BIODIVERSITY JOURNAL

2014,5 (2): 93-374

Quaternly scientific journal

edited by Edizioni Danaus,

viaV. Di Marco 41, 90143 Palermo, Italy

www.biodiversityjournal.com

biodiversityjournal@gmail.com

Official authorization no. 40 (28. 1 2.20 1 0)

ISSN 2039-0394 (Print Edition)

ISSN 2039-0408 (Online Edition)

Study of birds in Libya. The coast line of Libya has different kinds of wetlands such as

islands, lagoons, lakes, salt marshes and bays (Defos et al., 2001). These habitats provide

foraging sites and good shelters for migratory birds during their migration from Europe

and Asia to Africa and back. Moreover, some of these sites provide residential and nesting

ground for some species. However, Libya, with its relatively dry climate, is perceived as

having comparatively few wetlands and waterbirds. It is also, omithologically speaking,

the least known country of Mediterranean Africa (Smart et ah, 2006). Recent reviews list

3 1 7 species, of which approximately 25% are breeders (Toschi, 1 969; Bundy, 1 976). They

account for about 75% of the avifauna of Libya, passing from the western Palearctic

region to their southern winter quarters. Some of these also occur here as breeding species.

There are many other papers reporting about waterbirds (e.g. Gaskell, 2005; Smart et ah,

2006; Hering, 2009; Essghaier et ah, 2009; Etayeb & Essghaier, 2012; Etayeb et ah,

2013a, b and 2014), as well as some unpublished reports (e.g. Defos du Ran et ah, 2001,

Etayeb, 2002). However, there are only few experienced ornithologists in Libya, which

has resulted in deficiency of information on Libyan birds during last three decades.

However during the last decade, interest in the Libyan wetlands has increased. In the mid-

nineties the UNEP Mediterranean Action Plan (MAP), adopted a “Protocol concerning

Special Protected Areas (SPA) and Biological Diversity in the Mediterranean”. Annex II

of the Protocol includes a “List of Endangered or Threatened Species”, including 15

waterbirds, for which a Bird Action Plan has been prepared (UNEP MAPRAC/SPA,

2003). Libya was involved in the framework of the Barcelona Convention and has also

ratified the Ramsar Convention, and in 2002, two wetlands in the Jebel Akhdar area were

designated. In 2005, Libya signed the African-Eurasian Waterbird Agreement (AEWA),

under the umbrella of the Convention on Migratory Species (CMS) (Smart et ah, 2006).

Consequently, a regular wintering water-bird census for Libyan wetlands has launched in

year 2005.

Khaled S. Etayeb. Zoology Dept. Faculty of Science, Tripoli University-P.o.Box: 1 3227;

e-mail: khaledetayeb@yahoo.com

Reviewed by: Prof. Essghaier M. F. A.

Middle: Gaber Own, photo K.S. Etayeb. Down: Common Sandpiper, Aclitis hypoleucos

(Linnaeus, 1758), Lybia, Tripoli, Alhadba, photo Bouras. Cover: Little Ringed plover,

Charadrius dubius curonicus Gmelin, 1 789, Lybia, Tripoli, Alhadba, photo Bouras.

Biodiversity Journal, 2014, 5 (2): 95-96

Monograph

Preface

The Malacological Pontine Meeting, San Felice Circeo, Italy

Bruno Fumanti

Via del Villaggio 108, 04010 Sabaudia, Latina, Italy; e-mail: brano.fomanti@libero.it

Received 28.01.2014; accepted 21.03.2014; printed 30.06.2014

Proceedings of the Seventh Malacological Pontine Meeting, October 5 th - 6 th , 2013 - San Felice Circeo, Italy

As part of the knowledge of the biodiversity of

the pontine area and in particular of the Circeo Na-

tional Park, in 2007 the writer, then director of the

“Museo Civico del Mare e della Costa Marcello

Zei” of Sabaudia, thought appropriate to include as

one of the institutional activities of the museum the

deepening of research at malacological level, hith-

erto restricted only to a few scientific papers related

to molluscs of Mount Circeo (Lepri, 1909), of the

area of Terracina (Sacchi & Girod, 1968), with some

footnotes to broader works on the biodiversity of

coastal lakes of Latium, as in the case of the Paola

(or Sabaudia) Lake (Brunelli & Cannicci, 1934) in

which is also described a new species (Rissoa

sabaudiae Coen, 1934 never reported again) or

parts of research related to marine areas overlook-

ing the coast and the Pontine islands (Gravina et al.,

1992) (Figs. 1,2).

Equally sporadical are the reports on fossil mol-

luscs: regarding Mount Circeo, Strombus bubonius

Lamarck, 1822 and Tritonidea viverrata Kiener,

1757 were reported by Blanc (1940) in the Guttari

Cave and Durante & Settepassi (1974) have reported

some species found in the “Grotta delle Capre”. The

malacofauna of the fossil dunes of the islands of

Ponza and Ventotene (Pontine Archipelago) was

finally studied by Sacchi (1952).

With these premises in May 2007 was therefore

organized the “Prima Giomata di Studi Malacologici

Pontini” in which, among other things Danilo Vani

presented the first report of Gibbula nebulosa

(Philippi, 1841) in the Pontine coast. In subsequent

years, given the interest of Italian malacologists for

this initiative, the name of the event changed to

“Convegno Malacologico Pontino” arranged in two

days of study and funded by the Lazio Region

through the “Museo Civico del Mare e della Costa”

of Sabaudia, which remained until 2012, the place

of the event , replaced in 2013 (and since October

2014) both as funding agency as seat by the Munic-

ipality of San Felice Circeo, together with the Pontine

Naturalistic Malacological Association “Malakos

2002” as co-financing organization.

In the seven editions (2007-2013) of the Meet-

ing implemented to date, numerous speakers both

professionals of academic origin and amateurs have

dealed with the most important topics regarding

malacology from the point of view of systematics,

ecology, biogeography, molecular biology, paleon-

tology, etc..., by several reports, some of which

have resulted both in scientific and popular

publications.

There have been numerous reports that have

dealt with topics of systematics and biogeography

concerning the Pontine Area, and in particular: the

first report of Gibbida nebulosa (Philippi, 1841) in

the Pontine coast (Danilo Vani, VII Meeting) the

study of land molluscs of Mount Circeo (Alessan-

dro Hallgass & Angelo Vannozzi, II Meeting) and

of the islands Ventotene and Santo Stefano (Alessan-

Bruno Fumanti

Figure 1. Mount Circeo (photo S. Moncado).

Figure 2. Pontine Archipelago (photo S. Moncado).

dro Hallgass & Angelo Vannozzi, VI Meeting), the

study of molluscs of the pontine dunes (Antonio In-

candela, II Meeting), preliminary (Andrea Bassi, III

Meeting) and final (Bruno Fumanti, VII Meeting)

reports on the marine molluscs of Zannone, the dis-

tribution of the genus Onoba H. & A. Adams, 1852

(Rissoidae) in pontine waters (Bruno Amati & Italo

Nofroni, IV Meeting), a study of nudibranchs from

“Canale Romano” of the Paola Lake (Giulia Furfaro,

Armando Macali & Paolo Mariottini, V Meeting),

the study of molluscs in the sands of beach nour-

ishment on the coast of Terracina (Luigi Giannelli,

VII Meeting), the study of the Mollusca from the

“Secca dei Mattoni”, Pontine Archipelago (Fabio

Crocetta, Giuseppe Fasulo, Italo Nofroni and Arturo

Facente, IV Meeting) and the study of the Pleis-

tocene continental molluscs in the Pontine Plain

(Daniele Gianola, III Meeting).

It follows therefore that these regular meetings

have contributed in some way to increase our knowl-

edge of biodiversity, even if limited to malacofauna,

of this area, which includes among other things a

National Park in many ways unique in the world.

ACKNOWLEDGEMENTS

The Author and the organizers of the Seventh

Malacological Pontine Meeting wish to thank the

Municipality of San Felice Circeo and in particular

the Mayor Giovanni Petrucci, the Chairman of the

Education Egidio Calisi, the delegate of the Mayor

Franco Domenichelli and “Malakos 2002” (Asso-

ciazione Macologica Naturalistica Pontina) in the

person of its president Claudio Buccarella for the

financial support to the realization of the meeting.

REFERENCES

Blanc A.C., 1940. Relazione sull’attivita scientifica

dell’Istituto di Paleontologia Umana durante gli anni

1938-1940. Rivista di Antropologia, 33: 1-4.

Brunelli G. & Canniccci G., 1934. Notizie prelimi-

nari sulle caratteristiche chimiche e biologiche del Lago

di Sabaudia. Rendiconti della Reale Accademia Nazionale

dei Lincei, 19: 345-351.

Durante S. & Settepassi S., 1974. Livelli marini e

molluschi tirreniani alia Grotta delle Capre. Memorie

dell’Istituto Italiano di Paleontologia Umana, 2.

Gravina M.F., Smriglio C. & Ardizzone G.,1992.

Benthos di fondo mobile delle isole pontine 1 . Molluschi.

Oebalia suppl. 17: 355-357.

Lepri G., 1909. Contributo alia conoscenza dei mol-

luschi terrestri e d’acqua dolce del Lazio. Bollettino della

Societa Zoologica Italiana, 18: 347-444.

Sacchi C., 1952. I molluschi delle dune fossili nelle

isole ponziane nei rapporti con la malacofauna vivente.

Atti della Societa Italiana di Scienze Naturali e del

Museo Civico di StoriaNaturale di Milano, 91: 240-250.

Sacchi C. & Girod A., 1968. I molluschi d’acqua

dolce di alcune stazioni presso Terracina, ricerche eco-

logiche e faunistiche. Bollettino di Pesca, Piscicultura e

Idrobiologia, 23: 1-32.

Biodiversity Journal, 2014, 5 (2): 97-106

Monograph

Contribution to the knowlegde of the benthic molluscan

thanatocoenosis of Zannone Island (Pontine Archipelago,

Latium, Italy)

Bruno Fumanti

Via del Villaggio 108, 04010 Sabaudia, Latina, Italy; e-mail: brano.fumanti@libero.it

ABSTRACT During the period May 2008-September 2012 were investigated some sediment samples col-

lected by scuba diving at various depths in the waters surrounding the Island of Zannone (Pon-

tine Archipelago, Latium, Italy). Altogether 280 taxa belonging to 156 genera were identified.

KEY WORDS Mollusca; thanatocoenosis; Zannone Island; Italy.

Received 28.01.2014; accepted 21.03.2014; printed 30.06.2014

Proceedings of the Seventh Malacological Pontine Meeting, October 5 th -6 th , 2013 - San Felice Circeo, Italy

INTRODUCTION

This study originates from the collaboration of

many malacologists, professional and otherwise,

who have joined a project for the study of marine

Mollusca of the Island of Zannone. The project has

been realized on proposal of “Museo Civico del

Mare e della Costa” in Sabaudia with funding from

the “Regione Lazio”. The idea of studying the ma-

rine Mollusca of Zannone was pondered on occa-

sion of an excursion organized at the conclusion of

the "Second Malacological Pontine Meeting" held

in Sabaudia on May 2008.

A quick analysis of some sediment samples

taken during scuba diving carried out by some par-

ticipants revealed a malacofauna relatively rich so

much to suggest a project for a more comprehensive

and careful study. The result was the creation of a

working group organized and coordinated by some

members of MALAKOS 2002 (Associazione

Malacologica Naturalistica Pontina), who, thanks

to the funds received, have been able to carry out

several samplings of bottom sediments and then

distribute the collected material to all the research

participants.

The broad involvement of many malacologists,

professional and otherwise, who collaborated in this

research, even if preliminary, is the first concerning

an island in the Pontine Archipelago and this makes

us hope in the possibility of achieving further

malacological studies on the coasts, the lakes and

island of the Pontine region.

The writer of this work even participating ac-

tively in the organization of the research has merely

reorganized what elaborated by all the participants,

listed below in alphabetic order: Silvia Alfinito

(Sabaudia, Italy), Andrea Bassi (Ravenna, Italy),

Enzo Campani (Livorno, Italy), Luigi Giannelli

(Terracina, Italy), Saverio Moncado (Latina, Italy),

Italo Nofroni (Rome, Italy), Marco Oliverio (Rome,

Italy), Angela Pierullo (Rome, Italy), Ermanno

Quaggiotto (Vicenza, Italy), Carlo Sbrana (Pisa,

Italy), Peter Sossi (Mondovi, Italy), Piergiorgio

Trillo (Rome, Italy); Daniele Trono (Copertino,

Italy), Danilo Van (Rome, Italy), Angelo Vannozzi

(Rome, Italy).

Bruno Fumanti

MATERIAL AND METHODS

The Pontine Islands are an archipelago in the

Tyrrhenian Sea off the west coast of Italy. The

islands were collectively named after the largest

island in the group, Ponza; the other islands in the

archipelago are Palmarola, Zannone, Gavi, Ventotene

and Santo Stefano.

Zannone is the most northern island of the

archipelago. Uninhabited, since 1979 is part of

the National Park of Circeo. It is a “green island”,

covered with a typical Mediterranean maquis.The

Island of Zannone has an area of approximately 1 06

ha and is located 6 miles from the Island of Ponza,

on which it administratively depends. Altogether 8

grit samples were examined:

2008, May: 40°58’28” E; 13 o 09’49” N

-18 m, -20 m

2009, June: 40°57’55” E; 13 o 02’59” N

-20m, -26m, -50m

2011, October: 40°57’54” E; 13 o 03’10” N

-18m, -20 m

2012, September: 40°07’55” E; 13°02 , 46” N

-20 m

The nomenclature of the species has been

updated according to WoRMS Editorial Board

(2014).

RESULTS

The analysis of samples distributed to the research

participants led to the identification of 280 taxa

(Table 1).

Polypla-

cophora

Gastro-

poda

Bivalvia

Scapho-

poda

Total

Ordines

Familiae

Genera

110

156

Species

221

280

Table 1. Diversity of the fauna of molluscs of Zannone

Island (Ponthine Archipelago, Latium, Italy).

Taxonomic list

Classis POLYPLACOPHORA Gray, 1821

Ordo CHITONIDA Thiele, 1909

Familia CHITONIDAE Rafmesque, 1815

Genus Chiton Linnaeus, 1758

Chiton ( Rhyssoplax ) corallinus Risso, 1 826

Chiton (. Rhyssoplax ) olivaceus Spengler, 1797

Familia ISCHNOCHITONIDAE Dali, 1889

Genus Ischnochiton Gray, 1 847

Ischnochiton {Ischnochiton) rissoi (Payraudeau,

1826)

Familia AC ANTHO CHITONIDAE Pilsbry, 1893

Genus Acanthochitona Gray, 1821

Acanthochitona crinita (Pennant, 1777)

Acanthochitona fascicularis (Linnaeus, 1767)

Classis GASTROPODA Cuvier, 1797

Ordo PATELLOGASTROPODALindenberg, 1986

Familia PATELLIDAE Rafmesque, 1815

Genus Patella Linnaeus, 1758

Patella caerulea Linnaeus, 1758

Patella rustica Linnaeus, 1758

Patella ulyssiponensis Gmelin, 1791

Ordo VETIGASTROPODA Salvini-Plawen, 1980

Familia FISSURELLIDAE Fleming, 1822

Genus Diodora J.E. Gray, 1821

Diodora gibberula (Lamarck, 1822)

Diodora graeca (Linnaeus, 1758)

Genus Emarginula Lamarck, 1801

Emarginula adriatica O.G. da Costa, 1830

Emarginula huzardii Payraudeau, 1 826

Emarginula octaviana Coen, 1939

Emarginula punctulum Piani, 1980

Emarginula tenera Locard, 1892

Familia SCISSURELLIDAE Gray, 1847

The benthic molluscan thanatocoenosis ofZannone Island (Pontine Archipelago, Latium, Italy)

Genus Scissurella d’Orbigny, 1824

Scissurella costata d’Orbigny, 1824

Familia HALIOTIDAE Rafmesque, 1815

Genus Haliotis Linnaeus, 1758

Haliotis tuberculata lamellosa Lamarck, 1 822

Haliotis tuberculata tuberculata Linnaeus, 1758

Familia TROCHIDAE Rafmesque, 1815

Genus Clanculus Monfort, 1810

Clanculus corallinus (Gmelin, 1791)

Clanculus cruciatus (Linnaeus, 1758)

Clanculus jus sieui (Payraudeau, 1826)

Genus Jujubinus Monterosato, 1884

Jujubinus exasperatus (Pennant, 1777)

Jujubinus gravinae (Dautzenberg, 1881)

Jujubinus striatus (Linnaeus, 1758)

Genus Gibbula Risso, 1826

Gibbula ardens (Salis Marschlins, 1793)

Gibbula guttadauri (Philippi, 1836)

Gibbula rackctti (Payraudeau, 1 826)

Gibbula turbinoides (Deshayes, 1835)

Gibbida umbilicaris (Linnaeus, 1758)

Gibbida varia (Linnaeus, 1758)

Familia CALLIOSTOMATIDAE Thiele, 1924

Genus Calliostoma Swainson, 1840

Calliostoma zizyphinum (Linnaeus, 1758)

Familia TURBINIDAE Rafmesque, 1815

Genus Bolma Risso, 1 826

Bolma rugosa (Linnaeus, 1767)

Familia SKENEIDAE Clark, 1851

Genus Skenea Fleming, 1 825

Skenea catenoides (Monterosato, 1877)

Genus Dikoleps Hoisaeter, 1968

Dikoleps marianae Rubio, Dantart et Luque, 1998

Dikoleps templadoi Rubio, Dantart et Luque, 2004

Dikoleps umbilicostriata (Gaglini, 1987)

Genus Skeneoides Waren, 1992

Skeneoides exilissima (Philippi, 1 844)

Fig. 1 . Zannone Island (Ponthine Archipelago, Latium, Italy).

Familia CHILODONTIDAE Wenz, 1938

Genus Danilia Brusina, 1865

Danilia tinei (Calcara, 1839)

Familia PHASIANELLIDAE Swainson, 1840

Genus Tricolia Risso, 1 826

Tricolia pullus (Linnaeus, 1758)

Tricolia speciosa (von Miihlfeldt, 1824)

Tricolia tenuis (Michaud, 1 829)

Familia COLLONIIDAE Cossmann, 1917

Genus Homalopoma Carpenter, 1864

Homalopoma sanguineum (Linnaeus, 1758)

Ordo NERITIMORPHA Golikov et Starobogatov,

1975

F amilia NERITID AE Rafmesque, 1815

Genus Sm aragdia Issel, 1869

Smaragdia viridis (Linnaeus, 1758)

Ordo CAENOGASTROPODA Cox, 1960

Familia CERITHIIDAE Fleming, 1822

Genus Cerithium Bruguiere, 1789

Cerithium lividulum Risso, 1826

Cerithium renovatum Monterosato, 1884

Cerithium vulgatum Bruguiere, 1792

Genus Bittium Gray, 1 847

Bittium latreillii (Payraudeau, 1826)

100

Bruno Fumanti

Bittium reticulatum (da Costa, 1778)

Bittium submammillatum (De Rayneval et

Ponzi,1854)

Familia PLANAXIDAE Gray, 1850

Genus Fossarus Philippi, 1841

Fossarus ambiguus (Linnaeus, 1758)

Familia SILIQUARIIDAE Anton, 1838

Genus Tenagodus Guettard, 1770

Tenagodus obtusus (Schumacher, 1817)

Familia TURRITELLIDAE Loven, 1847

Genus Turritella Lamarck, 1799

Turritella communis Risso, 1826

Turritella turbona Monterosato, 1877

Familia TRIPHOR1DAE J.E. Gray, 1847

Genus Marshallora Bouchet, 1985

Marshallora adversa (Montagu, 1803)

Genus Monophorus Grillo, 1877

Monophorus perversus (Linnaeus, 1758)

Genus Metaxia Monterosato, 1884

Metaxia metaxa (Delle Chiaje, 1828)

Familia CERITHIOPSIDAE H. Adams et A.

Adams, 1853

Genus Cerithiopsis Forbes et Hanley, 1850

Cerithiopsis barleei Jeffreys, 1 867

Cerithiopsis jeffreysi Watson, 1885

Cerithiopsis tub ercularis (Montagu, 1803)

Genus Nanopsis Cecalupo et Robba, 2010

Nanopsis nana (Jeffreys, 1867)

Genus Krachia Baluk, 1975

Krachia cylindrata (Jeffreys, 1885)

Familia EPITONIIDAE Berry, 1910

Genus Epitonium Roding, 1798

Epitonium clathrus (Linnaeus, 1758)

Familia EULIMIDAE Philippi, 1853

Genus Ersilia Monterosato, 1872

Ersilia mediterranea (Monterosato, 1869)

Genus Pa rvi o ris Ware n , 1981

Parvioris ibizenca (Nordsieck, 1968)

Genus Sticteulima Laseron, 1955

Sticteulima jeffreysiana (Brusina, 1869)

Genus Vitreolina Monterosato, 1884

Vitreolina curva (Monterosato, 1874)

Vitreolina philippi (de Rayneval et Ponzi, 1854)

Familia LITTORINIDAE Children, 1834

Genus Mel arhaphe Menke, 1828

Melarhaphe neritoides (Linnaeus, 1758)

Familia CINGULOPSIDAE Fretter et Patil, 1958

Genus Eatonina Thiele, 1912

Eatonina ochroleuca (Brusina, 1 869)

Eatonina pumila (Monterosato, 1884)

Genus Tubbreva Ponder, 1965

Tubbreva micrometrica (Aradas et Benoit, 1876)

Familia RISSOIDAE J.E.Gray, 1847

Genus Rissoa Desmarest, 1814

Rissoa auriscalpium (Linnaeus, 1758)

Rissoa guerinii Recluz, 1 843

Rissoa lia (Monterosato, 1884)

Rissoa monodonta Philippi, 1836

Rissoa rodhensis Verduin, 1985

Rissoa scurra (Monterosato, 1917)

Rissoa similis Scacchi, 1836

Rissoa variabilis (von Miihlfeldt, 1 824)

Rissoa ventricosa Desmarest, 1814

Rissoa violacea Desmarest, 1814

Genus Pusillina Monterosato, 1884

Pusillina inconspicua (Alder, 1844)

Pusillina philippi (Aradas et Maggiore, 1844)

Pusillina radiata (Philippi, 1836)

Genus Setia H. Adams et A. Adams, 1852

Setia ambigua (Brugnone, 1873)

Setia maculata (Monterosato, 1 869)

The benthic molluscan thanatocoenosis ofZannone Island (Pontine Archipelago, Latium, Italy)

101

Genus Alvania Risso, 1826

Alvania cfr. aeoliae Palazzi, 1988

Alvania beanii (Hanley in Thorpe, 1844)

Alvania cimex (Linnaeus, 1758)

Alvania cancellata (da Costa, 1778)

Alvania clathrella (Seguenza, 1903)

Alvania claudioi Buzzurro et Landini, 2007

Alvania dictyophora (Philippi, 1844)

Alvania discors (Allan, 1818)

Alvania geryonia (Nardo, 1847)

Alvania hirta (Monterosato, 1844)

Alvania hispidula (Monterosato, 1844)

Alvania lineata Risso, 1826

Alvania lucinae Oberling, 1970

Alvania mamillata Risso, 1826

Alvania scabra (Philippi, 1844)

Alvania settepassii Amati et Nofroni, 1985

Alvania sororcula Granata-Grillo, 1877

Alvania subareolata Monterosato, 1869

Alvania subcrenulata (Bucquoy, Dautzenberg et

Dollfus, 1884)

Alvania tenera (Philippi, 1844)

Genus Crisilla Monterosato, 1917

Crisilla cfr. aartseni (Verduin, 1984)

Crisilla beniamina (Monterosato, 1884)

Crisilla marioni Fasulo et Gaglini, 1987

Crisilla semistriata (Montagu, 1808)

Genus Manzonia Brusina, 1870

Manzonia crassa (Kanmacher, 1798)

Genus Obtusella Cossmann, 1921

Obtusella intersecta (S.Wood, 1857)

Genus Onoba H. Adams et A. Adams, 1852

Onoba dimassai Amati et Nofroni, 1991

Genus Peringiella Monterosato, 1878

Peringella elegans (Locard, 1892)

Genus Rissoina d’Orbigny, 1840

Rissoina bruguieri (Payraudeau, 1 826)

Familia BARLEEIDAE J.F. Gray, 1857

Genus Barleeia W. Clark, 1853

Barleeia unifasciata (Montagu, 1803)

Familia CAECIDAE Gray, 1850

Genus Caecum Fleming, 1813

Caecum auriculatum de Folin, 1868

Caecum clarkii Carpenter, 1859

Caecum sub annul atum de Folin, 1870

Caecum trachea (Montagu, 1803)

Familia HYDROBIIDAE Stimpson, 1865

Genus Hydrobia Hartmann, 1821

Hydrobia acuta (Drapamaud, 1805)

Familia TORN ID AE Sacco, 1896

Genus Torn us Turton et Kingston, 1830

Tornus subcarinatus (Montagu, 1803)

Familia T RUN CAT ELLID AE J.E. Gray, 1840

Genus Truncatella Risso, 1 826

Truncatella subcylindrica (Linnaeus, 1767)

Familia VERMETIDAE Rafmesque, 1815

Genus Petaloconchus Lea, 1 843

Petaloconchus glomeratus (Linnaeus, 1758)

Genus Vermetus Daudin, 1800

Vermetus granulatus (Gravenhorst, 1831)

Vermetus rugulosus Monterosato, 1878

Vermetus triquetrus Bivona-Bemardi, 1832

Familia APORRHAIDAE J.E. Gray, 1850

Genus Aporrhais da Costa, 1778

Aporrhais pespelecani (Linnaeus, 1758)

Familia VANIKORIDAE J.E. Gray, 1840

Genus Megalomphalus Bmsina, 1871

Megalomphalus azonus (Brusina, 1865)

Megalomphalus disciformis (Granata-Grillo,

1877)

Familia CALYPTRAEIDAE Lamarck, 1809

Genus Crepidula Lamarck, 1799

Crepidula unguiformis Lamarck, 1822

Genus Calyptraea Lamarck, 1799

Calyptraea chinensis (Linnaeus, 1758)

102

Bruno Fumanti

Familia VELUTINIDAE J.E. Gray, 1840

Genus Lamellaria Montagu, 1815

Lamellaria perspicua (Linnaeus, 1758)

Familia CYPRAEIDAE Rafmesque, 1815

Genus Luria Jousseaume, 1884

Luria lurida (Linnaeus, 1758)

Familia NATICIDAE Guilding, 1834

Genus Euspira Agassiz in Sowerby, 1837

Euspira nitida (Donovan, 1804)

Genus Notocochlis Powell, 1933

Notocochlis dillwynii (Payraudeau, 1826)

Familia ATLANTIDAE Rang, 1829

Genus Atlanta Lesueur, 1817

Adanta brunnea J.E.Gray,1850

Adanta peronii Lesueur, 1817

Familia MURICIDAE Rafmesque, 1815

Genus Bolinus Pusch, 1837

Bolinus brandaris (Linnaeus, 1758)

Genus Hexaplex Perry, 1810

Hexaplex trunculus (Linnaeus, 1758)

Genus Ocinebrina Jousseaume, 1880

Ocinebrina aciculata (Lamarck, 1822)

Ocinebrina edwardsii (Payraudeau, 1 826)

Ocinebrina reinai Bonomolo et Crocetta, 2012

Genus Muricopsis Bucquoy et Dautzenberg, 1882

Muricopsis cristata (Brocchi, 1814)

Genus Coradiophila H. Adams et A. Adams,, 1853

Coradiophila meyendorffu (Calcara, 1845)

Familia MARGINELLIDAE Fleming, 1828

Genus Granulina Jousseaume, 1888

Granulina bouched Gofas, 1992

Granulina mediterranea Landau, La Perna et

Marquet, 2006

Granulina marginata (Bivona, 1832)

Granulina occulta (Monterosato, 1869)

Genus Volvarina Hinds, 1844

Volvarina mitrella (Risso, 1 826)

Familia CYSTISCIDAE Stimpson, 1865

Genus Gibberula Swainson, 1840

Gibberula miliaria (Linnaeus, 1758)

Gibberula philippii (Monterosato, 1878)

Gibberula turgidula (Locard et Caziot, 1900)

Familia MITRIDAESwainson, 1829

Genus Mitra Lamarck, 1798

Mitra cornicula (Linnaeus, 1758)

Familia CO STELL ARIID AE MacDonald, 1860

Genus Vexillum Roding, 1798

Vexillum ebenus (Lamarck, 1811)

Vexillum savignyi (Payraudeau, 1826)

Vexillum tricolor (Gmelin, 1791)

Familia BUCCINIDAE Rafmesque, 1815

Genus Euthria Gray, 1850

Euthria cornea (Linnaeus, 1758)

Genus Chauvetia Monterosato, 1884

Chauvetia mamillata (Risso, 1826)

Chauvetia procerula (Monterosato, 1889)

Chauvetia turritellata (Deshayes, 1835)

Chauvetia sp.

