows 17:17:15, smooth; cloacal

entire; ventral scales 3 + 167, subcaudals 42, divided.

Coloration in preservative: dorsal head with a

grey chevron; dorsal surface of body yellowish

brown with 38 grey, black-edged bands on body, 6

bands on tail; ventral surface cream with dark spots

(identification after Bourret, 1936; Smith, 1943).

Distribution. In Vietnam, this subspecies is

known from Thua Thien - Hue, Ba Ria - Vung Tau,

and Ho Chi Minh City (“Form IV” in Smith, 1943).

This is the first record of O. cinereus pallidocinctus

from Phong Nha - Ke Bang National Park, Quang

Binh Province.

Remarks. The male specimen differs from the

description of Smith (1943) in having more light

bands on body (38 versus 27-34) and more bands

on tail (6 versus 3-4).

Parahelicops annamensis Bourret, 1934

Annam Keelback / Ran binh mui trung bo (Fig. 11)

Examined material. Two males IEBR

A.20 1 3 . 1 6- A.20 13.17 collected on 23 July 20 1 1 by

TQN and others in Hoa Son Commune, Minh Hoa

District (17°42.612’N, 105°52.571’E, at an eleva-

tion of 537 m a.s.l.) and one female ZFMK 94255

collected on 16 September 2011 in Hoa Son Com-

mune, Minh Hoa District (17°42.208’N,

105°46.970’E, at an elevation of 641 m a.s.l.).

Morphological characters. SVF 358-430

mm in males, 455 mm in the female; TaF 165-177

mm in males, 102 mm in the female; head distinct

from neck; maxillary teeth 21/21; rostral flat,

broader than high; nostril in the nasal; intemasals

narrowed anteriorly; prefrontals 2/2, slightly

broader than long; frontal narrowed posteriorly,

about half the length of the parietals; loreal single,

elongated; preoculars 2/2, upper larger; postoculars

2/2; temporals 1 + 1; supralabials 8 or 9; fourth to

sixth (in one male) or fifth and sixth bordering orbit;

infralabials 10, first to fifth in contact with first chin

shield; first pair of chin shield shorter than second

pair, second pair divided by an elongated scale pos-

teriorly; body scales rows 19:17:17, median rows

strongly keeled posteriorly; dorsal scales on tail

strongly keeled, eight longitudinal keel rows at

base, decreasing to four rows distally; ventrals 167-

169; cloacal divided; subcaudals divided, 95 and

121 in two males, 51 in the female.

Coloration in preservative: dorsal head with ir-

regular brown markings; a yellow stripe present

from posterior margin of eye to neck, continuing

onto body as broken dorsolateral stripe, being less

distinct posteriorly; tail iridescent yellowish-brown;

ventral and subcaudal surface cream, outer margin

of ventrals dark brown (determination after Bourret,

1936; Stuart, 2006; Ziegler et al., 2007).

Distribution. In Vietnam, the species is only

known from Da Nang and Kon Turn provinces (see

Stuart, 2006; Nguyen et al., 2009). This is the first

record from Phong Nha - Ke Bang National Park as

well as from Quang Binh Province. Elsewhere, this

species is reported from Faos (Xe Kong) (Nguyen

et al., 2009).

Remarks. P. annamensis was previously listed

as Amphiesma sp. by Ziegler & Vu (2009).

Further new records of amphibians and reptiles from Phong Nha - Ke Bang National Park, Quang Binh Province, Vietnam 293

Rhynchophis boulengeri Mocquard, 1897

Rhinoceros snake / Ran voi (Fig. 12)

Examined material. Two males deposited in

PNKB.

Morphological characters. SVL 940-950

mm; TaL 350-360 mm; presence of a long pointed

nasal appendage covered with small scales; inter-

nasals much smaller than the prefrontals; frontal

narrowed posteriorly; loreal single, longer than

wide; preocular single; postoculars 2; temporals

2+3; supralabials 8 or 10, fourth to fifth or fifth to

seventh in contact with the eye; infralabials 11;

midbody scale rows 19, slightly keeled dorsally;

ventral scales 1 + 211 or 1 + 208; subcaudal scales

122 or 126, divided; cloacal plate divided.

Coloration in preservative: dorsum green, venter

paler; thin black stripe behind eye; interstitial skin

on the sides of the body bluish-black and white;

light stripes on subcaudal fold (determination after

Smith, 1943; Nguyen et al., 2011b).

Distribution. In Vietnam, R. boulengeri has

been recorded from the provinces of Son La, Thai

Nguyen, Vinh Phuc, Ha Noi, Quang Ninh, Hai

Phong, and Ha Tinh (Nguyen et al., 2009; Nguyen

et al., 2011b). This is a new record for Phong Nha -

Ke Bang National Park as well as for Quang Binh

Province. Elsewhere, this species is known only

from China (Nguyen et al., 2009).

ELAPIDAE

Sinomicrurus macclellandii (Reinhardt, 1844)

MacClelland’s Coral Snake/ Ran la kho thuong

(Figs. 13-14)

Morphological characters. Morphological

characters based on photographic record: vertebral

scales not larger than adjacent scales; subcaudals

divided; dorsal surface reddish brown with 3 1 black

cross-bands from behind head to tip of tail; head black

with a wide, white cross-band behind eyes; ventral

surface cream with black bands and black squarish

marks (determination after Ziegler et al., 2007).

Distribution. In Vietnam, this species is

known from Lao Cai and Cao Bang provinces in

the North southwards to Lam Dong and Dong Nai

provinces (Nguyen et al., 2009). This is the first

confirmed record for Phong Nha - Ke Bang Na-

tional Park. Elsewhere, this species is reported

from India, Nepal, Myanmar, Thailand, China,

Japan, Taiwan (Nguyen et al., 2009).

Remarks. S. macclellandii was mentioned as

unconfirmed record for Phong Nha - Ke Bang by

Ziegler et al. (2007) and thus was subsequently

removed from the herpetofaunal list of Phong Nha

-Ke Bang by Ziegler & Vu (2009).

VIPERIDAE

Protobothrops mucrosquamatus (Cantor, 1839)

Chinese Habu / Ran luc cuom (Fig. 15)

Examined material. One specimen was found

at night of 27 October 2009 by TZ and others in the

vegetation nearby a forest path in the Cha Noi

region and one male IEBR A.2013.18 (PN-KB

2011.51), collected on 16 July 2011 by TQN and

others in Thuong Hoa Commune, Minh Hoa Dis-

trict (17°40.405’N, 105°56.656’E, at an elevation

of 260 m a.s.l.).

Morphological characters. SVL 950 mm;

TaL 225 mm; head long, narrow; singe loreal pit;

supralabials 10; gular scales smooth; mental bor-

dering infralabial posteriorly; elongated subocular

scales, divided from supralabials; dorsal scales

strongly keeled, midbody scale rows 27; ventral

scales 3 + 216; subcaudals 98; cloacal undivided.

Coloration in preservative: dorsal head brown,

paler below; dorsum greyish brown, with a series

of large brown, dark-edged spots; ventral surface

cream, with light brown, light areas appearing as

squarish spots; dorsal tail light brown, with a series

of conspicuous black spots (determination after

Smith, 1943; Ziegler et al., 2007; Nguyen et al.,

2011b).

Distribution. In Vietnam, P. mucrosquamatus

has been recorded from Lao Cai, Ha Giang, Cao

Bang, Bac Kan, Lang Son, Thai Nguyen, Vinh

Phuc, Quang Ninh, Hai Phong, Hai Duong, Ha Noi,

Ninh Binh, Nghe An, Ha Tinh, Quang Tri, Thua

Thien-Hue, Kon Turn, and Gia Lai provinces

(Nguyen et al., 2009). This is the first record of the

species for Phong Nha - Ke Bang National Park and

for Quang Binh Province. Elsewhere, this species

is known from India, Bangladesh, China, Taiwan,

and Myanmar (Nguyen et al., 2009).

Remarks. The male specimen differs from the

description of Smith (1943) in having more subcau-

dals (98 versus 76-91).

294

Vinh Quang Luu etalii

Figure 10. Guenther’s Kukri Snake, Oligodon cinereus pallidocinctus. Figure 11. Annam Keelback, Parahelicops anna-

mensis. Photos: K. N. Dang & T. Q. Nguyen. Figure 12. Rhinoceros Snake, Rhynchophis boulengeri. Figures 13-14. Mac-

Clelland’s Coral Snake, Sinomicrurus macclellandii. Figure 15. Chinese Habu, Protobothrops macros quamatus . Photos:

K. N. Dang, T. Q. Nguyen & T. Ziegler.

Further new records of amphibians and reptiles from Phong Nha - Ke Bang National Park, Quang Binh Province, Vietnam 295

Check-list of amphibians and reptiles recorded

from Phong Nha - Ke Bang

Current check-list of amphibians and reptiles recorded

from Phong Nha - Ke Bang after Ziegler & Vu

(2009), including the herein listed new records (*),

additions (**) according to Ziegler et al. (2010),

Nguyen et al. (2011a), Hoang et al. (2012), and tax-

onomic reassignments (***) according to Blanck et

al. (2006), Stuart & Fritz (2008), Fritz et al. (2008,

2010), Inger & Stuart (2010), McLeod (2010), Yu

et al. (2010), David et al. (2011), Nguyen et al.

(2011b, ), Ohler et al. (2011), Rosier et al. (2011),

Kuraishi et al. (2012), Uetz (2013), Frost (2013),

Siler et al. (2013), and Patrick David (pers. comm,

to replace Amphiesma khasiense in central Vietnam

with A. boulengeri ); snake species which have been

previously listed but could not be confirmed as oc-

curring in Phong Nha - Ke Bang by Ziegler et al.

(2007) are excluded from the list: Typhlops diardi

Schlegel, 1839, Calamaria pavimentata Dumeril,

Bibron et Dumeril, 1854, C. septentrioncdis Bou-

lenger, 1890, Dendrelaphis pictns (Gmelin, 1789),

Lycodon septentrionalis (Gunther, 1875), Orthrio-

phis moellendorffi (Boettger, 1886), and Sibynophis

collaris (Gray, 1853).

AMPHIBIA

ANURA

MEGOPHRYIDAE

Brachytarsophrys intermedia (Smith, 1921)

Leptobrachium chapaense (Bourret, 1937)

Leptolalax aereus Rowley, Stuart, Richards, Phim-

machak et Sivongxay, 2010(***)

Ophryophryne hansi Ohler, 2003

Xenophrys major (Boulenger, 1908)

BUFONIDAE

Duttaphrynus melanostictus (Schneider, 1799)

Ingerophrynus galeatus (Gunther, 1864)

Ingerophrynus macrotis (Boulenger, 1887) (*)

HYLIDAE

Hyla simplex Boettger, 1901

MICROHYLIDAE

Kalophrynus interlineatus (Blyth, 1854)

Kaloula pulchra Gray, 1831

Microhyla berdmorei (Blyth, 1856)

Microhyla butleri Boulenger, 1900

Microhyla fissipes Boulenger, 1884

Microhyla heymonsi Vogt, 1911

Microhyla inornata (Boulenger, 1 890)

Microhyla marmorata Bain et Nguyen, 2004

Microhyla pulchra (Hallo well, 1861)

DICROGLOSSIDAE

Fejervarya limnocharis (Gravenhorst, 1829)

Hoplobatrachus rugulosus (Wiegmann, 1834) (***)

Limnonectes bannaensis Je, Fei et Jiang, 2007 (***)

Limnonectes gyldenstolpei (Andersson, 1916) (*)

Limnonectes limborgi (Sclater, 1892) (***)

Limnonectes poilani (Bourret, 1942)

Occidozyga lima (Gravenhorst, 1 829)

Occidozyga martensii (Peters, 1 867)

RANIDAE

Amolops cremnobatus Inger et Kottelat, 1998

Babina chapaensis (Bourret, 1937) (*)

Hylarana attigua (Inger, Orlov et Darevsky, 1999)

Hylarana guentheri (Boulenger, 1882)

Hylarana maosonensis Bourret, 1937

Hylarana nigrovittata (Blyth, 1856)

O dorr ana chloronota (Gunther, 1876)

Odorrana tiannanensis (Yang et Li, 1980)

Rana johnsi Smith, 1921

RHACOPHORIDAE

Chiromantis vittatus (Boulenger, 1887)

Gracixalus quyeti (Nguyen, Hendrix, Bohme, Vu,

et Ziegler, 2008)

Kurixalus banaensis (Bourret, 1939)

Kurixalus bisacculus (Taylor, 1962) (***)

Polypedates megacephalus Hallowell, 1861(***)

Polypedates mutus (Smith, 1940)

Rhacophorus annamensis Smith, 1924

Rhacophorus dennysi Blanford, 1881

Rhacophorus exechopygus Inger, Orlov et Darev-

sky, 1999

Rhacophorus kio Ohler et Delorme, 2006

Rhacophorus orlovi Ziegler et Kohler, 2001

Rhacophorus rhodopus Liu et Hu, 1960

Theloderma asperum (Boulenger, 1886)

Theloderma corticale (Boulenger, 1903) (*)

Theloderma stellatum Taylor, 1962 (*)

296

Vinh Quang Luu etalii

REPTILIA

TESTUDINES

PLATYSTERNIDAE

Platysternon megacephalum Gray, 1831

GEOEMYDIDAE

Cuora cyclornata Blanck, McCord et Le, 2006 (***)

Cuora galbinifrons Bourret, 1939

Cuora mouhotii (Gray, 1862)

Cydemys oldhamii Gray, 1863 (***)

Heosemys grandis (Gray, 1860)

Mauremys mutica (Cantor, 1842)

Ocadia sinensis (Gray, 1834)

Sacalia quadrioceUata (Siebenrock, 1903)

TESTUDINIDAE

Manouria impressa (Gunther, 1882)

TRIONYCHIDAE

Palea steindachneri (Siebenrock, 1906)

Pelodiscus cf. parviformis Tang, 1997 (***)

SQUAMATA: SAURIA

GEKKONIDAE

Cyrtodactylus cryptus Heidrich, Rosier, Vu, Bohme

et Ziegler, 2007

Cyrtodactylus phongnhakebangensis Ziegler, Ro-

sier, Herrmann et Vu, 2003

Cyrtodactylus roesleri Ziegler, Nazarov, Orlov,

Nguyen, Vu, Dang, Dinh et Schmitz, 2010 (**)

Gehyra mutilata (Wiegmann, 1834)

Gekko palmatus Boulenger, 1907

Gekko reevesii Gray, 1831 (***)

Gekko scientiadventura Rosier, Ziegler, Vu, Herrmann

et Bohme, 2004

Hemidactylus frenatus Dumerilet Bibron, 1836

Ptychozoon lionotum Annandale, 1905 (**)

AGAM1DAE

Acanthosaura lepidogaster (Cuvier, 1 829)

Calotes emma Gray, 1 845

Calotes versicolor (Daudin, 1802)

Physignathus cocincinus Cuvier, 1 829

ANGUIDAE

Dopasia gracilis G ray, 1845 (**)

VARANIDAE

Varanus salvator (Laurenti, 1768)

LACERT1DAE

Takydromus hani Chou, Nguyen et Pauwels, 2001

Takydromus kuehnei van Denburgh, 1909

Takydromus sexlineatus Daudin, 1 802

SCINCIDAE

Eutropis longicaudata (Hallowed, 1856)

Eutropis macularia (Blyth, 1853)

Eutropis multifasciata (Kuhl, 1 820)

Lygosoma boehmei Ziegler, Schmitz, Heidrich, Vu

et Nguyen, 2007

Lygosoma quadrupes (Linnaeus, 1766)

Plestiodon elegans (Boulenger, 1887)

Plestiodon quadrilineatus Blyth, 1853

Scincella melanosticta (Boulenger, 1887)

Scincella rufocaudata (Darevsky et Nguyen, 1983) (*)

Sphenomorphus indicus (Gray, 1853)

Sphenomorphus tetradactylus (Darevsky et Orlov,

2005) (***)

Tropidophorus cocincinensis Dumeril et Bibron, 1839

Tropidophorus noggei Ziegler, Vu et Bui, 2005

SQUAMATA: SERPENTES

TYPHLOPIDAE

Ramphotyphlops braminus (Daudin, 1803)

XENOPELTIDAE

Xenopeltis hainanensis Hu et Zhao, 1972

Xenopeltis unicolor Boie, 1 827

BOIDAE

Broghammerus reticulatus (Schneider, 1801) (***)

Python molurus (Linnaeus, 1758)

XENODERMATIDAE

Fimbrios smithi Ziegler, David, Miralles, Doan et

Nguyen, 2008

Further new records of amphibians and reptiles from Phong Nha - Ke Bang National Park, Quang Binh Province, Vietnam 297

COLUBRIDAE

Ahaetulla prasina (Boie, 1827)

Amphiesma andreae Ziegler et Le, 2006

Amphiesma boulengeri (Gressitt, 1937) (***)

Amphiesma leucomystax David, Bain, Nguyen,

Orlov, Vogel, Vu et Ziegler, 2007

Amphiesma stolatum (Linnaeus, 1758)

Boiga bourreti Tillack, Ziegler et Le, 2004

Boiga guangxiensis Wen, 1998

Boiga multomaculata (Boie, 1827)

Calamaria thanhi Ziegler & Le, 2005

Chrysopelea ornata (Shaw, 1802)

Coelognathus radiatus (Boie, 1827)

Cyclophiops major (Gunther, 1858)

Cyclophiops multicinctus (Roux, 1907)

Dendrelaphis ngansonensis (Bourret, 1935)

Dryocalamus davisonii (Blanford, 1878)

Enhydris plumbea (Boie, 1827)

Liopeltis frenatus (Gunther, 1858)

Lycodon fasciatus (Anderson, 1897

Lycodon futsingensis (Pope, 1928)

Lycodon paucifasciatus Rendahl, 1 943

Lycodon cf. rufozonatum Cantor, 1842 (***)

Lycodon ruhstrati (Lischer, 1886)

Oligodon chinensis (Gunther, 1888)

Oligodon cinereus pallidocinctus (Bourret, 1934) (*)

Oreocryptophis porphyraceus (Cantor, 1839)

Orthriophis taeniurus Cope, 1861

Parahelicops annamensis Bourret, 1934 (*)

P areas carinatus Wagler, 1830

Pareas hamptoni (Boulenger, 1905)

Pareas macularius Blyth, 1868

Pareas margaritophorus (Jan, 1 866)

Psammodynastes pulverulentus (Boie, 1 827)

Pseudoxenodon macrops (Blyth, 1854)

Ptyas korros (Schlegel, 1837)

Ptyas mucosa (Linnaeus, 1758)

Rhabdophis chrysargos (Schlegel, 1837)

Rhabdophis subminiatus (Schlegel, 1837)

Rhadinophis prasinus (Blyth, 1854) (***)

Rhynchophis boulengeri Mocquard, 1897 (*)

Sinonatrix percarinata (Boulenger, 1899)

Xenochrophis flavipunctatus (Hallowed, 1860)

ELAPIDAE

Bungarus candidus (Linnaeus, 1758)

Bungarus fasciatus (Schneider, 1801)

Naja cf. atra Cantor, 1 842

Ophiophagus hannah (Cantor, 1836)

Sinomicrurus macclellandii (Reinhardt, 1844) (*)

VIPER1DAE

Protobothrops cornutus (Smith, 1930)

Protobothrops mucrosquamatus (Cantor, 1839) (*)

Protobothrops sieversorum (Ziegler, Herrmann,

David, Orlov et Pauwels, 2000)

Trimeresurus albolabris (Gray, 1842) (***)

Trimeresurus truongsonensis (Orlov, Ryabov, Bui

et Ho, 2004) (***)

Trimeresurus vogeli (David, Vidal et Pauwels,

2001 ) (***)

DISCUSSION

This study brings the confirmed species number

of amphibians and reptiles recorded from Phong

Nha - Ke Bang National Park to 151, including 50

species of amphibians (5 Megophryidae, 3 Bu-

fonidae, 1 Hylidae, 9 Microhylidae, 8 Dicroglossi-

dae, 9 Ranidae, 15 Rhacophoridae), 12 species of

turtles (1 Platysternidae, 8 Geoemydidae, 1 Testu-

dinidae, 2 Trionychidae), 31 species of lizards (9

Gekkonidae, 4 Agamidae, 1 Anguidae, 1 Varanidae,

3 Lacertidae, 13 Scincidae), and 58 species of snakes

(1 Typhlopidae, 2 Xenopeltidae, 2 Boidae, 1 Xeno-

dermatidae, 41 Colubridae, 5 Elapidae, 6 Viperi-

dae), see also fig. 16. An updated list, including

recent taxonomic/nomenclatural changes since the

last review by Ziegler & Vu (2009), is provided.

The research history of the herpetofauna of Phong

Nha - Ke Bang clearly shows that even after more

than a decade of very intensive herpetological sur-

veys, additional species can be recorded or even

discovered as new to science. Since the description

of the first herpetological discovery from Phong

Nha - Ke Bang, Cyrtodactylus phongnhakebangen-

sis by Ziegler et al. (2002), a series of new species

descriptions took place to date, amongst others

eight snake and four gecko taxa, to mention only

the most outstanding examples (see overview in

Ziegler & Vu, 2009). And we are aware of further

new discoveries to be described from the region in

the near future. This does not only underline the

importance of Phong Nha - Ke Bang National Park

in a regional scale and the Truong Son Mountain

Range along the border between Vietnam and Laos

298

Vinh Quang Luu etalii

•^VVVVVV 0

Figure 16. Species richness of amphibian and reptile families from Phong Nha - Ke Bang National Park (new records are

marked by open rectangles).

in a wider geographical scale as centres of biodi-

versity and endemism, but also shows that longterm

biodiversity research is crucial for covering the

total species richness in tropical environment,

which is prerequisite for appropriate evaluation of

the conservation status and application of adequate

protection measures.

ACKNOWLEDGEMENTS

We are grateful to the directorates of Phong Nha

- Ke Bang National Park for the support of our field

work and issuing relevant permits. We thank H. T.

An and D. V. Nguyen (Hanoi) and H. V. Nguyen

(Quang Binh) for their assistance in the field. Field

surveys in Phong Nha - Ke Bang National Park and

extension area was supported by the Nature Con-

servation and Sustainable Natural Resource Man-

agement in Phong Nha - Ke Bang National Park

Region Project with funding from the Vietnam-Ger-

many Development Cooperation. We also acknowl-

edge the World Association of Zoos and Aquariums

(WAZA) for the support of biodiversity research

and conservation in Phong Nha - Ke Bang National

Park (WAZA branded project 07011). Research of

V. Q. Luu in Germany is funded by the Ministry of

Education and Training of Vietnam (MOET, Project

322) and the German Academic Exchange Service

(DAAD). Research of T. Q. Nguyen in Germany is

funded by the Alexander von Humboldt

Stiftung/Foundation (VIE 114344). For discussions

about the taxonomic status of some snake taxa we

are grateful to Patrick David (Paris). Thanks also

to Anna Rauhaus (Cologne Zoo) for preparing the

plates.

REFERENCES

Blanck T., McCord W. P. & Le M., 2006. On the vari-

ability of Cuora trifasciata (Bell, 1825), the redisco-

very of the type specimen, with descriptions of a new

Cuora species and subspecies, and remarks on the

distribution, habitat and vulnerability of these spe-

cies. Chimaira, Frankfurt am Main.

Bourret R., 1936. Les serpents de Flndochine, II, Cata-

logue systematique descriptif. H. Basuyau, Toulouse.

Bourret R., 1942. Les Batraciens de Flndochine [Hanoi]:

Institut Oceanographique de Flndochine.

Chuaynkern Y., Ohler A., Inthara C., Duengkae P.,

Makchai S. & Salangsingha N., 2010. A revision

of species in the subgenus Nidirana Dubois, 1992,

with special attention to the identity of specimens al-

located to Rana adenopleura Boulenger, 1909, and

Rana chapaensis (Bourret, 1937) (Amphibia: Anura:

Ranidae) from Thailand and Laos. Raffles Bulletin

of Zoology, 58: 291-310.

Darevsky I.S. & Orlov N. L., 2005. New species of limb-

reduced lygosomine skink genus Leptoseps Greer,

Further new records of amphibians and reptiles from Phong Nha - Ke Bang National Park, Quang Binh Province, Vietnam 299

1997 (Sauria, Scincidae) from Vietnam. Russian

Journal of Herpetology, 12: 65-68.

David P., Nguyen T.Q., Nguyen T.T., Jiang K., Chen T.

B., Teynie A. & Ziegler T., 2012. A new species of

the genus Oligodon Fitzinger, 1826 (Squamata:

Colubridae) from northern Vietnam, southern China

and central Laos. Zootaxa, 3498: 45-62.

David P., Vogel G. & Dubois A., 2011. On the need to

follow rigorously the Rules of the Code for the sub-

sequent designation of a nucleospecies (type species)

for a nominal genus which lacked one: the case of the

r \

nominal genus Trimeresurus Lacepede, 1804 (Rep-

tilia: Squamata: Viperidae). Zootaxa, 2992: 1-51.

Fritz U., Gong S., Auer M., Kuchling G., Schneeweifi N.

& Hunsdorfer A. K., 2010. The word’s economically

most important chelonians represent a diverse species

complex (Testudines: Trionychidae: Pelodiscus). Or-

ganisms Diversity & Evolution, 10: 227-242.

Fritz U., Guicking D., Auer M., Sommer R.S., Wink M.

& Hundsdorfer A.K., 2008. Diversity of the South-

east Asian leaf turtle genus Cyclemys : how many

leaves on its tree of life? Zoologica Scripta, 37: 367-

390.

Frost, D.R., 2013. Amphibian Species of the World: an On-

line Reference. Version 5.6 (9 January 2013). Electronic

Database accessible at http://research.amnh.org/herpe-

tology/amphibia/index.html. American Museum of Na-

tural History, New York, USA.

Glaw F. & Vences M., 2007. A field guide to the amphi-

bians and reptiles of Madagascar. Third Edition,

Frosch Verlag, Cologne, 496 pp.

Hendrix R., Nguyen T.Q., Bohme W. & Ziegler T., 2008.

New anuran records from Phong Nha - Ke Bang Na-

tional Park, Truong Son, central Vietnam. Herpetol-

ogy Notes, 1: 23-31.

Hoang N.T., Ho A.T. & Hoang Q.D., 2012. New record

of Ptvchozoon lionotum Amiandale, 1905 (Squamata:

Sauria: Gekkonidae) in Phong Nha - Ke Bang Na-

tional Park, Quang Binh Province. Pp. 255-259 (in

Vietnamese). In: Hoang X.Q. et al. (Eds.), Proceed-

ings of the second National Scientific Workshop on

Amphibia and Reptilia in Vietnam. Vinh University

Press.

Inger R.F. & Stuart B. L., 2010. Systematics of Lim-

nonectes (Taylorana) Dubois. Current Herpetology,

29: 51-68.

Inger R.F., Orlov N.L. & Darevsky I.S., 1999. Frogs of

Vietnam: A report on new collections. Fieldiana:

Zoology, New Series, 92: 1-46.

Kuraishi N., Matsui M., Hamidy A., Belabut D. M.,

Ahmad N., Panha S., Sudin A., Yong H.S.Y., Jiang

J.-P, Ota H., Thong H.T. & Nishikawa K., 2012. Phy-

logenetic and taxonomic relationships of the Polype-

dates leucomystax complex (Amphibia). Zoologica

Scripta, doi:10.1111/j.l463-6409.2012.00562.x

McLeod D.S., 2010. Of least concern? Systematics of a

cryptic species complex: Limnonectes kuhlii (Am-

phibia: Anura: Dicroglossidae). Molecular Phyloge-

netics and Evolution, 56: 991-1000.

Nguyen Q.T. & Nguyen V.K., 2008. First record of The-

loderma stellatum Taylor, 1962 from Phu Quoc Is-

land, Kien Giang Province, southern Vietnam.

Herpetology Notes 1: 61-62.

Nguyen Q.T., Bohme W., Nguyen T.T., Le K.Q., Pahl

K.R., Haus T. & Ziegler T., 2011a. Review of the

genus Dopasia Gray, 1853 (Squamata: Anguidae) in

the Indochina subregion. Zootaxa 2894: 58-68.

Nguyen V.S., Ho T. C. & Nguyen Q. T., 2009. Herpeto-

fauna of Vietnam. Edition Chimaira, Frankfurt am

Main, 768 pp.

Nguyen Q.T., Le D. M., Pham T.C., Nguyen T.T., Bon-

kowski M. & Ziegler T., 2012. Anew species of Gra-

cixalus (Amphibia, Anura, Rhacophoridae) from

northern Vietnam. Organisms Diversity and Evolu-

tion, doi: 10. 1007/s 13 127-0 12-0 11 6-0.

Nguyen Q.T., Schmitz A., Nguyen T.T., Orlov N.L., Bohme

W. & Ziegler T., 2011b. A review of the genus Sphe-

normorphus Fitzinger, 1843 (Squamata: Sauria: Scin-

cidae) in Vietnam, with description of a new species

from northern Vietnam and Hainan Island, southern

China and the first record of S. mimicus Taylor, 1962

from Vietnam. Journal of Herpetology, 45: 145-154.

Nguyen Q. T., Stenke R., Nguyen H.X. & Ziegler T.,

2011c. The terrestrial reptilian 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: 99-115.

Ohler A. & Dubois A., 1999. The identity of Elachy-

glossa gyldenstolpei Andersson, 1916 (Amphibia,

Ranidae), with comments on some aspects of statisti-

cal support to taxonomy. Zoologica Scripta, 28: 269-

279.

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.L., Dutta S.K., Ghate H.V. & Kent Y., 2006.

New species of Theloderma from Kon Turn Province

(Vietnam) and Nagaland State (India) (Anura: Rha-

cophoridae). Russian Journal of Herpetology 13:

165-175.

Phung M.T. & Ziegler T., 2011. Another new Gekko

species (Squamata: Gekkonidae) from southern Viet-

nam. Zootaxa, 3129: 51-61.

Rosier H., Bauer A.M., Heinicke M.P., Greenbaum E.,

Jackman T., Nguyen Q.T. & Ziegler T., 2011. Phy-

logeny, taxonomy, and zoogeography of the genus

300

Vinh Quang Luu etalii

Gekko Laurenti, 1768 with the revalidation of G. ree-

ves ii Gray, 1831 (Sauria: Gekkonidae). Zootaxa,

2989: 1-50.

Siler C.D., Oliveros C.H., Santanen A. & Browne R.M.,

2013. Multilocus phylogeny reveals unexpected di-

versification patterns in Asian wolf snakes (genus Ly-

codon). Zoologica Scripta, doi: 1 0. 1 1 1 1/zsc. 12007.

Smith M.A., 1943. The fauna of British India, Ceylon and

Burma, including the whole of the Indo-Chinese sub-

region. Vol. 3-Serpentes. Taylor and Francis, London.

Stuart B.L., 2006. A second specimen of Parahelicops

annamensis Bourret, 1934 (Serpentes: Colubridae:

Natricinae). Hamadryad, 30: 167-171.

Stuart B.L. & Emmett D.A., 2006. A collection of am-

phibians and reptiles from the Cardamom Mountains,

southwestern Cambodia. Fieldiana: Zoology, 109: 1-

27.

Stuart B.L. & Fritz U., 2008. Historical DNA from mu-

seum type specimens clarifies diversity of Asian leaf

turtles ( Cyclemys ). Biological Journal of the Linnean

Society, 94: 131-141.

Taylor E.H., 1962. The amphibian fauna of Thailand.

University of Kansas Science Bulletin, 63: 265-599.

Uetz P., 2013. The Reptile Database, http://www.reptile-

database.org, accessed 31 January, 2013.

Yu G.-H., Zhang M.-W. & Yang J.-X., 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.

Ziegler T., Hendrix R., Vu N.T., Vogt M., Forster B. &

Dang N.K., 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: i_40.

Ziegler T. & Herrmann H.-W., 2000. Preliminary list of

the herpetofauna of the Phong Nha - Ke Bang area in

Quang Binh province, Vietnam. Biogeographica, 76:

49-62.

Ziegler T., Herrmann H.-W., Vu N.T., Le K.Q., Nguyen

T.H., Cao X.C., Luu M.T. & Dinh H.T., 2004. The

amphibians and reptiles of the Phong Nha - Ke Bang

National Park, Quang Binh Province, Vietnam. Ha-

madryad, Tamil Nadu, 28: 19-42.

Ziegler T., Nazarov R., Orlov N., Nguyen Q.T., Vu N.T.,

Dang N.K., Dinh H.T. & Schmitz A., 2010. A third

new Cyrtodactylus (Squamata: Gekkonidae) from

Phong Nha - Ke Bang National Park, Truong Son,

Vietnam. Zootaxa, 2413: 20-36.

Ziegler T., Ohler A., Vu N.T., Le K.Q., Nguyen X.T.,

Dinh H.T. & Bui N.T., 2006. Review of the am-

phibian and reptile diversity of Phong Nha - Ke

Bang National Park and adjacent areas, central

Truong Son, Vietnam. In: Vences, M., Kohler, J.,

Ziegler, T. & W. Bohme (Eds.): Herpetologia Bon-

nensis II: 247-262. Proceedings of the 13th Ordinary

General Meeting of the Societas Europaea Herpeto-

logica, Bonn.

Ziegler T., Rosier H., Herrmann H.-W. & Vu N.T., 2002.

Cyrtodactylus phongnhakebangensis sp. n., a new

bent-toed gecko from the Annamitic karst forest mas-

sif, Vietnam. Herpetofauna, 24: 11-25.

Ziegler T. & Vu N.T., 2009. Ten years of herpetodiversity

research in Phong Nha - Ke Bang National Park, cen-

tral Vietnam. In: In: Vo V. T., Nguyen D. T., Dang K.

N. & Pham Y. H. T. (Eds.). Phong Nha - Ke Bang

National Park and Cologne Zoo, 10 years of coope-

ration: 103-124.

Biodiversity Journal, 2013, 4 (2): 301-326

First contribution to the reptile fauna of Quang Ngai Pro-

vince, central Vietnam

Liza Nemes 1 , Randall Babb 2 , Wa/ne Van Devender 2 , Khoi Vu Nguyen 2 , Quyet Khac Le 3 , Thanh NgocVu 4 ,

Anna Rauhaus 1 , Truong Quang Nguyen 5,6 &Thomas Ziegler 1,6 *

'AG Zoologischer Garten Koln, Riehler Strasse 173, D-50735 Cologne

2 Wildlife At Risk, 202/10 Nguyen Xi Street, Ward 26, Binh Thanh District, Ho Chi Minh City, Vietnam

’Department of Anthropology, University of Colorado, Boulder, CO 80309 USA

4 Faculty of Biology, University of Science, Vietnam National University, 334 Nguyen Trai Str., Thanh Xuan, Hanoi, Vietnam,

institute of Ecology and Biological Resources, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet, Hanoi, Vietnam.

6 Zoological Institute, University of Cologne, Ztilpicher Strasse 47b, D-50674 Cologne, Germany; e-mail: ziegler@koelnerzoo.de

‘Corresponding author

ABSTRACT Based on recent field surveys, we provide a first preliminary list of reptiles from Quang

Ngai Province, central Vietnam, comprising 35 recorded species, including 16 species of

lizards (Agamidae: 6, Gekkonidae: 2, Lacertidae: 1, Scincidae: 6, Anguidae: 1), 18 species

of snakes (Xenodermatidae: 1, Colubridae: 14, Viperidae: 3), and 1 species of turtle (Geoe-

mydidae: 1). The taxonomic status of Fimbrios cf. klossi from Quang Ngai needs further

examination.

KEY WORDS herpetofauna; taxonomy; distribution; Kon Turn Plateau.

Received 20.04.2013; accepted 08.06.2013; printed 30.06.2013

INTRODUCTION

According to the checklist of Nguyen et al.

