Vol. 23 (1)

REVISTA DE LA

SOCIEDAD ESPAÑOLA

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Oviedo, junio 2005

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Revista de la

SOCIEDAD ESPAÑOLA DE MALACOLOGÍA

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Universidad de Málaga, España

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Apdo. 156, Mieres del Camino, Asturias, España

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Embajada de España, Japón

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REVISTA DE LA

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Vol. 23 (1) Oviedo, junio 2005

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Dep. Leg. B-43072-81

ISSN 0212-3010

Diseño y maquetación: Gonzalo Rodríguez

Impresión: LOREDO, S. L. - Gijón

O Sociedad Española de Malacología —__——T— Iberus, 23 (1): 1-13, 2005

Toxic effects of taraxerol extracted from Codiaeum variega-

tum stem-bark on target vector snail Lymnaea acuminata

and non-target fish

Toxicidad del taraxerol extraído del tallo de Codiaeum variegatum

sobre el caracol Lymnaea acuminata y sobre peces

Ram P YADAV*, Sudhanshu TIWARÍ* and Ajay SINGH*!

Recibido el 26-V-2003. Aceptado el 7-IV-2004

ABSTRACT

The active compound taraxerol (triterpene) extracted from the stem-bark of Codiaeum var-

iegatum Blume (Euphorbiaceae) was administered for 24h or 96h to the freshwater snail

lymnaea (Radix) acuminata Lamarck in order to test its lethality. It was observed that the

molluscicidal activity of taraxerol was time as well as dose dependent for the snail at all

the exposure periods. There was a significant negative correlation between LC values of

taraxerol and exposure period, thus LCso values decrease from 1.69 mg/L (24h) to 0.74

mg/L (96h) against lymnaea (Radix) acuminata Lamarck. Toxicity experiments were also

carried out on non-target freshwater fish Channa punctatus (Bloch) (Channidae, Ophi-

cephalidae), which shares the habitat with snails, for environmental toxicity, if any.

Exposure of sub-lethal (40% and 80% of LC50)] doses of taraxerol over 24h caused signifi-

cant alterations in carbohydrates and nitrogenous metabolism in nervous, hepatopancreas

and ovotestis tissues of snail Lymnaea acuminata. Fish Channa punctatus also exposed to

sub-lethal doses of taraxerol (40% and 80% of LCso 24h of lymnaea acuminata) for 96h

periods in order to measure potential effects on non-target organisms , also show significant

alteration in carbohydrates and nitrogenous metabolism in muscle, liver and gonadal tis-

sues. This study also shows that there ¡is partial recovery in these parameters in both snail

and fish after the 7 day of withdrawal of the treatment, which supports the view that it is

safe to use plant products as molluscicides for controlling snails in aquatic bodies.

RESUMEN

Se administró el principio activo taraxerol (un triterpeno) obtenido de los tallos de Codiaeum

variegatum Blume (Euphorbiaceae) al caracol Lymnaea (Radix) acuminata Lamarck durante

24 0 96 horas, para probar su letalidad. La actividad molusquicida del compuesto depende

tanto del tiempo como de la dosis para ambos periodos de exposición. Hay una correlación

negativa significativa entre los valores LC del compuesto y el periodo de exposición, los val-

ores LCso descienden de 1,69 mg/L (24h) a 0,74 mg/L (96h) para los caracoles. También

se probó el compuesto en el pez Channa punctatus (Bloch) (Channidae, Ophicephalidae),

que comporte hábitat con el molusco, para determinar la posible toxicidad ambiental.

la exposición durante 24 horas a dosis subletales (40 y 80% de LC50) provocó

alteraciones en el metabolismo de carbohidratos y compuestos nitrogenados en tejidos

* Natural Product Laboratory, Department of Zoology, D.D.U. Gorakhpur University, Gorakhpur- 273 009

(U.P.) India.

'Corresponding Author: Dr. Ajay Singh, E-mail: ajay_sCsancharnet.in

Iberus, 23 (1), 2005

nervioso, del hepatopáncreas y ovotestis del caracol. Exposiciones de 96 horas de los

peces a las mismas dosis mostraron los mismos efectos sobre los tejidos muscular, hep-

ático y gonadal. El estudio muestra que hay una recuperación parcial de los parámetros

en ambos organismos al septimo día tras abandonar el tratamiento, lo que apoya la idea

de que es seguro el uso de productos vegetales como molusquicidas en medios acuáticos.

KEY WORDS: Lymnaea acuminata, Channa punctatus, molluscicides, taraxerol, metabolism.

PALABRAS CLAVE: Lymnaea acuminata, Channa punctatus, molusquicida, taraxerol, metabolismo.

INTRODUCTION

Once a pesticide is released into the

environment, chemical, physical, biolog-

ical and other allied factors determine

its fate and distribution in the ecosys-

tem. Heavy use of pesticides to control

pests results in the pollution of the envi-

ronment affecting both target species

snails, slugs etc. and many non-target

species viz. Phytoplankton, Zooplank-

ton, fish etc in freshwater ecosystems

(GOPAL, KHANNA, ANAND AND GUPTA,

1981). Pesticides interfere with the phys-

iological and metabolic function in ani-

mals, which sometimes result in death

(ARASTA, BAIS AND THAKUR, 1996).

With growing awareness of environ-

mental pollution caused by synthetic

molluscicides (RITCHIE, 1973; SRIVAS-

TAVA AND SINGH, 2001), efforts are being

made to find molluscicides of plant

origin. Being the product of biosynthe-

sis, they are highly toxic and easily

biodegradable in nature (MARSTON AND

HOSTETTMAN 1987; SINGH, SINGH,

MISHRA AND AGARWAL, 1996). Two dis-

eases - schistosomiasis and fascioliasis -

carried by aquatic snails, cause immense

harm to man and his domestic animals

(BALI, SINGH AND SHARMA, 1986;

AGARWAL AND SINGH, 1988). Large pop-

ulations of aquatic snails inhabiting

freshwater bodies cause serious destruc-

tion of freshwater vegetation which ulti-

mately affect the growth of organisms

feeding on them (REINERT, 1972).

Fascioliasis caused by Fasciola hepat-

ica, the large liver-fluke, is common in

sheep, cattle, goats and other herbivo-

rous animals throughout the world. The

freshwater snail L. acuminata is the inter-

mediate host of Fasciola hepatica and Fas-

ciola gigantica (AGARWAL AND SINGH,

1988; SINGH AND AGARWAL 1992; YADAV

AND SINGH 2001, 2002), which cause

great harm to domestic animals. SINGH

AND AGARWAL (1981) reported that 94%

of buffaloes slaughtered in Gorakhpur

of Uttar Pradesh, India were infected by

liver fluke Fasciola gigantica.

The aim of the present study is to

report on the molluscicidal activity of

taraxerol extracted from stem-bark of C.

variegatum , against the harmful snail L.

acuminata. lts effects on biochemical

parameters of target snail L. acuminata

and non-target fish Channa punctatus

were also studied. Freshwater fish

Channa punctatus is a common fish in

India and shares the habitat with the

snails.

MATERIALS AND METHODS

The stem-bark of C. variegatum was

collected locally from the Botanical

garden of D.D.U Gorakhpur University

Gorakhpur, where a voucher specimen

is deposited and identified by Prof. S.K.

Singh (taxonomist), Department of

Botany, D.D.U, Gorakhpur University,

Gorakhpur (U.P) India.

Extraction of active compounds from

stem-bark: Pure taraxerol was isolated

from the stem-bark of C. variegatum by

the method of CHATTERJEE AND BANER-

JEE (1977). The stem-bark of C. variega-

tum was dried in an incubator at 37"C

and dried stem-bark was powdered

with the help of a mechanical device.

The dried powdered stem-bark (2 kg) of

C. variegatum was extracted in Soxhlet

YADAV ET AL.: Toxicity of taraxerol from Codiaeum variegatum on Lymnaea acuminata

apparatus with petrol, for about 70

hours and a little amount of concen-

trated solution was obtained. After

evaporation of the solvent by vacuum

pump, the isolated compound in dried

form was obtained. The organic con-

stituents present in stem-bark ¡.e. taraxe-

rone-2, taraxerol, taraxeryl acetate-4 and

sitosterol were extracted with petrol.

Taraxerol is soluble in organic solvents

such as CHCl3 and CHCl-MeOH. Iden-

tification of the isolated compound was

further confirmed with an authentic

sample of taraxerol (C32H4809), supplied

by Sigma chemical Co. U.S.A. |

The extracted compound was stored

in an airtight desicator and used for tox-

icity experiments. Toxicity experiments

were performed by the method of SINGH

AND AGARWAL (1988). The freshwater

snail L. acuminata were exposed to four

different concentrations of taraxerol, 0.7

mg/L, 1.0 mg/L, 1.4 mg/L and 1.7

mg/L respectively for 24h, 48h, 72h and

96h exposure period. Mortality was

recorded every 24h up to 96h. Ten

snails were kept in 3L de-chlorinated

tap water. Control animals were kept in

similar conditions without any treat-

ment. Each set of experiments was repli-

cated six times. Effective doses (LC

values), upper and lower confidence

limits, slope value, t'ratio, “g' factor and

heterogeneity were calculated by Probit

log analysis method using POLO com-

puter programme of RUSSELL, ROBERT-

SON AND SEVIN (1977).

For environmental toxicity, if any,

the toxic effect of taraxerol was also

studied in mixed populations of target

organism snails and non-target organ-

ism fish. In this experiment, groups of

10 snail L. acuminata and 10 fish C. punc-

tatus were put together in 6L de-chlori-

nated tap water, which were exposed to

2.92 mg/L (LCo 24h of L. acuminata) of

taraxerol for 24h.

Treatment protocol for Dose-Response

relationship: L. acuminata was kept in

glass aquaria containing 6L de-chlori-

nated tap water. Each aquarium con-

tains 30 experimental animals. Snails

and fish were exposed for 24h in the

case Of L. acuminata and 96h for fish C.

punctatus to sub lethal doses 0.76 mg/L

and 1.35 mg/L (40% and 80% of LCso

24h of L. acuminata) of taraxerol. Control

animals were held in similar conditions

without any treatment. After completion

of treatment the test animals were re-

moved from aquaria, and washed with

freshwater. The nervous, hepatopan-

creas and ovotestis tissue of L. acuminata

and muscle, liver and gonadal tissue of

freshwater fish C. punctatus were

quickly dissected out in an ice tray and

used for biochemical analysis.

In order to see the effect oí with-

drawal from treatment, both the experi-

mental animals were exposed for 24h in

the case of L. acuminata and 96h for fish

C. punctatus to sub-lethal doses 1.35

mg/L (80% of 24h LCso of L. acuminata)

of taraxerol, following which the test

animals were transferred to freshwater.

This water was changed every 24h for

the next seven days. After this the test

animals were removed from aquaria,

washed with freshwater and nervous,

hepatopancreas and ovotestis tissue of

L. acuminata and muscle, liver and

gonadal tissues of freshwater fish C.

punctatus were quickly dissected out in

an ice tray and all the above mentioned

biochemical parameters were estimated.

Each experiment was replicated at

least six times and the values have been

expressed as mean +SE of six replicates.

Student's “t' test and analysis of vari-

ance were applied to locate significant

changes (SOKAL AND ROHLF 1973).

BIOCHEMICAL ESTIMATIONS

Protein: Protein levels were esti-

mated according to the method of

LOwRkrY, ROSENBROUGH, FARR, RANNDALL

(1951) using bovine serum albumin as

standard. Homogenates (5 mg/mL,

w/v) were prepared in 10% TCA.

Total free amino acids: Estimation of

total free amino acid was made accord-

ing to the method of SPICES (1957). Ho-

mogenates (10 mg/mL, w/v) were pre-

pared in 95% ethanol, centrifuged at

6000 xg and used for amino acid estima-

tion.

Iberus, 23 (1), 2005

Nucleic acids: Estimation of nucleic

acids (DNA and RNA) was performed,

by methods of SCHNEIDER (1957) using

diphenylamine and orcinol reagents,

respectively. Homogenates (Img/mL,

w/v) were prepared in 5% TCA at 900C,

centrifuged at 5000 xg for 20 min and

supernatant was prepared and used for

estimation. Both DNA and RNA have

been expressed as 1g/mg tissue.

Glycogen: Glycogen was estimated

by the Anthrone method of VAN DER

VIES (1954) as modified by MAHENDRU

AND AGARWAL (1982) for snail L. acumi-

nata. In the present experiment 50 mg of

tissue were homogenised with 5 mL of

cold 5%TCA. The homogenate was fil-

tered and 1.0 mL of filtrate was used for

assay.

Pyruvate: Pyruvate level was measured

according to FRIEDEMANN AND HAUGEN

(1943). Homogenate (50 mg/mL, w/v)

was prepared in 10% TCA. Sodium pyru-

vate was taken as standard.

Lactate: Lactate was estimated

according to BARKER AND SUMMERSON

(1941), modified by HUCKABEE (1961).

Homogenate (50 mg/mL, w/v) was

prepared in 10% cold TCA. Sodium

lactate was taken as standard.

Protease: Protease activity was esti-

mated by the method of MOORE AND

STEIN (1954). Homogenate (50 mg/mL,

w/v) was prepared in cold distilled

water. Optical density was measured at

570 nm. The enzyme activity was

expressed in qmol of tyrosine equiva-

lent/mg protein/h.

Acid and alkaline phosphatase: Activi-

ties of acid and alkaline phosphatase

were measured by the method of

BERGMEYER (1967) and modified by

SINGH AND AGARWAL (1983). Tissue

homogenate (2% w/v) was prepared in

ice cold 0.9% saline and centrifuged at

5000 xg at 0 *C for 15 min. Optical

density was measured at 420 nm against

a blank, prepared simultaneously. The

enzyme activity has been expressed as

amount of p-nitrophenol formed /30

min /mg protein in supernatant.

Lactic dehydrogenase: Lactic dehydro-

genase activity was measured according

to the method of ANONYMOUS (1984).

Homogenates (50 mg/mL, w/v) were

prepared in 1 mL of 0.1 M phosphate

buffer, pH 7.5 for 5 min in an ice bath.

Enzyme activity has been expressed as

nanomol of pyruvate reduced /min/mg

protein.

Succinic dehydrogenase: Succinic

dehydrogenase activity was measured

by the method of ARRIGONI AND SINGER

(1962). Homogenate (50 mg/mL, w/v)

was prepared in 1 mL of 0.5 M potas-

sium phosphate buffer, pH 7.6 for 5 min

in an ice bath. Optical density was mea-

sured at 600 nm. Enzyme activity has

been expressed as pumol dye

reduced /min/mg protein.

Cytochrome oxidase: Cytochrome

oxidase activity was measured accord-

ing to the method of COOPERSTEIN AND

LAZAROW (1951). Homogenates (50

mg/mL, w/v) were prepared in 1 mL of

0.33 M phosphate buffer (pH 7.4) for 5

min in ice bath. Enzyme activity has

been expressed in arbitrary

units /min/mg of proteins.

Acetylcholinesterase: Acetylcholin-

esterase was estimated by the method of

ELLMAN, (COURTNEY, ANDRES AND

FEATHERSTONE (1961) as in 0.1 M phos-

phate buffer in ice bath. Optical density

was measured at 412 nm at 25 *C.

Enzyme activity expressed in mol “SH”

hydrolysed /min/mg protein.

RESULTS

Molluscicidal activity: The toxicity of

taraxerol was also time and dose-depen-

dent for the freshwater snail L. acumi-

nata. There was a significant negative

correlation between LC5o values and all

the exposure periods for 24h or 96h

(Table ID). Thus with an increase in expo-

sure periods of taraxerol the LC50 values

show a significant decrease from 1.69

mg/L (24h);> 1.30 mg/L (48h);> 0.86

mg/L (72h);> to 0.74 mg/L (96h)

respectively, in case of freshwater snail

L. acuminata. (Table ID).

The active (moiety), of taraxerol

which were effective against freshwater

snail L. acuminata, would also cause

death amongst the fish. Consequently,

YADAV ET AL.: Toxicity of taraxerol from Codiaeum variegatum on Lymnaea acuminata

Table 1. Toxicity (LC1o, LCso and LCo0) of taraxerol against freshwater snail Lymnaea acuminata at

different time intervals.

Tabla I. Toxicidad (LCi0 , LCso y LC90) del taraxerol sobre el caracol Lymnaea acuminata a diferentes

intervalos de tiempo.

Exposure Effective dose Limits (mg/L) Slope value y factor “Prato Hetero-geneity

periods LCL UCL

24h 1C10=0.98 0.829 1.089

LC50=1.69 LSD NO 5.397+0.853 0.10 6.33 0.70

1C90=2.92 DIAS,

48h 1C10=0.68 0.494 0.818

1C50=1.30 lo O 4.629+0.649 0.13 7.14 0.50

[C90=2.46 SETS

72h [C10=0.49 0.381 0.586

1C50=0.86 AS US 5.347+0.693 0.07 1.66 0.34

1C90=1.49 1.348 1.759

96h LC10=0.41 0.294 0.514

1C50=0.74 0.640 0.822 5.15240.750 0.08 6.87 0.53

1C90=1.31 1.187 1.542

Botches of ten snails were exposed to four different concentrations of taraxerol.

Concentrations given are the final concentrations (w/v) in aquarium water.

Regression coefficient showed that there was significant (P<0.05) negative correlation between exposure time and different LC values.

LCL = Lower confidence limit; UCL = Upper confidence limit

mixed populations of 10 snails (L. acumi-

nata) and 10 fishes (C. punctatus) were

treated for 24h of the LCoo of taraxerol.

Up to the LCoo0 doses for snail L. acumi-

nata there was no mortality amongst the

freshwater fish C. punctatus. The doses,

which can be used for killing the snails,

are safe for fish. This is supported by

our observations in a mixed population.

Effect on freshwater target snail: Data

of sub-lethal doses of 40% and 80% of

LC5o (0.76 mg/L € 1.35 mg/L) taraxerol

exposure to freshwater snail L. acumi-

nata are given in Tables II and III. Expo-

sure of snails to sub-lethal doses of

taraxerol for 24h caused significant

alterations in nitrogenous and carbohy-

drate metabolism in different body

tissues of the freshwater snail L. acumi-

nata. Total protein and nucleic acids

(DNA and RNA) levels were signifi-

cantly reduced, while free amino acid

level was significantly enhanced after

the exposure to sub-lethal doses in all

the body tissues. Acid and alkaline

phosphatase activities were significantly

reduced, while protease activity was

increased after the exposure.

Total protein levels were reduced to

31%, 37% and 28% of controls after

exposure to sub-lethal doses of 1.35

mg/L of taraxerol respectively in the

nervous, hepatopancreas and ovotestis

tissue of L. acuminata, respectively. The

DNA level was reduced to 41%, 32%

and 28% of controls after treatment with

1.35 mg/L of taraxerol in nervous,

hepatopancreas and ovotestis tissue Of

L. acuminata, respectively. The RNA

level was reduced to 36%, 39% and 28%

of controls after treatment with sub-

lethal doses of 1.35 mg/L of taraxerol

respectively in nervous, hepatopancreas

and ovotestis of L. acuminata. Total free

amino acid levels were induced to 171%,

151% and 174% of controls after treat-

ment with sub-lethal doses of 1.35 mg

/L of taraxerol respectively in nervous,

hepatopancreas and ovotestis of L.

acuminata (Table II).

Activity of acid phosphatase was

inhibited to 75%, 82% and 77% of con-

Iberus, 23 (1), 2005

Table IL. Changes in total protein, total free amino acids, nucleic acid (DNA and RNA) (1g/mg)

level and activity of protease (umol of tyrosine equivalents/mg protein/h) and acid and alkaline

phosphatase (mol substrate hydrolysed/30 min/mg protein) in nervous (NT), hepatopancreas

(HP) and ovotestis (OT) tissues of Lymnaea acuminata after exposure to sub-lethal doses of 40%

and 80% (0.76 mg/L and 1.35 mg/L) of taraxerol after 24h.

Tabla II. Cambios en los niveles de proteínas, aminoácidos libres y ácidos nucléicos (ADN y ARN)

(uglmg), actividad de proteasa (umol de equivalentes de tirosinalmg proteínalh) y de fosfatasas ácida y

alcalina (umol de sustrato hidrolizado/30 min/mg proteína) en tejido nervioso (NT), hepatopáncreas

(AP) y ovotestis (OT) de Lymnaea acuminata tras exposición a dosis subletales de 40% y 80% (0,76

mg/L y 1,35 mg/L) de taraxerol tras 24h.

Pe 5 Conta 40% of LC5o (24h) 80% of LC5o (24H) 7 days of the

od ll (0.76 mg/L) (1.35 mg/L) withdrawal

Protein NT 64.0%0.18(100) 37.1240.24(58) 15.48+0.01(31) 156.80.71* (98)

HP. 66.0+2.61(100) 37.6240.91(57) 24.42+0.81 (37) 136.7+1.00* (97)

01 71.0x0.53(100) 38.34+0.87(54) 19.880.84(28) 131.140.45* (96)

Amino acid NT. 31.3+1.14(100) 51.64+1.15(165) — 53.52+0.28(171) 29.3+0.23* (103)

HP. 26.8+1.51(100) 38.86+0.38(145) 40.40+1.07(151) 23.7+0.38* (105)

0T 34.3+0.68(100) 57.62+0.15(168) 59.68+0.14(174) 21.4+0.10* (104)

DNA NT 76.6%0.29(100) 46.72+0.31(61) 31.40+0.21(41) 136.7+0.69* (96)

HP. 73.8+0.21(100) 49.44+0.33(67) 23.61+0.29(32) 134.4+0.51* (96)

01 81.5+0.89(100) 44.01+0.21(54) 22.82+0.51(28) 137.740.34* (95)

RNA NT 52.31+0.38(100) 26.67+0.27(51) 18.83+0.31(36) 99.9:0.28* (97)

HP. 50.21+0.81(100) 30.12+0.77(60) 19.58+0.22(39) 100.0+0.21* (96)

OT 55.33+0.41(100) 23.79+2.87(43) 15.49+0.58(28) 98.5+0.17* (93)

Protease NT 0.471%0.051(100) 0.664+0.050(141) 0.706*0.051(150) 0.639+0.061* (108)

HP 0.492+0.001(100) 0.6834+0.002(139) 0.713+0.001(145) 0.571+0.012* (94)

OT 0.474+0.007(100) 0.673+0.003(142) 0.725*0.004(153) 0.739%0.154* (106)

Acid phosphatase NT 0.281+0.008(100) 0.258+0.006(92) 0.210+0.004(75) 0.260+0.0123* (92)

HP. 0.275+0.007(100) 0.264+0.008(96) 0.225+0.009(82) 0.276+0.013* (93)

0T 0.280+0.008(100) 0.263+0.003(94) 0.215+0.005(77) 0.267+0.012* (93)

Alkaline phosphatase NT 0.481+0.021(100) 0.384+0.001(80) 0.307+0.003(64) 0.394+0.002* (91)

HP. 0.482+0.004(100) 0.404+0.002(80) 0.307+0.001(68) 0.426+0.003* (92)

OT 0.475+0.008(100) 0.384+0.001(81) 0.308%0.002(65) 0.403+0.004* (92)

* Significant (P<0.05). Student's *F' test was applied between 80% of LCso (24h) and withdrawal groups. Values are mean +SE of six replicates.

Values in parenthesis are percentage changes with control taken as 100%

trols after treatment with sub-lethal doses

of 1.35 mg/L of taraxerol respectively in

nervous, hepatopancreas and ovotestis.

The activity of alkaline phosphatase was

reduced to 64%, 68% and 65% of controls

after treatment with sub-lethal doses of

1.35 mg/L of taraxerol respectively in

nervous, hepatopancreas and ovotestis.

The protease activity was increased to

150%, 145% and 153% of controls after

treatment with sub-lethal doses of 1.35

mg/L of taraxerol respectively in the

nervous, hepatopancreas and ovotestis of

snail L. acuminata (Table ID.

Glycogen and pyruvate levels were

significantly reduced, while lactate level

was significantly enhanced after the

exposure to sub-lethal doses in all the

body tissues. Lactic dehydrogenase

(LDH), cytochrome oxidase and acetyl-

cholinesterase (ACHhE) activities were

significantly reduced, while succinic

dehydrogenase (SDH) activity was

increased after the exposure.

YADAV ET AL.: Toxicity of taraxerol from Codiaeum variegatum on Lymnaea acuminata

Table III. Changes in glycogen (mg/g), pyruvate (umol/g), lactate (mg/g) level and activity of

LDH (umol/mg protein/h), SDH (umol of dye reduced/min/mg protein), cytochrome oxidase

(arbitrary unit/min/mg protein) and ACE (pmol “SH” hydrolysed/min/mg protein) after 24h

exposure to sub-lethal doses of 40% and 80% (0.76 mg/L and 1.35 mg/L) of taraxerol in nervous

(NT), hepatopancreas (HP) and ovotestis (OT) tissues of snail Lymnaea acuminata after 24h.

Tabla II. Cambios en los niveles de glucógeno (mglg), piruvato (ymollg), lactato (mg/g) y actividad de

LDH (umol/mg proteína/h), SDH (pmol de tinción reducidalmin/mg proteína), citocromo oxidasa

(unidad arbitrarialmin/img proteína) y ACHE (umol “SH” hidrolizado/min/mg proteína) tras 24h de

exposición a dosis subletales al 40% y 80% (0,76 mg/L y 1,35 mg/L) de taraxerol en los tejidos nervioso

(NT), hepatopáncreas (HP) y ovotestis (OT) del caracol Lymnaea acuminata tras 24».

a E lissuES Cda 40% of LC5o (24h) 80% of LC5o (24H) 7 doys of the

(0.76 mg/L) (1.35 mg/L) withdrawal

Glycogen NT 6.8%0.02 (100) 3.12+0.03 (46) 2.31+0.01 (34) 7.340.03* (93)

HP. 7.2%0.10 (100) 3.67+0.01 (51) 2.80+0.03 (39) 6.8+0.02* (94)

01 9.1:0.08 (100) 4.09+0.03 (45) 3.00+0.01 (33) 7.3+0.04* (93)

Pyruvate NT 0.578%0.03 (100) 0.196+0.04 (34) 0.167+0.23 (29) 0.635+0.27* (91)

HP. 0.609+0.03 (100) 0.255*0.27 (42) 0.237+0.11 (39) 0.592+0.08* (90)

0T 0.581+0.04 (100) 0.180+0.18(31) 0.156+0.27 (27) 0.628+0.03* (92)

Lactate NT 3.11%0.07 (100) — 5.06+0.15(163) 5.56+0.19 (179) 2.4640.04* (113)

HP. 3.79+0.05 (100) 6.59+0.06 (174) 7.20+0.02 (190) 2.83+0.05* (118)

OT 3.95+0.07 (100) 6.47+0.08 (164) 7.11%-0.06 (180) 2.4640.03* (113)

LDH NT 0.084+0.004 (100) 0.068+0.001 (81) 0.043+0.002 (52 0.065+0.003* (90)

HP. 0.096%0.003 (100) 0.081%0.002 (85) 0.056+0.001 (59 0.068+0.004 ( (91)

OT 0.087%0.004 (100) 0.072+0.001 (83) 0.043%0.004 (5 0.070+0.003* (90)

SDH NT 31.01%0.11 (100) 39.69+0.25 (128) 51.16+0.31* (165) 18.38+0.84* (112)

HP. 43.21+0.13 (100) 53.58+0.29* (124) 65.24*0.27* (151) 15.63*0.18* (108)

) 34,42+0.30* (126) 162) — 20.43+0.20* (111)

5

6

(

(

01 27.32:0.10 (100

(16

(15

44.25+0.34* (16

(55

(02

)

)

0)

)

l

2

) 16.68+0.04*

)

0)

2

l

3

Cytochrome oxidase NT 18.23+0.12 (100) 11.84+0.16 (65) 10.02+0.18 (92)

HP. 716.21+0.14 (100) — 11.34+0.21 (70) 10.05+0.31 13.05+0.12* (90)

01 17.32+0.16 (100) — 11.77%0.15 (68) 10.04+ 0.27 (5 16.18+0.02* (94)

ACHE NT 0.068+0.0008 (100) 0.044+0.0003 (65) 0.028+0.0003 (92) 0.064+0.007* (90)

HP. 0.088%0.0002 (100) 0.060%0.0007 (69) 0.036%0.0002 (41) 0.085+0.002* (93)

OT 0.071:0.0002 (100) 0.046+0.0003 (66) 0.030%0.0004 (43) 0.063*0.007* (90)

Details are as given in Table ll

Glycogen level was reduced to 34%,

39% and 33% of controls after treatment

with sub-lethal doses of 1.35 mg/L

taraxerol respectively in nervous,

hepatopancreas and ovotestis tissues of

L. acuminata. Pyruvate level was

reduced to 29%, 39% and 27% of con-

trols after treatment with sub-lethal

doses of 1.35 mg/L taraxerol respec-

tively in nervous, hepatopancreas and

ovotestis tissues of L. acuminata. Lactate

level was increased to 179%, 190% and

180% of controls after treatment with

sub-lethal doses of 1.35 mg/L of taraxe-

rol respectively in nervous, hepatopan-

creas and ovotestis tissues of snail L.

acuminata (Table II.

Lactic dehydrogenase activity was

reduced up to 52%, 59% and 50% of con-

trols after treatment with sub-lethal

doses of 1.35 mg/L of taraxerol respec-

tively in nervous, hepatopancreas and

ovotestis tissue of snail L. acuminata.

Activity of cytochrome oxidase was

reduced to 55%, 62% and 58% of controls

after treatment with sub-lethal doses of

Iberus, 23 (1), 2005

Table IV. Changes in total protein, total free amino acids, nucleic acid (DNA 8 RNA) (1g/mg)

level and activity of protease (umol of tyrosine equivalents/mg protein/h) and acid and alkaline

phosphatase (umol substrate hydrolysed/30 min/mg protein) in muscle, liver and gonadal tissues

of freshwater fish Channa punctatus after 96h exposure to sub-lethal doses of 40% and 80% (0.76

mg/L and 1.35 mg/L) of taraxerol and 7 days after withdrawal.

Tabla IV. Cambios en los niveles de proteínas, aminoácidos libres, ácidos nucléicos (ADN y ARN)

(ug/mg), actividad de la proteasa (umol of equivalentes de tirosinalmg proteínalh) y fosfatasas ácida y

alcalina (umol sustrato hidrolizado/30 min/mg proteína) en los tejidos muscular, hepático y gonadal del

pez Channa punctatus tras 96h de exposición a dosis subletales al 40% y 80% (0,76 mg/L and 1,35

mg/L) de taraxerol y a los 7 días de retirar el tratamiento.

a his a 40% of LC5o (24h) 80% of LCso (24H) 7 days of the

Ab (0.76 mg/L) (1.35 mg/L) withdrawal

Protein Muscle 171.2+0.75(100) 104.3+0.22(61) 70.1+0.24(41) 162.6+0.68* (95)

Liver 151.0+0.65(100) 128.3+0.21(85) 119.2+0.20(79) 146.4+1.00* (97)

Gonadal 144.6+1.00(100) 112.7+0.12 (78) 76.4+0.07(53) 134.4+0.43* (93)

Amino acid Muscle 38.20+0.23(100) 42.7%0.26(112) 48.5+0.38(127) 42.2+0.20* (110)

Liver 22.6+0.40(100) 26.840.44(119) 31.840.40(141) 24.4+0.30* (108)

Gonadal 31.0+0.70(100) 42.4+0.02(137) 46.1+0.03(149) 35.6+0.08* (115)

DNA Muscle 152.44+0.73(100) 124.8+0.14(82) 94.4+0.14(62) 144.7+0.60* (96)

Liver 149.02*0.70(100) 129.8%0.07(87) 105.9+0.13(71) — 137.440.50* (92)

Gonadal — 143.00%0.73(100) 111.05+0.44(78) 82.94+0.23(58) 128.7+0.30* (90)

RNA Muscle 103.00+0.28(100) 84.41+0.03(82) 69.01%0.04(67) 99.90x0.28* (97)

Liver 100.0+0.29(100) 92.00%0.18(92) 72.00+0.18(72) 95.00%0.21* (95)

Gonadal 106.60+0.61(100) 83.21+0.40(78) 65.02+0.28(61) 98.07+0.16* (92)

Protease Muscle 0.598+0.011(100) 0.825+0.047(138) 0.926+0.015(155) 0.568+0.060* (92)

Liver 0.652+0.016(100) 0.796+0.017(122) 0.920%0.014(141) 0.046+0.010* (99)

Gonadal 0.601+0.016(100) 0.775+0.013(129) 0.949+0.014(158) 0.552+0.154* (92)

Acid phosphatose Muscle 0.302+0.012(100) 0.138+0.010(30) 0.090+0.013(30) 0.277+0.0122*(92)

Liver 0.292+0.014(100) 0.110*0.010(38) 0.078+0.009(27) 0.277+0.009* (95)

Gonadal 0.281+0.015(100) 0.092+0.007(33) 0.061+0.017(22) 0.275*0.010* (98)

Alkaline phosphatase Muscle 0.451%0.010(100) 0.193:<0.006(43) 0.121:0.03(27) 0.410:0.002* (91)

Liver 0.400%0.030(100) 0.148+0.006(37) 0.096+0.006(24) 0.368+0.002* (92)

Gonadal 0.438+0.012(100) 0.210+0.005 (48) 0.135+0.003(31) 0.420+0.003* (96)

*, Significant (P<0.05) when Student's *' test was applied between 80% of LC50 (24h) and withdrawal groups

Details are as given in Table ||

1.35 mg/L of taraxerol respectively in

nervous, hepatopancreas and ovotestis

of L. acuminata. Acetylcholinesterase

activity was reduced to 92%, 41% and

43% of controls after treatment with sub-

lethal doses of 1.35 mg/L of taraxerol

respectively in nervous, hepatopancreas

and ovotestis of snail L. acuminata. The

succinic dehydrogenase (SDH) activity

was increased to 165%, 151% and 162%

of controls after treatment with sub-

lethal doses of 1.35 mg/L of taraxerol

respectively in nervous, hepatopancreas

and ovotestis tissues of freshwater snail

L. acuminata (Table IID.

Effect on freshwater non-target fish:

Higher doses (LCso of snails) have no

apparent toxic effect on non-target fresh-

water fish C. punctatus after 24h exposure.

But exposure of fish to sub-lethal doses

(1.e. 40% and 80% of 24h LCso of snail) 0.76

mg/Land 1.35 mg/L of taraxerol for 96h

caused a significant alteration in nitroge-

nous and carbohydrates metabolism in

YADAV ET AL.: Toxicity of taraxerol from Codiaeum variegatum on Lymnaea acuminata

Table V. Changes in glycogen (mg/g), pyruvate (umol/g), lactate (mg/g) level and activity of LDH

(umol/mg protein/h), SDH (umol of dye reduced/min/mg protein), cytochrome oxidase (arbitrary

unit/min/mg protein) and ACHE (umol “SH” hydrolysed/min/mg protein) in muscle, liver and

gonadal tissues of Channa punctatus after 96h exposure to sub-lethal doses of 40% and 80% (0.76

mg/L and 1.35 mg/L) of taraxerol and 7 days after withdrawal.

Tabla V. Cambios en los niveles de glucógeno (mgle), piruvato (uymolle), lactato (mg/g) y actividad de

LDH (umol/mg proteínalh), SDH (umol de tinción reducidalminímg proteína), citocromo oxidasa

(unidad arbitrarialminimg proteína) y ACHE (pmol “SH? hidrolizado/min/mg proteína) en los tejidos

muscular, hepático y gonadal del pez Channa punctatus tras 96h de exposición a dosis subletales al

40% y 80% (0,76 mg/L and 1,35 mg/L) de taraxerol y a los 7 días de retirar el tratamiento.