Genus Pisania Bivona-Bemardi, 1832

Pis ania striata (Gmelin, 1791)

Genus Pollia J.E. Gray, 1834

Pollia dorbignyi (Payraudeau, 1 826)

Pollia scabra Locard, 1892

Pollia scacchiana (Philippi, 1 844)

Familia NASSARIIDAE Iredale, 1916

Genus Nassarius Dumeril, 1805

Nassarius nitidus Jeffreys, 1867

Genus Ciclope Risso, 1 826

Ciclope neritea (Linnaeus, 1758)

Familia COL UMBELLIDAE Swainson, 1840

The benthic molluscan thanatocoenosis ofZannone Island (Pontine Archipelago, Latium, Italy)

103

Genus Columbella Lamarck, 1799

Columbella rustica (Linnaeus, 1758)

Familia FASCIOLARIIDAE J.E. Gray, 1853

Genus Fusinus Rafmesque, 1815

Fusinus pulchellus (Philippi, 1 844)

Fusinus syracusanus (Linnaeus, 1758)

Familia CONIDAE Fleming, 1822

Genus Conus Linnaeus, 1758

Conus ventricosus Gmelin,1791

Familia FIORAICLAVIDAE Bouchet, Kantor,

Sysoev et Puillandre, 2011

Genus Haedropleura Bucquoy, Dautzenberg et

Dollfus, 1883

Haedropleura secalina (Philippi, 1 844)

Familia CLATHURELLIDAE H. Adams et A.

Adams, 1858

Genus Clathromangelia Monterosato, 1 844

Clathromangelia granum (Philippi, 1 844)

Familia MITROMORPHIDAE Casey, 1904

Genus Mitromorpha Carpenter, 1 865

Mitromorpha mediterranea Milfsud, 2001

Mitromorpha melitensis (Milfsud, 1993)

Familia MANGELIIDAE P. Fischer, 1883

Genus Bela J.E. Gray, 1847

Bela menkhorsti \ anAartsen, 1988

Genus Mangelia Risso, 1 826

Mangelia multilineolata (Deshayes, 1835)

Mangelia striolata Risso, 1 826

Mangelia taeniata (Deshayes, 1835)

Mangeria unifasciata (Deshayes, 1835)

Familia RAPHITOMIDAE Bellardi, 1875

Genus Raphitoma Bellardi, 1 847

Raphitoma densa (Monterosato, 1884)

Raphitoma echinata (Brocchi, 1814)

Raphitoma horrida (Monterosato, 1884)

Raphitoma leufroyi (Micaud, 1828)

Raphitoma linearis (Montagu, 1803)

Raphitoma pseudohystrix (Sykes, 1906)

Raphitoma purpurea (Montagu, 1803)

Ordo HETEROSTROPHAP. Fischer, 1885

Familia OMALOGYRIDAE G.O. Sars, 1878

Genus Omalogyra Jeffreys, 1859

Omalogyra simplex O.G. Costa, 1861

Familia CORNIROSTRIDAE Ponder, 1990

Genus Tomura Pilsbry et MacGinty, 1946

Tomura depressa (Granata-Grillo, 1877)

Familia RISSOELLIDAE Gray, 1850

Genus Rissoella Gray, 1847

Rissoella inflata (Monterosato, 1880)

Rissoella opalina (Jeffreys, 1848)

Familia PYRAMIDELLIDAE Gray, 1840

Genus Chrysallida Carpenter, 1856

Chrysallida emaciata (Brusina, 1866)

Chrysallida excavata (Philippi, 1836)

Chrysallida intermixta (Monterosato, 1884)

Chrysallida interstincta (J. Adams, 1797)

Chrysallida moolenbeeki Amati, 1987

Chrysallida penchynati (Bucquoi, Dautzenberg

et Dollfus, 1883)

Genus Odostomella Bucquoy, Dautzenberg et

Dollfus, 1883

Odostomella doliolum (Philippi, 1844)

Genus Euparthenia Thiele, 1931

Euparthenia humboldti (Risso, 1826)

Genus Eulimella Forbes et M’Andrew,1846

Eulimella acicula (Philippi, 1836)

Genus Megastomia Monterosato, 1884

Megastomia conoidea (Brocchi, 1814)

Genus Odostornia Fleming, 1813

Odostomia carrozai Van Aartsen, 1987

Odostornia eulimoides Hanley, 1844

Odostomia lukisii Jeffeys, 1859

Odostomia scalaris Mac Gillivray, 1 843

104

Bruno Fumanti

Odostomia turrita Hanley, 1844

Odostomia unidentata (Montagu, 1803)

Genus Ondina De Folin, 1870

Ondina vitrea (Brusina, 1866)

Genus Parthenina Bucquoy, Dautzenberg et

Dollfus, 1883

Parthenina dollfusi Kobelt, 1903

Genus Tnrbonilla Risso, 1826

Turhonilla pumila Seguenza G., 1876

Tnrbonilla striatula (Linnaeus, 1758)

Familia AMATHIN1DAE Ponder, 1987

Genus Clathrella Recluz, 1864

Clathrella clathrata (Philippi, 1 844)

Familia MURCHISONELLIDAE Casey, 1904

Genus Ebala Gray, 1 847

Ebala nitidissima (Montagu, 1803)

Ordo CEPHALAPSIDEA Fischer, 1883

Familia HAMINOEIDAE Pilsbry, 1895

Genus Haminoea Turton et Kingston in Carrington,

1830

Haminoea hydatis (Linnaeus, 1758)

Genus Weinkauffia Monterosato, 1884

Weinkauffia turgidula (Forbes, 1884)

Familia RETUSIDAE Thiele, 1925

Genus Retusa T. Brown, 1827

Retusa mammillata (Philippi, 1836)

Retusa truncatula (Bruguiere, 1792)

Retusa umbilicata (Montagu, 1803)

Ordo THECOSOMATA de Blainville, 1824

Familia CAVOLINIIDAE Gray, 1850

Genus Cavolinia Abildgaard, 1791

Cavolinia inflexa (Lesueur, 1813)

Familia CLIIDAE Jeffreys, 1869

Genus Clio Linnaeus, 1767

Clio pyramidata Linnaeus, 1767

Familia CRESEIDAE Rampal, 1973

Genus Creseis Rang, 1828

Creseis clava (Rang, 1828)

Genus Styliola Gray, 1 847

Styliola subula (Quoy et Gaimard, 1827)

Ordo UMBRACULIDA Dali, 1899

Familia TYLODINIDAE Gray, 1847

Genus Tylodina Rafmesque, 1814

Tylodina perversa (Gmelin, 1791)

Ordo PULMONATA Cuvier, 1817

Familia SIPHONARIIDAE Gray, 1827

Genus Williamia Monterosato, 1844

Williamia gussonii (O.G. Costa, 1829)

Familia TRIMU SCULID AE J.Q. Burch, 1945

Genus Trimusculus EC. Schmidt, 1818

Trimusculus mammilaris (Linnaeus, 1758)

Classis BIVALVIA Linnaeus, 1758

Ordo SOLEMYOIDA Dali, 1889

Familia NUCULIDAE Gray, 1824

Genus Austronucula Powell, 1939

Austronucula perminima (Monterosato, 1875)

Ordo ARCOID A Stoliczka, 1871

Familia ARCIDAE Lamarck, 1809

Genus Area Linnaeus, 1758

Area noae Linnaeus, 1758

Area tertragona Poli, 1795

Genus Asperarca Sacco, 1898

Asperarca nodulosa (O.F. Muller, 1776)

Asperarca secreta La Perna, 1998

The benthic molluscan thanatocoenosis ofZannone Island (Pontine Archipelago, Latium, Italy)

105

Genus Barbatia Gray, 1 842

Barbatia barbata (Linnaeus, 1758)

Familia NOETIIDAE Steward, 1930

Genus Striarca Conrad, 1862

Striarca lactea (Linnaeus, 1758)

Ordo MYTILOIDA Ferussac, 1822

Familia MYTILIDAE Rafmesque, 1815

Genus Mytilus Linnaeus, 1758

Mytilus galloprovincialis Lamarck, 1819

Genus Crenella T.Brown, 1827

Crenella arenaria Monterosato, 1875 ex H.

Martin, ms.

Genus Gregariella Monterosato, 1884

Gregariella semigranata (Reeve, 1858)

Genus Musculus Roding, 1798

Musculus costulatus (Risso, 1 826)

Musculus discors (Linnaeus, 1767)

Genus Lithophaga Roding, 1798

Lithophaga lithophaga (Linnaeus, 1758)

Genus Modiolus Lamarck, 1799

Modiolus barbatus (Linnaeus, 1758)

Genus Modiohda Sacco, 1897

Modiolula phaseolina (Philippi, 1844)

Genus Rhomboidella Monterosato, 1884

Romboidella prideauxi (Leach, 1815)

Ordo PECTINOIDA Gray, 1854

Familia PECTIN1DAE Rafmesque, 1815

Genus Talochlamys Iredale, 1935

Talochlamys multistriata (Poli, 1795)

Genus Flexopecten Sacco, 1 897

Flexopecten flexuosus (Poli, 1795)

Flexopecten hyalinus (Poli, 1795)

Familia PROPEAMUSSIIDAE Abbot, 1954

Genus Cyclopecten A.E.Verrill, 1897

Cyclopecten brundisiensis Smriglio et Mariot-

tini, 1990

Familia SPONDYLIDAE Gray, 1826

Genus Spondylus Linnaeus, 1758

Spondylus gaederopus Linnaeus, 1758

Ordo LIMOIDA Moore, 1952

Familia LIMIDAE Rafmesque, 1815

Genus Lima Bruguiere, 1797

Lima lima (Linnaeus, 1758)

Genus Limaria Link, 1807

Limaria hians (Gmelin, 1791)

Limaria tuberculata (Olivi, 1792)

Genus Limatula S.V. Wood, 1839

Limatula subauriculata Habe, 1958

Limatula subovata (Monterosato, 1875)

Ordo LUCINOIDA Gray, 1854

Familia LUCINIDAE Fleming, 1828

Genus Ctena Morch, 1860

Ctena decussata (O.G. Costa, 1829)

Genus Lucinella Monterosato, 1884

Lucinella divaricata (Linnaeus, 1758)

Ordo VENEROIDA Gray, 1854

Familia CHAMIDAE Lamarck, 1809

Genus Chama Linnaeus, 1758

Chama gtyphoides Linnaeus, 1758

Genus Pseudochama Odhner, 1917

P seudo chama griphina (Lamarck, 1819)

Familia LAS AEIDAE Gray, 1842

Genus Lasaea Brown, 1 827

Lasaea adansoni (Gmelin, 1791)

Familia NEOLEPTONIDAE Thiele, 1934

106

Bruno Fumanti

Genus Neolepton Monterosato, 1875

Neolepton sulcatulum (Jeffreys, 1859)

Familia CARDIIDAE Lamarck, 1809

Genus Parvicardium Monterosato, 1884

Parvicardium exiguum (Gmelin, 1791)

Parvicardium scriptum (Bucquoy, Dautzenberg

et Dollfus, 1892)

Genus Papillicardium Sacco, 1899

Papillicardium papillosum (Poli, 1791)

Familia TELLIN1DAE Blainville, 1814

Genus Moerella (Linnaeus, 1758)

Moerella donacina (Linnaeus, 1758)

Moerella pygmaea (Loven, 1 846)

Familia TRAPEZIDAE Lamy, 1920

Genus Coralliophaga Blainville, 1824

Coralliophaga lithophagella (Lamarck, 1819)

F amilia VENERID AE Rafmesque, 1815

Genus Venus Linnaeus, 1758

Venus casina Linnaeus, 1758

Venus verrucosa Linnaeus, 1758

Genus Timoclea T. Brown, 1 827

Timoclea ovata (Pennant, 1777)

Genus Gouldia C.B. Adams, 1847

Gouldia minima (Montagu, 1803)

Genus Irus F.C. Schmidt, 1818

Irus irus (Linnaeus, 1758)

Ordo CARDITOIDA Dali, 1889

Familia CARDITIDAE Ferussac, 1822

Genus Cardita Bruguiere, 1792

Cardita calyculata (Linnaeus, 1758)

Genus Gians Megerle von Miihlfeldt, 1811

Gians trapezia (Linnaeus, 1767)

Genus Centrocardita Sacco, 1899

Centrocardita aculeata (Poli, 1795)

Familia AS TARTIDAE d’Orbigny, 1844

Genus Astarte J.C. Sowerby, 1816

Astarte fusca (Poli, 1791)

Ordo MYOIDA Stoliczlca, 1870

Familia MYIDAE Lamarck, 1809

Genus Sphenia Turton, 1 822

Sphenia binghami Turton, 1822

Familia HIATELLIDAE Gray, 1824

Genus Hiatella Bose, 1801

Hiatella arctica (Linnaeus, 1767)

Hiatella rugosa (Linnaeus, 1767)

Ordo ANOMALODESMATA Dali, 1889

Familia THRACIIDAE Stoliczka, 1870

Genus Thracia Blainville, 1 824

Thracia distorta (Montagu, 1803)

Thracia villosiuscula (Mac Gillivray, 1 827)

Familia LYONSIIDAE P. Fischer, 1887

Genus Lyonsia Turton, 1 822

Lyonsia norwegica (Gmelin, 1791)

Classis SCAPHOPODABronn, 1862

Ordo DENTALIIDAE da Costa, 1776

Familia GADILINIDAE Chistikov, 1975

Genus Episiphon Pilsbry et Sharp, 1897

Episiphon fdum (G.B. Sowerby II, 1860)

REFERENCES

WoRMS Editorial Board, 2014. World Register of

Marine Species. Available from http://www.ma-

rinespecies.org at VLIZ. Accessed 2014-03-12.

Biodiversity Journal, 2014, 5 (2): 107-116

Monograph

The long journey of Fusinus rostratus (Olivi, 1 792) (Gastropoda

Fasciolariidae) from Portugal coasts to Venice Lagoon

Paolo Russo

Santa Croce 421, 30135 Venezia, Italy; e-mail: russorusso@v irg ilio .it

ABSTRACT In the present paper the following morpha of FusiflUS TOStVCltUS (Olivi. 1 792) (Gastropoda

Fasciolariidae) were investigated: Atlantic, Central and Southern Tyrrhenian Sea, Egadi Islands

and the Sicilian Channel, Coasts ofNorth Africa, the Central Adriatic Sea, Upper Adriatic Sea

and the Venice Lagoon. Each of these morpha shows such morphological characteristics to be

easily separated from the others. It is interesting to observe that the morphotype from the coast

of Portugal is by far morphologically the closest to that fro m Northern Adriatic. A feature com-

mon to all the described morphotypes, is the presence of secondary cords, regularly spaced

between the primary ones. The aim of this study is to split this species by geographical areas

in order to facilitate further studies.

KEY WORDS Fusinus rostratus ; Fasciolariidae; Mediterranean Sea; morphotipi.

Received 28.01.2014; accepted 2 1.03.20 14; printed 30.06.2014

Proceedings of the Seventh M alac o lo g ic al Pontine Meeting, September 9 th -10 th , 2013 - San Felice Circeo, Italy

INTRODUCTION

The Fusinus rostratus (Olivi, 1 792) (Gas-

tropoda Fasciolariidae) is a species distributed all

over the Mediterranean Sea.

It is more common in the Northern and Central

Adriatic and in Tyrrhenian Sea. It is also reported

for the Atlantic Coasts, Portugal (Hidalgo, 1917;

Barash & Danin, 1992), the Can ary Islands (Aradas

& Benoit, 1 870 Poppe & Goto, 1991; Barasch &

Danin, 1992). There are also records from Morocco

(P asteu r-H u m b ert, 1962; Barasch & Danin, 1 988,

1992; Ardovini & Cossignani, 2004) and M aurita-

nia (Lozet & D e j e an - A rre c g ro s , 1 977) but these

latters should be investigated; rare in the Aegean

Sea with records that require a careful study (per-

sonal observation).According to Mallard & Robin

(2005), F. WStratUS is endemic to the Mediter-

ranean Sea.

F. rostratus is an eurybates species, found in a

few centimeters of water in the Lagoon of Venice

(Buzzurro & Russo, 2001; Russo, 2012) to a max-

imum of detected depth of 8 2 3 m (D'Amico,

1912).

This species is related to soft sediments (Vio &

De Min, 1994, 1996), muddy (Monterosato, 1877),

debris and muddy (Coen & Vatova, 1 932), debris

and muddy-sandy (Vatova, 1943), muddy-sandy

(Vatova, 1 940; Taviani, 1 978).

F. rostratus also occurs in Peyssonnelia poly-

morpha facies and maerl(Jacquotte, 1962;Ledoyer,

1 969). It feeds on polychetes.

In the present paper the following morpha of F.

WStratUS were investigated: Atlantic, Central and

Southern Tyrrhenian Sea, Egadi Islands and the

Strait of Sicily, Coasts of North Africa, Central

Adriatic Sea, Northern Adriatic Sea and the Venice

Lagoon.

108

Paolo Russo

MATERIAL AND METHODS

Due to the considerable amount of available

material, it was possible to select a typical range

for all considered morpha. For the most part these

are from residues of fishing. The following loca-

tions were selected: Atlantic: coasts of Portugal,

Algarve from nets at 60 m; Central Thyrennian:

Tuscan Archipelago from fishing vessels at 100/300

m; Southern Thyrennian, Pozzuoli from fishing ves-

sels at 60 m; Strait of Sicily: Egadi Islands by fish-

in g vessels at unknown depths; Coasts of North

Africa: Algeria from creels at 60 m; Central Adri-

atic, Pescara from fishing vessels at 40-60 m; North-

ern Adriatic, Chioggia from nets for AequipeCten

opercularis (Linnaeus, 1 75 8 ) and Pecten jacobaeus

(Linnaeus, 1758) at 25-30 m; Venice Lagoon, Pellest-

rina Island, harvested by hand during low tide.

SYSTEMATICS

L am ilia LASCIOLARIID AE J.E. Gray, 1853

Subfamilia LUSININAE Wrigley, 1927

Genus Fusinus R afinesque, 18 15

Fusinus rostratus (Oiivi, 1792)

O rig in al description (O livi, 1 792): “M Strombo

di prima spezie di colore biondetto formato ad an-

goli, e tutto ricoperto di finissimi cordoncini, che

gli girano pel traverso. Gin. Adr. t. ii. Pag. 8 tav. 7

fig. 5 6.” (Fig.l).

A very brief description not easy to interpret. A

more accurate description can be found in D 'Ancona

(18 7 1): “ Conchiglia fusiforme allungata, acuminata

all’apice e terminata alia base da un canale dritto,

stretto, di poco piu corto della spira. Questa consta

di circa 9 giri convessi, carenati ad eccezione dei

primi tre o quattro, divisi da una sutura molto pro-

fonda, i quali portano otto o move coste longitudinal i

piuttosto grosse, rotondate, sporgenti, ristrette al

loro principiare verso la sutura superiore e piu larghe

al loro terminare verso quella inferiore. Tutti gli

anfratti sono divisi quasi nel mezzo in due porzioni

pressoche uguali (la superiore sovente maggiore) da

una carena rilevata, talora lamellosa e sfrangiata,

producendo in tal caso in corrispondenza delle coste

longitudinale delle punte molto ottuse e molto com-

press e dal basso alValto, come apparisce dalla fig.

9 (a, b) della Tav. 14. Numerosi sono i solchi ed i

cordoncini traversi, ravvicinati fra loro, rilevati,

rugosi e leggennente ondulati, i quali gradatamente

diminuendo di numero e di grossezza giungono fino

alia estremita del canale. Tali cordoncini sogliono

essere un poco meno grossi nella porzione superiore

dei giri, e nella inferiore si osserva ordinariamente

che nel solco che divide due di loro vi ha un sottile

filetto. Tutta la superficie della conchiglia e resa

scabra da numerosissime linee di accrescimento sot-

tilissime che rendono quasi granulosi i cordoncini

trasversali. L’apertura e piuttosto piccolo, ovale; il

labbro alquanto spesso, e acuto nel margine ed

internamente solcato; la lamina columellare perfet-

tamente liscia nella maggior parte dei casi si rialza

sul penultimo anfratto prolungandosi in questo

modo anche lungo il canale. 11 quale e mediocre-

mente lungo, stretto, dritto ed a per tor

In F. WStratUS the protoconch, always paucispi-

ral, cannot be considered a diagnostic element

(Buzzurro & Russo, 2007) as highly variable de-

pending on the population. For further clarity here

are illustrated the protoconchs of all the considered

morpha with the exception of specimens from

Egadi Islands due to lack of intact specime ns (Figs.

2-8 and Table 1). The presence of secondary cords,

regularly spaced between the primary ones, is an

element of diagnostic character (Fig. 9). Normally

the shells are 50-60 mm up to 87 mm high

(Donnarumma, 1 968), reaching 95 mm (Kaicher,

1 978, unverified). It counts 76 synonymies.

Algeria

9 1 4

364

Chioggia

907

392

Civitavecchia

892

478

Pescara

928

385

Portugal

735

32 1

Pozzuoli

7 1 4

52 1

Strait of Messina

642

228

Venice Lagoon

664

385

Table 1. Sizes of protoconchs and nuclei (expressed in

pm) of the described mo rp ho types.

The long journey of Fusinus rostratus (Olivi, 1 792) (Gastropoda Fasciolariidae) from Portugal coasts to Venice Lagoon 1 09

rostratus mfo,

M. Strombo di prima spezie di colore biondetto formato

ad angoli, e tutto ricoperto di finissimi cordoncini ,

che gli girano pel traverse . Gin* Adr* T * II* pag*

8, tav, 7. fig* $6*

Abka diversi fondi, e predilige git arenacei : Frequent* .

Si trova angora la Strombo di seconda i ’pegic rigato , e papi*

pjiato , di vostro ettrvo , t di colore che incline al carneo dei-

Jo stesso Ginanm tav, 7. fig. 570 ed urfaltra variety piii

ventricosa a coda replicata , e corta.

Figure 1. Originaldescription of FusiflUS VOStVOtUS Olivi, 1792. Figures 2-8. Protoconchs. Fig. 2: morphotype from Portugal.

Fig. 3: morphotype from Southern Tyrrhenian. Fig. 4: morphotype from Central Tyrrhenian. Fig. 5: morphotype from

North Africa. Fig. 6: morphotype from Central Adriatic. Fig. 7: morphotype from Northern Adriatic. Fig. 8: morphotype

from Venice Lagoon. Figure 9. Secondary cords in F. VOStvatUS.

Paolo Russo

Following is a summary of the main populations

of the Mediterranean Sea.

COAST OF PORTUGAL (Figs. 10-12)

Medium sized

Shell rather thick and solid

Siphonal canal of medium length and slightly

deviated

Teleoconch consisting of 7-7.5 whorls

Light brown or yellowish in color

Axial ribs not very prominent

Usually acarinate, sometimes the supramediane

cord of the body whorl is slightly raised

Protoconch diameter 735 pm, nucleus 321pm

CENTRAL TYRRHENIAN (Figs. 13-14)

M edium sized

S hell rather light

Siphonal canal long and straight

Teleoconch consisting of 7-7.5 whorls

Milk white, sometimes with pale yellow shades

Axialribs not very prominent, sometimes barely

hinted

Spiral cords rather thin and raised

A lw ay s ac arin ate

Protoconch diameter 892 pm, nucleus 478 pm

SOUTHERN TYRRHENIAN (Figs. 15-17)

Small and medium sized

S hell rather light

Siphonal canal long and straight

Teleoconch consisting of 7-7.5 whorls

Reddish brown

Axial ribs not very prominent

Spiral cords rather thin and raised

A lw ay s ac arin ate

Protoconch diameter 7 14 pm, nucleus 521 pm

EGADI AND STRAIT OF SICILY (Figs. 18-19)

M edium sized

S hell rather thick

Siphonal canal long and straight

Teleoconch consisting of 6-7 whorls

Whitish with pale yellow shades

Axial ribs little prominent, barely visible in the

adapical area of the body whorl

Spiral cords irregular and of discontinuous

thickness

Keel very raised

It was not possible to detect the protoconch for

lack of intact specimens

NORTH COAST OF AFRICA (Figs. 20-22)

M edium sized

Shell rather thick and solid

Siphonal canal of medium length and slightly

deviated to the left

Teleoconch consisting of 7 whorls

Pale yellow to light brown

Axial ribs normally rised

Spiral cords particularly evident and spaced

Generally has a rather evident keel

Presence of a rather evident columellar callus

Protoconch diameter 914 pm, nucleus 364 pm

C EN T R A L A D R IAT IC (Figs. 23-25)

Medium to large sized for the species

Shell thick and solid

Siphonal canal of medium length and often

tw isted

Teleoconch consisting of 7-7.5 whorls

W hitish in colour

Axial ribs not very raised

Spirals cords thin and of medium height

Whorls particularly inflated

Seldom a slight keel is present

Aperture particularly wide

Pro to conch diameter 925 pm, nucleus 385 pm

NORTHERN ADRIATIC (Figs. 26-28)

M edium sized

S hell rather thick

Siphonal canal of medium length and slightly

deviated

Teleoconch consisting of 7-7.5 whorls

Deep reddish brown to pale yellow and straw-

coloured

Axial ribs not very raised, sometimes lacking on

the last w horl

Spirals cords thin and of medium height

Often acarinate, sometimes the supramediane

cord of the last whorl slightly raised

Pro to conch diameter 907 pm, nucleus 392 pm

The long journey of Fusinus rostratus (Olivi, 1 792) (Gastropoda Fasciolariidae) from Portugal coasts to Venice Lagoon 1 1 1

Figures 10-12. Morphotype from Coast ofPortugal. Fig. 10. h: 39.3 mm, D: 14.5 mm. Fig. 11. h: 41.4 mm, D: 17.6 mm.

Fig. 12. h: 35.3 mm, D: 12.2 mm. Figures 13, 14. Morphotype from Central Ty rrh en ian . Fig. 13. h: 61.6 mm, D: 21.0 mm.

Fig. 14. h: 47.5 mm, D: 20.4 mm. Figures 15-17. Morphotype from Southern Ty rrh en ian .Fig. 15.h:50.5 m m , D : 16.3 mm.

Fig. 16. h: 36.4 mm, D: 12.7 mm. Fig. 17. h: 37.7 mm, D: 13.3 mm.