(2009) 368 species of reptiles are known to occur

in Vietnam. In the following year 16 species were

added to the country’s reptile fauna (see overview

in Ziegler & Nguyen, 2010), thus increasing the

number of Vietnamese reptile species to 384. Since

then new country records and new species descrip-

tions of reptiles were regularly published from Viet-

nam (e.g., Nguyen et al., 2011a, b; David et al.,

2012; Nazarov et al., 2012; Luu et al., 2013), show-

ing that the Vietnamese reptile fauna is still little

understood. One of the most poorly studied regions

in terms of herpetodiversity is Quang Ngai Province

at the south central coast of Vietnam (Fig. 1). Only

recently, a first preliminary list of amphibians oc-

curring in this province has been published by Tran

et al. (2010) comprising 16 species of anurans.

Nguyen et al. (2009) listed only two terrestrial

reptile species with definite records from Quang

Ngai Province, i.e., the lizard Leiolepis guenther-

petersi, and the snake Oligodon ocellatus. We

herein provide a first preliminary reptile inventory

for Quang Ngai Province, based on recent field re-

search in particular in the evergreen forest in a tran-

sitional area between the Kon Turn Plateau and the

lowlands.

MATERIALS AND METHODS

Preliminary field work was conducted by Quyet

Khac Le (QKL) and Khoi Le Vu (KLV) in the ever-

302

Liza Nemes et alii

green forest in the vicinity of Mo Nit Village, Son

Ky Town, Son Ha District, Quang Ngai Province,

Vietnam (14°51’N, 108°31’E, elevation 800 m

a.s.l.), from 12 to 21 July 2008 (Fig. 2). The survey

site is located in a transitional area between the

Kon Turn Plateau and the lowlands.

Additional surveys were conducted by Klioi Vu

Nguyen, Paul Moler, Randall Babb, Ab Abercrom-

bie, Chris Hope, and Wayne Van Devender

(hereafter KVN and others) in Ba To District, dur-

ing May and June 2011 (camp site 14°39’36.7”N,

108 o 36’27.0”E, elevation 750 m), in March 2012

(camp site 14°37 , 10.2 ,, N, 108°35’04.2”E, elevation

920 m), and from May to June 2012 (camp site

14°37’05.9 ,, N, 108°30’10.6”E, elevation 1,025 m).

The only specimens which were not collected at

the camps in the highlands are Hemidactylus fre-

natus (derive from human environment in low-

lands), Leiolepis guentherpetersi and Lygosoma

bowringii (collected at the beach east of Quang

Ngai), as well as Enhydris subtaeniata (fished in

lowlands). Specimens were photographed in the

field. A few individuals were collected by hand for

subsequent proper identification. Collected speci-

mens were anaesthetised and euthanized with ethyl

acetate, fixed in 80-90% ethanol and subsequently

stored in 70% ethanol. Specimens were deposited

in the following collections: Institute of Ecology

and Biological Resources (IEBR), Hanoi, Vietnam;

Zoological Museum, Vietnam National University

(VNUH), Hanoi, Vietnam; Zoologisches Forschun-

gsmuseum Alexander Koenig (ZFMK), Bonn, Ger-

many.

Taxonomic determination followed Pope

(1935), Smith (1935, 1943), Bourret (1936, 2009),

Gressit (1937), Taylor (1963), Musters (1983), Da-

revsky & Kupriyanova (1993), Murphy & Voris

(1994), Manthey & Grossmann (1997), Ziegler &

Vogel (1999), David et al. (2001, 2007), Ziegler

(2002), Leviton et al. (2003), Orlov et al. (2006),

Vogel & David (2006), Ziegler et al. (2006, 2007,

2010), Fritz et al. (2008), Rosier et al. (2008), Green

(2010), Grismer & Grismer (2010), Kams et al.

(2010), Nguyen et al. (2009, 2010, 2011a, b), and

Hartmann et al. (2013). Morphological characters

were given based on the recorded specimens, de-

viations from literature or in between literature

records were presented under remarks. Sex was

identified by examination of external characters

such as hemipenis swellings or through dissection

Figure 1. Map showing the survey site in Quang Ngai

Province, central Vietnam.

(inspection of gonads, retracted copulatory organs).

Scientific and common names (in English and Viet-

namese) as well as distribution records (except

otherwise noticed) followed Nguyen et al. (2009)

and David et al. (2011).

Measurements were taken with callipers to the

nearest 0. 1 mm. Dorsal scale row formula is given as

dorsal scales around body at one head length behind

head : at midbody : one head length before cloaca.

ABBREVIATIONS. HD = head depth at the

pari-etal region; HL = head length (from the pos-

terior edge of the comer of jaw to the snout tip);

HW = head width at the level of tympanum; O = di-

ameter of the orbit; SnL = snout length (from the

anterior edge of the orbital cavity to the snout tip);

SVL (snout-vent length) = distance between tip of

snout and vent; T = tympanum diameter; TAL = tail

length (measured from posterior margin of cloaca

to tip of tail; + = tail reduced or regenerated).

First contribution to the reptile fauna of Quang Ngai Province, central Vietnam

303

RESULTS

SQUAMATA SAURIA

AGAMIDAE

Acanthosaura lepidogaster (Cuvier, 1829)

Scale-bellied tree lizard; O ro vay

Examined material. Specimens examined (n =

2). One adult male collected by QKL and KLV

(ZFMK 94265), SVL: 74.1 mm, TAL: 119.6 mm);

one adult female coll, by KVN and others (IEBR

3277, SVL: 75.7 mm, TAL: 108.7 mm) (Fig. 3).

Morphological characters. Body laterally

compressed; head about 1 .5 times longer than wide;

snout shorter than orbit; frontal region deeply con-

cave; scales on upper head unequal, obtusely

keeled, much smaller in size in posterior region;

canthus rostralis and supraciliary edge strongly pro-

jecting; a spine present at the end of the super-

cilium; another one on the nape, mid- way between

the tympanum and the nuchal crest, with enlarged

scales at the base; ear opening distinct, visible;

supralabials 10-12; infralabials 11 or 12; gular sac

absent, gular scales strongly keeled, smaller than

ventrals; a strong oblique fold present in front of the

shoulder, almost extending across the throat; nuchal

crest composed of long, narrow, compressed spines,

with two rows of shorter ones at the base; dorsal

crest not continuous with the nuchal crest, forming

a prominent ridge, composed of broad, triangular

scales; dorsal scales very small, keeled, intermixed

with much larger, strongly keeled ones, upper scales

pointing upwards, lateral scales pointing backwards

and upwards, sometimes downwards; hind-limbs

reaching or almost reaching to snout tip; third and

fourth fingers equal or nearly so, fourth toe dis-

tinctly longer than third; femoral pores absent; tail

feebly compressed, subtriangular at the base, cov-

ered with equally keeled scales above, strongly

keeled and elongated scales below; ventrals as large

as the largest dorsal, strongly keeled (determination

after Smith, 1935; Ziegler, 2002; Bourret, 2009).

The colouration of the preserved specimens is

olive grey or reddish brown, with light and dark lat-

eral marbling; a large dark diamond- shaped patch

on the neck, the rest of the head being light brown;

tail alternately barred with light and dark; belly

whitish, uniform, or spotted with black; for coloura-

tion in life see figure 3.

Distribution. Vietnam: from the Chinese

border southwards to Lam Dong, Binh Phuoc, and

Dong Nai provinces (this is the first record of the

species from Quang Ngai Province); elsewhere:

South China (Guangdong, Guangxi, Yunnan,

Fujian, Hainan Island), southern Myanmar, Laos,

Thailand, Cambodia.

Remarks. There are some discrepancies re-

garding morphological characters given in the lit-

erature. According to Smith (1935) the head is

shorter (less than one and a half time its width),

whereas Bourret (2009) mentioned the head being

1 % longer than broad. Smith (1935) stated that the

dorsal crest forming a prominent ridge, whereas

such feature is absent in the description of Bourret

(2009). Whether this is due to imprecise descrip-

tions, individual variation or points to cryptic taxa

within A. lepidogaster (e.g., Wood et al., 2010)

must be dealt with in detail elsewhere.

Natural history notes. The testicles of the

male measured 7.3 x 4.7 mm (left side) and 7.5 x

4.9 mm (right side).

Acanthosaura nataliae Orlov, Nguyen et

Nguyen, 2006

Natalia's tree lizard; O ro na-ta-li-a

Examined material. Specimens examined (n =

3). One adult female collected by QKL and KLV

(ZFMK 94266, SVL: 120.1 mm, TAL: 212.1 mm);

one adult male collected by QKL and KLV (ZFMK

94267, SVL: 148.4 mm, TAL: 276.5 mm); one

adult male coll, by KVN and others (IEBR 3276,

SVL: 120.5 mm, TAL: 234.6 mm) (Figs. 4, 5).

Morphological characters. A large-sized

Acanthosaura , postorbital spine present; spine on

occiput between tympanum and nuchal crest absent;

both sexes with well-developed gular pouch; lateral

and dorsal surface of body with large keeled scales

intermixed with small scales; hindlimbs with het-

erogeneous imbricate scales intermixed with larger

scales; 2 or 3 distinct rows of papillary scales

present on the midline of belly (determination after

Orlov et al., 2006).

The colouration of the preserved specimens is

brownish grey, reddish brown, or bluish; upper head

light or dark brown; orbital region blackish brown,

a dark stripe extending from behind the eye to the

tympanum; labial region lighter; tail with broad, al-

304

Liza Nemes et alii

temating dark and light bands; belly yellowish

white; for colouration in life see figures 4, 5.

Distribution. Vietnam: Thanh Hoa, Nghe An,

Quang Tri, Thua Thien-Hue, Da Nang, Quang Nam,

Kon Turn, and Gia Lai provinces (this is the first

record of the species from Quang Ngai Province);

elsewhere: southern Laos (Saravane and Xe Kong

provinces).

Remarks. Enlarged scales in the cloacal region

as described by Orlov et al. (2006) were not seen in

the specimens from Quang Ngai.

Natural history notes. A total of 8 small and

1 0 larger oocytes were found in the female (ZFMK

94266), of which the larger eggs measured 6. 5 -7. 9

mm (left side) and 6.6-8 mm (right side). Testicle

sizes in the male (ZFMK 94267) were 14.2 x 9.5

mm (left side) and 16.8 x 8.3 mm (right side).

Calotes mystaceus Dumeril etBibron, 1837

Blue-crested lizard; Nhong xam

Examined material. Specimens examined (n

2). One adult was photographed by RB (Fig. 6);

another adult female, collected by KVN and

others (IEBR 3278, SVF: 82.7 mm, TAF: 206 mm;

see Fig. 7).

Morphological characters. Body com-

pressed; head more than 1.5 times longer than wide

(ratio HF/HW=1 .8); snout distinctly longer than the

orbit (10.9 versus 8 mm); frontal region feebly con-

cave; upper head-scales unequal, smooth or keeled;

canthus rostralis and supraciliary edge sharp; postor-

bital spine absent; two short, separated spines present

on each side of the nape, the lower one separated

from the tympanum by 4 or 5 scales; a row of 3 or 4

enlarged scales between the eye and the tympanum;

tympanum diameter about half of the orbit (ratio

T/O=0.4); supralabials 1 1 ; infr alabials 10 or 11; gular

pouch small, gular scales strongly keeled, mucronate,

larger than ventrals; an oblique fold present in front

of the shoulder, covered with small granular scales;

dorsal scales strongly keeled, pointing backwards

and upwards, nearly twice as large as ventrals; scales

around midbody 54; limbs moderate; third and fourth

fingers nearly equal; fourth toe distinctly longer than

third toe; hind-limb reaching to the posterior comer

of orbit; tail feebly compressed, covered with sube-

qual, keeled scales; ventrals strongly keeled (deter-

mination followed Smith, 1935, compared with

Hartmann et al., 2013).

The colouration of the preserved specimen is

greyish blue on body with six large chocolate-

coloured spots along the vertebral line; head brow-

nish, with thin dark lines radiating from the eye;

fold in front of the shoulder black; an eponymous

whitish stripe on upper lip extends from between

nostril and eye to the neck; tail pale blue; dirty

whitish below; for colouration in life see figures 6, 7.

Distribution. Vietnam: from Son Fa, Fang

Son, and Nghe An provinces in the North south-

wards to Kien Giang Province (this is the first

record of the species from Quang Ngai Province);

elsewhere: India, China, Myanmar, Faos, Thailand,

and Cambodia.

Draco indochinensis Smith, 1928

Indochinese flying lizard; Than lan bay dong duong

Examined material. Specimen examined (n =

1). One adult female collected by QKF and KLV

(ZFMK 94268, SVF: 110.7 mm, TAF: 205.5 mm)

(Figs. 8, 9).

Morphological characters. Body large; pata-

gial ribs 5; head moderate (HF/SVF 0.21 and

HD/HF 0.44); snout length 0.44 times of the head

length; snout longer than the diameter of the orbit;

nostrils directed almost straight upwards; ear

opening distinct; upper head-shields unequal,

strongly keeled, compressed and more or less

erected over the canthus rostralis; supralabials 10

or 11; two median teeth present on upper jaw; no

thornlike scale on the supraciliary edge, but a

rounded tubercle; gular appendage 0.76 times of the

head length, with enlarged scales; dorsal scales

173, unequal, smooth or faintly keeled, smaller than

the ventral scales; ventrals strongly keeled; a series

of enlarged scales present on the border between

body and patagium, widely separated from each

other, strongly keeled; subdigital lamellae under the

fourth finger 28 or 29, keeled, under the fourth toe

33-35; hindlimbs 0.5 times of the snout-vent length;

fore-limb reaching to beyond the snout, hindlimb

nearly reaching to the axilla; fringelike scales

present on posterior edge of thigh and on base of tail;

tail length 1.86 times the snout- vent length; ventrals

144, keeled, larger than dorsals (determination after

Smith, 1935; Musters, 1983; Bourret, 2009).

The colouration of the presereved specimen is

greyish or bronze and bluish above, with numerous

small black spots; wing-membranes reddish-brown

First contribution to the reptile fauna of Quang Ngai Province, central Vietnam

305

Figure 2. Primary forest in Quang Ngai Province, photo. Q.K. Le. Figure 3. Acanthosaura lepidogaster, adult female, photo

R. Babb. Figure 4. Acanthosaura nataliae, adult male, photo R. Babb. Figure 5. Acanthosaura nataliae, portrait of adult

male, photo Q.K. Le. Figure 6. Calotes mystaceus, uncollected adult, photo R. Babb.

306

Liza Nemes et alii

above, with four distinct, light-bordered, curved

transverse black bands which bifurcate as they

approach the body; ventral surface of patagium

with a black stripe along the outer margin; chin

spotted with black; throat blue, with a broad, black,

transverse bar extending to the inner sides of the

wattles; belly yellowish grey; for colouration in life

see figures 8, 9.

Distribution. Vietnam: Khanh Hoa, Kon Turn,

Lam Dong, Dong Nai, and Tay Ninh provinces (this

is the first record of the species from Quang Ngai

Province); elsewhere: Cambodia.

Remarks. The specimen from Quang Ngai

differs from the description provided by Musters

(1983) in some aspects: 10 or 11 instead of 8-10 su-

pralabials, 28 or 29 instead of 25 or 26 subdigital

lamellae under fourth finger, 33-35 instead of 29-

32 subdigital lamellae under fourth toe, 144 instead

of 123-129 ventrals, and 173 instead of 189-210

dorsals; however, it must be taken into account that

the description of D. blanfordii indochinensis in

Musters (1983) was based on only two specimens.

Natural history notes. The female contained

one egg on the left side (15.8 mm) and two eggs on

the right side (14.8-15.7 mm).

Leiolepis guentherpetersi Darevsky et Kupriya-

nova, 1993

Peter’s butterfly lizard; Nhong cat soc

Examined material. Specimen photographed (n

= 1): one adult was photographed by RB (Fig. 10).

Morphological characters. Identification of

the photographed specimen is based on character-

istic body shape and colour pattern (determination

after Darevsky & Kupriyanova, 1993; Nguyen et

al., 2009; Grismer & Grismer, 2010).

Distribution. Vietnam: Thua Thien-Hue, Da

Nang, Quang Nam, and Quang Ngai provinces;

elsewhere: this species is currently known only

from central Vietnam.

Physignathus cocincinus Cuvier, 1829

Indochinese water dragon / Rong dat

Examined material. Specimens photographed

(n = 2): two subadult to adult specimens were pho-

tographed by QKL and RB (Fig. 11).

Morphological characters. Identification of

the photographed specimens is based on character-

istic body shape and colour pattern, such as dorsal

scales small; distinctly enlarged scale row present

below infralabials; some scattered enlarged scales

present behind mandible; transverse skin fold

present in posterior gular region; nuchal, dorsal and

tail crests well developed; tail laterally com-

pressed; toes without distinct lobes (determination

after Ziegler, 2002).

Distribution. Vietnam: entire country (how-

ever, this is the first documented record of the

species from Quang Ngai Province); elsewhere:

Southern China, Myanmar, Laos, southeastern

Thailand, and Cambodia.

GEKKONIDAE

Cyrtodactylus pseudoquadrivirgatus Rosier,

Vu, Nguyen, Ngo et Ziegler, 2008

Pretended four-striped forest gecko; Thach sung

ngon gia bon vach

Examined material. Specimens examined (n =

3): two adult males (IEBR 3270, SVL: 67.4 mm,

TAL: 54.7 mm; IEBR 3272, SVL: 65.4 mm, TAL:

6.5+ mm); one adult female collected by KVN and

others (IEBR 3271, SVL: 57.4 mm, TAL: 61.3 mm).

Morphological characters. A medium-sized

Cyrtodactylus ; lateral fold narrow, with enlarged

lateral tubercles; interorbital scales 34-45; tubercles

present on dorsum of head, body, limbs and tail;

dorsal tubercles in 18 to 20 rows; lamellae under

first finger 11-13, lamellae under first toe 11-13,

lamellae under fourth toe 18-23; ventral scales at

midbody in 39-46 longitudinal rows; precloacal

pores 7 or 8, in angular series in males, absent in

adult female; enlarged precloacal scales 8-11,

behind angular pore series; enlarged femoral scales

absent; femoral pores and a precloacal groove

absent; transversely enlarged subcaudals absent (de-

termination after Rosier et al., 2008).

Colouration of the preserved specimens: dorsal

surface brown, mottled in dark brown; neck band

medially interrupted; tail with dark and light bands;

for colouration in life (uncollected specimens from

Quang Ngai Province) see figure 12.

Distribution. Vietnam: Quang Tri, Thua Thien-

Hue, Da Nang, and Kon Turn (this is the first record

of the species from Quang Ngai Province); elsewhere:

this species is currently known only from Vietnam.

First contribution to the reptile fauna of Quang Ngai Province, central Vietnam

307

Figure 7. Calotes mystaceus, female, photo R. Babb. Figure 8. Draco indochinensis, adult female in dorsal view, photo Q.K. Le.

Figure 9. Draco indochinensis, adult female in ventral view, photo Q.K. Le. Figure 10. Leiolepis guentherpetersi, photo R. Babb.

Figure 11. Physignathus cocincinus, photo R. Babb. Figure 12. Cyrtoclactylus pseudoquadrivirgatus, photo Q.K. Le.

308

Liza Nemes et alii

Hemidactylus frenatus Schlegel, 1836

Common house gecko; Thach sung duoi san

Examined material. Specimens examined (n =

2): two adult males collected by KVN and others

(ZFMK 94660, SVL: 49.6 mm, TAL: 56 mm;

ZFMK 94661, SVF: 44.8 mm, TAF: 9.1+ mm).

Morphological characters. Head large;

snout obtusely pointed, longer than the distance

between eye and ear-opening; snout with large

scales, posterior part of head covered with small

granules; dorsum with small granules intermixed

with scattered, rounded tubercles; nostril sur-

rounded by rostral, first labial, and three or four

small scales; rostral distinctly broader than high,

quadrangular, with median groove; supralabials 1 1

or 12; infralabials 9; ear-opening oval, oblique, less

than half of the eye diameter; mental large, sub-

triangular; postmentals two pairs, well-developed,

outer pair slightly smaller than the inner; gular re-

gion with small granular scales; fingers and toes

free of webbing; lamellae medially forked, 4 or 5

(some missing) under the first toe, 10 under the

fourth toe; hind-limb not reaching to the axilla; tail

feebly depressed, oval in section, verticillate, dor-

sal surface covered with small scales and series of

six enlarged pointed tubercles; ventrolateral skin

fold absent; belly with smooth, rounded, imbricate

scales; ventrals in 35 and 37 rows; males with 28-

32 precloacal-femoral pores, in continuous series;

postcloacal tubercles 2; ventral tail with a median

series of transversely enlarged scales (determina-

tion after Smith, 1935; Manthey & Grossmann,

1997; Ziegler, 2002; Bourret, 2009).

The colouration of the preserved specimens

is greyish-brown above, with indistinct darker

markings; a bright line along the side of the head;

yellowish white below.

Distribution. Vietnam: entire country; elsewhere:

worldwide in tropical and subtropical regions.

Remarks. The examined specimens differ from

the description by Manthey & Grossmann (1997) in

the number of ventral scales (35-37 versus 28-36).

FACERTIDAE

Takydromus sexlineatus Daudin, 1802

Asian grass lizard; Fiu diu chi

Examined material. Specimen examined (n =

1): one adult male collected by KVN and others

(ZFMK 94659, SVF: 63.5 mm, TAF: 253.7 mm).

Morphological characters. Head narrow and

elongate, pointed anteriorly; nasals just touching

each other behind the rostral; fronto-nasal single, as

long as wide; prefrontals in contact with each other;

supraoculars 3, in contact with the supraciliaries, the

first two the largest, anterior in contact with poste-

rior loreal; supraciliaries 3; interparietal about half

the size of fronto-parietals, larger than the occipital;

anterior loreal smaller than the posterior; temporal

scales strongly keeled, upper ones bordered with

parietal, enlarged; fifth supralabial in contact with

subocular; chin shields 3 pairs, the first pair in con-

tact, third pair largest in size; collar indistinct; gular

scales 23, in a line between chin shield and collar,

scales on the anterior half of the gular region more

elongate and considerably smaller than those on the

posterior half; dorsal plates truncate and shortly

mucronate behind, in 6 rows across the nape, in 4

across the back; lateral scales small, more or less

granular, bordered above and below by larger ones,

10 in a vertical series between the dorsal and ventral

plates; tail very long; caudal scales as large as dorsal

plates, strongly keeled and mucronate; ventral scales

strongly keeled and mucronate, in 12 longitudinal

series; scales between collar and groin 29; a single

femoral pore present on each side; precloacal scale

single (determination after Smith, 1935; Manthey &

Grossmann, 1997; Bourret, 2009).

The colouration of the preserved specimen is

brown to bluish-brown above, with metallic gloss;

a light, dorso-lateral stripe starting from above the

eye and extending on to the base of the tail, edged

above with black spots, below with a black line;

flanks with a series of white black-edged ocelli;

brown colour terminated half way down the flank

and bordered by a thin black stripe, starting from the

nostril to above the ear; upper head and tail brown;

venter and upper lip yellowish and bluish- white.

Distribution. Vietnam: from the Chinese bor-

der southwards to Dong Nai, Tay Ninh and Binh

Duong provinces (this is the first record of the

species from Quang Ngai Province); elsewhere:

India, southern China, Taiwan, Myanmar, Faos,

Thailand, Cambodia, Malaysia, and Indonesia.

Remarks. The specimen matches the description

of Smith (1935) except for only one pair of temporal

scales bordering the parietal being larger than the

First contribution to the reptile fauna of Quang Ngai Province, central Vietnam

309

others; also only the first pair of chin shields is in

contact in the examined specimen instead of the first

two pairs, as was mentioned by Smith (1935).

SCINCIDAE

Eutropis longicaudatus (Hallowell, 1856)

Long-tailed mabuya; Than lan bong duoi dai

Examined material. Specimen photographed

(n = 1): one individual was photographed by RB

(Fig. 13).

Morphological characters. Identification

of the photographed specimen is based on charac-

teristic body shape and colour pattern, such as

dorsal scales with two weakly developed longitu-

dinal keels; well developed limbs with five

fingers and toes each overlapping when laterally

adpressed to the body; ear opening immersed,

visible; supranasals present, touching each other;

postnasal present on each side (determination

after Ziegler, 2002).

Distribution. Vietnam: entire country; else-

where: Southern China, Taiwan, Laos, Thailand,

Cambodia, and Malaysia.

Eutropis multifasciatus (Kuhl, 1820)

Many-lined sun skink; Than lan bong hoa

Examined material. Specimen examined (n = 1):

one juvenile specimen collected by KVN and others

(ZFMK 94662, SVL: 39.5 mm, TAL: 33.1+ mm).

Morphological characters. Supranasals just

touching each other; fronto-nasal broader than long;

postnasal 1; anterior loreal as large as posterior

loreal; supraoculars 4, large; parietals divided by a

large interparietal; lower eyelid scaly, without disc;

supralabials 7, the fifth below the eye; infralabials

7; temporal scales smooth or feebly keeled; ear-

opening visible, moderately large, subcircular,

sunken, with small, pointed lobules anteriorly; nu-

chals in 1 pair; dorsal scales with 3, more or less

strongly keeled, lateral scales feebly keeled; scales

around the midbody 30; vertebrals between pari-

etals and tail base 45; digits moderately long, with

smooth lamellae, 19 beneath the fourth toe; limbs

well developed, hind-limb reaching to elbow when

adpresses along body; precloacals not enlarged (de-

termination after Smith, 1935; Manthey & Gros-

smann, 1997; Ziegler, 2002; Bourret, 2009).

The colouration of the preserved specimen is

uniform olive -brown above; upper parts of flanks

dark brown with white spots; light dorso-lateral

stripe present.

Distribution. Vietnam: entire country; else-

where: India, China, Taiwan, Myamnar, Laos, Thai-

land, Cambodia, Malaysia, Indonesia, Philippines,

and New Guinea.

Lygosoma bowringii (Guenther, 1864)

Bowring’s supple skink; Than lan chan ngan bao-ring

Examined material. Specimen photographed

(n = 1): one individual was photographed by RB

(Fig. 14).

Morphological characters. Identification of

the photographed specimen is based on characteristic

body shape and colour pattern, such as ground

colouration being brown above; each dorsal scale

generally with a darker spot forming more or less

continuous longitudinal lines; dark brown or black

dorsolateral stripe of variable thickness, edged in light

above; sides of neck and body usually thickly spotted

with black and white; white longitudinal stripe across

supralabials until ear opening; yellow below; sides of

neck, body and tail often red in life; fifth supralabial

longest, below the eye; ear opening subcircular, about

half as large as the eye, with one very small projecting

lobule anteriorly (determination after Smith, 1935;

Manthey & Grossmann, 1997).

Distribution. Vietnam: from Hai Duong Prov-

ince in the North to Kien Giang Province in the

South (this is the first record of the species from

Quang Ngai Province); elsewhere: India, China,

Myanmar, Laos, Thailand, Cambodia, Malaysia,

and Philippines.

Scincella melanosticta (Boulenger, 1887)

Black-spotted smooth skink; Than lan co dom den

Examined material. Specimens examined (n =

2): one adult female collected by QKL and KLV

(ZFMK 94391 , SVL: 5 1 .3 mm); one adult male col-

lected by KVN and others (IEBR 3274, SVL 37.4

mm, Fig. 15).

Morphological characters. Size small, tail

regenerated in both specimens; prefrontals in contact

310

Liza Nemes et alii

with each other; supraoculars 4; nuchals absent;

supralabials 7; infralabials 6; primary temporals 2;

secondary temporals 2, lower one overlapped by

upper one; external ear opening present, without lob-

les on anterior margin; tympanum deeply sunken;

midbody scales in 34 rows; dorsal scales smooth, in

8 rows across the dorsum; paravertebral scales 67,

not widened; subdigital lamellae under fourth toe 14-

15 (determination after Smith, 1935; Taylor, 1963).

The colouration of the preserved specimens is

bronze brown on back and dorsal tail base with a

row of large black spots; a dark stripe running from

snout to anterior comer of eye and from posterior

corner of eye to shoulder; upper lateral zone with

large black spots from axilla to tail base; chin with

some dark dots, throat and belly cream.

Distribution. Vietnam: Quang Binh, Lam Dong,

and Ba Ria-Vung Tau provinces (this is the first re-

cord of the species from Quang Ngai Province); else-

where: Myanmar, Laos, Thailand, and Cambodia.

Remarks. Specimens from Quang Ngai also

resemble S. rupicola, however, they were assigned

to S. melanosticta by having a higher number of

midbody scale rows (34 vs. 30) and vertebral row

of dark spots unpaired anteriorly (see Taylor, 1963).

Sphenomorphus indicus (Gray, 1853)

Indian forest skink; Than lan phe-no an do

Examined material. Specimens examined (n =

3): one adult male collected by KVN and others

(ZFMK 94663, SVL: 65.4 mm, TAL: 125.1 mm,

Fig. 16); one adult female collected by QKF & KLV

(ZFMK 94269, SVF: 75.6 mm, TAF: 56 + mm); one

subadult collected by QKF & KLV (ZFMK 94393,

SVF: 43.4 mm).

Morphological characters. Size large; nostril

in a big nasal; supranasals absent; prefrontals sepa-

rated from each other; supraoculars 4; loreals 2;

supraciliaries 8 or 9; lower eyelid scaly; primary

temporals 2; supralabials 7 or 8; external ear

present, with 2 or 3 small lobules on anterior margin;

tympanum deeply sunken; nuchals absent; midbody

scales smooth, in 30-34 rows; paravertebral scales

69, not widened; limbs pentadactyl; lamellae under

fourth toe 16-19 (determination after Smith, 1935;

Taylor, 1963; Ziegler, 2002; Nguyen et al., 2011b).

The colouration of the preserved specimens is

bronze-brown above with irregular black dots; light

dorsolateral stripe present on neck and shoulder;

upper lateral zone with a dark stripe from behind eye

to tail base; lower lateral zone with light bars; ventral

surface cream; for colouration in life see figure 16.

Distribution. Vietnam: from Fao Cai, Cao Bang,

and Bac Kan provinces in the North southwards to

Dong Nai Province (this is the first record of the

species from Quang Ngai Province); elsewhere:

India, Bhutan, China, Taiwan, Myanmar, Faos, Thai-

land, Cambodia, Malaysia, and Indonesia.

Natural history notes. The adult female con-

tained 17 oocytes, 0.5 -1.8 mm in size.

Tropidophorus cocincinensis Dumeril et Bi-

bron, 1839

Cochinchinese water skink; Than lan tai nam bo

Examined material. Specimens examined (n

= 3): three adult specimens collected by QKF and

TNV (ZFMK 94270, SVF: 87.5 mm, TAF: 81.6+

mm; ZFMK 94271, SVF: 76.3 mm, TAF: 73.3+

mm; ZFMK 94272, SVF: 73.4 mm, TAF: 1 14 mm),

see figure 17.

Morphological characters. Upper head-

shields strongly striated; frontonasal undivided;

prefrontals in broad contact, touching each other or

with a small azygous shield in between loreals 2

(posterior loreal fused with frontonasal on right side

in one specimen), separated from supralabials by a

series of small scales; superciliaries 7, superciliary

row not completed along the entire length of lateral

edge of supraoculars; supralabials 7 (8 in one

specimen), fifth largest, below the eye; infralabials

6-8; postmental undivided; nuchals in 3 or 4 pairs;

dorsals strongly keeled; paravertebral scales 41-44;

midbody scales in 30 or 32 rows; ventrals in 46 or

47 transverse rows; enlarged precloacal shields 2;

scale rows at position of tenth subcaudal on tail 14

or 15; dorsal keels on the tail forming strong ridges;

two series of moderately elevated spines along

middle of tail dorsum, continuous with those on

back; subdigital lamellae under fourth toe 18-20

(determination after Nguyen et al., 2010).

The colouration of the preserved specimens is

reddish-brown above with indistinct black spots or

transverse markings on back; large white spots

starting behind the head, extending over the flanks,

paler on the tail, arranged in 1 or 2 longitudinal

rows; belly brownish- white.

First contribution to the reptile fauna of Quang Ngai Province, central Vietnam

311

Figure 13. Eutropis longicaudata, photo R. Babb. Figure 14. Lygosoma bowringii, photo R. Babb. Figure 15. Scincella me-

lanosticta, photo R. Babb. Figure 16. Sphenomorphus indicus, photo R. Babb. Figure 17. Tropidophorus cocincinensis,

photo Q.K. Le. Figure 18. Dopasia gracilis, photo Q.K. Le.

312

Liza Nemes et alii

Distribution. Vietnam: Quang Binh, Quang

Tri, Thua Thien-Hue, Da Nang, Quang Nam, and

Kon Turn provinces (this is the first record of the

species from Quang Ngai Province); elsewhere:

Laos, Cambodia, and Thailand (see also Nguyen

et al., 2010).

Remarks. One specimen from Quang Ngai

shows 8 instead of 7 supralabials. Compared with

the diagnosis of T. cocincinensis in Nguyen et al.

(2010), the Quang Ngai specimens have 3 to 4 pairs

of nuchals (versus 1-3), and the number of scale

rows at position of tenth subcaudal on tail is 14 or

15 (versus 13 or 14).

Natural history notes. The females (ZFMK

94270-94271) contained about 10-20 follicles of up

to 2.1 cm in size on the left side and about 13-20

follicles of up to 2.6 cm in size on the right side,

respectively.

ANGUIDAE

Dopasia gracilis Gray, 1845

Asian glass lizard; Than lan ran

Examined material. Specimens examined (n =

2): one subadult male collected by QKL and KLV

(VNUH 2008.7.10, SVL: 105.0 mm, TAL: 212.0

mm) (Fig. 18).

Morphological characters. Body cylindrical,

without limbs; ear opening 2 times greater than

nostril; nasal separated from frontonasal by 3 scales;

supraoculars 5; supraciliaries 6; supralabials 10, first

and second bordering nostril; infralabials 9; dorsal

scales strongly keeled; longitudinal scale rows

between lateral folds 18; number of scales along la-

teral fold 81-87; ventral scales in 10 longitudinal

rows; vertebrae from atlas to remnants of hind limb

bones 47 (determination after Nguyen et al., 2011a).

The colouration of the preserved specimen is

brown or grey brown dorsally with longitudinal

series of small brown spots on anterior part of back;

two dark stripes present on each side of body:

dorsolateral stripe from one head-length behind ear

opening to tail tip, progressively broader poste-

riorly; ventrolateral stripe narrower but more

distinct than dorsolateral stripe, from angle of jaw

to vent, changing to a series of small spots on tail;

for colouration in life see figure 18.

Distribution. Vietnam: Cao Bang, Ha Giang,

Yen Bai, Quang Binh, and Quang Ngai provinces

(see Nguyen et al., 2011a); elsewhere: India, China,

Myanmar, Laos, and Thailand.

SQUAMATA SERPENTES

XENODERMATIDAE

Fimbrios cf. klossi Smith, 1921

Kloss' snake; Ran ma

Examined material. Specimen examined (n =

1): one adult male collected by KVN and others

(IEBR 3275, SVL: 203.1 mm, TAL: 51.3 mm; Fig.

19), ratio TAL/TL: 0.25.