E Tissue na 40% of LC5o (24h) 80% of LCso (24H) 7 days of the

(0.76 mg/L) (1.35 mg/l) withdrawal

Glycogen Muscle 2.20+0.001(100) 1.89+0.004(86) 1.380.04(63) 1.9140.02* (87)

Liver 2.980.002(100) 2.17%0.001(73) 2.62+0.03(68) 2.47+0.04* (83)

Gonadal 3.00+0.01(100) 2.61+0.02(87) 2.13+0.04(71) 2.73+0.04* (91)

Pyruvate Muscle 3.416%0.018(100) 1.702*0.013(50) 1.097%0.024(32) 3.050+0.014* (90)

Liver 4.076+0.018(100) 2.446+0.035(60) 1.548+0.007(38) 3.788+0.031* (93)

Gonadal 2.993+0.036(100) 1.705+0.014(57) 0.921*0.020(31) 2.721:+0.016* (91)

Lactate Muscle 3.816+0.018(100) 4.649+0.090(122) 6.846+0.060(179) 4.221%0.080* (111)

Liver 2.323+0.020(100) 3.205+0.020(138) 3.750%0.074 (172) 2.500%0.069* (112)

Gonadal 3.816+0.083(100) 4.502+0.088(118) 6.181+0.092(162) 4.159+0.043* (109)

LDH Muscle 425.3:0.88(100) 374.2:0.81(88) 259.4:0.83(61) 382.5+0.81* (90)

Liver 555.0+1.0(100) 521.7+0.80(94) 394.0+0.81(71) 516.1%0.76* (93)

Gonadal 467.1+0.84(100) 420.3*+0.70(90) 350.3+0.81(75) 448.4+0.78* (93)

SDH Muscle 59.4+0.21(100) 72.4:0.20(122) 81.3:0.27(137) 64.7+0.20* (109)

Liver 62.2+0.20(100) 712.240.20(116) 85.9+0.19(138) 69.30.20* (111)

Gonadal 64.4+0.26(100) 85.8+0.16(110) 95.9+0.10(149) 68.2+0.23* (106)

Cytochrome oxidase Muscle 28.9140.21(100) 24.86%0.20(86) 17.92*0.24(62) 27.17+0.30* (94)

Liver 23.1240.05(100) — 17.10%0.20(74) 15.95:0.24(69) 21.03+0.022* (91)

Gonadal 33.20+0.05(100) 32.20+0.10(87) 24.28:+0.13(73) 30.54+0.17* (92)

ACHE Muscle 0.091<0.0010(100) 0.045+0.0004(50) 0.034+0.0006 (38) 0.085+0.0002* (94)

liver 0.097+0.0009(100) 0.053+0.0004(55) 0.034+0.0003(36) 0.082+0.0001* (92)

Gonadal 0.088+0.0020(100) 0.043<0.0006(49) 0.024+0.0005(27) 0.079+0.0004* (90)

+, Significant (P<0.05) when Student's “Y test was applied between 80% of LC50 (24h) and withdrawal groups

Details are as given in Table ||

different body tissues of fish C. punctatus

(Tables IV and V).

Total protein and nucleic acids

(DNA and RNA) levels were signifi-

cantly reduced, while free amino acid

level was significantly enhanced after

the exposure to sub-lethal doses in all

the studied body tissues. Acid and alka-

line phosphatase activities were signifi-

cantly reduced, while protease activity

was increased after the exposure. Total

protein levels were reduced to 41%, 79%

and 53%; DNA level was reduced to

62%, 71% and 58% and RNA level was

reduced to 67%, 72% and 61% in muscle,

liver and gonadal tissue of freshwater

fish C. punctatus. Total free amino acid

levels were induced to 127%, 141% and

149% of controls after 96h treatment

with 1.35 mg/L of LCso of taraxerol in

muscle, liver and gonadal tissues,

respectively (Table IV).

Activity of acid phosphatase was

inhibited to 30%, 27% and 22% and

Iberus, 23 (1), 2005

activity of alkaline phosphatase was

reduced to 27%, 24% and 31% but Pro-

tease activity was increased to 155%,

141% and 158% of controls after 96h

treatment with 1.35 mg/L of taraxerol in

muscle, liver and gonadal tissues of

freshwater non-target fish C. punctatus,

respectively (Table IV).

Glycogen and pyruvate levels were

significantly reduced, while lactate level

was significantly enhanced after the

exposure to sub-lethal doses in the

studied body tissues. Lactic dehydroge-

nase (LDH), cytochrome oxidase and

acetylcholinesterase (ACHE) activities

were significantly reduced, while suc-

cinic dehydrogenase (SDH) activity was

increased after the exposure. Glycogen

level was reduced to 63%, 68% and 71%

and pyruvate level was reduced to 32%,

38% and 31% in muscle, liver and

gonadal tissue of fish. Lactate level was

increased to 179%, 172% and 162% of

controls after 96h treatment with 1.35

mg/L taraxerol in muscle, liver and

gonadal tissues of fish c. punctatus,

respectively (Table V).

Lactic dehydrogenase (LDH) activity

was reduced to 61%, 71% and 75% and

activity of cytochrome oxidase was

reduced to 62%, 69% and 73% and

acetylcholinesterase (ACHE) activity was

reduced to 38%, 36% and 27% in muscle,

liver and gonadal tissue of fish C. puncta-

tus, respectively. Succinic dehydrogenase

(SDH) activity was increased to 137%,

138% and 149% of controls after 96h

treatment with 1.35 mg/L LC5o taraxerol

in muscle, liver and gonadal tissues of

fish C. punctatus, respectively (Table V).

DISCUSSION

Itis clear from the results section that

the extracted compound taraxerol from

the stem-bark of Codiaeum variegatum has

potent molluscicidal activity against the

freshwater target snail L. acuminata.

More important is the fact that the

extracted compound is much more toxic

than synthetic pesticides. The present

study demonstrated that taraxerol has

higher molluscicidal activity than any of

10

the prevalent synthetic molluscicides

like carbamate, organophosphate and

synthetic pyrethroids. Thus, the 24h

LC5so of mexacarbamate (3.5 ppm),

aldicarb (30.00 ppm), farmothion (27.00

ppm), cypermethrin (2.5 ppm), perme-

thrin (0.82 ppm) and fenvalerate (2.5

ppm) against L. acuminata (SINGH AND

AGARWAL, 1981; SINGH AND AGARWAL,

1986; SINGH AND AGARWAL, 1988; SINGH

AND AGARWAL, 1991) is higher than that

of taraxerol (0.183 ppm), which is about

65 times stronger than the standard mol-

luscicide niclosamide (LCso 11.8 ppm)

SAHAY, SINGH AND AGARWAL (1991).

Statistical analysis of the data on tox-

icity brings out several important

points. The X? test for goodness of fit

(Heterogeneity) demonstrated that the

mortality counts were not found to be

significantly heterogeneous and other

variables (e.g. resistance) do not signifi-

cantly affect the LC5o values, as these

were found to lie within the 95% confi-

dence limits. The slope is thus, an index

of the susceptibility of the target animal

to the extract used. A steep slope is also

indicative of rapid absorption and onset

of effects. Since the LC5o of taraxerol is

within the 95% confidence limits, it is

obvious that in replicate tests of random

samples, the concentration response

lines would fall in the same range

(RAND AND PETROCELLI, 1988).

The depletion of protein fraction in

different tissues of snail and fish may

have been due to their degradation and

possible utilization of degraded prod-

ucts for metabolic purposes. Mom-

MENSEN AND WALSH (1992) reported that

proteins are mainly involved in the

architecture of the cell, which is the main

source of nitrogenous metabolism, and

during chronic periods of stress they are

also a source of energy. Increment in free

amino acids level was the result of

breakdown of protein for energy require-

ments and impaired incorporation of

amino acids in protein synthesis. Inhibi-

tions of DNA synthesis might affect both

protein as well as amino acid levels by

decreasing the level of RNA in protein

synthesis machinery (NORDENKJOLD,

SODERHALL AND MOLDEUS, 1979).

YADAV ET AL.: Toxicity of taraxerol from Codiaeum variegatum on Lymnaea acuminata

The increase in the protease activity

corroborates the enhancement in the FAA

(Free amino acids) level in tissues, the

formation of which might be the result of

protein hydrolysis in the tissues suggest-

ing stimulation during toxic stress.

Similar trend of results on protease activ-

ity were also reported by several workers

in different animals (IT. mossambica

(Peters), P. globosa (Swaimson) including

mammals (KABEER, SAHIB, SIVA PRASAD

AND SAMBASIVA RAO, 1984). SINGH AND

AGARWAL (1992) reported that several

euphorbious plants significantly reduced

the alkaline and acid phosphatase activ-

ity in nervous tissue of L. acuminata so the

reduction in protein level may be due to

the inhibition of alkaline phosphatase

activity, as it plays an important role in

protein synthesis (PILO, ASNANI AND

SHAH, 1972) and other secretory activities

(IBRAHIM, HIGAZI AND DEMIAN, 1974).

Carbohydrates reserves were

depleted to meet energy demand, thus

depletion of glycogen may be due to

direct utilization for energy generation, a

demand caused by active moiety-

induced hypoxia. The glycogenolysis

seems to be the result of increased secre-

tion of catecholamine due to stress.

Decrease in pyruvate level is due to

higher energy demand during exposure,

which suggests the possibility of a shift

towards anaerobic dependence due to a

remarkable drop in the amount of

oxygen. The increase in lactate also sug-

gests a shift towards anaerobiosis as a

consequence of hypoxia leading to respi-

ratory distress (SIvVA PRASADA RAO, 1980).

Lactic dehydrogenase (LDH) cat-

alyzes the inter-conversions of lactic

acid and pyruvic acid during anaerobic

conditions. Inhibition of lactic dehydro-

BIBLIOGRAPHY

AGARWAL, R. A., AND SINGH, D. K., 1988. Harm-

ful gastropods and their control. Acta Hy-

drochímica et Hydrobiologica 16: 113-138.

ANONYMOUS, 1984. Sigma diagnostics TM: Lac-

tic dehydrogenase (quantitative, colorimet-

ric determination in serum, urine and cere-

brospinal fluid) at 400-450 nm. Procedure

no. 500. St. Louis, U.S. A.

genase and cytochrome oxidase activity

indicates that taraxerol significantly

inhibits aerobic, as well as anaerobic

metabolism in exposed animals (EVERSE

AND KALPAN, 1973). Succinic dehydro-

genase (SDH) is one of the active regula-

tory enzymes of the TCA cycle. Inhibi-

tion in cytochrome oxidase activity by

taraxerol moieties supports that

Euphorbiales show a profound impact

on the oxidative metabolism.

Withdrawal experiments were per-

formed to see whether biochemical alter-

ation caused by taraxerol moiety would

return to normal, if the treatment were

discontinued. There was nearly com-

plete recovery of total protein, total free

amino acid, lactate, nucleic acid (DNA

and RNA), pyruvate level and in the

activity of cytochrome oxidase, succinic

dehydrogenase, protease, lactic dehydro-

genase, acetylcholinesterase and acid

and alkaline phosphatase but only a

partial recovery of glycogen level in the

different body tissues of freshwater snail

EL. acuminata and fish C. punctatus.

In conclusion, it is believed that

extracted compound of selected plants

may be used as a potent source of mol-

luscicides for addition; plant products

are less expensive, easily available,

easily soluble in water and safer for

non-target animals than synthetic mol-

luscicides.

ACKNOWLEDGEMENTS

One of the Authors (Ram P. Yadav) is

thankful to the Department of Environ-

ment, Ministry of Environment and Forest,

Govt. of India New Delhi (F-14/35/96-

MAB/RE) for financial Assistance.

ARASTA, T., BAIs, V. S. AND THAKUR, P., 1996.

Effect of Nuvan on some biological parame-

ters of Indian catfish Mystus vittatus. Journal

of Environmental Biology 17 (2): 167-169.

ARRIGONI, O., AND SINGER, T. P., 1962. Limita-

tion of the phenozine methosulphate assay

for succinic and related dehydrogenase. Na-

ture , 193: 1256-1258.

Iberus, 23 (1), 2005

BALI, H. S., SINGH, S. AND SHARMA, $S., 1986. The

distribution and ecology of vectors snails of

Punjab. Indian Journal Ecology 13: 31-37.

BARKER, S. B., AND SUMMERSON, W. H., 1941. The

colorimetric determination of lactic acid in bi-

ological materials. Journal Biological Chem-

istry. 138: 535-5427.

BERGMEYER, U. H., 1967. Method of Enzymatic

Analysis. Academic Press, New York. pp.,

DO:

CHATTERJEE, A., AND BANERJEE, A., 1977. Cro-

tocaudin; A rearranged labdane type nor-

diditerpines from Croton caudatus geisel.

Tetrahedran 33: 2407-2414.

COOPERSTEIN, S. J., AND LAZAROW, A., 1951.

Microspectrophotometric method for the de-

termination of cytochrome oxidase. Journal

of Biological Chemistry 189: 665-670.

ELLMAN, G. L., COURTNEY, K. D. ANDRES, V. J.

R. AND FEATHERSTONE, |. M., 1961. A new

and rapid colorimetric determination of

acetylcholinesterase activity. Biochemistry

Pharmacology 7: 88-95.

EVERSE, T., AND KAPLAN, N. O., 1973. Lactate

dehydrogenase: Structure and function. In

“Advances in Enzymology” (A. Meister, Ed.

) John Wiley, New York Vol. 27: 61-133.

FRIEDEMANN, T. E., AND HAUGEN, G. F., 1943.

Pyruvic acid. [. Collection of blood for the de-

termination of pyruvic acid and lactic acid.

Journal of Biological Chemistry 144: 67 - 77.

GOPAL, K., KHANNA, R. N. ANAND, M. AND

GUPTA, G. S. D., 1981. The acute toxicity of

endosulfan to freshwater organisms. Toxi-

cology Letter 7: 453-456.

HUCKABEE, W. E., 1961. In: Hawk's Physiolog-

ical Chemistry, 14th edition, (Ed. Oser, B. L.

), New Delhi, Tata McGraw-Hill, pp 1103.

IBRAHIM, A. M., HIGAZI, M. G. AND DEMIAN, S.,

1974. Histochemical localisation of alkaline

phosphatase activity in alimentary tract of the

snail Marisa cornuarietes (L.). Bulletin Zoolog-

ical Society Egypt, 26: 94-105.

KABEER, A., SAHIB, I. SIVA PRASAD, R. K. SAM-

BASIVA, R. K. R. AND RAMANA RAO, K. V.,

1984. Sub-lethal toxicity of malathion on the

protease and free amino acid composition

of the liver of the teleost Tilapia mossambica

(Peters). Toxicology Letter 20: 59-62.

LowkrY, O. H., ROSENBROUGH, N. . J., FARR A.

L AND RANDALL R. J, 1951. Protein mea-

surement with folin phenol reagent. Journal

of Biological Chemistry 193: 265-275.

MAHENDRU, V. K., AND AGARWAL, R. A., 1982.

Changes induced by phorate in the carbo-

hydrate metabolism of snail Lymnaea acumi-

nata. Pesticide Science 13: 611-616.

MOORE, S., AND STEIN, W. H., 1954. A modified

ninhydrine reagent for the photometric de-

termination of amino acids and related com-

pounds. Journal of Biological Chemistry 211:

907-913.

12

MOMMENSEN, T. P., AND WALSH, P. J., 1992.

Biochemical and metal perspectives on ni-

trogen in fishes. Experimentia, 48: 583-593.

MARSTON, AÁ., AND HOSTETTMAN, K., 1987. An-

tifungal, molluscicidal and cytotoxic com-

pounds from plants used in traditional med-

icine. Biologically active natural products ed

by Hostettmann K, Lea PJ Clarendon Press,

Oxford, 65-83.

NORDENSKJOLD, M., SODERHALL, J. AND

MOLDEUS, P., 1979. Studies on DNA strand

breaks induced in human fibroblasts by

chemical mutagens and carcinogens. Muta-

tion Research 63: 393-400.

PiLO, B., ASNANI, M. V. AND SHAH, R. V., 1972.

Studies on wound healing and repair in pi-

geon III. Histochemical studies on acid al-

kaline phosphatase activity during the

process. Journal of Animal Physiology 19: 205-

212:

RAND, G. M. AND PETROCELLI, S. R., 1988.

Fundamentals of aquatic toxicology. Hemi-

sphere Publishing Corporation, New York

pp1129:

REINERT, J. A., 1972. Snail control under aquatic

conditions. Proceeding of Florida State Horti-

culture Society 85: 380-384.

RITCHIE, L. S., 1973. Chemical control of snails.

In Epidemiology and control of schistoso-

miasis. ed by Ansari N, Darger S B, pp 458-

392:

RUSSELL, R. M., ROBERTSON, J. L. AND SEVIN, N.

E., 1977. A new computer Programme for

probit log analysis. Bulletin Entomological So-

ciety America 23: 209-213.

SAHAY, N., SINGH, D. K. AND AGARWAL, R. A.,

1991. Synergistic effect of pipernoyl butaox-

ide on the toxicity of synthetic pyrethroids

in the snail Lymnaea acuminata. Journal of Med-

ical and Applied Malacology 3: 107-111.

SCHNEIDER, W. C., 1957. Determination of nu-

cleic acid in tissue by pentose analysis In

Calowick S. P. and Kaplon N. O. (Eds. ) Aca-

demic press New York pp680.

SINGH, A., SINGH, D. K. MISHRA, T. N. AND

AGARWAL, R. A., 1996. Molluscicides of plant

origin. Biological Agriculture and Horticulture

1352057292

SINGH, A., AND AGARWAL, RR. A., 1992. Toxic-

ity of the latex of Euphorbiales: Effect on

acid and alkaline phosphatases of the snail

Lymnaea acuminata. Biological Agriculture and

Horticulture 8: 211-219.

SINGH, A., AND AGARWAL, R. A., 1988. Possi-

bility of using latex of euphorbiales for snail

control. The Science of the Total Environment

77:231-236.

SINGH, D. K., AND AGARWAL, R. A., 1983. In

vivo and in vitro studies on synergism with

anticholinestrerase pesticides in the snail

Lymnaea acuminata. Archives Environmental

Contamination and Toxicology 12: 483-487.

YADAV ET AL.: Toxicity of taraxerol from Codiaeum variegatum on Lymnaea acuminata

SINGH, D. K., AND AGARWAL, R. A., 1986. Piper-

onyl butaoxide synergism with two synthetic

pyrethroids against Lymnaea acuminata.

Chemosphere 15: 493-498.

SINGH, D. K., AND AGARWAL, R. A., 1987. Effect

of the synthetic pyrethroids permethrin on

the snail Lymnaea acuminata. The Science of

the total Environment 67: 263-267.

SINGH, D. K., AND AGARWAL, R. A., 1984. Cor-

relation of the anti- cholinesterase and mol-

luscicidal activity of the latex of Euphorbia

royleana Bioss. on Lymnaea acuminata. Journal

of Natural Product 47: 702-705.

SINGH, O., AND AGARWAL, R. A., 1981. Toxic-

ity of certain pesticides to two economic

species of snails in Northern India. Journal of

Economic Entomology 74: 568-571.

SIVA PRASADA RAO, K., 1980. Studies on some

aspects of metabolic changes with emphasis

on carbohydrate utility in the cell-free systems

of the teleost. T. mossambica (Peters) under

methyl parathion exposure. Ph. D. thesis, SV

University Tirupati, India.

SOKAL, R. R., AND ROHLE, F. J., 1973. Introduc-

tion of Biostatic WH Freeman and Company

San Francisco pp368.

SPICES, J. R., 1957. Colorimetric procedure for

amino acids. In Methods in enzymology, ed by

Calowick SP, Kalpan NO, Academic press

New York. 468 pp.

SRIVASTAVA, V. K., AND SINGH, A., 2001. Possi-

bility of using snails as bioindicator of pes-

ticidal pollution. Iberus 19 (1): 1-5.

VAN DER VIES, J., 1954. Two methods for the de-

termination of glycogen in liver. Journal of Bio-

chemistry 57: 410-416.

YADAV, R. P., AND SINGH, A., 2001. Environ-

mentally safe molluscicides from two com-

mon euphorbiales. Iberus, 19 (1): 65-73.

YADAV, R. P., AND SINGH, A., 2002. Toxic effects

of latex of Croton tiglium on Lymnaea acumi-

nata and Channa punctatus. Iberus 20 (2): 33-

44.

O Sociedad Española de Malacología —__——— Iberus, 23 (1): 15-24, 2005

Un avance sobre la composición y microestructura de la

concha de lberus gualtierianus morfotipo gualtierianus

(Linnaeus, 1758) (Gastropoda: Helicidae)

Notes on the composition and microstructure of the shell of /berus

gualtierianus morphotype gualtierianus (Linmaeus, 1758)

(Gastropoda: Helicidae)

Rocío MÁRQUEZ*, José Ramón ARRÉBOLA** y Rafael DELGADO*

Recibido el 20-V-2004. Aceptado el 5-VII-2004

RESUMEN

Se ha estudiado la concha de Iberus gualtierianus morfotipo gualtierianus (Linnaeus, 1758)

desde el punto de vista de su composición mineralógica (difracción de rayos X), composi-

ción química (fluorescencia de rayos X, microanálisis de rayos X y calcimetría) y microes-

tructura (microscopía electrónica de barrido, MEB). El mineral presente es aragonito (CaCOs)

(sobre 99%), con trazas de calcita. Los principales elementos químicos detectados, aparte

de Ca, € y O, son Si, Mg, Na y Sr, probablemente como sustituciones en la red cristalina

y/o contaminaciones. Los contenidos de CaCO3 están entre 97% y 99%. La concha tiene

un espesor de unos 300 pm, con microestructura en capas superpuestas. Dentro de cada

una de ellas, el tipo de microestructura se califica como laminar cruzada simple, compuesta

de láminas de tercer, segundo y primer orden y con orientaciones distintas de los cristales

de aragonito. Rasgos microestructurales destacables son el distinto espesor de las capas,

dependiendo de su situación bajo las costillas o estriaciones de la concha, y que el número

de capas disminuye en la última media vuelta de espira. La mayoría de estos caracteres son

comunes a otros moluscos y en especial a los gasterópodos terrestres, si bien hasta el pre-

sente no habían sido establecidos en /. gualtierianus morfotipo gualtierianus.

SUMMARY

The mineralogical composition of the shell of Iberus gualtierianus morphotype gualtierianus

(Linnaeus, 1758) was studied (X-ray diffraction), as was the chemical composition (X-ray

fluorescence, X-ray microanalysis and calcimetry) and the microstructure (scanning elec-

tron microscopy, SEM). The mineral present is aragonite (CaCO3) (around 99%) with tra-

ces of calcite. The main chemical elements detected, apart from Ca, C and O, are Si, Mg,

Na and Sr, probably substituting Ca in the crystalline structure and/or as impurities. The

CaCOz3 contents are between 97% and 99%. Shell thickness is around 300 pm with supe-

rimposed layers. In each of these the microstructure is classified as simple crossed lame-

llar, composed of third, second and first order lamellae with differing orientation of the

aragonite crystals. Noteworthy microstructural features are the different thicknesses of the

layers, depending on their position below the ribs or striations of the shell and the fact that

* Departamento de Edafología y Química Agrícola, Facultad de Farmacia, Universidad de Granada, Campus de

Cartuja s/n, 18071, Granada, España.

** Departamento de Fisiología y Zoología, Facultad de Biología, Universidad de Sevilla, Avda. Reina Mercedes,

6, 41012, Sevilla, España.

15

Iberus, 23 (1), 2005

the number of layers decreases in the last half turn of the whorl. Most of these characteris-

tics are common to other molluscs, especially to the terrestrial gastropods, although, until

now, these had not been identified in Iberus gualtierianus morphotype gualtierianus.

PALABRAS CLAVE: /berus gualtierianus morfotipo gualtierianus, concha, mineralogía, composición química,

microestructura, MEB.

KEY WORDS: /berus gualtierianus morphotype gualtierianus, shell, mineralogy, chemical composition, micros-

tructure, SEM.

INTRODUCCIÓN

La mayoría de los moluscos se carac-

terizan por presentar una concha, verda-

dero esqueleto externo que ellos mismos

segregan, que posee distintas funciones y

que está compuesta en un 95-99% por car-

bonato, esencialmente cálcico (CaCO»), y

materia orgánica, que representa entre el

5% y el 0,1% restante (HARE Y ABELSON,

1965).

El carbonato cálcico de las conchas

de los moluscos precipita como calcita o

aragonito, siendo la última especie

mineral el polimorfo más frecuente

(LOWENSTAM Y WEINER, 1989). Aún así,

algunos moluscos presentan simultáne-

amente los dos polimorfos en sus

conchas, bien como constituyentes de

una misma capa (MUTVEIL, DAUPHIN Y

Curr, 1985), o bien localizados en capas

diferenciadas (SABATIER, 1953; DAUPHIN,

CUIE, MUTVEI Y DENIS, 1989; LOWENSTAM

Y WEINER, 1989). Otras formas de CaCOz3

han sido también reconocidas en la com-

posición mineral de las conchas de los

moluscos, pero nunca como constitu-

yentes principales. Así por ejemplo, la

vaterita ha sido identificada en zonas

regeneradas de las conchas tras sufrir

éstas algún daño estructural (WILBURG,

1964; TAYLOR, KENNEDY Y HALL, 1969;

LOWENSTAM Y WEINER, 1989) o en los

márgenes de las conchas recientemente

mineralizados (LOWENSTAM Y WEINER,

1989; HAsseE, EHRENBERG, MARXEN,

BECKER Y EPPLE, 2000), aunque siempre

en cantidades inferiores al 1,5%.

En la concha de los moluscos apare-

cen también ciertos elementos traza,

cuyo contenido concreto puede dar

información acerca de las condiciones

ambientales y paleoambientales del

16

medio en el que se produjo la precipita-

ción del mineral. El análisis isotópico

(810 y 8C) de las conchas ha aportado,

igualmente, información de esa natura-

leza (KRANTZ, WILLIAMS Y JONES, 1987).

La microestructura de la concha de

los moluscos ha sido descrita y analizada

por muchos autores (MaAckay, 1952;

CARTER, 1980; MUTVEI ET AL., 1985; HE-

DEGAARD, 1997; CHATEIGNER, HEDEGA-

ARD Y WENK, 2000). BSGGILD (1930) rea-

lizó la primera clasificación de la micro-

estructura mediante descripciones con

microscopía de luz transmitida. Gracias

a la introducción de la microscopía elec-

trónica en estos estudios (GRÉGOIRE, Du-

CHATEAU Y FLORKIN, 1949), y principal-

mente desde la utilización del microsco-

pio electrónico de barrido (MEB) a

finales de la década de los sesenta, la mi-

croestructura de las conchas se ha defi-

nido más a partir de su micromorfología

que a partir de su mineralogía o cristalo-

grafía Óptica. Esta situación ha hecho

que las clasificaciones iniciales hayan

sido redefinidas (KOBAYASHI, 1964, 1971;

MACCLINTOCK, 1967; TAYLOR ET AL.,

1969, TAYLOR, KENNEDY Y HALL, 1973;

GRÉGOIRE, 1972), realizándose importan-

tes revisiones de la nomenclatura des-

criptiva aplicada a la microestructura y

ultraestructura (WILBUR Y SALEUDDIN,

1983; WATABE, 1984; LOWENSTAM Y WEI-

_NER, 1989). Cabe destacar, en este sen-

tido, los trabajos de CARTER Y CLARK

(1985) y CARTER (1990), en los que se in-

tegra la nomenclatura de la mineralogía

y la microestructura de la mayoría de los

grupos de invertebrados y vertebrados,

con la intención de poder hacer compa-

raciones entre distintos organismos.

MÁRQUEZ ET AL.: Composición y microestructura de la concha de /berus gualtierianus

El estudio de la microestructura de

la concha de los moluscos también se ha

empleado para el análisis de los proce-

sos y mecanismos que dan lugar a la

precipitación del carbonato cálcico y su

posterior organización ultraestructural

(WATABE Y WILBUR, 1961; WISE Y Hay,

1968).

Entre las distintas aplicaciones que

tiene el análisis de la microestructura de

las conchas se encuentra su utilidad en

estudios taxonómicos, filogenéticos y

evolutivos (MACKaAY, 1952; WIsE, 1970;

TAYLOR, 1973; PoPOv Y BARSKOV, 1978;

CARTER Y CLARK, 1985; SCHNEIDER Y

CARTER, 2001).

En este trabajo se estudia la compo-

sición mineralógica y química y la

microestructura de la concha de Iberus

gualtierianus morfotipo gualtierianus

(Linnaeus, 1758), una de las varias

formas conquiliológicas conocidas en

esta especie.

Iberus gualtierianus morfotipo gualtie-

rianus es un endemismo andaluz cono-

cido popularmente como “chapa”, que

vive en tres enclaves muy localizados:

Sierra Elvira en la provincia de Granada,

Sierra de Jaén en la de Jaén y Sierra de

Gádor en la provincia de Almería.

Habita en suelos xéricos, sobre calizas y

dolomías con modelado de tipo kárstico

y una vegetación correspondiente a

etapas de regresión del bosque climá-

cico. En este ambiente, es capaz de colo-

nizar zonas semidesérticas con poco

desarrollo de suelo y escasas precipita-

ciones anuales. Debido a las presiones de

diversa índole que le afectan desde hace

varias décadas, y en especial las relacio-

nadas con su interés gastronómico

(ARRÉBOLA, 2002), ha sido propuesta por

la Sociedad Española de Malacología

para su incorporación al Catálogo Nacio-

nal de Especies Amenazadas.

MATERIAL Y MÉTODOS

Cuatro conchas (A, B, C y D) de

Iberus gualtierianus morfotipo gualtieria-

nus fueron recogidas en Sierra Elvira

(UTM: VG32), al noroeste de la ciudad

de Granada.

La composición mineralógica de los

cuatro ejemplares ha sido analizada

mediante difracción de rayos X (XRD), con

un difractómetro Rigaku-Miniflex Ca 2005,

equipado con un filtro de Ni, sistema de

discriminación de impulsos y contador de

gases, según describe GÁMIZ (1987). La

composición química de los ejemplares A

y B se ha conocido mediante fluorescen-

cia de rayos X (XRF) (SORIANO, 1994) y

microanálisis de rayos X de energía dis-

persiva (EDR) (MÁRQUEZ, 2003). La

primera técnica se ha realizado con un

espectrómetro secuencial de longitud de

onda dispersiva, equipado con un ánodo

de Rh y un generador de rayos X de 4 kW

de potencia, Philips Magix Pro (PW-2440).

Por su parte, el análisis EDR se ha llevado

a cabo con un espectrómetro de rayos X

de energía dispersiva, Róntec, 288, M-Serie,

EDWIN, acoplado a un equipo de MEB

(se describirá posteriormente), que permite

analizar muestras de superficie rugosa.

Para este análisis las muestras fueron meta-

lizadas con grafito. Finalmente, los ejem-

plares € y D han sido utilizados para ana-

lizar el contenido de carbonatos (CaCO%3

equivalente) con el calcímetro de Bernard.

Para el análisis de la microestructura

de la concha de I. gualtierianus morfo-

tipo gualtierianus se han seleccionado,

del ejemplar A, muestras de las zonas

de crecimiento más reciente de las tres

últimas medias vueltas de espira de la

concha, ya que según LÓPEZ-ALCÁN-

TARA, RIVAS, ALONSO E IBÁÑEZ (1983)

cada media vuelta de espira se corres-

ponde con un estadio de crecimiento del

organismo. Se han observado fracturas

frescas de la concha, que facilitan la

observación de la microestructura en el

MEB (HEDEGAARD, LINDBERG Y BANDEL,

1997), en secciones paralelas y perpendi-

culares a las líneas de crecimiento. Los

fragmentos de concha fueron pegados

directamente sobre el portamuestras de

aluminio mediante pegamento de plata

coloidal. Posteriormente fueron metali-

zados con grafito para facilitar la obser-

vación y permitir el análisis EDR. La

observación se ha realizado en un

microscopio electrónico de barrido

Hitachi, modelo S-501 (voltaje de acele-

ración de 25 kV; imágenes de electrones

17

Iberus, 23 (1), 2005

Tabla I. Análisis por fluorescencia de rayos X (XRE, ejemplares A y B) y calcimetría (calcímetro de

Bernard, ejemplares C y D) de las conchas de /berus gualtierianus morfotipo gualtierianys.

Table I. X-ray fluorescence analysis (XRE specimens A and B) and carbonate content (Bernard calcime-

ter, specimens C and D) of the shells of lberus gualtierianus morphotype gualtierianus.

Resultados XRF

Elementos mayores como óxidos (%)

Al03 C00 Fez03 K20 MgO Noz20 P205 SiO2

Ei. A 0,05 55,72 0,06 0,07 0,12 0,08 0,02 0,25

Ei. B 0,06 55,44 0,02 0,07 0,08 0,08 0,01 0,30

10,9 624,7 10,5 43,13 99,50 99,40 98,0

21,1 579,2 10,9 43,23 99,29 98/90: 98,20-197 5299772

Colcimetría

Elementos menores (ppm) LOI'Y% Suma CaCOs CoCO3 equiva-

Sr dr (0%) (4) (0% (40 | lente (%)0

EC ED

(a) Pérdida de peso por calcinación (1000 *C), (b) Calculado a partir de los porcentajes de CO; % CaCOz = % CaO x Pm CaCO3/Pm Cad

(c) Calculado a partir de los porcentajes de LO; % CaCOz = % LOI x Pm CaCO3/Pm CO»; (Pm, peso molecular); (d) Calcímetro de Bernard

secundarios) con el programa de digita-

lización de imágenes Scan Vision.

RESULTADOS Y DISCUSIÓN

El análisis mineralógico pone de

manifiesto que las conchas de 1. gualtieria-

nus morfotipo gualtierianus están com-

puestas, mayoritariamente, por aragonito

(ejemplar A, 99,8%; ejemplar B, 99,4%;

ejemplar C, 99,5% y ejemplar D, 98,7%),

conteniendo cantidades traza de calcita

(ejemplar A, 0,2%; ejemplar B, 0,6%;

ejemplar C, 0,5%; ejemplar D, 1,3%).

El patrón de difracción de los dia-

gramas corresponde al del aragonito en

la carta JCPDS n* 5-453, con una

pequeña reflexión a 0,303 nm correspon-

diente a la calcita (carta JCPDS n* 5-586)

(JCPDS, 1974).

En este resultado, I. gualtierianus mor-

fotipo gualtierianus se asemeja a la mayoría

de los moluscos, cuya concha es aragoní-

tica (LOWENSTAM Y WEINER, 1989). La pre-

sencia de pequeñas cantidades de calcita

puede interpretarse de varios modos, tal

como indican BALMAIN, HANNOYER Y

López (1999). Cabe la posibilidad de que

esta especie mineral se concentre en deter-

minadas partes de la concha, lo que no

parece probable, ya que las proporciones

en las que aparece son distintas entre los

ejemplares, siendo éstos similares macro-

morfológicamente. Otra posibilidad es que

se trate de una fase secundaria a la for-

mación de la concha por un proceso de

contaminación natural, dado que estos

animales habitan ambientes carbonatados.

18

Por último, cabría el paso de un polimorfo

a otro (de aragonito a calcita) generado

por el calentamiento durante la molienda;

este proceso tampoco parece muy proba-

ble dada la escasa velocidad de la reac-

ción (CARLSON, 1983).

Los resultados del análisis XRF se

muestran en la Tabla I. Se observa que

entre los elementos mayores destaca por

su abundancia el calcio, constituyendo,

en óxidos, el 55,72% del ejemplar A y el

55,44% del ejemplar B. Las pérdidas por

calcinación (LOL) son elevadas, superiores

al 43%. Les siguen en proporción el silicio

(en óxidos, 0,25% para el ejemplar A y

0,30% para el ejemplar B), mientras que el

resto de los óxidos aparecen con valores

inferiores al 0,1% (salvo el de magnesio en

el ejemplar A, cuyo valor es 0,12%). De los

elementos menores, el estroncio es el que

presenta el porcentaje más alto en ambas

conchas (624,7 ppm en el ejemplar A y

579,2 ppm en el ejemplar B).

Las proporciones de CaO son las espe-

radas dada la mineralogía aragonítica, y

en menor medida calcítica (ambos poli-

morfos del CaCO»), de la concha. Con estas

proporciones de CaO, cabe calcular el con-

tenido de CaCO> presente, que resulta de

99,4% y 98,9% respectivamente para el

ejemplar A y B (Tabla 1). Lo mismo se

podría decir de las altas pérdidas por cal-

cinación que corresponderían al CO» gene-

rado en la descomposición térmica de los

carbonatos; en este caso el contenido de

CaCO> calculado a partir de ellas es de

98,0% y 98,2% respectivamente para el

ejemplar A y B (Tabla ]). El ajuste con las

anteriores cifras es aceptable.