112

Paolo Russo

Figures 18, 19. Morphotype from Egadi and S trait of Sicily. Fig. 18. h: 46.0 mm, D: 24.7 mm. Fig. 19. h: 44.0 mm, D: 27.6

mm. Fig u res 20-22. Morphotype from No rth coast ofAfrica. Fig. 20. h: 40.5 mm, D: 14.6 mm. Fig. 21. h: 34.0 mm, D:

14.2 mm. Fig. 22. h: 40.6 mm, D: 15.4 mm. Figures 23-25. Morphotype from CentralAdriatic. Fig. 23. h: 57.6 mm, D:

22.0 mm; Fig. 24. h: 59.0 mm, D: 22.2 mm; Fig. 25. h: 55.8 mm, D: 21.0 mm.

The long journey of Fusinus rostratus (Olivi, 1 792) (Gastropoda Fasciolariidae) from Portugal coasts to Venice Lagoon 1 1 3

Figures 26-28. Morphotype from Northern Adriatic. Fig. 26. h: 40.0 mm, D: 15.2 mm; Fig. 27. h: 41.5 mm, D: 15.7 mm.

Fig. 28. h: 61.0 mm, D: 21.4 mm. Fig u res 29-31. Morphotype from Venice Lagoon. Fig. 29. h: 27.4 mm , D: 12.0 mm; Fig.

30, h: 28.2 mm, D: 11.4 mm; Fig. 31. h: 25.2 mm, D: 11.0 mm. Figures 32-3 3. Comparison among the morphotype from

Portugal Coasts (Fig. 32) and that from N orthern A d riatic (Fig . 3 3).

114

Paolo Russo

VENICE LAGOON (Figs. 29-31)

Small sized

Shell rather thick and almost always eroded

Siphonal canal short

Teleoconch consisting of 5-6 whorls

From dark brown to almost black in colour

Axial ribs not very raised and often eroded

Spirals cords thin and of little raised

A lw ay s ac arin ate

In some areas of the Venice Lagoon, during the

low tide, it lives in absence of water

Protoconch diameter 664 pm, nucleus 3 85 pm

RESULTS AND DISCUSSION

Each of these morpha shows such morphologi-

cal characteristics to be easily separated from the

others, therefore, despite being F. WStratUS a poly-

morphic species, it is stable within the analyzed

m orpha.

It is interesting to observe that the morphotype

from the coast of Portugal is by farmorphologically

the closest to that from the Northern Adriatic (Figs.

32, 33). This may not be surprising when one con-

siders that the Northern Adriatic lagoon environ-

ments show, for concomitant geographical, climatic

and environmental factors, sub-Atlantic rather than

Mediterranean characteristics (Sacchi, 1 977, 1983;

Bianchi, 1983;Mizzan, 1999). Among the No rt hern

Adriatic malacofauna we can include at least two

other "cold" guests as LittOrinQ SQ.XQ.tUis (Olivi,

1 7 9 2) and CallioStoma vires cens (Coen, 1933).

A feature common to all the described morpho-

types, is the presence of secondary cords, regularly

spaced between the prim ary ones. It is believed that

this element is a diagnostic character (Merle, 2001,

2005; Crocetta et al., 2012; Russo, 2013) ( Fig. 9).

It can therefore be said that the alternation of

(primary cords, secondary cords) is a valuable char-

acter for the determ ination of F. WStVQtUS, or rather,

the presence of this sequence, excludes other species

with the exception of F. buZZUrroi Prkic et Russo,

Mnrphottpo

pa r:i metro

somm a

media h/d

Porto gait o

44.4

17.7

41.0

16.0

42,5

17,0

29.6

12.0

31.0

11.7

30.3

11.8

29,4

11,6

39,4

15.3

h/d

2.51

2,56

2.50

2.47

2.65

2.57

2.53

2.58

20.36

2.55

Medio Ttrreno

61.4

2!. 2

60.0

19,4

60,0

19,6

57.0

20.0

57.0

20.0

53.0

20.0

53,0

19,0

57,0

17.2

h/d

2.90

3.09

3.06

2.85

2.65

2.79

3.31

23.50

2.94

Basso Tirreno

53.0

27.6

50.3

16.6

36,4

12.0

37.6

13.3

36.3

12.4

41.3

13.8

33.4

12,8

28.0

10.7

h/d

1.92

3.03

3,03

2.83

2.93

2.99

2.61

2,62

21.96

2,74

Nord A frica

40.6 1

15.8

40.3

14.0

34.0

14.0

32.0

12.8

34.0

12.5

40.0

14,0

39.0

15.0

34.5

13.0

h/d

2.57

2.88

2,43

2.50

2.72

2.86

2.60

2,65

21.2)

2.65

Medio Adriadco

57.8

23.0

57.S

21.6

54,0

20,0

53.2

19.0

42.7

18.3

55.0

21.0

44.5

18,0

44,7

19.0

h/d

2.51

2.68

2.70

2.80

2.33

2.62

2.47

2.35

20.47

2.56

Alto Adriadco

43.6

18,4

46,2

16,8

40,0

15.0

46.0

17.8

37.4

13.0

450

18,0

48,0

18,0

49,0

19.0

h/d

2.37

2.75

2.67

2.58

2.88

2.50

2.67

2.58

20.99

2.62

Laguna di Venezia

25.0

10.6

27.4

12.0

26.0

1 1.2

282

1 1.9

25.0

10.8

21.5

10,0

31.0

12.3

26.8

12.0

h/d

2.36

2.28

2.32

2.37

2.31

2.15

2.52

2,23

18.55

2,32

Table 2. h/D ratio of the described morphotypes.

The long journey of Fusinus rostratus (Olivi, 1 792) (Gastropoda Fasciolariidae) from Portugal coasts to Venice Lagoon 1 1 5

2008 but this latter is easily distinguishable. These

observations we re made on a large amount of spec-

imens from the different localities.

To confute this thesis the observations were

extended also to a number of tropical species of the

genus FusiviUS , not useful to list here, confirming

that the presence of secondary cords is not occa-

sional and cannot be attributed to the single spec-

imen: some species possess them and others do not.

From the observation of several juveniles it can

be seen that for a height of 8 mm there are not yet

secondary cords for 3 whorls; for a height of 14 mm

and 4 whorls they appear on the body whorl; in spec-

imens with 5-5,5 whorls they appear also in the

penultimate whorl and in the larger ones with 7

whorls the secondary cords are present from the

third last w horl.

This preliminary study has not dealt with the

problem of a possible specific division of the dif-

ferent morph a of Fusinus YOStYCltUS. The current

state of the art considers them all belonging to the

sam e species.

The aim of this study is to split this species by

geographical areas in order to facilitate further

s tu d ie s .

ACKNOWLEDGEMENTS

The Author wishes to thank Dr. D. Scarponi and

Dr. G. Gasparotto, Department of Earth Sciences,

University of Bologna (Italy) for the SEM photo-

graphs implement and the friends Loris Perini and

M irco Vianello (Chioggia, Italy), Roberto Costan-

tini (Silvi Marina, Italy), Fabio Crocetta (Naples,

Italy), Arman do Verdasca, Carlos Afonso and Nel-

son Tiago (Portugal) for providing study material.

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Biodiversity Journal, 2014, 5 (2): 117-130

Monograph

Phenotypic diversity of Thuridilla hope/' (Verany, 1 853) (Gas-

tropoda Heterobranchia Sacoglossa). A DNA-barcoding ap-

proach

Giulia Furfaro 1 , Maria Vittoria Modica 2 , Marco Oliverio 2 ,Juan Lucas Cervera 3 & Paolo Mariottini 1

'Dipartimento di Scienze, Universita degli Studi di “Roma Tre”, Viale Marconi 446, 00146 Rome, Italy;

e-mail: giuliafurfaro@hotmail.it; paolo.mariottini@uniroma3.it

2 Dipartimento di Biologia e Biotecnologie “Charles Darwin”, Universita di Roma “La Sapienza”, Viale dell’Universita 32, 00185

Rome, Italy; e-mail: marco.oliverio@uniromal.it; mariavittoria.modica@uniromal.it

Departamento de Biologia, Facultad de Ciencias del Mar y Ambientales, Campus de Excelencia Internacional del Mar (CEUMAR,)

Universidad de Cadiz. Poligono Rio San Pedro, s/n, Ap.40. 11510 Puerto Real (Cadiz), Spain; e-mail: lucas.cervera@uca.es

■"Corresponding author

ABSTRACT The sacoglossan Thuridilla hopei (Verany, 1853) shows highly diverse chromatic patterns.

Based on the morphological examination of specimens from different Mediterranean locali-

ties, we have observed that in spite of this great variability in colours of T. hopei , two major

chromatic morphotypes are related to bathymetry. Specimens from deeper water exhibit blue

darker and more uniform patterns than individuals from shallower water, which show a more

variable, dashed and spotted arrangement of light blue, yellow, orange, white and black

pigmentation. A molecular genetic analysis using the mitochondrial COI and 16S rDNA

markers has confirmed that all these extremely different chromatic morphotypes belong to a

single specific entity, i.e. T. hopei, a sacoglossan with a wide distribution, from Macaronesia

in the Atlantic, to the easternmost Mediterranean Sea.

KEY WORDS Sacoglossa; Thuridilla hopei', colour morphotypes; Mediterranean Sea; Atlantic Ocean.

Received 28.01.2014; accepted 21.03.2014; printed 30.06.2014

Proceedings of the Seventh Malacological Pontine Meeting, September 9 th - 10 th , 2013 - San Felice Circeo, Italy

INTRODUCTION

The plakobranchid sacoglossan genus Thuri-

dilla Bergh, 1872 is represented in the northeastern

Atlantic and in the Mediterranean Sea by two

species, T. hopei (Verany, 1853) (Carmona et al.,

2011; Malaquias et al., 2012) and the amphiatlantic

T. mazda Ortea et Spinosa, 2000, recently recorded

from the Azores (Malaquias et al., 2012). The brightly

coloured T. hopei lives from the lower intertidal

down to about 35 m depth, and generally is found

crawling on hard substrate. It feeds suctorially

on photophilous algae, in particular on Derbesia

tenuis sima (de Notaris) Crouan et Crouanand

Cladophora vagabunda (Linnaeus) van den Hoek

(Marin & Ros, 2004; Handeler & Wagele, 2007;

Handeler, 2011) and it retains functional algal chloro-

plasts from its food for few days (Marin & Ros,

1989, 2004). The aposematic chromatic pattern of

this slug is related to the presence in its tissues of

toxic compounds like the diterpenoids thuridillin A,

B and C and nor- thuridillin (Gavagnin et al., 1993;

De Rinaldis, 2012).

During the last decade, the high levels of in-

traspecific chromatic variation of T. hopei has been

well documented in literature (Trainito, 2005;

Handeler, 2011; Carmona et al., 2011 and refer-

ences therein), as well as depicted in the Sea Slug

118

Giulia Furfaro et alii

Forum (Australian Museum, Sydney, Available

from http://www.seaslugforum.net/). It is worth of

mentioning that the Macaronesian specimens of T.

hopei , have long been ascribed to the Caribbean T.

picta (Verril, 1901), due to a similar colour pattern

(see Cervera et al., 2004; Malaquias et al., 2009),

until Carmona et al. (2011) demonstrated by genetic

data that they belong to T. hopei.

We have observed and documented the chro-

matic variability of T. hopei at different Mediter-

ranean localities, particularly along the Italian coast.

Interestingly, we observed that the distribution of

different chromatic phenotypes is related to ba-

thymetry, as previously reported by Cattaneo-Vietti

(1990) and Handeler (201 1). Specimens from deeper

water exhibit dark blue and more uniform patterns

(“bluish” form according to Handeler, 2011: 37)

than individuals from shallower water, which show

a more dashed and spotted arrangement of light

blue, yellow, orange, white and black pigmentation

(“rosy” form according to Handeler, 2011: 37). Dif-

ferent colour types have been usually considered as

chromatic moiphotypes of T. hopei , despite the con-

sistent chromatic differences linked to their ba-

thymetry (Carmona et al., 2011).

Therefore, we wanted to either confirm or deny

the conspecificity of “bluish” and “rosy” forms by

genetic data, being the significance of chromatic

differences difficult to evaluate based on morpho-

logical data alone. In this study we applied a molecu-

lar analysis, using two mitochondrial markers, the

barcode COI and the 16S rDNA, to the different

colour moiphotypes of T. hopei , to test the hypothe-

sis that “bluish” and “rosy” specimens of these

snails belong to the same species and confirm the

lusitanic distribution of this sacoglossan sea slug.

MATERIAL AND METHODS

Collection data, vouchers and accession num-

bers are listed in Table 1 . Italian specimens from

several localities were collected by hand during

SCUBA diving at different depth. Each specimen

was photographed in situ and/or in aquarium, then

fixed in ethanol.

A piece of tissue was dissected from the foot for

DNA extraction, and the remaining animal was de-

posited at the Department of Biology and Biotech-

nologies “Charles Darwin” (La Sapienza Rome

University). DNA was extracted using a standard

proteinase K phenol/chloroform method with

ethanol precipitation, as reported in Oliverio &

Mariottini (2001). A fragment of the mitochondrial

1 6S rDNA was amplified by PCR using the univer-

sal primers 16Sar-L and 16Sbr-H (Palumbi et al.,

2001), while a fragment of the mitochondrial cy-

tochrome oxidase I (COI) was amplified using the

universal primers LCO1490 and HC02198 (Folmer

et al., 1994); for PCR conditions see Prkic et al.

(2014). All amplicons were sequenced by Genechron

Centre of Sequencing, ENEA (La Casaccia, Rome,

Italy) or by European Division of Macrogen Inc.

(Amsterdam, The Netherlands), using the same

PCR primers.

Forward and reverse sequences were assembled

and edited, and the resulting consensus sequences

of each specimen were readily aligned by hand.

BLAST search was always conducted for each se-

quence. Published sequences (COI and 16S) of

Thuridilla were downloaded from theGenBank.

Although the definition of a phylogenetic hypothe-

sis for the genus Thuridilla was not within the aims

of this paper, nevertheless, phylogenetic relation-

ships among the Thuridilla sequences were inferred

to have a phylogenetic framework for the estima-

tion of the genetic distances: we used neighbour-

joining (NJ) and maximum likelihood (ML) (both

bootstrapped over 1000 replicates), by the software

MEGA 5.0 (Tamura et al., 2011) and Bayesian

Inference (BI) by the software MrBayes (with 5*10 6

generations, and 25% bumin) by MrBayes 3.2.2

(Ronquist et al., 2012). Sequences of the Atlantic

Elysia timida Risso, 1818, retrieved from the

GenBank, were used as outgroup. Nodes in the phy-

logenetic trees were considered ‘highly’ supported

with Bayesian posterior values >96% and bootstrap

values >80%; nodes with Bayesian posterior values

of 90-95% and bootstrap values of 70-79% were

considered ‘moderately’ supported (lower support

values were considered not significant).

Genetic divergence among the barcode COI

sequences was observed (p distance) and estimated

using the Kimura-2 -parameters nucleotide substi-

tution model (K2p distance).

RESULTS AND DISCUSSION

Animals were observed and sampled at seven

localities of the Tyrrhenian Sea (Table 1).

Phenotypic diversity of Thuridilla hopei (Gastropoda Heterobranchia Sacoglossa). A DNA-barcoding approach 119

Species

Voucher ID

Locality

COI

16S

References

Elysia timida

MNCN

15.05/53680

Menorca,

Spain (MED)

HQ6 16847

Carmona et al.,

2011

Elysia timida

MNCN

15.05/53680

Menorca, Spain (MED)

HQ616818

Carmona et al.,

2011

Thuridilla

albopustulosa

ZSM:20033615

NW-Sulawesi,

Indonesia

EU140889

Handeler &

Wagele, 2007

Thuridilla bayeri

DQ471279

DQ480208

Bass, 2006

Thuridilla bayeri

DQ480206

Bass, 2006

Thuridilla bayeri

DQ480207

Bass, 2006

Thuridilla bayeri

DQ480207

Bass, 2006

Thuridilla bayeri

ZSM:20033612

NW-Sulawesi,

Indonesia

EU140886

Handeler &

Wagele, 2007

Thuridilla carlsoni

Lizard Island,

Australia

HM 187640

Wagele et al.,

2010

Thuridilla carlsoni

Lizard Island,

Australia

EU 140877

Handeler &

Wagele, 2007

Thuridilla carlsoni

Lizard Island,

Australia

EU140878

Handeler &

Wagele, 2007

Thuridilla carlsoni

DQ480214

Bass, 2006

Thuridilla gracilis

Lizard Island,

Australia

HM 187641

Wagele et al.,

2010

Thuridilla gracilis

Hamahiga, Okinawa,

Japan

AB758972

Takano et al.,

2013

Thuridilla gracilis

Lizard Island,

Australia

EU140884

Handeler &

Wagele, 2007

Thuridilla gracilis

Lizard Island,

Australia

EU140885

Handeler &

Wagele, 2007

Thuridilla gracilis

Lizard Island,

Australia

EU140883

Handeler &

Wagele, 2007

Thuridilla gracilis

Hamahiga,

Okinawa, Japan

AB759041

Takano et al.,

2013

Thuridilla hoffae

ZSM:20060224

Samoa

EU140880

Handeler &

Wagele, 2007

Thuridilla hoffae

DQ480213

Bass, 2006

Thuridilla hopei

Elba Is., Italy (MED)

EU 140881

Handeler &

Wagele, 2007

Thuridilla hopei

CASIZ 184307

Erance (MED)

HQ6 16854

HQ616825

Carmona et al.,

2011

Table 1. Voucher ID, collection localities and sequence accession numbers of Thuridilla and Elysia specimens.

MED: Mediterranean; EA: Eastern Atlantic; WA: Western Atlantic (continued).

120

Giulia Furfaro et alii

Species

Voucher ID

Locality

COI

16S

References

Thuridilla hopei

MNCN

15.05/53682

Menorca, Spain (MED)

HQ6 16849

HQ6 16820

Carmona et al.,

2011

Thuridilla hopei

Mataro, Baretta del Abre,

Barcelona, Spain (MED)

EU140882

Handeler &

Wagele, 2007

Thuridilla hopei

MNCN/ADN

17015

Western Andalucia, Spain

(EA)

HQ616855

HQ6 16826

Carmona et al.,

2011

Thuridilla hopei

ZMBN 81680

Azores, Portugal (EA)

HQ616850

HQ6 16821

Carmona et al.,

2011

Thuridilla hopei

ZMBN 81680

Azores, Portugal (EA)

HQ658123

Carmona et al.,

2011

Thuridilla hopei

MNCN

15.05/53685

Madeira, Portugal (EA)

HQ616853

HQ6 16824

Carmona et al.,

2011

Thuridilla hopei

Blanes, Cala St. Francesc,

Spain (MED)

GQ996678

Handeler et al.,

2009

Thuridilla hopei

Giglio Is., Italy (MED)

KC573743

Krug et al., 2013

Thuridilla hopei

Mataro, Baretta del Abre,

Barcellona, Spain (MED)

GQ996677

Handeler et al.,

2009

Thuridilla hopei

AF249810

Wollscheid-Len-

geling et al., 2001

Thuridilla hopei

BAU 1651.1

Giglio Is., Italy,

42°22’27”N 10°52’47”E,

30 m depth (MED)

KJ397547

KJ363910

Present study

Thuridilla hopei

BAU1651.2

Giglio Is., Italy,

42°22’27”N 10°52 , 47”E,

30 m depth (MED)

KJ397548

KJ363911

Present study

Thuridilla hopei

BAU 1652

Le Formiche rocks, Italy,

42°34’28”N 10°52’58”E,

30 m depth (MED)

KJ397549

KJ363912

Present study

Thuridilla hopei

BAU 1653.1

Sant’Agostino, Italy,

42°08’45”N 11°43’48”E,

25 m depth (MED)

KJ397550

KJ363913

Present study

Thuridilla hopei

BAU1 653.2

Sant’Agostino, Italy,

42°08’45” N 11°43’48”E,

25 m depth (MED)

KJ397551

KJ363914

Present study

Thuridilla hopei

BAU 1654

Giannutri Is., Italy,

42°15’07”N 11°07’04”E,

20 m depth (MED)

KJ363915

Present study

Thuridilla hopei

BAU 1655.1

Ponza Is., Italy,

40°52’52”N 12°58’02”E,

25 m depth (MED)

KJ363916

Present study

Thuridilla hopei

BAU1 655.2

Ponza Is., Italy,

40°52’52”N 12°58’02”E,

25 m depth (MED)

KJ397552

Present study

Thuridilla hopei

BAU 165 6

Cape Circeo, Italy,

41°13’31”N 13°03’02”E,

14 m depth (MED)

KJ363917

Present study

Thuridilla hopei

BAU1 657.1

San Vito Lo Capo, Sicily,

38°10’02”N 12°46’11”E,

1 0 m depth (MED)

KJ397553

KJ363918

Present study

Table 1. Voucher ID, collection localities and sequence accession numbers of Thuridilla and Elysia specimens.

MED: Mediterranean; EA: Eastern Atlantic; WA: Western Atlantic (continued).

Phenotypic diversity of Thuridilla hopei (Gastropoda Heterobranchia Sacoglossa). A DNA-barcoding approach 121

Species

Voucher ID

Locality

COI

16S

References

Thuridilla hopei

BAU1 657.1

San Vito Lo Capo, Sicily,

38°10’02”N 12°46’11”E,

10m depth (MED)

KJ397553

KJ363918

Present study

Thuridilla hopei

BAU1 657.2

San Vito Lo Capo, Sicily,

38°10 , 02”N12°46 , 11”E,

1 0 m depth (MED)

KJ397554

KJ363919

Present study

Thuridilla kathae

Lizard Island, Australia

EU 140879

Handeler &

Wagele, 2007

Thuridilla

lineolata

Sulawesi, Indonesia

EU140887

Handeler &

Wagele, 2007

Thuridilla livida

Merizo Clay's backyard,

Guam

HM 187636

Wagele et al.,

2010

Thuridilla livida

Bile Bay, Guam

HM 187607

Wagele et al.,

2010

Thuridilla livida

DQ480211

Bass, 2006

Thuridilla mazda

UNAM 3027

Mexico

HQ616836

Carmona et al.,

2011

Thuridilla neona

Lord Howe Is., Australia

KC573747

Krug et al., 2013

Thuridilla neona

DQ480209

Bass, 2006

Thuridilla picta

ZMBN 83023

Bermuda (WA)

HQ6 16851

HQ6 16822

Carmona et al.,

2011

Thuridilla picta

ZMBN 83023

Bermuda (WA)

HQ658125

Carmona et al.,

2011

Thuridilla picta

MNCN

15.05/53683

Colombia (WA)

HQ616861

HQ6 16832

Carmona et al.,

2011

Thuridilla picta

MNCN

15.05/54991

Colombia (WA)

HQ6 16862

HQ6 16833

Carmona et al.,

2011

Thuridilla picta

MNCN/ADN

17016

Cuba (WA)

HQ6 16852

HQ6 16823

Carmona et al.,

2011

Thuridilla ratna

AF249256

Wollscheid-Lenge-

ling et al., 2001

Thuridilla

splendens

Kouri, Okinawa, Japan

AB758973

Takano et al.,

2013

Thuridilla

splendens

Kouri, Okinawa, Japan

AB759042

Takano et al.,

2013

Thuridilla undula

DQ480210

Bass, 2006

Thuridilla vatae

Vaisala lagoon, Savaii

Island, Samoa

HM187637

Wagele et al.,

2010

Thuridilla vatae

ZSM:20060088

Samoa

EU140888

Handeler &

Wagele, 2007

Thuridilla vatae

DQ480212

Bass, 2006

Table 1. Voucher ID, collection localities and sequence accession numbers of Thuridilla and Elysia specimens.

MED: Mediterranean; EA: Eastern Atlantic; WA: Western Atlantic.

122

Giulia Furfaro et alii

Figure 1. Thuridilla hopei “bluish” morphotype. Giannutri Is., Tuscany Archipelago, 42°15’07”N 11°07’04”E, Italy, 20 m

depth. Figure 2. T. hopei “bluish” morphotype. St. Agostino, Latium coast, 42°08’45”N 11°43’48”E, Italy, 25 m depth.

Figure 3. T. hopei “bluish” morphotype. Ponza is., Latium coast, 40°52’52”N 12°58’02”E, Italy, 25 m depth.

Phenotypic diversity of Thuridilla hopei (Gastropoda Heterobranchia Sacoglossa). A DNA-barcoding approach 123

Figure 4. Thuridilla hopei “rosy” moiphotype. Zannone Is., Latium coast, 40°57’ 1 9”N 1 3°03 ’ 1 9”E, Italy, 2 m depth. Figure

5. T. hopei “rosy” morphotype. San Vito Lo Capo, Sicily coast, 38°10’02”N 12°46’11”E, Italy, 10 m depth. Figure 6.

T. hopei intermediate morphotype. St. Agostino, Latium coast, 42°08’45”N 11°43’48”E, Italy, 25 m depth.

124

Giulia Furfaro et alii

According to the body colour, almost all speci-

mens could be split into two main colour forms, the

“bluish” (Figs. 1-3) and “rosy” (Figs. 4, 5) morpho-

types. According to our data and other sources (see

Table 2), slugs could be referred to: 1) the “bluish”

form, showing a dark blue colour on the upper part

of the rhinophores and living in deeper water

(usually deeper than -25 m); 2) the “rosy” form,

showing a wider white coloration of the rhinophores,

including the upper portion, is a shallower water

inhabitant (max depth recorded 15m). Interestingly,

the few specimens showing a colour pattern that was

not easy to assign unambiguously (an example is

depicted in Fig. 6), were collected at intermediate

depths, from -20 to -25 m (Table 2; Fig. 7).

The phylogenetic analysis based on the partial

sequences of COI and 16S mithocondrial genes,

yielded similar trees (Figs. 8, 9). Since this work

was not aimed to the definition of a molecular phy-

logenetic hypothesis for the entire genus Thuridilla,

we will not discuss the phylogenetic details of the

trees. We just notice that according to the relation-

ships among the sequences of specimens of the

complex of T. bayeri (Marcus, 1965), including also

the nominal taxa T. gracilis (Risbec, 1928), T. ratna

(Marcus, 1965), T. splendens (Baba, 1949), this

group is worthy of a genetic approach to clarify the

status of the various taxa. The sequences ascribed

to T. hopei and T. picta (Verrill, 1901) were more

closely related to T. neona Gosliner, 1995 than to

any other species of the genus (support range: 95-

98%), similarly to what found by Gosliner et al.

(1995) based on an anatomical dataset. The dif-

ference in our trees was that T. hopei and T. picta

formed two reciprocally monophyletic clades more

closely related each other (98-99%) than to T.

neona. In all phylogenetic analyses (Figs. 8, 9) the

specimens of T. picta from the Caribbean formed a

highly supported clade (98-99%). All specimens

from the eastern Atlantic also formed a highly sup-

ported clade (98-100%), corresponding to T. hopei

as conceived in Carmona et al. (2011).

Figure 7. Depth ranges in meters of the ‘rosy’, ‘bluish’ and intermediate morphotypes, summarized from the data of Table 2.