Morphological characters. Nostril in the

anterior part of a large concave nasal; rostral trian-

gular, not visible from above; suture between inter-

nasals shorter than that between prefrontals;

intemasals separated from rostral by a horizontal

skin fold; frontal broadly tmncate in front, broader

than long, as long as its distance from the tip of

snout, much shorter than parietals, about three times

as broad as the supraoculars; preocular small, just

touching the frontal; large quadrangular loreal in

contact with eye; postoculars 3; subocular 1; tem-

porals 5; supralabials 9 or 10, first 5 very small with

strongly raised edges, the last one more elongated;

infralabials 1 1 , first 7 very small with raised edges

like supralabials; mental absent; a pair of large chin

shields, in contact with the first ventral; dorsal scales

in 28 or 29 rows at anterior part of body; dorsal

scale rows: 28 or 29 : 30 : 31; interstitial skin partly

visible between anterior scales, scales feebly imbri-

cate posteriorly; ventrals 161; cloacal scale undi-

vided; subcaudals 62, unpaired (determination after

Bourret, 1936; Smith, 1943; Ziegler et al., 2008).

The colouration of the preserved specimen is

black above, brighter ventrally; edges of ventrals

and of subcaudals tinged with dark grey.

Distribution. Vietnam: Quang Tri, Thua Thien-

Hue, Kon Turn, Gia Lai, Dak Lak, and Lam Dong

provinces (this is the first record of the species from

Quang Ngai Province); elsewhere: Cambodia.

Remarks. The collected individual differs from

previous descriptions (Bourret, 1936; Smith, 1943;

Ziegler et al., 2008) by a lower number of infralabials

First contribution to the reptile fauna of Quang Ngai Province, central Vietnam

313

(11 versus 12), a higher number of temporals (5

versus 2-3) and subcaudals (62 versus 43-60), a less

developed scale imbrication, no mental shield, and a

different TAL/TL ratio (0.25 versus 0.126 to 0.2).

COLUBRIDAE

Ahaetulla prasina (Boie, 1827)

Oriental whip snake; Ran roi thuong

Examined material. Specimen examined (n = 1):

one specimen collected by KVN and others (ZFMK

94664, SVL: 584.3 mm, TAL: 319.2 mm; Fig. 20).

Morphological characters. Head shields

large; pupil horizontal; snout pointed, projecting,

without dermal appendage, snout more than twice

as long as eye diameter; canthus rostralis sharp;

nostrils lateral, not valvular; nasal in contact with

the labials; loreals 3, small; frontal followed by two

symmetric parietals of size equal to frontal;

preocular 1, in contact with frontal; postoculars 2;

temporals 1 or 2 + 2; supralabials 9, fourth to sixth

touching the eye, entire; infralabials 8, first 4 in

contact with anterior chin shields, shorter than the

posterior; mental groove present; dorsals smooth,

slightly keeled in the sacral region; dorsal scale

rows at midbody 15; ventrals 196; cloacal scale di-

vided; subcaudals 168, divided (determination after

Bourret, 1936; Smith, 1943; Manthey & Gros-

smann, 1997; Ziegler, 2002).

The colouration of the preserved specimen is

bluish-green above, with a yellowish-white line on

both sides of the ventral part; interstitial skin in the

neck region black and white.

Distribution. Vietnam: entire country; else-

where: India, Bangladesh, Bhutan, China, Myanmar,

Laos, Thailand, Cambodia, Malaysia, Singapore,

Bmnei Damssalam, Indonesia, and Philippines.

Amphiesma boulengeri (Gressitt, 1937)

Boulenger's keelback; Ran sai bau-len-go

Examined material. Specimens examined (n

= 2): one subadult specimen collected by KVN

and others (ZFMK 94665, SVL: 107.2 mm, TAL:

46.6 mm; see Fig. 21); one adult male collected by

QKL and KLV (ZFMK 94273, SVL: 370 mm,

TAL: 179 mm).

Morphological characters. Nostrils lateral;

intemasals truncated; preocular 1; postoculars 2;

temporals 1+2, anterior at least twice as long as

posterior; supralabials 9, fourth to sixth touching the

eye, seventh to ninth very large; infralabials 9 or 1 0,

first five in contact with anterior chin shields; ante-

rior chin shields two-thirds as long as posterior pair;

outermost dorsal scale row smooth, remainder finely

keeled; dorsal scale rows 19 : 19 : 17; ventrals 140-

145; subcaudals 89-98, divided (determination after

Gressit, 1937; David et al., 2007).

The colouration of the preserved specimen is

black dorsally; head brown above; anterior supral-

abials white, posterior black with a median elon-

gated cream colored streak, forming a postocular

stripe extending on the neck, followed by white

dorsolateral spots extending to the anterior part of

the tail; venter white.

Distribution. Vietnam: Vinh Phuc, Thua Thien-

Hue, Dak Lak, and Lam Dong provinces (this is the

first record of the species from Quang Ngai Prov-

ince); elsewhere: China.

Remarks. Both specimens differ from the

original description by Gressit (1937) in the number

of posterior temporals (2 instead of 1). The adult

specimen moreover shows a slightly lower number

of ventrals (140 vs. 143-147).

Boiga guangxiensis Wen, 1998

Guangxi cat snake; Ran rao quang Tay

Examined material. Specimen photographed

(n = 1): one adult specimen was photographed by

KVN (Fig. 22).

Morphological characters. Identification of

the photographed specimen is based on character-

istic body shape and colour pattern; single loreal;

postoculars 2; temporals 3 + 3; supralabials 8, third

to fifth touching the eye; dorsal scales smooth (de-

termination after Ziegler et al., 2006, 2010).

Distribution. Vietnam: from Lao Cai, Cao

Bang and Lang Son in the north southwards to

Dong Nai Province (this is the first record of the

species from Quang Ngai Province); elsewhere:

southern China, Laos.

Calamaria pavimentata Dumeril, Bibron et Du-

meril, 1854

Collared reed snake; Ran mai gam lat

314

Liza Nemes et alii

Examined material. Specimen examined (n =

1): one adult female collected by KVN and others

(IEBR 3292, SVL: 231.1 mm, TAL: 18.6 mm;

Fig. 23).

Morphological characters. Head covered

with large shields; pupil round; nostrils lateral, not

valvular; rostral wider than high, partly visible from

above, 0.6 times as long as interprefrontal suture;

supranasals absent; frontal longer than broad,

shorter than parietals, 2 times wider than supraocu-

lars, followed by two symmetrical parietals; loreal

absent; preocular 1 ; postocular 1 ; temporal absent;

supralabials 4, second and third touching the eye,

second and fourth largest; first pair of infralabials

in contact with each other behind the mental;

mental groove present; chin shields in 2 pairs, in

contact with each other, anterior longer than poste-

rior; middorsal scale rows 13, smooth; tail pointed;

ventrals 151, more than twice as broad as adjacent

dorsals; ventrals and subcaudals without notches;

cloacal scale entire; subcaudals 20, divided (deter-

mination after Bourret, 1936; Smith, 1943; Ziegler,

2002; Ziegler et al., 2007).

The colouration of the preserved specimen is

reddish-brown above, with six dark dorsolateral lines

and three paler ones, consisting of small spots; neck

dark brown; belly yellowish; tail ventrally yellowish,

with a dark line medially across the subcaudals.

Distribution. Vietnam: From Lao Cai and Son

La in the West eastwards to Quang Ninh and south-

wards to Quang Nam, and Lam Dong provinces (this

is the first record of the species from Quang Ngai

Province); elsewhere: China, Myanmar, Laos, Cam-

bodia, Thailand, Malaysia, Indonesia, and Japan.

Dendrelaphis ngansonensis (Bourret, 1935)

Nganson bronzeback; Ram leo cay ngan son

Examined material. Specimen examined (n =

1): one adult male collected by KVN and others

(IEBR 3287, SVL: 728.5 mm, TAL: 392.2 mm; Fig.

24), ratio TAL/TL: 0.35.

Morphological characters. Eye as long as

its distance from the nostrils; pupil round; nostrils

lateral, not valvular; rostral much broader than

high, visible from above; intemasals a little shorter

than prefrontals; frontal 1.4 times longer than

broad, as long as its distance from the tip of snout,

shorter than symmetrical parietals; loreal elon-

gated; preocular 1; postoculars 2; temporals 2 + 2;

supralabials 9, fourth to sixth touching the eye; first

five infralabials in contact with anterior chin

shields, anterior chin shields shorter than the pos-

terior; dorsals smooth, dorsal scale rows 15:15:

1 1 ; vertebral scales enlarged, variable in width, not

broader than the outermost scale row at midbody,

posterior margin rounded; ventrals 177, keeled;

cloacal scale divided; subcaudals 148, divided (de-

termination after Bourret, 1936; Smith, 1943; Zie-

gler & Vogel, 1999; Ziegler, 2002).

The colouration of the preserved specimen is

bluish and bronze-brown above; interstitial skin

black and white; dorsal head bronze brown; a black

stripe present along the side of the head, strongly

marked on the temple and extending to the neck;

supralabials yellow; cream below; for colouration

in life see figures 24, 25.

Distribution. Vietnam: From Ha Giang and

Cao Bang provinces in the North southwards to

Quang Nam Province (this is the first record of the

species from Quang Ngai Province); elsewhere:

Laos (Xieng Khouang Province).

Enhydris subtaeniata (Bourret, 1934)

Mekong mud snake; Ran bong me-kong

Examined material. Specimen examined (n =

1): one adult collected by KVN and others (IEBR

3289, SVL: 300 mm, TAL: 56.6 mm; Fig. 26), ratio

TAL/TL: 0.16.

Morphological characters. Body stout;

snout blunt, squarish; nasals contiguous; intemasal

2 times wider than long, in contact with triangular

loreal (loreal divided on left side); preocular 1;

postoculars 2; temporals 1+2; supralabials 8,

fourth in contact with eye; infralabials 3, in contact

with anterior chin shields; posterior pair of chin

shields longer than anterior pair, separated from

each other by small scales; dorsal scale rows 2 1 :

21 : 21; ventrals 143; subcaudals 45, divided (de-

termination after Bourret, 1936; Smith, 1943; Mur-

phy & Voris, 1994; Kams et al., 2010).

The colouration of the preserved specimen is

olive-grey above, with indistinct blackish spots;

ventrals and 3 outer scale rows yellowish; outer

margins of the ventrals and adjacent scale-rows

heavily margined with dark grey; a series of dark

spots forming a median ventral line at the posterior

First contribution to the reptile fauna of Quang Ngai Province, central Vietnam

315

21 22

Figure 19. Fimbrios cf. klossi, photo R. Babb. Figure 20. Ahaetulla prasina, photo R. Babb. Figure 21 . Amphiesma boulen-

geri, juvenile, photo R. Babb. Figure 22. Boiga guangxiensis, adult, photo K.V. Nguyen. Figure 23. Calamaria pavimentata,

photo R. Babb. Figure 24. Dendrelaphis ngansonensis, photo R. Babb.

316

Liza Nemes et alii

part of the body; a darker, continuous line medially

across the subcaudals.

Distribution. Vietnam: Kon Turn, Tay Ninh,

Ho Chi Minh City, Soc Trang, Kien Giang, and Ca

Mau provinces (this is the first record of the species

from Quang Ngai Province); elsewhere: Laos, Thai-

land, and Cambodia.

Remarks. Murphy (2007) revalidated Enhydris

subtaeniata, which was originally described as a

subspecies of E. enhydris (Bourret, 1934) and sub-

sequently synonymized with E. jagorii by Smith

(1943). According to Nguyen et al. (2009),

previous records of E. jagorii from Vietnam and

neighbouring countries should be assigned to E.

subtaeniata.

Oligodon chinensis (Gunther, 1888)

Chinese kukri snake; Ran khiem trung quoc

Examined material. Specimens examined (n =

2): one adult male collected by QKL and KLV

(ZFMK 94274, SVL: 514.2 mm, TAL: 115.7 mm);

one adult male collected by KVN and others (ZFMK

94672, SVF: 557.6 mm, TAF: 126.2 mm, Fig. 27).

Morphological characters. Nasal divided;

internasals 2; rostral partly visible from above, as

long as its distance from frontal; suture of inter-

nasals shorter than sutures of prefrontals; frontal

longer than its distance to the tip of snout, as long

as parietals; loreal somewhat longer than broad;

preocular 1; postoculars 2; subocular 1, temporals

1+2; supralabials 7 or 8, fourth and fifth touching

the eye; infralabials 8 or 9, fourth in contact with

anterior chin shields; anterior pair of chin shields

longer than posterior ones; dorsal scale rows 17 :

17 : 15; ventrals 170 or 172; cloacal scale undi-

vided; subcaudals 58 or 59 (determination after Bour-

ret, 1936; Smith, 1943; Ziegler, 2002; Green, 2010).

The colouration of the preserved specimens is

reddish-brown dorsally; with dark, rounded

blotches; blotches on the back with dark margins,

occupying about 3 to 4 scale rows along the

vertebral line, descending laterally; several similar

spots on the tail; the darker edges of the scales

between these spots forming less visible equidi-

stant fasciatures in each section; head olive brown

with a dark brown stripe across the eyes, edged in

black, running upwards to the posterior end of the

rostral and the upper lip; a long arrowhead-shaped

dark brown spot, edged in black, tip of arrow at

the posterior third of the frontal, the two rear ends

extending to the side of the neck, almost up to the

ventrals; temporal bars absent; ventral surface

white, with a longitudinal row of quadrangular

black blotches forming along each side, disappear-

ing gradually at the tail base.

Distribution. Vietnam: Fao Cai, Bac Kan,

Fang Son, Vinh Phuc, Quang Ninh, Hai Phong, Hai

Duong, Nghe An, Ha Tinh, Quang Binh, and Gia

Fai provinces (this is the first record of the species

from Quang Ngai Province); elsewhere: China.

P areas hamptoni (Boulenger, 1905)

Hampton's slug snake; Ran ho may ham-ton

Examined material. Specimen examined (n =

1): one adult male collected by KVN and others

(ZFMK 94666, SVF: 317.2 mm, TAF: 110.1 mm;

Fig. 28), ratio TAF/TF: 0.26.

Morphological characters. Body elongated,

strongly compressed; pupil vertical; snout short;

rostral wider than high, visible from above; intema-

sals about 0.6 times as long as prefrontals; prefron-

tals touching the eye; frontal somewhat longer than

broad, longer than its distance from the snout tip;

supraocular about half as wide as frontal; nasal

complete; loreal small, higher than long, separated

from the eye by 1 or 2 small preoculars; postocular

1, small; subocular 1, long and slender, separating

the eye from the labials; temporals 2 + 3, anterior

larger than posterior ones; supralabials 7, fourth and

fifth below the eye, seventh very long; mental

groove absent; chin shields in 3 pairs, large, anterior

ones longer than wide; dorsal scales smooth, 15

rows at midbody; ventrals 189, more than 2 times as

broad as adjacent dorsals; cloacal scale undivided;

subcaudals 78, divided (determination after Bourret,

1936; Smith, 1943; Ziegler, 2002).

The colouration of the preserved specimen is

light brown dorsally, with dorsolateral rows of

alternating spots, forming a zigzag line, spots absent

at the margin of the ventrals, not more than 2 scales

in length; two parallel brown lines, starting from the

posterior part of the supraoculars, bordering the

head and ending at the nape; two other bands from

behind the parietals forming a X on the neck; a few

larger spots forming a stripe from the eye to the oral

commissure.

First contribution to the reptile fauna of Quang Ngai Province, central Vietnam

317

Distribution. Vietnam: Lao Cai, Ha Giang, Bac

Kan, Thai Nguyen, Vinh Phuc, Hai Duong, Ha Tinh

southwards to Lam Dong and Dong Nai provinces

(this is the first record of the species from Quang

Ngai Province); elsewhere: China, Myanmar, Laos,

and Cambodia.

Remarks. In contrast to Bourret (1936) and

Smith (1943), the vertebral scales of the examined

specimen are not enlarged.

P areas margaritophorus (Jan, 1866)

White-spotted slug snake; Ran ho may ngoc

Examined material. Specimen examined (n =

1): one adult male collected by KVN and others

(IEBR 3294, SVL: 219.9 mm, TAL: 60 mm; Fig.

29), ratio TAL/TL: 0.21.

Morphological characters. Eye moderate,

diameter equal to its distance from the mouth;

preocular 1, triangular; postocular 1; subocular 1,

long, crescentic, united with postocular on left side;

temporals 2 or 3 + 1 , elongated; supralabials 7, not

touching the eye, fourth and fifth below the eye;

infralabials 7; chin shields in 4 pairs, large; dorsal

scale rows 15 at midbody, smooth, equal; vertebrals

not enlarged; ventrals 144, more than twice as broad

as adjacent dorsals; cloacal scale undivided;

subcaudals 55, divided (determination after Bour-

ret, 1936; Smith, 1943; Manthey & Grossmann,

1997; Ziegler, 2002).

The colouration of the preserved specimen is

dark grey above with irregular transverse rows of

black and white blotches, the scales white ante-

riorly and posteriorly; a white nuchal collar

present; ventral surface yellowish white, densely

spotted with black.

Remarks. In contrast to the description of Bour-

ret (1936), the specimen shows only 1 + 1 tempo-

rals on one side instead of 2 + 3, 4 pairs of chin

shields instead of 3 pairs.

Distribution. Vietnam: Vinh Phuc, Hai Duong,

Hoa Binh, Ha Tay, Ninh Binh, Ha Tinh, Quang

Binh, Quang Tri, Gia Lai, Lam Dong, Tay Ninh, Ho

Chi Minh City, and Kien Giang provinces (this is

the first record of the species from Quang Ngai

Province); elsewhere: China, Myanmar, Laos, Thai-

land, Cambodia, and Malaysia.

Pseudoxenodon macrops (Blyth, 1854)

Big-eyed bamboo snake; Ran ho xien mat to

Examined material. Specimen examined (n =

1): one adult male collected by KVN and others

(IEBR 3300, SVL mm: 587, TAL: 127.1 mm;

Fig. 30).

Morphological characters. Nostril large,

between two nasals; rostral slightly visible from

above; suture between internasals half as long as

suture between prefrontals; frontal 1.3 times longer

than broad, shorter than its distance from the snout

tip, shorter than parietals; loreal large, longer than

high; preocular 1, not touching the frontal; pos-

toculars 3; temporals 2 + 2 or 3; supralabials 8,

fourth and fifth in contact with the eye, seventh

highest; infralabials 9, first five in contact with

anterior chin shields; first pair of chin shields a little

shorter than the posterior; dorsal scales strongly

keeled except the outermost row, scale rows 19:17

: 15; ventrals 151; cloacal scale divided; subcaudals

63, divided (determination after Bourret, 1936;

Smith, 1943; Manthey & Grossmann, 1997).

The colouration of the preserved specimen is

greyish-brown and reddish-brown above, with a

vertebral series of large, pale-orange spots, edged

in black, extending over the whole body and tail;

neck with a yellowish white, chevron-shaped mark,

the tip pointing forward; ventral surface yellowish-

white, with large, blackish-brown, quadrangular

blotches anteriorly, blotches paler posteriorly.

Distribution. Vietnam: Lai Chau, Lao Cai,

Vinh Phuc, Nghe An, Ha Tinh, Quang Binh, Da

Nang, Kon Turn, and Lam Dong provinces (this is

the first record of the species from Quang Ngai

Province); elsewhere: India, Nepal, China, Myan-

mar, Laos, Thailand, and Malaysia.

Rhabdophis subminiatus (Schlegel, 1837)

Red-necked keelback; Ran hoa co nho

Examined material. Specimen examined (n =

1): one adult female collected by KVN and others

(ZFMK 94667, SVL: 494.9 mm, TAL: 48.6 mm;

Fig. 31), TAL/TL: 0.1.

Morphological characters. Body rather

stout; eye large; pupil round; nostrils lateral, not

valvular; mental groove present; rostral 2 times as

wide as high, visible from above; intemasals con-

joint, a little broader than long, truncated at the

318

Liza Nemes et alii

front, shorter than prefrontals; frontal 1.4 times

longer than wide, longer than its distance from the

snout tip, shorter than parietals; loreal as high as

long; preocular 1; postoculars 3; temporals 2 + 3;

supralabials 8, third to fifth touching the eye, se-

venth largest; infralabials 10, first 5 in contact with

anterior chin shields, anterior chin shields a little

shorter than posterior; dorsal scales in 19 rows at

midbody, strongly keeled, the outermost row

smooth; ventrals 161, more than twice as broad as

adjacent dorsals; cloacal scale divided; subcaudals

23 (tail tip lost), divided (determination after Bour-

ret, 1936; Smith, 1943; Manthey & Grossmann,

1997; Ziegler, 2002; Ziegler et al., 2007).

The colouration of the preserved specimen is

bluish above, head and neck greenish; body with

white, transverse spots; a black oblique bar radiated

from below the eye; venter yellowish anteriorly,

strongly spotted posteriorly.

Distribution. Vietnam: From Cao Bang Prov-

ince in the North westwards to Lao Cai and Son

La provinces, eastwards to Quang Ninh and south-

wards to Tay Ninh, Ho Chi Minh City, and Kien

Giang provinces (this is the first record of the

species from Quang Ngai Province); elsewhere:

India, Bangladesh, Nepal, Bhutan, China, Myan-

mar, Laos, Thailand, Cambodia, Malaysia, Singa-

pore, and Indonesia.

Sibynophis collaris (Gray, 1853)

Common black-headed snake; Ran rong co den

Examined material. Specimen examined (n =

1): one male specimen collected by KVN and

others (ZFMK 94668, SVL: 360 mm, TAL: 72

mm, Fig. 32).

Morphological characters. Preocular single,

postoculars 2; loreal 1; fourth to sixth supralabials

entering orbit, eighth very large, in contact with

anterior temporal; temporals 1+2, lower posterior

one divided vertically; parietal touching upper

postocular only; dorsal scales smooth, in 17 rows

at midbody; cloacal undivided (determination after

Pope, 1935; Smith, 1943; Ziegler et al., 2007).

The colouration of the preserved specimen is

greyish brown dorsally; venter whitish cream with

a black dot on the edge of each ventral; head olive

grey with dark markings in the snout region and two

black transverse bars (one behind the eyes and an-

other across the occiput); a distinct dark neck band

followed by a white transverse band; a black edged

white horizontal line present along the supralabials.

Distribution. Vietnam: from Dien Bien, Son La

and Ninh Binh southwards to Lam Dong and Dong

Nai provinces (this is the first record of the species

for Quang Ngai Province); elsewhere: India, Nepal,

China, Taiwan, Myanmar, Laos, Thailand, Cambo-

dia, and Malaysia.

Remarks. The specimen was damaged so that

only the above mentioned scale counts could be

taken.

Sinonatrix percarinata (Boulenger, 1899)

Mountain water snake; Ran hoa can van den

Examined material. Specimen examined (n =

1): one juvenile specimen collected by KVN and

others (ZFMK 94671, SVL: 198.9 mm, TAL: 68.4

mm; Fig. 33), ratio TAL/TL: 0.26.

Morphological characters. Body stout; eye

moderate; pupil round; nostrils directed slightly

upwards, not valvular; rostral 2 times wider than

high; nasal completely divided; internasals longer

than wide, distinctly narrowed anteriorly, longer

than the prefrontals; loreal as long as high; pre-

ocular 1; pos-toculars 3; suboculars 2; temporals 2

or 3 + 3; supralabials 9, fourth and fifth touching

the eye; dorsal scale rows: 19 : 17 : 17, keeled,

except outermost row; ventrals 136, more than

twice as broad as adjacent dorsals; cloacal scale di-

vided; subcaudals 76, paired (determination after

Bourret, 1936; Smith, 1943; Ziegler, 2002).

The colouration of the preserved specimen is

greyish brown above with 28 vertical black bars

on each side, not symmetrical, getting wider

towards the back, where all are confluent, dark

bars in anterior part of body ending in the middle

of ventrals, forming broken rings in width of 2 or

3 scale rows; upper head dark olive brown, supral-

abials white posteriorly; chin and venter white,

except for the black rings.

Distribution. Vietnam: From Lao Cai, Ha

Giang, and Cao Bang in the North, southwards to

Kon Turn, Gia Lai, and Dong Nai provinces (this is

the first record of the species for Quang Ngai

Province); elsewhere: India, China, Taiwan, Myan-

mar, Laos, and Thailand.

Remarks. The colouration of the specimen is

typical for young individuals (see Smith, 1943). Ac-

First contribution to the reptile fauna of Quang Ngai Province, central Vietnam

319

Figure 25. Uncollected Dendrelaphis ngansonensis, 23.VI.2009, feeding on Polypedates sp., photo T. Geismann. Figure 26.

Preserved Enhydris subtaeniata (IEBR 3289), photo R. Babb. Figure 27. Oligodon chinensis, photo R. Babb. Figure 28. Pareas

hamptoni, photo R. Babb. Figure 29. P. margaritophorus , photo R. Babb. Figure 30. Pseudoxenodon macrops, photo R. Babb.

320

Liza Nemes et alii

cording to the key of Ziegler et al. (2007), two

supralabials entering the eye are typical for Sino-

natrix percarinata, as well as vertical or Y-shaped

black bars on the body sides, whereas the number

of these dark body marks (fewer than 30) is typi-

cal for Sinonatrix aequifasciata (which, however,

in contrast normally has not more than one supral-

abial entering the eye, and which has broad, roun-

ded X-shaped dark marks on sides of body

instead).

Xenochrophis flavipunctatus (Hallowell, 1861)

Yellow- spotted keelback; Ran nuoc dom vang

Examined material. Specimen examined (n =

1): one adult male collected by KVN and others

(ZFMK 94669, SVL: 370 mm, TAL: 172.8 mm;

Fig. 34), ratio TAF/TF: 0.32.

Morphological characters. Body stout;

pupil round; eye shorter than its distance from the

nostril; nostril slightly directed upwards, not

valvular; rostral visible from above; contiguous in-

temasals strongly narrowed towards the front, sub-

triangular, with the anterior angle truncated, nearly

as long as prefrontals; frontal constricted in the

middle, 2.3 times as long as broad, longer than its

distance from the snout tip, a little shorter than

pari-etals; loreal as long as high; preocular 1;

postoculars 3; temporals 2 + 2 ; supralabials 9,

third to fourth or to fifth touching the eye; infral-

abials 5, in contact with anterior chin shields; men-

tal groove present; dorsal scales in 19 rows at

midbody, more or less strongly keeled, 3 outermost

rows smooth; ventrals 128, more than twice as broad

as adjacent dorsals; cloacal scale divided; subcau-

dals 83, divided (determination after Bourret, 1936;

Smith, 1943; Ziegler, 2002; Vogel & David, 2006).

The colouration of the preserved specimen is

pale olive above, with black spots, spots larger

laterally; a pale, dark V-shaped marking on nape;

two well-defined subocular streaks present on

neck, posterior one extending from the eye to the

corner of the mouth; ventral surface yellow,

ventral and subcaudal scales with entire, broad,

dark margins.

Distribution. Vietnam: entire country; else-

where: Bangladesh, China, Myanmar, Faos, Thai-

land, Cambodia, Malaysia, and Indonesia.

VIPERIDAE

Protobothrops mucrosquamatus (Cantor, 1839)

Brown-spotted pitviper / Ran luc cuom

Examined material. Specimen examined (n =

1): one subadult specimen collected by KVN and

others (ZFMK 94673, SVF: 437.3 mm, TAF: 90.6

mm; see Fig. 35), ratio TAF/TF: 0.17.

Morphological characters. Head elongate;

upper head scales unequal, extremely small, granu-

lar, smooth; snout 2.75 times as long as eye diam-

eter; rostral slightly broader than high; supraoculars

long and narrow, entire; intemasals separated from

each other by 3 small scales, separated from

supra-oculars by two enlarged scales; scales

between the supraoculars 15; postoculars 4, small;

subocular 1 , separated from the labials by 2 scale

rows; temporal scales 2 or 3 rows, enlarged, smooth;

supralabials 10, the first completely separated from

the nasal, the second forming the anterior margin of

the loreal pit, the third largest; dorsal scales keeled;

dorsal scale rows 31 : 28-30 : 24; ventrals 232;

cloacal scale undivided; subcaudals 96, divided (de-

termination after Bourret, 1936; Smith, 1943; Fe-

viton et al., 2003).

The colouration of the preserved specimen is

pale brown above, with a series of large, dark-

brown blotches; lower lateral with a series of

smaller blotches, all edged in black; a dark brown

line from the eye to the angle of the mouth, edged in

black; ventral surface brownish with white blotches.

Distribution. Vietnam: From Fao Cai, Ha

Giang, and Cao Bang provinces in the North east-

wards to Quang Ninh and southwards to Kon Turn,

and Gia Fai provinces (this is the first record of the

species from Quang Ngai Province); elsewhere:

India, Bangladesh, China, Taiwan, and Myanmar.

Remarks. In contrast to the descriptions of Smith

(1943) and Feviton et al. (2003) all the head scales of

the specimen are smooth. The number of dorsal scale

rows, ventrals, and subcaudals is somewhat higher

than in the previous descriptions. The ratio TAF/TF

is smaller than in Smith's description (1943). The

intemasals are only 2 times, not 5 to 10 times longer

than the adjacent scales (see Feviton et al., 2003).

Trimeresurus albolabris Gray, 1842

White-lipped pitviper / Ran luc mep trang

Examined material. Specimen examined (n =

First contribution to the reptile fauna of Quang Ngai Province, central Vietnam

321

Figure 31. Rhabdophis subminiatus, photo R. Babb. Figure 32. Sibynophis collaris, photo T. Ziegler. Figure 33. Sinonatrix

percarinata, photo R. Babb. Figure 34. Preserved Xenochrophis flavipunctatus (IEBR 3297), photo R. Babb. Figure 35.

Protbothrops mucrosquamatus , photo R. Babb. Figure 36. Trimeresurus albolabris , photo R. Babb.

322

Liza Nemes et alii

1) : one adult male collected by KVN and others

(ZFMK 94670, SVL: 485.2 mm, TAL: 135.9 mm;

see Fig. 36), ratio TAL/TL: 0.22.

Morphological characters. Pupil vertical;

snout 2.4 times as long as eye diameter; loreal pit

between nostril and eye; canthus rostralis sharp,

rostral wider than high; scales on top of head small,

smooth, subequal, subimbricate; scales between

supraoculars 12; intemasals about 3 times larger

than adjacent scales, in contact with each other;

preoculars 3; postoculars 2, small; subocular 1, in

contact with third labial; scale rows between sub-

ocular and labials 1-3; temporal scales feebly

keeled; supralabials 9-10, not in contact with the

eye, the first partially fused to the nasal, the second

bordering loreal pit, the third largest; infralabials

12-13; dorsal scales keeled; dorsal scale rows 20 :

21 : 15; ventrals 166, more than twice as broad as

the adjacent dorsals; cloacal scale entire; subcau-

dals 76, divided (determination after Bourret, 1936;

Smith, 1943; Manthey & Grossmann, 1997; Zie-

gler, 2002; Leviton et al., 2003).

The colouration of the preserved specimen is

blue above; supralabials pale blue; ventral surface

yellowish blue.

Distribution. Vietnam: From Lao Cai and Cao

Bang provinces in the North southwards to Dong

Nai, Tay Ninh, Kien Giang, and Ca Mau provinces

(this is the first record of the species from Quang

Ngai Province); elsewhere: Northeast India, China,

Myanmar, Laos, Thailand, and Cambodia.

Trimeresurus vogeli David, Vidal et Pauwels,

2001

VogeFs green pitviper; Ran luc von-gen

Examined material. Specimens examined (n =

2) : one adult female collected by QKL and KLV

(ZFMK 94275, SVL: 462 mm, TAL: 83 mm); one

adult male collected by KVN and others (IEBR

3305, SVL: 508.8 mm, TAL: 119 mm; Fig. 37), tail

average, ratio TaL/TL of 0.15 in the female.

Morphological characters. Dorsum green; a

white lateral stripe, edged in red below in males,

and a narrow yellow stripe without red in females;

a constant lack of red color in the postocular streak

in both males and females; not more than 25 % of

tail tip rusty red; tail average, with a ratio TaL/TL

of 0.15 in the female; labial region somewhat

lighter greenish than green head colouration; first

supralabial totally separated from the corresponding

nasal; intemasals separated from each other by 0-2

scales; dorsal scales 21 rows at midbody, strongly

keeled; ventrals 155-158, plus 2-3 preventrals;

cloacal entire; subcaudals paired, 60-67; hemipenis

short and strongly spinose (determination after

David et al., 2001; Ziegler et al., 2007).

Distribution. Vietnam: from Ha Tinh and

Quang Binh southwards to Dak Lak and Lam Dong

provinces (this is the first record of the species from

Quang Ngai Province); elsewhere: Laos, Thailand,

Cambodia.

Remarks. The specimens differ somewhat from

the original description of David et al. (2001) by

having a lower ventral count (155-158 versus 163-

173); the head streak is not distinct; the eyes are

greenish orange instead of greenish yellow to

yellow; the internasals in the male IEBR 3305 are

not separated (usually separated by 1-3 scales), and

the dorsum bears no faint dark crossbands and no

white vertebral spots.

TESTUDINES

GEOEMYDIDAE

Cyclemys pulchristriata Fritz, Gaulke et Lehr,

1997

Striped leaf turtle; Rua dua soc

Examined material. Specimens photographed

(n = 2): two subadults were photographed by QKL

and RB (see Figs. 38-40).

Morphological characters. Identification of

the photographed specimens is based on character-

istic scalation and colour pattern, such as plastron

entirely yellow or prevalent plastral colour yellow;

femoral mid-seam shorter than cloacal mid-seam;

cloacal notch small to wide and acute-angled to

obtuse-angled; throat uniformly white coloured;

light head and neck stripes (yellow to salmon in live

specimens) wide; if black radiating pattern present

on plastron, radiating lines short and stout (deter-

mination after Fritz et al., 2008).

Distribution. Vietnam: Quang Nam, Gia Lai, and

Lam Dong provinces (this is the first record of the

species from Quang Ngai Province); elsewhere: ac-

cording to Fritz et al. (2008) easternmost Cambodia.

First contribution to the reptile fauna of Quang Ngai Province, central Vietnam

323

Figure 37. Trimeresurus vogeli. Figure 38. Cyclemys pulchristriata. Figure 39. Cyclemys pulchristriata, portrait. Figure 40.

Cyclemys pulchristriata , ventral view. Photos R. Babb.

DISCUSSION

We herein provide a list of 35 identified reptile

species (16 lizards, 18 snakes, and 1 turtle) based

on our recent field surveys in Quang Ngai Province.

Fourteen of the recorded species belong to the

family Colubridae, which is the most species-rich

reptilian family in Quang Ngai, followed by 6

agamid and 6 scincid species. Nguyen et al. (2009)

also listed Oligodon ocellatus as occurring in

Quang Ngai Province, which brings the total

reptilian number known from that province to 36.

Nguyen et al. (2009) further list the terrestrial rep-

tilian species Gekko gecko , Coelognathus radiatus,

and Enhydris plumbea as occurring in the whole

country, so that there is high probability that they

also will be recorded from Quang Ngai Province in

the future. Regardless, G. gecko from Vietnam

recently has been revised, and it has not yet been

determined whether G. gecko or G. reevesii is

involved (see Rosier et al., 201 1). A further number

of species are listed in Nguyen et al. (2009) as oc-

curring along Vietnam’s central coasts, but without

definite records for Quang Ngai Province. The data

presented herein thus serve only as a preliminary

reptilian list, which must be extended in the future

based on further field research. Also the taxonomic

status of Fimbrios cf. klossi from Quang Ngai needs

further examination. There exist clear differences

between the available specimen and known varia-

tion of F. klossi. The combination of the number of

temporals, the lack of a mental, and the longer tail

324

Liza Nemes et alii

would suggest a distinct species. However, because

only one individual is available at this time, an

anomalous specimen cannot be excluded with

certainty. Together with the 1 6 amphibian species

recorded from this area (Tran et al., 2010), the her-

petofaunal list for Quang Ngai Province currently

covers 52 amphibian and reptile species. Compared

with the herpetofaunal composition of other central

Vietnamese provinces, such as Quang Binh Prov-

ince (see Ziegler & Vu, 2009), species numbers

could easily be doubled to tripled in the near future.