MÁRQUEZ ET AL.: Composición y microestructura de la concha de /berus gualtierianus

x103 Ca

Cuentas por segundo

2. .NNO ps

0,0 SS S IO MAAS

ON OSO ESAS SEO

Energía

Cuentas por segundo

SS Lore

Ss e se

0.0.0.3 1,0 1,53:20.25 510.35) 2401 172/

Energía

Figura 1. Espectros de microanálisis de rayos X (EDR) en fracturas frescas de conchas de /berus

gualtierianus morfotipo gualtierianus. Izquierda: ejemplar A; derecha: ejemplar B.

/

Figure 1. X-ray microanalysis spectra (EDR) in fresh fractures of the shells of Iberus gualtierianus morp-

hotype gualtierianus. Left: specimen A; right: specimen B.

Las proporciones de Si0O»2 (0,25% en

el ejemplar A y 0,30% en el ejemplar B) y

ADO (0,05% y 0,06% para el ejemplar A

y B, respectivamente), son imputables a

la contaminación de la concha por silica-

tos del suelo, tratándose de componen-

tes secundarios.

A pesar de que el aragonito es un

mineral bastante puro desde el punto de

vista químico (DEER, HOWIE Y ZUSSMAN,

1992), el Sr presente en las conchas se

incluiría en la red de este mineral susti-

tuyendo al Ca, lo cual es un hecho amplia-

mente descrito en la bibliografía sobre ara-

gonito biogénico (TUREKIAN Y ARMSTRONG,

1960; DoDD, 1967; MASUDA, 1976; MASUDA

E HIRANO, 1980). Lo mismo le ocurriría al

Mg, pudiendo ser también, en parte, pro-

ducto de contaminación de los materiales

dolomíticos del suelo.

El Na es un catión que también puede

estar presente en la red del aragonito,

incluso en cantidades elevadas (DAUPHIN

Y DENIS, 2000; DAUPHIN, GUZMAN, DENIS,

CUIF, Y ORTLIEB, 2003), aunque su propor-

ción es mayor en las conchas marinas

(WADA Y FUJINUKL, 1974). Otros autores

han considerado que la presencia del Na

en las conchas de los moluscos, se debe a

procesos de contaminación (por ejemplo,

EsTES, 1972), o al hecho de ser un catión

que forma parte de la composición

química de la matriz orgánica de las

conchas (LÉCUYER, 1996).

El análisis del contenido de carbonato

cálcico equivalente de las conchas (Tabla

D, registra los siguientes porcentajes:

97,52% para el ejemplar C y 97,72% para

el ejemplar D. Estos resultados son de mag-

nitudes similares a los obtenidos por aná-

lisis XRF ya expuestos (Tabla 1). Se puede,

por tanto, afirmar que los ejemplares de

[ gualtierianus morfotipo gualtierianus ana-

lizados tienen un porcentaje de CaCO3

entre el 97% y el 99% y que se encuentra,

de esta forma, dentro del rango de valores

conocido para la concha de los moluscos

(entre 95% y 99,9%); el resto son compo-

nentes orgánicos (HARE Y ABELSON, 1965).

El estudio de la composición se ha

completado con los resultados de micro-

análisis de rayos X (EDR) (Fig. 1) de la

superficie de las fracturas frescas de las

conchas. Se ha demostrado la composi-

ción carbonática al aparecer las líneas

correspondientes a los elementos carbono,

oxígeno y calcio. Hay, pues, una perfecta

coherencia con el resto de los resultados

expuestos. Los elementos químicos detec-

tados en XRF (Tabla D, distintos al carbono,

oxígeno y calcio, no se registran con EDR,

al encontrarse sus cantidades por debajo

de los límites de detección de esta técnica.

El estudio de la concha en fractura

fresca bajo el microscopio electrónico de

barrido (MEB) ha puesto de manifiesto,

en primer lugar, su constitución en capas

superpuestas (Figs. 2A, B). Dichas capas

presentan estructura laminar cruzada (se

describirá posteriormente) y distintas

orientaciones de los cristales de aragonito

(Fig. 2B). Esta configuración interna de la

1

Iberus, 23 (1), 2005

Tabla IT. Capas reconocidas, y espesores medios de las mismas (um), en una sección perpendicular

(A) y en otra paralela (B) a las líneas de crecimiento de la concha estudiada del ejemplar A de

Iberus gualtierianus morfotipo gualtierianus. Medidas realizadas en las observaciones con microsco-

pio electrónico de barrido (MEB). La capa a es la más interna de la concha y la e la más externa.

Table II. Layers observed and their mean thicknesses (um), in both perpendicular (A) and parallel (B)

sections to the growth lines of the studied shell of the specimen A of lberus gualtierianus morphotype

gualtierianus. The measurements were carried out using scanning electron microscopy (SEM). Layer a is

the most internal and layer e the most external.

A. Sección perpendicular a las líneas de crecimiento

Copos 0 b C d e > (espesor total)

Media vuelta

de espira esp ds nm. esp dsu nm esp ds. Mo sesp 05 Desp USES ds n

Última 132,39 123,14 8 132,60 43,03 8 101,84 14,331 4 366,83 60,16 20

Penúltima 101,14 15,17 8 55,00 16,22 8 61,47 748 8 30/46 9,57 8 55,91 18,93 8 303,98 13,47 40

Antepenúltima 113,24 34,56 8 67,50 36,29 8 46,32 885 8 36,91 911 8 3406 535 8 298,03 18,83 40

B. Sección paralela a las líneas de crecimiento

Copos 0 b C d e > (espesor total)

Media vuelta

de espira esp 05 nm. esp. 05 nm esp. ds n esp “05. .n €5p... 05 Mn esp

Último 250,972815 40 53131331054 304,10 21,03 8

Penúltima 77,91 3,91 8 55,57 10,07 8 66,56 7,73 8 5247 919 8 42,335 6,61 8 294,86 7,50 40

Antepenúltima 47,93 13,04 8 60,75 894 8 48,17 1238 8 26,59 5/04 8 24,89 7,52 8 208,33 9,38 40

esp: espesor medio (pm); ds: desviación típica; n: número de medidas realizadas

concha ha permitido establecer el número

y espesor de las capas en las últimas fases

del crecimiento del organismo (Tabla ID.

Destaca, como hecho observable, que el

espesor de las capas, en general, varía de-

pendiendo de la zona que se trate, au-

mentando bajo las costillas y decreciendo

en las estriaciones (Fig. 2A) presentes en

estas conchas (LÓPEZ-ALCÁNTARA, RIVAS,

ALONSO E IBÁÑEZ, 1985). Además, cabe

destacar que, en la mayoría de los casos,

los espesores medidos tienen un cierto

grado de espesores aparentes, ya que las

secciones observadas son irregulares al ser

fracturas frescas y no se disponen total-

mente perpendiculares al plano del ob-

(Página derecha) Figura 2. Imágenes de microscopía electrónica de barrido de la concha de l/berus

gualtierianus morfotipo gualtierianus. A: microestructura en corte fresco de una sección perpendicu-

lar a las líneas de crecimiento de la penúltima media vuelta de espira. Presencia de cinco capas con

estructura laminar cruzada y orientaciones distintas. La flecha indica una costilla y el incremento de

los espesores de las capas debajo de ella. B: detalle de la imagen anterior, demostrando las distintas

orientaciones de los cristales en las capas contiguas (por ejemplo, a con b). C: microestructura en

corte fresco de una sección paralela a las líneas de crecimiento de la penúltima media vuelta de espira.

Se muestran las varillas de aragonito (lt) que componen el tercer orden de la estructura laminar cruzada,

y los paquetes de varillas (ls) que constituyen el segundo orden de dicha estructura. D: microestruc-

tura en corte fresco de una sección paralela a las líneas de crecimiento de la última media vuelta de

espira. Se muestran las unidades estructurales de primer orden de forma tabular (Ip). Las flechas indican

las dos distintas orientaciones de los cristales en las láminas de primer orden. E: superficie interna de

la concha en la última media vuelta de espira. Se muestran, con flechas discontinuas, las ondulacio-

nes (od) y estrías finas (est). La flecha continua señala hacia la abertura de la concha. E: superficie

interna de la concha en la última media vuelta de espira. Se destacan con la flecha discontinua las

pequeñas protuberancias (pb). La flecha continua señala hacia la abertura de la concha.

20

MARQUEZ ET AL.: Composición y microestructura de la concha de /berus gualtierianus

NASA

E F

Figure 2. Scanning electron micrographs of the shell of Tberus gualtierianus morphotype gualtierianus.

A: microstructure in freshly cut section perpendicular to the growth lines of the penultimate half turn of

the whorl. Presence of the five layers with crossed. lamellar structure and different orientations. The

arrow shows a rib and the increase in thickness of the layers below it. B: detail of A showing different

orientations of the crystals in the contiguous layers (e. g. a with b). C: microstructure in freshly cut

section parallel to the growth lines of the penultimate half turn of the whorl. The aragonite rods (lt)

which constitute the third order of the crossed lamellar structure and the bundles of rods (ls) forming the

second order of the structure can be seen. D: microstructure in freshly cut section parallel to the growth

lines of the last half turn of the whorl. The first order structural units (lp) can be seen. The arrows show

the two different orientations of the crystals in the first order lamellae. E: internal surface of the shell in

the last half turn of the whorl. The dotted arrows show the ondulations (od) and fine striations (est).

The solid arrow points towards the shell aperture. E: internal surface of the shell in the last half turn of

the whorl. The dotted arrows show the small protuberances (pb). The solid arrow points towards the

shell aperture.

2]

Iberus, 23 (1), 2005

servador, a pesar de los esfuerzos reali-

zados en el microscopio para que así sea.

Por todo ello se aportan valores medios

(Tabla ID.

De estos datos se deduce, en primer

lugar, que en la sección perpendicular a

las líneas de crecimiento de la última

media vuelta de espira (Tabla IIA) se

reconocen tres capas (a, b y c), mientras

que en la sección paralela a dichas líneas

sólo se reconocen dos (Tabla IIB). Pensa-

mos que esto pueda deberse a la orien-

tación relativa de los cristales de arago-

nito en las láminas de las capas, de tal

forma que, en la sección paralela, dos de

ellas, las más internas, tienen una orien-

tación similar y se confunden.

El número de capas aumenta hasta

cinco en la penúltima y antepenúltima

medias vueltas de espira en ambas sec-

ciones, lo que supone admitir que la

concha va cambiando en su microestruc-

tura en el transcurso de las fases de

desarrollo del animal. |

Una cuestión interesante es el espesor

de las capas, variable de unas a otras (entre

24,89 um y 250,97 um) (Tabla II). Por otra

parte, el espesor total de la concha es de

unos 300 um, tendiendo a incrementarse

en la última media vuelta de espira res-

pecto a las anteriores, lo que apuntaría no

sólo a una reconstrucción en el número de

capas ya referido, sino también a una

pequeña reducción en el volumen de la

concha en las medias vueltas de espira

más internas.

Respecto a la microestructura en

detalle de las capas, hay que decir que

están constituidas por distintas unida-

des estructurales que crecen en comple-

jidad y que se denominan, respectiva-

mente, láminas de tercer, segundo y

primer orden, como corresponde con su

calificación de laminar cruzada

(BSGGILD, 1930; UOZUMI, IWATA Y TOGO,

1972; CARTER, 1990). Las unidades más

pequeñas o de tercer orden, tienen

forma de varilla y un espesor medio de

0,49 um (desviación típica, 0,14 um; n,

20) (Fig. 2C). Estas unidades son parale-

las entre sí y están unidas generando

pequeños paquetes (Fig. 2C) que consti-

tuyen las unidades estructurales inter-

medias O de segundo orden, con un

22

grosor medio de unos 5 um (4,91 um;

desviación típica, 1,16 um; n, 6). A su

vez, éstas están unidas entre sí dando

lugar a las unidades estructurales

mayores o de primer orden que presen-

tan una forma plana o tabular (Fig. 2D).

De acuerdo con los caracteres descri-

tos y según la terminología utilizada por

CARTER (1990), esta microestructura

podemos clasificarla como laminar

cruzada simple (“simple crossed lame-

llar”). La microestructura de tipo laminar

cruzada es la más extendida en las conchas

de los moluscos (B9GGILD, 1930), estando

constituida por aragonito salvo en algunas

especies (UOZUMI ET AL., 1972). Una carac-

terística de esta microestructura (BY9GGILD,

1930; CARTER, 1990) es que la orientación

de las unidades aciculares de aragonito

(láminas de tercer orden), así como los

paquetes que componen (láminas de

segundo orden), cambian de orientación

entre las láminas de primer orden adya-

centes (Fig. 2D).

Tanto la constitución en varias capas

superpuestas, como la microestructura

laminar cruzada, confieren a la concha

de 1. gualtierianus morfotipo gualtierianus

una gran resistencia mecánica.

La superficie interna de la concha es

prácticamente lisa (Fig. 2E), si bien en

algunas zonas presenta suaves ondula-

ciones (Fig. 2E), que pueden incluso llegar

a constituir pequeños montículos o pro-

tuberancias (Fig. 2F) y que parecen ser un

reflejo suavizado de la ornamentación

externa. También por zonas, la superficie

interna de la concha está microesculpida

con estriaciones muy finas (Fig. 2B).

AGRADECIMIENTOS

Al Centro de Instrumentación Cientí-

fica (CIC) de la Universidad de Granada,

en el que se ha realizado el análisis de

fluorescencia de rayos X (XRP).

Al Servicio de Microscopía Electró-

nica de Barrido y Microanálisis de la

Facultad de Farmacia (Universidad de

Granada) por la preparación de las

muestras, la realización de las fotogra-

fías y el estudio de microanálisis de

rayos-X de energía dispersiva (EDR).

MÁRQUEZ ET AL.: Composición y microestructura de la concha de /berus gualtierianus

BIBLIOGRAFÍA

ARRÉBOLA-BURGOS, J. R., 2002. Caracoles terres-

tres de Andalucía. Consejería de Medio Am-

biente, Junta de Andalucía, Manuales de

Conservación de la Naturaleza, n” 1, 64 pp.

BALMAIN, J., HANNOYER, B. Y LÓPEZ, E., 1999.

Fourier Transform Infrared Spectroscopy

(FTIR) and X-Ray Diffraction Analyses of

Mineral and Organic Matrix During Hea-

ting of Mother of Pearl (Nacre) from the Shell

of the Mollusc Pinctada maxima. Journal of

Biomedical Materials Research Part B: Applied

Biomaterials, 48: 749-754.

BOGGILD, O. B., 1930. The shell structure of the

mollusk. Det Kongelige Danske Videnskabernes

Selskabs Skrifter Naturvidenskabelig og Mathe-

matisk Afdeling, ser. 9, 2: 231-326.

CARLSON, W. D., 1983. The polimorphs of

CaCOz3 and the aragonite-calcite transfor-

mation. En: Reeder, R. J. (Ed.), Reviews in Mi-

neralogy, Vol. 11: Carbonates: Mineralogy and

Chemistry. Mineralogical Society of America,

Washington, D. C., pp. 191-225.

CARTER, J. G. Y CLARK Il, G. R., 1985. Classifi-

cation and phylogenetic significance of mo-

lluscan shell microstructure. En: Bottjer, D.

J., Hickman, C. S., Ward, P. D., Broadhead,

T. W. (Eds.), Mollusks: notes for a short course.

University of Tennessee, Department of Ge-

ological Sciences Studies in Geology, pp. 50-

EL.

CARTER, J. G., 1980. Guide to bivalve shell mi-

crostructures. En: Rhoads, D.C., Lutz, R. A.

(Eds.), Skeletal Growth of Aquatic Organisms.

Plenum, New York, pp. 645-673.

CARTER, J. G., 1990. Skeletal Biomineralization:

Patterns, Processes and Evolutionary Trends

Vol. I Van Nostrand Reinhold, New York, 823

bs

CHATEIGNER, D., HEDEGAARD, C. Y WENK, H. R,,

2000. Mollusc shell microstructures and crys-

tallographic textures. Journal of Structural Ge-

ology, 22: 1723-1735.

DAUPHIN, Y. Y DENSS, A., 2000. Structure and

composition of the aragonitic crossed lame-

llar layers in six species of Bivalvia and Gas-

tropoda. Comparative Biochemistry and Phy-

siology, Part A, 126: 367-377.

DAUPHIN, Y., CUTE, J. P., MUTVEL H. Y DENIS,

A., 1989. Mineralogy, chemistry and ultras-

tructure of the external shell-layer in ten

species of Haliotis tuberculata (Mollusca:

Archaeogastropoda). Bulletin of the Geologí-

cal Institutions of the Untversity of Uppsala,

15: 7-38.

DAUPHIN, Y., GUZMAN, N., DENIS, A., CUIF, J.

P. Y ORTLIEB, L., 2003. Microstructure, na-

nostructure and composition of the shell of

Concholepas concholepas (Gastropoda, Muri-

cidae). Aquatic Living Resources, 16: 95-103.

DEER, W. A., HOWIE, R. A. Y ZUSSMAN, J., 1992.

An Introduction to the Rock-Forming Minerals.

Longmans Scientific 8: Technical, England,

696 pp.

DODD, J. R., 1967. Manganesium and strontium

in calcareous skeletons. A review. Journal of

Paleontology, 41: 1313-1329.

ESTES, E. L. IHl, 1972. Diagenetic alteration of Mer-

cenaria mercenaria as determined by laser mi-

croprobe analysis. Thesis, University of North

Carolina, Chapel Hill.

GÁMIZ, E., 1987. Caracterización de caolines, tal-

cos y bentonitas españoles para su posible apli-

cación en Farmacia. Tesis Doctoral, Universi-

dad de Granada, 418 pp.

GRÉGOIRE, C., DUCHATEAU, G. Y FLORKIN, M.,

1949. Exame au microscope électronique de

la pellicule prénacrée et de la nacre décalci-

fiée de l'anodonte. Archives Internationales de

Physiologie, 57: 121-124.

GRÉGOIRE, C., 1972. Structure of the molluscan

shell. En: Florkin, M. €z Scheer, B. T. (Eds.),

Chemical zoology, vol. VII (Mollusca). Acade-

mic Press, New York, London, pp. 45-102.

HARE, P. E. Y ABELSON, P. H., 1965. Amino acid

composition of some calcified proteins. Car-

negie Institution Washington Yearbook, 64: 223-

SD

HAssE, B., EHRENBERG, H., MARXEN, J. C., BEC-

KER, W. Y EPPLE, M., 2000. Calcium carbonate

modifications in the mineralized shell of the

freshwater snail Biomphalaria glabrata. Che-

mistry-A European Journal, 6 (20): 3679-3685.

HEDEGAAROD, C., LINDBERG, D. R. Y BANDEL, K.,

1997. Shell microstructure of a Triassic pate-

llogastropod limpet. Lethaia, 30: 331-335.

HEDEGAARD, C., 1997. Shell structure of the re-

cent Vetigastropoda. Journal of Molluscan Stu-

dies, 63: 369-377.

(JCPDS) JornTr COMMITTEE ON POWDER DIF-

FRACTION STANDARDS AND AMERICAN SO-

CIETY FOR TESTING AND MATERIALS, 1974. Se-

lected Powder diffraction data for minerals.

JCPDS, Park Lane, Swarthmore, PA.

KOBAYASHI, I., 1964. Introduction to the shell

structure of bivalvian molluscs. Chikyu Ka-

SaKu, 73: 1512.

KOBAYASHI, IL, 1971. Internal shell microstruc-

ture of recent bivalvian molluscs. Science Re-

port of Nigata University, Series E (Geology

and mineralogy), 2: 27-50.

KRANTZ D. E., WILLIAMS, D. F. Y JONES, D. S.,

1987. Ecological and paleoenvironmental in-

formation using stable isotope profiles from

living and fossil mollusks. Palaeogeography,

Palaeoclimatology, Palaeoecology, 58: 249-266.

LÉCUYER, C., 1996. Effects of heating on the ge-

ochemistry of biogenic carbonates. Chemical

Geology, 129 (3-4): 173-183.

2)

Iberus, 23 (1), 2005

LÓPEZ-ALCÁNTARA, A., RIVAS, P., ALONSO, M.

R. E IBÁÑEZ, M., 1983. Origen de Iberus gual-

tierianus. Modelo evolutivo. Haliotis, 13: 145-

154.

LÓPEZ-ALCÁNTARA, A., RIVAS, P., ALONSO, M.

R. E IBÁÑEZ, M,, 1985. Variabilidad de Iberus

gualtierianus (Linnaeus, 1758) (Pulmonata,

Helicidae). Iberus, 5: 83-112.

LOWENSTAM, H. A. Y WEINER, S., 1989. On Bio-

mineralization. Oxford University Press, New

York, 324 pp.

MACCLINTOCK, C., 1967. Shell structure of pa-

telloid and bellerophontoid gastropods (Mo-

llusca). Bulletin of the Peabody Museum of Na-

tural History, 22: 1-140.

MAckKay, I. H., 1952. The shell structure of the

modern mollusks. Quarterly of the Colorado

School of Mines, 47: 1-27.

MÁRQUEZ, R., 2003. El cuarzo de la fracción arena

fina en suelos de la provincia de Granada. Me-

moria de Investigación Tutelada, Universi-

dad de Granada, 178 pp.

MASUDA, F. E HIRANO, M,, 1980. Chemical com-

position of some modern marine pelecypod

shells. Science Reports Institute of Geoscience

University of Tsukuba, Section B, 1: 163-177.

MASUDA, F, 1976. Strontium contents in shell of

Glycymeris. Journal of the Geological Society of

Japan, 82: 565-572.

MUTVEL H., DAUPHIN, Y. Y CUIE, J. P., 1985. Ob-

servations sur l'organisation de la couche

externe du test des Haliotis (Gastropoda):

un cas exceptionnel de variabilité mineralo-

gique et microstructurale. Bulletin du Mu-

séum National d Histoire Naturelle, Paris, 4 Sér.,

sect A. 7739

Popov, S. V. Y BARSKOV, I. S., 1978. Shell struc-

ture of mollusks and its value in phylogeny

and classification. Malacological Review, 11:

152-153.

SABATIER, G., 1953. Application de la diffraction

des rayon X a l'étude des coquilles de Mo-

llusques. Cahiers des naturalistes, Bulletin des

Naturalistes Parisiens, nouvelle série, 8: 97-102.

SCHNEIDER, J. A. Y CARTER, J. G., 2001. Evolu-

tion and phylogenetic significance of Car-

dioidean shell microstructure (Mollusca, Bi-

valvia). Journal of Paleontology, 75: 607-643.

SORIANO, M., 1994. Estudio geofarmacéutico de

polvos de talco. Primera aproximación a la far-

macopea internacional armonizada. Tesis Doc-

toral, Universidad de Granada, 619 pp.

TAYLOR, J. D., KENNEDY, W. J. Y HALL, A., 1969.

The shell structure and mineralogy of the

Bivalvia. Introduction. Nuculacea-Trigona-

cea. Bulletin of the British Museum of Natural

History (Zoology), Supplement, 3: 1-125.

24

TAYLOR, J. D., KENNEDY, W. J. Y HALL, A., 1973.

The shell structure and mineralogy of the

Bivalvia. II. Lucinacea-Clavagellacea. Con-

clusions. Bulletin of the British Museum of Na-

tural History (Zoology), Supplement, 22 (9):

253-294.

TAYLOR, J. D., 1973. The structural evolution of

the bivalve shell. Palaeontology, 16: 519-534.

TUREKIAN, K. K. Y ARMSTRONG, R. L., 1960.

Magnesium, strontium and barium concen-

trations and calcite-aragonite ratios of some

recent molluscan shells. Journal of Marine Re-

search, 18: 133-151.

UOzUMI, S., IwATA, K. Y TOGO, Y., 1972. The ul-

trastructure of the mineral in and the cons-

truction of the crossed-lamellar layer in mo-

lluscan shell. Contributions from the Department

of Geology and Mineralogy, Faculty of Science,

Hokkaido University, 1236: 447-477.

WADA, K. Y FUJINUKL, T., 1974. Physiological re-

gulation of shell formation in molluscs. 1.

Chemical composition of extrapallial fluid.

Bulletin of the National Pearl Research Labora-

tory, 18: 2085-2110.

WATABE, N. Y WILBURG, K. M., 1961. Studies on

shell formation. IX An Electron Microscope

Study of Crystal Layer Formation in the Oys-

ter. The Journal of Biophysical and Biochemical

Cytology, 9: 761-777.

WATABE, N., 1984. Mollusca: Shell. En: Bereiter-

Hahn, J., Matolsty, K. 8: Richards, S. K. (Eds.),

Biology of the integument, vol. I (Invertebrates).

Springer-Verlag, New York, pp. 448-485.

WILBUR, K. M., 1964. Shell formation and re-

generation. En: Wilbur, K. M. y Yonge, C. M.

(Eds.), Physiology of Mollusca. Academic Press,

London, pp. 243-282.

WILBUR, K. M. Y SALEUDDIN, A. S. M., 1983.

Shell formation. En: Saleuddin, A. S. M. éz

Wilbur, K. M. (Eds.), The Mollusca, Vol. 4,

Part 1. Academic Press, New York, 523 pp.

WISE, S. W. Jr. Y HaY, W. W., 1968. Scanning

Electron Microscopy of molluscan shell ul-

trastructures II. Observations of growth sur-

faces. Transactions of the American Microsco-

pical Society, 87: 419-430.

Wise, S. W. JR., 1970. Microarchitecture and

mode of formation of nacre (mother-of-pe-

arl) in Pelecypods, Gastropods and Cepha-

lopods. Eclogae Geologicae Helvetiae, 63: 775-

797.

O Sociedad Española de Malacología —__——T— Iberus, 23 (1): 25-31, 2005

Changes in phospholipid and lipid peroxidation levels due

to latex of Croton tiglium in freshwater snail Lymnaea

acuminata

Cambios en los niveles de peroxidación de fosfolípidos y lípidos por

efecto del latex de Croton tiglium sobre el molusco dulceacuícola

Lymnaea acuminata

Digvijay SINGH*!, Ram P. YADAV* and Ajay SINGH*

Recibido el 18-VII-2003. Aceptado el 13-1X-2004

ABSTRACT

Exposure to 40% and 60% of LCso of 48h of the latex extract of Croton tiglium Linn

(Euphorbiaceae) for 24h, 48h, 72h and 96h significantly reduced the endogenous levels

of phospholipid and increased the rate of lipid peroxidation in brain and foot tissue of

lymnaea (Radix) acuminata Lamarck. Alteration in the levels of phospholipid and rate of

lipid peroxidation were time and dose dependent. It is concluded that latex extract of Cro-

ton tiglium or its metabolites increase peroxidation of lipids in Lymnaea acuminata, which

brings about change in membrane permeability to various ions and may thus be the cause

of the toxicity of this plant extract.

RESUMEN

La exposición a LCso de 48 horas del extracto del latex de Croton tiglium al 40% y 60%

durante 24, 48, 72 y 96 horas redujo significativamente los niveles de fosfolípidos e

incrementó la tasa de peroxidación de lípidos en los tejidos cerebral y del pie de Lym-

naea (Radix) acuminata Lamarck. Ambos cambios dependen tanto del tiempo como de la

dosis. Esto confirma que dicho latex o sus metabolitos incrementa la peroxidación de lípi-

dos en el molusco, por medio de cambios en la permeabilidad de la menbrana a varios

iones. Esta puede ser la causa de la toxicidad del extracto de esta planta.

KEY WORDS: Lipid peroxidation, phospholipids level, latex of Croton tiglium.

PALABRAS CLAVE: Peroxidación de lípidos, niveles de fosfolípidos, latex de Croton tiglium.

INTRODUCTION

The snail Lymnaea acuminata is the SINGH AND SINGH, 2002; TRIPATHI AND

vector of liver flukes Fasciola hepatica SINGH, 2003; SINGH AND SINGH, 2003).

and Fasciola gigantica, which cause Heavy use of synthetic pesticides/ syn-

endemic fascialiasis in cattle and liver thetic pyrethroids for the control of

stock in northern parts of India (YADAV, aquatic vectors created other serious

* Natural Product Laboratory, Department of Zoology, D.D.U. Gorakhpur University, Gorakhpur — 273 009

(U.P.) INDIA 'Corresponding author: Dr. Digvijay Singh. e-mail: digvijaysinOrediffmail.com

P4S

Iberus, 23 (1), 2005

problems to the aquatic environment

(SASTRY AND SHUKLA, 1993; DEVI, 1997).

Their hazardous nature has prompted

scientists to find out non-disruptive,

suitable and newer options for the

control of weed and aquatic pests. In

recent times, the use of natural pesti-

cides has gained popularity all over the

world. These plant products are the

focus of attention as a suitable alterna-

tive to synthetic pesticides due to some

ideal properties i.e. low cost, easy avail-

ability and biodegradability in nature

(MARSTON AND HOSTETTIMAN, 1985;

SINGH AND SINGH, 2002).

A large number of plant pesticides are

lethal to aquatic snails (GODAN, 1983;

SINGH, SINGH, MISRA AND AGARWAL, 1996).

SINGH AND AGARWAL, (1984 a, b) for the

first time reported that the lattices of

euphorbiales are highly toxic to snails.

Later SINGH AND AGARWAL (1992; 1993

and 1995) reported several euphorbious

species (falta algo :p.ej.species) as plant

mollusicicides. It is known that the

primary target of these plant molluscicides

is the nervous system (SINGH AND

AGARWAL, 1984a, b). It was observed that

these plant molluscicides affect acetyl-

choline, dopamine and non-epinephrine

levels in nervous tissues of exposed

animals (SINGH AND AGARWAL, 1995). Evi-

dences for tissues injury induced by certain

anti cholinesterase (Anti-ACHE) pesticides

has been shown to be associated with

increased fragility of various biological

membranes and the structural lipids of

biomemebranes undergo peroxidation

decomposition (CHVAPIL, RYAN AND

BRADA, 1972).

YADAV AND SINGH (2001, 2002)

reported that latex extract of Codiaeum

variegatum and Croton tiglium have high

molluscicidal and anti-cholinesterase

activity against snails Lymnaea acuminata

and Indoplanorbis exustus. In the present

work the author is interested in

knowing the sub-lethal effects of the

latex extract of Croton tiglium on the

endogenous level of phospholipids and

rate of lipid peroxidation level in

nervous and foot tissues of the freshwa-

ter snail Lymnaea acuminata to see if it

can serve as effective molluscicides.

26

MATERIAL AND METHODS

Adult Lymnaea acuminata (2.4+0.4 cm

in length) were collected from local ponds

and acclimatized in the laboratory for 96h

in dechlorinated tap water. Batches of 50

snails were kept in 12L glass aquaria con-

taining 6L-dechlorinated tap water. Snails

were treated according to methods of

SINGH AND AGARWAL (1986). Experiments

were carried out at 24-26 *C. No food was

given during the course of the experiment.

There was no mortality in either controls

or experiments. Oxygen concentration was

normal in control animals as there was no

sign of oxygen deficiency stress. 40 snails

/ 5L water were used which caused no

oxygen stress in test snails. The snails were

exposed to 40% (0.016 mg/L) and 60%

(0.024 mg/L) of 48h LC5o of 0.04 mg/L for

24h, 48h, 72h and 96h. These doses were

based on 48h LCso values reported by

YADAV AND SINGH, (2001). After 24h, 48h,

72h or 96h the snails were removed from

the aquaria and rinsed in water. Control

groups were kept in de-chlorinated tap

water without any treatment. Endogenous

levels of phospholipids were determined

in nervous and foot tissue by the method

Of FISKE AND ROW, (1925) as modified by

DITTMER AND MICHAEL (1965), and mod-

ified for snails by SINGH, SINGH AND

AGARWAL (1993). The rate of lipid perox-

idation was determined in nervous and

foot tissues by the methods of (UTELY,

BERMHEIM AND HOCHTEIN, 1967).

Phospholipids levels estimation: Brain and

foot tissue were dissected out and placed

on filter paper for removal of adherent

water. 300 mg each of brain and foot

tissues were used for each replicate. Brain

tissue was pooled from eighty five snails.

The tissues were then homogenized in

chloroform/methanol mixture (2:1v/v)

using a Teflon pestle to give a 5% w/v

homogenate . Supernatants were filtered

and two drops of 0.05M NaCl solutions

was added to the supernatant and left for

24h. After 24h the upper layer was dis-

carded. 1 mL of lipid layer was placed in

a test tube and solvent was removed by

heating in a boiling water bath. 1 mL of

distilled water and 0.4 mL of 10%

trichloroacetic acid was added to this and

SINGH ET AL.: Latex of Croton tiglium changes lipid peroxidation in Lymnaea acuminata

Table 1. /n vivo effect of exposure to 40% and 60% of 48h LCso of latex of Croton tiglium on the

levels of phospholipids in the nervous and foot tissues of snail L. acuminata after exposure to 24h,

48h, 72h and 96h. Values are mean +SE of six replicates. Values in parenthesis indicate % of

control. Student test *t” test demonstrated that the treated values were significantly different from

control (P<0.05). Two-way analysis of variance demonstrated that the changes were dose and time

dependent. 300 mg nervous and foot tissues were taken for all estimations.

Table 1. Efecto in vivo de la exposición al 40% y 60% de 48h LC5o del latex de Croton tiglium en los

niveles de fosfolípidos en los tejidos nervioso y del pie de L. acuminata tras 24h, 48h, 72h y 96h de

exposición. Los valores son la media +SE de 6 réplicas. Valores en paréntesis indican % del control. El

test “t” de Student muestra que los valores tratados son significativamente diferentes de los del control

(P<0.05). El análisis de varianza de dos vías demuestra que los cambios dependen de la dosis y el

tiempo. Se usaron 300 mg de tejido nervioso y del pie en todas las estimaciones.

Tissues 40% of LCso (48h) 60% of LCso (48h)

Control Broin 23.26+0.15 (100) 23.2640.15 (100)

Foot 21.01%0.31 (100) 21.0140.31 (100)

24h Brain 17.45+0.18 (62.12) 13.26+0.22 (57.00)

Foot 18.00+0.18 (85.67) 16.30.17 (77.58)

48h Brain 14.19+0.39 (60.99) 13.520.7 (51.52)

Foot 14.95+0.26 (71.15) 13.22+0.19 (62.92)

72h Brain 13.02+0.19 (55.96) 11.00+0.25 (47.29)

Foot 11.29+0.33 (53.73) 10.64+0.39 (50.64)

96h Brain 9.2+0.23 (39.55) 8.67+0.20 (37.27)

Foot 9.63+0.34 (45.83) 8.23+0.28 (39.317)

heated. After this, 0.4 mL of 2.5 % Ammo-

nium molybdate solutions and 0.2 mL of

amino naphthosulphonic acid was added

and the mixture heated at 80*C for 15 min,

cooled and diluted with 4 ml distilled

water. After 5 minutes the absorbance was

read at 640 nm with the help of a spec-

trophotometer (No. 106 S.No. 3265 Sys-

tronics). The blank consisted of 1 mL dis-

tilled water, 0.4 mL 10% dichloroacetic

acid, 0.4 mL 2.5% ammonium molybdate

solution, 0.2 mL of aminonaphthosul-

phonic acid and 4.0 mL distilled water but

no tissue homogenate. Standards curves

were prepared by reading the absorbance

of different concentrations of monopotas-

sium phosphate solutions containing 0.008

to 0.04 mg of pure salt.

Lipid peroxidation estimation: Brain and

foot tissues (100 mg each) were homoge-

nized in chilled 0.15 M KCI solution using

a Teflon pestle to give a 10% w/v

homogenate. 1 ml aliquots of the

homogenate were incubated at 37*C for

2h, following which 1.0 ml of 10% (w/v)

trichoroacetic acid was added. After thor-

ough mixing, the mixture was centrifuged

at 2200g for 10 min and 1ml of supernatant

was then taken with an equal volume of

0.67% (w/v) 2-thiobarbituric acid (Sigma

Chemical Company, USA) and kept in a

boiling water bath for 10 min, cooled and

diluted with one ml of distilled water. The

absorbance read at 535 nm against a blank

containing only KCl solution and results

were expressed as mol of malonal-

dialdehyde formed per 30 min /mg tissue.