Phenotypic diversity of Thuridilla hopei (Gastropoda Heterobranchia Sacoglossa). A DNA-barcoding approach 125

T. hopei

morphotypes

N°

Specimens

Mediterranean records

Depth

(meter)

Bluish

Le Fonniche rocks, Italy, 42°34 , 28”N I0°52’58”E

Giglio Is., Italy, 42 0 22’27”N IG°52’47”E

Giannutri Is., Italy. 42 o l5’07’*N ll°07WE

St. Agostino, Italy, 42°08 , 45 ,, N ll°43 , 48' , E

25-30

MPA “Secche di Tor Patemo", Italy, 41°36W'N 12°19'00”E

25-35

Ponza Is., Italy, 40°52'52"N 12 o 58’02”E

25-30

Baiun Is., Croatia (http://www,seasIugforum.net/message/1934)

Area Marina Protetta “Secche di Tor Patemo”, Italy, 41°36 , 00 1, N

12°19’00”E (Alberto Altomare, personal communication)

20-25

Intermediate

Giannutri Is., Italy, 42°15’07”N ll o 07’04”E

St. Agostino, Italy, 42°08 1 45”N 1 1°43’48”E

Cap d' Antibes, France (http://www.seaslugforum.net/message/2l023)

Dive site “POceir, Cerbere, France

(http://www.seaslugforum.net/message/755 1 )

“Secchitelle” Torvaianica, Italy, 4l°34’I5”N 12°19W’E

10-14

Rosy

Torre Astura, Italy, 41 0 24’31”N ]2°45’54 ,4 E

0.5

off Cape Circeo, Italy, 4l°13’3r’N 13°03’02”E

Zannone Is., Italy, 40°57 , 19 ,1 N 13 o 03't9”E

Santa Caterina, Italy, 40 Q 08’23”N 17°59M2”E

San Vito Lo Capo, Sicily, 38°10'02"N l] e 46’H”E

“La digue”, Port-Leucate, France

{http://www.seaslugforum.net/message/ 1 4769)

Villafranche-sur-mer, France,

( http :// w w w. seas 1 ugforum . net/message/22 52 0 )

“Pierre qui tramole”. Cap Croisette, France,

{ http ://w w w. seas 1 ugforum , n et/message/73 04)

Kounoupeli beach, lleia, Greece

{ http : // w w w. se as l ug forum . n et/message/ 1 5 3 07 )

0.3

Bodrum, Turkey (http://w\vw.seaslugforum. net/message/ 1 7707)

Michmort beach, Israel (http://www.seaslugforum.net/message/20048)

Split. Croatia (Jakov Prkie, personal communication)

0-1

Murter Island - loc. Kosirina, Croatia ( Alen Petani, personal

communication)

0-1

Turanj, Croatia (Alen Petani, personal communication)

0-1

>300

Biograd - loc. Bosana, Croatia (Alen Petani, personal communication)

0-2

Zadar - loc. Karma, Croatia (Alen Petani, personal communication)

0.5-2

> 100

Zaton - loc, Bilotinjak, Croatia (Alen Petani. personal communication)

0-2

Vrsi, Croatia (Alen Petani, personal communication)

0-1

Nin - loc. Sabunike, Croatia (Alen Petani, personal communication)

0.5-1

Vir Island - loc. Pedinka, Croatia (Alen Petani, personal communication)

1-2

Dive site "Yellow' Wall", Capoliveri, Elba Is., Italy

( http://www .seaslugforum.net/message/ 1934)

Capo Figari, Sardinia, (Egidio Trainito, personal communication)

Divesite "Blue Grotto", Malta

(http://www.seaslugforum.net/message/1934)

“Seech iielle”, Torvajanica, Italy (Alberto Altomare, personal

communication)

10-1 5

Brindisi, Italy (Vincenzo Marra, personal communication)

5-15

Table 2. Mediterranean Records of Thuridilla hopei assigned to bluish-rosy or intermediate morphotypes, with their

locality and bathymetry. Original records by the authors, personal communications or retrieved from the Internet.

126

Giulia Furfaro et alii

Distribution of K2p distances

80-

eo-

0.00

0 02 0.04 0 06 0 0B 0 10

98/99/99

96/97/98

98/99/98

002

91/90/91

90/9 f/93

T hopei GQ 99667 8

-7! tope/ BAU 1657.1

X hopei BAU 1657.2

7: hopei HQ6 16849

1 00/99/tbj T - hopei KC573743

' X hopei BAU 1651 ,1

-7: hopei GQ 99 66 77

X hopei HQ616855

T hopei BAU 1652

C - X hopei AF249810

T hopei HQ61 6854

T. hope/ BAU1 653.1

X tope/ BAU1651 .2

X tope/ BAU 1653.2

— X tope/ HQ6 16853

X tope/ HQ61 6850

X hopei HQ 65 81 23

L X hopei BAU1 655.2

98/99/98

96/96/96

|— X picta HQ616861

X picta HG6 16862

- X picta HQ6581 25

[rX picta HG6 16851

T- X p/cfa HQ6 16852

X neona KC573747

X livida HM1 87636

X cadsoni HM 187640

X vatae HM1 87637

99/98/97

X splendens AB7 58973

— - X gracilis HM 187641

X gracilis AB758972

E. timida HG6 16847

Figure 8. Maximum Likelihood tree based on the COI dataset (K2p model of evolution). Numbers at nodes are bootstrap on

NJ and ML analyses (1000 replicates) and Bayesian posterior support (5* 10 6 generations and 25% bumin). Only moderately

or highly supported nodes are annotated. Scale is calibrated ML distance. The histogram shows the distribution of the pairwise

estimated genetic distances (K2p) in intraspecific (left, light grey) and interspecific (right, dark grey) comparisons.

Phenotypic diversity of Thuridilla hopei (Gastropoda Heterobranchia Sacoglossa). A DNA-barcoding approach 127

96/97/98

5(6/96/95

98/99/99

7 hopei BAU 1657.1

7 hopei BAU 1657.2

T. hopei BAU 1652

— 7 hopei BAU 165 1.1

1 1 hope i BAU 1654

^7 hopei BAU 1655.1

- 7 hopei HQ 616826

90/95/93

/jope/EU14Q882

7 hopei HQ6 16820

7 hopei EU 140881

7 hopei BAU 1656

L | 7 hopei HQ6 16825

p7 hopei BAU 165 1,2

L T. hopei BAU 1653.1

7 hopei BAU 1653.2

7 hopei HQ6 16824

7 hopei HQ6 16821

r 7I picta HQ616833

7 p/c/a HQ6 16832

7 p/cfa HQ6 16822

7 p/cfa HQ6 16823

4_ 7

^ 7 neona DQ480209

7 //Wda IHM 187607

100/99/99^^ //Wafa DQ480211

— 7 undula DQ480210

7 lineolata EU 140887

98/99/100

91/90/91

7 gracilis EU 140884

98/99/98 j-7 gracilis EU 140835

7 ratna AF249256

7 gracilis EU 140883

-7 bayeri DQ480207

-7 bayeri DQ480206

98/97/96j 7 gracilis AB759041

- 7 bayeri DQ480208

7 splendens A B 759 04 2

7 bayeri EU1 40886

99/100/100

99/98/97

7 vatae EU 140888

7 vatae DQ480212

■7 albopustulosa EU1 40889

[

■7 tnazda HQ6 16836

99/100/100

7 hoffae EU 140880

L 7 hoffae DQ480213

7 kathae EU 140879

7 carlsoni EU1 40878

100 / 100/100

- 7 carlsoni EU1 40877

'—T. carlsoni DQ48 02 14

— E. timida HQ616818

Figure 9. Maximum Likelihood tree based on the 16S dataset (K2p model of evolution). Numbers at nodes are bootstrap on

NJ and ML analyses (1000 replicates) and Bayesian posterior support (5* 10 6 generations and 25% bumin). Only moderately

or highly supported nodes are annotated. Scale is calibrated ML distance.

128

Giulia Furfaro et alii

Figure 10. Thuridilla hopei “bluish” morphotype and Felimare tricolor. St. Agostino, Latium coast, 42°08’45”N

11°43’48”E, Italy, 35 m depth. Figure 11. T. hopei “bluish” morphotype and Felimare fontandraui. Giglio Is., Tuscany

Archipelago, 42°22’27”N 10°52’47”E, Italy, 30 m depth.

Phenotypic diversity of Thuridilla hopei (Gastropoda Heterobranchia Sacoglossa). A DNA-barcoding approach 129

The interspecific K2p genetic distances esti-

mated on the COI sequence alignment (Fig. 8; dis-

tance matrices available on request from the

authors), excluding the comparisons with the out-

gtroup, ranged from 5.1% to 20.8% (mean 11.6%).

With the exception of the two specimens ascribed

to T. gracilis (from a complex in need of revision,

and with a distance of 6.8%), the K2p distances

(COI) ranged from 0.5% to 0.9% (mean 0.7%) in

the T. picta clade, and from 0% to 2.2% (mean

0.8%) in the T. hopei clade.

The largest distance values in the T. hopei clade

were observed between the Macaronesian and the

other eastern Atlantic and Mediterranean specimens

(0.8%-2.2%). This is perfectly fitting a pattern with

only two species involved in this clade, T. picta in

the western and T. hopei in the eastern Atlantic,

respectively, as previously proposed by Carmona et

al. (2011).

Furthermore, our results do not support any

taxonomic split of the “bluish” and “rosy” chro-

matic forms, which fall within the genetic variation

of T. hopei. Interestingly, the “bluish” morphotype

was often recorded at several Tyrrhenian localities

(Figs. 10,11), in the same spots where also species

of the family Chromodorididae of the blue chro-

matic group, such as Felimare tricolor (Cantraine,

1835) and F. fontandraui (Pruvot-Fol, 1951) were

observed. Mullerian mimicry complexes have been

described, involving similarly coloured toxic sea

slugs, but sometimes including also polyclads

(Gosliner, 2001: 166, fig. 1 ). In T. hopei the intraspe-

cific variability of colour patterns, with somehow

discrete morphotypes living at different depths, may

be partly driven by mimicry coevolution.

The deeper water “bluish” morphotype observed

in strict syntopy with the “blue” Felimare spp.,

might be a new Mullerian mimicry complex for the

Mediterranean, where the sympatric species may

have evolved the shared bright body colours and

patterns convergently in the deeper waters (Gosliner,

2001). Conversely, no evidence has been so far

collected for a similar mimicry complex involving

the “rosy” morphotype in shallow waters. It is

worth mentioning that the shallower specimens dis-

play a higher variability.

The Macaronesian specimens of T. hopei , could

represent a third colour morphotype, since their

colour pattern albeit similar to the “rosy” morpho-

type, yet shows a reduction of the dashed light-blue

spots at the edge of the parapodia, a larger black

portion without the basal white line, the terminal

portion of rhinophores with a coloured band con-

sisting of light blue, black and orange rings. This

“ringed-rhinophore” chromatism, which has been

so far documented in a single Mediterranean case,

from Israel (Table 2, http://www.seaslugfomm.net/mes-

sage/20048), may either represent a third discrete

colour morphotype or fall within the high variabil-

ity of the shallow water T. hopei.

Future studies will focus on the selective factors

acting at different depths, to maintain these two

colour forms in the Mediterranean Sea.

ACKNOWLEDGEMENTS

The authors gratefully acknowledge Egidio

Trainito (Olbia, Italy) and Jakov Prkic (Split,

Croatia) for their critical comments and helpful sug-

gestions on the present paper. We are indebted to

professional marine photographers Alessio Sera

(Rome, Italy), for the permission to use some T.

hopei photographs, and Marco Cesaroni (Rome,

Italy) for technical support. We thank people from

“Gruppo Malacologico Mediterraneo” (Rome,

Italy) for their assistance during samplings. PM

wishes to thank the University of Roma Tre for fi-

nancial funding.

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Biodiversity Journal, 2014, 5 (2): 131-140

Monograph

The molluscs found after the nourishment of the littoral of

Terracina (Latium, Italy)

Luigi Giannelli

Via A. Martucci 3, 04019 Terracina, Latina, Italy; e-mail: lgiannelli72@ gmail.com

ABSTRACT In the present paper molluscs found after the beach nourishment carried out in 2006 on the

coast of Terracina are reported. Altogether were identified 144 taxa, of which 105 Gastropoda,

37 Bivalvia and 2 Scaphopoda.

KEY WORDS Mollusca; beach nourishment; Terracina; Italy; Mediterranean Sea.

Received 28.01.2014; accepted 21.03.2 0 14; printed 30.06.2 0 14

Proceedings of the Seventh Malacological Pontine Meeting, October 5 th - 6 th , 2013 - San Felice Circeo, Italy

INTRODUCTION

In the years 1 950-60 the uncontrolled anthropic

alteration carried out on the sandy coasts had as a

consequence the modification of currents and tides,

thus increasing the beach erosion. The first methods

of consolidation of coasts were carried out in the

absence of standards and draft rules, thus causing

many environmental and aesthetic problems inter-

fering with the dynamics of the coastline. This sit-

uation has made it necessary to study and fund

several methods based on scientific and technical

assessments. The purpose of beach nourishment is

to rebuild the eroded beach through the use of suit-

able sand directly taken from the sea bottom and

afterwards released on the eroded beach (Gar bin et

al., 20 12).

At the beginning for the beach nourishment of

the littoral of Terracina was utilized a sandy lime-

stone from inland quarries, absolutely unsuitable.

This led to continuous washing out and narrowings

of the beach that caused with time the silting up of

the seabed. Such action, later revealed be ineffec-

tive and disastrous, led to the decision to utilize a

sand with morphological and granulometric charac-

teristics as similar as possible to those of the eroded

beach (Garbin et al., 2012).

In 2006 it was decided to carry out a kind of

"soft" beach nourishment taking directly the sand

from the so called "underwater quarry", located on

relatively deep seabed off the coast depositing it

directly on the eroded beaches (Figs. 1, 2). The

most promising underwater cave was discovered

about fifty miles north-west from Terracina, specif-

ically off of Torvajanica (Rome) named “Cava

sot to marina Torvaianica Sud Zona C 2”. This is just

one of the many quarries of the continental marine

shelf of Latium, used in the nourishment of other

coasts such as Anzio, Ostia, Montalto di Castro,

etc.. This site mainly differs from others in that the

storage m aterial w as aspirated and draw n at a greater

depth, around 110 meters depth.

This operation was divided into two different

stages, the first was effected in 2006 for about two

kilometers on the first half of the Ponente Beach

and the second, to be performed the following year,

on the other half up to the port. A fter the first stage

on the shoreline, with the disappointment of the

local community, the beach appeared covered with

a large amount of pebbles and shells, thus putting

132

Luigi Giannelli

at risk the long tourist season. As a consequence the

Genus Glbbula R is so , 1826

second stage was canceled (Giannelli et al., 2012).

Gibbula magus (Linnaeus, 1758)

A total of 1300 meters in length were interested

in the beach, were obtained 66,000 square feet of

Familia C A L L IO S T O M AT ID A E Thiele, 1924

new surface after the intervention with 460,000

(1847)

cubic meters of sand poured. From the malacologi-

cal point of view this large amount of m ate rial, both

Genus CalliostOma S w ainson, 1840

fossil and subfossil, is very interesting.

Calliostoma conulum (Linnaeus, 1758)

From the several collections made just after the

Calliostoma granulatum (Born, 1 1 1 8 )

first nourishment and during the immediately fol-

Calliostoma laugieri (Payraudeau, 1826)

lowing months, altogether 144 taxa have been iden-

Calliostoma zizyphinum (Linnaeus, 1758)

tified, of which 105 Gastropoda(73%),37 Bivalvia

(26%) and 2 Scaphopoda (1%).

Familia TURBINIDAE Rafinesque, 1815

MATERIAL AND METHODS

Genus Bolmd Risso, 1826

Bolma rugosa ( Linnaeus, 1767)

All species were collected directly along the

shore line. Several species (for example Neptunea

COfltrurid (L in n ae u s , 1771) are clearly fossils but

Genus Homalopoma Carpenter, 1864

Homalopoma sanguineum (Linnaeus, 1 7 5 8 )

for many others it was impossible to detect if it were

Familia P H A S I A N E L L ID A E Swainson, 1840

the state fossil or subfossil. The nomenclature of the

species has been updated according to WoRMS

Genus Tricolia R is so , 1826

Editorial Board (2014).

Tricolia pullus (Linnaeus, 1758)

RESULTS

Ordo CAENOGASTROPODA Cox, 1960

Taxonomic list

Familia CERITHIIDAE Fleming, 1822

Classis GASTROPODA Cuvier, 1797

Genus Ccrithium B ru g u iere , 1789

Cerithium alucaster (Brocchi, 1 8 1 4 )

Ordo V ETIG A STR O PO D A S alvini-Plaw en , 1980

Cerithium protractum (Bivona Ant. in Bivona

And., 1 83 8 )

Cerithium vulgatum (B ru g u iere , 1792 )

Familia F IS S U R E L L ID A E Fleming, 1822

Genus Diodora J.E.Gray, 1821

Diodora gibberula (L am ark, 1 8 2 2 )

Diodora graeca { Linnaeus, 1 758) (Fig. 3)

Diodora c ft demartinio rum b uzzurro et Russo

Familia S IL IQ U A R 1ID A E Anton, 1838

Genus TenagoduS G u ettard , 1770

Tenagodus obtUSUS (Schumacher, 1817)

2005 (Fig. 4)

Familia T U R R IT E L L ID A E Loven, 1847

Genus Einarginuld L am arck , 1801

Emarginula f is sura (Linnaeus, 1758)

Genus Turritella Lam arck, 1799

Turritella turbona (M onterosato, 1 877)

Familia TROCHIDAE Rafinesque, 1815

Familia TRIPHORIDAE Gray, 1847

Genus Clelandella W inckw orth, 1932

Clelandella miliaris (B roc chi, 1 8 1 4)

Genus MomphorUS Grillo, 1877

Monophorus perversus (Linnaeus, 1758)

The molluscs found after the nourishment of the littoral ofTerracina (Latium, Italy)

133

Fam ilia EPITO N IID A E B erry, 1910 (1812)

Genus Epitonium Rod in g , 1798

Epitonium clathrus (Linnaeus, 1758)

Epitonium turtonis (T u rto n , 1 8 1 9 )

Fam ilia EULIMIDAE Philippi, 1853

Genus EulimCl R i s s o , 1826

Eulima glabra (da Costa, 1 77 8 )

Familia RISSOIDAE Gray, 1847

G e n u s Alvania R is s o , 18 26

Alvania punctur a { Montagu, 1 8 o 3 )

Familia VERMET1DAE Rafinesque, 1815

Genus Thylacodes Guettard, 1770

Thylacodes arenarius (Linnaeus, 1 7 5 8)

Familia APORRHAIDAE Gray, 1850

Genus Aporrhais da C osta, 1 7 7 8

Aporrhais pespelecani (Linnaeus, 1758)

Familia C A LY P T R A E ID A E Lamarck, 1809

Genus Calyptraea L am arck, 1799

Calyptraea chinensis (Linnaeus, 1 758)

Figure 1. Study area: littoral ofTerracina, Latium, Italy (right); underwater quarry (left) . Figure 2. Littoral ofTerracina

(L atium , Italy).

134

Luigi Giannelli

Familia CAPULIDAE Fleming, 1822

Genus CcipuluS M ontfort, 18 10

Capulus ungaricus (Linnaeus, 1758)

Familia TRIVIIDAE Troschel, 1863

Genus EratO Risso, 1826

Erato voluta (Montagu, 1 803)

Genus Trivia Gray, 1837

Trivia multilirata (G.b. Sower by n, 1 8 7 o )

(Figs. 5, 6)

Familia CYPRAEIDAE Rafinesque, 1815

Genus Luria Jousseaume, 1884

Luria lurida ( Linnaeus, 1 75 8)

Genus Naria B ro d erip ,18 3 7

Naria spurca (Linnaeus, 1 758)

Genus Schilderia Tomlin, 1930

Schilderia achatidea ( Gray in G.B. Sowerby I,

1 837) (Figs. 7, 8)

Genus Zonaria Jousseaume, 18 84

Zonaria pyrum (Gmeiin, 1 7 9 1 ) (F ig s . 9 , 1 0 )

Familia OVULIDAE Fleming, 1822

Genus Pseudosimnia s child er, 1927

Pseudosimnia adriatica (g.b. Sowerby 1 , 1 8 2 8 )

(Figs. 11, 12)

Pseudosimnia camea (Poiret,i789) (Figs. 13 , 1 4 )

Genus Silfinia R is so , 18 26

Simnia spelta (Linnaeus, 1758) (Figs. 15, 16)

Familia NATICIDAE Guilding, 1834

Genus Euspira Agassiz in J. Sowerby, 1837

Euspira flisca (B lainville, 1 825)

Euspira guilleminii (Payraudeau, 1826)

Euspira intricata (Donovan, 1 8 0 4 )

Euspira macilenta (Philip pi, 1 8 4 4 )

Genus NaticariuS D urn eril, 1805

Naticarius hebraeus (Martyn, 1786)

Naticarius stercus muscarum (Gmeiin, 1 7 9 1 )

Genus Tectonatica Sacco, 1890

Tectonatica rizzae (Philippi, 1 8 44)

Familia CASSIDAE Latreille, 1825

Genus Galeodea Link, 1 807

Galeodea echinophora (L innaeus, 1758)

Genus Semicassis Morch, 1 8 5 2

Semicassis granulata undulata (G m eiin , 1 7 9 1 )

Familia RANELLIDAE Gray, 1854

Genus Cabestana Rod in g, 1798

Cabestana CUtaeea ( Linnaeus, 1 767) (Fig. 17)

Genus Charonia g isti, 1 84 7

Charonia lampas (L innaeus, 1758)

Genus Monoplex Perry, 1 8 1 0

Monoplex corrugatum (Lamarck, 1816)

Monoplex parthenopeum (Von Saiis, 1 7 9 3)

Genus Ranella l arnarck, 1816

Ranella olearium (Linnaeus, 175 8 )

Familia BURS1DAE Thiele, 1925

Genus Bursa Ro ding, 17 98

Bursa scrobilator (Linnaeus, 1 758) (Figs. 18, 19)

Familia MURICIDAE Rafinesque, 1815

Genus Babelomurex Coen, 1922

Babelomurex benoiti (Tiberi, 1 8 5 5 )

Genus Bolinus Pusch, 1837

Bolinus brandaris (Linnaeus, 1758)

Genus DermomureX M onterosato, 1890

Dermomurex scalaroides (B lainville, 1 829)

(Fig. 20)

Genus Hadriania Bucquoy et Dautzemberg, 1882

Hadriania craticula (Bucquoy, Dautzemberg et

Dollfus, 1 8 82) (Fig. 21)

Genus Hexaplex Perry, 18 10

Hexaplex trunculus (L innaeus, 1758)

The molluscs found after the nourishment of the littoral ofTerracina (Latium, Italy)

135

Fig. 3 . Diodora graeca, h : 2 5.6 mm. Fig. 4 . Diodora dr. demartiniorum, h: 25.3 mm. Figs. 5,6. Trivia multilirata, h : 1 2 .2

m m . Figs. 7 , 8 . Schilderia achatidea, h : 3 8 m m . Figs. 9, 10. Zonaria pvrum, h : 34.8 mm. Figs. 11. 12. Pseudosimnia adria-

tica, h : 2 3 . 1 mm. Figs. 13, 14. i 3 . carnea, h: 16.2 mm. Figs. 15, 16. Simnia spelta , h : 13.6 mm. Fig. 17. Cabestana cutacea,

h: 44.2 mm. Fig. is. 19. Bursa scrobilator, h: 49.2 mm. Fig. 20. Dermomurex scalaroides, h: 17.4 mm. Fig. 2i. Hadriania

craticula h : 3 1 .7 m m . F ig . 2 2 . HirtomureX squamosus, h : 2 6.5 m m . F ig . 2 3 . Murexul aradasii, h : 12 m m . F ig . 2 4 . Muricopsis

cristata, h : 2 1 m m . Fig. 25. Ocinebrina helleri, h : 16.3 mm.

136

Luigi Giannelli

Genus HirtOmureX Coen, 1922

Hirtomurex squamosus (Bivona Ant. in Bivona

And., 1 8 38 ) (Fig. 22)

Genus Murexul Iredale, 19 15

Murexul aradasii (M onterosato in Poirieri,

1 8 8 3 ) (Fig. 23)

Genus Muvicopsis Bucquoy et Dautzemberg, 1882

Muricopsis cristata (B ro c ch i, 1 8 1 4) (F ig . 2 4)

Genus Ocenebra g ray, 1 8 47

Ocenebra erinaceus (Linnaeus, i 75 8 )

Genus Ocinebrina j ousseaume, 1880

Ocinebrina edwardsi (Payraudeau, 1 8 2 6 )

Ocinebrina helleri (B r u s in a , 1 8 6 5) (F ig . 2 5 )

Genus Pagodula M onterosato, 1884

Pagodula echinata (Kiener, 1 8 40) (Fig. 26, 2 7)

Genus TwpHonopsis Bucquoy, Dautzemberg et

D o llfu s s , 18 8 2

Trophonopsis muricata (Montagu, 1 8 o 3 )

Genus TyphinelluS Jousseaume, 1880

Typhinellus labiatus (de Cristofori et Jan, 1 83 2)

(Fig. 28, 29)

Familia M A R G IN E L L ID A E Fleming, 1828

Genus Volvarina Hinds, 1844

Volvarina mitre lla (Risso, 1826)

Familia MITRIDAE Swainson, 1829

Genus Mitra L am arc k , 179 8

Mitra cornicula (Linnaeus, 1758)

Mitra zonata (M an-yat, 1 8 1 8)

Familia COSTELLAR1IDAE Mac Donald, 1860

Genus Vexillum Rod in g , 1798

Vexillum ebenus (Lamarck, 18 11)

Vexillum granum (Forbes, 1844)

Vexillum tricolor (G m elin , 1 7 9 1 )

Familia BUCCINIDAE Rafinesque, 1815

Genus Euthria G ray, 185 0

Euthria cornea (L innaeus, 1 75 8)

Genus Neptunea Rod in g, 1798

Neptunea contraria (Linnaeus, 1 7 7 1 )

Familia NASSARIIDAE Iredale, 1 9 1 6 (1 835)

G enus NaSSaUUS D um eril, 1805

Nassarius corniculum (O liv i, 1792)

Nassarius incrassatus (S tro m , 1 7 6 8 )

Nassarius nitidus (Jeffreys, 1 8 67)

Nassarius pygmaeus (Lamarck, 1 8 2 2 )

Familia COLUMBELLIDAE Swainson, 1840

Genus Mitrella R is so, 1826

Mitrella coccinea (Philippi, 1 8 36)

Mitrella minor (Scacchi, 1836)

Mitrella scripta (Linnaeus, 1758)

Familia FASCIOLAR1ID AE Gray, 1853

Genus EusinUS Rafinesque, 1815

Eusinus rostratus { oiivi, 1792) (Fig. 30)

Familia C L AT H U R E L L ID A E H. Adams et A.

Adams, 1858

Genus ComarmOlldia M onterosato, 1884

Comarmondia gracilis (Montagu, 1803)

Familia M ITROM ORPHID AE Casey, 1904

Genus Mitromorpha Carpenter, 1865

Mitromorpha karpathoensis (Nordsiek, 1969)

Mitromorpha mediterranea (M ifsud, 200 1 )

Familia M ANGEL1ID AE P. Fisher, 1883

Genus Bela Gray, 1 84 7

Bela nebula (M ontagu, 1 803)

Genus Mangelia R is so , 1826

Mangelia cost at a (Pennant, 1777)

Mangelia costulata (Risso, 1 8 2 6 )

Mangelia sp .