Such further research also will be essential for

evaluating the conservation status of Quang Ngai

forested areas, in particular as protected areas have

not yet been established in this province until today.

ACKNOWLEDGEMENTS

Field surveys in Quang Ngai Province were

assisted by Khoi Le Vu, Tuan Thanh Nguyen, Tran

Anh Vu, Ab Abercrombie, Chris Hope, Paul Moler,

and Bill Turner. We are grateful to the directorates

of the Forest Protection Department of Quang Ngai

Province for the support of our field work and is-

suing relevant permits. For help with species iden-

tification we are indebted to Patrick David, Paris

(. Amphiesma , Fimbrios ), and Ulrich Manthey, Ber-

lin ( Calotes ). For the loan of specimens, we thank

Canh Xuan Le, Thai Huy Tran, Cuong The Pham

(IEBR, Hanoi) and Tao Tien Nguyen (VNMN,

Hanoi). Thanks to Eleanor Sterling (New York) and

Kevin Koy (Berkeley) for providing the map, and

to Thomas Geismami for providing figure 25. Field

surveys in Quang Ngai were partially supported by

the Douc Langur Foundation (DLF) and Wildlife at

Risk (WAR).

REFERENCES

Bourret R., 1936. Les serpents de FIndochine. I. Etudes

sur la faune. Henry Basuyau et Cie, Toulouse, 505 pp.

Bourret R., 2009. Les lezards de FIndochine. Edition

Chimaira, Frankfurt am Main, 624 pp.

Darevsky, I.S. & Kupriyanova L.A., 1993. Two new all-

female lizard species of the genus Leiolepis Cuvier,

1829 from Thailand and Vietnam (Squamata: Sauria:

Uromastycinae). Herpetozoa, 6: 3-20.

David P., Bain R.H., Nguyen, T.Q., Orlov N.L., Vogel G.,

Vu T.N. & Ziegler T., 2007. A new species of the

natricine snake genus Amphiesma from the Indochi-

nese Region (Squamata: Colubridae: Natricinae).

Zootaxa, 1462: 41-60.

David R, Nguyen T.Q., Nguyen T.T., Jiang K., Cchen T.,

Teynie A. & Ziegler T., 2012. A new species of the

genus Oligodon Fitzinger, 1826 (Squamata: Colubri-

dae) from northern Vietnam, southern China and

central Laos. Zootaxa, 3498: 45-62.

David P., Vidal N. & Pauwels O.S.G., 2001. A morpho-

logical study of Stejneger's pitviper Trimeresurus

stejnegeri (Serpentes, Viperidae, Crotalinae), with the

description of a new species from Thailand. Russian

Journal of Herpetology, 8: 205-222.

David P., Vogel G. & Dubois A. 201 1 . On the need to fol-

low rigorously the Rules of the Code for the subse-

quent designation of a nucleospecies (type species)

for a nominal genus which lacked one: the case of the

r \

nominal genus Trimeresurus Lacepede, 1804 (Rep-

tilia: Squamata: Viperidae). Zootaxa, 2992: 1-51.

Fritz U., Guicking D., Auer M., Sommer R.S., Wink M.

& Hundsdorfer A.K., 2008. Diversity of the South-

east Asian leaf turtle genus Cyclemys : how many

leaves on its tree of life? Zoologica Scripta, 37: 367-

390.

Green M.D., 2010. Molecular phylogeny of the snake

genus Oligodon (Serpentes: Colubridae), with an

annotated checklist and key. Master thesis, University

of Toronto, 161 pp.

Gressit J.L., 1937. Anew snake from southeastern China.

Proceedings of the Biological Society of Washington,

50: 125-128.

Grismer J.L. & Grismer L.L., 2010. Who’s your

mommy? Identifying maternal ancestors of asexual

species of Leiolepis Cuvier, 1 829 and the description

of a new endemic species of asexual Leiolepis

Cuvier, 1829 from Southern Vietnam. Zootaxa, 2433:

47-61.

Hartmann T., Geissler P., Poyarkov N.A., Ihlow F., Ga-

loyan E.A., Rodder D. & Bohme W., 2013. A new

species of the genus Calotes Cuvier, 1817 (Squamata:

Agamidae) from southern Vietnam. Zootaxa, 3599:

246-260.

Kams D.R., Lukoschek V., Osterhage J., Murphy J.C. &

Voris H.K., 2010. Phylogeny and biogeography of the

Enhydris clade (Serpentes: Homalopsidae). Zootaxa,

2452: 18-30.

Leviton A.E., Wogan G.O.U., Koo M.S., Zug G.R.,

Lucas R.S. & Vindum J.V., 2003. The dangerously

venomous snakes of Myanmar. Illustrated checklist

with keys. Proceedings of the California Academy of

Sciences, ser. 4, 54: 407-460.

Luu V.Q., Nguyen T.Q., Pham C.T., Dang K.N., Vu T.N.,

Miskovic S., Bonkowski M. & Ziegler T., 2013. No

end in sight? Further new records of amphibians and

reptiles from Phong Nha - Ke Bang National Park,

First contribution to the reptile fauna of Quang Ngai Province, central Vietnam

325

Quang Binh Province, Vietnam. Biodiversity Journal,

4: 285-300.

Manthey U. & Grossmann W., 1997. Amphibien und

Reptilien Sudostasiens. Natur und Tier-Verlag, Mun-

ster, 512 pp.

Murphy J.C., 2007. Homalopsid snakes: evolution in the

mud. Krieger, 249 pp.

Murphy J. C. & Voris H. K., 1994. A key to the homa-

lopsine snakes. The Snake, 26: 123-133.

Musters C.J.M., 1983. Taxonomy of the genus Draco L.

(Agamidae, Lacertilia, Reptilia). Zoologische Ver-

handelingen, 199: 1-120.

Nazarov R., Poyarkov N.A., Orlov N.I., Phung T.M.,

Nguyen T.T., Hoang D. M. & Ziegler T. 2012. Two

new cryptic species of the Cyrtodactylus irregularis

complex (Squamata: Gekkonidae) from southern

Vietnam. Zootaxa, 3302: 1-24.

Nguyen S.V., Ho C.T., Nguyen T.Q., 2009. Herpetofauna

of Vietnam. Edition Chimaria, Franfkurt am Main,

768 pp.

Nguyen T.Q., Bohme W., Nguyen T.T., Le Q.K., Pahl K.

R., Haus T. & Ziegler T., 2011a. Review of the genus

Dopasia Gray, 1853 (Squamata: Anguidae) in the In-

dochina subregion. Zootaxa, 2894: 58-68.

Nguyen T.Q., Nguyen S.V., Orlov N.L., Hoang T.N.,

Bohme W. & Ziegler, T., 2010. A review of the

genus Tropidophorus (Squamata, Scincidae) from

Vietnam with new species records and additional

data on natural history. Zoosystematics and Evolu-

tion, 86: 5-19.

Nguyen T.Q., Schmitz A., Nguyen T.T., Orlov N.L.,

Bohme W. & Ziegler T., 20 1 lb. 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 Tay-

lor, 1962 from Vietnam. Journal of Herpetology, 45:

145-154.

Orlov N.L., Truong N.Q. & Sang N.V., 2006. A new

Acanthosaura allied to A. capra Gunther, 1861 (Aga-

midae, Sauria) from central Vietnam and southern

Laos. Russian Journal of Herpetology, 13: 61-76.

Pope C.H., 1935. The reptiles of China. Turtles,

crocodilians, snakes, lizards. Natural History of cen-

tral Asia. Vol. X. American Museum of Natural Hi-

story, New York, 604 pp.

Rosier H., Bauer A.M., Heinicke M.P., Greenbaum E.,

Jackman T., Nguyen T.Q. & Ziegler T., 2011. Phy-

logeny, taxonomy, and zoogeography of the genus

Gekko Laurenti, 1768 with the revalidation of G. ree-

vesii Gray, 1831 (Sauria: Gekkonidae). Zootaxa,

2989: 1-50.

Rosier H., Vu T.N, Nguyen T.Q., Ngo T.V. & Ziegler T.,

2008. Anew Cyrtodactylus (Squamata: Gekkonidae)

from central Vietnam. Hamadryad, 33: 48-63.

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, including the whole of the Indo-Chinese

Subregion. Reptilia and Amphibia. Vol. III. Ser-

pentes. Taylor and Francis (London), 583 pp.

Taylor E.H., 1963. The lizards of Thailand. University of

Kansas Science Bulletin, 44: 687-1077.

Tran D. T. A., Le K. Q„ Le K. V., Vu T. N., Nguyen T.

Q., Bohme W. & Ziegler T., 2010. First and prelim-

inary frog records (Amphibia: Anura) from Quang

Ngai Province, Vietnam. Herpetology Notes, 3:

111-119.

Vogel G. & David P, 2006. On the taxonomy of the Xe-

nochrophis pis cat or complex (Serpentes, Natricidae).

In: Vences M., Kohler J., Ziegler T. & Bohme W.

(Eds.). Herpetologia Bonnensis II: Proceedings of the

13th Congress of the Societas Europaea Herpetolog-

ica, Bonn, pp. 241-246.

Wood P.L., Grismer L.L., Grismer J.L., Neang T., Chav

T. & Holden J., 2010. Anew cryptic species of Acan-

thosaura Gray, 1831 (Squamata: Agamidae) from

Thailand and Cambodia. Zootaxa, 2488: 22-38.

Ziegler T., 2002. Die Amphibien und Reptilien eines Tie-

flandfeuchtwald-Schutzgebietes in Vietnam. Natur &

Tier Verlag, Munster, 342 pp.

Ziegler T., David P., Miralles A., Dang K.N., Nguyen

T.Q., 2008. A new species of the snake genus Fim-

brios from Phong Nha - Ke Bang National Park,

Truong Son, central Vietnam (Squamata: Xenoder-

matidae). Zootaxa, 1729: 37-48.

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: i_40.

Ziegler T. & Nguyen T.Q., 2010. New discoveries of am-

phibians and reptiles from Vietnam. Bonn zoological

Bulletin, 57: 137-147.

Ziegler T., Ohler A., Vu T.N., Le Q. K., Nguyen T.X.,

Dinh T.H. & Bui T.N, 2006. Review of the amphib-

ian and reptile diversity of Phong Nha - Ke Bang Na-

tional Park and adjacent areas, central Truong Son,

Vietnam. In: Vences M., Kohler J., Ziegler T. &

Bohme W. (Eds.): Herpetologia Bonnensis II. Pro-

ceedings of the 13th Ordinary General Meeting of

the Societas Europaea Herpetologica, Bonn, pp. 247-

262.

Ziegler T., Orlov N.L., Giang T., Nguyen T.Q., Nguyen

T.T., Le Q.K., Nguyen K.V. & Vu T.N., 2010. New

provincial records of cat snakes, Boiga Fitzinger,

1826 (Squamata: Serpentes: Colubridae), from Viet-

nam, inclusive of an extended diagnosis of Boiga

326

Liza Nemes et alii

bourreti Tillack, Le & Ziegler, 2004. Zoosystematics

and Evolution, 86: 263-274.

Ziegler T. & Vogel G., 1999. On the knowledge and spe-

cific status of Dendrelaphis ngansonensis (Bourret,

1935) (Reptilia: Serpentes: Colubridae). Russian

Journal of Herpetology, 6: 199-208.

Ziegler T. & Vu T.N., 2009. Ten years of herpetodiversity

research in Phong Nha - Ke Bang National Park, cen-

tral Vietnam. In: Vo Van Tri, Nguyen Tien Dat, Dang

Ngoc Kien & Pham Thi Hai Yen (Eds.): Phong Nha -

Ke Bang National Park and Cologne Zoo, 10 years of

cooperation (1999-2009). Quang Binh, pp. 103-124.

Biodiversity Journal, 2013, 4 (2): 327-334

Foraging behaviour of three primate species in a Costa Rican

coastal lowland tropical wet forest

Kirsty E. Graham 1 *, Megan J. Bulloch 2 &Todd R. Lewis 3

'Quest University Canada, 3200 University Blvd, Squamish, BC, V8B 0N8, Canada; email: kirsty.graham@questu.ca

2 Quest University Canada, 3200 University Blvd, Squamish, BC, V8B 0N8, Canada; email: megan.bulloch@questu.ca

3 Estacion Biologica Cano Palma, Refugia Barra del Colorado, Tortuguero, Costa Rica; email: ecolewis@gmail.com

^Corresponding author

ABSTRACT Primates are predominantly distributed across tropical regions, many of which are threatened

by deforestation. Removal of mature trees can harm primate populations by reducing avail-

able food resources. Understanding the dietary requirements of primates at local levels can

help identify key habitats to conserve, and protect plant species on which primates rely. Little

is known about local diets of Alouatta palliata (mantled howler monkey), Ateles geoffroyi

(black-handed spider monkey), and Cebus capucinus (white-faced capuchin) in Costa Rica's

lowland tropical wet forests. Because diet and activity levels are closely connected, studies

examining one provide insight into the other. We used group scan sample methods to record

activity and diet, identifying all plant species on which the primates fed. We identified nine

families of plants eaten by Ateles geoffroyi , four families eaten by Alouatta palliata, and

two families eaten by Cebus capucinus. Activity budgets demonstrated that Alouatta palliata

was the least active species and Cebus capucinus the most active. We also found differences

in the type of plant parts consumed by the three primate species; Alouatta palliata and Ateles

geoffroyi fed mostly on fruit and new leaves, whereas Cebus capucinus fed on fruit and in-

sects. The nine families of plants identified in this study are potentially important for all

three primate species locally, and warrant conservation.

KEY WORDS Primate; Activity budget; Foraging; Lowland wet forest; Costa Rica.

Received 7.04.2013; accepted 24.05.2013; printed 30.06.2013

INTRODUCTION

New World primates have evolved to fill spe-

cific niches through variable body size, activity lev-

els, and dietary preference (Rosenberger, 1992).

However, these distinct niches may be contingent

on intact forest. Timber extraction increases inter-

species competition by narrowing the number of

preferred plant species, in addition to reducing the

total biomass of food resources. Studying basic pri-

mate natural history by recording aspects such as

diet can assist conservation land management pre-

scriptions at a local scale by identifying key habitats

and plants that primates use, and protecting them as

food resources.

In recent history, Costa Rica has suffered habitat

loss of up to 4% forest cover per annum, mostly

from expanding agriculture and human frontiers

(Myers et al., 2000; Sanchez- Azofeifa et al., 2001;

Achard et al., 2002). The country sought to redress

this issue by the formation of national parks, and

has recently expanded these to include wildlife cor-

ridors that connect reserve land to benefit primates,

among a myriad of diverse wildlife (Canet-Desanti

& Finegan, 2010). Costa Rica is biologically di-

328

K.E. Graham, M.J. Bulloch &T.R. Lewis

verse, and has four species of primates that live in

its tropical forests; Saimiri oerstedii (Reinhardt,

1872) (Central American squirrel monkey), Cebus

capucinus (Linnaeus, 1758) (white-faced capuch-

in), Alouatta palliata (Gray, 1849) (mantled howler

monkey), and Ateles geoffroyi (Kellogg et Gol-

dman, 1944) (Geoffroy's or black-handed spider

monkey). Despite their iconic status within Costa

Rica’s eco-tourism industry, surprisingly little is

known about the diet of A. palliata, Ateles geoffroyi,

and C. capucinus among these reserve lands, espe-

cially at local levels.

Group size and social structure can mitigate

intra- and inter-species competition for food re-

sources. Mantled howler monkeys form large, sta-

ble groups ranging from 2-39 individuals (Crockett

& Eisenberg, 1987). They can live in large groups

without intense food competition because their fit-

ness is limited by access to mates rather than food;

leaves are an abundant and easily accessible food

source (Di Fiore & Campbell, 2007). Species with

a higher proportion of fruit in their diet often use

grouping strategies that allow access to this widely

dispersed food source. Black-handed spider mon-

keys have a fission- fusion social structure (Di Fiore

& Campbell, 2007), and live in groups of 15-25 in-

dividuals, forming sub-groups of 3 individuals (Ro-

binson & Janson, 1987). Fission- fusion social

structures may be a way of avoiding direct compe-

tition over food resources with conspecifics, by dis-

persing in small groups to scattered fruit sources

(Di Fiore & Campbell, 2007). White-faced ca-

puchins also feed predominantly on fruit, and form

mixed-sex groups of 10-35 individuals (Robinson

& Janson, 1987; Jack, 2007). Farge group size

might be a way to increase frequency of detecting

fruit sources and defending these sources against

other groups (Janson, 1988). Capuchins’ smaller

body size means that they have to intake fewer calo-

ries, mitigating competition within each group.

This study was performed at Cano Palma Bio-

logical Station, 8 km north of the coastal village of

Tortuguero, in NE Costa Rica. Cano Palma is a

small 5,000 m 2 reserve of mixed lowland tropical

wet forest with dominant stands of Manicaria and

Raphia palms (Fewis et al., 2010). The main objec-

tive of this study was to identify plant species and

parts consumed by A. palliata, Ateles geoffroyi, and

C. capucinus. In addition, each primate species was

monitored for activity budget and food competition,

to create an ethogram to determine whether they fed

on overlapping plant species and plant parts.

MATERIALS AND METHODS

The study period lasted 28 survey days from 8

April to 20 May 2011. Each day, the study site was

surveyed commencing at 06:00. The first located

species of primate was followed and monitored

through the day until nightfall or until they became

inaccessible. Observational frequency of all three

primate species was adjusted in the field to ensure

equal time budgets were allocated to each species.

Namely, if one species had been followed more

often than another, but was the first to be located

during a survey, this species was immediately fol-

lowed until an encounter with a less frequently ob-

served species. For each data point, GPS coordinates

were recorded to create distributional maps of each

primate species across the site.

Activity budget data were collected using a

group scan sample method (Martin & Bateson,

1993). At two-minute intervals the activity of each

individual was recorded as one of four categories;

inactive, travelling, feeding, and social interaction

(Pavelka & Knopff, 2004). The age and sex of each

individual was also recorded. Juveniles were not di-

vided by sex, and adults of unknown sex were not

included in analysis. Infants were excluded from ac-

tivity budgets because of dependence on mothers

for food (Pavelka & Knopff, 2004). When an indi-

vidual was feeding, the species of plant and/or plant

part was recorded. The five categories of plant parts

were new leaf, mature leaf, unknown leaf, fruit, and

stalk (Pavelka & Knopff, 2004). Plant parts that

could not be identified were recorded as 'unknown'.

Unknown plant species were included in the analy-

sis of activity budget and plant parts, but discounted

in analysis of plant species. For insectivores, such

as C. capucinus, the number of insect feeding bouts

was recorded, but the species of insect was not iden-

tified due to the complexity of deciphering specific

species for each insect.

Data analyses of activity budgets and food bud-

gets were conducted using R (version 2.11.1). Food

budgets were sub-divided into separate analyses for

plant species and plant parts. Individual identifica-

tion of primates was not possible, and therefore, re-

peat sampling of some groups likely occurred. We

Foraging behaviour of three primate species in a Costa Rican coastal lowland tropical wet forest

329

attempted to control for repeat sampling by using

group scan sampling. The time budget spent for

each activity was measured as a proportion of the

time observed (Pavelka & Knopff, 2004). There-

fore, time spent by a primate on an activity, the de-

pendent variable, was measured as a percentage.

Activity budget analyses used a 3 (species) x 2

(age) x 2 (sex) three-way ANOVA on mean fre-

quency of each activity. Variation in plant parts con-

sumed was analyzed by a 2 (sex) x 2 (age) x 3

(species) three-way ANOVA on mean frequency of

type of plant part consumed.

RESULTS

The dataset comprised 28 survey days, of which

monkeys were observed on 1 7 days, for a total of

approximately 49 hours. Totals of 933 A. palliata ,

368 Ateles geoffroyi , and 156 C. capucinus obser-

vations were recorded. Individual monkeys often

provided multiple observations upon encounter.

Population and Distribution

A. palliata were found in groups of 3-4 individ-

uals, Ateles geoffroyi in groups of 3-7 individuals,

and C. capucinus in groups of 12-20 individuals.

Distribution maps for each primate species indicated

that most groups were located, and foraged, in edge

and transitional (occasionally flooded) habitat (Fig.

1). Only A. palliata were found farthest north of the

reserve among Manicaria swamp habitat (Fig. 1).

Activity Budgets

Overall activity budget for A. palliata, including

all age and sex groups, indicated they spent 77% of

time inactive, 14% moving, 7% feeding, and 2%

engaging in social behaviour. Ateles geoffroyi spent

41% of time moving, 31% inactive, 20% feeding,

and 8% engaging in social behaviour. C. capucinus

spent 76% of time moving, 10% engaging in social

behaviour, 8% feeding, and 6% inactive (Fig. 2). A

3 (species) x 2 (age) x 2 (sex) three-way ANOVA

on mean frequency of each activity produced a sta-

tistical interaction between activity and primate

species. The percentage of time spent on each activ-

ity differed significantly between primate species,

F (6, 192) = 8.2069, P = 0.001 (Fig. 2).

Diet

For time engaged in feeding, A. palliata spent

35% feeding on new leaves, 32% on fruit, 30% on

unknown leaves, 2% on stalks, and 2% on unknown

plant parts. Ateles geoffroyi spent 44% feeding on

new leaves, 28% feeding on fruit, 17% on unknown

leaves, 11% on plant stalks, and 1% on mature

leaves. C. capucinus spent 77% of the time feeding

on fruit, 15% on insects, and 8% on unknown parts

(Fig. 3). A 2 (sex) x 2 (age) x 3 (species) three-way

ANOVA on mean frequency of type of plant part

consumed produced an interaction between plant

part and primate species. Primate species ate differ-

ent plant part, F (10, 291) = 2.347, p = 0.0123, but

not different plant families (Table 1).

N ine families of plants were identified as food for

A. geoffroyi ; Anacardiaceae, Araceae, Arecaceae,

Clusiaceae, Melastomaceae, Moraceae, Myrtaceae,

Rubiaceae and Tiliaceae. A. palliata fed on four plant

families (Anacardiaceae, Araceae, Arecaceae, Clu-

siaceae), and C. capucinus fed on two plant families

(Anacardiaceae and Melastomaceae) (Table 1).

DISCUSSION

Population and Distribution

All three species of primate were observed

around Cano Palma’s reserve land without disturb-

ing normal activity. This suggested that all three

species were habituated to human presence, likely

due to the long-term presence of researchers at

Cano Palma Biological Station and regular eco-

tourism to the area.

Cano Palma Biological Station’s reserve land

features lowland tropical wet forest edge, lowland

tropical wet transitional (occasionally flooded)

forest, and Manicaria swamp forest (Lewis et al.,

2010). All three species of primates in this study

were located, and observed foraging among edge

and transitional vegetative zones within Cano Pal-

ma’s reserve land. Only A. palliata was located in

Manicaria swamp forest habitat. This is possibly

due to the higher diversity of food providing trees

and high vegetation within the transitional forest

zone (Lewis et al., 2010). The observations of group

numbers of A. palliata, Ateles geoffroyi, and C. ca-

330

K.E. Graham, M.J. Bulloch &T.R. Lewis

Population Distribution Over Time

Legend

Ecoregion

Edge Habitat

Manicaria Forest

Occasional Flooding

Primate Distribution over Time (days)

earliest data

mid data

latest data

0 75 150

Meters

Cebus capucinus

Ateles geoffroyi

Alouatta palliata

Figure 1 . Distribution of each primate species across the study site as a function of time.

pucinus concurred with known group numbers for

each species. A. palliata were found in groups of 3-

4 individuals, within the reported range of 2-39 in-

dividuals (Crockett & Eisenberg, 1987). Ateles

geoffroyi were found in small groups of 3-7 indi-

viduals that were possibly sub-groups, although

they have been observed at Cano Palma Biological

Station in larger groups (P. Grant & R. Ballard, pers.

comm.). Ateles geoffroyi are usually found in fis-

sion-fusion structured social groups of 15-25 indi-

viduals, and form sub-groups of up to three

individuals (Robinson & Janson, 1987; Di Fiore &

Campbell, 2007). These fission- fusion social struc-

tures may be a way of avoiding direct competition

over food resources with conspecifics (Di Fiore &

Campbell, 2007). C. capucinus groups ranged from

12-20 and are known to form mixed-sex groups of

10-35 individuals (Jack, 2007; Robinson & Janson,

1987). C. capucinus were only observed on 3 non-

consecutive days of 28 survey days, travelling

through the study site. It is possible that Cano Pal-

ma's reserve is only a portion of this group's range.

Foraging behaviour of three primate species in a Costa Rican coastal lowland tropical wet forest

331

Activity budget for three monkey species

100

■AP

D AG

°CC

s, so

Feeding Inactive Moving

Activity

Social

Food budget

100

Plant Part

■Mantled Howler

Monkeys

0 Black-handed

Spider Monkeys

°White-faced

Capuchins

J] .0 0

Figure 2. Activity budget by species (Ateles geoffroyi -

black, Alouatta palliata - white, Cebus capucinus - grey);

percentage of total time observed performing each activity.

The percentage of time spent on each activity differed sig-

nificantly between primate species, F (6, 192) = 8.2069,

P = 0.001.

Figure 3. Food budget by species ( Ateles geoffroyi - black,

Alouatta palliata - white, Cebus capucinus - grey); percent-

age of total time observed feeding on each plant part.

Table 1. Food budget by species, age, and sex; percentage

of total time observed feeding on each plant species.

Species

Ateles geoffroyi

Alouatta palliata

Cebus

capucinus

Age

All

Sex

All

Unk

All

Unk

All

Unk

Anacardiaceae

Araceae

Arecaceae

Clusiaceae

Melastomataceae

100

Moraceae

Myrtaceae

Rubiaceae

Tiliaceae

Unknown

100

Insect

During this study, little interspecific competition

was observed between primate species despite their

overlapping distribution. This is possibly because

each species foraging strategy led them to feed on

different plant parts and plant species. It is possible

that each primate species filled a specific niche, al-

though a year-long study would clarify this concept

and determine whether competition increases dur-

ing periods of lower food availability.

Activity Budgets and Food Choice

A. palliata , Ateles geoffroyi, and C. capucinus

exhibited different activity budgets, and fed on dif-

ferent plant parts to sustain activity. Differing activ-

ity budgets are known for all three species

(Schoener, 1971; Crockett & Eisenberg, 1987; Di

Fiore & Campbell, 2007). A. palliata was observed

to consume mostly leaves and fruit, and did not

332

K.E. Graham, M.J. Bulloch &T.R. Lewis

travel as much as Ateles geoffroyi or C. capucinus.

These findings are consistent with the hypothesis

that A. palliata exhibits a strategy of energy mini-

mization. Namely, they receive less energy from the

leaves they eat, and therefore exert less energy (Di

Fiore & Campbell, 2007). A. palliata is primarily fo-

livorous, with a diet similar to colobines and in-

driids, but without a specialized digestive tract to

efficiently digest fibre (Milton, 1979). Although

their capacious gut allows A. palliata to digest

leaves, they do not extract the same amount of nu-

trients as their old world relatives (Milton, 1979).

Across primate species, diets high in leaves correlate

to low levels of activity (Dasilva, 1992). At Cano

Palma, A. palliata exhibited a preference for young

leaves, which may be because they contain higher

protein content, and are generally lower in fibre, thus

easier to digest (Milton, 1979). A. palliata can re-

main in stable social groups sharing the same area

without intense competition because leaves are

abundant and readily available in high density (Di

Fiore & Campbell, 2007). At Cano Palma we did not

observe intraspecific competition for food resources

either within or between howler monkey groups.

Ateles geoffroyi are known to move to specific

areas to exploit seasonally ripe fruit and can spend

between 18.9% and 50.5% of their day feeding (Ro-

binson & Janson, 1987). Spider monkeys in this

study ate fruit and leaves from a wide variety of plant

species, and were more active than A. palliata. Un-

like A. palliata , Ateles geoffroyi live in fission-fusion

societies, possibly as a response to their dietary pref-

erence for ripe fruit (Di Fiore & Campbell, 2007).

Because fruit is a densely but sparsely distributed re-

source, spider monkeys often divide into sub-groups

to avoid competition while foraging (Di Fiore &

Campbell, 2007). This was evident within Cano Pal-

ma’s spider monkey groups from the low group sizes

encountered. Ateles geoffroyi maximize energy in-

take by moving quickly to sources of high-energy

foods (Di Fiore & Campbell, 2007). To gain the

energy required to travel and forage; they must eat

fruit and, synchronously to eat fruit, they must travel

to foraging areas. At Cano Palma, Ateles geoffroyi

ranged constantly in the field as was observed in their

distribution. Ateles geoffroyi do not have the capa-

cious gut of A. palliata and therefore possibly cannot

extract sufficient calories from leaves alone.

C. capucinus can spend 70-80% of their day for-

aging and move seasonally to find ripe fruit (Robin-

son & Janson, 1987). Our observations showed C.

capucinus to be more active than Ateles geoffroyi ,

eating mainly fruit, as well as occasional insects. This

diet of high energy fruit and insects could allow

higher activity levels for capuchins. According to op-

timal foraging theory, individuals should select fruit

based on caloric/nutritional profitability even when

other familiar food sources are available (Schoener,

1971). Considering this, C. capucinus should con-

sume the available fruit with the highest energy con-

tent. A study to investigate capuchin seasonal diet at

Cano Palma would confirm this aspect. Diets bet-

ween neighbouring groups of C. capucinus can vary

and such differences in diet between groups at Cano

Palma could be cultural, intraspecific, or due to food

profitability (Chapman & Fedigan, 1990).

Diet Composition

In this study, the three primate species fed on a

total of nine families of plants, some of which over-

lapped across two, or all three species. The differ-

ence in plant families consumed may be explained

by the preferred plant parts of each primate species,

and possibly by plant distribution. A. palliata has

been known to consume mostly young leaves and

fruit, and the morphology of their teeth suggests

they have a predominantly folivorous diet (Crockett

& Eisenberg, 1987; Di Fiore & Campbell, 2007).

Past studies have shown that the proportion of fruit

in the diet of A. palliata affects their activity levels,

because fruit is a higher energy source than leaves

(Pavelka & Knopff, 2004). At Cano Palma, A. pal-

liata confirmed their known ecological behaviour

trait as inactive frugivore-folivores by their selec-

tion of vegetative food (Table 1).

Our observations of Ateles geoffroyi diet con-

firm their ecological behaviour as active frugivore-

folivores that prefer ripe fruit (Robinson & Janson,

1987; Di Fiore & Campbell, 2007). At Cano Palma,

Ateles geoffroyi were seen consuming Philodendron

sp. (Araceae). Philodendron is a genus known for

its toxic properties due to the presence of calcium

oxalate crystals (Genua & Hillson, 1985). Calcium

oxalate crystals are a secondary compound that de-

fends certain plant species against herbivores (Fran-

ceschi & Horner, 1980). Other mammal species,

with ruminating or complex guts, have been known

to eat high concentrations of oxalates (Freeland &

Janzen, 1974). Humans also consume some oxalate

Foraging behaviour of three primate species in a Costa Rican coastal lowland tropical wet forest

333

rich foods such as rhubarb, spinach, nuts, and tea,

and over-consumption can lead to adverse health ef-

fects (Massey et al., 1993). Ateles geoffroyi may

possess a tolerance to oxalate crystals that allow

them to consume calcium oxalate rich plants, such

as Philodendron spp., in conjunction with other

plant foods when preferred sources are scarce

(Franceschi & Homer, 1980). This food choice

could also be an act of self-medication (Huffman,

1997). Ateles geoffroyi in certain parts of Costa Rica

have lower parasite loads due to the ingestion of fig

leaves (Glander, 1994), and it is possible that cal-

cium oxalate acts as a similar vermicide.

C. capucinus confirmed their ecological behav-

iour as very active frugivore-insectivores (Robinson

& Janson, 1987). Their diet is known to be variable,

comprising mostly seasonal fruit or insects, and they

are known to hunt small vertebrates (Jack, 2007). At

Cano Palma, C. capucinus consumed species of plant

not atypically described for the species at other sites

across Costa Rica and the neotropics (Jack, 2007).

Habitat and Conservation for Primates at

Cano Palma

Factors such as variable body size and feeding

specializations may help New world primate species

to mitigate interspecific competition. Additionally,

each species' social structure can alleviate pressure

from intraspecific competition and resource scarcity

(Rosenberger, 1992). This self-regulating behaviour

among and within species of primate highlights the

importance of protecting areas that contain preferred

food plants, especially in connected habitats like

Cano Palma that buffer national reserves.

However, despite laws to regulate selective log-

ging within the Barra del Colorado Refuge, unmoni-

tored logging still occurs (Lewis et al., 2010; R.

Ballard, pers. comm.). We recommend that any se-

lective logging should avoid plant families and com-

munities of local provenance to maintain the integrity

of food resources for primates. Notwithstanding, reg-

ulation of selective logging on the vast Barra del Col-

orado Refuge is not always pragmatic. By studying

and considering local foraging by primates, a strategy

to conserve connected areas around Cano Palma

could be realised, and is suggested. Further study of

territory and distribution combined with disseminat-

ing information, and involvement of local commu-

nities, support this recommendation.

Figure 4. White faced capuchin. Copyright Jamie Bowkett.

Figure 5. Mantled Howler Monkey. Copyright Michelle

Haines. Fig. 6 Black-handed Spider Monkey. Copyright

Paul Grant.

334

K.E. Graham, M.J. Bulloch &T.R. Lewis

ACKNOWLEDGEMENTS

We thank the Canadian Organization for Tropi-

cal Education and Rainforest Conservation for al-

lowing use of the study site, particularly Dr.

Kymberley Snarr for her support of this project.

Ana Maria-Monge and Elena Vargas of the Mini-

sterio de Ambiente, Energia y Telecomunicaciones

permitted the study under license ACTo-GASP-

PIN-06-2011. Special thanks are extended to Ross

Ballard for plant identification and K. Julian Hoc-

kin-Grant for GIS plotting. We also thank the stu-

dents and volunteers who helped to collect data, and

the many friends involved in the editing process.

REFERENCES

Blanck Achard F., Eva H.D., Stibig H.-J., Mayaux P.,

Gallego J., Richards T. & Malingreau J.-P., 2 0 0 2 .

Determination of deforestation rates of the world’s

humid tropical forests. Science, 297: 999-1002.

Canet-Desanti L. & Finegan B., 2010. Bases de conoci-

miento para la gestion de corredores biologi-

cos en Costa Rica, Foundations of knowledge for the

management of biological corridors in Costa Rica.

Meso Americana, 14: 11-24.

Chapman C.A. & Fedigan L.M., 1990. Dietary differ-

ences between neighboring Cebus capucinus groups:

local traditions, food availability or responses to food

profitability. Folia Primatologica, 54: 177-186.

Crockett C.M. & Eisenberg J.F., 1987. Howlers: varia-

tions in group size and demography. In: Smuts B.B.,

Cheney D.L., Seyfarth R.M., Wrangham R.W. &

Struhsaker T.T. (Eds.), Primate societies. University

of Chicago Press, Chicago, pp. 54-68.

Dasilva G.L., 1992. The western black-and-white Colo-

bus as a low-energy strategist: activity budgets,

energy expenditure and energy intake. Journal of Ani-

mal Ecology, 61: 79-9.

Di Fiore A. & Campbell C.J., 2007. The Atelines: varia-

tion in ecology, behavior, and social organization. In:

Campbell C.J., Fuentes A., MacKinnon K.C., Panger

M. & Bearder S. K. (Eds.), Primates in perspective.

Oxford University Press, Oxford, pp. 155-185.

Franceschi V.R. & Homer H.T., 1980. Calcium oxalate

crystals in plants. Botanical Review, 46: 361-427.

Freeland W.J. & Janzen D.H., 1974. Strategies in her-

bivory by mammals: the role of plant secondary com-

pounds. American Naturalist, 108: 269-289.