The extinction coefficient was 1.56x105 as

described by UTELY ET AL., (1967). The

methods of SOKAL AND ROHLF (1973) were

applied for analysis of students “t” test

and two-way analysis of variance (P<0.05).

RESULTS

The endogenous levels of phospho-

lipids in the brain and foot tissues were

23.26 and 21.01 mg/g, respectively. In

vivo 24h, 48h, 72h and 96h exposure to

27

Iberus, 23 (1), 2005

Table IL. Zn vivo effect of exposure to 40% and 60% of 48h LCso of Croton tiglium on the rate of

lipid peroxidation in the nervous and foot tissues of snail L. acuminata after exposure to 24h, 48h,

72h and 96h. Values are mean +SE of six replicates. Values in parenthesis indicate % of control.

Student test “t” test demonstrated that the treated values were significantly different from control

(P<0.05). Two-way analysis of variance demonstrated that the changes were dose and time depen-

dent. 600 mg nervous and foot tissues were taken for all estimations.

Table 11. Efecto in vivo de la exposición al 40% y 60% de 48h LC5o del latex de Croton tiglium en los

niveles de fosfolípidos en los tejidos nervioso y del pie de L. acuminata tras 24h, 48h, 72h y 96h de

exposición. Los valores son la media +SE de 6 réplicas. Valores en paréntesis indican % del control. El

test “t” de Student muestra que los valores tratados son significativamente diferentes de los del control

(P<0.05). El análisis de varianza de dos vías demuestra que los cambios dependen de la dosis y el

tiempo. Se usaron 600 mg de tejido nervioso y del pie en todas las estimaciones.

Tissues 40% of LCso (48h) 60% of LCso (48h)

Control Brain 2.49+0.24 (100) 2.49+0.24 (100)

Foot 1.61+0.06 (100) 1.61%0.06 (100)

24h Brain 3.00+0.07 (120.48) 3.33+0.13 (133.73)

Foot 1.89+0.02 (117.39) 2.41+0.04 (149.68)

48h Brain 3.28+0.065 (131.72) 3.61+0.053 (144.97)

Foot 2.01+0.06 (124.84) 2.64+0.05 (163.97)

12h Brain 3.64+0.060 (146.18) 4.19+0.058 (168.27)

Foot 3.16+0.039 (196.27) 3.75+0.11 (232.91)

96h Brain 5.18+0.04 (208.03) 6.42+0.07 (257.83)

Foot 4.73+0.17 (293.78) 6.40%0.30 (397.51)

40% and 60% LCso (48h) of C. tiglium

caused a significant change in levels of

phospholipids (Table I) and activity of

lipid peroxidation (Table II) in the

nervous and foot tissues of the snails.

Treatment with 40% LCso of latex extract

of C. tiglium for 24h, 48h, 72h and 96h

decreased the levels of phospholipids in

the brain tissues up to 62.1, 60.99, 55.96

and 39.55 % of the control, respectively

(Table I). Likewise in foot tissue, it

decreased to 85.67, 71.15, 53.73, and 45.83

% of the control respectively (Table ID).

With exposure to a higher dose i.e. 60% of

48h LCso for 24h, 48h, 72h and 96h, there

was a further decrease in the endogenous

levels of phospholipids (Table ID.

The rate of lipid peroxidation in the

brain and foot tissues of control snails

was 2.53 and 1.66 mol malonaldialde-

hyde formed/ 30min/ mg, respectively.

Exposure to the two doses of lattices of

C. tiglium increased the rate of lipid per-

oxidation in both the nervous and foot

tissues. Exposure to 40% of LC5o for 24h,

28

48h, 72h and 96h increased this to

120.48, 131.72, 146.18 and 208.03 % of

control in brain tissue and to 117.39,

124.84, 196.27 and 293.78 % of control in

foot tissue, respectively. Exposure to

60% of LCso (48h) caused a greater

increase in the rate of lipid peroxidation

(Table ID). Analysis of variance demon-

strated that the increase in the rate of

lipid peroxidation was time as well as

dose dependent (P<0.05).

DISCUSSION

The data obtained indicate that

exposure of Lymnaea acuminata to latex of

C.tiglium, increased the rate of lipid per-

oxidation along with decrease in the

endogenous levels of phospholipids in

the nervous and foot tissue of the snail.

That these effects were indeed caused

by lattices of C. tiglium is evidenced by

the fact that both changes were time as

well as dose dependent.

SINGH ET AL.: Latex of Croton tiglium changes lipid peroxidation in Lymnaea acuminata

Biological membranes are known to

be rich in polyunsaturated fatty acids. In

fact, lipids may account for 28% to 79%

of the mass of cell membranes. While

the inner mitochondrial membrane con-

tains about 60% proteins and only 20%

lipids, the myelin sheath of the brain

contains as much as 80% lipids and only

20% protein. Most snail membranes,

however, contain nearly equal amounts

of protein and fat. Membrane lipids are

mainly phospholipids and most of them

are polyglycerides. Up to 20% of the

phospholipids are acidic which are neg-

atively charged and are associated with

membrane proteins due to lipid protein

interaction (LEHNINGER, 1987).

Several workers reported that syn-

thetic / plant pesticides cause repetitive

discharges due to prolongation of sodium

current and increased depolarization

(NARAHASHI, 1983; UPRETI ET AL, 1991;

SIDDIQUI, SAYEED, ZAFAR AND ISLAM, 2002).

The site of action is most likely to be

within the lipophilic environment of the

membrane in the neighbourhood of

sodium proteins (OSBORNE AND SMALL-

COMBE, 1983; PATEL, FULLONE AND

ANDERS, 1992). Any change in the struc-

ture of the concentration of lipids would

radically alter the structure of membranes,

bringing about a change in their perme-

ability characteristic. Indeed, lipid perox-

idation has been reported to play an

important role in a wide variety of patho-

logical and degradative conditions (Bus

AND GIBSON, 1979).

During lipid peroxidation, oxygen

reacts with polyunsaturated lipids to

form lipid radicals and semistable hydro-

peroxides. During this process hydrogen

is abstracted from a carbon-carbon bond

of unsaturated lipid. This process

requires an initiator radical, which may

BIBLIOGRAPHY

Bus, J. S. AND GIBSON, J. E., 1979. Lipid perox-

idation and its role in toxicity. In: Hodgson,

E., Bend, J. R. and Philpot, R. M. (eds) Review

in biochemical Toxicology 1: 125-149.

come from the metabolic degradation of

an external agent (TAPPEL, 1973; ZIA AND

IsLam, 2000). According to BUS AND

GIBSON (1983) certain classes of xenobi-

otics such as parquets and inorganic sub-

stances such as carbon tetrachloride lose

a single electron but under aerobic condi-

tions get re-oxidized immediately to their

original compound with the concurrent

formation of a super-oxide radical. Toxic-

ity of these compounds is due to their

ability to initiate the generation or induc-

tion of this superoxide anion radical. A

similar mechanism may be possible in

the case of L. acuminata treated with lat-

tices of C. tiglium. Formation of activated

oxygen can have extremely detrimental

consequences not only for phospholipids

but also for proteins, nucleic acids and

polysaccharides (BUS AND GIBSON, 1983;

SINGH ET AL., 1996). Even through the

mechanism of formation of superoxide in

the case of C. tiglium is not known yet, it

is possible that the metabolic degradation

of this extract produces an initiator

organic radical which is capable of start-

ing the peroxidation reaction. It is clear

that the latex extract of Croton tiglium is

highly effective at a lower dose and may

eventually prove to be a very useful mol-

luscicide .. Their effectiveness / attractive-

ness as molluscicides in comparison to

synthetic pesticides lie in their plant

origin, which makes them inexpensive

and more easily biodegradable and safe

for non-target organismos.

ACKNOWLEDGEMENTS

One of the authors (Digvijay Singh)

is thankful to CSIR, New Delhi (San.

Letter No. 9/57 (163)/2K2/ EMR - I

dated 4.9.2002) for financial support.

Bus, J. S. AND GIBSON, J. E., 1983. Oxygen acti-

vation and lipoperoxidative metabolism of

toxicity of pesticides and other xenobiotics.

In: Miyamoto, J. and Kerney, P. C. (eds) Pes-

ticides Chemistry. Human welfare and Envi-

ronment. Pergmon Press, London € New

York, 3 Pp-457 - 62.

29

Iberus, 23 (1), 2005

CHVAPIL, M., RYAN, J. N. AND BRADA, Z., 1972.

Effect of selected chelating agents and met-

als on stability of liver lÍysosome. Biochemistry

and Pharmacology 21:1097-1105.

Devi U., 1997. Heavy metal toxicity to gastro-

pod, Morula granulata: tolerance to copper,

mercury, zinc and cadmium. Journal of En-

vironmental Biology 18 (3): 287-290.

DITTMER, J. C. AND MICHAEL, A. W., 1965.

Quantitative and qualitative analysis of

lipids and lipid components. In: methods

of Enzymology. Colowick, S. P. and

Kaplam, N. O. (eds). Vol XIV. Academic

Press. Pp - 483.

FiskE, C. H. AND ROW, Y., 1925. Quantitative and

qualitative analysis of lipids and lipid com-

ponents. In: methods of Enzymology. Colow-

- ick, S. P. and Kaplam, N. O. (eds). Vol. XIV.

Academic Press. Pp 483.

GODAN, D., 1983. Pest sludge and snail, Biology

and control (ed., Dora Godan) translated by

Sheila Gruber, Springer verlag, Berlin, Hei-

delberg, New York.

LEHNINGER, A. L., 1987. Principles of Bio-

chemistry. 2nd Indian reprints. CBC Pub-

lisher and Distributors Delhi Pp - 1011

MARSTON, A. AND HOSTETTMANN, K., 1985.

Plant molluscicide. Phytochemistry 24: 639-

652.

NARAHASHI, T., 1985. Nerve membrane ionic

channels as the primary target of pyrethroids.

Neuro toxicology 6: 3-22.

OSBORNE, P. AND SMALLCOMBE, A., 1983. Site

of actions of pyrethroid insecticides In:

Miyamoto, J. and Kerney, P. C. (eds). Human

welfare and Environment. Pergmon Press,

London and New York Pesticides Chemistry

NON Z:

PATEL, N., FULLONE, J. AND ANDERS, M. W.,

1992. Brain uptake and metabolism of S-(1,2-

dichlorovenyl) glutathione (DCVG) and S-

(1,2-Dichlorovinyl)-L cystine (DCVC). Toxi-

cology 12: 343-346.

SASTRY, K. V. AND SHUKLA, V., 1993. Uptake and

distribution of cadmium in tissue of Channa

marulius. Journal of Environmental Biology 14

(2): 137 - 142.

SIDDIQUI, A., SAYEED, L, ZAFAR, K. S. AND ISLAM,

F., 2002. Argemone oil augmented oxidative

stress in discrete areas of rat brain. Bulletin

of Environmental Contamination and Toxicology

69: 734 - 740.

SINGH, A. AND AGARWAL, R. A., 1992. Mollus-

cicidal activity of euphorbiales against the

snail Indoplanorbis exustus. Acta Hydrochimica

et Hydrobiogica 20: 262- 264.

SINGH, A., SINGH, D. K. AND AGARWAL, R. A.,

1993. Effect of cypermethrin, mexacarbate

and phorate on phospholipid and lipi-

deroxidation in the snail Lymnaea acuminata.

Bulletin of Environmental Contamination and

Toxicology 51: 68-71.

30

SINGH, A. SINGH, D. K., MISRA, T. N. AND AGAR-

WAL, R. A., 1996. Molluscicides of plant or-

gin. Biological Agriculture and Horticulture 13:

205-252.

SINGH, D. AND SINGH, A., 2002. Piscicidal effects

of some common plants of India used in

freshwater bodies against target animals.

Chemosphere 49 (1): 45-49.

SINGH, D. K. AND AGARWAL, R. A., 1986. Piper-

onyl butaoxide synergism with two synthetic

pyrethroids against Lymnaea acuminata.

Chemosphere 15: 493-498.

SINGH, D. K. AND AGARWAL, R. A., 1984a. Al-

teration of biogenic level in the snail Lymnaea

acuminata by the latex of Euphorbia royleana.

Toxicological Letters 21: 309-314.

SINGH, D. K. AND AGARWAL R. A., 1984b. Cor-

relation of the anti-cholinesterase and mol-

luscicidal activity of the latex of Euphorbia

royleana Bioss. on Lymnaea acuminata. Journal

of Natural Product 47: 702-705.

SINGH, A. AND AGARWAL, R. A., 1990. Mollus-

cicidal and Anti-Cholinesterase activity of

euphorbiales. Biological Agriculture and Hor-

ticulture 7: 81-91.

SINGH, D. K., SINGH, A. AND AGARWAL, R. A.,

1993. Nerium indicum as a potent molluscicide

of plant origin. Journal of Medical and Applied

Malacology 5: 93-95.

SINGH, K. AND SINGH, D. K., 1995. Effect of

Azadirachta indica (Neem) on biochemical pa-

rameters in ovotestis of Lymnaea acuminata.

Malaysian Applied Biology 24: 7-11.

SINGH, S. K. AND SINGH, A., 2003. Effect of the

plants Thevetía peruviana and Alstonia schol-

aris (Family: Apocynaceae) on acetyl-

cholinesterase activity of Lymnaea acuminata

snails. Egyptian Journal of Schistosomiasis In-

fectious Ey Endemic Disease 25:

SOKAL, R. R. AND ROHLE, F. J., 1973. Introduc-

tion to bio-statistic edited by Freeman, M. N.,

San Francisco. Pp. 368.

TAPPEL, A. L., 1973. Lipid peroxidation damage

to cell component. Federation Proceedings 32:

1870-1874.

TRIPATHI, P. K. AND SINGH, A., 2003. Toxic ef-

fects of dimethoate and carbaryl pesticides

on protein metabolism of freshwater snail

Lymnaea acuminata. Bulletin of Environmental

Contamination Toxicology 70: 146-152.

UPRETIL, K. K., DAS, M. AND KHANNA, $. K.,

1991. Biochemical toxicology of Argemone

oil: effect on hepatic cyt P-450 and Xenobi-

otic metabolizing enzymes. Journal of Applied

Toxicology 11: 203-209.

UTELY, H. C., BERMHEIM, F. AND HOCHTEIN, P.,

1967. Effects of sulfhydryl reagent on per-

oxidation in microsome. Achieves of Bio-

chemistry and Biophysics 188: 29-32.

YADAV, R. P. AND SINGH, A., 2001. Environ-

mentally safe molluscicide from two common

Euphorbiales. Iberus 19 (1): 65-73.

SINGH ET 4L.: Latex of Croton tiglium changes lipid peroxidation in Lymnaea acuminata

YADAV, R. P. AND SINGH, A., 2002. Toxic effect ZIA, S. AND ISLAM, F., 2000. Selenium altered the

of Croton tiglium on L. acuminata and non-tar- levels of lipids, lipid peroxidation and

get fish Channa punctatus. Iberus 20 (2): 31-44. sulfhydryl group in straitum and thalamus

YADAV, R. P., SINGH, S. K. AND SINGH, A., 2002. of rat Biology of Trace Element Research 77:

Molluscicidal activity of Codiaeum variega- 251-259.

tum, effects on snail metabolism. Journal of

Ecophysiology and Occupational Health 2: 73 -

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O Sociedad Española de Malacología —__—_——T— Iberus, 23 (1): 33-42, 2005

Cephalopod assemblages caught by trawling along the

Southern Tyrrhenian Sea (Central Mediterranean)

Asociaciones de cefalópodos capturados por la pesca de arrastre

comercial en el Tirreno meridional (Mediterráneo central)

Daniela GIORDANO*, Teresa BOTTARI* and Paola RINELLI*

Recibido el 26-11-2004. Aceptado el 27-IX-2004

ABSTRACT

The teuthofauna assemblages of the Southern Tyrrhenian Sea are identified utilising data from

five trawl surveys. A total of 139 hauls were carried out in late spring or the middle of sum-

mer between 1995 and 1999, and 25 cephalopod species were found between 18 and

652 m of depth. The assemblages were analysed with the Bray-Curtis similarity index. Depth

showed a positive correlation with cephalopod distribution. Four main cephalopod groups

were defined: inshore (10-100 m), shelf (101-200 m), slope (201-600 m) and midslope (350-

600 m). Loligo media was the main species for the first group, lllex coindetii for the second,

Todaropsis eblanae for the third group and Todarodes sagittatus for the last group.

RESUMEN

Se identifican las asociaciones de teutofauna capturadas por los arrastreros comerciales en

el Tirreno meridional. Se analiza la fauna de cefalópodos recogida en 139 lances realiza-

dos en primavera o mediados del verano durante cinco campañas de pesca de arrastre rea-

lizadas entre 1995 y 1999. Se identificaron 25 especies de cefalópodos capturadas entre

18 y 652 m de profundidad. El análisis de las asociaciones faunísticas se realizó mediante

el índice de similitud de Bray-Curtis. La profundidad tiene una correlación positiva con la

distribución de los cefalópodos, pudiéndose definir cuatro grupos principales: costero o lito-

ral (10-100 m), de la plataforma (101-200 mj), del talud (201-600 m) y del talud medio

(350-600 m). Loligo media fue la principal especie del primer grupo, lllex coindetiidel segundo,

Todaropsis eblanae del tercero, y Todarodes sagittatus del último grupo.

KEY WORDS: Biogeography, cephalopods, trawl fishery, Tyrrhenian Sea, faunal assemblages, Mediterranean Sea.

PALABRAS CLAVE: Biogeografia, cefalópodos, pesquería de arrastre, asociación faunistica, Mar Tirreno,

Mediterráneo.

INTRODUCTION

The geographic and bathymetric dis- QUETGLAS AND SÁNCHEZ, 1998; SORIANO,

tributions of demersal cephalopods have SÁNCHEZ LIZASO AND GUERRA, 2003; VIL-

been studied in detail in different Mediter- LANUEVA, 1995); the coasts of Libya and

ranean areas: Western Mediterranean Sea Tunisia (BONNET, 1973); Marmara Sea

(SARTOR, BELCARI, CARBONELL, GONZÁLEZ, (KATAGAN, SALMAN AND BENLI, 1993;

* Istituto per Ambiente Marino Costiero IAMC-CNR, Messina Section, Spianata San Raineri 86, 98122

Messina, Italy. Corresponding author: Daniela Giordano. E-mail: daniela.giordanoCiamc.cnr.it

33

Iberus, 23 (1), 2005

UNSAL, UNSAL, ERK AND KABASAKAL,

1999); Eastern Mediterranean Sea (RUBY

AND KNUDSEN, 1972; KNUDSEN, 1981;

SALMAN, KATAGAN AND BOLETZKY, 1999);

Ligurian Sea (RELINI, DE RossI, PIANO AND

Z_AAMBONI, 2002; BERTULETTI AND ORSI

RELINI, 1986; ORSI RELINI AND BERTULETTI,

1989; WURTZ, 1979; OrSI RELINI, 1995); Sar-

dinian Sea (BONNET, 1965; CUCCU, ADDIS,

DAMELE AND MANFRIN PICCINETTI, 2003);

Tyrrhenian Sea (BERDAR, POTOSCHI, CAV-

ALLARO, CAVALIERE AND LI GRECI, 1983;

BERDAR AND CAVALLARO, 1975; LUMARE

1968, 1970; BELCARI, BIAGI, BIAGI, DE

RANIERI, MORI AND PELLEGRINI, 1986;

MANNINI AND VOLPI, 1989; BELCARI AND

SARTOR, 1993; BELCARI, SARTOR AND DE

RANIERI, 1998; WURTZ, MATRICARDI AND

BELCARI, 1992; SPEDICATO, MINETTI,

REBORA, MATRICARDI AND WURTZ, 1990;

GIORDANO AND PERDICHIZZI, 1998; BELLO

AND ARCULEO, 1994; GIORDANO AND CAR-

BONARA, 1999); Sicilian Channel (JEREB

AND RAGONESE, 1991, 1994; RAGONESE,

JEREB AND DI STEFANO, 1992); Adriatic Sea

(BELLO AND MOTOLESE, 1983; GUESCINI

AND PICCINETTI MANERIN, 1986; GAMULIN-

BRIDA AND ILIJANIC, 1972; CASALI,

MANFRIN PICCINETTI AND SORO, 1998;

BELLO, 1990; PASTORELLI, VACCARELLA AND

DE Zio, 1995); lonian Sea (PANETTA,

D'ONGHIA, TURSI AND CECERE, 1986; 'TURSI

AND D'ONGHIA, 1992; D'ONGHIA, MATAR-

RESE, TURSI, MAIORANO AND PANETTA,

1995); Catalan Sea (MANGOLD-WIRZ, 1963;

SÁNCHEZ , BELCARI AND SARTOR, 1998);

and Aegean Sea (D'ONGHIA ET AL., 1995;

SALMAN, KATAGAN AND BENLI, 2002).

In particular, assemblages of

cephalopod communities have been

studied in different areas of this basin

(UNGARO, MARANO, MARSAN, MARTINO,

MARZANO, STRIPPOLI AND VLORA, 1999;

TSERPES, PERISTERAKI, POTAMIAS AND

TSIMENIDES, 1999; (GONZÁLEZ AND

SÁNCHEZ, 2002; QUETGLAS, CARBONELL

AND SÁNCHEZ, 2000). Information about

spatio-temporal distributions of these

species and their linkage to environ-

mental factors is necessary for the study

of many commercial species. The

studies focused on demersal assem-

blages and in particular on their persis-

tence, such as the capability to maintain

34

the same specific composition during a

lag time are useful even for manage-

ment purposes (BIAGI, SARTOR, ARDIZ-

ZONE, BELCARI, BELLUSCIO AND SERENA,

2002).

In this respect, this work analyses

the structure of cephalopod assem-

blages, providing information concern-

ing their distribution for all four groups

defined: inshore, shelf, slope and mids-

lope.

MATERIAL AND METHODS

The study area is situated in the

Southern part of the Tyrrhenian Sea

(central Mediterranean) between Suvero

Cape (Calabria) and San Vito Cape

(Sicily). The data here reported come

from five trawl surveys carried out from

1995 to 1999, during the MEDITS project

(International bottom trawl survey in

the Mediterranean Sea). The experimen-

tal surveys were always conducted

during late spring or the middle of

summer. Hauls were carried out from

one hour before dawn until one hour

after sunset. A fishing vessel equipped

with an experimental trawl net with 20

mm stretched mesh size in the cod-end,

and 2-2.5 m of vertical opening (FIOREN-

TINI, COSIMI, SALA, LEONORI AND

PALOMBO, 1999) was used. A total of 139

hauls were carried out, randomly allo-

cated into five bathymetrical strata: “A”:

10-50 m; “B”: 51-100 m; “C”: 101-200 m;

“D”:> 201-500. m; +*E*%%9 5UESO0B

(BERTRAND, GIL DE SOLA, PAPACONSTAN-

TINOU, RELINI AND SOUPLET , 2002).

The duration of each haul was 30

minutes at depths less than 200 m and

one hour at greater depths. All the data

were standardised to one hour haul

duration. All cephalopods were identi-

fied and counted on board. A multivari-

ate approach, on the basis of the calcula-

tion of a triangular similarity matrix

(group-average linkage), by depth, year

and abundance (number of speci-

mens/hour), according to BRAY AND

CURTIS (1957), with relative dendrogram

and non metric plan Multi Dimensional

Scaling (MDS) was elaborated (CLARKE

GIORDANO £7 AL.: Cephalopod assemblages along the Southern Tyrrhenian Sea

0 Bray-Curtis Similarity

20

40

60

80

100

AAA AA

100 m 200 m

litoral shelf slope

Figure 1. Dendrogram showing similarities between hauls for the 1996 surveys. Mean depth of

each haul is presented.

Figura 1. Dendrograma que muestra las similitudes entre los arrastres de 1996. Se indica la profundi-

dad media de cada arrastre.

AND WARWICK, 1994). Because the

annual variations of cephalopod distrib-

ution were not significant, the five

surveys were pooled together to calcu-

late the total cluster. Hauls in which

only one species was caught and the

species with abundance values lower

than 3 were not considered for the

analysis. To establish which taxa con-

tributed most to the separation of one

group from another, the SIMPER routine

was used (CLARKE, 1993). Abundance

values were also analysed by means of

univariate indices, in relation to the four

assemblages evidenced by cluster analy-

sis: total mumber of taxa (S), total

number of individuals (N), richness of

Margalef (d), Shannon-Wiener diversity

(H”) and Pielou's evenness (J) indices.

The cumulative abundances were illus-

trated in order of dominance as K-domi-

nance curves.

RESULTS

Twenty-five species of cephalopods

were collected belonging to three orders

and eight families. In Table I the system-

atic list of species, drawn up according

to BELLO (1986), within the bathymetri-

cal range of each species, is reported. In

particular, Onychoteuthis banksii,

Ancistroteuthis lichtensteini and Abralia

veranyi were caught only once during

the period studied.

The cephalopods collected were dis-

tributed between 18 and 652 m of depth.

The more coastal species was repre-

sented by Sepia officinalis, confined

within the depth of 68 m. Fifty percent

of the species showed a wide bathymet-

rical range that included the shelf and

the beginning of the slope. The widest

distribution was shown by Eledone cir-

rhosa (72-584 m), Scaeurgus unicirrhus

(38-549 m), Todaropsis eblanae (61-613 m)

and Pteroctopus tetracirrhus (118-633 m).

Alloteuthis subulata, Octopus vulgaris and

Eledone moschata were found only on the

continental shelf. On the contrary,

Octopus salutii and Rossia macrosoma

were caught at over 200 m of depth. His-

tioteuthis bonnellii was found exclusively

in the deepest stratum.

The dendrogram of all surveys (Fig.

1) showed three principal clusters: l,

inshore (<100 m); IL, shelf (80 — 200 m)

and III, slope (200 — 600 m). The coastal

group was characterized by Loligo media

39

Iberus, 23 (1), 2005

Table I. Cephalopod species captured from 1995 to 1999 off the Southern Tyrrhenian Sea, showing

their depth range and mean depth of occurrence with the associated standard deviation (SD).

Tabla 1. Especies de cefalópodos capturadas desde 1995 a 1999 en el mar Tirreno meridional, indicando

su rango de distribución batimétrica, profundidad media y la desviación estándar asociada (SD).

Min Max Media SD

Class CEPHALOPODA Cuvier, 1798

Subclass COLEOIDEA Bather, 1888

Order SEPIOIDEA Naef, 1916

Family Sepiidae Leach, 1817

Genus Sepia Linnaeus, 1758

Sepia officinalis Linnaeus, 1758 40 68 495 12.66

Sepia orbignyana Férussac in Orbigny, 1826 61 432 185.5 96.87

Sepia elegans Blainville, 1827 40 345 137,3 14.02

Family Sepiolidae Leach, 1817

Subfamily Sepiolinae Leach, 1817

Genus Sepiola Leach, 1817

Sepiola rondeletii Leach, 1817 110 395 276 115

Genus Rondeletiola Naef, 1921

Rondeletiola minor (Naef, 1912) 117 297 207 127.28

Subfamily Rossiinae Appellof, 1898

Genus Rossia Owen in Ross, 1835

Rossia macrosoma (Delle Chiaie, 1830) 2d 596 484.5 115.56

Genus Neorossia Boletzky, 1971

Neorossia caroli (Joubin, 1902) 123 608 309 195:9

Order TEUTHOIDEA Naef, 1916

Suborder MYOPSIDA Orbygny in Férussac 8. Orbigny, 1840

Family Loliginidae Lesueur, 1821

Genus Loligo Lamarck, 1798

Loligo vulgaris Lamarck, 1798 18 29 75.72 12

Loligo forbesi Steenstrup, 1856 12. 505 134.2 127.3

Loligo media (Linnaeus, 1758) 18 E al 15.43

Genus Alloteuthis Wulker, 1920

Alloteuthis subulata (Lamarck, 1798) 16 143 109,5 47.4

Suborder OEGOPSIDA Orbigny, 1845

Family Enoploteuthidae Pfeffer, 1900

Subfamily Enoploteuthinae Pfeffer, 1900

Genus Abralia Gray, 1849

Abralia veranyi (Ruppell, 1844) 310 ES6

Family Onychoteuthidae Gray, 1847

Genus Onychoteuthis Lichtenstein, 1818

Onychoteuthis banksii (Leach, 1817) 358 358

Genus Ancistroteuthis Gray, 1849

Ancistroteuthis lichtensteinii (Férussac, 1839) 562 562

Family Histioteuthidae Verrill, 1881

Genus Histioteuthis Orbigny, 1840

Histioteuthis bonnellii (Férussac, 1834) 53] 645 565.5 53.33

Histioteuthis reversa (Verrill, 1880) 383 652 565.3 89.4

Family Ommastrephidae Steenstrup, 1857

Subfamily Illicinae Posselt, 1890

Genus /llex Steenstrup, 1880

Illex coindetii (Verany, 1839) 60 NES 128.7

36

GIORDANO ET AL.: Cephalopod assemblages along the Southern Tyrrhenian Sea

Table I. Continuación.

Tabla I. Continuation.

Genus Jodaropsis Girard, 1890

Min Max Media SD

Todaropsis eblonae (Ball, 1841) 61 0135.3235] 152.2

Sufamily Todarodinae Adam, 1960

Genus Jodarodes Steenstrup, 1880

lodarodes sagittatus (Lamarck, 1798) 114 593 438.5

Order OCTOPODA Leach, 1818

Suborder INCIRRATA Grimpe, 1916

154.69

Family Octopodidae Orbigny in Férussac 8. Orbigny, 1840

Subfamily Octopodinae Orbigny in Férussac 8. Orbigny, 1840

Genus Octopus Cuvier, 1798

Octopus vulgaris Cuvier, 1798

Octopus salutii Vérany, 1836

Genus Scaeurgus Troschel, 1857

27 114 62.86 25.85

MIL ESTA 10.87

Scaeurgus unicirrhus (Delle Chiaje in

Férussac 8. Orbigny, 1840) 00 549 1747 110.8

Genus Pteroctopus P. Fischer, 1882

Pteroctopus tetracirrhus (Delle Chiaje, 1830) UA A 147.83

Subfamily Eledoninae Grimpe, 1921

Genus Eledone Leach, 1817

Eledone moschata (Lamarck, 1798) 18 143 115.4 92.82

Eledone cirrhosa (Lamarck, 1798) IDEA ZOO OY

and Loligo vulgaris. In the shelf group

(Table lla) Illex coindetíí and Loligo media

prevailed, followed by Sepia orbignyana,

Sepia elegans and Scaeurgus unicirrhus.

The slope group (Table Ilb) was charac-

terized by Todaropsis eblanae. The Multi-

dimensional Scaling analysis (30%:

Stress 0.15) also showed four groups,

well discriminated by depth (Fig. 2).

There is also a midslope group (Fig. 3),

limited to a few hauls, in which Todaro-

des sagittatus was the prevalent species.

Analysing the univariate indices

(Table IMM), elaborated for the four

groups, the highest biodiversity was

observed in the shelf. In this macrostra-

tum, the highest values of Margalef and

Shannon-Wiener indices were recorded

as well as the highest number of species

and individuals. The highest values of

H' found at this bathymetrical level can

be explained by the relative homogene-

ity in abundance of the most frequent

species.

However, a trend was observed

according to the depth as evidenced also

by the cumulative abundance curve, in

which the shelf group showed the

highest biodiversity and the midslope

the lowest. In this last group in fact both

d and H' values recorded the lowest

values (0.973 and 1.063, respectively).

Finally the Evenness (J') values

ranged from 0.616 to 0.661. The higher

value was recorded for the last cluster.

Also in the second cluster this value is

quite high.

DISCUSSION AND CONCLUSIONS

The 25 cephalopod species distrib-

uted in 8 families recorded in this study

account for 42.4% of species reported

from the Mediterranean (MANGOLD AND

BOLETZKY, 1988; GUERRA, 1992). Absent

were certain species of the Sepiolidae

family and pelagic species. In compari-

37

Iberus, 23 (1), 2005

Table Ia. Indicator species and related data from the SIMPER analysis. Abbreviations: AA, average

abundance, contribution of each species in each group; AT, average term, average Bray-Curtis con-

tribution of each species to distinguish between groups. The ratio (AT/SD), the percentage contri-

bution to the separation (%), and the cumulative percentage (Cum %) are shown for each group

comparison.

Tabla lla. Especies indicadoras y datos relacionados, según el análisis SIMPER. Abreviaturas: AA,

abundancia promedio, contribución de cada especie en cada uno de los grupos; Average Term (AT), con-

tribución promedio de Bray-Curtis de cada especie en la diferenciación de los grupos. Cada grupo viene

definido, además, por la relación AT/SD (ratio), la contribución porcentual (%), y el porcentaje acumu-

lativo de cada especie (Cum %).

Average dissimilarity: 84.1 inshore AA shelf AA

l. vulgaris 106.88 0.90

[. coindetii 0.58 26.62

L. vulgaris 18.88 0.02

S. elegans 0.05 0.41

S. orbignyana 0.00 8.86

S. unicirrhus 0.00 0.29

E. cirrhosa 0.05 0.13

Average dissimilarity: 99 inshore AA shelf AA

L. media 106.88 + 0.00

L. vulgaris | 18.88 0.00

T. eblanae 0.00 0.46

P tetracirrhus 0.00 2.74

0. vulgaris 0.09 0.00

S. orbignyana 0.00 2.89

l. coindetii 0.04 1.67

E. moschato 2.00 0.19

Average dissimilarity: 92.7 inshore AA shelf AA

L. media 21.43 0.00

l. coindetii 26.62 1.67

S. elegans 757 0.00

S. orbignyana 8.86 2.89

T. eblanae 1.14 10.56

S. unicirrhus 6.54 0.85

E. cirrhosa 3.30 0.67

P tetracirhus 0.22 2.14

son, TURSI AND D'ONGHIA (1992) caught

24 species of cephalopods in the lonian

Sea, D'ONGHIA ET AL. (1995) 29 species

in the North Aegean Sea, QUETGLAS ET

AL. (2000) found 30 species in the

Balearic Sea, SÁNCHEZ ET AL. (1998)

found 36 species in the northern

Tyrrhenian Sea and 47 in the Catalan

Sea and SORIANO ET AL. (2003) found 18

species in the upper continental slope of

Alicante (western Mediterranean).

38

AT Ratio % Cum (%)

34.64 1.39 41.18 41.18

0.50 0.67 13.74 54.92

0.47 1.03 13.22 68.13

0.28 0.71 7.70 15.83

5.69 0.53 6.17 82.6

0.21 0.6 5,45 88.05

0.13 0.6 3.57 91.62

AT Ratio % Cum (9)

47.82 1.48 48.3 48.3

0.80 1.07 19.25 67.56

9.70 0.64 98 17.35

3.8] 0.67 3.05 81.2

0.13 0.63 3.17 84.37

2.68 0.32 1 87.08

2.43 0.56 2.46 89.54

2.37 0.76 2.39 91.93

AT Ratio % Cum (9%)

19.36 1.03 20.89 20.89

17.55 0.82 18.94 39.83

10.85 0.82 11.71 MD

10.51 0.69 11.34 62.88

9.67 0.70 10.43 13.31

117 0.69 8.38 81.69

4.95 0.63 5.34 87.04

3.47 0.83 35 90.79

In previous surveys of the study

area before 1995 (GIORDANO AND

PERDICHIZZI, - 1998), 32. speciesimolk

cephalopods were identified. In general,

a similar cephalopod distribution

pattern was observed from 1994-1995

trawl surveys, but the sampling was

carried out at a different period (in

autumn) and with a different net (with

36 mm of stretched mesh). The species

caught in the previous faunistic list in

the same area were: Sepiola affinis,

GIORDANO ET AL.: Cephalopod assemblages along the Southern Tyrrhenian Sea

Table ITb.

Tabla I1b.