Familia DRILL IID A E Olsson, 1964

Genus CraSSOpleura M onterosato, 1884

Crassopleura maravignae (B ivona Ant. in

BivonaAnd., 1 8 3 8 )

The molluscs found after the nourishment of the littoral ofTerracina (Latium, Italy)

137

Familia CLAVATULIDAE Gray, 1853

Genus Fusiturris Thiele, 1929

Fusiturris similis (Bivona Ant. in Bivona And.,

1 8 3 8 )

Familia RAPHITOMIDAE Bellardi, 1875

Genus RaphitOlTlCl B e Hard i, 1847

Raphitoma cfr. atropurpurea (Fig. 3D

Raphitoma cfr. echinata (Fig. 3 2)

Raphitoma leufroyi (M ichaud, 1 8 2 8) (Fig. 33)

Raphitoma s p . l (F ig . 3 4 )

Raphitoma sp. 2

Ordo HETEROSTROPHA P. Fischer, 1885

Familia A R C H IT E C T O N 1C ID A E J.E. Gray in

M .E. Gray, 1850

Genus DisCOtectonica Marwick, 1931

Discotectonica discus (Philippi, 1 844) (Figs.

35, 36)

Genus Heliacus d ’O 1-big ny, 18 42

Heliacus fallaciosus {Tihexi, 1 8 7 2 ) (Fig. 3 7)

Genus Pseudotorinia Sacco, 1892

Pseudotorinia architae (O.g. Costa, 1 8 4 1 )

(Figs. 38-40)

Familia M AT HIE DID A E Dali, 18 89

Genus Mathilda s ernper, 1865

Mathilda quadricarinata (B ro c ch i, 1 8 1 4 )

Familia PYRAM IDELLID AE Gray, 1840

Genus Euparthenia Thiele, 1 9 3 1

Euparthenia bulinea (Lowe, 1 8 4 1 ) (F ig s . 4 1 , 4 2 )

Familia ACTEONIDAE d’Orbigny, 1843

G e n u s ActeOU M o n tfo rt, 18 10

Acteon tornatilis (Linnaeus, 1758)

Familia RINGICULIDAE Philippi, 1853

Genus Ringicula Deshayes,1838

Ringicula auriculata (M en aid de la Groye,

18 11 )

Ordo CEPH ALASPIDEA P. Fischer, 1883

Familia CYLICHNIDAE FI. Adams et A. Adams,

1854

Genus Cylichna L o v e n , 1846

Cylichna cylindracea (Pennant, 1777)

Familia S C A P H A N D R ID A E G.O. Sars, 1878

Genus Scaphander m ontfort, 1 8 1 o

Scaphander lignarius (Linnaeus, 1758)

Ordo UMBRACULIDA Odhner, 1939

Familia UMBRACULIDAE Dali, 1 8 89 (1 827)

Genus Umbraculum S chum acher, 1817

Umbraculum umbraculum (Light foot, 1786)

C lassis B IV A LV IA

Ordo SOLEMYOIDA Dali, 1889

Familia NUCULIDAE Gray, 1824

Genus Nucula Lamarck, 1799

Nucula nucleus (Linnaeus, 1 75 8 )

Nucula sulcata (Bronn, 18 3 1)

Familia NUCULANIDAE H. Adams et A . Adams,

1858 (1854)

Genus Nuculana Link, 1 8 07

Nuculana pella (L innaeus, 1767)

Ordo ARCOIDA Stoliczka, 1871

Familia ARCIDAE Lamarck, 1809

Genus Anadara Gray, 1 847

Anadara polii (Mayer, 1 868)

Genus AfCd Linnaeus, 1758

Area tetragona ( p o n , 1795)

Familia GLY CYMERIDIDAE Dali, 1908 (1847)

138

Luigi Giannelli

Genus Glycymeris da Costa, 177 8

Glycymeris glycymeris Linnaeus (1758)

O rd o PEC TIN 0 ID A Gray, 1854

Familia PECTIN IDAE Rafinesque, 1815

Genus Aequipecten P. Fisher, 1 8 8 6

Aequipecten commutatus (Monterosato, 1 8 1 5 )

(Fig. 43)

Aequipecten opercularis (L innaeus, 1758)

(Fig. 44)

Genus Manupecten Monterosato, 1872

Manupecten pesfelis (Finnaeus, 1 7 5 8 )

Genus Mimachlamys 1 redale, 1929

Mimachlamys varia (Finnaeus, 1 7 5 8 ) (Fig. 45)

G enus Pecten O .F. M u ller, 1776

Pectenjacobeus ( Finnaeus, 1758) (Fig. 46)

Pecten maximus (Finnaeus, 1 75 8) (Fig. 47)

Genus PseudaniUSSium Morch, 1853

Pseudamussium clavatum (Poii, i795)(Fig.48)

Genus SunilipeCten W inckw orth, 1932

Similipecten similis (Faskey, 1 8 1 1 )

Genus Talochlamys Iredale, 1935

Talochlamys multistriata (P oii, 1795) (Fig. 49)

Familia SPONDYLIDAE Gray, 1826

Genus SpO ndyluS L in n aeu s, 1758

Spondylus gaederopus (Finnaeus, 1758)

Or do OSTREOIDA Ferussac, 1822

Familia OSTREIDAE Rafinesque, 1815

Genus Ostrea l innaeus, 175 8

Ostrea edulis (Linnaeus, 1758)

O rdo LU C IN O ID A Gray, 1854

Fam ilia FUCINIDAE Fleming, 1828

Genus Ludnella Monterosato, 1884

Lucinella divaricata (Finnaeus, 1758)

Genus Lucinoma d ail, 1901

Lucinoma borealis (Finnaeus, 1 767)

Genus Myrtea Turton , 1822

Myrtea spinifera (Montagu, 1 8 o 3 )

Ordo VENEROIDA Gray, 18 54

Familia CHAMIDAE Famarck, 1809

Genus Chama L in n aeu s, 1758

Chama gryphoides { Linnaeus, 1 758)

Familia CARDITIDAE Ferussac, 1822

Genus Centrocardita Sacco, 1 8 9 9

Centrocardita aculeata (P o ii, 1795)

Familia ASTARTIDAE d’Orbigny, 1 844 (1 840)

Genus Astarte J.de c . s owerby, 1816

Astarte Jus ca (Poll, 1795)

Familia CARD1IDAE Famarck, 1809

Genus Laevicardium s w ainson,l 840

Laevicardium oblongum (G m eiin, 1 7 9 1 )

Genus Papillicardium Sacco, 1899

Papillicardium papillosum (P o n, 1795 )

Genus Parvicardium m on tero s a to , 18 8 4

Parvicardium minimum (P h iiip pi, 1 8 3 6 )

Familia M ACTRIDAE Famarck, 1809

Genus Lutraria Lamarck, 1799

Lutraria lutraria (Linnaeus, 1 7 5 8 )

Lutraria oblonga (Gmeiin, 1 7 9 1 )

Familia TEFFINIDAE Blainville, 1814

Genus Tellina L in n aeu s , 1758

Tellina serrata (B ro c ch i, 1 8 1 4 )

Familia SOLECURTIDAE d’Orbigny, 1846

The molluscs found after the nourishment of the littoral ofTerracina (Latium, Italy)

139

Figs. 26, 27. Pagodula echinata, h: 1 9 .8 m m . Figs. 28, 29. TyplTmeUuS labiatlis, h : 18.1 m m . Fig. 30. Fusinus rostratus, h: 36.2

in ill . F ig . 3 1 . R. c fr. atropurpurea, h : 1 5 .4 m m . F ig . 3 2 . R. c fr. echinata , h : 2 1 .6 mm. Fig. 33 . R. leufroyi, h: 22 111 m . F ig . 3 4 . Rci-

phitoma sp., h: 22.7 m m . Figs. 35 , 36. Discotectonica discus, h: 26.6 mm . Fig. 37 . Heliacus fallaciosns, h : 16.3 mm.FigS.38-

4 o . Pseudotorinia architae, h : 8 . 7 m m . f ig s . 4 1 , 4 2 . Euparthenia bulinea, h : 1 3 .9 m m . Fig . 4 3 . Aequipecten commutatus, h : 24 .8

m m . Fig. 44. A. Opercularis, h: 3 3 .9 mm . Fig. 45. MimachlamyS Varia, h: 26.4 m m . Fig. 46. Pecten jaCobeUS, h: 6 1 m m . Fig. 47.

Pecten maximus, h: 74.9 m m . Fig. 48. Pseudamussium clavatum, h: 30.4 mm. Fig. 49 . Talochlamys multistriata, h: 24.7 m m .

140

Luigi Giannelli

Genus SoleCUVtUS B lain v ille , 1824

Solecurtus S copula (Tu r ton, 1 822 )

Genus Cuspidaria Nardo, 1840

Cuspidaria cuspidata (O liv i, 1792 )

Familia VENERIDAE Rafinesque, 1815

Genus ChcUTieleCl Morch, 1853

Chamelea striatula (da Costa, 1 7 7 8 )

Classis SCAPHOPODA

Ordo DENTALIIDA Starobogatov, 1974

Genus CldUSinella Gray, 1851

Clausinella fasciata { da Costa, 177 8)

Familia DENTAL1IDAE Children, 1834

Genus GlobiveUUS Coen, 1934

Globivenus ejfossa (Philippi, 1 8 36)

Genus Antalis H. Adams et A. Adams, 1854

Antalis dentalis (Linnaeus, 1 758)

Antalis inaequicostata (Dautzenberg, 1 8 9 1 )

Genus PitCIV Romer, 1857

Pitar rudis (P oii, 1795 )

REFERENCES

Genus TitflOClsCl B row n , 1827

Timoclea ovata { Pennant, 1 777)

Garbin F., Ginanni Corradini R. & Tramonti L., 2012.

Problem atiche costiere e ripascim enti: il caso della

Genus VeUUS Linnaeus, 1758

Venus nux ( Gmelin, 1791)

spiaggia di Terracina. Geologia dell’Ambiente,

SIGEA, 2: 11-15.

Giannelli L., Fanelli C ., Pace D.S.. Pellegrini D. & Pie-

rullo A., 2012. I R ipascim enti. A rgonauta suppl. 7:

1-209.

Ordo AN 0 M ALO DESM ATA Dali, 1889

WoRMS EditorialBoard (2014).World Register of M a-

rine S pecies. Available from http ://w w w .m arinespe-

Familia CUSPID ARIIDAE Dali, 1886

cies.org a t V L IZ . A c c e s s e d 20 14-03-1 2.

Biodiversity Journal, 2014, 5 (2): 141-146

Monograph

On the systematic position of “C/mo” melitensis Mifsud, 1998,

with erection of the new genus Mifsudia (Heterobranchia

Cimidae)

Paolo Mietto 1 , Italo Nofroni 2 & Ermanno Quaggiotto 3

1 Univcrsita degli stiidi di Padova, Dipartimento di Geoscienze, via Gradenigo 6, 35 13 1 Padova, Italy; e-mail: paolo.mietto@unipd.it

2 Via B. Croce 97, 00152 Roma, Italy; e-mail: italo.nofroni@uniromal.it

3 Via Secula 13, 36023 Longare, Vicenza, Italy; e-mail: ermanno@quaggiotto.net

^Corresponding author

ABSTRACT Based on teleoconch and, especially, protoconch features, the new genus Mifsudia is erected

for Cima melitensis Mifsud, 1998 and placed in the family Cimidae. The protoconch is hyper-

strophic, as in the other cimids. At least, two European fossil species ( Cima gantensis Bandel,

2005, from the Middle Eocene of Hungary and Murchisonella cf. obtusa Gougerot & Le

Renard, 1978 from Early Oligocene of France) are also included in the new genus. Mifsudia

melitensis (Mifsud, 1998) comb, nov., originally described from Malta, is here recorded for

the first time from Lampedusa Island, Alboran Sea and the coasts of Mauritania (West Africa).

KEY WORDS Gastropoda; Cima; Murchisonella; Mifsudia; Mediterranean Sea; Western Africa.

Received 28.01.2014; accepted 21.03.2014; printed 30.06.2014

Proceedings of the Seventh Malacological Pontine Meeting, September 9 th - 10 th , 2013 - San Felice Circeo, Italy

INTRODUCTION

During the in progress revision of the genus

Cima Chaster, 1896 (Heterobranchia Cimidae) in

the Mediterranean Sea, we examined several speci-

mens of Mediterranean and Atlantic “Czraa” and

recorded notable differences on the protoconchs.

This drove us to investigate in depth this complex

group of Heterostropha and allowed recognizing

two different typologies of protoconchs: the first one

is globular, tipical of Cima sensu strictu (e.g. Cima

minima Jeffreys, 1858, Fig. 5), the second one is

clearly hyperstrophic (e.g. Cima melitensis Mifsud,

1998, Figs. 1- 4). This latter type, resembles the pro-

toconch of some Murchisonella Mork, 1875

(Waren, 2013) but is clearly distinguishable. This

double typology of protoconch can be recognized in

the fossil species referred to “CYma” (Waren, 2013).

The known species attributable to the genus

Cima, for the Mediterranean and European sea

waters (C. minima; C. cylindrica Jeffreys, 1856;

C. cuticulata Waren, 1993; C. inconspicua Waren,

1993; C. apicisbelli Rolan, 2003) consist of a

morphologically homogeneous group. One excep-

tion is “C/ma” melitensis, described by Mifsud

(1998) from a limited number of specimens com-

ing from 80-100 m deep Malta’s waters, and

lacking of soft tissue. This species is characterized

by a pyramidelliform teloconch, surmounted by a

protoconch that seems truncated; at a first sight the

shell resembles closely Odostomia Fleming, 1813

or Liostomia G.O. Sars, 1878.

142

P. Mietto, I. Nofroni & E. Quaggiotto

The analysis at SEM of some shells highlighted

a very peculiar protoconch, certainly different

from Pyramidellidae: after an extremely small nu-

cleus (about 27 pm), the protoconch unwinds on

an horizontal axis for half a rotation; then it raises,

creating a small prominence easy visible in lateral

view (Fig. 4); then it goes down and continues

with a normal dextrorse envelopment. This kind

of protoconch is called hyperstrophic and is typical

of some families of Heterobranchia (Heterostropha)

such as Architectonicidae, Murchisonellidae, etc.

Basing on these morphological features we believe

that the collocation of this species into the genus

Cima is incorrect.

Chaster (1896) instituted the subgenus Cima (ex

Monterosato) without any description, just declar-

ing that “... which Jeffreys described as Odostomia

minima, for which species and the closely allied blit

quite distinct Jeffreysia cylindrica Jeffr, Mon-

terosato proposes the sub-genus Cima, a separation

with which I entirely concur ”

One first problem is to determine which is the

type species of Cima. Following Waren (1993) this

is Odostomia minima Jeffreys, 1858, while Bandel

(2005) indicates as type species Jeffreysia cylin-

drica Jeffreys, 1856. The work of Waren (1993)

is earlier and then the correct type species for

Cima is Odostomia minima.

Van Aartsen (1981) validates the separation be-

tween Cima and Pherusina Norman, 1888 ( =Aclis

Foven, 1 846), proposed by Monterosato, but, bas-

ing on morphological features such as the shape of

peristome, the embrional whorls, the clear flexuous

growth lines, he considers valid the position of the

genus in Aclididae. The same opinion is shared by

Fretter & Graham (1982), that nevertheless do not

exclude to put this genus in a new, to be created,

family Cimidae. Afterwards Graham (1988) pre-

serves the collocation of Cima in Aclididae.

Waren (1993), mainly on anatomical base, puts

Cima in the new monogeneric family Cimidae, into

the subclass Heterobranchia.

Bouchet & Rocroi (2005) maintain Cima in the

Cimidae family, into the Heterobranchia, but with-

out further collocation. Bandel (2005) discusses the

collocation of many genera, now included in Het-

erostropha, such as Aclis Foven, 1846, Hemiaclis

G.O. Sars, 1878, Graphis Jeffreys, 1867, Cima ,

Murchisonella Morch, 1875, and Ebala Gray,

1847. The result, based on anatomy, shell morphol-

ogy and evolutionary trend, is that all these genera

have to be included in different families. In partic-

ular, Cima is re-positioned in Streptacididae, a fam-

ily that includes other fossil genera from the

Paleozoic. Murchisonella is included in the Donal-

dinidae family, including fossil genera from Car-

boniferous. The Ebalidae (= Anisocyclidae) is

considered separated family.

Recently, Penas & Rolan (2013) reviewed the

genus Murchisonella and proposed using the

genus Pseudoaclisina Yoo, 1994 for the species

with rounded coils; in the same year Waren (2013)

published a study about Murchisonellidae, where

he analyzed this family and other similar ones,

basing on genetic, anatomical and palaeontologi-

cal features, providing guidelines on these small

Heterobranchia for future studies.

The morphological differences that all the Au-

thors recognize between Cima and Murchisonella

regard the shape of the protoconch and the presence

of growth lines: Cima does not have the deep sinus

close to the suture that characterizes Murchisonella.

Moreover, Murchisonella has a scalariform profile,

while in Cima it is rounded.

Considering the Cima species, both the type

species Odostomia minima Jeffreys, 1858 and then

Jeffreysia cylindrica Jeffreys, 1856 present a globu-

lar and slightly inclined protoconch (Figs. 5, 7, 8),

completely different from those of melitensis.

The features of the protoconch of C. melitensis

bring the species close to the genus Murchisonella ,

whose type species Murchisonia {Murchisonella)

spectrum Mork, 1875 (Fig. 9), comes from the Car-

ibbean area (Redfem, 2001). This genus is charac-

terized by an aclidiform shell, densely striated in

the middle and inferior part of the whorls, with a

sinus in the upper part of the external peristome

edge, and hyperstrophic protoconch.

The only species attributed to this genus, reported

in the Mediterranean, is Murchisonella mediterranea

Penas & Rolan, 2013 (= Murchisonella columna

Auctores not Hadely, 1807).

With both these species C. melitensis shares

only the protoconch and not the growth style or the

shape of the whorls that are in common with Cima.

Thus, we think that C. melitensis has to be attrib-

uted to a distinct genus, but none of the known

ones, both from fossil and living records, seems

suitable. So we believe it is necessary to institute a

new genus.

On the systematic position of “Cima”melitensis, with erection of the new genus Mifsudia (Heterobranchia Cimidae) 143

Figure l. Mifsudia melitensis, Mauritania, -80/100 m (1.1 mm). Figure 2. Mifsudia melitensis, Mauritania, -80/100 m (0.98

mm). Figure 3. Mifsudia melitensis, Mauritania, -80/100 m, protoconch of shell in Fig 2. Figure 4. Mifsudia melitensis, pro-

toconch, lateral view, Mauritania, -80/100 m. Figure 5. Cima minima, protoconch, Giannutri Island, Grosseto, Italy, -18 m.

Figure 6. Cima sp., Getares, Algeciras, Spain, beach (0.86 mm). Figure 7. Cima cf. cylindrica, Sorrento, Naples, Italy,

-50/60 m (1.48 mm). Figure 8. Cima cylindrica, Levanzo Island, Trapani, Italy, -31 m (1.45 mm). Figure 9. Murchisonella

spectrum, Varadero, Bahia de Cochinos (Cuba), -10 m (1.4 mm).

144

P. Mietto, I. Nofroni & E. Quaggiotto

ACRONYMS. Franco Gubbioli collection,

Marbella, Malaga, Spain = FG. Paolo Mietto col-

lection, Vicenza, Italy = PM. Italo Nofroni collec-

tion, Rome, Italy = IN. Ermanno Quaggiotto

collection, Longare, Vicenza, Italy = EQ.

RESULTS

SYSTEMATIC

Class GASTROPODA Cuvier, 1797

Subclass HETEROBRANCHIA Gray, 1840 (unas-

signed)

Infraclass HETROBRANCHIA Gray, 1840

Family CIMIDAE Waren, 1993

Genus Mifsudia n. gen.

Type species. Cima melitensis Mifsud, 1998:

Figs. 4, 5.

Examined material. “C/mu” melitensis. Lampe-

dusa Island, Cala Calandra, -30 m, legit M. Oliv-

erio, 1 shell (IN). Between Estepona (Malaga,

Spain) e Tetuan (Morocco), -25/35 m, 1 shell (IN).

Mauritania (West Africa, Atlantic Ocean), - 80/100

m, more than 40 shells, legit F. Gubbioli, (FG,

PM, IN, EQ). All inedited reports. Malta, Golden

Bay, -100/120 m, legit F. Carmona, 1 shell (EQ).

Cima minima : more than 60 shells from all the

Mediterranean Sea, from 0 to 180 m of depth

(PM, IN, EQ).

Cima cylindrica. More than 60 shells from all the

Mediterranean Sea, from 0 to 100 m of depth (PM,

IN, EQ).

Cima sp. 2 shells from Getares (Algeciras. Spain),

beach, inedited report (IN).

Murchisonella spectrum. Varadero (Cuba, Caraib

Sea), beach, legit C. Petrella, 1 shell (EQ). Bahia

de Cochinos (Cuba, Carribean), debris -10 m, legit

M. Chiodi, 7 shells (IN).

Murchisonella mediterranea. Aydincik (Turkey),

-27 m, , legit M. Oliverio, 1 shell (IN).

Murchisonella sp. Watamu (Kenia, Indian Ocean),

-32 m, legit L. Contessini, 1 shell (IN). Shaiab

Rumi (Sudan, Red Sea), - 60 m, legit M. Oliverio,

1 shell (IN).

Description. Small shell, white, bright, lacking

in the spiral sculpture, widely umbilicate, with the

shape similar to Odostomia; rounded whorl with

growth lines flexuous but lacking in the subsutural

sinus. External peristome edge thin and sharp,

lacking in sinus. Hyperstrophic protoconch with

probable planctotrophic development.

Etimology. The name has been coined in honor

of Constantine Mifsud, the well known Maltese mala-

cologist, discoverer of C. melitensis.

Remarks. Composition of the genus:

Mifsudia melitensis Mifsud, 1998 - living, Mediter-

ranean Sea and Atlantic Ocean

Mifsudia gantensis Bandel, 2005 - fossil, Middle

Eocene, Hungary (= Cima gantensis)

Mifsudia sp. (= Murchisonella n. sp.? pro Murchi-

sonella cf. obtusa Gougerot & Le Renard, 1978)

fossil, Early Oligocene, France, see below.

DISCUSSION

As previously discussed, the new genus Mifsudia

n. gen. differs from Cima for the shape of the pro-

toconch, hyperstrophic and not globular; the shape

of the growth lines, sinuous (sigmoids) but always

lacking in the subsutural sinus. The rounded shape

of the whorls is similar in both genera.

Mifsudia n. gen. and Murchisonella share the

same typology of hyperstrophic protoconch but not

the shape of the whorls, that in the latter is clearly

angular, nor the shape of the growing lines that in

Murchisonella is sigmoid and characterized by a

deep sinus in the subsutural area.

These differences occur also in the fossil forms,

at least from Lutetian (Middle Eocene).

Thus, to be included in Mifsudia there are:

1) Cima gantensis Bandel, 2005 from the Middle

Eocene of Hungary.

2) Murchisonella n. sp.? pro Murchisonella cf. ob-

tusa (in Gougerot & Le Renard, http.//www.so-

mali.asso.fr/fossils/biotaxis.php, fische batch LR-

71951) from the Early Oligocene (Stampiano Auct.)

of France.

According to Le Renard (http://www.somah.asso.fr/

fossils/biotaxis.php), JanssenA.W. (1984), because of

On the systematic position of “Cima”melitensis, with erection of the new genus Mifsudia (Heterobranchia Cimidae) 145

the features of the protoconch and of the growth

lines, have to be referred to Cima the fossil species

Cima gougeroti Le Renard, in schedis, from the

Lutetian of the Paris Basin (http.//www.

somali.asso.fr/fossils/biotaxis.php, batch 60859 and

61595), Cima microscopica Le Renard, in schedis,

from the Lutetian and “Biarritzian” of the Paris

Basin (http.//www.somali.asso.fr/fossils/biotaxis.

php, batch 73241), Aclis fStilbe ) proneglecta R.

Janssen, 1978 from the Upper Oligocene of Glim-

merode (Germany), Adis ( Stilbe ) neglecta A.W.

Janssen, 1969 from the Miocene of Dingden (Ger-

many).

According to Pachaud & Le Renard (1995)

should be referred to Murchisonella the fossil species

Adculina emarginata Deshayes, 1861, Murchi-

sonella densesulcata Gougerot, 1966 and M. obtusa

Gougerot & Le Renard, 1978, all from the Lutetian

of the Paris Basin.

The presence of clear distinctive character-

istics among these three genera, highlighted since

the Lutetian, supports the validity of the new

genus Mifsudia.

About what concerns the systematic collocation

of the new genus, we have to confess some embar-

rassment because the previous Authors used sev-

eral and different criteria for the collocation of the

genera at the family level. We think that what pro-

posed by Bandel (2005) is not completely share-

able because we separate Mifsudia from Cima due

to the protoconch shape, without considering other

anatomical characteristics.

Without starting a systematic discussion, it has

to be considered that Bandel (2005) and Waren

(2013) used as criterion the teloconch feature rather

than the protoconch. Following this rule, the ab-

sence of the characteristic sinus in the growing

lines, typical of Murchisonella and Pseudoa-

clisina, gives credit for the collocation of Mifsudia

in Cimidae.

ACKNOWLEDGEMENTS

A special thanks to all the friends, particularly

to Franco Gubbioli (Marbella, Malaga, Spain) that

gave the malacological samples used in this study,

to Matteo Belvedere (Padova, Italy) to Stefano

Meggio (Vicenza, Italy) for english translation and

to Stefano Bartolini (Firenze, Italy) for providing

us some colour photos (Fig. 1 and Fig. 8).

REFERENCES

Aartsen J.J. van, 1981. European marine Mollusca: notes

on less well-known species II. The genus Cima

Chaster, 1896. Basteria, 45: 117-119.

Bandel K., 2005. Living fossils among tiny Allogas-

tropoda with high and slender shell from the reef en-

vironment of the Gulf of Aqaba with remarks on fos-

sil and recent relatives. Mitteilungen aus dem

Geologisch-Palaontologischen Institut der Universitat

Hamburg, 89: 1-24.

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

clator of Gastropod Families. Malacologia, 47: 1-

397.

Chaster G.W., 1896. Some new marine Mollusca from

Tangier. Journal of Malacology, 5: 1-4.

Fretter V. & Graham A., 1982. The Prosobranch Mol-

luscs of Britain and Denmark. Part 7 - “Heterogas-

tropoda” (Cerithiopsacea, Triforacea, Epitoniacea,

Eulimacea). Journal ofMolluscan Studies, 11 suppl.:

363-434.

Graham A., 1988. Molluscs: Prosobranch and Pyra-

midellid Gastropods. The Linnean Society of London

and the Estuarine and Brackish- Water Sciences As-

sociation. Synopses of the British fauna (NS), 2: 1-

662.

Janssen A. W., 1984. Mollusken uit het mioceen van Win-

terswijk-Miste. Een inventarisatie, met Geshrijvingen

eu afbeeldingen vom alle aangetroffen soorten. Konin-

klijke Nederlandse Naturhistorische Vereninging,

Amsterdam, 1 : 45 1 + 82 platen.

Le Renard P. http://www.somali.asso.fr/fossils/biotaxis.

Php

Mifsud C., 1998. Pseudographis cachiai n. gen. e n. sp.,

e Cima melitensis n. sp.: due specie nuove di Het-

erostropha (Mollusca, Gastropoda) dell’arcipelago

maltese. La Conchiglia, 286: 25-29.

Pachaud J.M. & Le Renard P, 1995. Revision des mol-

lusques paleogenes du Bassin de Paris. IV - Liste

systematique actualise. Cossmanniana, 3 : 151-187.

Penas A. & Rolan E., 2013. Revision of the genera

Murchisonella and Pseudoaclisina (Gastropoda, Het-

erobranchia, Murchisonellidae). Vita Malacologica,

11: 15-64.

Redfem C, 200 1 . Bahamian Seashells. A thousand species

from Abaco, Bahamas. Bahamianseashells.com, Inc:

Boca Raton, Florida. 280 pp., +124 pis.