Genua J.M. & Hillson C.J., 1985. The occurrence, type

and location of Calcium oxalate crystals in the leaves

of fourteen species of Araceae. Annals of Botany, 56:

351-361.

GlanderK.E., 1994. Nonhuman primate self-medication

with wild plant foods. In: Etkin N.L. (Ed.), Eating on

the wild side: The pharmacologic, ecologic, and so-

cial implications of using noncultigens. University of

Arizona Press, Tuscon, pp. 239-256.

Huffman M.A., 1997. Current evidence for self-medica-

tion in primates: a multidisciplinary perspective.

Yearbook of Physical Anthropology, 40: 171-200.

Jack K.M., 2007. The Cebines: toward an explanation of

variable social structure. In: Campbell C.J., Fuentes

A., MacKinnon K.C., Panger M. & Bearder S. K.

(Eds.), Primates in perspective. Oxford University

Press, Oxford, pp. 107-121.

Janson C.H., 1988. Intra- Specific Food Competition and

Primate Social Stmcture: A Synthesis. Behaviour,

105: 1-17.

Lewis T.R., Grant P.B.C., Quesada M.G., Ryall C. &

LaDuke T.C., 2010. A botanical survey of Cano

Palma Biological Station (Estacion Biologica Cano

Palma), Tortuguero, Costa Rica. Brenesia, 73-74:

73-84.

Martin P. & Bateson P., 1993. Measuring behaviour: An

introductory guide, second edition. Cambridge Uni-

versity Press, Cambridge, 176 pp.

Massey L.K., Roman-Smith H. & Sutton R.A.L., 1993.

Effect of dietary oxalate and calcium on urinary ox-

alate and risk of formation of calcium oxalate kidney

stones. Journal of the American Dietetic Association,

93: 901-906.

Milton K., 1979. Factors influencing leaf choice by

howler monkeys: a test of some hypotheses of food

selection by generalist herbivores. American Natura-

list, 114:362-378.

Myers N., Mittermeier R.A., Mittermeier C.G., da Fon-

seca G.A.B. & Kent J., 2000. Biodiversity hotspots

for conservation priorities. Nature, 40: 853-858.

Pavelka M.S.M. & Knopff K.H., 2004. Diet and activity

in black howler monkeys ( Alouatta pigm ) in southern

Belize: does degree of fmgivory influence activity

level? Primates, 45: 105-111.

Robinson J.G. & Janson C.H., 1987. Capuchins, squirrel

monkeys, and Atelines: socio-ecological convergence

with Old World primates. In: Smuts B.B., Cheney

D.L., Seyfarth R.M., Wrangham R.W. & Struhsaker

T.T., (Eds.). Primate societies. University of Chicago

Press, Chicago, pp. 69-82.

Rosenberger A.L., 1992. Evolution of feeding niches in

new world monkeys. American Journal of Physical

Anthropology, 88: 525-562.

Sanchez-Azofeifa G.A., Harriss R.C. & Skole D.L.,

200 1 . Deforestation in Costa Rica: a quantitative ana-

lysis using remote sensing imagery. Biotropica, 33:

378-384.

Schoener T.W., 1971. Theory of feeding strategies. An-

nual Review of Ecology and Systematics, 2: 369-404.

Biodiversity Journal, 2013, 4 (2): 335-342

Diversity of mosquitoes (Diptera Culicidae) in protected na-

tural parks from Valencian Autonomous Region (Eastern

Spain)

Alberto Bernues-Baneres* & Ricardo Jimenez-Peydro

Entomology and Pest Control Laboratory, Cavanilles Institute for Biodiversity and Evolutionary Biology, University of Valencia,

Official Post 22085, 46071 Valencia, Spain

* Corresponding author, email: alberto.bernues@uv.es

ABSTRACT Several larval samplings of mosquitoes (Diptera Culicidae) were carried out between 2008-

2011, throughout very diverse larval biotopes located in thirteen protected natural parks from

Eastern Spain, offering new information about the faunistic diversity of mosquitoes in these

protected areas. Biodiversity was analyzed in terms of alpha, beta and gamma components,

with the aim of comparing mosquito diversity according to the typology of the natural parks

under study. A total of 15355 specimens belonging to 25 different mosquito species and 6

genera were collected and identified. Diversity analysis indicated higher diversity for Inland

Mountainous Areas (IMAs) with a low degree of interspecific dominance in these communi-

ties, while Coastal Wetlands and Marshes registered the lowest observed diversity and a high

degree of interespecific dominance. The cluster analysis revealed the relationship between

the categories (IMA, CMA, CWM), while the Principal Components Analysis proved the re-

lationship between larval abundance and the categories studied.

KEY WORDS Mosquitoes; Culicidae; biodiversity; natural parks; Eastern Spain.

Received 29.04.2013; accepted 29.05.2013; printed 30.06.2013

INTRODUCTION

Mosquitoes (Diptera Culicidae) are considered

one of the most relevant group of arthropods in the

public health field (Schaffner et al., 2001; Becker

et al., 2010) and, like other organisms, show a di-

rect relation to different factors such as environ-

mental and habitat heterogeneity or host

preferences (Zhong et al., 2003). Unfortunately,

since the eradication of malaria in Spain (Bueno

Mari & Jimenez Peydro, 2008), there have been

few scientific studies aimed to increase the knowl-

edge about mosquito diversity and the factors that

regulate its change and their populations in our

country. From this point of view, the comparison

of mosquito diversity (alpha diversity) and the

structure of the communities in wich they are inte-

grated (beta diversity) can provide us with a pow-

erful tool for the implementation of more effective

and efficient population control programs, accord-

ing to the structure of the landscape (Wittaker,

1972; Magurran, 1988).

We can define alpha diversity (a) as the spe-

cific richness of a community that we consider ho-

mogeneous. Beta diversity (ft) refers to the

replacement degree in the specific composition

between different communities of a landscape.

And, finally, we can define gamma diversity (y) as

the specific richness of the grouped communities

that form a landscape, resulting from both alpha

and beta diversities interaction (Magurran, 1988).

This method of biodiversity analysis is useful not

336

Alberto Bernues-Baneres & Ricardo Jimenez-Peydro

only to explore the climatic, physical or biological

influences on biodiversity, but also to study the ef-

fects of human pressure on biodiversity (Halffter,

1998; Moreno, 2001). Taking into account these

considerations, the aim of this study was to ana-

lyze the diversity of Culicidae present in the natu-

ral areas considered, as well as the differences on

the faunistic composition of mosquito species in

function of the climatic and ecological features of

each natural park.

MATERIALS AND METHODS

Study area

To develop the study, we selected thirteen natu-

ral parks belonging to the Valencian Autonomous

Region (Spain): eight of them belonging to inland

mountainous areas, two belonging to coastal moun-

tainous areas and, finally, three belonging to coastal

wetlands and marshes (Fig. 1). Due to the climatic

variability recorded in the Valencian Autonomous

Region, it is possible to observe large differences

in the average temperatures and precipitacions reg-

istered between the different categories of the natu-

ral parks here studied. According to this, we can

define our study area as follows (GVA, 2003):

• Inland Mountainous Areas (IMA). Character-

ized by a tipical Mediterranean climate, but influ-

enced by continental climate, these are the only areas

where it is possible to find Supramediterranean

(mean temperature range between 13-8 °C) and Oro-

mediterranian (mean temperature range between 8-

4 °C) termotypes. During the study period, maximum

average temperatures of 26.5 °C and minimum aver-

age temperatures of 2.5 °C were recorded, with an

avergage precipitation of 36.7 1 (Fig. 2).

• Coastal Mountainous Areas (CMA). Very

similar to the IMA, are characterized by being

classified as Termomediterranean (mean tempera-

ture range between 19-17 °C) and Lower Me-

somediterranean (mean temperature range between

17-13 °C) termotypes, which is why the average

temperature is higher than in the previous class.

During the study period, maximum average tem-

peratures of 27.8 °C and minimum average tem-

peratures of 5.0 °C were recorded, with an avergage

precipitation of 29.2 1 (Fig. 3).

• Coastal Wetlands and Marshes (CWM). CWM

are the most common coastal environments of the

Valencian territory, registering the highest average

values of temperature (Inframediterranean termo-

type, mean temperature >19 °C). Rainfall is

strongly influenced by seasonality, characterized by

a severe drought time during the summer months.

During the study period, maximum average tem-

peratures of 29.1 °C and minimum average tem-

peratures of 5.5 °C were recorded, with an average

rainfall of 27.8 1 (Fig. 4).

Figure 1. Study area and the classification of each natural park studied by category. IMA (Inland Mountainous Areas),

CMA (Coastal Mountainous Areas) and CWM (Coastal Wetlands and Marshes).

Diversity of mosquitoes (Diptera Culicidae) in protected natural parks from Valencian Autonomous Region (Eastern Spain) 337

Sampling methods and taxonomic identification

A simple random sampling method was car-

ried out across the study area by selecting all suit-

able biotopes to accomodate immature forms of

mosquitoes. In this way, many different points

were sampled by using “dipping” technique (Ser-

vice, 1993) over 4 consecutive years (2008-2011).

Mosquito species were identified according to the

keys of Encinas Grandes (1982), Darsie & Sami-

nadou Voyadjoglou (1997) and Schaffner et al.

( 2001 ).

Diversity studies and statistical analysis

Diversity studies (alpha diversity) were con-

ducted separately for each natural park category

(IMA, CMA and CWM) by calculating classic di-

versity indexes like Margalef’s (D Mg = (S- 1 )/ln7V)

(Simpson, 1949; Magurran, 1988; Moreno, 2001)

and Simpson’s Indexes (2 = Zpf , where = nfN

[uf, relative abundance of the species calculated as

the proportion of individuals of a given species

against the total number of individuals of a commu-

nity, N]). Shannon diversity idex (H’ = -\L(pj •ln/; / -)])

is commonly used to characterize species diversity

in a community, accounting for both abundance and

evenness of the species present (Shannon & Wea-

ver, 1949). Species richness (5) is the number of

species present in a community while species even-

ness («/’) indicates the distribution of individuals

within the species and it’s calculated by using Pie-

lou’s Index formula ( J ’ = //77/ max , where H max =

In (5)) (Magurran, 1988; Moreno, 2001).

On the other hand, to calculate beta diversity, a

variety of similarity/dissimilarity indexes were used,

both qualitative (Jaccard’s Index, L = c/[ a+b-c]) and

quantitative (Sorensen’s Index, I s = [2 pN]/[aN+bN]),

as well as Whittaker’s (wich calculates the

species replacement according to the expression (i w

= <Sy[(2<2+Z?+c)-l]) and Complementarity Idex (C AB

= [S a + Sb-2VabV[Sa+S b -V ab ], where V AB repre-

sents the number of common species to both sites

A and B) (Magurran, 1988; Moreno, 2001).

The calculation of gamma diversity, was carried

out by using the classic proposal of Schluter and

Ricklefs (Schluter & Ricklefs, 1993) ([average a

diversity] [average (3 diversity] [sample size(7V’)]),

as well as the modification made by Lande

(1996) ([average a diversity] [/? diversity], where

Figure 2. Characteristic climogram (average maximum

and minimum temperatures and precipitations) of IMAs

natural parks for the study period (2008-2011). Figure 3.

Characteristic climogram (average maximum and mini-

mum temperatures and precipitations) of CMAs natural

parks for the study period (2008-2011). Figure 4. Charac-

teristic climogram (average maximum and minimum tem-

peratures and precipitations) of CWMs natural parks for

the study period (2008-2011).

f) = Zjqj[S r Sj |) to calculate the contribution made

by alpha and beta diversity to gamma (Moreno,

2001). Finally, to calculate the ecological distance

between different environments, a cluster (based on

338

Alberto Bernues-Baneres & Ricardo Jimenez-Peydro

IMA

Relative

abundance

CMA

Relative

abundance

CWM

Relative

abundance

Aedes

Aedes vexans

0.16

0.00

Aedes vittatus

0.00

1.37

0.00

Anopheles

Anopheles atroparvus

0.23

0.00

Anopheles claviger

0.32

0.00

Anopheles maculipennis s.s.

0.36

0.00

Anopheles marteri

0.20

0.00

Anopheles petragnani

1069

11.23

1.77

0.00

Culex

Culex hortensis hortensis

1216

12.77

0.20

0.00

Culex impudicus

697

7.32

1.23

0.00

Culex laticinctus

1265

13.29

1462

40.96

0.00

Culex mimeticus

583

6.12

1.15

0.00

Culex modestus

0.18

0.00

2.08

Culex pipiens

1935

20.32

708

19.84

1138

50.24

Culex territans

246

2.58

0.00

Culex theileri

0.02

0.00

Culiseta

Culiseta annulata

0.65

0.00

0.49

Culiseta longiareolata

2151

22.59

1195

33.48

168

7.42

Ochlerotatus

Ochlerotatus berlandi

0.15

0.00

Ochlerotatus caspius

0.00

658

29.05

Ochlerotatus detritus

0.00

223

9.85

Ochlerotatus echinus

0.98

0.00

Ochlerotatus geniculatus

0.35

0.00

Ochlerotatus gilcolladoi

0.12

0.00

Ochlerotatus pulcritarsis

0.07

0.00

Uranotaenia

Uranotaenia unguiculata

0.00

0.88

TOTAL COUNT

9521

62.00%

3569

23.24%

2265

14.75%

Table 1. Number of specimens captured for each environmental category (IMA, CMA and CWM).

Diversity of mosquitoes (Diptera Culicidae) in protected natural parks from Valencian Autonomous Region (Eastern Spain) 339

Jaccard’s Index) and a principal components analy-

ses (PCA) were made, offering the cophenetic cor-

relation value for the Jaccard cluster to calculate the

degree of reliability of the classification system

used. PAST software (Paleontological Statistics

Software Package) was used to carry out all calcu-

lations developed (Hammer et al., 2001).

RESULTS AND DISCUSSION

Faunistic and systematic results

A total of 15,355 mosquito larvae were collected

from 285 sampling points, obtaining a total of 900

samples. The systematic study showed a total of 25

species belonging to 6 different genera (Table 1)

wich represents an 86.21% of the maximum spe-

cific richness calculated for the Valencian Autono-

mous Region (Bueno Mari, 2011). The complete

catalogue of species collected is listed below: Aedes

vexans (Meigen, 1830); Aedes vittatus (Bigot,

1861); Anopheles atroparvus Van Thiel, 1927; Ano-

IMA

CMA

CWM

Abundance

9521

3569

2265

Specific richness ( S)

Margalef index (Dy[g)

2.183

0.856

0.777

Simpson index (2)

0.149

0.320

0.353

Shannon index ( H 3

2.101

1.301

1.274

Evenness of Pielou index (J r )

0.690

0.626

0.655

Table 2. Alpha biodiversity estimators for each environ-

mental category (IMA, CMA and CWM).

1 MA-

GMA

IMA-

CWM

CMA-

CWM

Jaccard index (Ip

0.32

0.17

0.15

Sorensen index C ^Squant >

0.51

0.23

0.30

Whittaker index (fi ^ )

0.52

0.71

0.73

Complementarity (C^g%)

68.18

83.33

84.62

Table 3. Beta biodiversity estimators for each environmen-

tal category (IMA, CMA and CWM)

pheles claviger (Meigen, 1804); Anopheles maculi-

pennis s.s. Meigen, 1818; Anopheles marteri Sene-

vet et Pmnelle, 1927; Anopheles petragnani De

Vecchio, 1939; Culex hortensis hortensis Ficalbi,

1889; Culex impudicus Ficalbi, 1890; Culex laticin-

ctus Edwards, 1913; Culex mimeticus Noe, 1899;

Culex modestus Ficalbi, 1889; Culex pipiens Lin-

naeus, 1758; Culex territans Walker, 1856; Culex

theileri Theobald, 1903; Culis eta annul ata ( Schrank,

1776); Culiseta longiareolata (Macquart, 1838);

Ochlerotatus berlandi (Seguy, 1921); Ochlerotatus

caspius (Pallas, 1771); Ochlerotatus detritus (Hali-

day, 1833); Ochlerotatus echinus (Edwards, 1830);

Ochlerotatus geniculatus (Olivier, 1791); Ochlero-

tatus gilcolladoi (Sanchez-Covisa, Rodriguez et

Guillen, 1985); Ochlerotatus pulcritarsis (Rondani,

1872) and Uranotaenia unguiculata Edwards, 1913.

Mosquito species richness and evenness

According to the analysis of a biodiversity in-

dexes (Table 2), it is possible to observe that IMA en-

vironments are the most diverse (N =21; D Mg = 2.1 83),

Figure 5. Cluster analysis based on Jaccard’s distance; co-

phenetic correlation rc=0.9975.

340

Alberto Bernues-Baneres & Ricardo Jimenez-Peydro

while CWM are the least diverse (S= 7; D Mg = 0.777).

Simpson and Shannon indexes highlight that in

CWMs (2 = 0.353; H’ = 1.274) species such as Cx.

pipiens (50.24%) and O. caspius (29.05%) strongly

dominate the rest of species present in the community.

Something similar occurs in the case of CMAs (2 =

0.320; H’ = 1.301), where Cx. laticinctus (40.96%),

Cs. longiareolata (33.48%) and Cx. pipiens (19.84%)

develop a strong influence. Finally, IMAs (2 = 0.149;

// ’ = 2.101) are the category of natural park where a

greater evenness degree can be ob served, because the

most dominant species do not show such a strong in-

fluence as in the two other cases.

These observations can be explained according

to the bioclimatic characteristics of each natural

park category. The IMAs record a greater rainfall

abundance (Fig. 2) as well as a higher variety of en-

vironments that are able to be colonized by mosqui-

toes than in other categories of natural parks. That

means a greater amount of larval biotopes available

to be exploited by different communities of culicids

along the year. The CMAs, can be defined as tran-

sitional environments between IMAs and CWMs

categories since rainfall regime is more heteroge-

neous, focusing on specific periods throughout the

year (Fig. 3). This factor, in combination with times

of severe drought during the summer months, de-

termines a population dynamics feature which is re-

flected in the diversity observed in the natural parks

grouped in this category. Finally, CWMs represent

the most extreme type of environment analyzed,

being the driest (Fig. 4) and most homogeneous in

terms of water bodies typology, which acts as limit-

ing factor in the diversity observed in that category.

Similarity and dissimilarity analysis

The analysis of fi biodiversity (Table 3), indicates

that IMAs and CMAs are the closest categories in

their specific composition (7 ; - = 0.32; I Squant = 0.5 1),

showing the lower replacement degree (fiw = 0.52)

between pairs analyzed, observations also supported

by the complementarity index (C IMA -C MA =

68.18%). With the aim of representing the informa-

tion provided by the Jaccard index, a cluster analysis

based on Jaccard’s distance was carried out, cor-

roborating the same conclusions already given be-

fore (Fig. 5). The high value of Jaccard distance

cophenetic correlation (r c = 0.9975) indicates a high

correlation level between the ecological distance ob-

served in the study and the distance predicted by the

hierarchical configuration of the cluster.

As a result of the PCA, two principal compo-

nents were extracted explaning a 93.06% of the

total variability observed (PCj = 83.55%; PC 2 =

9.51%) (Fig. 6). It is worth pointing out the large

dispersion exhibited by Cx. pipiens, Cs. longiareo-

lata and Cx. laticinctus, which are the most abun-

dant and dominant species in all studied

communities. It is also interesting to mention the

close relationship existing between IMAs (the most

diverse and heterogeneous) and species such as Cx.

mimeticus, Cx. impudicus and An. petragnani, wich

define perfectly the main vector of this natural park

category. Specially significant is the case of Cx.

hortensis hortensis, which has been described as

one of the regular members of the most biodiverse

Culicidae communities in our region (Bueno Mari

& Jimenez Peydro, 2011).

In the case of CWMs, note the relationship with

typical halophilic species, being O. caspius and O.

detritus the ones that best define the axis of this cat-

egory, despite the fact that Cx. pipiens was the most

common and dominant species in CWMs.

Integrated landscape biodiversity

As mentioned above, gamma (y) diversity was

calculated following Schluter & Ricklefs (1993)

and Lande (1996) criteria. As a result, 7(s c hluter and

Ricklefs) was 25.7 1 , virtually identical to the value of

the total specific richness (S = 25) evidenced in the

study area. On the other hand, y (Lande) was 19.99

with an a contribution of 60.05% and a /> contribu-

tion of 39.95%, demonstrating that the alpha diver-

sity of the richest community (IMAs) contributes to

a greater extent of the gamma diversity (60.05 %),

which implies a low level of complementarity be-

tween categories and a high proportion of exclusive

species in each category studied.

CONCLUSIONS

Due to the limited existence of studies based on

mosquito diversity in Spain (Demba et al., 2005;

Bueno Mari, 2011; Bueno Mari & Jimenez Peydro,

2011), the results of our study represent an inter-

esting contribution to the general knowkedge about

Culicidae diversity in our country. According to

Diversity of mosquitoes (Diptera Culicidae) in protected natural parks from Valencian Autonomous Region (Eastern Spain) 341

1 . 8 -

l.J-

1 , 2 -

0,9-

0 . 6 -

0.3-

-0.3-

-0.6-

-0.9-

Ae. vittatus

CMA

Cx.laticinctus

Cs. longiareolata

IMA

Cx.mimelicus/territans

Cx, pipiens

An.petragnani 1

*Cs. annulata/subochrea

Cx. hortensis hortensis’

Cx. impudicus

An. maculipennis s.s.

CWM

Cx. modestus

-i-

-0.4

U. unguiculata

O. detritus

O, caspius

- 0.8

0.4

1.2

1.6

Axis 1

2.4

Figure 6. Principal Components Analysis (PCA) based on the larval abundance within each natural park category.

Margalef Index and specific richness, IMAs show

the highest diversity observed in the Valencian Au-

tonomous Region’s natural parks, probably due to

various factors that should be taken into account. In

first place, these are the most extensive areas, fa-

cilitating landscape heterogeneity and larval

biotopes diversity present within their limits. These

conditions favor their colonization and increase the

likelihood of the presence of suitable host on which

to feed. Another aspect to consider is the water qual-

ity sampled (low levels of eutrophy), the high level

of replacement and longer periods of stay (higher

precipitation regimes favor both aspects) (Rivas

Martinez, 2004), aspects strongly related with the

presence of species such as Cx. hortensis hortensis

and An. petragnani. CMAs recorded a lower level

of diversity, a fact that relates to their lower surface

and larger coastline influence. These conditions lead

to a decrease in turnover rate and retention of water

(high droguht times) (Rivas Martinez, 2004), favor-

ing an ecological filter toward less sensitive species

and better adapted to strong changes such as Cs. lon-

giareolata and Cx. laticinctus (Becker et al., 2010).

CWMs are the natural parks that suffer from the

stronger influence by the closeness of the sea, show-

ing a greater homogeneity of larval habitats, a lower

level of replacement and water permanence (Rivas

Martinez, 2004), promoting the presence of heavily

adapted species (such as O. caspius , O. detritus and

U. unguiculata ) or highly plastic ones (such as Cx.

pipiens). On the other hand, the concentration of

human population in coastal zones (INE, 2011) and

the higher level of anthropogenic influence has

been linked to the decline of diversity in other areas

of our territory (Bueno Mari et al., 2010). In this

case, it has been observed a pattern of loss of diver-

sity from inland areas (IMAs) toward coastal ones

(CWMs), coinciding with the assessments of Bueno

Mari & Jimenez Peydro (2011), who mantain that

a high level of anthropization does not imply a re-

duction of mosquito populations in urban environ-

ment, but rather an ecological selection filter that is

only surpassed by a few species.

Finally, it is particularly interesting to note the

high level of correlation observed between IMAS

and a small group of species (An. petragnani, Cx.

hortensis hortensis, Cx. impudicus and Cx. mimeti-

cus ), which behave as indicators of this category and,

therefore, can be related to a high degree of conser-

vation of the environment for future studies on

bioindicator species (Dorville, 1996, Montes, 2005).

To conclude, IMAs have a higher diversity of

Culicidae with a lesser degree of dominance and a

greater intraspecific evenness. On the other hand,

CWMs represent the less diverse and uniform com-

munities, with a greater degree of interspecific

dominance. These communities are composed of a

few abundant and a high number of rare species,

342

Alberto Bernues-Baneres & Ricardo Jimenez-Peydro

establishing a clear relationship between Culicidae

abundance and the prevaling climatic conditions.

ACKNOWLEDGEMENTS

We are grateful to the Regional Ministry of Ge-

neralitat Valenciana (Conselleria de Medi Ambient,

Aigua, Urbanisme i Habitatge) for the grant of the

insects capture permissions in the protected enclaves.

We also want to point out that this article was par-

tially funded by Research Project CGL 2009-11364

(BOS), supported by the Ministry of Science and In-

novation of Spain (Ministerio de Ciencia e Innova-

cion del Gobierno de Espana).

REFERENCES

Becker N., Petrie D., Zgomba M., Boase C., Madon M.,

Dahl C. & Kaiser, A., 2010. Mosquitoes and their

control. Springer, London, 577 pp.

Bueno Mari R., 2011. Bioecologia, diversidad e interes

epidemiologico de los culicidos mediterraneos (Dip-

tera: Culicidae). Servei de Publicacions de la Univer-

sitat de Valencia, Valencia, 423 pp. Available online

at: http://hdl.handle.net/10803/52158.

Bueno Mari R. & Jimenez Peydro R., 2008 Malaria en

Espana: Aspectos entomologicos y perspectivas de fu-

turo. Revista Espanola de Salud Publica, 82: 161-163.

Bueno Mari R. & Jimenez Peydro R., 2011. Diferences

in mosquito (Diptera: Culicidae) biodiversity across

varying climates and land-use categories in Eastern

Spain. Entomologica Fennica, 22: 190-198.

Bueno Mari R., Corella Lopez E. & Jimenez Peydro R.,

2010. Culicidofauna (Diptera: Culicidae) presente en

los distintos enclaves hidricos de la ciudad de Valen-

cia (Espana). Revista Colombiana de Entomologia,

36: 235-241.

Darsie R.F. & Saminadou Voyadjoglou A., 1997. Keys

for the identification of the mosquitoes of Greece.

Journal of the American Mosquito Control Associa-

tion, 13: 247-254.

Demba Sy M., Remus Zamfirescu S., Thiam N. & Palanca

Soler A., 2005. Diversity of mosquitoes in a semiarid

environment from San Juan del Flumen (Los Mone-

gros, Huesca, Spain) (Diptera, Culicidae). Boletin de

la Asociacion Espanola de Entomologia, 29: 23-33.

Dorville L.F.M., 1996. Mosquitoes as bioindicators of

forest degradation in southeastern Brazil, a statistical

evaluation of published data in the literature. Studies

on Neotropical Fauna and Environment, 31: 68-78.

Generalitat Valenciana (GVA), 2003. Habitats prioritarios

de la Comunidad Valenciana. Conselleria de Territori

i Habitatge, Valencia, 222 pp.

Halffter G., 1998. A strategy for measuring landscape

biodiversity. Biology International, 36: 3-17.

Hammer 0., Harper D.A.T. & Ryan P.D., 2001. PAST:

Paleontological statistics software package for ed-

ucation and data analysis. Palaeontologia Electronica,

4: 9. Available online in: http://palaeo-electronica.

org/ 2001_l/past/ issuel_01.htm.

Instituto Nacional de Estadistica (INE), 2011. Cifras ofi-

ciales de poblacion resultantes de la revision del pa-

dron municipal (01/01/2011). Available online at:

http:// www.ine.es/iaxi/tabla.do. Accessed 3 March

2013.

Lande R., 1996. Statistics and partitioning of species di-

versity and similarity among multiple communities.

Oikos 76: 5-13.

Magurran A.E., 1988. Ecological diversity and its mea-

surement. Princeton University Press, New Jersey,

179 pp.

Montes J., 2005. Culicidae fauna of Serra da Cantareira,

Sao Paulo, Brazil. Revista de Saude Publica Sao

Paulo, 29: 578-584.

Moreno C., 2001. Metodos para medir la biodiversidad.

Manuales & Tesis de la SEA. Editorial Cyted, Orcyt-

Unesco & SEA, Zaragoza, 84 pp.

Rivas Martinez S., 2004. Centro de Investigaciones Fi-

tosociologicas. Global Bioclimatics Org. Available

online at: http://www.ucm.es/info/cif. Accessed 3

March 2013.

Schaffner F., Angel G., Geoffroy B., Hervy J.O. & Rha-

eim A., 2001. The mosquitoes of Europe/Les mousti-

ques d’ Europe [computer program], France: IRD

Editions and EID Mediterranee, Montpellier.

Schluter R.E. & Ricklefs R.E., 1994. Species diversity: an

introduction to the problem, in: Ricklefs, R.E. &

Schluter, R.E. (Eds.), Species diversity in ecological

communities: historical and geographical perspectives.

The University of Chicago Press, Chicago, pp. 1-10.

Service M.W., 1993. Mosquito Ecology. Field Sam-

pling Methods. Elsevier Science Publishers, Lon-

don, 988 pp.

Shannon C.E. & Weaver W., 1949. The Mathematical

Theory of Communication. The University of Illinois

Press, Urbana, 111, 144 pp.

Simpson E.H., 1949. Measurement of diversity. Nature,

163:688-688.

Wittaker R.H., 1972. Evolution and measurement of

species diversity. Taxon, 21: 213-251.

Zhong H., Yan Z., Jones F. & Brock C,, 2003. Ecological

analysis of mosquito light trap collections from West

Central Florida. Environmental Entomolgy, 32: 807-

815.

Biodiversity Journal, 2013, 4 (2): 343-354

Phenotypic diversity, taxonomic remarks and updated distri-

bution of the Mediterranean Jujubinus baudoni (Monterosato,

1891) (GastropodaTrochidae)

Paolo Mariottini 1 *, Andrea Di Giulio 1 , Massimo Appolloni 2 & Carlo Smriglio 1

‘D ipartim ento di Scienze, Universita degli Studi di Roma Tre, Viale Marconi 446, 00 146 Rome, Italy; e-mails:

paolo.mariottini@ unir oma3.it; andrea.digiulio@ unir oma3.it; csmriglio@ alice.it

; Museo Civico di Zoologia, Sezione Molluschi, Via Ulisse Aldrovandi, 1 8, 00 197 Rome, Italy; e-mail:

m assim o.appolloni@ com une.ro m a. it

* C orresponding author

ABSTRACT Jujubinus baudoni (M onterosato, 1891) shows highly diverse chromatic and morphological

patterns. Based on the examination of the type material and series of specimens from private

collections, and of recent findings from Sardinia, we reviewed and updated the distribution

of this species, and figured the three representative shell colour morphs which are constant

in local populations. A comparative SEM analysis did not outline significant differences

among shells with greatly different colour patterns, thus justifying their belonging to a single

specif ic entity. The survey of the material of J. baudoni did not show a strict correspondence

between Corse and Sardinian shell colour morphs and their geographical distribution, not al-

lowing to clearly state trends and relationships among island populations. On the contrary

the colour pattern of the continental shells of J. baudoni seems to be constant and different

from the ones of the island shells, indicating a well separated colour morph. This study con-

firmed that the distribution of J. baudoni is limited to the north-western Mediterranean Sea,

with its eastern range widened to the La Maddalena Archipelago (Sardinia). The record of

this species from Azores is due to a misidentification with J. pS6lldogravina6 Nordsieck, 1973,

which is also figured for comparison. A lectotype and p arale c to ty p e s of TvOChuS baudoni are

herein designated .

KEY WORDS Trochidae; JujllbinUS baudoni', Monterosato collection; colour morph; Mediterranean Sea.

Received 1 6.05.20 1 3; accepted 20.06.20 1 3; printed 30.06.20 1 3

INTRODUCTION

The genus Jujubinus Monterosato, 1 8 84 was

nominally based on TvochllS YUCltOYli Montagu, 1803

and it is represented by a small group of marine gas-

tropods living mostly in the interdital zone down to

about 80 m, invariably associated with photophilic

algal vegetation and/or marine p h an ero g am e s . The

alp h a -tax o n o m y of this genus still has to be com-

pletely solved, but it is currently accepted that sev-

eral different species occur worldwide albeit,

mainly on European waters, the exact number at the

moment amounts to 29 according to WoRMS

(World Register of Marine Species available at

http ://w w w ,m arinespecies.org/index .php ; A ppeltans

et al., 2012). In particular, 18 taxa are listed by

CLEMAM (at http:// w ww.somali. asso.fr/clem am /

index. php; Gofas & Le Renard, 2013) as European

species, most of them restricted to the Mediter-

ranean basin. Among the Mediterranean taxa be-

344

P. Mariottini, A. Di Giulio, M.Appolloni & C. Smriglio

longing to this genus, JujubinUS baudoni (Monte-

rosato, 1891) was revaluated as bona species by Cu-

rini Galletti (1982). J. baudOYli was described under

the genus TvochuS Linnaeus, 1758 by Monterosato

(1891) with a very scanty description: “II T. Bau-

doni, H. Martin mss., e una piccola forma grosse-

sculpta, vivente in abbondanza sulle coste di

Provenza” (“The T. Baudoni, H. Martin mss., is a

small form heavily sculptured, abundant and living

along the coast of Provence”). Since the paper of

Curini Galletti (1982), only two further reports we re

recently published dealing with this species: by Sca-

pe rrotta et al. (2011), who figured shells from C orse,

and by Spanu (2011) who reported specimens from

the northwestern Sardinian coast. J. bcilidofli is

present on the infralittoral zone and is considered

locally common of the Catalan coast, Gulf of Lion,

Corse and North-Western Sardinia (Curini Galletti,

1 982; Scaperrotta et al., 2011; Spanu, 2011). The

Azores Islands cannot be considered in the J. bau-

doni distribution, since the materialexamined in the

Monterosato collection and labelled as CallioStoma

baudoni coming from these islands, tentatively iden-

tified as J. pseudogravinae N ordsieck , 1973 by Cu-

rini Galletti (1982), has been confirmed to belong to

this latter taxon. We have analyzed the type material

of Trochus baudoni m on ter os a to, 1891 and 7. bau-

doni var. incomparabilis Locard et Caziot (1901)

(reported by clem am as J. baudoni incompara-

bilis Ghisotti et Melone, 1975), and series of speci-

mens of J. baudoni from private collections. A

lectotype of J. baudoni is herein designated. We

found that this species shows a great mo rphological

variability, mainly concerning its colour pattern,

which is very constant within the different popula-

tion examined.A survey of colour mo rphs from dif-

ferent localities is here presented and, as far as to

our knowledge, this study confirmed that the distri-

bution of J. baudoni is lim ited to a restricted area of

the North-Western Mediterranean basin, being

slightly enlarged to east with the sampling of shells

from the La Maddalena Archipelago (Sardinia).

Shells from M o ntero sato ’s collection labelled as J.

baudoni and J. pseudogravinae fr o m the Azores are

figured for comparison.

ACRONYMS AND ABBREVIATIONS. Carlo

Smriglio and Paolo Mariottini collections, Rome,

Italy (CS-PM); Monterosato (MTS); Museo Civico

di Zoologia, Rome, Italy (MCZR); Scanning Elec-

tron Microscopy (SEM).

MATERIALS AND METHODS

Examined Material. We could examine the

type material of TvochuS baudoni in the Montero-

sato collection at the MCZR, consisting of empty

shells: 134 exx from Paulilles (France); 30 exx from

Roussillon (France); 12 exx from Departement de

l’Aude, Pyrenees Orientales (France); 20 exx J.

baudoni var. incomparabilis from Pietranera,

(Corse). A lectotype from the Paulille lot has been

designated with the number MCZR00218 and the

remaining material have been designed as paralec-

totypes M C Z R 0 0 2 1 9 /I - 1 9 5 . Furthermore, 18 exx

from Azores (Portugal) MCZR 11761, named as

Calliostoma baudoni but not related to this species,

have been examined.