Average dissimilarity: 100 inshore AA midslope AA

L. media 106.88 0

L. vulgaris 18.88 0

Í. sagittatus 0 /

0. vulgaris DS 0

S. orbignyana 0 3.6

E. moschata 2 0

Average dissimilarity: 97.18 inshore AA midslope AA

L. media 21.43 0

Í. coindefii 26.62 0

S. elegans 9.57 0

S. orbignyana 0.86 3.6

Í. sagittatus 0.16 /

S. Unicirrhus 6.54 0.2

E. cirrhosa 3 0

E. moschato 0.22 0

Average dissimilarity: 92.38 inshore AA midslope AA

T. eblanae 10.56 0

/. sagittatus 0.3 /

P tetracirrhus 2.81 0.4

S. orbignyana 2.22 3.6

R. macrosoma 0.19 l

l. coindefii 1.67 0

S. unicirrhus 0.85 0.2

E. cirrhosa 0.67 0

Sepiola ligulata, Sepiola robusta, Sepietta

oweniana, Sepietta neglecta (Sepiolidae),

Chiroteuthis veranii (Chiroteutidae) and

Bathypolypus sponsalis (Octopodidae).

On-board observation of catches indi-

cated that the demersal fauna sampled

is strongly influenced by depth. The

AT Ratio % Cum (%)

50.79 1.52 50.79 50.79

ES 1.08 21.3 72.09

9.23 0.85 9.23 81.32

3.63 0.62 3.63 84.94

3.5] 0.42 3.5] 88.46

2.61 0.74 2.61 91.07

AT Ratio % Cum (%)

20.96 1.05 21.57 ES

18.87 0.81 19.42 40.98

11.87 0.84 12.21 53.19

11.6 0.72 11.93 65.13

8.66 0.99 8.9] 74.04

8.4] 0.68 8.66 82.7

5.64 0.61 5.8 88.5

2.65 0.64 Di 91.22

AT Ratio % Cum (0)

3805 0.95 DS IS

NDS 18 23.33 48.5

11.48 0.96 12.42 60.92

10.36 0.57 11.21 712.14

6.78 0.6 7.34 79.48

5.67 0.6 6.14 85.61

3.49 0.57 3.78 89.39

2.93 0.46 35 92.56

depth range of each species with mean

depth and standard deviation are

reported in Table I. Eledone cirrhosa (32-

584 m), Scaeurgus unicirrhus (38-549 m),

Todaropsis eblanae (61-613 m), Sepia

elegans (40-432 m), Loligo media (18-383

m) and Illex coindeti (60-593 m) showed

Table III. Univariate diversity indexes for each cluster. Abbreviations: S, number of species; N,

number of specimens; d, richness of Margalef; J, Pielow's evenness; H”, Shannon-Wiener diversity.

Tabla UI Índices univariados de diversidad de cada grupo. Abreviaturas: S, número de especies; N,

número de ejemplares; d, riqueza según Margalef; J, uniformidad según Pielou; H”, diversidad de

Shannon- Wiener.

Cluster S N

inshore 8 709

shelf 15 2267

slope 12 529

midslope 5 61

d J H

1.066 0.616 1.28

1.812 0.652 1.765

1.754 0.019 1.539

0.973 0.661 1.063

9

Iberus, 23 (1), 2005

Stress: 0.15

MX 18-80 m

V 196-283 m

O 204-600 m

<> 366-585 m

Figure 2. Multidimensional scaling ordination analysis for the five surveys.

Figura 2. Análisis de ordenación de escalamiento multidimensional para las cinco campañas.

a wide distribution range, extending up

to the continental slope.

During the present study four main

cephalopod groups were defined:

inshore (10-100 m), shelf (101-200 m),

slope (201-600 m) and midslope (350-

600 m). Loligo media was the main

species for the first group, Illex coindetit

for the second, Todaropsis eblanae for the

BIBLIOGRAPHY

BELCARI, P., BIAGI, F., BIAGI, V., DE RANIERL S.,

MORI, M. AND PELLEGRINI, D., 1986. Obser-

vations about Cephalopods distribution in the

north Tyrrhenian sea. Rapport de la Commis-

sion Internationale pour l'exploration scientifi-

que de la Mer Méditerranée, 30 (2): 246.

BELCARI, P. AND SARTOR, P., 1993. Bottom traw-

ling teuthofauna of the North Tyrrhenian

sea. Scientia Marina, 57 (3): 145-152.

BELCARI, P., SARTOR, P. AND DE RANIERL S., 1998.

I cefalopodi nello sbarcato commerciale con

reti a strascico nel Mar Tirreno settentrionale.

Biologia Marina Mediterranea, 5 (2): 318-325.

BELLO, G., 1986. Catalogo dei molluschi cefa-

lopodi viventi nel Mediterraneo. Bollettino

Malacologico, 22: 197-214.

BELLO, G., 1990. The Cephalopods fauna of the

Adriatic. Acta Adriatica, 31: 275-291.

BELLO, G. AND ARCULEO, M., 1994. I cefalopodi

dei fondi strascicabili del Golfo di Caste-

llammare. Bollettino Malacologico, 30 (5-9):

173-181.

40

third group and Todarodes sagittatus for

the last group.

ACKNOWLEDGEMENTS

We are grateful to Dr. Patrizia Jereb

for helpful comments on the manu-

script.

BELLO, G. AND MOTOLESE, G. 1983. Sepiolids

from the Adriatic Sea (Mollusca, Cephalo-

poda). Rapport de la Commission Internatio-

nale pour l'exploration scientifique de la Mer

Méditerranee, 28 (5): 281-282.

BERDAR, A., POTOSCHI, A., CAVALLARO, G., CA-

VALIERE, A. AND LI GRECI, F., 1983. Su alcuni

cefalopodi spiaggiati e pescati nello Stretto

di Messina. Memorie di Biologia Marina ed Oce-

anica, 13 (2): 115-127.

BERDAR, A. AND CAVALLARO, G., 1975. Ulte-

riore contributo alla conoscenza dei Cefalo-

podi spiaggiati lungo la costa siciliana dello

Stretto di Messina. Memorie di Biologia Marina

ed Oceanica, 5 (5):121-138.

BERTRAND, J. A., GIL DE SOLA, L., PAPACONS-

TANTINOU, C., RELINI, G. AND SOUPLET, A.,

2002. An international bottom trawl in the

Mediterranean: the MEDITS programme.

1995 ICES Annual Science Conference, CM

1997 /y:03: 16 pp.

GIORDANO ET AL.: Cephalopod assemblages along the Southern Tyrrhenian Sea

BERTULETTI, M. AND ORSI RELINI, L., 1986. No-

tes on the presence and distribution of the Se-

piolinae in the Ligurian Sea. Rapport Com-

mission Internationale pour l'exploration Scien-

tifique de la Mer Méditerranée, 30 (2): 247.

BIAGI F., SARTOR, P., ARDIZZONE, G., BELCARI,

P., BELLUSCIO, A. AND SERENA, F., 2002.

Analysis of demersal assemblages off the

Tuscany and Latium coasts (north-western

Mediterranean). Scientia Marina, 66 (supp!l. 2):

233-242.

BONNET, M., 1973. Les céphalopodes capturés

par la Thalassa en Novembre 1969 au large

de la Libye et de la cóte orientale tunisienne.

Revue des travaux de l'Institut des Péches Ma-

ritimes, 37 (2): 253-256.

BONNET, M., 1965. Remarques sur l'ecologie

des Céphalopodes des cotes de Sardegne et

de Corse capturés par la “Thalassa” en no-

vembre et dicembre 1963. Rapport de la Com-

mission Internationale pour l'exploration scien-

tifique de la Mer Méditerranee, 18: 234-240.

BRAY, J. R. AND CURTIS, J. T., 1957. An ordina-

tion of the upland forest communities of the

Southern Wisconsin. Ecological Monographs,

27: 325-349

CASALI, P., MANFRIN PICCINETTI, G. AND SORO,

S., 1998. Distribuzione di cefalopodi in alto

e medio Adriatico. Biologia Marina Medite-

rranea, 5 (2) : 307-317.

CLARKE, K. R., 1993. Non parametric multiva-

riate analyse of changes in community struc-

ture. Australian Journal of Ecology, 18: 117-

143. ,

CLARKE, K. R. AND WARWICK, R. M., 1994.

Change in marine communities: an approach to

statistical analysis and interpretation. Plymouth

Marine Laboratory, Plymouth, 144 pp.

CuUccU, D., ADDIS, P., DAMELE, F. AND MANFRIN

PICCINETTI, G., 2003. Primo censimento de-

lla teutofauna dei mari circostanti la Sar-

degna. Biologia Marina Mediterranea, 10

(2):795-798.

D'ONGHIA, G., MATARRESE, A., TURSI, A., MAIO-

RANO P. AND PANETTA, P., 1995. Osserva-

zioni sulla teutofauna epi e mesobatiale nel

Mediterraneo Orientale (Mar lonio e Mar

Egeo). Biologia Marina Mediterranea, 2 (2): 199-

204.

FIORENTINI, L., COsIMI, G., SALA, A., LEONORI,

I. AND PALOMBO, V., 1999. Efficiency of the

bottom trawl used for Mediterranean inter-

national trawl survey (MEDITS). Aquatic Li-

ving Resources, 12 (3): 187-205.

GAMULIN-BRIDA, H. AND ILIJANIC, V., 1972. Con-

tribution a la connaissance des Céphalopodes

del'Adriatique. Acta Adriatica, 14 (6) : 12 pp.

GIORDANO, D. AND CARBONARA, P., 1999. Nota

sulla distribuzione di molluschi cefalopodi

nel Tirreno centro-meridionale. Biologia Ma-

rina Mediterranea, 6 (1): 573-575.

GIORDANO, D. AND PERDICHIZZI, F., 1998. Os-

servazioni sulla teutofauna nel Tirreno me-

ridionale da Capo Suvero (Calabria) a Capo

S. Vito (Sicilia). Biologia Marina Mediterranea,

5 (1): 807-810.

GONZÁLEZ, M. AND SÁNCHEZ, P., 2002. Cepha-

lopod assemblages caught by trawling along

the Iberian Peninsula Mediterranean coast.

Scientia Marina, 66 (Suppl. 2): 199-208.

GUERRA, A., 1992. Mollusca Cephalopoda. En:

Fauna Ibérica, vol. 1. Ramos, M. A. et al. (Ed.).

Museo Nacional de Ciencias Naturales, CSIC,

Madrid, 327 pp.

GUESCINL, A. AND PICCINETTI MANFRIN, G., 1986.

Distribuzione di Sepiolidi nell'Adriatico cen-

tro-settentrionale. Nova Thalassia, 8, Suppl. 3,

pp. 513-518.

JEREB, P. AND RAGONESE, S., 1991. On the asso-

ciation of the squid Illex coindetíi (Mollusca:

Cephalopoda) with the target species traw-

led in the Sicilian Channel. Note Tecniche e Re-

prints — Istituto di Tecnología della Pesca e del

Pescato, 32: 3 p.

JEREB, P. AND RAGONESE, S., 1994. The Medite-

rranean teuthofauna: Towards a biogeo-

graphical coverage by regional census. II.

Strait of Sicily. Bollettino Malacologico, 30 (5-

9): 161-172.

KATAGAN, T., SALMAN, A. AND BENLI, H. A.,

1993. The cephalopod fauna of the Sea of

Marmara. Israel Journal of Zoology, 39: 255-

261.

KNUDSEN, J., 1981. Three Sepiolidae to the

eastern Mediterranean (Mollusca: Cepha-

lopoda). Argamon, Israel Journal of Malaco-

logy, 7: 45-50.

LUMARE, F., 1968. Ricerche sui Cefalopodi

dell'alto Tirreno. Commissione Studi Oceano-

grafici e di Limnologia, Programma di ricerca

sulle risorse Marine e del fondo Marino, C. N. R.

B., 24: 5-29.

LUMARE, F., 1970. Nota sulla distribuzione di al-

cuni cefalopodi del mar Tirreno. Bollettino

Pesca Piscicoltura e Idrobiologia, 25 (2): 313-

344.

MANGOLD, K. AND BOLETSKY, S., 1988. Medite-

rranean Cephalopod Fauna. In: M. R. Clarke

and E. R. Trueman (eds.), Ethology and Ne-

ontology of Cephalopods, The Mollusca, 12:

315-330. Academic Press, San Diego.

MANGOLD-WIRz, K., 1963. Biologie des Cep-

halopodes benthiques et nectoniques de la

Mer Catalane. Vie et Milieu, 13: 1-285.

MANNINI, P. AND VOLPL C., 1989. Nota sulla pre-

senza e distribuzione di alcuni cefalopodi

del Tirreno settentrionale. Oebalia, 25-2, N. S.:

693-701.

OrsI RELINI, L., 1995. Notes on midwater co-

llections of Heteroteuthis dispar (Cephalopoda,

Sepiolidae). Bulletin de l'Institut océanograp-

hique Monaco, n. special 16 : 63-72.

41

Iberus, 23 (1), 2005

ORSI RELINL, L. AND BERTULETTI, M., 1989. Se-

piolinae (Mollusca, Cephalopoda) from the

Ligurian Sea. Vie et Milieu, 39 (3/4):183-

190.

PANETTA, P., D'ONGHIA, G., TURSI, A. AND CE-

CERE, E., 1986. Il ruolo dei Molluschi Cefalo-

podi nella valutazione delle risorse marine

dello Jonio (campagne di pesca 1985). Nova

Thalassia, 8, Suppl. 3:523-527.

PASTORELLI, A. M., VACCARELLA, R. AND DE

Z10, V., 1995. Distribuzione dei Cefalopodi

commerciali del basso Adriatico. Biologia Ma-

rina Mediterranea, 2 (2): 501-502.

QUETGLAS, A, CARBONELL, A. AND SÁNCHEZ,

P., 2000. Demersal continental self and up-

per slope cephalopods assemblages from

the Balearic Sea (North-Western Medite-

- rranean). Biological aspects of some deep-

sea species. Estuarine, Coastal and Shelf

Science, 50: 739-749.

RAGONESE, S., JEREB, P. AND DI STEFANO, L.,

1992. Occurrence of the unicorn octopus Sca-

eurgus unicirrhus in the Sicilian Channel. L

Spatial distribution and abundance. II: Bathy-

metric distribution. Rapport Commission In-

ternationale pour l'exploration Scientifique de la

Mer Méditerranée, 33: 306-307.

RELINI, G., DE ROSSI, C., PIANO, T. AND ZAMBONI

A., 2002. Osservazioni sui Cefalopodi dei

fondi strascicabili liguri. Biologia Marina Me-

diterranea, 9 (1): 792-795.

RELINI, G. AND ORSI RELINI, L., 1984. The role

of cephalopods in the inshore trawl fishing

of the Ligurian Sea. Oebalia, 10: 35-58.

RESTUCCIA, V. AND RAGONESE, S., 1986. Cefa-

lopodi dei fondali strascicabili della Sicilia

nord-orientale: nota preliminare. Nova Tha-

lassia, 8, Suppl. 3:543-549.

RUBY, G. AND KNUDSEN, J., 1972. Cephalopoda

from the Eastern Mediterranean. Israel Jour-

nal of Zoology, 21: 83-97.

SALMAN, A., KATAGAN, T. AND BENLI, H. A.,

2002. Cephalopod Fauna of the Eastern Me-

diterranean. Turkish Journal of Zoology, 26:

47-52.

SALMAN, A., KATAGAN, T. AND BOLETZKY, S. V.,

1999. New cephalopod molluscs in the Eas-

tern Mediterranean: previously unnoted spe-

cies or Lessepsian migrants? Vie et Milieu,

49 (1): 11-17.

SÁNCHEZ, P., BELCARI, P. AND SARTOR, P., 1998.

Composition and spatial distribution of cep-

halopods in two north-western Mediterra-

nean areas. South African Journal of Marine

Science, 20: 17-24.

42

SARTOR, P., BELCARI, P., CARBONELL, A., GON-

ZÁLEZ, M., QUETGLAS, A. AND SÁNCHEZ, P.,

1998. The importance of cephalopods to

trawl fisheries in the western Mediterra-

nean. South African Journal of Marine

Science, 20: 67-72.

SORIANO, S., SÁNCHEZ LIZASO, J. L. AND GUE-

RRA, A., 2003. Cephalopod species collected

in the upper continental slope off Alicante

(Western Mediterranean). Iberus, 21: 57-66.

SPEDICATO, M. T., MINETTI, D., REBORA, F., MA-

TRICARDI, G. AND WURTZ, M. 1990. Distribu-

zione di Cefalopodi Decapodi nel medio-

basso Tirreno. Oebalia, Suppl., 26 (2): 761-766.

TSERPES, G., PERISTERAKI, P., POTAMIAS, G. AND

TSIMENIDES, N., 1999. Species distribution in

the southern Aegean sea based on bottom-

trawl surveys. Aquatic Living Resources, 12

(3): 167-175.

TursI, A. AND D'ONGHIA, G., 1992. Cephalopods

in the lonian Sea (Mediterranean sea). Oeba-

lía, 18: 25-43.

UNGARO, N., MARANO, C. A., MARSAN, R., MAR-

TINO, M., MARZANO, M. C., STRIPPOLI, G. AND

VLORA, A., 1999. Analysis of demersal spe-

cies assemblages from trawl surveys in the

South Adriatic sea. Aquatic Living Resources,

12 (3): 177-185.

UnsaL, I., UNSAL, N., ERK, M. H. AND KABA-

SAKAL, H., 1999. Demersal cephalopods from

the Sea of Marmara with remarks on some

ecological characteristics. Acta Adriatica, 40:

105-104.

VILLANUEVA, R., 1995. Distribution and abun-

dance of bathyal sepiolids (Mollusca: Cep-

halopoda) in the northwestern Mediterra-

nean. Bulletin de l'Institut océanographique,

Monaco, n* special 16: 19-26.

VOLPI, C., AUTERI, R., BORRI, M. AND MANNINI,

P., 1990. Distribution and reproduction of

Sepia elegans in the North Tyrrhenian Sea.

Rapport Commission Internationale pour l'ex-

ploration Scientifique de la Mer Méditerranée, 32

(1): 244.

WURTZ, M., 1979. I Cefalopodi raccolti in Mar

Ligure durante la campagna di pesca batiale

1977/1978. Atti della Societá Toscana di Scienze

Naturali, memorie, ser. B, 86: 374-377.

WURTZ, M., MATRICARDI, G. AND BELCARI, P.,

1992. Distribution and abundance of the Oc-

topus Eledone cirrhosa in the Tyrrhenian sea,

central Mediterranean. Fishery Research, 13 (1):

53-66.

O Sociedad Española de Malacología Iberus, 23 (1): 43-48, 2005

Onchidoris neapolitana (Delle Chiaje, 1844) (Gastropoda:

Nudibranchia: Onchidorididae): una nueva especie de

molusco para la fauna andaluza

Onchidoris neapolitana (Delle Chiaje, 1844) (Gastropoda:

Nudibranchia: Onchidorididae): a new molluscan species for the

Andalousian fauna

Alma SÁNCHEZ-SANTOS*

Recibido el 6-VI1-2004. Aceptado el 24-1-2005

RESUMEN

En este trabajo se redescribe el doridoideo Onchidoris neapolitana (Delle Chiaje, 1844)

a partir de varios ejemplares recolectados en la zona del Estrecho de Gibraltar. Este

hallazgo constituye la primera cita de esta especie en el litoral andaluz y amplía su área

de distribución hacia el oeste.

ABSTRACT

In this paper, the doridoidean nudibranch Onchidoris neapolitana (Delle Chiaje, 1844) is

redescribed on the basis of several specimens collected from the Strait of Gibraltar. This

record constitutes the first one of this species from the Andalousian littoral and extends the

known range of this species to the western.

PALABRAS CLAVE: Nudibranchia, Onchidoris neapolitana, Estrecho de Gibraltar, Sur de España.

KEY WORDS: Nudibranchia, Onchidoris neapolitana, Strait of Gibraltar, Southern Spain.

INTRODUCCIÓN

El género Onchidoris está represen-

tado en el litoral peninsular ibérico por

nueve especies, cuya distribución cono-

cida en dicho ámbito geográfico es reco-

gida por CERVERA, TEMPLADO, GARCÍA-

GÓMEZ, BALLESTEROS, ORTEA, GARCÍA,

Ros Y LUQUE (1988).

Una de estas especies es Onchidoris

neapolitana, descrita por vez primera en

las costas de Italia (localidad tipo

Nápoles) y con una distribución medite-

rránea. En el mar Mediterráneo, además

de la costa italiana (golfo de Nápoles,

SCHMEKEL, 1968; SCHMEKEL Y PORT-

MANN, 1982), las otras citas conocidas

proceden de las costas de Francia

(Banyuls, PRUVOT-FoL, 1951) y España

(Ros, 1975, 1978, 1985; Ros Y ÁLTIMIRA,

1977; PEREIRA Y BALLESTEROS, 1979; ALTI-

MIRA, HUELIN Y ROs, 1981; ORTEA Y

BALLESTEROS, 1982; HUELIN Y Ros, 1984;

BALLESTEROS, 1985), donde todas las

referencias de esta especie restringen su

área de distribución al litoral catalán.

Según CERVERA ET AL. (1988), el litoral

catalán constituye el límite más occiden-

* Urb. San García, C/ Trucha, 85. 11207 Algeciras (Cádiz), Spain; e-mail: almasanchez830 hotmail.com

Á3

Iberus, 23 (1), 2005

conducto uterino

vagina conducto

deferente

ampolla

hermafrodita

ampolla

na B hermafrodita

próstata Y glándula femenina

receptáculo

seminal

bolsa copulatriz

3

Figura 1. Anatomía externa de un ejemplar de 8 mm de Onchidoris neapolitana. A: vista dorsal; B:

vista ventral; C: detalle de un rinóforo; D: disposición de las hojas branquiales y tubérculos intra-

branquiales. Figura 2. Fotografía al microscopio electrónico de barrido de la rádula de un ejemplar

de 7 mm (fijado). Escala: 30 pm. Figura 3. Sistema reproductor de un ejemplar de 8 mm (fijado).

Figure 1. External anatomy of one specimen of 8 mm of Onchidoris neapolitana. A: dorsal view; B:

ventral view; C: detail of a rinophore; D: detail of the arrangement of the branchial leaves and intra-

branchial tubercles. Figure 2. Scanning electron micrographs of the radula of one specimen of 7 mm long

(preserved). Scale bar: 30 ym. Figure 3. Reproductive system of' one specimen of. 8 mm (preservea).

1 mm

tal conocido de su área de distribución.

Así, Ros (1975) cita por primera vez

para el litoral peninsular ibérico a O.

neapolitana como Lamellidoris (Atalodoris)

neapolitana (Delle Chiaje, 1844) en base a

dos ejemplares recolectados en Cada-

qués e Islas Medas (Gerona), asociados

al briozoo incrustante Schizobrachiella

sanguinea (Norman, 1868). Posterior-

mente, PEREIRA Y BALLESTEROS (1979)

citan nuevamente esta especie como L.

(A.) neapolitana en base a un ejemplar

recolectado en Tossa de Mar (Gerona)

sobre el alga rodofícea Sphaerococcus

coronopifolius Stackhouse, 1797, no apor-

tando ninguna información anatómica y

cromática. Años más tarde, ORTEA Y

44

BALLESTEROS (1982), en su estudio

monográfico del género Onchidoris,

incluyen una descripción de la anatomía

externa y coloración de O. neapolitana a

partir de un ejemplar de 9 mm de longi-

tud, recolectado también en Tossa de

Mar y sobre el alga rodofícea Sphaerococ-

cus coronopifolius. En este mismo trabajo,

los autores consideran dudosa la cita de

Ros (1975), debido a que los caracteres

anatómicos externos en base a los cuales

se llevó ha cabo la diagnosis diferencial

específica, son del todo genéricos.

En el océano Atlántico, esta especie

ha sido citada en Gran Bretaña por

PruvoT-FoL (1954) y España (Asturias)

(ORTEA, 1976). Sin embargo, las diagno-

SÁNCHEZ-SANTOS: Redescripción de Onchidoris neapolitana

sis específicas del material en los que

ambos registros atlánticos fueron

basados han sido materia de controver-

sia. Así, mientras que ORTEA Y BALLES-

TEROS (1982) consideraron la cita de

PrRUuvoT-FoL (1954) como muy dudosa,

CATTANEO-VIETTI Y THOMPSON (1989),

basándose en ella, incluyeron a O. neapo-

litana entre las especies con distribución

atlanto-mediterránea. En cuanto a la

presencia de O. neapolitana en la costa

atlántica de la Península Ibérica, ORTEA

Y URGORRI (1979) y BALLESTEROS (1985),

tomando como referencia la cita de

ORTEA (1976), consideraron también

para esta especie una distribución

atlanto-mediterránea. Sin embargo,

ORTEA Y BALLESTEROS (1982), en su revi-

sión del género Onchidoris en el litoral

ibérico, establecen como únicas referen-

cias válidas las procedentes de la costa

mediterránea española.

En este trabajo se redescribe esta

especie a partir de ejemplares proceden-

tes del Sur de España (Estrecho de

Gibraltar), así como algunos datos bioló-

gicos inherentes a su hábitat y la puesta.

MATERIAL Y MÉTODOS

En junio de 2003 se recogieron varios

ejemplares en la zona del Estrecho de

SISTEMÁTICA

Gibraltar que atribuimos a Onchidoris

neapolitana y que constituyen el objeto

del presente trabajo. Se redescribe la

especie a partir de dichos ejemplares y

se aportan algunos datos biológicos

referidos a su hábitat y a la puesta.

Los animales, tras 24 horas de con-

gelación en agua de mar, fueron preser-

vados en etanol al 96%. Para el estudio

anatómico interno, dos ejemplares

fijados, de 7 y 8 mm respectivamente,

fueron disecados por incisión dorsal

prestando especial atención a la morfo-

logía del sistema reproductor y de la

rádula. La genitalia fue examinada y

dibujada bajo un microscopio óptico

con cámara clara. Para estudiar la mor- :

fología radular, el aparato bucal fue

extraído y mantenido en una solución

de KOH al 10% con el objeto de disol-

ver el tejido orgánico circundante. La

rádula, así aislada, fue sumergida en

agua, secada y finalmente montada

para su examen con microscopía elec-

trónica de barrido.

El material examinado se encuentra

depositado en el Natural History

Museum of Los Angeles County

(LACM), en el Departamento de Biolo-

gía de la Universidad de Cádiz (UCA) y

en el Departamento de Biología Animal

y Ecología de la Universidad de

Granada (UGR).

DORIDOIDEA Pelseneer, 1894

Familia ONCHIDORIDIDAE Alder y Hancock, 1845

Género Onchidoris Blainville, 1816

Onchidoris neapolitana (Delle Chiaje, 1844) (Figs. 1-3)

Idalia neapolitana Delle Chiaje, 1844, Napoli 8: appendice prima tomo 2: 5-12 [Localidad tipo:

Nápoles]

Anatomía externa: Cuerpo muy apla-

nado. El noto es espiculoso, ovalado y

sobresale ampliamente al pie (Fig. 14).

De los tres tipos de disposición espicu-

lar característicos del género, sólo se

pudo apreciar la disposición radial, en

el margen del noto, y la transversal en la

zona central del mismo. La oblicua no

pudo ser observada. La superficie dorsal

del noto presenta papilas digitiformes

(Fig. 14). Las de la zona central son más

cortas que las laterales y marginales,

proyectándose estas últimas más allá

del margen notal claramente, mientras

que del área intrabranquial son más

cortas que las laterales y marginales. El

45

Iberus, 23 (1), 2005

número de papilas presentes en el área

intrabranquial oscila entre 5 y 8. El

número de laminillas rinofóricas oscila

entre 6 y 7 (Fig. 1C). Orificios rinofóricos

lisos y sin vaina. La branquia está cons-

tituida por hojas branquiales unipinna-

das alineadas alrededor del ano for-

mando un óvalo abierto posteriormente

(Fig. 1D). El tamaño de las hojas decrece

desde la central hasta las posteriores. El

número de hojas branquiales oscila

entre 12, en ejemplares de 5, 6 y 7 mm

de longitud, y 14 en un ejemplar de 9

mm de longitud.

Coloración: El noto es de color rojo-

anaranjado con pequeñas manchas

pardas de geometría irregular, más den-

samente concentradas en la zona central

del noto. En el borde del noto se apre-

cian bandas de color pardo grisáceo en

disposición radial con manchas naranja

intercaladas entre ellas. Las papilas son

de color pardo claro translúcido. Las

áreas que rodean los orificios rinofóricos

y branquial están despigmentadas. Los

rinóforos son gris oscuro, mientras que

las branquias son de color naranja. La

superficie del área intrabranquial está

despigmentada con papilas ligeramente

más claras que en el resto del dorso. El

pie y el velo bucal son naranja claro.

Anatomía interna: La rádula es bise-

riada, con ausencia de diente raquídeo.

El diente lateral más interno (Fig. 2) es

de gran tamaño, tiene una placa basal

cuadrangular grande con el borde inte-

rior curvado y orientado hacia fuera, el

borde exterior presenta una cúspide

prominente y alargada en cuyo borde

interno presenta 7-8 dentículos bien

desarrollados. Asimismo, destaca la pre-

sencia de una prolongación posterior

por debajo de la cúspide. El diente más

externo está formado por una placa

basal pequeña y cóncava en cuyo borde

inferior posee una pequeña cúspide

rudimentaria (Fig. 2). La fórmula

radular de un ejemplar de 7 mm es 22 x

1.1.0.1.1. La armadura labial está com-

puesta por uncinos simples.

El sistema reproductor (Fig. 3) pre-

senta un conducto hermafrodita que se

continua en una ampolla hermafrodita

pequeña y curvada. El conducto defe-

46

rente es alargado y no presenta región

prostática morfológicamente diferen-

ciada, desembocando en el atrio genital.

El pene es inerme. El conducto vaginal,

más corto y delgado que el conducto

deferente, conecta con una bolsa copula-

triz esférica. El receptáculo seminal tiene

forma de saco curvado y se conecta a la

vagina cerca de la bolsa copulatriz. La

bolsa copulatriz, el receptáculo seminal

y la ampolla hermafrodita son de

tamaño similar. Un corto y muy delgado

conducto uterino conecta directamente

con la bolsa copulatriz en un punto

adyacente a la conexión de esta con la

vagina.

Discusión: La coloración y la anato-

mía externa de los ejemplares del Estre-

cho de Gibraltar son muy similares a las

que muestran SCHMEKEL Y PORTMANN

(1982) en los ejemplares recolectados en

el golfo de Nápoles. Uno de los ejempla-

res estudiados por dichos autores pre-

senta una rádula con un diente lateral

interno con 4-6 dentículos. Las diferen-

cias respecto al número de dentículos de

nuestro material probablemente se

deban a una cierta variabilidad en este

carácter. Asimismo, a nivel del sistema

reproductor hemos observado diferen-

cias entre ambos materiales respecto al

tamaño y la forma del receptáculo

seminal y la bolsa copulatriz, así como

la ausencia de un conducto uterino cla-

ramente definido en el esquema que

presentan SCHMEKEL Y PORTMANN (1982,

pág. 118, fig. 7.28a). Sin embargo, el

esquema de estos autores es una repre-

sentación idealizada del sistema repro-

ductor, y no un estudio morfológico

detallado. Por ello, tales diferencias con

nuestras observaciones no deben consi-

derarse significativas.

Respecto al ejemplar del litoral

catalán descrito por ORTEA Y BALLESTE-

ROS (1982), las semejanzas entre ambos

materiales son grandes. La variación en

el número de laminillas rinofóricas y

hojas branquiales observada en nuestros

ejemplares se debe probablemente a la

variabilidad intraespecífica de la

especie. En cuanto a la coloración, la

única diferencia observada es la presen-

cia en todos los ejemplares del Estrecho

SÁNCHEZ-SANTOS: Redescripción de Onchidoris neapolitana

de Gibraltar de una zona despigmen-

tada alrededor de los orificios rinofóri-

cos y branquial. ORTEA Y BALLESTEROS

(1982) no describen la anatomía interna

de su material, por lo que no hemos

podido compararla con la del nuestro.

De las especies de Onchidoris citadas

en las costas de la Península Ibérica,

únicamente Onchidoris depressa (Alder y

Hancock, 1842) y Onchidoris tridactila

Ortea y Ballesteros, 1982, ambas con dis-

tribución atlántica, comparten con O.

neapolitana la presencia de papilas en su

noto. Sin embargo, desde el punto de

vista del aspecto externo, O. depressa

difiere de esta última por su modelo cro-

mático y disposición en forma de herra-

dura de las hojas branquiales (ORTEA Y

BALLESTEROS, 1982; THOMPSON Y

BROWN, 1984; PICTON Y MORROW, 1994)

y O. tridactila por su diseño cromático y

la presencia de una vaina rinofórica pro-

vista de tubérculos y por diferencias en

la puesta (ORTEA Y BALLESTEROS, 1982).

Desde el punto de vista de la anato-

mía interna, la morfología radular de O.

depressa, O. tridactila y O. neapolitana es

muy similar, si bien, los datos existentes

de estas tres especies, revelan pequeñas

diferencias en el número de dentículos

en el diente lateral más interno. Así, en

O. neapolitana este sería de 4-8 (SCHME-

KEL Y PORTMANN, 1982; presente

trabajo), en O. depressa, sería de 4-5

(THOMPSON Y BROWN, 1984) y en O. tri-

dactila es de 3-4 en el holotipo de la

especie (ORTEA Y BALLESTEROS, 1982). El

segundo diente lateral es idéntico en las

tres especies. Dada la similitud en la

morfología radular, las diferencias

observadas en el número de dentículos

pueden ser debidas a una variación

intra-específica u ontogenética de esta

estructura. Esto último, unido al hecho

de que la captura de O. neapolitana en

aguas del Estrecho de Gibraltar indica

que las tres especies muestran áreas de

distribución solapadas en el Atlántico,

sugieren la posibilidad de que sean

sinónimas. Para aclarar este aspecto

habría que examinar el material tipo de

las tres especies.

En cuanto al sistema reproductor,

estas tres especies no pueden ser com-

paradas ya que el único conocido es el

de O. neapolitana.

Existe otra especie mediterránea no

citada en las costas ibéricas, también con

papilas en su noto, Onchidoris bouvieri

(Vayssiere, 1919). Sin embargo difiere de

O. neapolitana por la presencia de una

espícula en el interior de las proyeccio-

nes papilares y la disposición en círculo

de las hojas branquiales (ORTEA Y

BALLESTEROS, 1982).

En cuanto a la biología de O. neapo-

litana, según la literatura, esta especie

se encuentra asociada a varias especies

de briozoos incrustantes de diversos

géneros (Chorizopora Hincks, 1879;

Cribrilaria Silén, 1941; Escharina Milne-

Edwards, 1836; Microporella Hincks,

1877; Parasmittina Osburn, 1952; Schis-

mopora MacGillivray, 1888; Schizobra-

chiella Canu y Bassler, 1920; Schizoma-

vella Canu y Bassler, 1917; Schizoporella

Hincks, 1877; Smittoidea Osburn, 1952;

Tubulipora Lamarck, 1816) de las que se

alimentaría (Ros, 1975, 1978). Todos

nuestros ejemplares fueron siempre

encontrados asociados a formaciones

del briozoo incrustante Schizobrachiella

sanguinea, sobre el que mostraban una

acusada homocromía. La puesta de

algunos ejemplares pudo ser observada

en el medio siempre depositadas sobre

S. sanguinea, y consistía en una cinta

transparente enrollada en espiral de

dos vueltas con los huevos de color

naranja.

El hallazgo de O. neapolitana en

aguas del estrecho de Gibraltar consti-

tuye la primera cita de esta especie para

el litoral andaluz. Asimismo, en las

colecciones malacológicas del Museo

Nacional de Ciencias Naturales de

Madrid existe un ejemplar de esta

especie (con el n* de catálogo

15.05/804), capturado en el Peñón del

Cuervo (Málaga), el 18 de julio de 1985

(colectado e identificado por José Tem-

plado). Esta nueva adición a la fauna de

gasterópodos opistobranquios del litoral

andaluz supone una considerable

ampliación de su área de distribución,

cuyo límite más occidental en el Medite-

rráneo estaba situado en el litoral

catalán.

47

Iberus, 23 (1), 2005

AGRADECIMIENTOS

Muy especialmente deseo expresar

mi más profunda gratitud al Dr. Angel

Valdés (Natural History Museum of Los

Angeles County, LACM) por su decisiva

ayuda en la realización del estudio ana-

tómico interno y por la revisión crítica

del manuscrito. Asimismo, mi agradeci-

miento a los Drs. Luis Sánchez-Tocino

(Universidad de Granada) y Juan Lucas

BIBLIOGRAFÍA

ALTIMIRA, C., HUELIN, M. F. Y Ros, J. D., 1981.