Waren A., 1993. New and little known Mollusca from

Iceland and Scandinavia. Part 2. Sarsia, 78: 159—

201, Bergen.

146

P. Mietto, I. Nofroni & E. Quaggiotto

Waren A., 2013. Murchisionellidae: who are they, poda, lowermost Heterobranchia). Vita Malacolo-

where are they and what are they doing? (Gastro- gica, 11: 1-14.

Biodiversity Journal, 2014, 5 (2): 147-150

Monograph

Natural values, coastal and marine ecosystems of the Circeo

National Park: conservation priorities

Stefano Raimondi

Circolo Laras, Legambiente Volontariato di Sabaudia. Strada Piscinara destra 2, 04100 Latina, Italy; e-mail: s.raimondi@legambiente.it

ABSTRACT The variety of environments that characterizes the Circeo National Park must also take into

account, in addition to the terrestrial natural values that are present, even the importance of

marine and coastal stretches that currently do not benefit from a similar regime of protection,

preserving instead important elements of wealth for marine biodiversity. This added value is

represented in a particular way by the presence of Posidonia oceanica (L.) Delile, 1813, habitat

of Community interest. The proposal of the Plan of the Park to extend to the sea The Circeo

National Park would help to protect and enhance areas for the most part already included in

the Natura 2000 network that could be handled in a unified manner by the Park providing for

their conservation through various management interventions. Another proposal involves,

instead of creating a true marine protected area, encompassing the whole Posidonia meadows

present both in the northern section of the coast and in the south, between San Felice and

Terracina and, hopefully also the stretch of sea that surrounds the island of Zannone (therefore

including the SPAs area regarding the Pontine Archipelago).

KEY WORDS Circeo National Park; marine protected area; Posidonia oceanica', Natura 2000 Network.

Received 28.01.2014; accepted 21.03.2014; printed 30.06.2014

Proceedings of the Seventh Malacological Pontine Meeting, October 5 th - 6 th , 2013 - San Felice Circeo, Italy

INTRODUCTION

The Circeo National Park, established by Law

No. 285 of 25 January 1934 in order to "... protect

and improve the flora and fauna, preserving the

special geological formations and the beauty of the

landscape and promote the development of

tourism," after have suffered several perimetral

changes over the years is now extended for a little

less than 9000 ha., while protecting currently only

a strip of land characterized by the presence of five

different environmental situations that make up the

rich mosaic that characterizes it: the plain forest, the

promontory, the coastal dune, coastal lakes and the

adjoining wetlands, the Island of Zannone part of

Pontine Islands.

The importance and value of the protected area

is especially evidenced by the layering of several

legislative instruments at regional, national, Com-

munity and international level that protect the ter-

ritory and biological forms and abiotic ones

preserved in it.

Regulatory instruments at the regional, national,

European and international level

Institution of the CNP with Law no. 285 of January

25, 1934

Institution of five natural reserves of the State

(1971-1979)

Bonn Convention - conservation of migratory

wildlife species

148

Stefano Raimondi

Berne Convention - Wildlife and conservation of

the natural environment in Europe

Ramsar Convention - conservation of wetlands of in-

ternational interest (presence of 3 sites included)

CITES - regulating international trade in endange-

red species of flora and fauna

Directive 79/409/EEC (Directive 2009/147/EC

“Birds Directive”)

“Habitats Directive” (92/43/EEC)

Framework law on protected areas 394/91

Act No 157 of 11 February 1992 (Omeoterma

wildlife protection and hunting)

Regional Law 18/88 (protection of minor fauna)

Presence of 2 IB A sites (Birdlife International)

SPAIT6040015 Parco Nazionale del Circeo

SPAIT6040019 Isole di Ponza, Palmarola, Zannone,

Ventotene e S. Stefano

SCI IT6040013 Lago di Sabaudia

SCI IT6040012 Laghi di Fogliano, Monaci, Capro-

lace e Pantani dell'Inferno

SCI IT6040014 Foresta Demaniale del Circeo

SCI IT6040016 Promontorio del Circeo (Quarto

Caldo)

SCI IT6040017 Promontorio del Circeo (Quarto

Freddo)

SCI IT6040018 Dune del Circeo

SCI IT6040020 Isole di Palmarola e Zannone

The Circeo National Park includes 5 nature

reserves established from 1971 to 1979. Among the

international conventions ratified by our country

that involve it directly, we can mention the Bonn

Convention on the Conservation of Migratory

Species of wildlife, the Beme Convention for the

conservation of the natural environment in Europe

and the wild species, the Ramsar Convention for

wetlands of international value, the Washington

Convention (CITES) for the regulation of interna-

tional trade of endangered species of flora and

fauna, the Habitats and Birds Directives, the frame-

work law on protected areas 394/91, the law num-

ber 157 of 1992 for the protection of homeothermic

wildlife from hunting, and even the regional law

18/88 for the protection of minor fauna. Finally, we

must remember both the presence of two Important

Bird Areas as well, with regard to Community

directives, the two SPAs and the seven SCIs that

insist throughout the area going to generate a multi-

layered system of protection at different levels both

normative and territorial, which bear witness to the

importance of these places. Nevertheless, thought

it was the first, and for a long time the only coastal

national park in Italy, the sea has always been

regarded in second place in the environmental pro-

tection policy.

DISCUSSION

In spite of the attention has always been focused

solely on terrestrial environments of the National

Park of Circeo, of great importance is also the

stretch of sea between Capo Circeo and the Pontine

Islands. In this area were reported indeed marine

mammals as bottlenose dolphin ( Tursiops truncatus

Montagu, 1821), striped dolphin ( Stenella coemleoalba

Meyen, 1833), common dolphin ( Delphinus delphis

Linnaeus, 1758), sperm whale ( Physeter macro-

cephalus Linnaeus, 1758), and Globicephala spp.

while in recent years are occurring ever more fre-

quent sightings (and strandings) of the sea turtle

Caretta caretta Linnaeus, 1758.

In the sea in front of the National Park, it is also

noted the presence of the following further species

present in the annexes of the Habitats Directive (in

general, some of the marine species listed in the

"Report of the Italian fauna protected" written by

the MATT, a document that lists protection status

and health of every animal species and related legis-

lation, are present along the coasts of Circeo):

Corallium rubrum Linnaeus, 1758. Red Coral,

Habitats Directive, Annex V - reported presence at

the site IT6040019

Pinna nobilis Linnaeus, 1758. Fin noble,

Habitats Directive, Annex IV - Reported presence

at the site IT6040020 and IT6000013

Scyllarides latus Latreille, 1802. Slipper

Lobster, Habitats Directive, Annex V - Reported

presence at the sites IT6040019 and IT6000013

Petromizon marinus Linnaeus, 1758. Sea

lamprey, Habitats Directive, Annex II - Reported

presence with a not very significant population at

the site 1T6040019

Aphanius fasciatus Valenciennes, 1821. Killfish,

Habitats Directive, Annex II - Reported presence at

the sites IT6040012 and IT6040013

Natural values, coastal and marine ecosystems of the Circeo National Park: conservation priorities

149

The Lazio Region, within the programs Beachmed

(Interreg) and ICZM (Integrated Coastal Zone

Management), has produced two publications and

analysis on the priorities for the conservation of

coastal and marine natural values of the region

(BEACHMED, 2004; DECOS, 2006; 2007). The

coast of Circeo (particularly the area in front of

Torre Astura, the area in front of the lakes and the

one between Capo Circeo and Terracina) turns out

to be one of the most interesting especially for the

presence of Posidonia, which performs nursery

function for fish and benthos. It should be recalled

in this connection that the seagrass meadows are

considered a priority habitat of interest at European

level; in Italy, are further protected and secured

since 2001 (Decree Law No. 93 of March 2001).

“La Sapienza” University of Rome has recently

developed on behalf of the city of San Felice

Circeo, a project for the knowledge and conserva-

tion of Posidonia meadows in the area of Circeo

(Universita degli studi di Roma “La Sapienza”,

Regione Lazio, 2008; Nascetti & Martino, 2009).

According to this work, the current distribution of

grasslands appears to be profoundly changed com-

pared to what is reported in the works described

above. Between Capo Portiere and Torre Astura

there is a compact Posidonia meadow, with a rela-

tively high density and settled on "matte", less com-

pact due to the presence of large areas of erosion in

slightly more depth and patches up to 31-32 m

deep; some areas of this grassland show signs of an

advanced state of regression while in others are

present rock structures. Posidonia is present along

the shoreline in front of Fogliano Lake, while it is

absent between the headland and the mouth of

Caprolace lake where it is present Cymodocea no-

dosa (Ucria) Asch., 1870 (Ardizzone & Belluscio,

1996; Diviacco et al., 2001)

The Posidonia meadow placed in front of the

promontory of Circeo is what seems to be the least

changed over the years, probably because localized

further out than the others, and therefore less

influenced by the contributions of continental

waters, but also thanks to the presence of rocky sub-

strates that probably have limited the activities of

illegal trawling. In fact, the most western front of

the Circeo promontory, has a meadow in good con-

dition, settled on the rock in the middle part and on

“matte” all around. A narrow zone with isolated

bundles of Posidonia on dead "matte" is present in

the vicinity of the bottom margin (Bouchette et al.,

2008; Ardizzone et al., 2009; Nascetti & Martino,

2009). The central meadow, between Cape Circeo

and Terracina, presents the most important regres-

sion of this stretch, especially evident with the re-

treat of the lower limit. This stretch of coastline was

subject to heavy changes in the coastline due to the

increased human pressure, with negative influences

on both the quality of water on the grain size of the

seabed. Most of the area is therefore occupied by a

meadow extremely rarefied with large areas of dead

"matte" (Bouchette et al., 2008; Ardizzone et al.,

2009; Nascetti & Martin, 2009).

Summing up therefore the expeditious visual

analysis of environmental emergencies and major

areas of interest of the Latium coasts, we can iden-

tify two major areas of interest in offshore and sub-

coast. Among the areas of greatest interest for the

establishment of Marine Protected areas there is the

offshore area in front of the Circeo; among the

inshore areas the stretch between Capo Circeo and

Terracina is particularly interesting for its fertility

due to its geological features.

The Plan of the Park proposes a widening in the

sea of the National Park that would include as well

an extraordinary variety of plant and animal spe-

cies, protecting and enhancing the features of the

marine and coastal biodiversity, also and above all

through environmental restoration measures. For

these reasons could be implemented programs of

study, monitoring and scientific research in the

fields of natural science and environmental protec-

tion, with the aim of ensuring the systematic

knowledge of the area, but also for the promotion

of sustainable development of the environment,

with particular emphasis on promoting traditional

activities of local cultures, tourism and environmen-

tally friendly use.

The areas involved by the proposal for extension

of the CNP to the sea are largely already included

in the Natura 2000 Network. With this proposal,

according to the editors, the management of the off-

shore part of the “SPA IT6040015 Parco Nazionale

del Circeo” and marine SCI that face the sea coast

of the Park and of Zannone (SPAs "Pontine

Islands") may be carried out in a unified manner by

Park Authority that can ensure their preservation

through various management interventions. This

solution, also, would fully respond to international

commitments in the European context for the

150

Stefano Raimondi

management of such sites, providing them with

concrete organizational, and financial skills that

would ensure the achievement of the objectives of

the Birds and Habitats Directives. The proposal put

forward at the time by Legambiente was, instead of

setting up a real marine protected area, to include,

rather, the entire complex of the seagrass meadows

present both in the northern section of the coast and

in the south ( between San Felice and Terracina).

Indeed, the marine area should not only encompass

the entire SPA Circeo National Park, but we imag-

ine that, facing if necessary with any reluctance of

local municipalities, the park could also be a pro-

moter of an extension of the area to the sea area

surrounding the island of Zannone (therefore com-

prising part of the second SPA of the Park, the one

concerning the archipelago of the Pontine Islands).

The ideal reference is to the “5 Terre National

Park” where the synergy between marine protected

area (institute aimed to the protection of marine

environments) and the national park (which protects

instead the ground part) has created an enviable

model of resource management and, at the same

time, of valorization of the tourist routes on which

the socio-economic system of the district is largely

based.

CONCLUSIONS

The coastal stretch in front of the National Park

of Circeo between Capo Circeo and Terracina and

between Astura and Capo Circeo, is one of the

coastal areas of most interest throughout the region

of Lazio in consideration of the presence of impor-

tant Posidonia oceanica meadows. The importance

of submerged marine vegetation, especially of the

systems of marine phanerogams, was now scien-

tifically recognized as crucial for their contribution

to the maintenance of infralittoral ecosystems. This

aspect suggests a priority in the preservation of the

natural values of the coastal and marine areas.

REFERENCES

Ardizzone G.D. & Belluscio A., 1996. Le praterie di Po-

sidonia oceanica delle coste laziali. In: il Mare del

Lazio. Regione Lazio - Universita degli Studi di

Roma “La Sapienza: 194-217.

Ardizzone G.D., Belluscio A., Barani P. & Criscoli A.,

2009. Rilievo e caratterizzazione delle praterie di

Posidonia antistanti le coste della regione Lazio e dei

principali popolamenti marini costieri per la realiz-

zazione di una cartografia dei fondali della regione

Lazio e la predisposizione di un atlante degli habitat

marini. Convenzione La Sapienza Universita di

Roma, Dipartimento di Biologia Animale e dell'Uomo

- Regione Lazio, Direzione regionale ambiente e

cooperazione tra i popoli. Rapporto II fase.

BEACHMED, 2004. II Progetto Beachmed: Recupero

ambientale e manutenzione dei litorali in erosione,

mediante l'impiego dei depositi sabbiosi marini,

Primo quademo tecnico fase A, seconda edizione.

www.beachmed.it

Bouchette F., Denamiel C., Lamberti A., Yorgos S.,

Deserti M., Ardizzone G.D. & Belluscio A., 2008.

Caratterizzazione delle condizioni idrometeoro-

logiche in zona litorale e analisi dei rischi costieri, del

comportamento delle opere di difesa e della dinamica

delle praterie di Posidonia oceanica, sottoprogetto 2.2

NAUSICAA, progetto Beachmed-e: 47-59.

DECOS, 2006. Azione 1.1.7 Programma integrato di in-

terventi per lo sviluppo del litorale del Lazio “Speri-

mentazione ICZM in aree pilota” Stato di

avanzamento del progetto, progetto ICZM, fase 1,

working paper

DECOS, 2007. Azione 1.1.7 Programma di interventi per

lo sviluppo del litorale del Lazio; Identificazione dei

criteri di scelta delle aree pilota ed analisi di dettaglio,

fase 2, working paper.

Diviacco G.D., Spada E. & Lamberti C., 2001. Le fa-

nerogame marine del Lazio. Descrizione e cartografia

delle praterie di Posidonia oceanica e dei prati di

Cymodocea nodosa. Ed. ICRAM, 113 pp.

Nascetti G. & Martino S., 2009. Valutazione dello stato

di conservazione delle aree marine della Regione

Lazio e Analisi di fattibilita per l'istituzione di aree

marine protette o di tutela biologica a livello re-

gionale. Rapporto prima fase. Universita degli Studi

della Tuscia, Dipartimento di Ecologia e Sviluppo

Economico e Sostenibile (DECOS) Regionale Lazio.

Universita degli studi di Roma “La Sapienza”, Regione

Lazio, 2008. Rilievo e caratterizzazione delle praterie

di Posidonia antistanti le coste della regione Lazio e

dei principali popolamenti marini costieri per la

realizzazione di una cartografia dei fondali della Re-

gione Lazio e predisposizione di un atlante degli habitat

marini. Rapporto prima fase.

Biodiversity Journal, 2014, 5 (2): 151-164

Monograph

The continental molluscs from Mount Circeo (Latium,

Italy)

Alessandro Hallgass * 1 & Angelo Vannozzi 2

'Via della Divina Provvidenza 16, 0 0166 Roma, Italy; e-mail: hallgass@hotmail.com

"Via Pietro de Cristofaro 46. 0 0 136 Roma. Italy; e-mail: ang.vannozzi@gmail.com

ABSTRACT This paper is the second step in a process that aims to asses biodiversity of land and freshwater

molluscs fauna of Mount Circeo (Latium, Italy). Forty species of land and freshwater molluscs

are listed, three more than in the previous work. A species of OxychiluS Fitzinger, 1 8 3 3 and

two species of himQX L in n ae u s , 1 75 8 remain undetermined, to date. The presence of PleU-

rodiscus bcilmei bcilmei (Potiez et m ichaud, 1 8 3 8 ) and Siciliaria gibbula honii (O . b oettger,

1 8 79) are confirmed and this is the known northern limit of their distribution areas in Italy.

M oreover, the presence of some species of biogeographical interest has allowed us to formulate

some hypotheses on the origin of this fauna, in the light of the most recent theories on the

formation of the Italian peninsula.

KEY WORDS Continental molluscs; M ount Circeo; checklist.

Received 28.01.2014; accepted 21.03.2 0 14; printed 30.06.2014

Proceedings of the Seventh Malacological Pontine Meeting, October 5 tll -6‘ h , 20 13 - San Felice Circeo, Italy

INTRODUCTION

The 2 nd Malacological Pontine Conference of

2008, gave us the opportunity to presented a first

contribution on the biodiversity of land molluscs

of Mount Circeo (Hallgass & Vannozzi, 2008). The

research carried out in this territory, which pro-

gressed in the recent last years, allowed us to up-

date our knowledge of this fauna which is reported

in th e fo llo w in g .

Study area

M ount Circeo is a promontory composed mostly

of marl and sandstone from the Paleogene and of

limestone from the lower Early Jurassic, which is

different from that constituting the nighbouring

Ausoni Mountains (Fig. 1). The shape of the pro-

montory is elongated in east-west direction with a

length of about 6 km , and a maximum heightof541

m above sea level (reached at the “Pizzo diCirce”);

otherimportant heights are the “Semaforo” (412 m)

and the “Le Cro cette” (3 5 2 m).

The geographical position of the promontory

contributes to create some sub-environments in

relation to their different exposure:

• “Quarto Fred do” with northern exposure.

• “Quarto Temperato” with western exposure; it is

a transition from the south side to the north side,

therefore it has no particular interest.

• “Quarto Caldo” with southern exposure.

• “Quarto Comunale” with eastern exposure; it is

anthropized and altered and therefore of little natu-

ralistic interest.

152

Alessandro Hallgass & Angelo Vannozzi

Our interest was, therefore, focused on the two

slopes, the northern and the southern ones. The veg-

etation in Mount Circeo is typically Mediterranean

but is very different by structure and composition

in the two mains slopes.

Quarto freddo. The vegetation on the north-

ern slope of the mountain is made up of a dense for-

est of tall trees dominated by holmoak. At the

baseline, however, there are scattered examples of

Italian oak, oak, and hornbeam, which represent the

penetration of the neighboring plainforest.

Other trees or shrubs species well represented

are: mock privet, ash, arbutus, heather and buck

thorn. Towards the plain, however, the beautiful

“Sughereta di Mezzomonte”, whose underwood is

characterized by fallen branches of which soon

remains only the thick bark that provides a shelters

for many species of molluscs (Figs. 3, 6). At about

200 meters above the sea level, we observed there

forestation of conifers that made the canopy of the

forest higher; the oaks adapted and have a very slen-

der truncks while the cork oaks were not able to

reach the canopy and are found as skeletons in the

underw ood. A t the top of the mountain on both lopes

there is the limestone exposed, which is the exclu-

sive habitat of strictly calciphilous species.

Quarto caldo. In the medium and high area of

the mountain, the vegetation is very compact and

consists of high and low maquis species dominated

by arborescent layer of oak with plenty of ash, arbu-

tus, mastic, heather and broom. The low maquis is

prevalent mostly in the lower part of the promontory

and is dominated by mastic, phillyrea, myrtle, and

holm oak which, in addition, shows a bushy shape.

Very interesting is the presence of several specimens

of the dwarf palm (ChdJfU 26 WpS Huiflilis L .), the only

spontaneous palm in Italy. On all the rock sand cliffs

overlooking the sea, we found a discontinuous

grouping characterized by lavender pillows placed

near to the ground, sea fennel and compact clumps

of weeds beaches, sometimes associated to the enula.

In more humid valley sholm returns to be dominant

in th e tree shape.

On the promontory several species of m o llu s c s

from different neighboring environments can be

found. The thermophilic species can reach the

promontory from the adjacent dune and colonize

hot-arid environments. From the adjacent plain

forest are the species characteristic of a moist en-

vironment; however, much more interesting are

the strictly c a lc io p h ilo u s species that could be the

remains of the old fauna of the promontory. For

some species, we cannot rule out an anthropogenic

input.

MATERIAL AND METHODS

The research of terrestrial molluscs was made

by manual sampling for most species. The list of

localities under consideration is shown in Table 1.

Collected samples were deposited in the rnala-

cological collection of the “Museo del M are e della

Costa” of Sabaudia. Some very small species, isolated

Mount

Circeo

Circeo National

Park

Sabaudia •

San

Felice

Circeo

San Felice

Circeo

Tyrrhenian

Sea

Figure 1. Study area: Mount Circeo, Latium, Italy.

The continental molluscs of Mount Circeo (Latium, Italy)

153

locality

slope

coordinates

altitude

a.s.l. (m)

type of

environments

From the road to “le Crocette” at

the beginning of the trail to “la

Guardia di Orlando”

Q u arto fre d d o

4 1 ° 1 4 ' 1 5 ” N

1 3°04'57”E

192

Ilex (H oily) trees

Top trail between “il Semaforo”

and “ il Fortino Rosso”

Q u arto fre d d o

4 1 0 1 3 ’ 5 5 ” N

1 3°04'09”E

352

High m aquis

Source of “Mezzo monte”

Q u arto fre d d o

4 1 ° 1 4 ' 2 3 ” N

1 3°04' 1 3 ”E

Dense woods of Ilex

(Holly) and cork oak trees

“La Cava”

Q u arto fre d d o

4 1 ° 1 4 ' 3 6 ” N

1 3°04'55”E

drystony ground and

limestone wall exposed

Beginning of trail 1 (Torre Paola)

Q u arto fre d d o

4 1 ° 1 4 ’ 4 8 ” N

1 3 ° 0 2 ’ 2 3 ”E

H oily trees

Crossroads between“del Faro”

road and “del Sole” road

Q uarto caldo

4 1 ° 1 3 ’ 2 5 ” N

1 3°03’59”E

limestone wall exposed

and low m aquis

On the walls of the houses of

“S. Felice al Circeo”

Quarto comunale

4 1 ° 1 3 ’ 6 0 ” N

13°05’16”E

ruderal

Table 1. List of sampled localities of Mount Circeo (Latium, Italy).

species

figs.

Platyla c f . microspira (P in i, 1 8 8 4 )

Pomatias elegans (0 .f. m uiier, 1 7 74)

V V V

V V

2, 12

Pseudamnicola moussonii (C aicara, 1 8 4 1 )

Islamia pusilla (Piersanti, 1 952)

Galba truncatula to .f. m uiier, l 774)

Carychium tridentatum ( r i s s o , l 8 2 6 )

Rupestrella philippii (C an train e, 1840)

V V V

1 3

Granopupa granum (Draparnaud, 18 0 1)

1 4

Acanthinula aculeata (O .f. m uiier, l 774)

Lauria cylindracea (Da Costa, 1 7 7 8 )

V V 1

1 5

Pleurodiscus balmei { Potiez et Michaud, 1838)

C C

1 8

Table 2 (continued). List of continental molluscs found in Mount Circeo (Latium, Italy) in the seven localities reported in

Table 1. Additional species collected by M.Bodon (pers. comm.) are listed in coloumn B. Abbreviations, v: living specimen;

c: empty shell. Single specimen: +; some specimens (2-4): + +; several specimens: + + +. In the case of many living speci-

mens, data on empty shells have been omitted. 1 in wall; 2 ilex wood; 3 juv.; 4 on road cut rocks.

154

Alessandro Hallgass & Angelo Vannozzi

species

figs.

Chondrula tridens (O.f. m uiier, 1774 )

c c

1 7

Punctum pygmaeum (Draparnaud, 18 0 1)

Discus rotundatus (O.F.Muiier, 1774 )

V V V

V V

Vitrea contracta ( w e s teriu n d , 1 8 7 1 )

1 6

Oxychilus sp .

c c

V -c c

c c

c c 2

1 9

Daudebardia rufa maravignae (Pirajno, 1 8 40)

Tandonia sowerbyi (Ferussac, 1 8 2 3 )

v v 3

Umax s p . 1

v v 3

V V

Umax sp. 2

V V

Cecilioides acicula (0 .f. m uiier, 1774 )

Rumina deco llata (Linnaeus, 1758)

c c

Leucostigma candidescens (Rossmassier, 1 8 3 5 )

v v v4

V V V

v v v 4

8, 22

Cochlodina incisa (Kuster, 1 8 7 6)

V V

20,2 1

Siciliaria paestana (Philippi, 1 8 3 6 )

V V V

V V

V V V

c c 2

9,25,26

Siciliaria gibbula honii (0 . Boettger 1 8 7 9 )

V V

Papillifera bideUS (Linnaeus, 1 7 5 8 )

V V V

23,24

Xerotricha apicina (Lamarck, 1822 )

V -c c c

Xerotricha conspurcata (Draparnaud, 1801)

V V -c c c

3 1

Hygromia cinctella (Draparnaud, 1801)

Cernuella cisalpina (Rossmassier, 1 837)

V V -c c c

Trochoidea trochoides (Poiret, 1 789)

V V V

Cochlicella acuta (0 .f. m uiier, 1 774 )

V V V

Cochlicella conoidea (D raparnaud, 1801)

V V

Chilostoma planospira (Lamarck, 1 8 2 2 )

V V

10,36,37

Marmorana muralis (0 .f. m uiier, 1774 )

c c

Marmorana fuscolabiata circeja (Kobeit, 1 9 0 3 )

v v-cc4

V V -c c

11,39

Eobania vermiculata (O.f. m uiier, 1774 )

c c

V -c c c

c c

Cantareus apertus (Born, 1 1 1 8 )

c c

Cantareus aspersus (0 .f. m uiier. 1 774)

c c

Table 2 . List of continental molluscs found in Mount Circeo (Latium, Italy) in the seven localities reported in Table 1.

Additional species collected by M. Bodon (pers. comm.) are listed in coloumn B. Abbreviations, v: living specimen; c:

empty shell. Single specimen: + ; some specimens (2-4): + + ; several specimens: + + + . In the case of many living specimens,

data on empty shells have been omitted. 1 in wall; 2 ilex wood; 3 juv.; 4 on road cut rocks.

The continental molluscs of Mount Circeo (Latium, Italy)

155

Figs. 2-9. Land snails from Mount Circeo. Fig. 2. PomatidS elegans. Fig. 3. DisCUS WtWldatUS . Fig. 4. Tandonia SOWerbyi.

Fig. 5. Limaxsp. 1 . Fig. 6. Limaxsp. 2 . Fig. 7 . Limax maximus from Terracina. Fig. 8. Leucostigma candidescens. Fig. 9 .

SicUiaria paestana.

156

Alessandro Hallgass & Angelo Vannozzi

by sieving litter and soil, and some freshwater

species found in the source Coppelia in San Felice

Circeo, were collected by M . Bodon and have been

added to the list of species found in this study

(Table 2). It was not possible to carry out surveys

of freshwater species in the source “M ezzomonte”,

the main source of the promontory, which provides

water to the town of San Felice Circeo and now

completely captured and inaccessible.

For the systematic nomenclature we referred

mainly to the Checklist of the species of the Italian

Fauna (Bodon etal., 1995;Manganellietal., 1995);

for the supra-generic systematic see the “Classifi-

cation and nomenclator of gastropod families”

(Bouchet & Rocroi, 2005).

All specimens illustrated are from the Mount

Circeo, unless otherwise stated.