Other empty shells of J. baudoni examined: 2

exx from Cadaques (Spain), collected at a depth of

4 m; 3 exx from Capo Caccia, Sardinia (Italy),

depth of 25 m. From La Maddalena Archipelago,

Sardinia (Italy) further empty materialwas gathered

among the shell grit collected handily by SCUBA

diving: 59 exx from Caprera Is., depth of 25 m; 91

exx from La Maddalena Is., depth of 25 m; 62 exx

of Spargi Is., depth of 20 m. M aterial referring to J.

pseudogravinae : 8 exx from Azores (Portugal), dea-

ler source. SEM photographs were carried out at the

Interdepartmental Laboratory of Electron Mi-

croscopy (LIME), Universita Roma Tre, Rome, Italy.

RESULTS AND DISCUSSION

We analysed the type m aterial of T. baudoni, in-

cluding the J. baudoni v ar. incomparabilis shells,

stored at the MCZR and we figured some shells

and original labels (Figs. 1 a - 1 2b). The shells of J.

baudoni V ar. incomparabilis w ell fit the description

given by Locard and Caziot (1901): “Nous revelon

une tres belle var. incomparabilis (M tr.), d’une

coloration verte et rouge”. A lectotype and paralec-

totypes of T. baudonihav e been herein designated.

From the examination of these shells, and the ac-

companying labels, as well as of shells from pri-

vate collections, we could confirm the systematic

status of this species and derived its distribution.

In fact, M onterosato’s material labelled as T. bau-

doni from Azores (Figs. 1 3 - 1 5 d ) resulted to be J.

pseudogravinae N ordsieck, 1 97 3 after a literature

research (Avila et al., 2004; Avila et al., 2007;

Phenotypic diversity, taxonomic remarks and updated distribution of the Mediterranean Jujubinus baudoni

345

Avila et al., 2011) and a direct comparison with

shells of this endemic species (Figs. 16a-d, 29a-n).

Most probably the bad conservation of the shells

(sent by Dautzenberg, see handwriting label ofFig.

14) has induced Monterosato to identify wrongly

these specimens, as already suggested by Curini

Galletti who examined this material (1982). As far

as we know, there are no reliable records of J. bau-

doni from A lb o ran Sea or Atlantic Ocean, in-

cluding Azores islands. On the contrary, from lit-

erature and personal sampling (Figs. 17a-24b) the

distribution of this species seems to be rather con-

fined to the north-western Mediterranean basin

(Ghisotti & Melone, 1 975; Curini Galletti, 1 982;

Scaperrotta et al., 2011; Spanu, 2011).

The distribution of J. baudoni has been slightly

enlarged to east with the shells collected by

SCUBA diving in the La Maddalena Archipelago

(Sardinia); in particular material was collected in

the infralittoral zone from three islands of the Ar-

chipelago (Caprera, La Maddalena, Spargi) (Figs.

2 2 a-2 4 b ; Table 1).

Regarding the morphological variability of this

taxon, interestingly we found different phenotypes,

being the shells (colour morph A) from the conti-

nental coast stouter, slightly more coarsely sculp-

tured, showing prosocline reddish-brown and

milky-white flam m ules, higher in number on the

basal cordlet (Figs. la-8c;17a-18c), in respect to the

specimens from Corse and Sardinia. In turn, these

latter can be divided in two different colour patterns

regarding the teleoconch coloration: 1) the typical

J. baudoni var. incomparabilis coloration (colour

morph B), which consists in an emerald green back-

ground interrupted by bright orange and/or red

flammules and milky-white prosocline stripes, with

white spots on the basal cordlet (Figs. 9a, 12b, 19a-

20b); 2) colour morph C, almost entirely brownish-

green or with prosocline faint whitish flammules

(Figs. 22a, 24b).

We recall that all J. baudoni shells invariantly

show protoconch and initial teleoconch whorls of

red colour. It is worth to mention that this feature

has induced several authors in the past to erro -

neousiy identify J. baudoni as J. exasperatus coralli-

HUS (Monterosato, 1884) (Figs. 21a-c-30a-e), as re-

called by Curini Galletti (1 982) and Spanu (2011).

Since it is possible to easily separate the three colour

patterns, as well as the continental shells from the

Corse and Sardinian ones, due to their slight dif-

ferences in the teleoconch sculpture, a comparative

SEM analysis was carried out. The examination of

the SEM photographs did not reveal any significant

protoconch and teleoconch sculpture differences

among all the shells analysed (Figs. 25a-28g), in

spite of their diverse colour patterns, thus indicating

that we are probably facing a single specific entity.

It is a matter of fact, the colour pattern of the conti-

nental shells of J. baudoni seems to be constant and

different from the ones of the island shells, indicating

a well separated phenotype. On the contrary, shells

of J. baudoni from C orse and S ardinia did not show

a strict correspondence between colour morphs and

their geographical distribution, not allowing to

clearly state trends and relationships among island

populations. In many marine gastropods a direct ge-

netic control of shell coloration has been demon-

strated, and in several cases variation in shell colour

has been considered to be an adaptive value related

to environmental variables such as climate, diet,

habitat choice, insolation, salinity, visual predation

and wave exposure (Terreni, 1981; Byers, 1989; So-

kolova & Berger, 2000 and references therein; Miura

et al., 2007). Shell colour may have three functions:

communication, crypsis and thermoregulation, and

it has been suggested that correlation between indi-

vidual physiology and shell colour polym orphism is

a result of pleiotropic effects of genes responsible for

the shell colour or a linkage between them and genes

determining certain physiological features (Sokolova

& Berger, 2000 and references therein; Miura et al.,

2007). The factors that maintain the color polymor-

phism, and those that contribute to among-site varia-

tion in color frequencies, re m a in unknow n. A lthough

the color polym orphism may have an underlying ge-

netic component, the regional-scale variation in color

frequency observed in J. baudoni could be ecologi-

cally controlled.

Interestingly, another member of the genus Ju-

jubinus, J. geographicus Poppe, Tagaro et D ekker,

20 0 6 shows a parallel shell colour variability (Figs.

31a, 32), including one very similar to the morph

B (red-green pattern, Figs. 3 1 a-c ) and one entirely

red (Fig. 32). In particular, green is a rare colour

occurring in marine shell coloration and the occur-

rence of this colour in two different species so dis-

tantly geographically separated (Mediterranean and

Philippines) could be due to a common genetic

background the phenotype of which is selected by

similar ecological constrains.

346

P. Mariottini, A. Di Giulio, M.Appolloni & C. Smriglio

Figure 1 . Tvochus bciudoni. MTS collection, lectotype MCZR00218, H = 6.5 x D = 5.2 mm, Pa u lilies (France). Figures 2 -

4. MTS collection, original labels. Figure 5. T. bciudoili. MTS collection, 6.3 x 5.0 mm, Roussillons (France). Fig u res 6,7.

MTS collection, original labels. Figure 8. T. baudoni. MTS collection, 5.7 x 4.7 mm, Department de l’Aude (France).

Phenotypic diversity, taxonomic remarks and updated distribution of the Mediterranean Jujubinus baudoni

347

Figure 9. JlljllbinUS baudoni var. incomparabilis. MTS collection, H = 6.0 X D = 4.9 mm, Pietranera (Corse). Figure 10.

MTS collection, original label. Fig u re 11./. baudoni var. inCOfnpCirClbilis . MTS collection, 6.1 x 5.0 mm, Pietranera (Corse).

Figure 12. J. baudoni v ar. incomparcibilis. MTS collection, 5.4 x 4.8 mm, Pietranera (Corse). Figure 13. MTS collection,

original label. Figure 14. J. pS € U d()g m villClC. MTS collection, 2.8 x 2.6 mm, Azores Islands.

348

P. Mariottini, A. Di Giulio, M.Appolloni & C. Smriglio

Figure 15. JujubinUS pseudo gravinae. MTS collection, H = 2.7 X D = 2.6 mm, Azores Islands, MTS collection. Figure 16.

J. pseudogravinae. CS-PM collection, 6.9 X 4.8 mm, Cais da Ribeirinha, S. Miguel Is. (Azores Islands), on stones, 10 m

depth. Figure 17. JujubinUS baudoni. CS-PM collection, 5.5 x 4.4 mm, Cadaques (Spain), 4 m depth. Figure 18. J. Baudoili.

CS-PM collection, 3.9 x 3.1 mm, Cadaques (Spain), 5 m depth.

Phenotypic diversity, taxonomic remarks and updated distribution of the Mediterranean Jujubinus baudoni

349

Figure 19. JujubinUS baudoni w ar. incomp ar abilis . CS-PM collection, H = 5.9 mm X D = 4.6 mm, Capo Caccia (Sardinia),

15 m depth. Figure 20. idem, 4.1 mm x 3.7 mm . Figure 21.7. CXClSpCVCltUS COTallinUS . CS-PM collection, 6.2 x 4.9 mm. La

Maddalena Is. (Sardinia), 25 m depth. Figure 22. J. bcilldoni. CS-PM collection, 7.1 x 4.8 mm, La Maddalena Is., Sardinia,

25 m depth. Figure 23. idem, 7.1 x 4.9 mm, Caprera Is. (Sardinia), 30 m depth. Figure 24. idem, 6.0 x 4.2 mm.

350

P. Mariottini, A. Di Giulio, M.Appolloni & C. Smriglio

Figure 25. Tvochus baudoYli. MTS collection, H = 4.7 x D = 4.2 mm, Paulilles (France). Figure 26. JujubifflAS baudoni v ar.

incomparabilis. MTS collection, 6.1 X 4.8 mm, Pietranera (Corse). Figure. 27. J. bciudoili { Monterosato, 189 1); specimen

of figure 2 2. Figure 28. J. baudoni. CS-PM collection, 4.6 x 3.4 mm. Cap re r a Is. (Sardinia), 30 m depth.

Phenotypic diversity, taxonomic remarks and updated distribution of the Mediterranean Jujubinus baudoni

35 1

Figure 29. Jujubinus pseudogravinae N ordsieck, 1 973. CS-PM collection, H = 4.5 x D = 3.0 mm, Cais da Ribeirinha, S.

Miguel Is. (Azores Islands), on stones, 10 m depth. Figure 30. J. CXCLSpemtUS COValUnUS. CS-PM collection, 6.8 x 4.6 mm,

Bosa Marina (Sardinia), 30 m depth.

352

P. Mariottini, A. Di Giulio, M.Appolloni & C. Smriglio

Figure 3 1. JlijubinUS geOgVOphicilS. Flolotype, FI = 4.9 x D = 3.9 mm, Punta Engano, Mactan Island (Philippines), 80-150

m d ep th . Fig u re 3 2. J. geOgraphicUS . Topotype, 4.8 x 3.3 mm, Punta Engano, Mac tan Island (Philippines), 80-150 m depth.

Fig . 3 3. J. baudofli. MorphotypesA-C distribution.

Phenotypic diversity, taxonomic remarks and updated distribution of the Mediterranean Jujubinus baudoni

353

COAST

LOCALITY

REFERENCES

SPAIN

La Escala, Girona, Palamos

Curini Galletti, 1 982; G ian n u z zi-S a v elli

et al., 1994

Port Lligat, Cadaques

Spanu, 2011; this work

FRANCE

Pa u lilies, Roussillon; Dupott de l’Aude,

Pyrenees Orientales

M onterosato, 1891

Lerin Islands, Provence

Spanu, 2011

CORSE

P ietran era

M onterosato, 1891

Ajaccio; Ile-Rousse; Barcaggio; B astia

Locard & Caziot, 1901

R e v ellata

Scaperrotta et al., 2011

SARDINIA

Capo C accia, A lghero (S assari); A sinara

Island, Punta Aguada (Sassari); Costa

Paradiso, Trinita d’Agultu (O lbia-Tem pio)

Spanu, 2011; this work

La M addalena Archipelago (Olbia-Tempio)

This w o rk

Table 1. Records o f JujuhinUS baudoni fro m the North-Western Mediterranean area (from literature and this work).

CONCLUSIONS

Type materialof TrochuS baudoni and Jujubinus

baudoni var. incomp arabilis, together with other

shells of this species have been analysed and fig-

ured in the present work. A lectotype and paralec-

to types of T. baudoni have been designated. J.

baudoni resulted to be represented by highly di-

verse chromatic shells, and three distinct colour

morphs (A-C) have been recognized and figured,

but comparative SEM analyses demonstrated that

there are no significant differences in the shell

sculpture, suggesting that they belong to a single

specific entity. Since members of genus Jujubinus

graze on a variety of micro-algal food sources, such

as diatoms and green algal epiphytes (Peduzzi,

1987), we can infer that the different colour patterns

observed in J. baudoni are influenced by diverse

environmental factors. This could explain the oc-

currence of the same colour pattern in disjointed

populations of the same geographical area, like the

case of the colour morph B of Pietranera (Corse)

and Capo Caccia (Sardinia), which are separated by

the colour morph C (Fig. 33 and Tablel). However,

we cannot rule out a different scenario, where those

phenotypes correspond to a mosaic of sibling

species. In the present study, we prefer to be con-

servative in non-assigning a taxonomic status to

these different chromatic patterns. Genetic analyses

are required to verify our current interpretation and

further studies are necessary to reveal mechanisms

maintaining shell colour polymorphism among

populations of J. baudoni and understand if there is

a physiological selection as a driving force shaping

the p h e n o -(g e n o - ) ty p ic structure of these popu-

lations along the continental and island coasts. As

far as we know, the distribution of J. baudoni is lim-

ited to the North-Western Mediterranean Sea, with

its eastern range widened to the La M addalena Ar-

chipelago (Sardinia) (Fig. 33). On the contrary, this

species does not occur in the Azores, having been

m isidentified with J. pSCudogravinae.

ACKNOWLEDGEMENTS

W e are indebted to M arco C urini G alletti (D ipar-

timento di Scienze della Natura e del Territorio, U ni-

versita di Sassari, Italy), Franco Rapetti (Societa

Toscana di Scienze Naturali, Pisa; Italy) and Bruno

Amati (Rome, Italy) who kindly provided bibliogra-

phy related to the Jujubinus genus. Sincere thanks

are due to Sergio Bartolini (Firenze, Italy), and to

Guido and Stefano Poppe (Lapu Lapu City, Cebu,

354

P. Mariottini, A. Di Giulio, M.Appolloni & C. Smriglio

Philippines) for the permission to use a J. baudoni

and J. geOgraphicUS photographs, respectively.

REFERENCES

Appeltans W., Bouchet P., Boxshall G.A., De Broyer C

de Voogd N.J., Gordon D.P.. Hoeksema B.W., Horton

T., Kennedy M., Mees J., Poore G.C.B., Read G.,

Stohr S., Walter T.C . & Costello M .J., 2012.

World Register of Marine Species. Available at

http://www.marinespecies.org (accessed 8 May 2013).

Avila S.P.. Cardigos F. & Santos R.S., 2004. D. Joao de

Castro Bank, a shallow water hydrothermal-vent in

the Azores: checklist of the marine molluscs. Arqui-

pelago. Life and Marine Sciences, 21A: 75-80.

Avila S.P., Cardigos F. & Santos R.S., 2007. Comparison

of the community structure of the marine molluscs

of the “Banco D. Joao de Castro” seamount (Azores,

Portugal) with that of typical inshore habitats on the

Azores archipelago. Helgol Marine Research, 61:

43 -5 3.

Avila S.P.. Borges J.P. & De Frias Martins A.M., 2011.

The littoral Trochoidea (M ollusca: Gastropoda) of the

Azores. Journal of Conchology, 40: 408-427.

Byers B.A., 1989. Habitat-choice polymorphism asso-

ciated with cryptic shell-color polymorphism in the

lim pet Lottici digitalis. T he Veliger, 32: 394-402.

Cur ini Galletti M., 1982. Note ai Trochidae, VI. Jlljllbi-

flUS Bcilldoni H . Martin in Monterosato, 189 1. A tti

della Societa Toscana di Scienze Naturali, Memorie,

serie B , 8 9: 75 -85.

Giannuzzi-Savelli R., Pusateri F., PalmeriA. & Ebreo C.,

1 994. Atlas of the Mediterranean Sea Shells. “La

Conchiglia” Ed., Roma, 125 pp.

Ghisotti F. & M clone G., 1975. Catalogo il lustra to delle

conchiglie marine del M ed iterran e o , 5. Conchiglie,

1 1 , suppl. 5 : 1 47-208.

Go fas S. & Le Renard J., 2013. CLEMAM: Check List

of European Marine Mollusca. Available at

http://www.som ali.asso.fr/clem am /index.clem am .htm 1

(accessed 8 May 2012).

Locat'd A. & Caziot E., 1901. Les coquilles marines des

cotes de Corse. Annales de la Societe Linneenne de

Lyon. 47: 158-294.

M iura O., N ishi S. & Chiba S., 2007. Tern peratu re-related

diversity of shell colour in the intertidal gastropod Bci-

tillaria. Journal of M oil u scan Studies, 73: 235-240.

Monterosato T. di Maria di, 1891. Molluschi fossili qua-

ternari di S. Flavia. II N aturalista sic ilia no, 10: 96-104.

Peduzzi P.. 1 987. Dietary preferences and carbon absorp-

tion by two grazing gastropods Gibblllci IWlbiUcClris

(Linne") and JujllbhlUS StvidtUS. Marine Ecology, 8:

359-3 70.

Scaperrotta M., Bartolini S. & Bogi C., 2011. Stages

of growth of marine molluscs of the Mediterrranean

Sea. Vol. III. L’lnform atore Pice no Ed., Ancona,

184 pp.

Sokolova I.M . & Berger V.Ja., 2000. Physiological vari-

ation related to shell colour polymorphism in White

Sea Littorina scixatilis. Journal of Experim ental Ma-

rine Biology and Ecology, 245: 1 -23.

Spanu M .T., 2011. Prim a segnalazione di JujubinilS bcill-

dOTli ( Monterosato, 189 1 ex H. Martin ms) (Gastro-

poda: Trochidae) per la Sardegna e le acque italiane.

Bollettino Malacologico,47: 1 35-1 37.

Terreni G., 1981. Molluschi conchiferi del mare anti-

stante la costa toscana (Gastropoda, Scaphopoda,

Amphineura, Bivalvia, Cephalopoda). Livorno,

10 6 p p .

Biodiversity Journal, 2013, 4 (2): 355-358

New and poorly-known harvestmen (Opiliones) from Sardinia

(Italy). Part I: Lacinius horridus (Panzer, 1 794), new to Sardinia

(Phalangiidae Oligolophinae)

Salvatore Canu 1 & Lars Friman 2

'Via Uri 6, 1-07049 Usini, Sassari, Italy; e-mail: SalvatoreCanul@tin.it

2 Larchenweg 28, D- 14806 Bad Belzig, Germany; e-mail: LF0107@t-online.de

"■Corresponding author

ABSTRACT The first occurrence of Lacinius horridus (Panzer, 1794) (Phalangiidae Oligolophinae) on

Sardinia is decribed and the general biogeographical distribution of Opiliones in Italy is

discussed.

KEY WORDS Opiliones; Sardinia; Lacinius horridus ; biogeography.

Received 16.05.2013; accepted 22.06.2013; printed 30.06.2013

INTRODUCTION

Lacinius horridus (Panzer, 1794) (Phalangiidae

Oligolophinae) is a well-known species found in

nearly all areas of Southern, Central and Northern

Europe (Martens, 1978; Blick & Komposch, 2004)

with the exception of Spain (Prieto, 2008), the

Netherlands (Wijnhoven, 2009) and Norway (Stol,

2007). In Italy L. horridus has been found on the

mainland and also in Sicily (Stoch, 2003), but not

undisputedly in Sardinia.

ABBREVIATIONS. S-Fri = Sardinian specimen;

G-Fri = german specimen; Fe = femur; Pt = patella;

Ti = tibia; Mt = metatarsus; Ta = tarsus;

Lacinius horridus (Panzer, 1794) from Sardinia

Examined material. Coordinates in WGS84.

Usini, Sassari, 31.04.2012, subadult female,

40°39'49" / 08°29'49", 78 m a.s.l., legit S. Canu (S-

Fri 01-099 Rif 71/8). Usini, Sassari, 05.04 2012,

subadult female, 40°39'46" / 08°32'41", 206 m a.s.l.

legit S. Canu (S-Fri 02-010 Rif 81/2). Usini, Sas-

sari, 29.04.2012, juvenile female, 40°39'44" /

08°32'44", 206 m a.s.l., legit S. Canu (coll. “Schon-

hofer”). Usini, Sassari, 04.06.2012, subadult fe-

male, 40°39'46" / 08°32'43", 206 m a.s.l., legit S.

Canu (coll. “Schonhofer”) (Fig. 1).

Remarks. For this first reliable record of the

thermophile L. horridus all specimens were col-

lected by hand. The localities where L. horridus was

found, can be described as typical “mediterran

semi-natural dry open grasslands”, mostly on cal-

careous substrates (Fig.l; Martens 1978:325; Kom-

posch & Gruber, 2004: 516). Two of the specimens

(Nr. Rif7 1/8, = Fri 01-099 and Nr. Rif 81/2, = Fri

02-010) were selected for further examination and

have been retained as voucher specimens in collec-

tions of Department of Agriculture, Plant Pathology

and Entomology Section, University of Sassari

(Italy). In Table 1 (measurements) we have added

4 reference-specimens, 3 from Germany/Branden-

burg (G) and 1 from Martens (1978, lit.).

Without any evidence other than the ominous ci-

tation in Costa (1 885: 1 1 , but as Acantholophus hor-

356

Salvatore Canu & Lars Friman

Specimens-nrs.

body length/breadth(l/b)

leg II (Fe, Pt, Ti, Mt, Ta)

Fe I / Fe II / Fe III / Fe IV

S-Fri 01-099 Rif 71/8

2.8/ 2.0 Fb = 1.4

1.6 / 0.5 / 1.6 / 0.8 / 3.5 = 7.4

0.5 / 1.7 / 0.5 / 1.3

S-Fri 02-010 Rif 81/2

2.8/ 1.9 1/b = 1.5

1.4 / 0.5 / 1.2 / 0.7 / 2.2 = 6.0

0.4 / 1.4 / 0.4 / 1.2

G-Fri 03-030

2.0/ 1.5 1/b = 1.3

1.3 / 0.5 / 1.2 / 0.5 / 2.0 = 5.5

0.3 / 1.3 / 0.3 / 1.2

G-Fri 03-031

5.5/ 3.5 1/b = 1.6

4.5 / 1.0 / 3.7 / 1.9 / 5.8 = 16.9

1.5/ 4.5 / 1.4 / 2.9

G-Fri 03-053

6.0/ 4.6 1/b = 1.3

4.8/ 1.0/ 3.7/ 2.4/ 6.0= 17.9

1.7/ 4.8 / 1.9 / 3.7

Martens (1978) ad. females

6.0/ 3.5 1/b = 1.7

5.2/1.5/4.1/3.1/7.3 = 21.2

- / 5.2 / - / -

Table 1. Measurements of specimens in mm. G-Fri 03-030, BadBelzig, 03.09.2010, juvenile female, 52°13'21" 12°34'57",

98 m a.s.l. G-Fri 03-031, Bad Belzig, 29.10.2012, subadult female, 52°13'25" 12°34'57", 98 msm. G-Fri 03-053, Bad

Belzig, 21.08.2009, adult females, 52°13'22" 12°35'03", 98 msm.

ridus Bose, not Panzer, 1794), Marcellino (e.g.

1973: Tab. 1. as “sistema sardo-corso”, p. 7,

1975:163 and 1980: 337, Tab. 1) includes the

species Lacinius horridus as known for Sardinia

(contra Roewer, 1924, 1957 and Trossarelli, 1943;

Stoch 2003). Consequently, this unverified state-

ment by Marcellino (e.g. 1980:337: “Non ho tro-

vato specifiche menzioni per la Sardegna, salvo

Tunica citazione di Costa ...”) was not followed by

Chemini (1995) or Stoch (2003).

It is nonetheless worth mentioning that Sardinia

plays an important role in the biogeographical

distribution of Opiliones in Italy. Chemini (1996)

already observed that Italy can be divided into five

faunal territorial regions: northern, middle, and

southern peninsula; Sicily and Sardinia. Further-

more (Chemini, 1996) there is a strong latitudinal

effect (North-South-Island) that can be seen in the

increasing number of Mediterranean species from

11% (Northern Italy) to 63% in Sardinia and the

decreasing number of European species from 59%

(Northern Italy) to 0% in Sardinia. A similar

tendency can be observed in the increasing number

of endemic species from the north (25%) to 41% in

Sicily (Chemini, 1996).

In contrast, the so-called widespread species

(Chemini, 1996, but not Marcellino, 1973: “geone-

mia europea”), which include L. horridus , Mitopus

morio (Fabricius, 1799), Trogidus nepaeformis

(Scopoli, 1763 ), Phalangium opilio Linnaeus, 1761

and Odiellus spinosus (Bose, 1792) are also wide-

spread in Italy and make up about 4-5% of all

opilionid species in all regions of Italy.

Marcellino (1980: 325, Tab. 1) listed 29 species

of Opiliones for Sardinia, of which 16 species were

recognized as valid by Stoch (2003). Now with the

new species L. horridus and the long "overseen"

Mitostoma sabbadinii (Tedeschi et Sciaky, 1997,

indicated with *) the number of Opiliones on Sar-

dinia increases to 18.

Family SIRONIDAE

Parasiro minor Juberthie, 1958

Family TRAVUNIIDAE

Buemarinoa patrizii Roewer, 1956

Family PHALANGODIDAE

Ptychosoma vitellinum Soerensen, 1873

Scotolemon terricola Simon, 1872

Family NEMASTOMATIDAE

Histricostoma argenteolunulatum (Canestrini, 1875)

Mitostoma patrizii Roewer, 1953

Mitostoma sabbadinii Tedeschi et Scia, 1997 *

Family DICRANOLASMATIDAE

Dicranolasma soerenseni Thorell, 1876

New and poorly-known harvestmen (Opiliones) from Sardinia (Italy). Part I: Lacinius horridus, new to Sardinia 357

Figure 1. Lacinius hor-

ridus (coll. “Schonho-

fer”, subadult female;

Usini,: 04.06.2012) and

its habitat (Photo S.

Canu).

Family TROGULIDAE

Anelasmocephalus brignolii Martens et Chemini, 1988

Family ISCHYROPSALIDIDAE

Ischyropsalis adamii Canestrini, 1873

Family PH ALAN GIID AE

Metaphalangium propinquum (Lucas, 1847)

Dasylobus argentatus argentatus (Canestrini, 1871)

Dasylobus gestroi (Thorell, 1876)

Phalangium opilio Linnaeus, 1758

Phalangium targionii (Canestrini, 1871)

Lacinius horridus (Panzer, 1794)

Family SCLEROSOMATIDAE

Metasclerosoma sardurn (Thorell, 1876)

Nelima doriae (Canestrini, 1871)

ACKNOWLEDGEMENTS

We like to thank Prof. Dr. A. Kury (Rio de

Janeiro, Brazil) and Dr. C. Komposch (Graz,

358

Salvatore Canu & Lars Friman

Germany) for reading the manuscript and Prof. Dr.

R. Pantaleoni (Sassari, Italy) for his help and for the

Costa-papers. The English text was corrected by

Kevin Pfeiffer (Berlin, Germany).

REFERENCES

Barnabe Blick T. & Komposch C., 2004. Checkliste der

Weberknechte Mittel- und nord-europas (Checklist

of the harvestmen of Central and Northern Europe).

Arachnida: Opiliones. Version 27. http://www.ara-

ges.de/checklist/checklist04 opiliones.html

Chemini C., 1995. Arachnida, Scorpiones, Palpigradi,

Solifugae, Opiliones, Fascicolo 21. In: Minelli, A.,

Ruffo, S. et La Posta, S. (Ed.). Checklist delle Specie

della Fauna Italiana. Ministero dell'Ambiente e Co-

mitato Scientifico per la Fauna d'ltalia, pp. 1-8.

Chemini C., 1996. Biogeographic patterns in the opil-

ionids of Italy. (Arachnida: Opiliones). Bollettino del

MUseo Civico di Storia Naturale di Verona, 30:531-

538.

Costa A., 1885. Notizie ed osservazioni sulla Geo-Fauna

sarda. Memoria Quinta. Risultamento delle ricerche

fatte nel mese di maggio. Rendiconti della Reale Ac-

cademia delle Scienze Fisiche e Matematiche di Na-

poli, Vol. 1, Ser. 2,1-31.

Komposch C. & Gruber J., 2004. Weberknechte oster-

reichs. Denisia 12 (Neue Serie), 14: 485-534.

Marcellino I., 1973. Opilioni (Arachnida) delle Isole

Eolie ed Egadi. Lavori della Societa Italiana di Bio-

geographia; Nuova Serie, 3: 329-339.

Marcellino I., 1975. Opilioni Italiani del gener eLacinius

THORELL, 1876, (Arachnida, Opiliones, Phalangii-

dae). Fragmenta entomologica, 11: 149-168.

Marcellino I., 1980. Opilioni di Sardegna (Arachnida,

Opiliones). Lavori della Societa Italiana di Biogeo-

grafia, Nuova Serie, 8: 323-345.

Martens J., 1978. Die Tierwelt Deutschlands 64. Teil,

Weberknechte, Opiliones. VEB Gustav Fischer

Verlag, Jena, 464 pp.

Prieto C. E., 2008. Updating the checklist of the Iberian

opiliofauna: corrections, suppressions and additions.

Revista Iberica de Aracnologia, 16: 49-65.

Roewer F., 1924. Opilioniden von der Insel Rhodos, Ita-

lien und Sardinien sowie der Cyrenaica. Bollettino

dei Musei di Zoologia e di Anatomia Comparata della

Reale Universita di Torino, 39: 1-7.

Roewer C.F., 1957. Uber Oligolophinae, Caddoinae,

Sclerosomatinae, Leiobuninae, Neopilioninae und

Leptobuninae (Phalangiidae, Opiliones Palpatores).

(Weitere Weberknechte XX). Senckenbergiana bio-

logica, 38: 323-358.

Trossarelli F., 1943. Contributo alio studio degli Oplio-

nidi Italiani. Bollettino della Societa Entomologica

Italiana, 5: 49-54.

Stoch F., 2003. In: Zuklat P., (Ed.), Checklist of the spe-

cies of the Italian fauna. Italian Ministry of Environ-

ment - Direction for Nature Protection (online

version 2.0). http://www.faunaitalia.it/checklist/intro-

duction.html

Stol I., 2007. Checklist of Nordic Opiliones. Norwegian

Journal of Entomology, 54: 23-26.

Wijnhoven H., 2009. De Nederlandse hooiwagens

(Opiliones). Entomologische Tabellen, Nr. 23.

Biodiversity Journal, 2013, 4 (2): 359-364

The potentially invasive opisthobranch Polycera hedgpeth Er.

Marcus, 1 964 (Gastropoda Nudibranchia), introduced in a Me-

diterranean coastal lagoon

Salvatore Giacobbe 1 ' & Sergio De Matteo 2

'Department of Biological and Environmental Sciences, Viale Ferdinando Stagno d’Alcontres 31, 98166 S. Agata-Messina, Italy;

email: sgiacobbe@unime.it

2 Mare Nostrum Italia, Via A. Sforza compl. Poker, pal. G. 1-98125 Messina, Italy

* Corresponding author

ABSTRACT The non-native opisthobranch Polycera hedgpethi Er. Marcus, 1964 is reported from the

coastal lagoon of Capo Peloro (Central Mediterranean Sea) with notes on its habitat typology

and feeding behavior. The recently published opinion that P. hedgpethi established in the

Mediterranean Sea is disputed, since records of adult specimens and egg laying were not

accompanied by the presence of juveniles and/or larvae. Mussels imported from Atlantic and

North- Adriatic lagoons are hypothesized to be the source of incoming populations in the

Peloro Lagoon.

KEY WORDS Opisthobranchs; Alien species; Sicily; Biopollution; Transitional waters.

Received 27.05.2013; accepted 26.06.2013; printed 30.06.2013

INTRODUCTION

Human activities are responsible of increasing

introductions of non-indigenous species (NIS)

worldwide. The rate of NIS introductions is notably

high in the Mediterranean Sea due to the strong

anthropogenic pressure, but also favoured by the

marked longitudinal and latitudinal gradients in

physical factors, and highly diversified habitat

typologies. Since investigation efforts are mainly

focused on those organisms having a potential

impact on the human economy, taxa with indirect

or weak interaction with anthropogenic activities

are less known. The opisthobranchs, as highly

diversified taxa including both generalist and

specialized species, which may display invasive

behaviors, constitute an ideal model to study colo-

nization processes. Some examples are Aplysia

dactilomela (Rang, 1 828), Bursatella leachii leachii

(Blainville, 1817), and Melibe viridis (Kelaart, 1858)

, whose spread throughout the Mediterranean was

carefully monitored (Cinar et al., 2006; Daskos &

Zenetos, 2007; Borg et al., 2009).

Polycera hedgpethi is a small sized nudi-

branch, which has been considered an invasive

species because of its worldwide distribution

(Wilson, 2006). This species, that is known from

the Mediterranean since 1988, has recently colo-

nized transitional and coastal environments in the

northern Adriatic (Keppel et al., 2012) and south

Tyrrhenian Sea.

In this note, the occurrence of P. hedgpethi in

the Lago Faro, coastal lagoon of Capo Peloro,

Sicily, is reported. The aim of this paper is to con-

tribute to the knowledge of the spread of P.

hedgpethi in the Mediterranean, as an example of

a species that could potentially become invasive.

360

Salvatore Giacobbe & Sergio De Matteo

METHODS

Study area

The Lago Faro (38 o 16’07”N, 15 0 38’13”E),

north-eastern Sicily, is a temperate body of water

(average temperature of shallow waters 20.4°C)

characterized by a meromictic regime which deter-

mines anoxic conditions below 10 m depth. Faro

is a mesotrophic lagoon (Sacca et ah, 2008) with

high levels of microbial productivity (Leonardi et

ah, 2009). A moderate anthropogenic contamina-

tion was attributed to the extensive shoreline ur-

banization (Giacalone et al., 2004; Minutoli et al.,

2008), but the impact of molluscan farms has not

been evaluated to date.

Molluscan assemblages have been scarcely in-

vestigated in the past (Spada, 1969; Parenzan,

1979), but recent data suggest the anthropogenic

introduction of some extra-Mediterranean species,

together with the progressive rarefaction of en-

demic taxa (Giacobbe, 2012). Alien molluscs, crus-

taceans and annelids from the Faro Lagoon were

also reported by Cosentino et al. (2011), Cosentino

& Giacobbe (2011), Crocetta (2012) and Gian-

grande et al. (2012).

Sampling

This research has been carried out in the

framework of the PRA 2008/2009 program: “Set-

tlement dynamics and colonization of allochthonous

assemblages in the Capo Peloro lagoon”. Quali-

tative surveys were carried out monthly since

March 2009, along the oxygenated shallower

lake-floors (0-4 m depth). Quantitative samples

were collected in 16 stations in Spring 2012, and

replicated in Autumn 2012 and Spring 2013. Each

station, covering a 100 m 2 surface, was explored

by means of a 5x5 m grid (four replicates). The

most easily recognizable opisthobranchs were

identified in situ, photographed and counted.

Some specimens were also collected to confirm

their identification, and preserved in ethanol 95%,

or formaldehyde 4%, or frozen at -24°C, accord-

ing to the different analytical requirements. Col-

lected specimens are currently deposited at the

Department of Biological and Environmental

Sciences, Messina, Italy.