Molluscs bentónics de les illes Medes (Gi-

rona). I. Sistemática. Butlleti de la Institucio Ca-

talana d'Historia Natural, 47 (Seccio de Zoo-

logia 4): 69-75.

BALLESTEROS, M., 1985. Contribución al conoci-

miento de los Sacoglossos y Nudibranquios (Mo-

llusca: Opisthobranchia). Estudio anatómico, sis-

temático y faunístico de las especies del Medite-

rráneo español. Resumen de la Tesis

Presentada para aspirar al grado de Doctor

en Ciencias Biológicas, Universitat de Bar-

celona. Centre de Publicaciones Intercanvi

Cientific I Extensio Universitaria, 46 pp.

CATTANEO-VIETTI, R. Y THOMPSON, T. E., 1989.

Mediterranean opisthobranch molluscs: a

zoogeographic approach. Bollettino Malaco-

logico 25(5-8): 183-204.

CERVERA, J. L., TEMPLADO, J., GARCÍA GÓMEZ,

J. C., BALLESTEROS, M., ORTEA, J. A., GARCÍA,

F. J., ROS, J. Y LUQUE, A. A., 1988. Catálogo

actualizado y comentado de los opistobran-

quios (Mollusca, Gastropoda) de la Península

Ibérica, Baleares y Canarias, con algunas re-

ferencias a Ceuta y la Isla de Alborán. Iberus,

suplemento 1.: 1-84.

DELLE CHIAJE, S., 1844. Descrizione e notomia

degli animali invertebrati della Sicilia citeriore

osservati vivi negli anni 1822-1830, Napoli 8:

appendice prima tomo 2, pp. 5-12.

HUELIN, M. F. Y Ros, J., 1984. Els molluscs ma-

rins de les Illes Medes, pp. 457-504. In: J. Ros,

I. Olivella, € J. M. Gili (eds.) Els sistemes na-

turals de les Illes Medes, Institut d'Estudis Ca-

talans.

ORTEA, J. A., 1976. Catálogo brevemente comen-

tado de la fauna de moluscos marinos gaste-

rópodos y bivalvos existentes en el estuario

de Villaviciosa. Asturnatura, 3, pp. 109-120.

ORTEA, J. A. Y URGORRI, V., 1979. Una nueva es-

pecie de Onchidoris (Moluscos, opistobran-

quios, Doridáceos) del Norte de Noroeste de

España, Onchidoris cerviñoi n. sp. Cahiers de

Biologie Marine, 20: 507-513

48

Cervera Currado (Universidad de

Cádiz) por sus valiosos comentarios

sobre el manuscrito. Por último, a los

alumnos de doctorado Manuel Caballer

Gutiérrez (Universidad de Cantabria) y

Leopoldo Moro Abad (Museo de Cien-

cias Naturales, Tenerife) por facilitarme

gran parte de la información bibliográ-

fica.

ORTEA, J. A. Y BALLESTEROS, M., 1982. Sobre al-

gunos Onchidoris Blainville, 1816 (Mollusca,

. Opisthobranchia, Doridácea) del Litoral Ibé-

rico. Investigación Pesquera, 46 (2): 239-254.

PEREIRA, F. Y BALLESTEROS, M., 1979. Gasteró-

podos del litoral mediterráneo español. II.

Tossa de Mar. Gerona. Primer Simposio de

Bentos Marino, 223-235 pp.

PIcTON, B. E. Y MORROW, C. C., 1994. A Field

Guide to the Nudibranchs of the British Isles,

Immel Publishing, 143 pp.

PRUVOT FoL, A., 1951. Etude des nudibranches

de la Méditerranée (2e partie). Archives de

Zoologie Experimentale et Generale. Paris. 88 (1):

1-80, pls. 1-4.

PRUVOT FoL, A., 1954. Mollusques Opisthobran-

ches. Faune de France, Paris (Lechevalier) 58:

1-460.

Ros, J. D., 1975. Opistobranquios (Gastropoda:

Euthyneura) del litoral ibérico. Investigación

Pesquera., 39 (2): 269-372.

Ros, J. D., 1978. La alimentación y el sustrato

en los opistobranquios ibéricos. Oecologia

Aquatica, 3: 153- 166.

Ros, J., 1985. Els poblaments d'Opistobranquis

de coves submarines mediterranies: noves da-

des i comentaris sobre llur afinitat faunis-

tica. Butlleti de la Institucio Catalana d'Histo-

ria Natural, 52 (Seccio de Zoologia, 6): 87-94.

Ros, J. D. Y ALTIMIRA, C., 1977. Comunidades

bentónicas de sustrato duro del litoral NE es-

pañol. V. Sistemática de moluscos. Miscelá-

nea Zoológica, 4 (1): 43-45.

SCHMEKEL, R. L., 1968. Ascoglossa, Notaspidea

und Nudibranchia im Litoral des Golfes von

Neapel. Revue Suisse de Zoologie, 75(6): 103-

15:

SCHMEKEL, R. L. Y PORTMANN, A., 1982. Opist-

hobranchia des Mittelmeeres. Nudibranchia und

Sacoglossa. Springer-Velarg. Berlin, 410 pp.

THOMPSON, T. E. Y BROwN, G. H., 1984. Biology

of opisthobranch molluscs, Vol. 2. The Ray So-

ciety, London, 229 pp.

O Sociedad Española de Malacología Iberus, 23 (1): 49-65, 2005

Contribution to the knowledge of the family Caecidae: 16.

Revision of the Caecidae of HEaster Island (Chile)

(Caenogastropoda: Rissooidea Gray J. E., 1847)

Contribución al conocimiento de la familia Caecidae: 16. Revisión

de los Caecidae de la Isla de Pascua (Chile) (Caenogastropoda:

Rissooidea J. E. Gray, 1847)

Bret RAINES* and Mauro PIZZINI**

Recibido el 26-X-2004. Aceptado el 11-11-2005

ABSTRACT

With the exception of two species, members of the family Caecidae from Easter Island have

been previously neglected. Based on type and additional material, a revision of the species

known to date from Easter ls. is herein proposed, with the description of 5 new species:

Caecum rehderi spec. nov., C. heterochromum spec. nov., C. pascuanum spec. nov., C.

rapanviense spec. nov., C. campanulatum spec. nov.

RESUMEN

Con la excepción de dos especies, los miembros de la familia Caecidae de la Isla de

Pascua han sido previamente desatendido. Basado en el tipo y en el material adicional, se

propone aquí dentro, una revisión de las especies conocidas hasta la fecha de la Isla de

Pascua, y incluye la descripción de 5 nuevas especies: C. rehderi esp. n., C. heterochro-

mum esp. n., C. pascuanum esp. n., C. rapanuiense esp. n., C. campanulatum esp. n.

KEY WORDS: Mollusca, Caenogastropoda, Rissooidea, Caecidae, taxonomy, new species, Easter Island,

Western Pacific.

PALABRAS CLAVE: Mollusca, Caenogastropoda, Rissooidea, Caecidae, taxonomia, nuevas especies, Isla de

Pascua, Pacífico Occidental.

INTRODUCTION

Easter Island (Fig. 1) is found in a to-

tally isolated position approximately

3,500 Km from the coast of Chile. From a

zoogeographical point of view, Easter ls.

is in a very peculiar area within the east-

ern Indo-Pacific region, and to the point

that, in 1965, Schilder proposed the Ra-

panuian province as a separate biogeo-

graphical province from the Polynesian

province. The submarine seascape fea-

tures widely scattered corals affixed to

the rugged volcanic substrate. The de-

pauperate benthic community employs a

variety of adaptive strategies for survival

in an environment stressed by waves,

currents and the absence of mineral nu-

* Research Associate, Natural History Museum of Los Angeles County , P.O. Box 612 Victorville, CA 92393

USA. e-mail: rainesbkPyahoo.com

** Largo della Caffarelletta, 6, 00179 ROME (Italy). e-mail: ma.pizziniClibero.it

AS

Iberus, 23 (1), 2005

trients. Most of the corals and other bot-

tom invertebrates are typical of the Indo-

Pacific reefs, but reefs have not formed.

The circulation pattern, and espe-

cially the marked upwelling between

the South Pacific and Mentor currents,

contributes to the isolation of the area.

In terms of marine ecosystems, this in-

sularity has also produced a high degree

of radiation in many groups, and partic-

ularly in the caenogastropods. Thus the

fauna can essentially be described as

typical Pacific fauna with a relatively

high number of endemic species. The is-

land's marine benthic fauna is generally

characterised by a high degree of

species diversity and a low abundance,

and this is true for both hard and soft

bottoms. The meiobenthic family Caeci-

dae also follows this pattern.

The Caecidae of the Easter Island

have been scarcely studied in the past,

with the exception of the work of

REHDER (1980). Past surveys have

reported endemicity rates within the

molluscan fauna ranging from 37% to

42% (REHDER, 1980, DISALVO, RANDALL

AND CEA, 1988, RAINES, 2002). Rehder

also indicates that some species appear

to have a dual relationship with certain

species from Hawaii, as well as species

from the Kermadec Islands. In prepara-

tion for his 1980 publication, Rehder

reviewed all previous studies and expe-

ditions. In addition to examining all the

Easter Is. material in US museums, he

also examined the full store of material

housed in the Museo National de Histo-

ria Natural in Santiago, Chile. In all,

Rehder examined over 7,000 specimens,

of which 3480 were collected during his

trip to the island in 1974.

In the mid 1980's, DiSalvo and his

team conducted a comprehensive survey

of Easter Island's sublittoral marine

environment. The authors of the present

work had the good fortune of having

access to examples of all the molluscan

material collected during DiSalvo's

investigation. Other than the few publi-

cations that we have been working on

(OLIVER, 1915, SCHILDER, 1965, REHDER,

1980, DISALVO ET AL., 1988, RAINES, 2002)

no other serious work on molluscs has

50

been completed during the last ten

years. With regard to Caecidae, during

the three trips to Easter Island, the

authors of the present work collected

more than 350 specimens of this family.

Rehder mentions only two species:

Caecum cf. solitarium Oliver, 1915, and C.

amydroglyptum Rehder, 1980. In all other

publications, which we have reviewed,

Caecidae are listed simply as “Caecum

species”. No other descriptions or illus-

trations were provided.

So we present herein a revision of

the species known to date from Easter

Is., based on type and additional mater-

ial, with the description of five new

species.

Abbreviations used:

AMS: Australian Museum Sydney,

Sydney (Australia)

CM: Canterbury Museum, Christchurch

(New Zealand)

LACM: Natural History Museum of Los

Angeles County, Los Angeles (U.S.A.)

MNHN: Museum National d'Histoire

Naturelle, Paris (France)

MPR: Mauro Pizzini collection.

NHML: Natural History Museum,

London (U.K.)

NMNZ: Museum of New Zealand Te

Papa Tongarewa, Wellington (New

Zealand)

NSMT: National Science Museum,

Tokyo (Japan)

USNM: National Museum of Natural

History, Washington D.C. (U.S.A.)

WAM: Western Australian Museum,

Perth (Australia)

Terminology:

Abapical: towards the apices (of the

septum)

Adapical: opposite to the apices direc-

tion (of the septum)

Aperture, apertural end: the round ante-

rior opening of the shell.

Apex, apical end: the smaller, narrower,

closed posterior end of the tube.

Cutting plane: the plane individuated

by the edge of the shell at the apex

(excluding septum and mucro).

RAINES AND PIZZINI: Revision of the Caecidae of Easter Island (Chile)

UNITED STA

SMAN $ ¡

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3 2 NEW NORTH Y

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Figure 1. Biogeographical provinces within the tropical eastern Indo-Pacific as proposed by

SCHILDER (1965), and as illustrated by REHDER (1980): Micronesian: M; Hawaiian: H; Fijian: E;

Polynesian: P; Rapanuian: R; Kermadec Islands: K (added by the authors).

Figura 1. Las provincias biogeographicas dentro del Indo-Pacífico oriental tropical propuestas por

SCHILDER (1965) e ¿ilustradas por REHDER (1980): Micronesian: M; Hawaiano: H; Fijian: E: Poline-

sio: P; Rapanuzan: R; Kermadec Islands: K (añadida por los autores).

Interspace: area between rings, with /

without microsculpture.

Meiobenthic: referred to all interstitial

molluscs living in sediment of

varying granule size.

Microsculpture: usually visible at very

high magnification or under SEM can

be transverse, longitudinal or both.

Mucro: small to large prong projecting

from the septum.

RESULTS

Rings, annular sculpture: transverse,

raised sculpture (equivalent to the

axial sculpture of the normally

coiled gastropods).

Septum: closure of the shell at the apex,

as it sheds earlier stages.

Shell(s): the shell, beached without gas-

tropod.

Spm(s): live collected specimen(s), with

soft parts and /or operculum(a).

Superfamily RIsso0IDEA Gray J. E., 1847

Family CAECIDAE Gray J. E., 1850

Genus Caecum Fleming, 1813

Diagnosis (BANDEL, 1996): “The shell of the teleoconch is a small, slightly curved tube orna-

mented only with growth lines, numerous ring-like collabral lirae ad /or axial ribs. The poste-

rior end of the tube is closed by a conical septum. The protoconch is trochospirally or planispi-

rally coiled. Uncoiling of the shell begins after metamorphosis”.

Type species (BANDEL, 1996): Dentalium imperforatum Kanmaker, 1798 (= trachea Montagu, 1803)

from Europe, Mediterranean Sea and Atlantic to southern England.

dl

Iberus, 23 (1), 2005

100 yum

Figure 2. Caecum rehderi spec. nov. A: holotype LACM 3019, gold coated, length 2.08 mm; B:

detail of septum; C: microsculpture; D: detail of aperture. SEM imaging by D. Geiger.

Figura 2. Caecum rehderi spec. nov. A: holotipo LACM 3019, metalizado en oro, longitud 2,08 mm,

B: detalle del septo; C: microescultura; D: detalle de la abertura. Imágenes al MEB por D. Geiger.

Caecum rehderi spec. nov. (Fig. 2)

Caecum cf. solitarium; Rehder, 1980: 31-32, pl. 5, fig. 11.

Type material: Holotype, LACM 3019; 1 paratype, LACM 3020; 1 paratype, USNM 756269; 1

paratype, MNHN.

Material examined: 1 specimen, Onetea, Hotuiti (length: 2.42 mm, USNM 756269) (Oct. 1974, leg.

H. Rehder); 4 specimens, Punta Rosalia, east of Anakena (Apr. 1998, leg. B. Raines),

Type locality: In sand collected along the base of cliffs at 10-20m, off Punta Rosalia, east of

Anakena, Easter 1s., Chile. 27? 04' 18” S, 109% 19 45” W.

Description: Shell small (holotype

measures, length: 2.08 mm; width: 0.42

mm), tube-like, slender, gently arched,

semi-translucent to opaque white. Tube

seemingly smooth almost glassy, sub-

cylindrical, with posterior end only

slightly smaller than anterior end.

Microsculpture nearly obsolete, with

only fine annular growth lines sometimes

present under magnification. Anterior

end somewhat flared just above aperture

92

with incised annular rings. Aperture cir-

cular, but slightly constricted. Posterior

end with tapered rim. Septum not

retracted, subquadrate lateral outline

inclined with elevated edge slightly right

of center when viewed frontally. Opercu-

lum and soft parts unknown.

Original description of C. cf. solitar-

ium Rehder, 1980: “Diagnosis. Shell

small, 2.4 to 2.7 mm in length, glassy,

grayish-white to whitish, slender, gently

RAINES AND PIZZINI: Revision of the Caecidae of Easter Island (Chile)

Figure 3. Caecum cf. solitarium Rehder (1980). A: USNM 756269, uncoated, length 2.42 mm; B:

detail of septum; C: microsculpture; D: detail of aperture. SEM imaging by D. Geiger.

Figura 3. Caecum cf. solitarium Rehder (1980). A: USNM 756269, no metalizado, longitud 2,42 mm;

B: detalle del septo; C: microescultura; D: detalle de la abertura. Imágenes al MEB por D. Geiger.

curved, diameter at posterior end only

slightly smaller than at anterior end,

where the aperture is slightly con-

stricted and somewhat opaque above

the aperture; the sculpture consists of

fine, rather crowded, subobscure

(worn?) annular riblets that gradually

and slightly increase in strength toward

the aperture; septum exserted, sub-

quadrate with a slightly convex surface

inclined from an elevated edge at the

right dorsal sector to the edge of the

posterior rim of the shell at the left

ventral sector.

Range. Kermadec Islands (and

Easter Island ?).

Material. 1 specimen from sta E-27A,

USNM 756269.

Measurements (mm). USNM 756269:

length, 2.42; diameter at anterior end, 0.4”.

Discussion: Rehder reported a caecid

from Easter Island, which he tentatively

identified as Caecum cf. solitarium Oliver,

1915. However, it seems that Rehder un-

fortunately overlooked several key char-

acteristics within Oliver”s description.

The first being the septum of C. solitar-

tum, which OLIVER (1915) described as

“....hemispherical, making an abrupt

shoulder at the junction of the shell”;

even the septum of Rehder”s specimen

(Fig. 3) could be associated to a tale ty-

pology. Oliver also mentions, that the

sculpture of C. solitarium consists of sim-

ple growth lines, while Rehder refers to

the sculpture as consisting of suboscure

(worn?) annular riblets that gradually

and slightly increase in strength toward

the aperture. Other main difference be-

tween C. solitarium Oliver, 1915 and C.

solitarium sensu Rehder is that the first

has a nearly uniform diameter, while the

second shows the diameter at posterior

end only slightly smaller than at ante-

93

Iberus, 23 (1), 2005

Figure 4. Caecum solitarium Oliver, 1915. A: Holotype CM M2867, uncoated, length 1.68 mm;

B: detail of aperture; C: microsculpture; D: detail of posterior. SEM imaging by N. Andrews.

Figura 4. Caecum solitarium Oliver, 1915. A: Holotipo CM M2867, no metalizado, longitud 1,68 mm;

B: detalle de la abertura; C: microescultura; D: detalle trasero. Imágenes al MEB por N. Andrews.

rior end. Furthermore, it appears Re-

hder did not examine Oliver”s type ma-

terial, because if he would have done,

he would have noted that the two speci-

mens have significantly different ante-

rior ends (Figs. 3D, 4B), and that the

holotype of C. solitarium is badly broken

and lacks the entire posterior end (Fig.

4). The damage to the holotype is old

and worn, and possibly occurred in situ

suggesting that Oliver may have chosen

an imperfect specimen as the type and

actually described another in his de-

scription. (Scofield, 2002, pers. commu-

nication). Rehder”s specimen is, how-

ever, consistent with C. rehderi, and

therefore, has been designated as the

paratype. Although Rehder broke the

anterior end of his specimen while mea-

suring it, all the pieces were available

for examination. The specimens which

we found, actually showed a series of

54

small rings along the entire tube; and

apart from the relativity of the term's

significance, we hold that the difference

between our specimens and those de-

scribed by the two authors falls within

the species” range of variability, in light,

above all else, of the high degree of

adaptation of the local molluscs to the

island's distinguishing geo-climatic con-

ditions. It is known that a number of

species of Caecidae (i.e. C. lightfootae

Pizzini, Nofroni and Oliverio, 1994),

though they have the same general

shape (septum, tube and aperture),

could show a very wide range of vari-

ability in terms of the type of sculpture.

Remarks: Caecum rehderi seems to be

an unusually fragile species. Of the five

known specimens, Rehder chipped the

aperture of his specimen (USNM

756269) while measurinyg it, the holotype

has a small longitudinal crack toward

RAINES AND PIZZINI: Revision of the Caecidae of Easter Island (Chile)

Figure 5. Caecum amydroglyptum Rehder, 1980. A: holotype USNM 757977, uncoated, length 1.67

mm; B: detail of septum; C: microsculpture; D: detail of aperture. SEM imaging by D. Geiger.

Figura 5. Caecum amydroglyptum Rehder, 1980. A: holotipo USNM 757977, no metalizado, longitud

1,67 mm; B: detalle del septo; C: microescultura; D: detalle de la abertura. Imágenes al MEB por D. Geiger.

the aperture end, the other paratype has the Kermadec Islands, while Rehder

a Chip in the aperture, and the junior

author completely crushed another

specimen while examining it.

Geographical distribution: Oliver

described Caecum solitarium only from

tentatively indicated the species as

being from Easter Is. In our opinion,

according to the present knowledge, C.

solitarium is restricted to the Kermadec

Is.

Caecum amydroglyptum Rehder, 1980 (Figs. 5, 6)

Caecum amydroglyptum; Rehder, 1980: 32, pl. 5, fig. 12.

Type material: Holotype, USNM 757977; 1 paratype, USNM 757978

Material examined: Original types. Holotype, USNM 757977; paratype, USNM 757978, (Oct. 1974,

leg. H. Rehder). 132 specimens in sand collected along the base of cliffs at 10-20m, off Punta Rosalia,

east of Anakena, 27” 04 18” S, 109* 19 45” W (Apr. 1998, leg. B. Raines).

Voucher material: 4 specimens were deposited in each of the following institutions: LACM; USNM

1018792; MNHN; NHML; NMNZ M.273207; NSMT Mo 73562; AMS C.205278; WAM 513783, and

6 specimens (3 adults /3 juveniles), MPR. 13 shells, beach of Anakena Bay, on the northern coast of

Easter Is., picked up among the rocky bottom on the west side of the bay at low tide, among com-

munities of Dictyotales, with Galaxaura obtusata. (12-1-1995, leg. E. Rolán), MPR.

Type locality: Station E-27A, Onetea, Hotuiti: in patch of sand above high tide level.

II

Iberus, 23 (1), 2005

B

100 um

100 um

Figure 6. Caecum amydroglyptum Rehder, 1980. A: Voucher specimen LACM, gold coated length

1.53 mm; B: detail of septum; C: microsculpture; D: detail of aperture. SEM imaging by D.

Geiger.

Figura 6. Caecum amydroglyptum Rehder, 1980. A: El espécimen del vale LACM, metalizado en oro,

longitud 1,53 mm, B: detalle del septo; C: microescultura; D: detalle de la abertura. Imágenes al MEB

por D. Getger.

Original description: “Shell small,

from 1.3 to 1.7 mm in length, curved,

rather evenly cylindrical with the ante-

rior end in fully grown specimens

slightly swollen above the aperture;

glassy grayish white to light orange

yellow in color; sculpture consists of

rather strong, somewhat distantly

spaced annular ribs that become more

or less obscure in the middle part of the

shell, with microscopic, longitudinal

wavy striae that are obscure at the ante-

rior and posterior ends; septum, low,

dome-shaped. ”

Additional description: Shell small

(mean length: 1.7 mm; width: min. 0.3

mm, max 0.4 mm), curved, colour

grayish white. The tube is perfectly

cylindrical, except near the aperture,

and its sculpture consists of about 36-40

rings, with some in the middle part of

56

the shell being less raised and changing

their shape, until they resemble very

fine growth lines. Microsculpture

formed by longitudinal worm-like striae

visible at enlargement of at least 180x.

Septum dome-shaped, slightly raised

over the cutting plane. Aperture consist-

ing of a large protuberance crossed by

slightly raised rings. Operculum and

soft parts unknown.

Remarks: We agree totally with

Rehder's conclusions; because we have

”...been unable to identify this species with

any published taxon” from either the

Indo-Pacific Provinces, the Panamic

Prov. or the Chilean Prov. We have

found only one species that resembles C.

amydroglyptum, which is C. vertebrale

Hedley, 1899, from Funafuti Is. It is

quite similar to amydroglyptum in terms

of the sculpture of the tube, longitudinal

RAINES AND PIZZINI: Revision of the Caecidae of Easter Island (Chile)

100 um

100 um

Figure 7. Caecum heterochromum spec. nov. A: holotype LACM 3021, gold coated, length 1.42

mm; B: detail of septum; C: microsculpture; D: detail of aperture. SEM imaging by D. Geiger.

Figura 7. Caecum heterochromum spec. nov. A: holotipo LACM 3021, metalizado en oro, longitud 1,42

mm); B: detalle del septo; C: microescultura; D: detalle de la abertura. Imágenes al MEB por D. Getger.

microsculpture and septum, but the tube, a very strong microsculpture

Rehder's species shows a much greater

swelling of the tube above the apertural

end, which is crossed by small sculp-

tured rings, while the adapical part of

vertebrale has almost the same diameter

of the tube. In addition, C. amydroglyp-

tum presents, along the entire length of

consisting of worm-like, longitudinal

striae, covering also the top of the rings,

while that of vertebrale is an indistinct

microsculpture not surely comparable to

a real striation (pers. observ.).

Geographical distribution: This species

would appear to be limited to Easter Is.

Caecum heterochromum spec. nov. (Figs. 7, 8)

Type material: Holotype, LACM 3021; 6 paratypes, LACM 3022; 6 Paratypes, USNM 1018789; 6

Paratypes, MNHN; 6 Paratypes, NHML,; 6 Paratypes, NMNZ M.273205; 6 Paratypes, NS5MT Mo

73560; 6 Paratypes, AMS C.205275,; 6 Paratypes, WAM S13780; 9 Paratypes, MPR.

Material examined: 168 specimens: off Hanga Nui; and 8 specimens off the western coastline

near Tahai (Dec. 2000, leg. B. Raines). 39 shells, beach of Anakena Bay, on the northern coast of

Easter Is., picked up among the rocky bottom on the west side of the bay at low tide, among

communities of Dictyotales, with Galaxaura obtusata. (12-1-1995, leg. E. Rolán) MPR.

Voucher material: 39 shells, beach of Anakena Bay, on the northern coast of Easter Is., picked up

among the rocky bottom on the west side of the bay at low tide, among communities of Dicty-

otales, with Galaxaura obtusata. (12-1-1995, leg. E. Rolán) MPR.

7)

Iberus, 23 (1), 2005

100 um

100 um

Figure 8. Caecum heterochromum spec. nov. A: paratype from lot LACM 3022, gold coated, length

1.53 mm; B: detail of septum; C: microsculpture; D: detail of aperture. SEM imaging by D. Geiger.

Figura 8. Caecum heterochromum spec. nov. A: paratipo de la porción LACM 3022, metalizado en oro,

longitud 1,53 mm; B: detalle del septo; C: microescultura; D: detalle de la abertura. Imágenes al MEB

por D. Getger.

Type locality: In sand collected along the base of cliffs at 20m off Hanga Nui, Easter Is., Chile.

27” 07' 46” S, 109* 16' 35” W.

Derivation of the name: From the greek etepoc: other and xpopo: colour.

Description: Shell small (mean

length: 1.6 mm; mean width: 0.4 mm)

with the tube subcylindrical in shape in

the abapical part and cylindrical up to

the vicinity of the aperture (Fig. 8),

where there is a slight swelling, fol-

lowed by a narrowing of the tube; the

aperture is perfectly circular, simple and

rimmed by a very slight flaring towards

the outside. The septum is dome-shaped

and slightly raised over the cutting

plane. Its sculpture is extremely vari-

able, ranging from specimens with

approximately 50 small raised rings that

are separated by interstices of corre-

58

sponding breadth and depth, particu-

larly in the upper portion of the tube

and near the aperture, to others that lack

any type of sculpture; these two extreme

represent the limits of the species” range

of variability, given that they were

found in intermediate specimens whose

rings are barely visible. The microsculp-

ture also presents a wide range of vari-

ability, with some specimens not

showing any trace of microsculpture,

while the surface of other specimens, at

an enlargement of 30x, presents a

microsculpture consisting of a large

number of very fine, worm-like striae

RAINES AND PIZZINI: Revision of the Caecidae of Easter Island (Chile)

100 um

Figure 9. Caecum pascuanum spec. nov. A: holotype LACM 3023, gold coated, length 1.64 mm;

B: detail of septum; C: microsculpture; D: detail of aperture. SEM imaging by D. Geiger.

Figura 9. Caecum pascuanum spec. nov. A: holotipo LACM 3023, metalizado en oro, longitud 1,64

mm; B: detalle del septo; C: microescultura; D: detalle de la abertura. Imágenes al MEB por D. Geiger:

that follow its axial direction, while

others have a rough surface. C. hete-

rochromum has an extremely variable

colouring and pattern; white, cream-

coloured shell with a pattern consisting

of brown, zigzagging lines running

almost parallel in a horizontal direction

(axial). There are also some specimens

showing a brown irregular stripe in the

middle portion of the shell. The colour-

ing is again a creamy white, with an

irregular vertical design consisting of

unequal spots. Operculum corneous,

light brown; its external side consists of

a smooth central nucleus and 5-6 con-

centric rings that run from this nucleus

up to the external border. Soft parts

unknown.

Remarks: Caecum heterochromum,

despite the limited nature of the name,

it is, in absolute terms, the most variable

of the species to be found on Easter ls.

In fact, its range of variability involves

not only its colouring and patterns, but

also major morphological characteris-

tics, such as microsculpture and sculp-

ture. Nevertheless, the silhouette, the

form of the septum and that of the tube,

as well as the apertural end, which,

when taken as a whole, constitute the

general form, are a constant that we

consider to be a distinguishing charac-

teristics of this species.

Geographical distribution: The species

is currently noted only in relation to

Easter ls.

59

Iberus, 23 (1), 2005

Figure 10. Caecum rapanuiense spec. nov. A: holotype LACM 3025, gold coated, length 1.58 mm;

B: detail of septum; C: microsculpture; D: detail of aperture. SEM imaging by D. Geiger.

Figura 10. Caecum rapanuiense spec. nov. A: holotipo LACM 3025, metalizado en oro, longitud 1,58

mm, B: detalle del septo; C: microescultura; D: detalle de la abertura. Imágenes al MEB por D. Geiger.

Caecum pascuanum spec. nov. (Fig. 9)

Type material: Holotype, LACM 3023; 2 paratypes, LACM 3024; 1 paratype, USNM 1018790; 1

paratype, MNHN,; 1 paratype, NHML; 1 paratype, AMS C.205276; 1 paratype, WAM 513781; 1

paratype, MPR.

Material examined: 9 specimens, Hanga-Teo on the northern coastline (Dec. 2000, leg. B. Raines).

Type locality: In silty mud collected at 15m in cave off Hanga-Teo on the northern coast, Easter Is.,

Chile, 27” 03” 37” S, 109* 21' 58” W.

Derivation of the name: The name of this new species comes from a latinized adjective formed

from the Spanish name of the island, “Isla de Pascua”.

Description: Shell small (holotype

measures, length: 1.96 mm; width: 0.36

mm), gently curved. The tube, evenly

cylindrical for almost its entire length,

presents a slightly smaller diameter only

in the abapical part. Towards the adapical

part, the tube widens visibly in a large

varix crossed by 5-6 rings that are sizable

but raised to various degrees, being sepa-

rated by interspaces that also vary in

terms of their depth and width. The

60

septum protrudes to a significant extent

over the cutting plane with an unguiform

mucro, visible to a greater or lesser extent

and oriented towards the dorsal side of

the tube. Frequently visible on the cutting

plane are residues of what may be a tem-

porary septum (PIZZINL, NOFRONI AND

OLIVERIO, 1998). Even under intensive

enlargement, no microsculptures are

visible. Circular aperture. Operculum

and soft parts unknown.

RAINES AND PIZZINI: Revision of the Caecidae of Easter Island (Chile)

Figure 11. Caecum crystallinum Folin, 1879. A: holotype NHML 1887.2.9.2363; B: detail of aper-

ture. SEM imaging by K. Way.

Figura 11. Caecum crystallinum Folin, 1879. A: holotipo NHML 1887.2.9.2363; B: detalle de la

abertura. Imágenes al MEB por K. Way.

Remarks: Caecum pascuanum presents

numerous morphological analogies with

C. rehderi, including the shape of the

tube and the septum, though it is set

apart by a large varix crossed by rings,

which is completely absent in the other.

In fact, this varix, though it can be

accentuated to a greater or lesser extent

- for that matter, consistently within the

range of variability - is always present

and would appear to represent the dis-

tinguishing morphological characteris-

tics of this species.

Geographical distribution: The species

is currently known only in its typical

location.

Caecum rapanuiense spec. nov. (Fig. 10)

Type material: Holotype, LACM 3025; 2 paratypes, LACM 3026; 1 paratype, USNM 1018791; 1

paratype, MNHN; 1 paratype, NHML; 1 paratype, NMNZ M.273206; 1 paratype, NSMT Mo 73561;

1 paratype, AMS C.205277; 1 paratype, WAM 513782; 1 paratype, MPR.

Material examined: 12 specimens, off the western coastline near Tahai (Dec. 2000, leg. B. Raines).

Type locality: Dredged at 30m in fine sand off the western coastline near Tahai, Easter Is., Chile,

ZIAOIF207 5, 109926305 WN:

Derivation of the name: This species take its title from the ancient name for Easter Is., which was

Rapa Nui.

Description: Shell small (dimensions

of the spm no. 3 from Easter ls., length:

1.5 mm; width: 0.35 mm), slightly

curved. Tube completely smooth

showing only a microsculpture consist-

ing of very weakly defined growth lines.

Septum dome-shaped, with the mucro

reduced to a small pedunculum, resem-

bling a squashed ball, found on the exte-

rior and oriented to the right. Aperture

simple, with no varix and only slight

swelling: further on the swelling tends

to contract, with a slightly reflected lip.

Growth stage and soft parts unknown.

Remarks: Following an initial exam-

ination, we tentatively classified this

species as C. crystallinum Folin, 1879,

despite the absence of the mucro in the

original type (Fig. 11), but it is

straighter, and the texture of the shell

is different, exhibiting under the

microscope fine longitudinal striations

(Fig. 11B), while C. rapanuiense shows

only very fine growth lines (Fig. 10C).

It also resembles C. glabriforme

Carpenter, 1857, but this species has

fairly strong microsculpture and a

large well developed septum. In terms

61

Iberus, 23 (1), 2005

Figure 12. Caecum campanulatum spec. nov. A: holotype LACM 3027, gold coated, length 1.96

mm; B: detail of septum; C: microsculpture; D: detail of aperture. SEM imaging by D. Geiger.

Figura 12. Caecum campanulatum spec. nov. A: holotipo LACM 3027, metalizado en oro, longitud 1,96

mm, B: detalle del septo; C: microescultura; D: detalle de la abertura. Imágenes al MEB por D. Geiger.

of the general shape of the tube and while our species is completely

septum, it resembles C. neocaledonicum smooth.

Folin, 1868, but the latter shows some Geographical distribution: Caecum

raised rings in the abapical part of the

tube near the aperture (PIZZINI, 1998),

rapanutense is actually known only from

Easter Is., the type locality.

Caecum campanulatum spec. nov. (Fig. 12)

Type material: Holotype, LACM 3027; 1 paratype, LACM 3028; 1 paratype, USNM 1019067.

Material examined: 6 specimens, off Hanga Nui (Dec. 2000, leg. B. Raines); 2 specimens, Punta

Rosalia, east of Anakena (Apr. 1998, leg. B. Raines).

Type locality: In sand collected along the base of cliffs at 20m, off Hanga Nui, Easter Is., Chile,

27” 07” 46” S, 109” 16' 35” W.

Derivation of the name: The name of the new species comes from the latinized adjective campan-

ulatus, which refers to the bell-shaped form of the apertural end.

Description: Shell small (holotype's

slightly near the apertural end, with a

dimensions, length: 1.64 mm; width:

silhouette that closely resembles that of

0.31 mm). Tube slightly arched, slender,

with it's abapical part only slightly

smaller then adapical. The tube widens

62

a bell, quickly narrowing itself once

again and ending in a sharp edge. The

microsculpture is quite obsolete and

RAINES AND PIZZINI: Revision of the Caecidae of Easter Island (Chile)

A 333 pm

Figure 13. Strebloceras subannulatum Folin, 1879. A: syntype NHML 1887.2.9.2308-2310, length

3.0 mm; B: detail of septum. SEM imaging by K. Way.