RESULTS

At present, 40 species of continental molluscs

are known from Mount Circeo, the mostinteresting

of which will be brie fly commented on below. The

full list is shown in Table 2.

Taxonomic list

Farn ilia A C IC U LID A E

Platyla c f . microspira (P in i, 1 8 8 4)

This species is morphologically referable to P.

microspira and is also known from the rear Ausoni

Mountains (Bodon & Cianfanelli, 2008). There still

remains, however, uncertain ty concerning the id en -

tity of the species, taken into account the geo-

graphic isolation from the typical populations of

Lombardy and Liguria.

Lam ilia CHONDRINIDAE

Rupestrella philippii ( C a n tr a in e , 1840)

Two empty shells of this small species (Lig. 13)

were found among the rocks of a road cut along

with Granopupa granum (D raparnaud, 180 1). The

association of these two species has already been

documented by Giusti (1970) for the Pianosa Is-

land. It would be extremely useful to study these

two small species from a genetic stand point, being

almost impossible to understand the relationships

among their populations either by anatomical or

morphometric analysis.

Granopupa granum (D raparnaud, 1 8 0 1 )

This small species (Pig. 14) is widespread in

Italy and in H o lo m e d ite r r a n e a n - M a c a r o n e s ia n -

Turanian region, albeit d isc o n tin u o u s ly, since it is

closely associated with a limestone substrate. It is

common in Mount C ire e o and th e shell is morpho-

logically little variable.

Pamilia PL E U R O D IS C ID A E

Figs. io, ii. Land snails from Mount Circeo. Fig. to. Chilostoma planospira. Fig. li. Mamiorana fiiscolabiata circeja.

The continental molluscs of Mount Circeo (Latium, Italy)

157

Pleurodiscus balmei balmei (Potiez et M ichaud,

1 8 3 8)

Only some empty shells of this species were

found (Fig. 18); however, one of them was a very

fresh shell, and we believe that this species actually

lives in Mount Circeo. This is the most northern

record of the species in the Italian peninsula. Re-

cently it has also been reported for Apulia (Ferreri

et al., 2005; Bassi, 2007). Instead, it is well known

for southern Calabria, Sicily and M alta (Giusti et al.,

1 995). The population of Mount Circeo could be a

relict population, but we cannot exlude a passive in-

troduction by man, as balmei is frequent in ruderal

habitats, gardens etc. (Kerney & Cameron, 1 979).

Fa m ilia D IS C ID A E

Discus rotundatus (O f. m oiler, 1774 )

A very common species and widely distributed

throughout Italy and Europe, D. VOtUYldcitUS is char-

acteristic of the underbrush (Fig. 3). The diameter

of the shell does not exceed 7 mm. In the forests

of the “Quarto Fred do” the thick bark of fallen

branches of cork oaks creates an ideal habitat for

this small gastropod.

Familia P R I S T I L O M A T I D A E

Vitrea contracta (w esteriund, 1 87 1 )

It has been found only a shell of this European

widespread species (Fig. 16). In Italy it is found at

low altitudes along the coast, while at higher al-

titudes it is replaced by other congeneric species

(G iusti et al., 1 9 8 5).

Familia OXY CHILIDAE

Oxychilus sp.

One living specimen and some shells of this

species were found (Fig. 1 9 ). A lthough it is rnorpho-

logically similar to a small form of O. dvapaVYiaudi

(Beck, 1837), it will be necessary to investigate

anatomical features in detail to determine its spe-

cific identity.As already pointed out by Manganelli

& Giusti (2001), species of OxycHHuS Fitzinger,

1 8 3 3 with well characterized genitalia may show

id e n tic al shells.

F am ilia M IF A C ID A E

Tandonia sowerbyi (Ferussac, 1 8 2 3 )

The species of TaYldOYlia Fessona et Pollonera,

1 882 genus are necrophagous and carnivorous mol-

luscs. Only some juveniles were observed in mount

Circeo. Externally it is distinguished from con-

geners by the dorsal hull orange and the clear sole

(Fig. 4). It is widely spread throughout Italy, prob-

ably dispersed by man.

F am ilia F IM A C ID A E

Limax sp . 1

This species (Fig. 5) is of medium size for the

genus, about 10-12 cm long; it is characterized by

colour, o c h e r- y e llo w , with a lighter uniform sole,

colour similar to LimaX bivOtlCie L e s so n a et Pollo-

nera, 1882, repor ted for north-eastern Sicily (Reitano

et al., 2007). It was sampled at different localities

of the promontory.

Limax sp . 2

Same size as the previous one, with darker color

and a pattern of small irregular spots limited to the

clypeus, looking as a variation of L. maximUS Fin-

naeus, 1 75 8. Only two specimens were found in the

cork trees of M ezzomonte (Fig. 6). It is currently not

possible to determine whether LiffiaX sp. 1 and

Limax sp. 2 belong to a single or tw o distinct species.

For the external morphology, both species are at-

tributable to the group of L. maximUS. However a

few m iles from these findings, out of the promontory,

towards Terracina, we observed L. maximUS speci-

mens with the characteristic spotted livery (Fig. 7).

Familia CFAUSIFIIDAE

Leucostigma candidescens (Rossm assler, 1835)

This species (Figs. 8, 22) is strictly c ale iop h ilo u s

158

Alessandro Hallgass & Angelo Vannozzi

Figs. 1 2 - 1 9. Land shells from M ount C irceo . Fig . 12. PomatiaS elegCUlS. Fig. 13. Rupestrdla philippU. Fig. 14. Granopupa

granum. Fig. 15. Lauria cylindracea. Fig. 16. Vitrea contracta. Fig. 17. Chondrula tridens. Fig. 18. Pleurodiscus balmei.

Fig. 19. Oxychilus sp.

The continental molluscs of Mount Circeo (Latium, Italy)

159

Figs. 20-3 3. Land shells from Mount Circeo. Fig. 20, 21. Cochlodina indsa. Fig. 22. LeUCOStigfiia CandideSCenS. Figs. 23,

2 4 . Papillifera bidens. Figs. 25 , 26 . SicUiaria paestana. Fig. 27 . S. gibbula honii. Fig. 28 . Rumina decollata. Fig. 29 . Xero-

tricha apicina. Fig. 30 . Trochoidea trochoides. Fig. 3 1 . X. conspurcata. Fig. 32 . Cochlicella conoidea. Fig. 33 . C. acuta

160

Alessandro Hallgass & Angelo Vannozzi

Figs. 34-39. Land shells from M ount Circeo. Fig. 34. Cemuella cisalpina. Fig. 35. Hygromid ciriCtella. Fig. 36. CllUoStOma

planospira. Fig. 37 . C. planospira-. protoconch with numerous tubercles. Fig. 38. Marmorana muralis . Fig. 39 . Marmorana

fuscolabiata circeja.

The continental molluscs of Mount Circeo (Latium, Italy)

161

and is distributed in the Italian peninsula (Umbria,

Latium , A bruzzo and Campania). Itis characteristic

of exposed walls however, it can also be found in

the woods, always on the rocky walls. The popula-

tions of Mount Circeo, not particularly abundant,

are of medium-small size for the species (h: 12-17

mm). It is morphologically similar to the southern

morphs of the species, i.e. whitish with smallpapil-

lae, and almost indistinguishable from the forms

present on Ausoni Mountains at Terracina. In the

nearby Lepini Mountains is dominant LdiCOStigma

leuco Stigma w ith purplish-brown shell and papillae

more evident. In Latium there are populations of

giant specimens, that exceed 22 mm.

Cochlodina incisa (K lister, 1 8 7 6 )

Species widely distributed in Italian peninsula at

lo w altitu des.AbovelOOO m it is replaced by C. lam-

inata (Montagu, 1803), a species widespread all over

Europe. In the contact areas the two species are found

in sym pa try. C. indsa from M ount Circeo (Figs. 20,

21) compared to the Apennine populations, shows a

mo re obese shell, som etim es w ith numerous in term e-

diate palatal plicae. As far as converns its anatom ical

features, no differences w ere observed. C. incisa w as

found, w ith the sam e character, even in the littoral ilex

trees wood of M acchiagrande (Rome).

This species is closely related to C. kuestevi

(Rossm assler, 1836) and C. meisneriana (Shuttle-

worth, 1 843) which are found in Sardinia and Cor-

sica, respectively.

Siciliaria paestana (Philippi, 1 8 3 6 )

A common species (Figs. 9, 25, 26) in the coastal

area of the Tyrrhenian coast, in alluvial plain be-

tween the ancient Pliocene coast line and the present

coast, from low lands behind the dunes to the inner

areas with low vegetation, never in very humid en-

vironments. The southern boundary for this species

is Paestum, its locus typicus. In the south and east

part of the species range (Campania and Basili-

cata), there are some forms of uncertain taxonomic

status.

Siciliaria gibbula honii (O. Boettger, 1 8 7 9 )

from the Vesuvian area to southern Latium . It is gen-

erally uncommon, abundant only on the Ventotene

island where it is the dominant species.

Nordisieck (2013) considers doubtful the sub-

specific relationship with S. gibblila. Further re-

search is needed to clarify the relationship of S. gib-

blila honii with some form of S. gibblila in Calabria,

the Aeolian Islands and with Siciliaria Vlilcanica

(Paulucci, 1878) from M ount Etna.

Papillifera bidens (Linnaeus, 1758)

Synonyms: Papillifera papillaris (O.F. Muller,

1 774). It is one of the species of Clausiliidae most

widely dispersed by man. It is common on old

walls. In Italy, P. bideUS lives in natural habitats

only in Apulia, Basilicata, Calabria, Sicily and Sar-

dinia. W ithin the MountCirceo it was found in the

locality known as "La Cava" and on the walls of the

town of San Felice (Figs. 23, 24) with ManflOrana

muralis (O.F. Muller, 1774 ).

F am ilia HY GROM IIDAE

Xerotricha apicina (Lamarck, 1 8 2 2 )

Species common (Fig. 29) in uncultivated lands

to the low lands behind the dunes. It is widespread

in Italy up to the Mediterranean coast of France.

The shell of the juvenile specimens has hairs on the

periostracum that do not persist in the adult.

Xerotricha conspurcata (Draparnaud, 1801)

This species is similar to X. apidna, however,

is smaller and with hairs of periostracum per-

sisting in adults (Fig. 31). It p refers areas less exposed

respect to the congener. Widespread throughout

Italy.

Hygromia cinctella (Draparnaud, 1801)

Species common in humid environments of the

Italian peninsula and Sicily. In the Mount Circeo

(Fig. 35) as found only one shell, not very fresh, in

the cork oak of Mezzo monte.

Poorly known species (Fig. 27), widespread Familia HELICIDAE

162

Alessandro Hallgass & Angelo Vannozzi

Chilostoma planospira (Lamarck, 1 8 22)

This species lives under brushes among the

rocks (Fig. 36). Anatomical examination of a living

specimen revealed an asymmetry in the morphol-

ogy of the two mucous glands, one being simple

and the other one bifid. This feature is not men-

tioned by any authors and could fall within the vari-

ability of the species. The protoconch is adorned

with numerous tubercles (Fig. 37).

Marmorana muralis (O .f. m mier, 1774 )

This species has a Sicilian distrib u tio n, but it has

been dispersed by man in many site in the western

Mediterranean area. In the Italian peninsula it is

known only in anthropic environments, with the ex-

ception of two localities in Umbria and Calabria. In

Sicily it is rather widespread e ith er in n atu ral or ru d -

eral environments. In Mount Circeo (Figs. 10, 38)

it was found on the walls of the town of San Felice

w ith Papillifera bidens.

Marmorana fuse olabiata circeja (K o b e it, 1 9 o 3 )

Marmorana fuscolabiata { Rossmasier, 1 842) is

widespread in the Southern Apennines. Geographi-

cal isolation of the population of Mount Circeo

(Figs. 11, 39) and the relative gene tic distance com-

pared to other neighboring population it seems to

justify the use of the rank of subspecies (Oliverio

et al., 1992). In recent years, probably because of

the drought, there has been a substantial decrease

in the number of individuals.

DISCUSSION AND CONCLUSIONS

Mount Circeo has long been an island; indeed a

sea bottom that does not exceed 300 m below the

sea level joins it to the Pontine Islands. This is a

group of volcanic islands, but still retains, in the is-

land of Zannone, a portion of the limestones that

possibly were part of a bigger platform.

The theory of Alvarez et al. (1 974) explains the

formation of Italy and of the western Mediterranean

by the counter clockwise rotation of the Sardo-

Corsican plate. Originally joined to the European

plate in correspondence of Provence, during its rota-

tional movement eastward, fragments detached to

form Balearic Islands, Cor sic a and Sardinia. Of great

importance is the C alab ro -P eloritan micro-plate that

detached from the whole S ardo -C orsic an complex

giving rise to the Tyrrhenian Sea. This theory has

allowed us to explain the fragmented distribution of

species with very low dispersal ability, including

Papillifera solida and the species of the genus So~

latopupa Pilsbry, 1917 as well as the close relation-

ship that occurs between Cochlodina kuesteri and C.

indsa (Giusti, 1 9 7 6 ; Ketmaier et al., 2006).

The distribution of Cochlodina Ferussac, 1821

species may be explained by the fragmentation of

the Alps occurred during the Oligocene. It is possible

that the distribution area of one species of Cochlo-

dina living with in the entire Alpine region was

fragmented and that the various isolated popula-

tions differentiated in different species over time.

In fact, we note that the center of diffusion of the

genus Cochlodina is the Alpine region, where there

are a dozen species, occurring - in addition to the

Alps and the Italian mainland - also in Corsica with

C. meisneriana , in S ard in ia w ith C. kuesteri , and in

Algeria (Kabilya) with C. bavayana Hagenm tiller,

1884 (Nordsieck, 1969), all fragments of the origi-

nal Alps, as well as it could have occurred in the

micro-plates that huddled to form the Apennines.

With the change of latitude the "Tyrrhenian"

Cochlodina adapted to warmer climates. It is there-

fore possible that the populations of C. laiTlinata th at

live at high altitudes in the Apennines are a relict

fauna that came down from the north in the colder

periods and, subsequently, C. indsa , com ing from the

west, colonized habitats at lower altitudes. However,

genetic studies are needed to test this hypothesis.

However, although the theory of Alvarez et al.

(1974) is now universally accepted, itleaves a large

margin of uncertainty as to how and when the var-

ious plates did move. D uerm eijeret et al. (1997) believe

that the separation between the S ardo-C orsican

complex and the C alabro-P eloritan microplate oc-

curred between 8.6 and 7.8 Mya, with the opening

of the Tyrrhenian Sea. The study on enzyme poly-

morphisms on the genus Marmorana ( Ambigua )

(O liv erio et al., 1992) partly confirms and p artly is in

contrast with this opinion. In fact, these times are well

suited for the separation between the TyrrheniberUS

ridens (M arte ns, 1884) from Sardinia and the Mar-

morana signata group (Ferussac, 1821), but do not

justify neither the much lower distance between

Tyrrheniberus and the Marmorana fuscolabiata

The continental molluscs of Mount Circeo (Latium, Italy)

163

group nor the proximity of this latter to MavniOVana

S axe tana (Paulucci, 1 8 8 6 ) occurring in the Argen-

tario. The genetic distance between TywhenibeVUS

and these latter two species would suggest a sepa-

ration occurred between 4 and 5 Mya.

A model that could justify these inconsistencies

would require the sep a ratio n, at different tim e s, of the

various plates travelling separately and subsequently

fused to form the current Italian peninsula. The

C alabro-Peloritan m icroplate at the tim e of separation

from the whole S aid o -C o rsic an complex might ap-

pear as an island very elongated in north-south direc-

tion that, for a certain tim e, traveled as a whole block.

During the movement from north-west to south-

east this microplate broke in several pieces, some

of which moved away to form the Argentario and

the calcareous parts of the Tuscan Archipelago, re-

spectively; this event could might have occurred

between 3.5 and 3 Mya.

In more recent times there w as a sliding move-

ment of the plate to the south (Van Dijk & Scheep-

ers, 1995) thatmighthave caused a new division of

the plate from which it broke away the part that we

find today as Mount Circeo and the limestone de-

bris of Zannone Island. This event may be dated

between 1 and 0.7 M a. This model could explain

the presence of MarmOranajliSCOlabiata on Mount

Circeo, the relatively close genetic distance be-

tween M. fuscolabiata and M. saxetana and the

greater genetic distance that separates these two

species from M. signata of Ausoni Mountains, less

than 15 km far from Mount Circeo, and could also

explain other b io g e o g r ap h ic mysteries such as the

presence of Pegea Camea (Risso, 1 826) in P an telle -

ria Island (Sparacio, 1997), the Aeolian Islands and

the Tuscan Archipelago and the presence of Pleu-

rodiscus balvnei at M ount C irceo. Only an additional

contribution of genetic data related to groups and

species strictly calciphilous with very low dispersal

abilities may corroborate or refuse this hypothesis.

ACKNOWLEDGEMENTS

We are grateful to Marco Bodon (Genoa, Italy) who

put at our disposal data of his samplings on the

promontory. We are also grateful to Aldo Marinelli

and M auro Grano (both Rome, Italy) for support in

the field and for the permission to publish the pho-

tos of figures 4 and 10, 11, 20 and 35,respectively.

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Biodiversity Journal, 2014, 5 (2): 165-174

Urban ecology: comparison of the effectiveness of five traps

commonly used to study the biodiversity of flying insects

Cedric Devigne 1,2 & Jean-Christophe De Biseau 3

'Universite de Lille nord de France, 59000 Lille, France

2 UCLILLE, FST - Laboratoire Ecologie & Biodiversite - Faculte des Sciences et Technologies / ICL, 41 me du Port, 59016 Lille

cedex, France

3 Universite Libre de Bruxelles, Service Evolution Biologique et Ecologie, CP 160/12, 50 av. F. Roosevelt, 1050 Bmssells, Belgium

"■Corresponding author, email: cedric.devigne@icl-lille.fr

ABSTRACT In this paper, we compare five different types of traps currently used in biodiversity studies to

collect flying insects. Our aim is to evaluate the potentials and the limits of these traps in the

assessment of insect biodiversity. Hence, we compared the diversity of insects caught by a

malaise trap, a yellow pan trap, a blue pan trap, a suction trap and a light trap in six different lo-

cations in Brussels. We showed that these traps caught nearly only insects: more than 98.3% of

all collected organisms were insects. Only the blue pan trap caught, in higher proportions, other

arthropods such as isopods or spiders. The Malaise trap was generally the most effective trap

capturing the majority of Homoptera, Heteroptera, Psocoptera, Diptera, Trichoptera and Hy-

menoptera. The yellow pan trap was often the second most effective trap particularly for Hy-

menoptera, Diptera and Homoptera. Without surprise, the light trap caught nearly all Lepidoptera

(Heterocera). Some combinations of two different traps were very effective. However, none of

these combinations were the most effective for all families of insects. Moreover, the combination

of the two most effective traps (Malaise and yellow pan traps) was not the best combination.

We discuss about the effectiveness of traps and the usefulness of their association. Finally, we

raise the particular case of urban environment which needs the use of discreet traps.

KEY WORDS Malaise trap; pan traps; suction trap; light trap; complementary traps; biodiversity.

Received 04.02.2014; accepted 23.04.2014; printed 30.06.2014

INTRODUCTION

Since the Earth summit in 1992, the conservation

of nature has taken more and more importance in the

world. The creation of an international day for bio-

logical diversity is a symbolic fact of the communi-

cation of the problem of the loss of biodiversity. At

the same time, many actions were developed to fend

off this trend. For example, the involvement to halt

the loss of biodiversity by 2010 shows that more

people feel concerned about the conservation of

natural heritage (Delbaere, 2004; EEA, 2007).

However, it is often difficult to explain why bio-

diversity is important and why we should be both-

ered with conserving it. Moreover, conservation of

biodiversity creates constraints for people who can

not see immediate outcomes. Some new political

decisions, aimed decreasing the discrepancy between

the knowledge of scientists and general understand-

ing of people, have been introduced (in France,

scientific foundation for biodiversity was launched

in 2008). It can be very difficult for a non-specialist

to understand that in order to make a success of the

166

Cedric Devigne & Jean-Christophe De Biseau

big challenge to halt the loss of biodiversity by

2010, the first step of these studies includes the re-

quirement to kill organisms. Indeed, to analyse bio-

diversity, scientists have to make inventories of

organisms in each locality. For numerous classes of

small animals (for example insects), the making of

such inventories implies that organisms have to be

killed. During such research, the killing of animals

is generally not specific and is indiscriminate thus

there is a risk that rare species may be destroyed.

Moreover, the traps designed to kill insects could

potentially kill other animals (Pendola & New,

2007). Fortunately, many species can be inventoried

without killing (generally the vertebrates, and some

insects like Orthoptera). However for the majority

of insects, death is unavoidable during capture. Be-

sides, insects are often used in ecological studies as

indicators species of biodiversity (Duelli et al., 1999;

Duelli & Obrist, 2003; EEA, 2007), of fragmenta-

tion or urbanisation of an enviromnent (Kremen et

al., 1993, Abensperg-Traun et al., 1996; Bolger et

al., 2000; Nelson, 2007). Consequently, biodiversity

studies are often confronted by this paradox:

to study biodiversity in order to improve our

knowledge, and thus to increase our abilities to pro-

tect and conserve it, specimens have to be killed. In

the extension of this paradox, some papers asked for

the development of ecological ethics or raised in-

teresting ideas about the ethics of killing organisms

for the purposes of scientific studies (Lockwood,

1987; Lockwood, 1988; Minteer & Collins, 2005a;

Minteer & Collins, 2005b).

Most studies concerning biodiversity need not

collect every species in a location, hideed, researchers

have developed several methods and strategies. In

this respect, different methods are available to esti-

mate species richness in an area: the use of a corre-

lation with determination level (Andersen, 1995;

Oliver & Beattie, 1996a; Oliver & Beattie, 1996b;

Andersen, 1997), the use of indicator species

(Rodriguez et al., 1998 but see McIntyre et al.,

2001; Kotze & Samways, 1999; Osborn et al.,

1999), the use of statistical methods to infer the species

richness from a sample (Colwell & Coddington,

1994). However, if the aim is to take inventory of

animals in a location then the observation and the

capture of at least one organism of each species is

necessary. Several ways are possible to limit the

death of insects. One of the ways is the use of

effective traps to limit the sampling frequencies. In

this respect, some studies were carried out to evaluate

the best trap design (Abensperg-Traun & Steven,

1995; Wang et al., 2001; Koivula et al., 2003; Pendola

& New, 2007), the best number of necessary traps

(Brose, 2002) or to compare effectiveness and the

complementarities of different traps (Lewis, 1959;

Obrist & Duelli, 1996; Duelli et al., 1999; Agosti et

al., 2000; Campbell & Hanula, 2007). However,

many of these studies focused on one or two species

and were not dedicated to global biodiversity esti-

mation (Brunner et al., 2007; Hossain et al., 2007;

Hardwick & Harens, 2007; Magina et al., 2007; Wu

et al., 2007; Blackmer et al., 2008). The studies

aiming at studying trapping methods in biodiversity

evaluation are marginal compared to the literature

about biodiversity generally. In this paper, we seek

to compare 5 traps commonly used in biodiversity

studies to collect flying insects in order to evaluate

their potential and their limits in the assessment of

insect biodiversity. In this paper, a trap was con-

sidered as most effective when it captures more

number of insects or number of families of insects.

Hence, it is attempted to define the effectiveness of

a trap as a function of their captures (abundance of

insects) and not from an economic point of view.

MATERIAL AND METHODS

The locations of trapping

The study was carried out in 6 locations in Brus-

sels. These sites were chosen according to their bio-

logical potential, it means their assumed probability

to have a high biodiversity. Three categories of bio-

logical potential were defined as a function of the

urban location of the site (if it is at the urban pe-

riphery or not), the management of the site (strong

human impact or not) and the previous estimation

and information given to us by the IBGE (Institut

Bruxellois pour la Gestion de l’Environnement - the

Brussels institute for the environmental manage-

ment). Consequently, 2 locations were supposed to

have a great biological potential and hence a high

biodiversity (the Massart botanic Garden, and the

Zavelenberg area), 2 locations, a mean biological

potential (the Tenbosch park and an abandoned pri-

vate garden at Simonis street) and 2 locations, a

poor biological potential (the highly maintained

Urban ecology: comparison of five traps commonly used to study the biodiversity of flying insects

167

garden of the Palais des academies and a very urban

private garden at Berceau street).

It is apparent that these sites are not directly

comparable. Indeed, the private gardens are very

small compared with the urban parks. However, the

aim of this study was to compare the effectiveness

of different kin ds of traps and not to compare bio-

diversity of different locations. Moreover, the use

of different urban green spaces should allow the

study to conclude about a potential generalisation

of the results.

Traps

In each site studied, 5 different traps, designed

to preferentially capture flying insects, were utilised

and compared:

- A suction trap was used (Fig. 1) for 30 minutes

in daylight (unfortunately, due to the noise of such

a trap, we were not able to use it during the night).

This trap consisted of a leaf blower/vacuum (PART-

NER-BV24 / nominal air flow = 0.142m 3 /s) direc-

ted toward the sky. A 2.5 metre high pipe (12 cm

diameter) was adapted in order to increase the

height of capture. A funnel with a collecting bottle

was inserted into the pipe to allow the collection

of insects.

- A Malaise trap was used for 24 hours. This was

a classic 2 metre high Malaise trap (S&S entrap net

company: http://www.geocities.com/ssentrap/).

- A light trap, put directly on the ground, for 7

hours during the night was utilised (Vermandel Ento-

mologie Speciaalzaak: http://www.vermandel.com/).

Attention was paid for the light trap so that it was

not impaired by artificial lighting.

- 2 coloured pan traps (15x12x5cm) were utilised:

one yellow and one blue were put directly on the

ground in each site for 48 hours. In these pans,

soapy water was used to kill insects.

This study was carried out during a hot, sunny

week from 3 to 6 of September 2002. The nights

were dark since the new moon was the 7th of

September 2002.

Hence, each location was sampled once with

eveiy trap. On each site, traps were used simulta-

neously but with different durations of working. In

this respect, it was possible to compare different

traps with a particular methodology.

Figure 1. Suction trap. This trap consisted of a leaf

blower/vacuum (1) (PARTNER-BV24) directed toward the

sky. A 2.5 metre high pipe (2) was adapted (12cm diameter)

in order to increase the height of capture. A funnel (made

with a piece of net (3)) with a collecting bottle (4) was in-

serted into the pipe to allow the collection of insects.

RESULTS

Global analysis

First of all, the results demonstrate large dif-

ferences in the effectiveness of the traps (Table 1).

If the effectiveness of a trap is considered as a func-

tion of the number of organisms caught, it can be

concluded that the Malaise trap is the most effective

trap. Indeed, this trap caught twice more animals

than the second most effective trap, the yellow pan

trap (Table 1). Nevertheless, the durations of work-

ing of traps were different. Hence, if the number of

capture per hour of working is observed, the suction

trap is by far the most effective trap (Table 1).

Secondly, regardless of the type of trap used, more

than 86% of organisms caught were insects (Table

1). Furthermore, most traps caught only insects, for

example Malaise traps seem specific for insects

(Table 1). However there are a higher number of

168

Cedric Devigne & Jean-Christophe De Biseau

other arthropods (mainly isopods and spiders) in

blue pan traps.

During the period of this study, 1746 specimens

were collected. Among these it was possible to

determine the families of 1597 specimens. Due to

deterioration or difficulty of determination, the re-

maining 149 specimens were identified as 15 Lepi-

doptera, 64 Trichoptera, 2 Diptera, 5 Hymenoptera,

1 8 Heteroptera, 44 Homoptera and 1 totally unde-

termined. This corresponded to a 5 to 13% of the

total insects collected in four traps. Only in the light

trap, 23% of the insects collected were assigned to

the order mainly due to the difficulty of determina-

tion of Lepidotera (Heterocera). Indeed, in this last

trap, many slugs were caught and their mucus dam-

aged the collected insects.