RESULTS AND DISCUSSION

A total of eight P. hedgpethi specimens were

found between April 2012 and May 2013 inside the

Faro Lagoon and adjacent channels. The specimens

were recorded in the stations shown in figure 1 , and

the data are summarized in table 1 . Four specimens

were found along the channel that connects the

Messina Strait with the lagoon (stations K14, K15,

K16). A mussel farm is located close to the inner

mouth. Mussel fanning is currently practiced in the

proximity of the K4 station, whereas stations K5

and K3 are not located near aquaculture farms.

All specimens were recorded on fouled hard

substrata, less than 1 m depth, except for the first

specimen, found at 1.5 m depth and the fourth

specimen, which was sampled on floating green

algae, Ulva rigida C. Agardh 1823 (Fig. 2). All the

specimens were found in association with bry-

ozoans, Bugula neritina (L., 1758), 3 specimens,

and Zoobotryon verticillatum (Delle Chiaje, 1822),

5 specimens. All the specimens, whose size was

comprised between 15 mm and 35 mm, showed the

characteristic chromatic pattern consisting in a

whitish background color, largely covered by dense

grey dots, with yellow-orange bands on propodial,

tentacles, rhinophores, branchial plumes and related

lateral processes, as accurately described by Keppel

et al. (2012) for Mediterranean specimens. A de-

scription of anatomical features was provided by

Cervera et al. (1988), who comprehensively con-

firmed previous records, except for some details in

the reproductive system.

Wilson (2006) indicated that P. hedgpethi is

native to California, where it is widely distributed

and common, and it was also recorded in several

localities both in Pacific and Atlantic oceans. The

species, that might be cryptogenic in the Caribbean,

was introduced to South Africa (Gosliner, 1982),

Australia (Wilson, 2006), New Zealand (Miller,

2001), Japan (Keiu, 2000), the Atlantic coasts of the

Iberian Peninsula (Caballer & Ortea, 2002), and the

Mediterranean Sea, where it was first reported from

the Fusaro Lake, south Tyrrhenian Sea (Cervera et

al., 1988). Subsequently, the species was found in

the coastal lagoon of Thau, Southern France

(http://www.seaslugfomm.net/fmd.cfin?id= 10791),

and more recently, in the North-Western Adriatic

Sea and the Venice Lagoon (Keppel et al., 2012),

the northernmost limit of its range.

The potentially invasive opisthobranch Polycera hedgpethi, introduced in a Mediterranean coastal lagoon

361

P. hedgpethi has a planktotrophic development

which favours natural dispersal (Goddard, 2004).

However, natural larval fluxes between the small

and sparsely situated Italian lagoons are difficult,

whilst human-mediated connections are frequent.

Because of the disjunct distribution of this species,

ship fouling was suggested as the most probable

vector (Ryland et al. 201 1) as its favourite prey, the

bryozoan Bugula neritina and other congeneric

species (McDonald & Nybakken, 1978; Gosliner,

Venice Lagoon

' N 4

Marina di Ravenna

Rimini

1 •'£&. v _ :,

1 VT?: V

; . -

Adriatic

Sea

Fusaro Lake

Tyrrenian

Sea

Sicily

Faro Lake

Ionian

Sea

at*.

Figure 1 . Distribution of P. hedgpethi in Italian waters. Faro Lake location and related sampling stations are reported. Figure

2. On-field photo of P. hedgpethi on floating green algae. Figure 3. On-field photo of P. hedgpethi preying upon Bugula ne-

ritina, with egg ribs.

362

Salvatore Giacobbe & Sergio De Matteo

1982) often grow on ship hulls. Nevertheless, P.

hedgpethi has been never reported from commercial

harbors in Mediterranean, while it occurs in moder-

ately polluted water bodies whose common trait is

the presence of mussel and oyster farms (Sacchi &

Renzoni, 1962; Ceccherelli & Barboni, 1983;

Malet et al., 2012; Giacobbe, 2012). (Fig. 1). The

regular occurrence in the Faro Lagoon of B. neritina

on mussels and oysters imported from both Atlantic

and Adriatic Sea, together with the Adriatic record

of P hedgpethi “on a mussel bed” (Keppel et al.,

2012), support the hypothesis of a mussel-mediated

spreading in the Mediterranean lagoons.

Spawning is documented in Spring (May 2012)

for the Faro Lagoon (Fig. 3), whilst in the Venice

Lagoon, Keppel et al. (2012) found egg masses in

Autumn, as reported by Gosliner, (1982) for South

hemisphere. Keppel et al. (2012) also found a juve-

nile specimen “which did not survive” and docu-

mented egg laying in the laboratory, followed by

larval hatch seven days later. Nevertheless, egg

laying is a common stress-induced response in opi-

sthobranchs, which does not necessarily involve

reproductive success. The low number of specimens

recorded in the Faro Lagoon confirms that P. hedg-

pethi introductions do not result in dense popula-

tions outside its native range (Cervera et al., 1988;

Caballer & Ortea, 2002; Cervera et al., 2010),

contrary to typical invasive behavior. Moreover, P.

hedgpethi was recently considered as casual in the

Mediterranean (Gofas & Zenetos, 2003; Zenetos et

al., 2010; Cervera et al., 2010; Occhipinti-Ambrogi

et al., 2011; Crocetta, 2012) until the recent records

from the Venice Lagoon, that were considered

consistent to an established population (Keppel et

al., 2012). Keppel et al. (2012) opinion was accep-

ted in the most recent revision by Crocetta et al.

(2013), because of the increased frequency of

specimens and egg depositions in different areas of

the Mediterranean, despite the fact that it was not

supported by the detection of a planktonic larvae

supply for self-sustaining populations. For a species

to be considered established, it is necessary to pro-

vide evidence of “self-maintaining and self-perpet-

uating populations” (European Commission, 2004).

In the case of P. hedgpethi such evidence is lacking

since no generation replacement has been observed.

On the contrary, the hypothesis of recurrent

pseudo-populations supported by external supplies

appears to agree with both the spatial-temporal dis-

continuity of records and disappearance of the species

from early-colonized areas (Villani, pers. comm.).

CONCLUSIONS

The present record of P hedgpethi increases the

number of NIS that have recorded in the Faro La-

goon, which might be considered a Mediterranean

hotspot for species introductions, similarly to the

larger and better known Venice Lagoon (Keppel et

al., 2012). The Mediterranean range of P hedgpethi

Specimen

n° 1

n° 2

n° 3

n° 4

n° 5

n° 6

n° 7

n° 8

Date

04/12/2012

05/16/2012

05/24/2012

02/09/2013

05/15/2013

Station

K13

K14

K15

K16

Lat. N

38°16'12.2"

38°16’15"

38°15'59.78"

38°15'57.40"

38°15'54.39"

30°16'9.4"

38°15'54.39"

Long. E

15°38'2.9"

15°38'10"

15°38'22.24"

15°38'28.04"

15°38'35.01"

15°38'2.1"

15°38'35.01"

Depth

1.5 m

0.5 m

Substratum

iron

builders

beton

green algae

builders

green algae

Association

Zoo. vert.

Bug. nerit.

Zoo. vert.

Bug. nerit.

Zoo. vert.

Table 1. Sampling data for the eight collected specimens. The associated preys, Zoobothryon verticillatum {Zoo. vert.)

and Bugula neritina {Bug. ner.), are indicated.

The potentially invasive opisthobranch Polycera hedgpethi, introduced in a Mediterranean coastal lagoon

363

suggests that its introduction in the Faro Lagoon

might be mediated by mussel and oyster fouling,

confirming the role of aquaculture as an important

pathway for introduction and/or secondary spread

into Mediterranean semi-enclosed water bodies.

Similarly to other non- indigenous and introduced

species, P hedgpethi needs to be carefully monitored

for a better management and spread prevention.

Nevertheless, this species does not appear to show

invasive characteristics despite its rapid worldwide

spread. P. hedgpethi might be recently established in

the Venice Lagoon, but this needs to be confirmed

with further evidence of the existence of self-main-

taining and self-perpetuating populations. The Venice

Lagoon might be the source of incoming populations

in the Faro Lagoon by mussel trade, but a direct in-

troduction from Atlantic farms cannot be discarded.

ACKNOWLEDGEMENTS

We tanks A. Valdes for manuscript critical review,

G. Villani for its suggestions and unpublished no-

tices, the “Provincia Regionale di Messina” and the

“Farau” mussel farm for permissions and facilities.

REFERENCES

Borg J.A., Evans J. & Schembri P.J., 2009. Occurrence

of the alien nudibranch Melibe viridis (Kelaart, 1858)

(Opisthobranchia,Tethydidae), in the Maltese Islands.

Mediterranean Marine Science, 10: 131-136.

Caballer M. & Ortea J.A., 2002. Primera cita de Polycera

hedgpethi Marcus, 1964 (Mollusca: Opisthobranchia)

para la Peninsula Iberica. Noticiario SEM, 37: 55-56.

Ceccherelli V.U. & Barboni A., 1983. Growth, survival

and yield of Mytilus galloprovincialis Lamk. on fixed

suspended culture in a bay of the Po River Delta.

Aquaculture, 34: 101-114.

Cervera J.L., Garcia-Gomez J.C., Toscano F. & Garcia

F.J., 1988. Polycera hedgpethi Marcus, 1964 (Gastro-

poda: Nudibranchia), an Indo-Pacific species dis-

covered in the Mediterranean Sea. Iberus, 8: 225-231.

Cervera J.L., Tamsouri N., Moukrim A. & Villani G.,

2010. New records of two alien opisthobranch

molluscs from the north-eastern Atlantic: Polycera

hedgpeti and Godiva quadricolor. Marine Biodiver-

sity Records, 3; e51: 4 pp.

Cinar M.E., Bilecenoglu M., Oztiirk B. & Can A., 2006.

New records of alien species on the Levantine coast

of Turkey. Aquatic Invasions, 1: 84-90.

Cosentino A., Giacobbe S. & Potoschi A., 2011. The CSI

of the Faro Lake (Messina): a natural observatory for

the incoming of marine alien species. Biologia Ma-

rina Mediterranea, 16: 132-133.

Cosentino A. & Giacobbe S., 20 1 1 . The new potential in-

vader Linopherus canariensis (Polychaeta: Amphino-

midae) in a Mediterranean coastal lake: Colonization

dynamics and morphological remarks. Marine Pollu-

tion Bulletin, 62: 236-245.

Crocetta F., 2012. Marine alien Mollusca in Italy: a

critical review and state of the knowledge. Journal of

the Marine Biological Association, 92: 1357-1365.

Crocetta F., Macali A., Furfaro G., Cooke S., Villani G.

& Valdes A., 2013. Alien molluscan established

along the italian shores: an update, with discussions

on some Mediterranean “alien species” categories.

ZooKeys, 277: 91-108.

Daskos A. & Zenetos A., 2007. Additions to the knowl-

edge of alien Opisthobranchia of Greece. Aquatic

Invasions, 2: 258-260.

European Commission, 2004. Environment Directorate-

General LIFE Focus I Alien species and nature con-

servation in the EU. The role of the LIFE program.

Luxembourg: Office for Official Publications of the

European Communities, 55 pp.

Giacalone A., GianguzzaA., Mannino M.R., Orecchio S.

& Piazzese D., 2004. Polycyclic aromatic hydrocar-

bons in sediments of marine coastal lagoons in Mes-

sina, Italy: extraction and GC/MS analysis,

distribution and sources. Polycyclic Aromatic

Compounds, 24: 135-149.

Giacobbe S., 2012. Biodiversity loss in Sicilian transi-

tional waters: the molluscs of Faro Lake. Biodiversity

Journal, 3: 501-510.

Giangrande A., Cosentino A., Lo Presti C. & Licciano

M., 2012. Sabellidae (Annelida) from the Faro

coastal lake (Messina, Ionian Sea), with the first

record of the invasive species Branchiomma bairdi

along the Italian coasts. Mediterranean Marine

Science, 13: 283-293.

Goddard J.H.R., 2004. Develpmental mode in benthic

opisthobranch molluscs from the northeast Pacific

Ocean: feeding in a sea of plenty. Canadian Journal

of Zoology, 82: 1954-1968.

Gofas S. & Zenetos A., 2003. Exotic molluscs in the

Mediterranean basin: current status and perspectives.

Oceanography and Marine Biology: an Annual

Review, 41: 237-277.

Gosliner T.M., 1982. A new record of the nudibranch

gastropod Polycera hedgpethi Marcus, from the

Indian Ocean of South Africa. Journal of Molluscan

Studies, 48: 30-35.

Keppel E., Sigovini M. & Tagliapietra D., 2012. Anew

geographical record of Polycera hedgpethi Er. Mar-

cus, 1964 (Nudibranchia: Polyceridae) and evidence

364

Salvatore Giacobbe & Sergio De Matteo

of its established presence in the Mediterranean Sea,

with a review of its geographical distribution. Marine

Biology Research, 8: 969-981.

Keiu S., 2000. Opisthobranchs of Izu Peninsula, TBS-

Britannica Co., Ltd, Tokyo. 184 pp.

Leonardi M., Azzaro F., Azzaro M., Caruso G., Mancuso

M., Monticelli L.S., Maimone G., La Ferla R., Raffa

F. & Zaccone R., 2009. A multidisciplinary study of

Cape Peloro brackish area (Messina, Italy): charac-

terizzation of tropic conditions, microbial abundances

and activities. Marine Ecology, 30: 32-42.

Malet N., Pastoureaud A., Vaquer A., Quere C. & Du-

broca L., 2012. Marine diatoms sustain growth of

bivalves in a Mediterranean lagoon. Journal of Sea

Research, 68: 20-32.

McDonald G.R., & Nybakken J.W., 1978. Additional

notes on the food of some California nudibranchs

with a summery of known food habits of California

species. Veliger, 21:110-118.

Miller M.C., 2001. Descriptions of the dorid nudibranchs

Polycerci hedgpethi Marcus, 1964 and P.fujitai Baba,

1937 in New Zealand. Journal of Molluscan Studies,

67: 491-499.

Minutoli R., Fossi M.C., Zagami G., Granata A. & Gu-

glielmo L., 2008. First application of biomarkers

approach in the zooplanktonic copepod Acartia lati-

setosa for the early management and conservation of

transitional waters ecosystem. Transitional Water

Bulletin, 1: 45-52.

Occhipinti-Ambrogi A., Marchini A., Cantone G., Castelli

A., Chimenz C., Cormaci M., Froglia C., Furnari G.,

Gambi M.C., Giaccone G., Giangrande A., Gravilli

C., Mastrototaro F., Mazziotti C., Orsi-Relini L. & Pi-

raino S., 2011. Alien species along the Italian coasts:

An overview. Biological Invasions, 13: 215-237.

Parenzan P., 1979. Fauna malacologica dei laghi di

Ganzirri e del Faro (Messina). Thalassia Salentina,

9: 67-78.

Ryland J.S., Bishop J.D.D., De Blauwe H., El Nagar A.,

MinchinD., Wood C. A. &Yunnie A.L.E., 2011. Alien

species of Bugula (Bryozoa) along the Atlantic coasts

of Europe. Aquatic Invasions, 6: 17-31.

Sacca A., Guglielmo L. & Bruni, V., 2008. Vertical and

temporal microbial community patterns in a meromic-

tic coastal lake influenced by the Straits of Messina

upwelling system. Hydrobiologia, 600: 89-104.

Sacchi C.F. & Renzoni A., 1962. L'ecologie de Mytilus

galloprovincialis (Lam.) dans Te tang littoral du Fu-

saro et les rythmes annuels et nyctemeraux des

facteurs environnants. Pubblicazioni della Stazione

Zoologica di Napoli, 32 (Suppl.): 255-293.

Spada A., 1969. Breve esplorazione malacologica del

lago di Ganzirri e del Pantano di Torre del Faro

presso Messina. Conchiglie, 5: 60-65.

Terrence M.G., 1982. Anew record of the nudibranch ga-

stropod Polycerci hedgpethi Marcus, from the Indian

Ocean of South Africa. Journal of Molluscan Studies,

48: 30-35.

Wilson N., 2006. New record of the nudibranch Poly-

cera hedgpethi Er. Marcus, 1964, in South Australia,

with a discussion on its occurrence in Australia. Re-

cords of the Western Australian Museum, Suppl.

69:137-140.

Zenetos A., Gofas S., Verlaque M., £inar M.E., Garcia-

Raso J.E., Bianchi C.N., Morri C., Azzurro E., Bile-

cenoglu M., Froglia C., Siokou I., Violanti D., Sfriso

A., San Martin G., Giangrande A., Katagan T., Bal-

lesteros E., Ramos-Espla A., Mastrototaro F., Ocana

0. , Zingone A., Gambi M.C. & Streftaris N., 2010.

Alien species in the Mediterranean by 2010. A

contribution to the application of European Union’s

Marine Strategy Framework Directive (MSFD). Part

1. Spatial distribution. Mediterranean Marine Science,

11: 381-493.

Biodiversity Journal, 2013, 4 (2): 365-378

Fossil starfishes (EchinodermataAsteriidae) and paleontolo-

gical analysis of the Pliocene of Cherasco, Piedmont region

(NW Italy)

Giovanni Repetto 1 & Erica Bicchi 2

'Museo civico «Federico Eusebio», via Vittorio Emanuele II n. 19 - 12051 Alba, Cuneo, Italy; e-mail: gianrepetto@yahoo.it

2 Pole Sciences de l'environnement, Groupe Esaip, St Barthelemy d' Anjou Cedex, France; e-mail: ebicchi@esaip.org

■"Corresponding author

ABSTRACT The remarkable discovery of two fossil starfishes in the Pliocene clay of Cherasco (NW

Italy), allowed to analyze in detail the sediment and paleontological material associated

(malacofauna and foraminifers assemblages), aiming at reconstructing paleoenvironmental

conditions in which these organisms lived. Furthermore biostratigraphic analysis based on

planktonic foraminiferal assemblages, allowed the recognition of the MP12 biozone of lower

Pliocene.

KEY WORDS Astropecten; malacofauna; Foraminifera; paleoenvironmental reconstruction; Pliocene.

Received 02.06.2013; accepted 22.06.2013; printed 30.06.2013

INTRODUCTION

The structure of starfish bodies is maintained by

soft tissues that rapidly disaggregate after death and

the discovery of remains relies on exceptional con-

ditions of preservation. In the Pliocene clay of Che-

rasco (Cuneo, Piedmont region, NW Italy), these

exceptional conditions allowed the discoveiy of two

fossil starfishes, attributed to Asteroidea. The fossil

record of starfishes is very sparse in Piedmont re-

gion. In the literature only two other discoveries of

Asteroidea are known in the area albese-braidese:

Astropecten bispinosus (Otto, 1823), reported by

Sacco (1893) for the Pliocene of Bra, and a print,

unfortunately devoid of significant details, illus-

trated by Cavallo et al. (1986), found in the Torto-

nian marls at Alba.

The remarkable discovery allowed to be analyzed

in detail the sediment and paleontological material

associated. The analysis focused first on the paleon-

tological study of fossil starfish, and in a second step

on malacofauna and foraminiferal assemblages in

order to characterize these clays of Cherasco from

paleocologic and biostratigraphic points of view, for

which no specific data exist in the literature.

MATERIAL AND METHODS

Cherasco (Cuneo, Piedmont region, NW Italy)

(Fig. 1) is located in a plain extending towards the

South West, bordered in the North West by the

Stura of Demonte river and in the East by the Ta-

naro river. In the western part of Cherasco, the

plain is strongly incised by “Rii di San Giovanni”

(30-40 meters depth). The fossil starfishes have

been discovered on the left side of the Rio Crosio,

one of the “Rii di San Giovanni” (3.5 km long).

The outcrop (20 m long) is located at 240 m a.s.l.

(44°38’48” N; 7°51’7” E), and it is composed of

grey-blue muddy clays which are continuous and

homogenous, 15 m thick (Figs. 2, 3). The Pliocene

366

Giovanni Repetto & Erica Bicchii

Figure 1. Location of Pliocene outcrop near Cherasco (Piedmont region, NW Italy). Figures 2, 3. Outcrop view of the

Pliocene clays.

muddy clays are outcropped by quaternary allu-

vions (see geological map of Italy, foglio 80

Cuneo).

The two fossils were found in the upper meter

of the outcrop. The oral ventral sides of the two

specimens were visible and in discrete preserva-

tion. The better preserved specimen has a diameter

of 60 mm (Fig. 4), while the other one is 55 mm

(Fig. 5). The muddy clay matrix, including speci-

mens, was disaggregated and observed under a mi-

croscope to analyze the malacofauna and

foraminiferal assemblages. For the malacofauna,

3 liters of sediment were observed after being

water sieved.

For foraminiferal assemblages, a sample of 300

g was previously disaggregated in hydrogen perox-

ide (120% by volume), for about 1 hour. Later it

was divided into four size fractions: >315 pm, 1 50-

315 pm, 125-150 pm, 63-125 pm. For the >315

pm, 150-315 pm, 125-150 pm fractions, a quanti-

tative study was performed on a volume of residue

after drying, split with an Otto microsplitter (Scott

Fossil starfishes and paleontological analysis of the Pliocene of Cherasco, Piedmont region (NW Italy) 367

et al., 1980) in order to obtain a representative quan-

tity of the particle size fraction. This allowed to be

evaluated the abundance of the residue, the state of

preservation of tests of foraminifera and the pres-

ence of any other organogenic and inorganic com-

ponents. The foraminiferal assemblages were

subsequently the subject of specific determination

and quantitative analysis. For the fraction 63-125

pm, which consists almost exclusively of biogenic

fragments and minute foraminifera shells, that only

in rare cases allow attribution of precise species

(Schonfeld et al., 2012 ), specific determinations

were not performed.

RESULTS

ECHINODERMATA ASTERIIDAE

Astropecten cf. irregularis pentacanthus

(Delle Chiaje, 1825)

The starfish, belonging to the same species,

were attributed without doubt to the genus Astro-

pecten Gray, 1 840; this systematic group currently

includes, worldwide, more than a hundred species

living in muddy and sandy bottoms of the littoral

system, with the exception of some species which

are bathyfile (Tortonese, 1965).

The comparison, as far as possible, with the

species of Astropecten currently present in the

Mediterranean, is particularly difficult in the ab-

sence of the clear visibility of anatomical structures

useful for this puipose. Nevertheless the Astropecten

analyzed can be attributed to the species A. irregu-

laris (Pennant, 1777) and more precisely to the sub-

species A. irregularis pentacanthus (Delle Chiaje,

1825). Fossil specimens from Cherasco, in addition

to the general morphological characteristics, showed

strong similarities with the subspecies mentioned

in: a) the apices of the arms, more or less rounded;

b) the size of the individuals close to the dimension

of individuals currently living in the Mediterranean,

which generally have a diameter of 80-120 mm; c)

the ratio R/r 4.4 to 4.7 which seems consistent with

their stage of growth, considering that it tends to be

greater in the larger specimens (Borghi & Bajo

Campos, 2008); and finally, d) because this subspe-

cies, which in the Mediterranean appears to be the

most frequent Asteroidea, is present on the muddy

or sandy-clay seabed at depths from 2 to 200 m

(Riedl, 1991). The brick-red color of the fossil spec-

imens is probably due to the processes of oxidation

of organic tissues.

Unfortunately, the clay investigated did not re-

turn ossicles of Asteroidea which, if present, could

possibly provide additional information for a more

exact specific determination.

Figure 4. Astropecten cf. irregularis pentacanthus, D 60 mm, from Cherasco, ventral side. Figure 5. Other specimen of A.

cf irregularis pentacanthus, D 55 mm, from Cherasco, ventral side.

368

Giovanni Repetto & Erica Bicchii

MALACOFAUNA

Given the extreme fragility of the shells, it was

hardly possible to isolate whole shells: even the

species of a few millimeters not resisted to the usual

treatments of water clay filtration, so it can be re-

trieved only in fragments. In association with bi-

valve fragments, have been found carbon wood,

parts of claws and fragments of crustaceans de-

capods arts and a dozen calcareous tubes of Ditrupa

sp. A list in systematic order of the species is found

below (Figs. 6-19)

Turritella spirata (Brocchi, 1814)

Examined material. 1 specimen (teleoconch al-

most complete) (Fig. 14).

Remarks. Although it is sometimes present in

shallower environments and not in pelitic facies,

generally outcrops where the species is found indi-

cate a muddy circalittoral-bathyal environment

(Borghi & Vecchi, 2005). This species, in fact, is

characteristic of shale and marl facies of the Mio-

cene and Pliocene (Malatesta, 1974).

Aporrhais uttingerianus peraraneosus (Sacco,

1893)

Examined material. 2 specimens (fragments).

Remarks. The fragments of Aporrhais Da Costa,

1778 found, show long and thin digits, which allow

to assign them to the subspecies peraraneosus. The

presence of this subspecies in the Rio Crosio is al-

ready mentioned by Charrier (1953). Probably the

long and thin digits are an adaptive character that

promotes the activities of these animals in the

particular seabed in which they live. In fact, the large

surface area between the coil and the digits tips line,

and the thinness of digits in proportion to their very

low weight, allows the bivalve, during their repta-

tion (movements), to "float" more smoothly on the

soft mud of seabed (Damarco, 1994). The ecological

niche of A. uttingerianus peraraneosus is now oc-

cupied by A. serresianus (Michaud, 1828), species

originating from the Atlantic Ocean and which oc-

casionally penetrated into the Mediterranean during

the Pliocene (Forli, 1989). This species developed

and proliferated later in the Quaternary (Ruggieri,

1971), and it is currently living in the mud-detritic

bottoms of the circalittoral zone also deep (Terreni,

1981; Brunetti & Forli, 2013).

Euspira sp.

Examined material. 1 specimen, in poor preser-

vation.

Remarks. The specimen found was too deterio-

rated and specific determination was impossible.

Phalium saburon (Bruguiere, 1792)

Examined material. 1 specimen (protoconch

more than 3 whorls of teleoconch).

Remarks. It is a euribate species, being corre-

lated to the deep muddy bottoms, in the circalittoral

and in the bathyal zones (settings) (Peres & Picard,

1964). In fact it lives in the Mediterranean Sea on

sandy muddy bottoms of the circalittoral zone

(Chirli, 2008).

Nassarius elatus (Gould, 1845)

Examined material. 2 specimens (1 juvenile)

(Figs. 6, 7). In the protoconch of the figure 7, the

slight rib characterizes the first whorls of teleoconch.

Remarks. The species has been reported by

Adam & Glibert (1975) for a dozen Italian fossil lo-

calities, both of the Lower and Upper Pliocene and

it can be confused with N. semistriatus (Brocchi,

1814). It still lives in the Mediterranean sandy-

muddy infra-circalittoral seabed.

Bela nitida Pavia, 1976

Examined material. 2 specimens (1 juvenile)

(Fig. 8).

Remarks. According to Pavia (1976) this species

is present in the lower and middle Pliocene of the

Piedmont (Monteu Roero, Valle Botto-Asti) and of

the Liguria (Rio Torsero-Ceriale).

Raphitoma leufroyi (Michaud, 1828)

Examined material. 2 specimens (1 juvenile).

(Figs. 9, 10). In figure 10 is illustrated planktotrophic

multispiral with 3 whorls, the first decorated by mi-

crogranules arranged in spirals, while the subsequent

whorls show sculpture canceled trending sigmoidal.

Fossil starfishes and paleontological analysis of the Pliocene of Cherasco, Piedmont region (NW Italy) 369

Figure 6. Nassarius elatus, h 1 1 .2 mm. Figure 7. Protoconch of N. elatus, magnification 9x. Figure 8. Bela nitida, h 3 mm, predation

hole on the penultimate whorl. Figure 9. Raphitoma leufroyi, h 2.3 mm. Figure 10. Protoconch of R. leufroyi, magnification 37x.

Figure 11. Odostomia conoidea, h 1.4, Figure 12. Idem, h 1.8 mm. Figure 13. Protoconch of O. conoidea, magnification 29x.

Figure 14. Turritella spirata, h 23 mm. Figure 15. Korobkovia oblonga, right valve, D 24.6 mm. Figure 16. Myrtea spinifera, D

6.9 mm, external and internal view of left valve, with predation hole. Figure 1 7. Neopycnodonte navicularis, external view of left

valve D 62.8 mm. Figure 18. Idem, right valve D 45.4 mm. Figure 19. Corbula gibba, D 4.4 mm, external view of right valve.

370

Giovanni Repetto & Erica Bicchii

Remarks. The specimen sampled, with two

whorls of teleoconch, is rather young, but its

preservation is excellent. This species lives on all

types of bottoms, from a few meters to about 150

m in depth (Poppe & Goto, 1991).

Odostomia conoidea (Brocchi, 1814)

Examined material. 2 specimens (Figs. 11-13).

The protoconch of the figure 13 consists of 2 whorls,

the first of which is wound in a spiral with the axis

approximately at right angles to the axis of the shell,

well represents the protoconch of “type A”, accord-

ing to van Aartsen (1987).

Remarks. According to Scaperrotta et al. (2011)

and Chirli & Micali (2011), this species lives in the

Mediterranean sea on the muddy-sandy bottoms of

the infra-circalittoral zone.

Philine cf. scabra (Muller O.F., 1784)

Examined material. 1 specimen (fragments).

Remarks. Fiving on sandy and mud-sandy bot-

toms at lower depths (Thompson, 1976).

Roxania utriculus (Brocchi, 1814)

Examined material. 1 specimen (juvenile).

Remarks. Fiving on the mud bottoms of the cir-

calittoral zone (150 m depth) (Ardovini & Cossi-

gnani, 1999).

Yoldia nitida (Brocchi, 1814)

Examined material. 1 specimen: fragments of

the right valve.

Remarks. The genus Yoldia is found on the

muddy-sandy bottoms of the infralittoral zone.

Aequipecten angelonii (De Stefani et Pantanelli,

1878)

Examined material. 1 specimen: fragments of

the left valve.

Remarks. It seems that this species was found

exclusively in clay-marl outcrops attributed to the

Pliocene: in the “turquoise clays“ of Colle Melame-

renda of coralligenous area (De Stefani & Panta-

nelli, 1878); in clays of Castelnuovo d'Asti, in

Fomaci di Savona and in Piacenza area (Sacco,

1897); in the marl of Rio Torsero in Savona (Andri

et al., 2005).

Flabellipecten bosniasckii (De Stefani et Pan-

tanelli, 1880)

Examined material. 1 specimen: fragments of

left and right valves.

Remarks. The Flabellipecten Sacco, 1897 con-

stitute a group adapted to live on soft bottoms,

therefore characterize all sandy-clayey facies of the

Upper Miocene and Pliocene (Malatesta, 1974).

Mimachlamys cf. varia (Linnaeus, 1758)

Examined material. 3 partials valves, 1 right.

Remarks. The species is ubiquitous: it lives on

rocky-gravelly-muddy bottoms, often with fine

bissus attached to the sponges (Malatesta, 1974).

Some authors (Tebble, 1966; Poppe & Goto,

1993), referring especially to the Atlantic Ocean,

reported M. varia at 83 m maximum depth, and

Terreni (1981) reported this species in Mediter-

ranean area living on the muddy-sandy bottoms in

the infra-circalittoral zone. Parenzan (1974) re-

ports it from 1350 m deep, but adds that the nor-

mal environment does not exceed perhaps a

hundred meters.

Korobkovia oblonga (Philippi, 1844)

Examined material. 1 specimen: right valve

(Fig. 15).

Remarks. This species is restricted to the silts

and fine-grained calcarenites of the base of the

Pliocene (Jimenez et al., 2009).

Neopycnodonte navicularis (Brocchi, 1814)

Examined material. 7 right valves (3 partials

and 1 juvenile) and 2 left valves (only 1 with um-

bonal area) (Figs. 17, 18).

Remarks. It seems certain, as reported by Ma-

latesta (1974), that the actual N. cochlear (Poli,

1795) result from a genetic mutation of Neopycno-

donte navicularis extinct in the late Pliocene;

regarding N. cochlear , Parenzan (1974) and Terreni

(1981) reported that it lives from a hundred feet to

over 1 000 m depth, on the muddy-debris bottoms

of the deep circalittoral zone.

Fossil starfishes and paleontological analysis of the Pliocene of Cherasco, Piedmont region (NW Italy) 371

Myrtea spinifera (Montagu, 1803)

Examined material. 1 specimen: left valve

(Fig. 16).

Remarks. In the Mediterranean it has been found

in the biocenosis of unstable bottom sediments (Ma-

latesta, 1974); in the British Islands it has been found

to a maximum depth of 1 1 0 m (Tebble, 1 966), on the

Tuscan coast (Italy); it lives in the detritic mud-bot-

tom of infra-circalittoral zone (Terreni, 1981).

Clausinella fasciata (Da Costa, 1778)

Examined material. 1 specimen: fragments of

the left valve.

Remarks. The species lives burrowing into the

gravelly, gravelly-stony, stony or sandy-muddy sed-

iment, up to about 110 m depth (Tebble, 1966).

According to Scaperrotta et al. (2009), it prefers wa-

ters with coarse gravel or gravel and mud, from a

few meters to over 1 00 m in depth.

Timoclea ovata (Pennant, 1777)

Examined material. 3 valves (partials)

Remarks. Euribathyal species, reported for the

Mediterranean from a few meters up to 700 meters

depth, on the muddy-debris bottoms (Malatesta,

1974).

Corbula gibba (Olivi, 1792)

Examined material. 1 right valve (partial) e 3

left (juveniles and partial). The external view of

right valve (Fig. 1 9) is characterized by a large ventral

rib concentric sculpture.

Remarks. In the Mediterranean it is found in the

biocenosis of the seabed debris (80-130 m) of the

infralittoral zone; with the polychaete Ditrupa arie-

tina (O.F. Muller, 1776) this species is characteristic

of the biocenosis of instable bottoms (Malatesta,

1974). This species seems to prefer muddy or

sandy-muddy bottoms (Scaperrotta et al., 2009).

The polychaete Ditrupa sp., of which we found

a dozen specimens, also lives in sandy-muddy bot-

toms and constitutes an indicator of the depth range

of substrate. It is a genus characteristic of biocoeno-

sis of unstable bottom which Peres & Picard (1964)

placed at depths between 35 to just over 100 m.

FORAMINIFERA

The residue of the sample is quantitatively

scarce. For the three fractions considered, the inor-

ganic materials are predominantly made up of abun-

dant particles of iron oxides, frequent angular

particles of quartz and lamellae of muscovite, some

rare granule mineralized with pyrite and glauconite.

The percentage of the inorganic fraction constitutes

60% of the residue for the >315 pm fraction, it

tends to decrease to about 40% in the 150-315 pm

fraction, to increase up to 80% of the residue in the

125-150 pm fraction. This increase is marked by an

abundance of muscovite flakes lamellae that be-

come prevalent.

The organogenic residue consists mainly of

abundant fragments of pyritized and partially oxi-

dized plant frustules, frequent fragments of echino-

derm spines and prevailing benthic foraminifera,

generally very well preserved. Sometimes there are

rare individuals mineralized with pyrite and glau-

conite. The planktonic foraminifera, in all fractions

analyzed, were extremely rare.

Benthic foraminifera

The benthic foraminiferal assemblages (44

species, Table 1) were represented principally by

hyaline species. Only two agglutinated species were

present, but rare ( Bigenerina nodosaria and Textu-

laria sp.). The porcellanaceous species were absent.

Species more abundant than 4% of the assemblage

of at least one station were described and consid-

ered as dominant: Bolivina anaeriensis, B. pun-

ctata :, Bulimina aculeata, B. lappa , B. inflata,

Stainforthia complanata, Uvigerina peregrina and

Valvulineria bradyana (Figs. 20-35; Fig. 36). In the

coarser fraction (>315 pm) Uvigerina peregrina

(-34%) and Valvulineria bradyana (-20%) pre-

vailed. Bulimina aculeata was almost frequent

( — 11%), while the species of genus Bolivina d'Or-

bigny, 1839 and Stainforthia complanata were rare

(<3%) (Fig. 36). In the 150-315 pm fraction the

percentage of Uvigerina peregrina was similar to

the coarser fraction, while the percentage of Valvu-

lineria bradyana decreased considerably (-13.5%).