Figura 13. Strebloceras subannulatum Folin, 1879. A: sintiyo NHML 1887.2.9.2308-2310, longi-

tud 3,0 mm; B: detalle del septo. Imágenes al MEB por K. Way.

scarcely visible, even under intensive

optical enlargement, while the sculpture

would appear to consist of rings, which

are also barely observable, though they

are more visible near the aperture. The

septum is dome-shaped and scarcely

raised over the cutting plane. The aper-

ture is circular. Colour translucent, with

faint axial brown wavy lines. Opercu-

lum and soft parts unknown.

Remarks: Caecum campanulatum

shows some similarities with Caecum

amydroglyptum about the general shape

of the tube and the mucro, but the latter

has a longitudinal microsculpture cover-

ing all the tube, while the first one pre-

sents only a microsculpture consisting

of a scarcely visible growth striation.

Besides the species closely resembles

another new sp. currently being studied

(PIZZINI AND NOFRONI, submitted).

Endemic to the Fiji Is., it is also present

in other Indian-Pacific zones.

Geographical distribution: Caecum cam-

panulatum is currently known only on

Easter Is., its typical location.

Superfamily RISSOOIDEA Gray J. E., 1847

Family CAECIDAE Gray J. E., 1850

Genus Strebloceras Carpenter, 1859 [1858]

Diagnosis (BANDEL, 1996): “The protoconch is trochospirally coiled, and the teleoconch is

uncoiled forming a slightly curving tube with slowly increasing diameter. Protoconch and

teleoconch remain together during the whole life-time”.

Type species (BANDEL, 1996): According to COSSMANN (1896) Caecum edwarsil Deshayes, 1864 [Oligocene

of France]; according to GOUGEROT AND LE RENARD (1981), Strebloceras lituus Deshayes, 1861.

Strebloceras subannulatum Folin, 1879 [not Caecum (Brochina) subannulatum

Folin, 1870 (Mediterranean Sea)] (Figs. 13, 14)

Type material: 2 syntypes, NHML 1887.2.9.2308-2310.

Material examined: Original types, 2 syntypes NHML 1887.2.9.2308-2310; 23 specimens, Punta

Rosalia, east of Anakena (Apr. 1998, leg. B. Raines).

Voucher material: 2 specimens were deposited in each of the following institutions: LACM;

USNM 1018793; MNHN; NHML; NMNZ M.273208; NSMT Mo 73563; AMS C.205279; WAM

63

Iberus, 23 (1), 2005

Bo 100 um

Figure 14. Strebloceras subannulatum Folin, 1879. A: Voucher specimen LACM, gold coated, length

[»)

200 pm

3.3 mm; B: detail of septum; C: microsculpture; D: detail of aperture. SEM imaging by D. Geiger.

Figura 14. Strebloceras subannulatum Folin, 1879. A: voucher espécimen LACM, metalizado en oro, lon-

gitud 3,3 mm; B: detalle del septo; C: microescultura; D: detalle de la abertura. Imágenes al MEB por D. Geiger.

513784; and MPR. 4 shells, beach of Anakena Bay, N. of Easter Is.: almost exposed coast, picked

up among rocky bottom on the west side of the bay, among communities of Dictyotales, with

Galaxaura obtusata. (12-1-1995, leg. E. Rolán) MPR.

Type locality: Reefs of Honolulu, 40 fms (73m).

Original description: “Minute, double-

curved head, vitreous, diaphanous and

clear; oblique nucleus of spirals; an-

fractibus duobus: postea testa tubularia,

latitudine acrescens, curvam duplicem

sequens, transversim subannulata, an-

nulis latis, minutissime expressis, suba-

cutis, late separatis: oblique aperture.

Long.: 3 mm, lat.: 0 mm 5” (FoLin, 1879).

Additional description: Shell small

(average length: 3.5 mm, min. diam. 0.18

mm, max diam. 0.7 mm), whitish; larval

shell (diam. of the last whorl about 0.23

mm) is slightly trochospiral, consisting

of roughly 2 and a half whorls. The tube

is separated from the protoconch by an

incision which, when seen in side-view,

64

is horseshoe shaped (Fig. 14B); the tube

has a double curve that forms itself on

two different levels and is crossed by a

microsculpture whose abapical portion

consists of fine, sinuous growth stria-

tions that gradually transform them-

selves, as they grow, into fairly clear-cut,

visible rings on the adapical portion of

the tube. The aperture is perfectly circu-

lar, with an almost sharp, oblique edge.

The operculum and the soft parts are

unknown.

Geographical distribution: Described

from Honolulu, its distribution has now

extended to the Easter Is. as well.

Remarks: The specimens found on Easter

Is. are wholly identical to the specimens of

RAINES AND PIZZINI: Revision of the Caecidae of Easter Island (Chile)

the typical series (Figs. 13, 14) and to the

specimens from Hawaii (MPR), despite the

fact that the tube of the former is slightly

CONCLUSIONS

As a result of the present comprehen-

sive review and revision of Easter Is. Cae-

cidae, we discovered an unusual

anomaly regarding the family's endemic-

ity rate. Unlike other regions, where the

rate is fairly low among this family, over

71% of the known caecid species are

endemic to the island. This is the highest

rate ever observed within any region.

ACKNOWLEDGEMENTS

We would like to thank Luis DiSalvo

(Coquimbo, Chile) for providing material

collected during his survey of the island's

marine environment; Jerry Harasewych

(National Museum of Natural History)

for his support and loan of Rehder”s type

and voucher material; Paul Scofield and

Neil Andrews (Canterbury Museum) for

BIBLIOGRAPHY

BANDEL, K., 1996. Philogeny of the Caecidae.

Mitteilungen adem Geologischen und Paláon-

tologischen Institut der Untversitát Hamburg. 79:

539-115.

COSSMANN, A. M., 1896. Appendix no.2 au ca-

talogue illustre des coquilles fossiles de

lÉocéne des environs de Paris. Annales de la

Societe Malacologique de Belgique, 31: 1-94.

DIsALVvO, L. H., RANDALL, J. E. AND CEA, A,,

1988. Ecological Reconnaissance of Easter

Island Sublittoral Marine Environment.

National Geographic Research. 4: 451-473.

FOLIN, A. G. L. DE, (1879) 1880. On the Mo-

llusca of the H.M.S. Challenger Expedition.

The Caecidae, comprising the Genera Paras-

trophia, Watsonia, and Caecum.. Proceedings of

the Zoological Society of London (16 dec. 1879):

52; 806-812 [no pls].

GOUGEROT, L. AND LE RENAROD, J., 1981. Clefs

de determination de petites especes de gas-

teropodes de l'Éocéne du Bassin Parisien. 18.

Le genre Tenuiscala. Cahiers des Naturalistes,

37 (3): 61-68.

OLIVER, W. R. B., 1915. The Mollusca of Ker-

madec Islands. Transactions and Proceedings

of the New Zealand Institute, 47: 509-568.

wider than that of the latter; the length is

also greater, but this is due to the fact that the

specimens in question are adult.

their comments and support in providing

SEM images of Oliver's type material;

Kathie Way (British Museum of Natural

History) for providing SEM images of the

Folin type material; Daniel Geiger (Santa

Barbara Museum of Natural History) for

his time and effort on the remaining SEM

work; Lindsey Groves (Natural History

Museum of Los Angeles County) and

Michel Garcia (Sociedad de Explotación y

Exploración Marítima Orca, Ltda.) for

their continued support; Emilio Rolán

(Spain) for sending some specimens. We

also wish to thank both Ms. Caterina Ciu-

ferri and Ms. Mary Taylor, who helped us

with captions in Spanish, and Ms.

Viviana Meyohas for her linguistic assis-

tance in English, which proved necessary,

on occasion, in the exchange of corre-

spondence between the first and the sec-

ond author.

PIZZINL, M., NOFRONI, Í. AND OLIVERIO, M., 1998.

Contribution to the knowledge of the family

Caecidae. 4. The temporary septum forma-

tion of some caecid species (Caenogas-

tropoda: Rissooidea), Iberus, 16 (1): 133-140.

PIZZINI, M., 1998. Contribution to the knowledge

of the family Caecidae. 7. C. fulvum Kisch,

1959 a junior synonym of C. neocaledonicum

Folin, 1868 (Caenogastropoda: Rissooidea

Gray J. E., 1847). Argonauta, 11 (2): 33-38.

PIZZINL, M. AND NOFRONI, l. Revision of The

family Caecidae in the South-West Pacific

ocean with description of 23 new species

(Mollusca: Gastropoda) (submitted).

RAINES, B. K., 2002. Contributions to the knowl-

edge of Easter Island Mollusca. La Conchiglia.

304: 11-40.

REHDER, H. A., 1980. The marine mollusks of

Easter Island (Isla de Pascua) and Sala y

Gómez. Smithsonian Contribution to Zoology,

289: 1-167.

SCHILDER, F. A., 1965. The Geographical Dis-

tribution of Cowries (Mollusca: Gastropoda).

The Veliger, 7 (3): 171-183.

65

O Sociedad Española de Malacología —__——T— Iberus, 23 (1): 67-76, 2005

The molluscs of the intertidal algal turf in the Azores

Los moluscos del cesped algal intermareale en Azores

Sérgio P. ÁVILA* **, Ana C. SANTOS*, Ana M. PENTEADO*, Ana M.

RODRIGUES*, Inés QUINTINO* and Maria Inés MACHADO*

Recibido el 25-VII1-2004. Aceptado el 1-IV-2005

ABSTRACT

The molluscan fauna of the high intertidal algal turf on the island of Pico (Azores) was

studied in order to complement existing descriptions of the intertidal biota on these islands.

A total of 15,275 specimens belonging to 19 species were found. Five species, the gas-

tropods Alvania mediolittoralis Gofas, 1989, Omalogyra atomus (Philippi, 1841), Pisinna

glabrata (Megerle von Muhlfeld, 1824) and Skeneopsis planorbis (Fabricius O., 1780),

and the bivalve Lasaea adansoni (Gmelin, 1791), account for 98% of the total number of

specimens, P. glabrata itself being responsible for 46% of all specimens.

In places where patellid limpets are absent, the algal turf extends higher in the intertidal

zone. In such places, three very abundant molluscan species characterize molluscan

assemblages in the algal turf: Lasaea adansoni, Skeneopsis planorbis and Pisinna

glabrata. Lasaea adansoni and Skeneopsis planorbis virtually disappear in the transition

from algal turf to algal fronds, the only (very) abundant species being P. glabrata and,

with less importance, Omalogyra atomus and Sinezona cingulata (Costa O. G., 1861).

RESUMEN

Se estudia la fauna malacológica del cesped algal del intermareal superior en la isla de

Pico (Azores), con el fin de complementar los trabajos ya existentes sobre la biota inter-

mareal de estas islas. Se encontraron un total de 15275 ejemplares de 19 especies distin-

tas. Cinco, los gasterópodos Alvania mediolittoralis Gofas, 1989, Omalogyra atomus

(Philippi, 1841), Pisinna glabrata (Megerle von Muhlfeld, 1824) y Skeneopsis planorbis

(Fabricius O., 1780), y el bivalvo Lasaea adansoni (Gmelin, 1791), suman el 98% del

total de ejemplares, de ellas P. glabrata representa al 46%.

En aquellos lugares donde están ausentes los patélidos, las algas se extienden hasta nive-

les superiores en el intermareal. En estas zonas hay tres especies de moluscos que carac-

terizan las asociaciones malacológicas algales: Lasaea adansoni, Skeneopsis planorbis y

Pisinna glabrata. lasaea adansoni y Skeneopsis planorbis desaparecen en la transición

entre el cesped algal y la zona de algas frondosas. La única especie abundante aquí es P.

glabrata, y en menor medida Omalogyra atomus y Sinezona cingulata (Costa O. G.,

1861).

KEYWORDS: micromolluscs, intertidal, algal turf, Azores.

PALABRAS CLAVE: micromoluscos, intermareal, cesped algal, Azores.

* Departamento de Biologia, Universidade dos Agores, Rua da Máe de Deus, PT-9500 Ponta Delgada, Acores,

Portugal.

** CIRN (Centro de Investigacio de Recursos Naturais), Departamento de Biologia, Universidade dos Acores,

PT 9500 Ponta Delgada — Acores. E-mail: avilanotes.uac.pt

67

Iberus, 23 (1), 2005

INTRODUCTION

Located in the middle of the northern

Atlantic and therefore relatively isolated,

the oceanic islands that form the Azores

archipelago (36” 55' to 39% 45” N, 24? 45'

to 31? 17 W) are a “live-laboratory”

where patterns and processes of disper-

sion, colonization and speciation can be

studied and ecological, evolutionary and

biogeographical theories be tested.

Lajes do Pico, a small town located

in the south coast of Pico island, Azores

(Fig. 1), is one of the most interesting

places in the littoral of the Azores,

because of its very diverse marine fauna

and flora. As a consequence of a number

of biological studies (AZEVEDO, 1990,

SANTOS, 1992; AZEVEDO, RODRIGUES,

MENDIZABAL AND ARRUDA, 1995;

MORTON, BRITTON AND MARTINS, 1996,

1998: ÁviLA, 1998), this area was chosen

as worthy of protection to conserve bio-

logical diversity; for a detailed review

see ÁVILA, ELIAS AND MEDEIROS (2000).

Rocky shores in the Azores are

usually covered by an intricate mixture

of small-sized species of algae forming a

characteristic algal turf (HAWKINS,

BURNAY, NETO, CUNHA AND MARTINS,

1990; NETO, 1992; NETO AND TITTLEY,

1995). This turf is particularly effective

for a number of small animal species,

protecting them from wave exposure,

excessive temperatures and desiccation

(AZEVEDO, 1992). CHAPMAN (1955) was

the first author to study the fauna asso-

ciated to this algal turf, mainly com-

posed of Corallina spp. and he was sur-

prised by the great abundance of mol-

luscs found in a sample of 10 x 10 cm,

collected at Faial Island. AZEVEDO (1992)

studied the molluscan species composi-

tion, abundance, diversity, seasonal

variations and the effect of differences in

wave exposure on the high intertidal

algal turf of the Azores, on a temporal

scale at Sáo Miguel Island. He con-

cluded that floristic composition and

biomass of the algal turf are fundamen-

tal for the molluscan communities in

this particular habitat.

According to NETO (1992), the “algal

turf” is divided into two distinct zones,

68

differentiated by their species-composi-

tion and morphology. The higher part is

characterized by a “dense and short

tangle forming a mat, almost impossible

to separate into components” whereas

the lower part is composed of frondose

and larger algae. We follow her designa-

tion and restrict most of this study to

the upper part of the “algal turf”.

The aim of this study is to incorpo-

rate the vertical distribution of micro-

molluscs in the general zonation pattern

already described for the intertidal of

the Azores.

MATERIAL AND METHODS

Between August 7 and 11, 1995, a 30

m long transect was examined on the

gently sloping rocky shore of Lajes do

Pico, near “Poca do Pano”, in a moder-

ately exposed site (Fig. 1). The shore”

profile was drawn, following the spirit

leveling method (EMERY, 1961; HAWKINS

AND JONES, 1992) and all elevations were

related to Chart Datum, Azores (CD), by

using sea-level at the time of predicted

low-tide. Tidal range in the Azores is

small (less than 2 m), for which reason

the transect location was carefully

chosen. The selected site presents a very

gentle slope and, as a result, the transect

was 30 m long, a distance very seldom

found in Azorean intertidal shores. This

procedure minimized problems derived

from the blurring effect that occurs in

the zonation of organisms, when tran-

sects are made in sites with steeper

slopes.

Five quadrats of 25 x 25 cm were col-

lected at 5, 10, 20, 25 and 30 m along one

transect (Fig. 2). The highly uniform

species composition of the algal turf and

the large area used (25 x 25 cm instead

of the usual 10 x 10 cm) (BuLLOCck, 1995)

minimizes possible sampling problems

derived from a single transect without

replicates. After the littorinid zone that

extended from 0 to 15 m along the tran-

sect, and the barnacle zone (10-15 m), 3

quadrats of 25 x 25 cm were scraped

from the “algal turf” at 20, 25 and 30 m

and the material collected (all of the

ÁVILA ET AL.: The molluscs of the intertidal algal turf in the Azores

Graciosa

Sáo Jorge

Azores Archipelago

sáo Migue!

Atlantic

Ocean

Poca do

Pano

Figure 1. Azores archipelago (top), Pico island (bottom, left) and detail of location of Poga do

Pano at Lajes do Pico village (Pico Island).

Figura 1. Archipiélago de las Azores (arriba), isla de Pico (abajo, izquierda) y detalle de la localización

de Poga do Pano y la villa de Lajes do Pico (isla de Pico).

algae and contained sediment) put into

labelled bags. In the laboratory, this

material was washed several times and

the animals removed from the algae.

Samples were then labelled and pre-

served in 70% ethanol. The live-col-

lected molluscs were sorted, identified

and counted under a binocular dissect-

ing microscope. Dominant algae were

identified and algal dry weight (g) was

determined for each quadrat, after

drying for 48 hours at 60 *C. Abundance

of molluscs was expressed as density

(n/m?) as well as n/100 g ADW (algal

dry weight), where n is the number of

specimens of the ith species in a

quadrate.

Species authorities and synonymy of

mollusc species follow the CLEMAM

database.

69

Iberus, 23 (1), 2005

0 5 10 15

Distance (m)

SHA Littorina striata

de Melarhapbe neritoides dh Chthamalus stellatus

wo Algal turf

“ LWST

20 23 30

Figure 2. Transect performed at Poga do Pano (Lajes do Pico, Pico island) and vertical distribution

of rocky shore organisms. HWST: mean high water level at spring tides; LWST: mean low water

level at spring tides.

Figura 2. Transecto realizado en Poga do Pano (Lajes do Pico, isla de Pico) y distribución vertical de

organismos de costa rocosa. HWST: nivel medio superior del agua en mareas de primavera: LWST: nivel

medio inferior del agua en mareas de primavera.

RESULTS

Zonation: The “littoral fringe” is

bound at the top by a littorinid zone that

extends for 15 m along the transect, with

a vertical range of 1.7 m, between 0.9 m

and 2.6 m above chart datum. This is fol-

lowed by a barnacle zone (Chthamalus

stellatus (Poli)) with an extension of about

5 m along the transect (between 10 and

15 m), free of molluscs and with a vertical

range of about 0.5 m (Fig. 2). After this,

an algal turf, dominated by Corallina offic-

inalis L., covers the rocky substrate

entirely. No limpets were found in this

zone, nor molluscs that were common

elsewhere, like Stramonita haemastoma

(Linnaeus, 1766) or Mitra cornea Lamarck,

1811 (HAWKINS, CORTE-REAL, PANNACCI-

ULLI, WEBER AND BISHOP, 2000). Immedi-

ately after LWST (mean low water level

at spring tides), frondose algae (mainly

Pterocladiella capillacea (S.G. Gmelin) and

Enteromorpha muscoides (Clem.) Cre-

mades, in Cremades et Perez-Cirera

replace the coralline turf.

Molluscs: A total of 15,275 specimens

belonging to 19 species were found in the

intertidal algal turf at “Poca do Pano”

(Lajes do Pico) (Table I and Fig. 5). Five

7O

species, the gastropods Omalogyra atomus

(Philippi, 1841), Skeneopsis planorbis

(Fabricius O., 1780), Alvania mediolittoralis

Gofas, 1989 and Pisinna glabrata (Megerle

von Múhlfeld, 1824) (=P. punctulum) and

the bivalve Lasaea adansoni (Gmelin,

1791), account for 98% of the total num-

ber of specimens, P. glabrata itself being

responsible for 46% of all specimens

(Table 1). No molluscs were collected in

the middle of the barnacle zone. In

quadrats 3 and 4, located at 20 and 25 m

in the transect, three species dominated:

L. adansoni, S. planorbis and P. glabrata

(Table II). In the lowermost quadrat, lo-

cated at LWST, P. glabrata was clearly the

most abundant species, representing

about 90% of all specimens in that

quadrat. Species density and number of

specimens /100g ADW (algal dry weight)

generally decreased towards LWST, with

the exception of O. atomus and Sinezona

cingulata (Costa O. G., 1861) (=Schismope

fayalensis Dautzenberg, 1889) (Figs. 3, 4).

DISCUSSION

The site studied broadly conforms to

the zonation pattern described from the

ÁVILA ET AL.: The molluscs of the intertidal algal turf in the Azores

Table 1. Specific composition of the molluscan fauna inhabiting the intertidal algal turf and total

number of specimens collected in each quadrat. Quadrats 1-5 collected at, respectively, 5, 10, 20,

25 and 30 m along transect (see Figure 2 for further details).

Tabla I. Composición específica de la fauna de moluscos del cesped algal intermareal y número total de

ejemplares recogido en cada cuadrante. Cuadrantes 1-5 obtenidos a 5, 10, 20, 25 y 30 m a lo largo del

transecto, respectivamente (ver la Figura 2 para más detalles).

—

TaxaX Quadrats

Alvania mediolittoralis Gotas, 1989

Bittium latreillii (Payraudeau, 1826)

Bothryphallus ovummuscae (Gotas, 1990)

Cardita calyculata (Linnaeus, 1758)

Cingula trifasciata (Adams J., 1798)

Gregariella semigranata (Reeve, 1858)

(= Trichomusculus semigranatus (Reeve, 1858))

Lasaea adansoni (Gmelin, 1791)

Littorina striata King and Broderip, 1832

Manzonia unifasciata Dautzenberg, 1889

Melarhaphe neritoides (Linnaeus, 1758)

Odostomia sp.

Omalogyra atomus (Philippi, 1841)

Pisinna glabrata (Megerle von Múhlfeld, 1824)

Rissoella cf. diaphana (Alder, 1848)

Runcina cf. adriatica Thompson, 1980

Setia subvaricosa Gotas, 1990

Setia sp.

Sinezona cingulata (Costa O. 6., 1861)

Skeneopsis planorbis (O. Fabricius, 1780)

Total

O OOOO O

Y OOOO OOOOOouomnN O

rocky shores of the Azores by HAWKINS

ET AL. (1990) (Caloura, Sáo Miguel Island)

and NETO AND AZEVEDO (1990) for Flores

Island (see also NETO, 1992, and MORTON

ET AL., 1998). These authors divided the

vertical zonation of the organisms into

three main zones: an upper one, the

splash and spray zone, dominated by lit-

torinids, lichens and ephemeral (sea-

sonal) algae; a middle zone located

between HWST (mean high water level

at spring tides) and LWST (mean low

water level at spring tides), occupied by

filter-feeding barnacles in its upper levels

and by the algal turf in the lower levels;

and the lowermost zone, characterized

by the appearance of algal fronds. To this

scheme, we may add now the vertical

distribution of other molluscs besides the

littorinids. In places where patellid

limpets are absent, possibly due to

2 3 4 5 Total %

0 40 325 22 387 2,5

0 0 l 6 ] 0.0

0 | 0 0 ] 0.0

0 36 47 6 89 0.6

0 | 0 0 | 0.0

0 0 | 0 | 0.0

DAT | 4229 JE)

6 0 0 0 8 0.]

0 0 0 | | 0.0

3 0 0 0 8 0.]

0 0 3 0 3 0.0

0 9 35 156 287 1.9

DAA OSOS 6,993 45.8

0 3 0 0 3 0.0

0 0 20 0 20 0.1

0 0 0 25 25 0.2

0 0 2 12 14 0.]

0 2 ' 18 86 0.6

DA OA0OS 2 EZ 20.4

9 4246 9169 1,844 15275510010

human overexploitation (HAWKINS ET

AL., 1990; HAWKINS ET AL., 2000), the algal

turf, usually coralline dominated by

Corallina, Jania, Amphiroa or Haliptylon

spp. directly attached to the rocky sub-

strate (NETO AND TITTLEY, 1995) extends

higher in the intertidal zone, a situation

also encountered by NETO (1992). In such

places of the Azorean rocky shores, three

very abundant micromolluscan species

characterize and further define the algal

turf located in the intertidal zone: Lasaea

adansoni, Skeneopsis planorbis and Pisinna

glabrata. In the transition from algal turf

to algal fronds, Lasaea adansoni and Ske-

neopsis planorbis virtually disappear, the

only (very) abundant species being P.

glabrata and, with less importance, Omal-

ogyra atomus and Sinezona cingulata.

Most of the species found in the inter-

tidal algal turf at Lajes do Pico do not

71

Iberus, 23 (1), 2005

Table II. Density (n/m?) and number of specimens per 100 g algal dry weight (n/100g ADW) of

the molluscan fauna inhabiting the intertidal algal turf at Lajes do Pico. For location of quadrats 1,

2, 3, 4 and 5, please refer to Figure 2.

Tabla 1]. Densida (n/m?) y número de ejemplares por 100 g de peso seco de algas (n/100g ADW) de la

fauna de moluscos del cesped algal intermareal en Lajes do Pico. Para la localización de los cuadranets 1

a 5, ver la Figura 2.

Density (n/m?) n/1009 ADW

Taxa A Quadrats 2 3 4 5 3 4 5

Alvania mediolittoralis Gotas, 1989 0 0 640 57 30% 208 618 57

Bittium latreillii (Payraudeau, 1826) 0 0 0 16 9% 0 2 16

Bothryphallus ovummuscae (Gotas, 1990) 0 0 16 0 0 5 0 0

Cardita calyculata (Linnaeus, 1758) 0 0 576 152 96 188 89 16

Cingula trifasciata (Adams J., 1798) 0 0 16 0 0 5 0 0

Rissoella cf. diaphana (Alder, 1848) 0 0 48 0 0 16 0 0

Lasaea adansoni (Gmelin, 1791) 0 0 28,976 38,672 16 91432 4,595 3

Littorina striata King and Broderip, 1832 32 9% 0 0 0 0 0 0

Manzonia unifasciata Dautzenberg, 1889 0 0 0 0 16 0 0 3

Melarhaphe neritoides (Linnaeus, 1758) 80 48 0 0 0 0 0 0

Omalogyra atomus (Philippi, 1841) 0 0 1536 560 2,496 500 67 407

Pisinna glabrata (Megerle von Múhlfeld, 1824) 0 O 18,384 68,944 24,560 5,984 8,192 4,008

Runcina cf. adriatica Thompson, 1980 0 0 0 320 0 0 38 0

Setia subvaricosa Gotas, 1990 0 0 0 0 400 0 0 65

Setia sp. 0 0 0 32 192 0 4 31

Sinezona cingulata (Costa 0. 6., 1861) 0 0 32 96 1,248 10 11 204

Skeneopsis planorbis (0. Fabricius, 1780) 0 0 17712 32,048 32 5,766 3,808 5

Gregariella semigranata (Reeve, 1858) 0 0 0 16 0 0 2 0

(= Trichomusculus semigranatus (Reeve, 1858))

Table II. Maximum densities (n/m?) recorded for the Azores. 1: Feteira, Faial Island; 2: Piscina de

Santa Cruz, Flores Island, sheltered; 3: Ponta Delgada, Flores Island, exposed; 4: Caloura, Sáo

Miguel Island, intertidal, sheltered; 5: Caloura, Sío Miguel Island, intertidal, exposed; 6: Caloura,

Sao Miguel Island, infralittoral, sheltered; 7: llhéu de Vila Franca, Sio Miguel Island; 8: *Poga da

Barra”, Lajes do Pico, Pico Island, intertidal, sheltered; 9: “Poca do Pano”, Lajes do Pico, Pico

Island, intertidal, moderately exposed.

Tabla III. Densidades máximas (n/m?) encontradas en las Azores. 1: Feteira, isla de Faial; 2: Piscina de

Santa Cruz, isla de Flores, protegido; 3: Ponta Delgada, isla de Flores, expuesto; 4: Caloura, isla de Sáo

Miguel, intermareal, protegido; 5: Caloura, isla de Sáo Miguel, intermareal, expuesto; 6: Caloura, isla

de Sáo Miguel, infralitoral, protegido; 7: llhéu de Vila Franca, isla de Sáo Miguel; 8: “Poga da Barra”,

Lajes do Pico, isla de Pico, intermareal, protegido; 9: “Poga do Pano”, Lajes do Pico, isla de Pico, inter-

mareal, moderadamente expuesto.

Chapman, 1955 Neto and Azevedo, 1990 Azevedo, 1991 Bullock, 1995 Ávila, 1998 — This work

Alvania mediolittoralis 64 (2) 14 (4) 54,400 (7) 160(8) 5,200 (9)

Lasaea adansoni 12,200 (1) 17,888 (3) 724 (4) 1,674,400 (7) 1,088 (8) 38,672 (9)

Omalogyra atomus 68,046 (6) 16 (8) 2,496 (9)

Pisinna glabrata 5,975 (4) 104,100 (7) 32 (8) 68,944 (9)

Skeneopsis planorbis 2,000 (1) 4,304 (5) 129,400 (7) 16 (8) 32,048 (9)

2

ÁVILA ET AL.: The molluscs of the intertidal algal turf in the Azores

Table IV. Geographical ranges of the most abundant molluscan species inhabiting the intertidal

algal turf at Lajes do Pico (see ÁVILA, 2000 and references therein). np: non-planktotrophic type

of development (it includes direct development and lecithotrophic development); br: brooding.

Tabla IV. Distribución geográfica de las especies de moluscos más abundantes en el cesped algal interma-

real de Lajes do Pico (ver ÁVILA, 2000 y las referencias allí citadas). np: desarrollo no planctotrófico

(incluyendo desarrollos directo y lecitotrófico); br: desarrollo planctotrófico.

Species Type of development

Alvania mediolittoralis np

Omalogyra atomus np

Skeneopsis planorbis np

Pisinna glabrata np

Lasaea adansoni br

extend their distribution to the sublit-

toral. ÁviLA (2003) has established the

molluscan vertical distribution between 3

and 30 m depths for the Azorean rocky

shores covered by algal fronds. He found

that the endemic rissoids Alvania angioyi

van Aartsen, 1982, Manzonia unifasciata

(Dautzenberg, 1889) and Rissoa guernel

Dautzenberg, 1889, the also endemic

trochid Gibbula delgadensis Nordsieck,

1982, the Macaronesian Anachis avaroides

Nordsieck, 1975 and the small bivalve

Parvicardium vroomi van Aartsen,

Menkhorst and Gittenberger, 1984 were

only abundant in shallow water (3 to 5-6

m), whereas Alvanía sleursi (Amati, 1987)

was especially abundant below 20 m

depth. Bittium latreillii (Payraudeau,

1826), Tricolia pullus azorica (Dautzenberg,

1889) and Jujubinus pseudogravinae Nord-

sieck, 1973 were found in large numbers

along the whole depth-range, albeit

slightly more abundant in the lower

levels (ÁviLa, 2003). Thus, as expected

because of the different spatial architec-

ture of algae, there is a marked difference

between the molluscan species composi-

tion of the algal turf and that present in

algal fronds. In fact, none of the most

abundant molluscan species of the algal

turf is present in high numbers in algal

fronds. The total number of specimens of

Alvania mediolittoralis, Lasaea adansoni,

Geographical range

Azores, Madeira

Scandinavia, British Isles, Bay of Biscay, Portugal, Mediterranean,

Azores, Madeira, Canary Islands, Cape Verde and Ascension Island

Scandinavia, British Isles, Bay of Biscay, Portugal, Mediterranean,

Azores, Madeira, Canary Islands and Caribbean

Azores, Canary Islands and Mediterranean

Scandinavia, British Isles, Bay of Biscay, Portugal, Mediterranean,

Azores, Madeira, Canary Islands, Cape Verde, Ascension Island,

Saint Helena and Caribbean

Omalogyra atomus, Pisinna glabrata and

Skeneopsis planorbis collected in 51

quadrats (50 x 50 cm) between 3 and 30

m depth, was only 73 individuals, corre-

sponding to about 0.20% of all specimens

collected (ÁviLA, 2003). The only species

that apparently extends its vertical distri-

bution from the lower levels of the inter-

tidal algal turf down to the shallow sub-

littoral algal fronds (2-3 m depth) is the

minute Omalogyra atomus, which is the

most abundant species at 2-3 m depth

throughout the year (AZEVvEDO, 1991).

However, this species is very uncommon

at greater depths (ÁviLa, 2003). The

abundances of the algal-turf associated

molluscan fauna found at Lajes do Pico

are in the range of those found by other

authors for other islands of the archipel-

ago, being intermediate between the very

high densities found by BULLOCK (1995)

in a very sheltered place (Ilhéu de Vila

Franca, Sáo Miguel Island) and the densi-

ties found by NETO AND AZEVEDO (1990)

and AZEVEDO (1991) (see Table III).

Some species found in the quadrats

are accidental (e.g. Botryphallus and

Cingula) which live under intertidal

boulders and not in algal turf.

It is noteworthy that 4 out of the 5

most abundant molluscan species in the

intertidal of the Azores (the minute Ske-

neopsis planorbis and Omalogyra atomus,

TAS

Iberus, 23 (1), 2005

Distance (m)

1000 4------

1004

10

Number per 100g algal dry weight

20 25

Distance (m)

Alvania mediolittoralis

Cardita calyculata

Lasaea adansoni

Omalogyra atomus

Pisinna glabrata

Sinezona cingulata

A

Skeneopsis planorbis

30

Figures 3, 4. Abundance of the molluscs collected in the algal turf at “Poga do Pano” (Lajes do

Pico, Pico island). 3: n/m?; 4: n/100 g ADW. ADW: algal dry weight.

Figuras 3, 4. Abundancia de moluscos recogidos en el cesped algal en “Poga do Pano” (Lajes do Pico, isla

de Pico). 3: n/m?; 4: n/100 g ADW. ADW. peso seco de algas.

the rissoid Alvania mediollitoralis and the

anabathrid Pisinna glabrata) all have a

non-planktotrophic type of develop-

ment (ÁvILA, 2000). This type of devel-

opment is usually associated with a

restricted geographical range (SCHEL-

TEMA, 1978; JABLONSKI, 1986), which

clearly is not the case (with the excep-

tion of A. mediolittoralis) (see Table IV).

Perhaps the small size of these gastro-

pod species is an advantage for disper-

sal, as well as their location in the inter-

tidal, therefore having higher possibili-

ties of rafting.

CONCLUSIONS

In the studied algal turf located

between the barnacle zone (above) and

the frondose algae (below), three species

of molluscs are very common: Lasaea

74

adanson1, Skeneopsis planorbis and Pisinna

glabrata. In the transition from algal turf

to frondose algae, Lasaea adansoni and

Skeneopsis planorbis virtually disappear,

the only (very) abundant species being

P. glabrata and, with less importance,

Omalogyra atomus and Sinezona cingulata.

ACKNOWLEDGEMENTS

We thank Manuel Campos Marques

and Rui Pedro Ávila Marques for field

assistance. We are also grateful to Frias

Martins for correcting the manuscript.

Pedro “Bué” Cerqueira and António

Moniz (Seccáo de Geografia do Departa-

mento de Biologia da Universidade dos

Acores) are acknowledged for Figure 1.

The author also thanks the comments of

two anonymous referees. S.P. Ávila was

supported by grant SFERH/BD /5115/2001.

ÁVILA ET AL.: The molluscs of the intertidal algal turf in the Azores

Y / Í

Y a

> a

So ¿E

100 um

Figure 5. Common molluscs of the intertidal zone in the Azores. A: Alvania mediolittoralis Gotas,

1989; B: Bothryphallus ovummuscae (Gofas, 1990); C: Manzonia unifasciata Dautzenberg, 1889;

D: Cingula trifasciata (Adams J., 1798); E: Setía subvaricosa Gofas, 1990; E: Pisinna glabrata

(Megerle von Muúhlfeld, 1824); G: Bittium latreillii (Payraudeau, 1826); H: Stramonita haemas-

toma (Linnaeus, 1766); L, J: Skeneopsis planorbis (O. Fabricius, 1780; K: Melarhaphe neritoides

(Linnaeus, 1758); L, M: Cardita calyculata (Linnaeus, 1758).

Figura 5. Moluscos comunes en la zona intermareal de las Azores. A: Alvania mediolittoralis Gofas,

1989; B: Bothryphallus ovummuscae (Gofas, 1990); C: Manzonia unifasciata Dautzenberg, 1889;

D: Cingula trifasciata (Adams J., 1798); E: Setia subvaricosa Gofas, 1990; F: Pisinna glabrata

(Megerle von Múbhlfeld, 1824); G: Bittium latreillii (Payraudeau, 1826); H: Stramonita haemastoma

(Linnaeus, 1766); L, J: Skeneopsis planorbis (O. Fabricius, 1780; K: Melarhaphe neritoides (Lin-

naeus, 1758); L, M: Cardita calyculata (Linnaeus, 1758).

ES

Iberus, 23 (1), 2005

BIBLIOGRAPHY

ÁviLA, S. P., 1998. Zonacáo intertidal de uma

comunidade malacológica numa lagoa

costeira localizada na costa Sul da ilha do

Pico, Acores. Acoreana, 8(4): 436-486.

ÁviLA, S. P., 2000. Shallow-water marine mol-

luscs of the Azores: biogeographical rela-

tionships. Arquipélago. Life and Marine Sci-

ences. Supplement 2 (Part A): 99-131.

ÁviLA, S. P., 2003. The littoral molluscs (Gas-

tropoda, Bivalvia and Polyplacophora) of

Sáo Vicente, Capelas (Sáo Miguel Island,

Azores): ecology and biological associations

_ to algae. Iberus, 21(1): 1-23.

ÁVILA, S. P., ELIAS, R. B. AND MEDEIROS, J., 2000.

Parque Natural Regional da Plataforma Costeira

. das Lajes do Pico (Agores). Proposta de imple-

mentacáo. Amigos dos Acores, Ponta Del-

gada, 48 pp.

AZEVEDO, J. M. N., 1990. Microgastrópodes.

Expedicáo Acores 89. Ecologia e Taxonomia do

Litoral Marinho. Relatório Preliminar, 1: 54-59.

AZEVEDO, J. M. N., 1991. Estudo das comunidades

malacológicas fitais do litoral em Sáo Miguel,

Acores. MSc. Thesis, Universidade dos Acores.

AZEVEDO, J. M. N., 1992. Algae-associated ma-

rine molluscs in the Azores. Biological Jour-

nal of the Linnean Society, 46: 177-187.

AZEVEDO, J. M. N., RODRIGUES, J. B., MEN-

DIZABAL, M. AND ARRUDA,L. M., 1995. Study

of a sample of Dusky groupers, Ephinephelus

marginatus (Lowe, 1834) caught in a tide pool

at Lajes do Pico, Azores. Boletim do Museu

Municipal do Funchal, Suplemento 4: 55-64.

BULLOCK, R. C., 1995. The distribution of the

molluscan fauna associated with the intertidal

coralline algal turf of a partially submerged

volcanic crater, the Ilhéu de Vila Franca, Sáo

Miguel, Azores. In Martins, A. M. de F. (Ed.):

The marine fauna and flora of the Azores.

Proceedings of the Second International

Workshop of Malacology and Marine Biol-

ogy, Vila Franca do Campo, Sáo Miguel,

Azores. Acoreana, Suplemento 4: 9-55.

CHAPMAN, G., 1955. Aspects of the fauna and

flora of the Azores. VI. The density of animal

life in the Coralline alga zone. Annals and

Magazine of Natural History, 12 (8): 801-805.

CLEMAM. Check List of European Marine Mo-

llusca. Unitas Malacologica, Internet Re-

sources for Malacologists (http: //

www.mnkhn.fr /base /malaco.html) [last ac-

cessed 10th of August, 2004].

EMERY, K. O., 1961. A simple method of mea-

suring beach profiles. Limnology and Oceanog-

raphy, 6: 90-93.

HAWKINS, S. J. AND JONES, H. D., 1992. Marine

Field Course Guide. 1. Rocky Shores. Marine

Conservation Society. Immel Publishers, Lon-

don, 144 pp.

7Ó

HAWKINS, S. J., BURNAY, L. P., NETO, A. L,

TRISTAO DA CUNHA, R. AND MARTINS, A. M.

de F., 1990. A description of the zonation

patterns of molluscs and other important

biota on the south coast of Sáo Miguel,

Azores. In Martins, A. M. de F. (Ed.): The ma-

rine fauna and flora of the Azores. Proceed-

ings of the First International Workshop of

Malacology Sáo Miguel, Azores. Acoreana,

Suplemento 2: 21-38.

HAWKINS, S. J., CORTE-REAL, H. B. S. M., PAN-

NACCIULLI, F. G., WEBER, L. C. AND BISHOP,

J. D. D., 2000. Thoughts on the ecology and

evolution of the intertidal biota of the

Azores and other Atlantic islands. In: Jones,

M. B., Azevedo, J. M. N., Neto, A. L, Costa,

A. C. and Martins, A. M. de F. (Eds.): Is-

land, Ocean and Deep-Sea Biology. Hydro-

biología, 440: 3-17.

JABLONSKI, D., 1986. Larval ecology and

macroevolution in marine invertebrates. Bul-

letin of Marine Science, 39(2): 565-587.

MORTON, B., BRITTON, J. C. AND MARTINS, A. M.

DE F., 1996. The Lajes do Pico marsh: a fur-

ther case for coastal conservation in the

Acores. Acoreana, 8(2): 183-200.

MORTON, B., BRITTON, J. C. AND MARTINS, A. M.

DE F., 1998. Ecología Costeira dos Acores. So-

ciedade Afonso Chaves, Ponta Delgada, 249

PB:

NETO, A. LI, 1992. Contribution to the taxonomy

and ecology of the Azorean benthic marine

algae. Biological Journal of the Linnean Society,

46: 163-176.

NETO, A. L, AND AZEVEDO, J. M. N., 1990. Con-

tribuicáo para o estudo dos padróes de zon-

acáo litoral da ilha das Flores. Relatórios e Co-

municacóes do Departamento de Biologia. Ex-

pedicáo Científica Flores/89 (Relatório

Preliminar), 18: 89-102.

NETO, A.I.,, AND TITTLEY, L, 1995. Structure and

zonation of algal turf communities on the

Azores: a numerical approach. Boletim do

Museu Municipal do Funchal, Suplemento 4:

487-504.

SANTOS, R. S., 1992. Proteccáo e conservacáo do

meio marinho nos Acores. In Saldanha, L.,,

Ré, P. and Martins, A. M. de F. (Eds.): Cen-

tenaire de la Derniere Campagne

Océanographique du Prince Albert de

Monaco aux Acores á bord de L'Hirondelle.

Acoreana, Suplemento 3: 107-122.

SCHELTEMA, R. S., 1978. On the relation between

dispersal of pelagic larvae and the evolution

of marine prosobranch gastropods. In

Battaglia, B. and Beardmore, J. A. (Eds.): Ma-

rine Organisms: Genetics, Ecology, and Evo-

lution. NATO Conference Series. Series IV:

Marine Sciences. Plenum Press, New York.

O Sociedad Española de Malacología Iberus, 23 (1): 77-82, 2005

Calyptraea capensis Tomlin, 1931 (Gastropoda, Calyp-

traeidae), a valid species from South Africa

Calyptraea capensís "Tomlin, 1931 (Gastropoda, Calyptraeidae), una

especie válida de Sudáfrica

Emilio ROLÁN*

Recibido el 14-VII1-2004. Aceptado el 14-IV-2005

RESUMEN

La especie del género Calypiraea de Sudáfrica considerada hasta la actualidad como C. chinen-

sis se estudia y compara con otras especies de Africa occidental y meridional y de Europa. Se con-

cluye que se trata de una especie diferente, cuyo nombre debe ser Calyptraea capensis Tomlin, 1931.

ABSTRACT

The species of the genus Calyptraea from South Africa up to now considered to be C. chinen-

sis is studied and compared with other European and West and South African species. lt is con-

cluded that it is a different species whose name should be Calyptraea capensis Tomlin, 1931.

KEY WORDS: Calyptraeidae, Calyptraea capensis, Calyptraea chinensis, South Africa.

PALABRAS CLAVE: Calyptraeidae, Calyptraea capensis, Calyptraea chinensis, Sudáfrica.

INTRODUCTION

The study of the genus Calyptraea in

the Eastern Atlantic (ROLÁN, 2004) has

shown the existence of three species. In

this work it was mentioned that the

South African species known as Calyp-

traea chinensis Linné, 1758 could proba-

bly be a different species.

TOMLIN (1931) described C. capensis.

Agreeing with Tomlin, TURTON (1932:

154) comments that the South African

species called C. chinensis by several

authors, should be called C. capensis.

Nevertheless, this species has been

recorded from South Africa in the most

recent literature under the name C. chi-

nensis (BARNARD, 1963; KENSLEY, 1973;

KILBURN AND RIPPEY, 1982; STEYN AND

Lussi, 1998, among others.

In order to check if this species was dif-

ferent from those previously described in

ROLÁN (2004), material from the Natal

Museum was examined and compared

with that previously studied and illustrated

from the West African coasts. The conclu-

sion is that it is a different species and

which should not be called C. chinensis but

C. capensis. The diagnostic characters of

this species are shown in the present work.

Abbreviations:

NMW: National Museum of Wales,

Cardiff

NM: Natal Museum, Pietermaritzburg

sp: specimen with soft parts

s: Shell

e Museo de Historia Natural, Campus Universitario Sur. 15782 Santiago de Compostela.

e-mail: emiliorolanGinicia.es

M7

Iberus, 23 (1), 2005

RESULTS

Family CALYPTRAEIDAE Lamarck, 1809

Genus Calyptraea Lamarck, 1799

Calyptraea capensis Tomlin, 1931 (Figs. 1-13, 17-19)

Calyptraea capensis Tomlin, 1931. Annals of the Natal Museum, 6: p. 428, pl. 36, fig. 6. [Type locality:

Jeffreys Bay, South Africa].

Calyptraea chinensis auct. non Linné, 1758.

Type material: Holotype (NMW (1955.158.01006, Figs. 7, 8).

Material studied (all from NM): W1093, 1 sp, S. of Cape St. Blaize, 34? 43.5” S 22” 09 E, 88 m,

NMDP (Africana) St. A16562D; S8593, 8 s, False Bay, off Falk Bay (34* 08.5” S 18* 28” E), 18 m,

sand with Pyura, NMDP CD29 (10.1V.1991); 7316, 1 s, Port Elizabeth; Amsterdam Neek, on

Zwartkops River; B2887, 47 s, Pondoland Coast, H. Becker coll (1978); B2886, 13 s, Port Alfred, H.

Becker coll. (1978); D2211, 2 sp, Algoa Bay; 1 sp, living inside outer lip of hermit crab shell

Amalda obtusa, 49 m, W of Stilbaai, 34” 29” S 21” 16' E; V603, 2 sp, W of Stilbaai, (34? 29” S; 21” 16'

E), 49 m, B2886, 13 s, Port Alfred; 6025, 16 s, Saldanha Bay; 7316, 1 s, Port Elisabeth, Amsterdam

Nock, Zwartkops River; E6457, 3 s, Cape Agulhas, Struinbari, Caravan Park; S8121, 2 s, SW of

Struis Bay (34” 46.6” S; 20? 09.4” E), 34 m, S3557, 1 s, False Bay (34* 11.2” S; 18” 35.8” E), 40 my;

B8224, 5 s, E. Cape, off East London (33* 04.9” S; 27” 54” E), 70 m; A3454, 4 s, False Bay, Muizen-

berg; B7690, 1 s, Port Elisabeth; B7677, 1 s, Port Elisabeth, Humewood Beach, S6353, 3 s, Agulhas

Bank, W of Martha Point (34? 29.5” S; 20? 33.3” E, 28 m, V186, 3 s, 3 s, Agulhas Bank, W of Martha

Point (34? 24.8” S; 20? 53.4 E, 31 m; C4677, 1 s, Transkei, of Qolora (32? 39.7" S; 28” 28.2' E), 50 m;

B7870, 1 s, E. Cape, off East London (33* 01.0” S; 27” 57.3” E), 30 m,; B8466, 7 s, E. Cape, off East

London (33* 06.2” S; 27” 52.4” E), 70 m; E8887, 2 s, Zululand, off Matigulu River mouth (29* 21.6'

S; 31? 57.1” E), 300 m, S9109, 1 s, SW Cape, SE of Cape Infanta (34” 24.8” S; 20? 53.4” E), 31 m;

C4309, 1 s, Transkei, off Stony Point (32* 32.8” S; 28” 38.2 E), 70 m; C3467, 1 s, Transkei, Kei River;

S3624, 3 s, Cape, off Simonstown (34* 10.7” S; 18” 28.4” E), 31 m; E8940, 1 s, Zululand, off

Matigulu River mouth (29? 22.2” S; 31? 57.2” E), 350 m, A3456, 4 sp, Fish Hock; 9068, 2 s, Jeffreys

Bay; B7657, 2 s, Durban; A3455, 44 s, False Bay, Simonstown dredgings; B2887, 47 S, Pondoland

Coast; S8443, 1 sp, False Bay, SE Seal Island (34* 11.5' S; 18? 37.1” E), 43 m; W1094, W1096, 2 sp

with eggs, E. Cape, off East London (33* 06.8” S; 27” 51.4' E), 10 m.

Description: The shell (Figs. 1-8) has

been described in TOMLIN (1931) and

KILBURN AND RIPPEY (1982). The color

may be white, pink, light brown or

violet, and the surface is usually

smooth, but some shells from deeper

water from Zululand have some

spicules. Sometimes, its external surface

is undulating towards the sutural

depression. Protoconch (Figs. 12, 13)

with about 1*/2 whorls, a diameter of

about 900 um, and a nucleus with a

diameter of 85-100 um, the protoconch

surface is totally smooth. A few radial

lines appear in the teleoconch of some

shells, near the protoconch. Dimensions:

up to 29.0 mm (in lot 9068).

Soft parts: No live material was

examined, but some lots had been pre-

served in alcohol. Two males (10.3, 11.1

78

and 10.3 mm) with dry soft parts were

hydrated in water with soap, and show

that the animal (Figs. 17, 18) has a

similar aspect to that of C. chinensis from

Europe. Only the penis (Fig. 19) seems

to be different: it was placed behind the

right tentacle, being elongated, appar-

ently flat, and placed on the dorsum of

the animal and towards the posterior

part. The tip is a little wider and finishes

in a fine prolongation. Two females of

13 and 15 mm examined, have a small

vestigial penis, elongated and sharp

pointed without any wider part.

Radula (Figs. 9-11) taenioglossate;

rachidian tooth narrow and with a

prominent central cusp on the edge,

with 4, sometimes 5-6, smaller cusps at

each side. Lateral tooth with a promi-

nent and a little wider cusp at the con-

ROLÁN: Calyptraea capensis Tomlin, 1931, a valid species from South Africa

Figures 1-8. Calyptraea capensis. 1-3: 13.4 mm, Cape St. Blaize, 88 m (NM W1093); 4-6: 13.5

mm, Saldanha Bay (NM 6025); 7, 8: holotype, 10.2 mm (MNW).

Figuras 1-8. Calyptraea capensis. 1-3: 13,4 mm, Cabo St. Blaize, 88 m (NM WI1093); 4-6: 13,5

mm, Bahía Saldanha (NM 6025); 7, 8: holotipo, 10,2 mm (MNW).

YO

Iberus, 23 (1), 2005

Figures 9-11. Radula of Calyptraea capensis. 9: female, specimen of 13.8 mm (NM W1093); 10,

11: males of 11.1 and 10.3 mm (NM S8593). Figures 12-16. Protoconchs. 12, 13: C. capensis, off

East London; 14: C. chinensis, Vigo; 15: C. imexpectata, Guinea-Conakry; 16: C. africana, Ivory

Coast.

Figuras 9-11. Rádula de Calyptraea capensis. 9: hembra, ejemplar de 13,8 mm (NM W1093); 10,

11: machos de 11,1 and 10,3 mm (NM $8593). Figuras 12-16. Protoconchas. 12, 13: C. capensis,

costa de East London; 14: C. chinensis, Vigo; 15: C. inexpectata, Guinea Conakry; 16: C. africana,

Costa de Marfil.

80

ROLÁN: Calyptraea capensis Tomlin, 1931, a valid species from South Africa

Figure 17-19. Drawings of soft parts. 17: detail of the head of a female with a 15 mm shell, ventral

view; 18: a male of 10 mm with penis, dorsal view; 19: detail of the penis.

Figuras 17-19. Dibujo de las partes blandas. 17: detalle de la cabeza de una hembra con una concha de

15 mm, en visión ventral; 18: macho de 10 mm con el pene, en visión dorsal; 19: detalle del pene.

fluence of the internal and the external

edges. Internal edge with few wide

cusps (usually 1-2), while the external

has between 5-7. Marginal teeth elon-

gate, curved, sharp pointed, the internal

ones with two edges with 4-6 cusps,

while the external teeth have only 1-2

cusps on their internal part.

Spawn: several spawns were exam-

ined. They have a similar disposition to

that of C. chinensis, the capsules being

more irregular. A specimen of 15 mm

diameter had a spawn of 12 capsules,

each one between 2-3 mm, with

between 20-60 embrionary larvae inside.

Remarks: The holotype has an eroded

protoconch but all other characters fit

well with most of the studied material.

Some variations were found in the

numerous shells examined: most shells

are depressed, but some are extremely

elevated, even higher than wide. We

could not determine if these elevated

forms are a different species, due to the

lack of protoconch in the specimens

examined.

The shell of C. capensis is rather dif-

ferent from the West African species. C.

chinensis is less pointed at the apex of

the shell, and has external prominences

more frequently. The penis is different,

with a spoon shape and a wide prolon-

gation in opposition. The rachidian and

marginal teeth of the radula are very

similar, but the lateral teeth have more

cusps (3-4) on the internal side; the pro-

toconch (Fig. 14) is wider, shorter (1

against 1!/2 whorls) and the nucleus

larger; furthermore, the protoconch of C.

capensis is smooth and the beginning of

the teleoconch has some inconspicuous

radial lines never present in C. chinensis.

C. africana Rolán, 2004 has a larger

shell, more depressed, a little ovoid,

usually translucent white, frequently

with elevated external prominences; the

penis is bilobed at its extreme, with one

of the lobes formed by two masses

together, and a narrow worm-like fila-

ment near the tip; the protoconch (Fig.

16) is rather similar, but the nucleus is

somewhat smaller. The radula has more

cusps in all the teeth.

C. inexpectata Rolán, 2004 has a

smaller shell, sometimes with a squa-

mous sculpture, white or brown in

colour, but never violet; the protoconch

(Fig. 15) is smaller, with a wider nucleus

and several evident sulcii at the begin-

ning of the teleoconch. All the radular

teeth have more cusps.

There are no problems in separating

the remaining South African species of

Calyptraea: C. helicoidea Sowerby, 1883,

has a subscalariform profile, with a

narrow but distinct umbilicus and the

dorsal surface has strong, narrow, rather

oblique radial ribs. C. aurita Reeve, 1859

also has an umbilicus, the columella is

widely reflexed, and the protoconch has

clear radiating spiral lirae (see

BARNARD, 1963: 73).

81

Iberus, 23 (1), 2005

ACKNOWLEDGEMENTS

The author wishes to thank the fol-

lowing: D. G. Herbert of the Natal

Museum, who loaned the material

studied in this work; Harriet Wood of

the National Museum of Wales, Cardiff,

for the information and photographs of

the holotype of C. capensis; Jesús

BIBLIOGRAPHY

BARNARD, K. H., 1963 “1962”. Contribution to

the knowledge of the South African marine

Mollusca. Part III. Gastropoda: Proso-

branchiata: Taenioglossa. Annals of the South

African Museum, 47(1): 1-199.

KENSLEY, B., 1973. Sea-shells of Southern Africa.

Gastropods. Maskew Miller Ltd., Cape Town,

236 pp.

KILBURN, R. AND RIPPEY, E., 1982. Sea Shells of

Southern Africa. MacMillan South Africa, Jo-

hannesburg, 249 pp.

ROLÁN, E., 2004. The genus Calyptraea (Gas-

tropoda, Caenogastropoda, Calyptraeidae) in

the East Atlantic. Iberus, 22(2): 51-79.

82

Méndez of the Centro de Apoyo Cientí-

fico y Tecnológico a la Investigación

(CACTI) of Vigo University, who made

the SEM micrographs, and Jesús S. Tron-

coso of the Department of Ecology of

the University of Vigo, for the digital

optical photographs.

STEYN, G. S. AND Lusst, M., 1998. Marine Shells

of South Africa. Ekogilde, Hartebeespoort,

264 pp.

TOMLIN, J. R. LE B., 1931. On South African ma-

rine mollusca, with descriptions of new gen-

era and species. Annals of the Natal Museum,

6: 415-450.

TURTON, D. $. O., 1932. The marine shells of Port

Alfred South Africa. Oxford University Press,

London, 331 pp., 70 pls.

O Sociedad Española de Malacología

Iberus, 23 (1): 83-86, 2005

Notas breves

Additional information on the biology of Argonauta argo

(Cephalopoda: Octopoda) in the Mediterranean Sea from

gastrointestinal contents of Risso's dolphin

Aportación a la biología de Argonauta argo (Cephalopoda:

Octopoda) en el mar Mediterráneo a partir del contenido gastroin-

testinal del delfín de Risso

Carmen BLANCO*, Ángeles RADUÁN* and Juan Antonio RAGA*

Recibido el 27-VII1-2004. Aceptado el 14-IV-2005

KEY WORDS: Argonauta argo, Grampus griseus, cephalopod biology, Mediterranean.

PALABRAS CLAVE: Argonauta argo, Grampus griseus, biología cefalópodos, Mediterráneo.

INTRODUCTION

Argonguta argo Linnaeus, 1758 is a

cosmopolitan epipelagic octopod living

in tropical and subtropical waters. It is

sexually dimorphic, with dwarf males

and females lodged in calcareous shells

that serve as brood chambers (GUERRA,

1992). Despite its abundance, whole

animals are rarely caught, so that their

life cycle is not completely known. The

reproduction of this species shows a

protracted and continuous spawning

which has been defined as asynchro-

nous ovulation with monocycle spawn-

ing reproductive strategy (ROCHA,

GUERRA AND GONZÁLEZ, 2001).

Living individuals of this species are

rarely found in the Mediterranean Sea

(BOLETZKY AND CENTELLES, 1979; BELLO

AND RIZZI, 1990; POPPER, BARASH AND

GALIL, 1990). Continuous spawning is

believed to take place in the eastern

Mediterranean Sea (LAPTIKHOVSKY AND

SALMAN, 2003), but the life span and

period of reproduction of the species

continue to be unknown. The aim of this

paper is to provide some data on the

biology of this little known Mediter-

ranean species by means of remains col-

lected from the gastrointestinal contents

of Risso's dolphins, Grampus griseus

(Cuvier, 1812) (Cetacea: Delphinidae), a

teuthivorous predator (KRUSE, CALD-

WELL AND CALDWELL, 1999).

MATERIAL AND METHODS

The data analysed in this paper

came from gastrointestinal contents of

eleven Rissos dolphins, Grampus

griseus, stranded on the west Mediter-

ranean coast, between 40" 25' N, 00% 32*

* Cavanilles Institute of Biodiversity and Evolutionary Biology, University of Valencia, P. O. Box 22085, 46100

Valencia, Spain. E-mail: carmen.blancoCuv.es

83

Iberus, 23 (1), 2005

Ve Zea da Má as

61171 3,49

O A. argo

=0— 6. griseus 3

0

10, AL A2

Figure 1. Annual distribution of specimens of Argonauta argo from gastrointestinal contents of

Mediterranean Risso's dolphin. N: number of A. argo; n: number of dolphins.

Figura 1. Distribución anual de los ejemplares de Argonauta argo obtenidos del contenido gastrointesti-

nal del delfín de Risso en el Mediterráneo.

W and 37* 35' N, 00% 45 E, collected

from April 1987 to January 2003. All gas-

trointestinal tracts were stored deep-

frozen (-20%C) and their contents subse-

quently washed through a 0.2 mm mesh

sieve and preserved in 70% ethanol;

glycerine was added to the preservative

solution for cephalopod beaks.

The beaks were identified according

to SMALE, CLARKE, KLAGES AND

ROELEVELD (1993). Cephalopod mantle

length was estimated from hood length

of lower beak (SMALE ET AL., 1993), due

to the frequent rotten condition of the

crest. Some underestimating of hood

length and, hence, mantle length may be

assumed because of the digestive action.

RESULTS

In the gastrointestinal contents of

Risso's dolphins, 336 lower beaks of

Argonauta argo were found, hood length

range: 1.31-5.87 mm. Distribution

through the time period for which data

were available (see Figure 1) shows a

higher number in early spring. The esti-

mated mantle lengths are shown in

Figure 2. Significant differences were

found in the size of specimens through-

out the year (nested ANOVA (Fos25 =

11.1, P < 0.001). The frequency distribu-

tion of beak length (HL) in the months

of maximum abundance, March and

84

April, is shown in Figure 3; significant

length distribution differences were

found between these two months (Kol-

mogorov-Smirnov test, maximum dif-

ferences between samples= 0.268, N

(192, 99), P< 0.001), 25.6-28.7 mm and

31.8-35.0 mm being the estimated

mantle length of modal class in March

and April, respectively. In all available

samples there were individuals whose

mantle length was longer than 35 mm.

An oviscapte with eggs (modal value

size 1.22-1.42 mm length) was found in

the intestine of a dolphin stranded in

January; this value may be underesti-

mated due to the possible shrinking

action of preservation in alcohol.

DISCUSSION

The high number of beaks from A.

argo gathered in this study, in contrast

with the scarcity of their catches (see the

Introduction), corroborates the effective-

ness of teuthophagous predators as

cephalopod collectors (BELLO, 1996).

All remains of A. argo found in this

study, apart from one collected in April,

were ascribed to female specimens

based on their size, since dwarf males

only reach 10 mm of dorsal mantle

length (GUERRA, 1992). In accordance

with the smallest female size at maturity

in the Mediterranean, i. e. 32 mm ML

BLANCO ET AL.: Argonauta argo in gastrointestinal contents of Risso's dolphin

3

El March

£

>

> April

LAS DORADO A

Months

EDO LAS E SA l,

Beak length (HL mm)

Figure 2. Annual length distribution of estimated mantle length (ML) of Argonauta argo from gas-

trointestinal contents of Mediterranean Risso's dolphin. Box-and-whisker-plot (TUKEY, 1977).

Figure 3. Frequency distribution of lower beak length (HL) of Argonauta argo in March and April.

Figura 2. Distribución anual de la longitud estimada del manto (ML) de Argonauta argo a partir del

contenido gastrointestinal del delfín de Risso en el Mediterráneo. Gráfico “Box-and-whisker” (TUKEY,

1977). Figura 3. Distribución de la frecuencia de la longitud de la mandíbula inferior (HL) de Argo-

nauta argo en marzo y abril.

(LAPTIKHOVSKY AND SALMAN, 2003), ma-

ture females were present in all sampled

seasons. Such a find and the occurrence

of fertilized eggs in the brooding cham-

ber from January, in addition to brood-

ing females collected in the Adriatic in

November (BELLO AND RIZZ1, 1990) and

mature females found in the eastern

Mediterranean Sea during winter (LAp-

TIKHOVSKY AND SALMAN, 2003), show

that the spawning period for Mediter-

ranean A. argo extends longer than pre-

viously believed (from May to October

according to GUERRA, 1992). Differences

BIBLIOGRAPHY

BELLO, G., 1996. Teuthophagous predators as

collectors of oceanic cephalopods: the case of

the Adriatic Sea. Bollettino Malacologico, 32 (1-

4): 71-78.

BELLO, G. AND RIZZI E., 1990. Comportamiento

di tre femmine di Argonauta argo in acquario

(Cephalopoda: Argonautidae). Atti della Societa

Italiana di Scienze Naturali e del Museo Cívico

di Storia Naturale di Milano, 131: 450-452.

BOLETZKY, S. V. AND CENTELLES, J., 1979. Argo-

nauta argo (Mollusca, Cephalopoda) dans la

région de Banyuls-sur-mer. Vie et Milieu, 28-

29 (4): 659-660.

GUERRA, A., 1992. Mollusca. Cephalopoda. In

Ramos, M. A. et al. (eds.): Fauna Ibérica, vol.

1. Museo Nacional de Ciencias Naturales,

CSIC, Madrid, 327 pp.

in the size of specimens through the

year, especially in the two consecutive

months could indicate either seasonality

or a preferential period of spawning

time.

ACKNOWLEDGEMENTS

This study was supported by Con-

sellería de Territorio y Vivienda, Gener-

alitat Valenciana, and Dirección General

de Conservación de la Naturaleza, Min-

isterio de Medio Ambiente.

KRUSE, S., CALDWELL, D. K. AND CALDWELL, M.

C., 1999. Risso's dolphin Grampus griseus (G.

Cuvier, 1812). In Ridgway, S. H. and Harrison,

R. (eds.): Handbook of Marine Mammals, Vol.

6, pp. 183-212. Academic Press, London.

LAPTIKHOVSKY, V. AND SALMAN, A., 2003. On

reproductive strategies of the epipelagic oc-

topods of the superfamily Argonautidea

(Cephalopoda: Octopoda). Marine Biology,

142: 321-326.

PopPPER, D., BARASH, A. AND GALIL, B. S., 1990.

Argonauta argo — A rare occurrence off the

shores of Israel. Israel Journal of Zoology, 37:

51-53.

ROCHA, F., GUERRA, A. AND GONZÁLEZ, A. F,,

2001. A review of reproductive strategies in

cephalopods. Biological Review, 76: 291-304.

85

Iberus, 23 (1), 2005

SMALE, M. J., CLARKE, M. R., KLAGES, T. W. TUKEY, J. W. 1977. Bob-and-whisker-Plot. In

AND ROELEVELD, A. C., 1993. Octopod beak Explanatory data analysis, pp. 39-43. Addison-

identification — resolution at a regional level Wesley, Massachusetts.

(Cephalopoda, Octopoda: Southern Africa).

South African Journal of Marine Science, 13:

269-293.

86

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Dendrodoris limbata (Cuvier, 1804)

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* Proofs will be sent to the author for correcting errors. At this stage no stylistic changes will be accepted. Pay special

attention to references and their dates in the text and the Bibliography section, and also to numbers of Figures and

Tables appearing in the text.

* Fifty reprints per article will be supplied free of charge. Additional reprints must be ordered when the page proofs

are returned, and will be charged at cost. NO LATER orders will be accepted.

LA SOCIEDAD ESPAÑOLA DE MALACOLOGÍA

Junta Directiva desde el 14 de noviembre de 2000

Presidente Emilio Rolán Mosquera

Vicepresidente Diego Moreno Lampreave

Secretario Luis Murillo Guillén

Tesorero Jorge J. Otero Schmitt

Avda. de las Ciencias s/n, Campus Universitario, 15706 Santiago

de Compostela, España

Editor de Publicaciones Gonzalo Rodríguez Casero

Apartado 156, 33600, Mieres del Camino, Asturias, España

Bibliotecario Rafael Araujo Armero

Museo Nacional de Ciencias Naturales, CSIC, c/ José Gutierrez

Abascal 2, 28006 Madrid, España

Vocales Ramon M. Álvarez Halcon

Benjamín Gómez Moliner

Eugenia María Martínez Cueto-Felgueroso

Jesús Souza Troncoso

José Templado González

La Sociedad Española de Malacología se fundó el 21 de agosto de 1980. La sociedad se registró como una aso-

ciación sin ánimo de lucro en Madrid (Registro N* 4053) con unos estatutos que fueron aprobados el 12 de

diciembre de 1980. Esta sociedad se constituye con el fin de fomentar y difundir los estudios malacológicos

mediante reuniones y publicaciones. A esta sociedad puede pertenecer cualquier persona o institución interesada

en el estudio de los moluscos.

SEDE SOCIAL; Museo Nacional de Ciencias Naturales, c/ José Gutierrez Abascal 2, 28006 Madrid, España.

CUOTAS PARA 2005:

Socio numerario (en España): 40 euros

(en Europa) 40 euros

(fuera de Europa): 48 euros

Socio estudiante (en España): 23 euros

(en el extranjero): 29 euros

Socio Familiar: (sin recepcion de revista) 4 euros

Socio Protector: (mínimo) 48 euros

Socio Corporativo (en Europa): 48 euros

(fuera de Europa): 54 euros

INSCRIPCIÓN: 6 euros, además de la cuota correspondiente.

A los socios residentes en España se les aconseja domiciliar su cuota. Todos los abonos deberán enviarse al

Tesorero (dirección reseñada anteriormente) el 1 de enero de cada año. Los abonos se harán sin recargos para la

sociedad y en favor de la Sociedad Española de Malacología y no de ninguna persona de la junta directiva. Aque-

llos socios que no abonen su cuota anual dejarán de recibir las publicaciones de la Sociedad. Los bonos de ins-

cripción se enviarán junto con el abono de una cuota anual al Tesorero.

A los residentes en el extranjero se les ruega que abonen su cuota mediante giro postal en euros (internatio-

nal postal money orders in euros sent to the Treasurer). Members living in foreing countries can deduce 6 euros

if paid before 15 April.

Cada socio tiene derecho a recibir anualmente los números de /berus, Reseñas Malacológicas y Noticiarios que

se publiquen.

UI UOAIAN

ÍNDICE

Iberus | 23 (1) 2005

YADAV, R. P, TIWARI, S. AND SINGH, A. Toxic effects of taraxerol extracted from Codiaeum varie-

gatum stem-bark on target vector snail Lymnaea acuminata and non-target fish

loxicidad del taraxerol extraído del tallo de Codiaeum variegatum sobre el caracol Lymnaea

ACUIMINALA Y SODIO PECES A 1-13

MÁRQUEZ, R., ARRÉBOLA, J. R. Y DELGADO, R. Un avance sobre la composición y microestruc-

tura de la concha de /berus gualtierianus morfotipo gualtierianus (Linnaeus, 1758) (Gastro-

poda: Helicidae)

Notes on the composition and microstructure of the shell of Iberus gualtierianus morphotype

gualtierianus (Linnaeus, 1758) (Gastropoda: Helicidae) ici 15-24

SINGH, D., YADAV, R. P. AND SINGH, A. Changes in phospholipid and lipid peroxidation 15%

due to latex of Croton tiglium in freshwater snail Lymnaea acuminata

Cambios en los niveles de peroxidación de fosfolípidos y lípidos por efecto del latex de Croton

tiglium sobre el molusco dulceacuícola Lymnaea acuminata ooo cicioccccccnoos 25:34

GIORDANO, D., BOTTARI, T. AND RINELLI, P. Cephalopod assemblages caught by trawling along

the Southern Tyrrhenian Sea (Central Mediterranean)

Asociaciones de cefalópodos capturados por la pesca de arrastre comercial en el Tirreno meridio-

nal (Mediterráneo central) 0 do AI aO Ea EN 33-42

SÁNCHEZ-SANTOS, A. Onchidoris neapolitana (Delle Chiaje, 1844) (Gastropoda: Nudibranchia:

Onchidorididae): una nueva especie de molusco para la fauna andaluza

Onchidoris neapolitana (Delle Chiaje, 1844) (Gastropoda: Nudibranchia: Onchidorididae):

a new molluscan species for the Andalousian fauna ooo .... 43-48

RAINES, B. AND PIZZINL M. Contribution to the knowledge of the family Caecidae: 16. Revision

of the Caecidae of Easter Island (Chile) (Caenogastropoda: Rissooidea Gray J. E., 1847)

Contribución al conocimiento de la familia Caecidae: 16. Revisión de los Caecidae de la Isla de

Pascua (Chile) (Caenogastropoda: Rissooidea J. E. Gray 1847) ici 49-65

ÁVILA, S. P, SANTOS, A. C., PENTEADO, A. M., RODRIGUES, A. M., QUINTINO, L. AND MACHA-

DO, M. I. The molluscs of the intertidal algal turf in the Azores

Los moluscos del cesped algal intermareale en Azores iii 67-76

ROLÁN, E. Calyptraea capensis Tomlin, 1931 (Gastropoda, Calyptraeidae), a valid species from

South Africa j

Calyptraea capensis Tomlin, 1931 (Gastropoda, o una especie válida de

SUAAICA da AU 77-82

Notas breves

BLANCO, C., RADUÁN, A. AND RAGA, J. A. Additional information on the biology of Argonauta

argo (Cephalopoda: Octopoda) in the Mediterranean Sea from gastrointestinal contents ef

Risso's dolphin

Aportación a la biología de Argonauta argo (Cephalopoda: Octopoda) en el mar Mei

a partir del contenido gastrointestinal del delfín de RissO lll 83-86

ISSN 0212-3010