The species of genus Bulimina d'Orbigny, 1826

showed a variable trend. Bulimina aculeata de-

creased (5%), while Bulimina lappa increased

strongly (-8%). Lenticulina calcar, absent in the

372

Giovanni Repetto & Erica Bicchii

Benthic foraminifera

>315p

150-315p

125-150p

>315p

150-315p

125-150p

Split

.1/2

.1/8

Hyaline species

Ammonia tepicla (Cushman, 1926)

0.0

0.4

Amphicoryna scalaris (Batsch, 1791)

0.0

0.7

0.0

Asterigerinata mammilla (Williamson, 1848)

0.0

0.4

Bolivina punctata d'Orbigny, 1839 (Fig. 22)

2.9

9.7

2.3

Bolivina albatrossi Cushman, 1922 (Fig. 21)

0.0

2.3

Bolivina alata (Seguenza, 1862)

0.0

3.4

Bolivina anaeriensis (Costa, 1856) (Fig. 23)

1.4

2.9

4.1

Bolivina dil alata Reuss, 1850

0.0

2.6

Bolivina spathulata (Williamson, 1858) (Fig. 24)

0.4

0.7

2.6

Bulimina aculeata d'Orbigny, 1826 (Fig. 25)

10.8

5.4

1.5

Bulimina lappa Cushman et Parker, 1937 (Fig. 20)

1.4

8.2

7.5

Bulimina inflata Seguenza, 1862 (Fig. 26)

4.7

3.9

5.3

Cassidulina carinata Silvestri, 1 896

0.0

0.4

Chilostomella oolina Schwager, 1878

1.1

0.0

Cibicides kullenbergi Parker, 1953

2.5

0.0

Cibicides sp.

0.4

0.0

3.4

Dentalina leguminiformis (Batsch, 1791)

0.4

0.0

0.4

Florilus boueanum (d'Orbigny, 1 846)

0.4

1.1

0.4

Globobulimina affinis (d'Orbigny, 1839)

1.4

0.4

0.0

Gyroidina altiformis (R.E. et K.C. Stewart, 1930)

0.4

0.0

Gyroidinoides umbonatus (Silvestri, 1898)

0.0

1.8

3.0

Heterolepa floridana (Cushman, 1922)

1.8

0.0

Lenticulina calcar (Linnaeus, 1767) (Fig. 28)

0.4

6.8

0.8

Marginulina hirsuta d'Orbigny, 1826 (Fig. 31)

0.0

1.1

0.0

Marginulina costata (Batsch, 1791) (Fig. 32)

2.2

0.0

Melonis padanum (Perconig, 1952) (Fig. 29)

2.9

0.0

Oridorsalis umbonatus (Reuss, 1851)

1.4

1.1

Pandaglandulina dinapolii Loeblich et Tappan, 1955

0.7

0.0

Planularia sp.

0.0

0.4

Fossil starfishes and paleontological analysis of the Pliocene of Cherasco, Piedmont region (NW Italy) 373

Pullenia bulloides (d'Orbigny, 1846)

1.8

2.2

1.1

Rectuvigerina bononiensis (Fomasini, 1888)

0.4

0.0

Rectuvigerina siphogenerinoides (Lipparini, 1932)

0.0

0.4

Saracenaria italica Defrance, 1824

0.0

0.4

0.0

Sphaeroidina bulloides d'Orbigny, 1826

1.4

0.0

Stainforthia complanata (Egger, 1893) (Fig. 27)

0.4

1.1

4.9

Stilostomella advena (Cushman et Laiming, 1931)

2.2

1.8

0.8

Stilostomella sp.

0.4

0.0

Trifarina angulosa (Williamson, 1858)

0.0

0.4

Uvigerina auberiana d'Orbigny, 1839

0.0

0.4

1.1

Uvigerina peregrina Cushman, 1923 (Fig. 33)

34.3

35.1

10.2

Valvulineria bradyana/ complanata (Fig. 30)

104

20.9

13.6

39.1

Agglutinated species

Bigenerina nodosaria d'Orbigny, 1 826

0.0

0.7

0.0

Textularia sp.

0.7

0.4

0.0

Total number of specimens

277

279

266

100.0

Table 1. Benthic foraminifera from Pliocene of Cherasco, Piedmont region (NW Italy).

Planktonic foraminifera

>315p

150-315p

125-150p

Split

.1/2

.1/8

Globigerina bulloides d'Orbigny, 1826

Globigerina falconensis Blow, 1959

Globigerina glutinata (Egger, 1 893)

Globigerinoides obliquus extremus Bolli et Bermudez, 1965 (Fig. 35)

Globigerinoides elongatus (d'Orbigny, 1826)

Globorotalia cf. magaritae (Bolli et Bermudez, 1965)

Neogloboquadrina acostaensis Blow, 1959

Orbulina universa d'Orbigny, 1839 (Fig. 34)

Sphaeroidinellopsis dehiscens (Parker et Jones, 1865)

Total number of specimens

Table 2. Planktonic fora min ifera from Pliocene of Cherasco, Piedmont region (NW Italy).

374

Giovanni Repetto & Erica Bicchii

Figure 20. Bulimina lappa. Figure 2F Bolivina albatrossi. Figure 22. Bolivina punctata. Figure 23. Bolivina anaeriensis.

Figure 24. Bolivina spathulata. Figure 25. Bulimina aculeata. Figure 26. Bulimina inflata. Figure 27. Stainforthia com-

planata. Figure 28. Lenticulina calcar. Figure 29. Melonis padanum. Figure 30. Valvulineria bradyana. Figure 3F Mar-

ginulina hirsuta. Figure 32. Marginulina costata. Figure 33. Uvigerina peregrina. Figure 34. Orbulina universa. Figure

35. Globigerinoides obliquus extremus. Each scale bar represents 100 |im.

Fossil starfishes and paleontological analysis of the Pliocene of Cherasco, Piedmont region (NW Italy) 375

coarser fraction, was present with a value of ~5%.

The percentage of Bulimina inflata was constant in

the three fractions considered (4-5%) (Fig. 37). In

the 125-150 pm fraction Uvigerina peregrina de-

creased with a minimum of 10%, while Valvuline-

ria bradyana increased with a maximum of 39%.

Bulimina aculeata decreased strongly (~1%); Bu-

limina lappa , on the contrary, increased (5%). Len-

ticulina calcar was still present, but with an

extremely low percentage (-0.5%). The genus Bo-

livina and Stainforthia, due to their natural small

dimensions, were more frequent in the finer frac-

tion (Fig. 37).

In the thanathocenosis the species allochthounous

were rare. Among these Ammonia tepida, Asteri-

gerinata mammilla have been recognized and in-

dividuals belonging to the genus Cibicides de

Montfort, 1808, typical of infralittoral zone. It

showed clearly signs of transport, highlighted by

the poor preservation of the shell. The autochthonous

fauna was represented by species typical of an

outer continental shelf and an upper continental

slope. For example the agglutinated Bigenerina

nodosaria, Uvigerina peregrina, Valvulineria

bradyana, Bulimina spp., Lenticulina and Boli-

vina spp. Uvigerina peregrina, a shallow infaunal

species living close to the sediment-water inter-

face, is characteristic of mesotrophic to eutrophic

continental slope settings (Lutze & Coulbourn,

1984; Corliss, 1985, 1991; Lutze, 1986; Corliss &

Emerson, 1990; Mackensen et al., 1995; Schmiedl

& Mackensen, 1997; Jannik et al., 1998; Kuhnt et

al., 1999; De Rijk et al., 2000; Schmiedl et al.,

2000; Morigi et al., 2001; Huang et al., 2002; Fon-

tanier et al., 2003).

Valvulineria bradyana is a typical species of

pelitic-sandy bottoms, characterized by low oxygen

content and rich in organic matter (Bergamin et al.,

1999). B. aculeata is generally also considered as a

eutrophic species, typical of low-oxygen environ-

ments (Murray, 1991). The Bolivina species (ex. B.

aenariensis and B. punctata), infaunals, are charac-

teristics of muddy bottoms and they are particularly

tolerant to reduced oxygenation levels, in environ-

ments such as meso-eutrophic (Blanc-Vemet, 1969;

di Napoli Alliata et al., 1970; Van der Zwaan, 1983;

Murray, 1991; Barmawidjaja et al., 1992; Sen

Gupta & Machain-Castillo, 1993).

Planktonic foraminifera

Planktonic foraminiferal assemblages were ex-

tremely poor in the three fractions considered (9

species, Table 2). Individuals, sometimes strongly

pyritizided or glauconitized of Orbulina universa

were found in association with rare Globigerina bul-

loides, G. falconensis and G. glutinata, as well as rare

Globigerinoides obliquus extremus, G. elongatus e

Sphaeroidine- llopsis dehischens. Only one specimen

of Globorotalia cf. margaritae was found in the

coarser fraction. The genus Neogloboquadrina

Frerichs et Vincent, 1 967 was also extremely rare. In-

deed only two specimens of N. acostaensis were iden-

tified. Globigerinoides obliquus extremus and Or-

bulina universa are represented in the figures 34, 35.

Figure 36. Total percentage of benthic fora min iferal dominant

species.

Figure 37. Percentage of benthic foraminifera dominant species

for all studied fraction (>315 pm, 150-315 pm, 125-150 pm).

376

Giovanni Repetto & Erica Bicchii

DISCUSSION AND CONCLUSION

In an outcrop of Pliocene clays near Rio Crosio

(Cherasco, Piemonte, NW Italy) two specimens of

fossil starfish were found, attributed to Astropecten

irregularis pentacanthus (Delle Chiaje, 1825).

The analysis of the malacofauna and the benthic

foraminiferal assemblages has provided paleoeco-

logical and paleoenvironmental data indicating a

circalittoral deep environment, little more than 100

m depth and characterized by a muddy bottom; the

trophic conditions were from meso to eutrophic,

with a rather low level of oxygenation, which thus

enabled the survival of upper and intermediate in-

faunals tolerant species, such as Uvigerina pere-

grina, Valvulineria bradyana, various species of the

genus Bulimina and genus Bolivina. On the base of

planktonic foraminiferal assemblages ( Globigeri -

noides obliquus extremus, G. elongatus, genus

Sphaeroidinellopsis Banner et Blow, 1959 and Glo-

borotalia margaritae ), the samples are attributed to

biozone MP12 of lower Pliocene (Zanclean) which,

according to Violanti (2012) in the NW Italy, corre-

spond to 5.08 and 4.52 Ma. The absence of species

such as Globorotalia inflata and Bulimina margi-

nata, very frequent in the upper Pliocene (Lentini,

1969), allows to confirm this datation.

ACKNOWLEDGEMENTS

We are sincerely grateful to B. Tibaldi and G.

Grisotto, for providing the samples for this study

and the data about outcrop; R. Brancato, L. Lacroce

and E. Borghi for the useful data provided. We also

thanks E. Ferrero Mortara for promoting the collab-

oration of authors and for your helpful suggestions

and advice; A. Pagano and S. O’Brien for revision

of the english version and S. Cavagna for the fora-

miniferal SEM photos.

REFERENCES

Aartsen J.J. van, 1987. European Pyramidellidae: III.

Odostomia and Ondina. Bollettino Malacologico, 23:

1-33.

Adam W. & Glibert M., 1975. Contribution a la connais-

sance de Nassarius semistriatus (Brocchi, 1814)

(Mollusca: Gastropoda). Bulletin de l’lnstitut Royal

des Sciences Naturelles de Belgique, 50: 1-78.

Andri E., Tagliamacco A., Testa M. & Marchini A., 2005.

Le malacofaune fossili del Rio Torsero. Regione Li-

guria, Catalogo dei beni naturali, 5 . Nuova Editrice

Genovese, Ceriale, 286 pp.

Ardovini R. & Cossignani T., 1999. Atlante delle conchi-

glie di profondita del Mediterraneo. L’lnformatore

Piceno Ed., Ancona, 111 pp.

Barmawidjaja D.M., Jorissen F.J., Puskaric S. & Van der

Zwaan G.J., 1992. Microhabitat selection by benthic

foraminifera in the northern Adriatic sea. Journal of

Foraminiferal Research, 22: 297-317.

Bergamin L., Di Bella L. & Carboni M.G.,1999. Valvu-

lineria bradyana (Fornasini) in organig matter-en-

riched environment (Ombrone river mouth, Central

Italy). II Quaternario, Italian Journal of Quaternary

Sciences, 12: 51-56.

Blanc- Vernet L., 1969. Contribution a E etude de forami-

niferes de Mediterranee. Recueil des Travaux de la-

Station Marine d'Endoume, 64: 1-315.

Borghi E. & Bajo Campos I., 2008. Asteroidei fossili del-

l’Emilia. Notiziario della Societa Reggiana di

Scienze Naturali, 13-29.

Borghi M. & Vecchi G., 2005. La malacofauna plio-plei-

stocenica del torrente Stirone (PR). Cerithiidae - Tur-

ritellidae. Parva Naturalia, 7: 3-46.

Brunetti M.M. & Forli M., 2013. The genus Aporrhais

Da Costa, 1778 (Gastropoda Aporrhaidae) in the kal-

ian Plio-Pleistocene. Biodiversity Journal, 4: 183-

208.

Cavallo O., Macagno M. & Pavia G., 1986. Fossili del-

l'Albese: aspetti geologici, e paleontologici delle Lan-

ghe e del Roero. Famija Albeisa, Alba, 223 pp.

Charrier G., 1953. Brevi note sulle forme del gen. Apor-

rhais ( Chenopus ) della fauna pliocenica di San Gau-

denzio (Lessona) nel Biellese. Rivista Italiana di

Paleontologia e Stratigrafia, Milano, 59: 195-202.

Chirli C., 2008. Malacofauna pliocenica toscana. Neo-

taenioglossa. Vol. 6. Grafiche PDB, Tavernelle, 128

pp.

Chirli C. & Micali P., 2011. Malacofauna pliocenica to-

scana. Pyramidelloidea. Vol. 8. Grafiche PDB, Taver-

nelle, 132 pp.

Corliss B.H., 1985. Microhabitats of benthic foraminifera

within deep-sea sediments. Nature, 314: 435-438.

Corliss B.H., 1991. Morphology and microhabitat prefer-

ences of benthic foraminifera from the northwest

Atlantic Ocean. Marine Micropaleontology, 17: 195-

236.

Corliss B.H. & Emerson S., 1990. Distribution of Rose

Bengal stained deep-sea benthic foraminifera from

the Nova Scotian continental margin and Gulf of

Maine. Deep-Sea Research, 37: 381-400.

Damarco P., 1994. Alcuni esemplari eccezionali di Apor-

rhaidae fossili del Pliocene piemontese. World Shells,

3:28-30.

Fossil starfishes and paleontological analysis of the Pliocene of Cherasco, Piedmont region (NW Italy) 377

De Rijk S., Jorissen F.J., Rohling E.J. & Troelstra S.R.,

2000. Organic flux control on bathymetric zonation

of Mediterranean benthic foraminifera. Marine Mi-

cropaleontology, 40: 151-166.

De Stefani C. & Pantanelli D., 1878-80. Molluschi plio-

cenici dei dintomi di Siena. Bullettino della Societa

malacologica italiana, 4 (1-6): 5-48 [1878], 4 (7-14):

49-112 [1879], 4 (15-20): 113-160 [1879], 4 (21-30):

161-215 [1880],

Di Napoli Alliata E., Stefanini S. & Fioramonti G., 1970.

Etude de quelques carottes du plateau et du talus con-

tinental de la Sardaigne meridionale. Bollettino della

Societa Geologica Italiana, 89: 209-232.

Fontanier C., Jorissen F.J., Chaillou G., David C., An-

schutz R & Lafon V., 2003. Seasonal and interannual

variability of benthic foraminiferal faunas at 550 m

depth in the Bay of Biscay. Deep-Sea Research, Part

I: Oceanographic Research Papers, 50: 457-494.

Forli M., 1989. Considerazioni filogenetiche su alcune

specie del genere Aporrhais da Costa, 1778 (Mollu-

sca: Gastropoda) (ultima parte). Argonauta, 5: 49-54.

Huang B., Jian Z., Cheng X. & Wang P, 2002.

Foraminiferal responses to upwelling variations in

the South China Sea over the last 220000 years. Ma-

rine Micropaleontology, 47: 1-15.

Jannik N.T., Zachariasse W.J. & Van der Zwaan G.J.,

1998. Living (Rose Bengal stained) benthic

foraminifera from the Pakistan continental margin

(northern Arabian Sea). Deep-Sea Research Part I.

Oceanographic research Papers, 45: 1483-1513.

Jimenez A.P., Aguirre J. & Rivas P, 2009. Taxonomic

study of scallops (Pectinidae: Mollusca, Bivalvia)

from Pliocene deposits (Almeria, SE Spain). Re vista

Espanola de Paleontologia, 24: 1-30.

Kuhnt W., Hess S. & Jian Z., 1999. Quantitative compo-

sition of benthic foraminiferal assemblages as a

proxy indicator for organic carbon flux rates in the

South China Sea. Marine Geology, 156: 123-157.

Lentini F., 1969. Sezioni stratigrafiche plioceniche nella

Val d’Agri presso Gannano (Matera). Atti dell'Acca-

demia Gioenia di Scienze Naturali di Catania, 20

(Suppl. Sc. Geolog.): 19-79.

Lutze G.F., 1986. Uvigerina species of the eastern North

Atlantic. In: Van der Zwaan G.J., Jorissen F.J., Ver-

hallen P.J.J.M. & von Daniels C.H. (Eds.), Atlantic -

European Oligocene to Recent Uvigerina. Utrecht

Micropaleontological Bulletins, 35: 21-46.

Lutze G. & Coulbourn W., 1984. Recent benthic

foraminifera from the continental margin off north-

west Africa: community structure and distribution.

Marine Micropaleontology, 8: 361-401.

Mackensen A., Schmiedl G., Harloff J. & Giese M.,

1995. Deep-sea foraminifera in the South Atlantic

Ocean: ecology and assemblage generation. Micropa-

leontology, 41: 342-358.

MalatestaA., 1974. Malacofauna pliocenica umbra. Me-

morie per servire alia descrizione della carta geolo-

gica d'ltalia: 13: XI+498.

Morigi C., Jorissen F.J., Gervais A., Guichard S. & Bor-

setti A.M., 2001. Benthic foraminiferal faunas in sur-

face sediments off NW Africa: relationship with the

organic flux to the ocean floor. Journal of Foramini-

feral Research, 31: 350-368.

Murray J.W., 1991. Ecology and distribution. Benthos

90, Tokai University Press, Sendai.

Parenzan P, 1974. Carta d’identita delle conchiglie del

Mediterraneo. Vol. II. Bivalvi (prima parte). Bios

Taras Editrice, Taranto, 280 pp.

Pavia G., 1976. I Molluschi del Pliocene inferiore di

Monteu Roero (Alba, Italia NW). Bollettino della So-

cieta Paleontologica Italiana, 14: 99-175.

Peres J.M. & Picard J., 1964. Nouveau manuel de Bio-

nomie benthique de la Mer Mediterranee. Recueil des

Travaux de la Station Marine d’Endoume, 31: 1-137.

Poppe G.T. & Goto Y., 1991. European Seashells. Vol.

1 (Polyplacophora, Caudofoveata, Solenogastra,

Gastropoda). Verlag Christa Hemmen, Wiesbaden,

352 pp.

Poppe G.T. & Goto Y., 1993. European Seashells. Vol. 2.

(Scaphopoda, Bivalvia, Cephalopoda). Verlag Christa

Hemmen, Wiesbaden, 221 pp.

Riedl R., 1991. Fauna e Flora del Mediterraneo. Franco

Muzzio Editore, Padova, 777 pp.

Ruggieri G., 1972. Considerazioni su alcune Aporrhais

del Neogene italiano e sul loro significato stratigra-

fico. Rivista Mineraria Siciliana, Palermo, 22: 14-20.

Sacco F., 1893. Sopra alcuni Asteroidei fossili. Atti Regia

Accademia delle Scienze, Torino, 28: 739-745.

Sacco F., 1897. I Molluschi dei terreni terziarii del Pie-

monte e della Liguria. Parte XXIV. (Pectinidae).

Carlo Clausen, Torino, 84 pp., 21 pis.

Scaperrotta M., Bartolini S. & Bogi C., 2009. Accresci-

menti. Stadi di accrescimento dei molluschi marini

del Mediterraneo. Vol. I. L’lnformatore Piceno Ed.,

Ancona, 167 pp.

Scaperrotta M., Bartolini S. & Bogi C., 2011. Accresci-

menti. Stadi di accrescimento dei molluschi marini

del Mediterraneo. Vol. III. L’lnformatore Piceno Ed.,

Ancona, 184 pp.

Schmiedl G. & Mackensen A., 1997. Late Quaternary pa-

leoproductivity and deep water circulation in the east-

ern South Atlantic Ocean: evidence from benthic

foraminifera. Palaeogeography, Palaeoclimatology,

Palaeoecology, 130: 43-80.

Schmiedl G., De Bovee F., Buscail R., Charriere B.,

Hemleben C., Medernach L. & Picon P, 2000. Tro-

phic control of benthic foraminiferal abundances and

microhabitat in the bathyal Gulf of Lions, western

Mediterranean Sea. Marine Micropaleontology, 40:

167-188.

378

Giovanni Repetto & Erica Bicchii

Schonfeld J., Alve E., Geslin E., Jorissen F., Korsun S.,

Spezzaferri S. and members of the FOBIMO group,

2012. The FOBIMO (FOraminiferal BIo-MOnitor-

ing) initiative - Towards a standardised protocol for

soft-bottom benthic foraminiferal monitoring studies.

Marine Micropaleontology, 94-95: 1-13.

Scott D.B., Schafer C.T. & Medioli F.S., 1980. Eastern

Canadian estuarine foraminifera: a framework for

comparison. Journal of Foraminiferal Research, 10:

205-234.

Sen Gupta B.K. & Machain-Castillo M.F., 1993. Benthic

foraminifera in oxygen-poor habitats. Marine Micropa-

leontology, 20: 183-201.

Tebble N., 1966. British Bivalve Seashells. The British

Museum of Natural History, Fondon, 212 pp.

Terreni G., 1981. Molluschi conchiferi del mare anti-

stante la costa toscana (Gastropoda, Scaphopoda,

Amphineura, Bivalvia, Cephalopoda). Tip. Benvenuti

& Cavaciocchi, Fivorno, 106 pp.

Thompson T.E., 1976. Biology of Opisthobranch Mol-

luscs. Vol. I. The Ray Society, Fondon, 206 pp.

Tortonese E., 1965. Echinodermata. Fauna d’ltalia, Vol.

VI. Calderini Ed., Bologna, 422 pp.

Van der Zwaan G.J., 1983. Quantitative analyses and the

reconstruction of benthic foraminiferal communities.

Utrecht Micropaleontological Bullettin, 26: 49-69.

Violanti D., 2012. Pliocene Mediterranean Foraminiferal

Biostratigraphy: A Synthesis and Application to the

Paleoenvironmental Evolution of Northwestern Italy,

Stratigraphic Analysis of Fayered Deposits. 123-160.

Dr. Omer Elitok (Ed.), InTech, Available from:

http://cdn.intechopen.com/pdfs/3632 1/InTech-

Pliocenejnediterranean foraminiferal biostratigra-

phy_a_synthesis_and_application_to_the_paleoenvi-

ronmental_evolution_of_northwestern_italy.pdf

Biodiversity Journal, 2013, 4 (2): 379-382

A remarkable new flightless Madrasostes Paulian, 1 975 from

Vietnam (Coleoptera Scarabaeoidea Hybosoridae Cerato-

canthinae)

Alberto Ballerio

Viale Venezia 45, 25123 Brescia, Italy, alberto.ballerio.bs@numerica.it

ABSTRACT Madrasostes bartolozzii n. sp. is described from Vietnam (Ba Be National Park) on the basis

of two females. Some remarks on this unusual flightless species are provided. Distinguishing

characters between Madrasostes Paulian, 1975 and Besuchestostes Paulian, 1972 are briefly

discussed.

KEY WORDS Ba Be National Park; Ceratocanthinae; Flightlessness; Madrasostes', new species.

Received 10.06.2013; accepted 26.06.2013; printed 30.06.2013

INTRODUCTION

The last (and only) synopsis of Vietnamese Ce-

ratocanthinae (Coleoptera Scarabaeoidea Hyboso-

ridae) was the volume by Renaud Paulian (1945)

on the Coleoptera Scarabaeoidea of French Indo-

china, which listed four species: Madrasostes ton-

kinense (Paulian, 1945), Pterorthochaetes incertus

Gestro, 1899, Pterorthochaetes coomani Paulian,

1945 and Pterorthochaetes armatus Paulian, 1945.

No additions have been recorded since then (Pau-

lian, 1978; Ocampo & Ballerio, 2006). Paulian's

work was based exclusively on material from Hoa

Binh region (Northern Vietnam), and the country

still awaits a proper exploration in order to inven-

tory its Ceratocanthine fauna, which will probably

score a final number several times larger than the

four species listed above (for instance the study of

the P incertus from Hoa Binh kept in the Museum

National d'Histoire Naturelle collection in Paris, in

the course of an ongoing revision of the genus Pte-

rorthochaetes Gestro, 1899, revealed that P. incer-

tus does not occur in Vietnam and that the alleged

P. incertus specimens actually represent three un-

described species). It is not surprising therefore that

an expedition organized in 20 1 1 by the team of the

Museo Zoologico La Specola of Florence discov-

ered a new species of the genus Madrasostes Pau-

lian, 1975, remarkable by its unusual morphology,

which is herein described.

METHODS

I refer to Ballerio et al. (2011) and references

therein quoted for methods and terminological

conventions. Habitus photographs were taken

with a Canon Eos D5 Mil with a macro objective

MP 65 mm, all photos were then mounted with

the Zerene Stacker software and cleaned and un-

masked using a photo processing software.

ABBREVIATIONS. EL = maximum elytral

length; EW = maximum total elytral width;

HL = maximum head length; HW maximum head

width; PL = maximum pronotal length at middle;

PW = maximum pronotal width at middle; W/L

ratio width length.

380

Alberto Ballerio

Madrasostes bartolozzii n. sp.

Examined material. Holotypus, female, in

Coll. Vietnam National Museum of Nature, Hanoi,

Vietnam: Northern Vietnam, Bac Kan Province, Ba

Be National Park (-350 m), 22°25'0.69"N

105°37'53.16"E, 3-8.VI.2011, L. Bartolozzi, S.

Bambi, F. Fabiano & E. Orbach leg. Paratypus, fe-

male, in Coll. Museo Zoologico La Specola, Fi-

renze: same data as holotypus.

Description of holotypus. HL =0.69 mm;

HW = 1.44 mm; PL = 1.38 mm; PW = 2.33 mm;

EL = 2.36 mm; EW = 2.27 mm.

Small sized Madrasostes (Figs. 1-4), flightless,

shiny dark-brown with green metallic sheen on re-

liefs, antennae, tarsi and sternum reddish-brown.

Glabrous (45 x). Head subpentagonal, wider than

long (W/L ratio= 1.26). Clypeal extremity pointed,

distinctly protruding forwards and apically slightly

bent upwards (probably a sexually dimorphic cha-

racter). Fore side of clypeus serrate, clypeopleuron

short, genae slightly protruding outwards, genal

canthus indistinct, dorsal ocular area absent. Head

dorsal surface completely covered by dense im-

pressed comma-shaped punctures, punctures lar-

ger on disc and frons and smaller on head base,

centrifugally oriented, each one bearing a simple

puncture next to inferior side, punctures dense, al-

most touching each other. Eyes small, longitudi-

nally elongate. Antennae 10-segmented, scape

long (about half the total length of antenna), sli-

ghtly clavate distad, pedicellus cupuliform, flagel-

lum short, made of short articles distinctly wider

than long, antennal club three-segmented, unifor-

mly hairy.

Mouthparts. Labrum wide and short, hemicircu-

lar, surface finely wrinkled, bearing medially a trans-

verse row of about ten long erect fine setae and

distally fringed by a dense fine setation directed for-

ward. Distal epipharynx bisinuate, longitudinally di-

vided by a strong anterior median process; median

brush and corypha absent; apical fringe made of

long, fine setae, absent in the middle. Mentum ven-

trally flat, widely emarginated in the middle,

emargination regularly wide-U-shaped; labial palpi

(including palpiger) four-segmented, first segment

short and transverse, segment two short, segment

three longer and plumper than preceding two toge-

ther, segment four subconical, about as long as seg-

ment three, apically bearing some short sensilla, all

segments, apart from the last one, fringed with long

setae. Maxillae with an elongate single membranous

lacinia, covered with fine long setae, monolobed

galea proximally sclerotized and distally clothed

with coarse long, fine setae with distinctive comb-

like tip (galeal brush), maxillary palpi (including

palpiger) four-segmented, palpiger small, segment

two wide and short, segment three short, segment

four long and subconical, about as long as preceding

two together, apically bearing some short sensilla.

Mandibles short, regularly curved, apicalis with

pointed apical tooth very short and blunt, not pro-

truding over mesal brush, lateral sclerite of apicalis

bearing a distinct large pore, mesal brush wide and

well developed, basalis with molar lobe strong.

Pronotum wider than long (W/L ratio= 1 .6), fore

angles truncate and sinuate externally, fore margin

bisinuate, with visible bead, lateral and basal margin

with a thick bead not visible from above, basal mar-

gin slightly swollen upwards, pronotal sculpturing

as follows: whole surface, apart from area near fore

angles, uniformly covered by veiy fine impressed

simple punctures, their distance being one or two

times their diameter. Area near fore angles covered

by a mix of coarse shallow comma-shaped and ocel-

late punctures. Two large longitudinal irregular

raised cariniform processes on disc, convergent

anteriad, basally, next to each cariniform process,

there is a large blunt tubercle, and next to the tuber-

cle there are two transversal processes subparallel to

each other, the superior one is an irregular carina,

while the inferior is a pair of tubercles; surface of

carinae and tubercles smooth, apart from a few

sparse large impressed simple punctures.

Scutellum wider than long (W/L ratio=l .4), sides

proximally subparallel and distinctly notched by ely-

tral articular process, then convergent to form a

triangle with elongate acute apex and sides slightly

curved inward. Surface slightly depressed in the

middle, covered by irregular impressed comma-

shaped punctures. Elytra slightly longer than wide

(W/L ratio=0.98), sutural interstria raised and

cariniform, another longitudinal strongly raised

complete carina next to sutural carina, one further

longitudinal complete raised carina, starting from

the end of proximal third and ending at the

beginning of the distal third, one fourth short raised

carina occupying most of median third and finally a

fifth longitudinal raised carina starting at humerus

A remarkable new flightless Madrasostes from Vietnam (Coleoptera Scarabaeoidea Hybosoridae Ceratocanthinae) 381

Figures 1-4. Madrasostes bartolozzii n. sp. Fig. 1: Holotypus dorsal view. Fig. 2: rolled up holotypus dorsal view. Fig. 3:

rolled up holotypus ventral view. Fig. 4: rolled up holotypus lateral view.

(and inglobating humeral callus) and ending near

apex, interruped medially in correspondence with

the fourth carina. Surface of carinae smooth, surface

between carinae covered by two irregular rows of

longitudinally oriented elongate and narrow shallow

horseshoe-shaped punctures, with opening back-

wards and having a simple puncture in the middle.

Pseudoepipleure absent, marginal area broad,

expanded outwards, coarsely wrinkled, inferior su-

tural stria not present, articular area not visible.

Fore tibiae ending with two outer teeth, outer mar-

gin coarsely serrate, apical spur long and broadly

curved downwards. Meso- and metatibiae ending

with two apical spurs.

Variability. The paratypus does not show any

appreciable variation compared to the holotypus in

size, colour, shape and sculpturing.

Etymology. I am pleased to dedicate this re-

markable new species to Dr. Luca Bartolozzi

(Museo Zoologico La Specola, Firenze), who col-

lected the type series.

Distribution and Biology. Known only from

the type locality, Ba Be national Park in Northern

Vietnam. The area falls within the South China-

Vietnam Subtropical Evergreen Forest ecoregion

(Wikramanayake et al., 2002). The type series was

collected by sifting leaf litter collected in the rain-

forest near the park headquarters (Bartolozzi, pers.

comm.).

Comparative notes. M. bartolozzii n. sp. can be

easily distinguished from all other Madrasostes be-

cause of the following combination of characters:

head with serrate clypeal margin, genal canthus in-

distinct without any dorsal ocular area, pronotum

with surface covered by a dense very fine punc-

turation, elytra with five strongly raised longitudi-

nal carinae, flightless.

Remarks. This new species strongly diverges

morphologically from all other known Madraso-

stes. No other known Madrasostes displays such a

combination of characters and at present it is not

382

Alberto Ballerio

possible even to find its closest relatives. The un-

usual puncturation pattern of pronotum, made of

very fine simple punctures is not found in any other

known Madrasostes and is unique even within Ce-

ratocanthinae. The combination of flightlessness, in-

distinct eye canthus and dorsal ocular area absent and

strong carinate elytra could lead to place this species

in the genus Besuchetostes Paulian, 1972, according

to the criteria set by Paulian (1978). However a re-

definition of the genus Besuchetostes in preparation

will soon circumscribe the genus to the Indian and

Sri Lankan species ascribed to it (Ballerio, in prep.).

Madrasostes bartolozzi n. sp. shares some characters

typical of the Perignamptus genus group, as defined

by Ballerio (2009), and absent in true Besuchetostes ,

such as the shape of epipharynx and the presence of

a basal pore in the mandibles. Indistinct eye canthus

and dorsal ocular area absent as well as strong cari-

nate elytra and the pattern of distribution of tubercles

on pronotum are probably the result of convergent

evolution, due to flightlessness and strong adaptation

to life in leaf litter.

Therefore the species is here attributed to Ma-

drasostes, even if as previously highlighted in Bal-

lerio (2009), this placement must be seen as

provisional, due to the unsatisfactory current de-

finition of the genus Madrasostes and its allies

within the Perignamptus generic group.

REFERENCES

Ballerio A., 2009. Unusual morphology in a new genus

and species of Ceratocanthinae from New Guinea

(Coleoptera: Scarabaeoidea: Hybosoridae). The Co-

leopterists Bulletin, 63: 44-53.

Ballerio A., B.D. Gill & V. Grebennikov. 2011. Illustrated

overview and identification key to Cameroonian Ce-

ratocanthinae beetles (Coleoptera: Scarabaeoidea:

Hybosoridae) with description of four new species.

Zootaxa2892: 1-24.

Ocampo F.C. & Ballerio A., 2006. Catalog of the subfa-

milies Anaidinae, Ceratocanthinae, Hybosorinae, Li-

parochrinae, and Pachyplectrinae (Scarabaeoidea:

Hybosoridae). In Ocampo F.C. (Ed.) Phylogenetic

analysis of the scarab family Hybosoridae and mono-

graphic revision of the New World subfamily Anaidi-

nae (Coleoptera: Scarabaeoidea). Bulletin of the

University of Nebraska State Museum, 19: 178-209.

Paulian R., 1945. Coleopteres Scarabaeides de l’lndo-

chine. Premiere partie. Faune de l’Empire Frangais

III. Librarie Larose, Paris, 228 pp.

Paulian R., 1978. Revision des Ceratocanthidae [Col.

Scarabaeoidea] II - Les especes orientales et austra-

liennes. Annales de la Societe Entomologique de

France, 14: 479-514.

Wikramanayake E., Dinerstein E., Loucks C.J., Olson

D.M., Morrison J., Lamoreux J., McKnight M. &

Hedao P., 2002. Terrestrial Ecoregions of the Indo-

Pacific. A Conservation Assessment. Island Press,

Washington, 643 pp.