Henceforth, when the study refers to insect

order, the number of specimens considered is 1745

and when the study refers to insect families, the

number of specimens considered is 1597.

During the trapping period, 72 insect families

were collected and determined (Table 2). The re-

sults show that the Malaise trap caught more than

76% of the families collected (Table 2). The second

most effective trap is the yellow pan trap which

caught 61.1% of the families which is not signifi-

cantly different to the Malaise trap (Fisher exact

test, p= 0.072). The other traps caught less than

50% of the families determined which is signifi-

cantly different from the two other traps (Fisher

exact test, p<0.02). Both traps, Malaise and yellow

pan, caught together 87.5% of the families. Accord-

ing to our results, to capture 90% of the families the

use of three complementary traps would be neces-

sary: the combination Malaise + yellow pan + blue

pan traps (90.3%) or the combination Malaise +

yellow pan + suction traps (93.1%).

The Malaise trap also seemed to show more

specificity in captures since 12.5% of the families

were caught by this trap only (Table 2). In contrast,

less than 5% of the families were caught in each

other trap individually. However, due to the weak

level of specificity of the traps, no significant

statistical results was found (Fisher exact test,

p>0.07). More precisely, Diptera (Stratiomyidae,

Empididae, Tanipezidae, Pipunculidae, Lonchaeidae,

Sciomyzidae, Tephritidae), Hymenoptera (Halictidae)

and Psocoptera (Stenopsocidae) were only caught

by the Malaise trap (Table 3).

Generally, the Malaise trap is the most effective

for Homoptera, Heteroptera, Psocoptera, Diptera.

Malaise and yellow pan traps are equally the most

effective in the capture of Hymenoptera (Fig. 2).

For Coleoptera and Lepidoptera, the light trap was

the most effective. Due to the small number of

captures, the statistical analyses was possible only for

Homoptera, Hymenoptera, Diptera and Trichoptera.

There is a high significant statistical difference

between the traps for these 4 orders of insects (% 2

test, p< 0.005 for each order).

In the captures, diversity in families was very

small for Orthoptera, Thysanoptera and Homoptera

with only one family determined and for Psocoptera

with only 3 families. Hence, at the family level,

Type of traps

Number of insects caught

Number of non -insects

caught

Total number

of insects and

non -insects

caught

Proportion of

insects caught

(%)

Average

number of

insects caught

per hour

Light trap

129

131

98.5

18.43

Blue pan

104

86.5

1.88

Yellow pan

350

355

98.6

7.29

Malaise trap

939

940

99.9

39.13

Suction trap

238

246

96.7

476

Total

1746

1776

98.3

Table 1 . Number of individuals caught in the different traps used.

Urban Ecology: comparison of five traps commonly used to study the biodiversity of flying insects

169

Light trap

Blue pan Yellow pan

trap trap

i Malaise Suction

trap trap

Total number of

families caught

Proportion of families

caught by traps

26.4 % b

40.3 % b 61.1 % a

76.4 % a 36.1 % b

Proportion of families

caught by one trap only

4.2%

2.8% 2.8%

12.5% 4.2 %

Table 2. Specificity of each trap and effectiveness of traps that means proportion of families caught by the different traps.

Proportions with different letter were significantly different (p<0.05; Fisher exact test). Due to small number of families,

statistical test was impossible for the proportion of families caught by one trap only.

Diptera

Hymenoptera

Psocoptera

Coleoptera

Malaise traps

Stratiomyidae

Empididae

Tanipezidae

Pipunculidae

Lonchaeidae

Sciomyzidae

Tephritidae

Halictidae

Stenopsocidae

Yellow traps

Ceraphronidae

Curculionidae

Blue traps

Platypezidae

Hydrophilidae

Light traps

Chaoboridae

Nitidulidae

Smicrinidae

Suction traps

Scatopsidae

Torymidae

Scelionidae

Table 3. Specificity of capture. Insect families only caught in one kind of trap.

comparisons between traps are possible for

Coleoptera (9 families caught), Diptera (38 fami-

lies) and Hymenoptera (19 families).

Light traps caught a higher proportion of cole-

opteran families but it is not significant (Table 4).

Malaise traps caught the majority of Diptera and

Hymenoptera with 89.5% and 68.4% of families, re-

spectively (% 2 test; x 2= 40.7, df=4, p<0.0001 and

X 2 =18.7, df=4, p<0.001, respectively). For each of

these three groups, the yellow pan trap was the second

most effective (Table 4). Different combinations of

traps improved the captures. For example, the com-

bination light trap/yellow pan trap caught all the 9

families of Coleoptera together. For Diptera, the com-

bination light trap/Malaise trap or the combination

suction trap/Malaise trap caught 94.7% of families.

The combination of the two most effective traps:

yellow pan trap/Malaise trap was not the best since it

caught 89.5% of captures (difference was not signif-

icant). For Hymenoptera, the best combination was

suction trap/Malaise trap with a total of 94.7% of

captures; the second most effective combination was

suction trap/yellow pan trap with a total of 89.5% of

captures. Once more, the combination yellow pan

170

Cedric Devigne & Jean-Christophe De Biseau

trap/Malaise trap was not the best since with 84.2%

of catch that is the third combination (difference was

not significant). 100% of captures of Hymenoptera

families were obtained with the combination of three

traps: Malaise trap, suction trap and yellow pan trap.

Analysis of trapping constancy

The comparison of effectiveness of traps

between sites enables to check if the observations

were constant from one site to another, and hence

if some generalisations could be possible.

In proportions of insects caught during the trap-

ping period (Fig. 3), the Malaise trap is the most

effective for 4 out of 6 sites studied (x 2 goodness-

of-fit test; all p<0.001). For the two other sites, the

Yellow pan trap is the most effective (x 2 goodness-

of-fit test; all p<0.001). However, the results did not

show significant constancy in the proportion of in-

sects caught by the different traps between the sites

(heterogeneity x 2 analysis, x 2 = 236.5, df =20,

pO.OOOl).

In proportions of families, the Malaise trap is the

most effective since it caught a higher proportion of

100

o'- 70

3 60

■*-> 50

~ 40

O 30

10 ■

N = 15

□ Light trap

□ Blue pan trap

■ Yellow pan trap

■ Malaise trap

m Suction trap

★ ★★

N= 248

N = 3

Orthoptera Lepidoptera Coleoptera Thysanoptera Trichoptera Diptera Hymenoptera Psocoptera heteroptera Homoptera

Figure 2. Proportion of insects caught by the different traps. N=total number of insects caught in each order, yj goodness-

of-fit tests were earned out for Trichoptera, Diptera, Hymenoptera and Homoptera (the number of insects in the other orders

was too small to permit the analysis) to determine the highlight the differences between the traps. ** = p<0.005, *** = pO.OOl.

Light trap

Blue pan

trap

Yellow pan

trap

Malaise

trap

Suction

trap

Total number of

families caught

Coleoptera

55.6%

11.1%

44.4 %

33.3 %

11.1%

Diptera

26.3%

47.4%

60.5 %

89.5%

28.9 %

Hymenoptera

10.5%

36.8%

68.4%

68.4 %

57.9%

Table 4. Effectiveness of traps that means the proportion of families caught by each traps.

Urban ecology: comparison of five traps commonly used to study the biodiversity of flying insects

171

families in 5 out of 6 sites studied (Fig. 4). The yel-

low pan trap is the second most effective trap in

these 5 sites and the most effective in the sixth one.

In each location, the blue pan trap and light trap

were the least effective in the collecting of numerous

families of insects. In this respect, from one site to

another one, no differences were found between pro-

portion of families caught by each trap (heteroge-

neity x 2 analysis, x 2 = 9.6, df =20, p>0.05). Hence,

the results showed constancy in the proportion of

families caught by the traps in each location.

More precisely, concerning the constancy of cap-

tures, the results show that only Diptera were caught

in every site by each trap. Hymenoptera and Ho-

‘k'Jc'k

N = 624

"kkk

N = 190

•kick

N= 121

k'k'k

N = 339

□ Light trap

□ Blue pan trap

■ Yellow pan trap

■ Malaise trap

H Suction trap

Zavelenberg Massart

Simonis

Teenbosch Academie

Berceau

Figure 3. Proportion of insects caught by the different traps as a function of the trapping sites, x 2 goodness-of-fit tests

were carried out for each location to highlight the differences between the traps in each site. *** = p<0.001.

N = 33

Zavelenberg Massart

Simonis

Teenbosch Academie

Berceau

□ Light trap

□ Blue pan trap

■ Yellow pan trap

■ Malaise trap

□ Suction trap

Figure 4. Proportion of families caught by the different traps as a function of the trapping sites, x 2 goodness-of-fit tests were

carried out for each location to highlight the differences in between the traps in each site. NS = Not Significant ** = p<0.01,

*** =p<0.001.

172

Cedric Devigne & Jean-Christophe De Biseau

moptera were collected in every site by Malaise, yel-

low pan and suction traps. Coleoptera were caught

in 3 sites by Malaise, yellow pan and light traps. With

the exception of one specimen caught in the Malaise

trap and another in the blue pan trap, only light traps

caught Lepidoptera (Heterocera).

Malaise traps were particularly effective in col-

lecting some Diptera: Sciaridae, Phoridae, Muscidae

and Sphaeroceridae, and some Homoptera: Aphidi-

dae. Indeed, these 5 families were found in the malaise

trap in each location. In contrast, the other traps

showed less consistency. Indeed, only two families

(Hymenoptera: Pteromalidae and Mymaridae) were

found in each suction trap. Similarly, only one family

of Hymenoptera (Braconidae) and one family of

Diptera (Ceratopogonidae) were caught in every lo-

cation by yellow pan trap and light trap respectively.

DISCUSSION

In this study, the Malaise trap was most effective

in terms of the largest catch of individuals and fami-

lies when compared to the other traps. Moreover,

more than 1 0% of insect families were caught by the

Malaise trap only whereas the specificity of the other

traps reached at maximum of 4% of insect families.

Therefore, the better trap seems to be the Malaise

trap. However, this trap is big and highly visible

which could impede its use in urban locations be-

cause of potential vandalism acts. The same problem

occurs with the light trap which could improve the

trapping of nocturnal insects (for example Lepi-

doptera) but which is by definition visible because it

is illuminated. The suction trap is very effective since

it caught nearly 30% of Diptera and 60% of Hy-

menoptera while it was only used for 30 minutes. In

a way, it can be concluded that this trap is the most

effective since it captured a lot of insects in a very

short period. However, this trap, in its design, is very

noisy and thus could be difficult to use for a long

time in urban areas. In summary, in some circum-

stances (e.g. residential urban areas), researchers may

wish to choose the less effective traps as there is less

risk from vandalism or causing disturbance.

In this study, different traps were used with very

different durations of working. Indeed, the suction

trap was working only 30 minutes whereas the pan

traps were left on the ground during 48 hours. That

could be a problem in analysis of their comparison.

However, the comparison of different traps with

same duration would not be useful. For example if

the aim is to compare the effectiveness between the

Malaise trap and the suction trap, the use of Malaise

trap during 30 minutes would be useless, or in con-

trast, the use of suction trap during 24h would be

too damaging for the environment by depleting use-

lessly the abundance of insects. Hence, for further

studies it is suggested that it is better to compare

different traps used in their best methodology

(Campbell & Hanula, 2007; Hardwick & Harens,

2007; Pendola & New, 2007; Blackmer et al., 2008)

and not necessarily in the same methodology.

The results show that no trap alone is able to

catch all insect families that we have captured dur-

ing the trapping period. Indeed even if the suction

trap is really effective it will capture only small

airborne insects constituting the aerial plankton.

However, some combinations are potentially very

useful to improve trapping. Indeed, the combina-

tion suction trap/Malaise trap caught more than

94% of families of Diptera and Hymenoptera. The

combination of the two most effective traps did not

give better results, proving that same families

could be caught by these two traps. More studies

would be necessary to compare the numerous trap

systems (Southwood, 1978) and the effectiveness

of their association.

This study can be considered as very limited

since the determination was carried out only at a fam-

ily level. Hence, further studies are needed to verify

whether the same holds true at genus and species

level. However, the advantage of such broad gener-

alizations is that trends can be quickly identified

(Gaston & Williams, 1993; Andersen, 1995). Indeed,

the study seems to indicate that some families (which

could represent several hundred of species) are only

caught by one trap. For example, if Malaise traps

were not used in our samplings, more than 10% of

insect families captured during the study could not

be observed. Scientists and technical professionals

need to have standardised observation methods

(Agosti et al., 2000 but see Melbourne, 1999).

This standardisation should allow comparison

to be made between sites, at the same sites at dif-

ferent time periods and by different people. Indeed,

it must be kept in mind that results from one kin d

of habitat could be different from another. For

example, the results present some differences with

those from other studies: the captures by the blue

Urban ecology: comparison of five traps commonly used to study the biodiversity of flying insects

173

pan traps are very poor whereas these traps could

be highly effective in catching Hymenoptera polli-

nators (Campbell & Hanula, 2007). Therefore,

every site has specific attributes and the choice of

traps based on a range of features (e.g. trapping ef-

ficiency but also resistance to the deterioration)

could not be prone to “blind“ standardisation

methods. On the basis of the results of this study, it

would be extremely beneficial to continue studies

comparing the effectiveness of traps to help im-

prove monitoring of insect biodiversity.

ACKNOWLEDGEMENTS

We warmly thank Andy Thomson and Peter

Roche for revising the English manuscript. We also

thank two anonymous referees for helpful com-

ments on the manuscript. We also thank the Institut

Bruxellois pour la Gestion de l’Environnement

(IBGE-BIM) that have financed this project.

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Biodiversity Journal, 2014, 5 (2): 175-196

Lectotype designation and descriptions of two new subspecies

of Amphidromus (Syndromus laevus Muller, 1 774 (Gastropoda

Pulmonata Camaenidae)

Jeff Parsons

47 Elizabeth Street, Aitkenvale, Queensland, Australia 4814; e-mail: jefFonese@yahoo.com.au

ABSTRACT Amphidromus ( Syndromus ) laevus Muller 1774 (Gastropoda Camaenidae) was described

without a type locality. Sampling made in Indonesia over the last 20 years has confirmed the

presence of this species on Kisar Island (Pulau Kisar) and the Leti Islands (Kepulauan Leti)

of the southwestern Maluku Province. Similar shells have also been found at Tutuala, Timor

L’este. However, none of these specimens represents the nominal taxon and so its type locality

is still to be determined. Sampling made in recent years on Roma (Pulau Romang) has not

located any specimens of A. (5.) laevus romaensis Rolle, 1903. Herein a lectotype for A. ( S .)

laevus is designated and two new subspecies are described from the Leti Islands: A. ( S .) laevus

janetabbasae n. ssp. from Western Moa Island (Pulau Moa) and A. (S.) laevus nusleti n. ssp.

found on Leti Island (Pulau Leti).

KEY WORDS Amphidromus ; subspecies; lectotype; taxonomy.

Received 22.02.2014; accepted 27.04.2014; printed 30.06.2014

INTRODUCTION

It has been 240 years since Amphidromus ( Syn-

dromus ) laevus Muller, 1774 was first described,

and its type locality still has not been determined.

Previous authors usually stated imprecise localities

for specimens of A. (S.) laevus (sensu lato), such as

islands of Moluccas (Malaku Islands) (Fulton,

1 896). Prof, von Martens (1867) said he had obtained

this species while on Amboina (Ambon) from Mr.

Hoedt, and other collectors indicated to him they

had found it on the Tenimber (Tanimbar) Islands.

In 1877 von Martens stated that Captain Schulze

found it on Keffmg Island close to Ceram (Seram),

Moluccas. In addition to these imprecise or generic

localities, Laidlaw & Solem (1961) added Java,

Timor and Singapore; although they considered

Keffmg as a doubtful locality. No specimens of A.

(iS.) laevus (s.l.) have been found on Ambon, Seram

and the Tanimbar Islands in recent years, and here

considered as doubtful localities until von Martens’

specimens can be located and studied. This study

has determined that this species does not live on

both Java and Singapore, and so they are erroneous

localities.

Specimens of A. (S.) laevus (s.l.) deposited in

the Field Museum of Natural History (FMNH,

Chicago, Illinois, USA) are from field trips in

1998. This confirms that this species lives on the

islands of Leti, Moa and Lakor of the Leti Group

(Kepulauan Leti), and Kisar Island (Pulau Kisar),

southwestern Maluku Province (Provinsi Maluku)

of Indonesia (Sevems, 2006). Shells comparable to

this species, A. (S.) cf. laevus , live near the eastern

most point of Timor L'este at Tutuala, which is

close to Leti and Kisar Islands (collections of

176

Jeff Parsons

Figure 1. Map showing the approximate positions of the type localities for Amphidromus ( Syndromus ) laevus misled n. ssp.

on Leti (black star) and A. (A.) laevus janetabbasae n. ssp. on Moa (black diamond) (modified from a map of “Palau Moa”

including nearby islands: Army Map Service, Corps of Engineering (U.S. Army) 1963).

FMNH and JP). Field collections by John Abbas

confirm A. ( A .) laevus (s.l.) lives on Kisar (2008-

2012 ).

Misidentification of other species as A. ( A .) lae-

vus (s.l.) has certainly occurred and led to erroneous

locations. Misidentified species and shells labelled

as “Timor” in museums are still under investigation

and not discussed here.

In 2012, John Abbas organised field trips to

Leti and Western Moa (see map, Fig. 1). These

confirmed A. (A) laevus (s.l.) lives on Leti and dis-

covered a distinct population on Moa. Moa is the

largest of the three main islands in the western

part of the Leti group, separated by narrow chan-

nels from both Leti to the west (Moa Strait or

Selat Moa) and Lakor to the east (Lakor Strait or

Selat Lakor). A syntype of Helix laeva Muller,

1774 is selected as the lectotype of A. (A.) laevus

and a detailed description is given. Detailed de-

scriptions are also given for A. (A) laevus ro-

maensis Rolle, 1903 (lectotype) and A. (S.) laevus

kissuensis Rolle, 1903 (lectotype), in addition two

new subspecies are herein described as A. (A.) lae-

vus janetabbasae n. sp. from Moa and A. (A.) lae-

vus nusleti n. sp. from Leti.

MATERIAL AND METHODS

In the absence of preserved anatomical material

or living animals for study, the descriptions are

based on the morphological analyses of dry empty

shells. Shells available for the present study are

stored in the author’s private collection (including

a single shell from Tutuala) (JP) and supplied by

John Abbas (JA). Comparisons of some subspecies

could only be made using digital images, supplied

by museum staff, syntype of A. (A.) laevus , anony-

mous sources, A. (A.) cf. laevus from Tutuala and

Lakor, and from online access of museum collec-

tions (lectotypes of A. (A.) laevus romaensis and A.

(A.) laevus kissuensis). Muller’s measurement of

“lin.” is assumed as the obsolete Danish linie

(English line), which is 2.18 mm (Stover, 2001)

and his measurements are converted accordingly.

Relative sizes of shells for the subgenus Syndromus

mentioned: small <30 mm; medium 30-45 mm;

large > 45 mm.

Shell sculpture was examined under low magni-

fication (lOx) using a jeweller’s loupe. All shells

examined had formed a lip; those with a thickened

lip were determined as adult and those with a thin

Lectotype designation and descriptions of two new subspecies of Amphidromus (S.) laevus (Pulmonata Camaenidae) 177

lip as subadult. The digital vernier calliper used to

measure all shells has a resolution of 0.01 mm

(Table 1). Measurements for specimens in digital

images were calculated with the included scale bar.

The five shell dimensions measured are (Figs. 2, 3):

shell height (H), maximum shell width (D) and

aperture height (AH) (measured including the lip);

shell width above the aperture (d) and umbilical

width across the opening (U). The columella angle

(CA) was measured by placing a protractor over

digital images of shells, with the shell axis as the

zero degree point (Figs. 2-4). Number of whorls

(N), include the apex and the teleoconch and counted

precise to 0.125 (% whorl) as per Haniel (1921)

(Fig. 4). Measurement of the perch angle (PA) fol-

lows Dharma (2007). The ratios of shell height to

shell width (H/D) and aperture height to shell height

(AH/H) were calculated as indices of shell shape.

In order to make a comparison of banding pat-

terns in A. (5.) laevus, von Martens (1867) counted

only the dark bands on the last whorl and utilized a

similar numbering system to that used for studying

banding in pentataeniate (five-banded) Helicids

(e.g. Taylor, 1914; Cook & King, 1966; and Cook,

1967). Modifying his method for a pattern of six

dark bands: the count starts below the suture from

the highest of the superior bands (1 and 2), through

the peripheral bands (3 and 4) to the lowest or basal

bands (5 and 6). Each band present is given a num-

ber when present and a zero when absent. When all

six bands are present, this gives a formula of

123456. Numbers placed in rounded brackets show

two or more bands that have fused to form a wider

single band, e.g. 1234 (56). Two numbers under-

lined represent a band pair that has connected to

form a two-toned band, e.g. 123456. Indistinct and

partial bands are shown by a colon, e.g. 1 :3456. Haniel

(1921) used a slightly modified method by describ-

ing the pattern as seen on the last half whorl.

Two independent spiral band networks combine

with a band around the umbilicus (circumumbilical

band) to form the basic shell pattern as seen on the

last whorl. Each network has three band zones oc-

curring in alternate positions, with these named by

position starting from below the suture working an-

teriorly toward the umbilicus. The first network has

Figures 2, 3. Shell dimensions, see Material and Methods for explanation of abbreviations, and Figure 4: method of whorl

count (N) (Haniel, 1921)

178

Jeff Parsons

Measurement

Species

Amphidromus (Syndromus) laevus

Subpecies

laevus

(L)

cf. laevus

janetabbasae

kissuensis

nusleti

romaensis

(L)

Locality

unknown

Tutuala

Moa

Kisar

Leti

Roma

Count

(mm)

range

12.06-17.04

12.26-15.47

12.36-16.59

average

11.83

18.54

15.05

14.03

14.04

18.27

AH/H

(ratio)

range

0.36-0.47

0.38-0.44

0.41-0.47

average

0.45

0.42

0.43

0.41

0.44

0.41

(degrees)

range

5 to 20

6 to 18

5 to 20

average

10.56

11.47

10.11

range

5.625-7

5.25-6.875

5.375-6.5

average

5.50

6.5

6.25

5.888

(degrees)

range

16-25

18-24

average

19.72

21.67

21.68

(mm)

range

27.59-44.21

29.57-40.52

27.07-36.30

average

26.26

44.23

35.19

34.45

31.72

45.06

(mm)

range

15.35 19.69

14.93 20.41

14.77-21.26

average

14.69

21.78

17.73

17.68

16.89

22.95

(mm)

range

12.30-16.50

13.09-16.91

10.73-17.51

average

12.71

18.19

14.92

15.12

13.62

18.70

H/D

(ratio)

range

1.78-2.43

1.77-2.38

1.69-2.11

average

1.79

2.03

1.98

1.95

1.88

1.96

(mm)

range

0.66 1.84

0.47-1.10

0.54-1.34

average

0.71x1.48

1.20

0.93

0.98

Table 1. Comparative shell measurements for the subspecies of Amphidromus ( Syndromus ) laevus. Data given as aperture

height (AH); aperture height to shell height ratio (AH/H); columella angle (CA), offset to shell axis; whorl count (N),

including apex (to nearest % whorl) ;perch angle (PA); shell height (H); shell width (D); shell width above the aperture (d);

shell height to shell width ratio (H/D); and umbilical width (U). Note: L = lectotype; count is the number of specimens

measured; and the umbilicus of A. (5.) cf. laevus (s.s.) from Tutuala is elongated whorl count (N) (Haniel, 1921)

Lectotype designation and descriptions of two new subspecies of Amphidromus (S.) laevus (Pulmonata Camaenidae) 179

generally pale coloured bands found below the

suture (subsutural), middle of the upper half whorl

(supermedial) and middle of the lower half whorl

(submedial). The second network usually has dark

coloured bands between the subsutural and super-

medial bands (superior), near or on the periphery

(peripheral) and between the submedial and circum-

umbilical bands (basal).

According to the International Commission on

Zoological Nomenclature (ICZN) code (1999),

Article 72.4, Muller’s type series includes one spec-

imen (alpha) examined by him from the Spengler

collection (Museo Spengleriano) and a cited speci-

men (beta) described and illustrated by Lister

(1685). The colour variant beta described by Muller

was unavailable for morphological examination or

comparison through a photo.

A large part of Spengler ’s collection is de-

posited in the Zoological Museum, University of

Copenhagen, Copenhagen, Denmark (ZMUC).

Thanks to the help of Danny Eibye- Jacobsen, the

specimen examined by Muller has been located in

their museum. This shell was compared to the orig-

inal description (Muller, 1774) and opinions of

Chemnitz (1786), using photos supplied (ZMUC).

This shell matches the dimensions given by Muller

and the illustrations given by Martini (1777) and

Chemnitz (1786), and is here considered as the

actual shell seen by Muller. In absence of any

other typical material with a known locality, under

the terms of the ICZN code (1999), Article 74.7,

this shell is here designated as the lectotype and

discussed below.

Some taxonomic comments are required. Firstly,

there has been some confusion made recently that

the shells from Leti are a rediscovery of the nominal

taxon (John Abbas pers. comm.). Those particular

shells do not match Muller’s type shell and they are

herein described as a new subspecies. The nominal

taxon remains an unlocalised species, since the

labels found with the lectotype shed no light on the

type locality. Secondly, certain shells from Kisar

have been circulating among collectors incorrectly

labelled as A. ( S .) laevus romaensis Rolle, 1903

(John Abbas pers. comm.), which is a distinct form

of its own and discussed below.

ABBREVIATIONS. Type material of the herein

newly described subspecies of A. ( S .) laevus have

been deposited in the Australian Museum, Sydney,

New South Wales, Australia (AM); the Natural

History Museum, London, England, UK

(NHMUK); and Aquazoo/Lobbecke-Museum,

Diisseldorf, Germany (LMA). Some additional

type specimens belong to the private collection of

John Abbas (JA) and private collection of the author

(JP). Digital images of other type shells studied

are from Senckenberg Naturmuseum, Frankfurt,

Germany (SMF) and Zoological Museum, Univer-

sity of Copenhagen, Copenhagen, Denmark

(ZMUC).

SYSTEMATICS

Family CAMAENIDAE Pilsbry, 1895

Genus Amphidromus Albers, 1850

Subgenus Syndromus Pilsbry, 1900

Amphidromus ( Syndromus ) laevus laevus

Muller, 1774

Helix laeva Muller, 1774: pp. 95-96, No. 293 (not

illustrated)

Examined Material. Digital images of a single

syntype specimen from the ZMUC (Danny Eibye-

Jacobsen), ex. Spengler Collection, and this is herein

designated as the lectotype ZMUC-GAS-274 (Figs.

5, 6); dimensions: H 26.26 mm; D 14.69 mm; d

12.71 mm; and H/D of 1.79; type locality: unknown,

original labels are without locality data (Figs. 8, 9).

Description of the lectotype. Shell small

(26.26 mm high), sinistral, obliquely perforate (col-

umella hollow) and relatively solid. Shape ovate-

conic with a moderately elongated spire. Surface

is scarcely glossy; without macrosculpture. Micro-

sculpture (as determined from digital images):

protoconch (embryonic whorls) smooth; teleo-

conch (post-embryonic whorls) with numerous

very fine growth threads with microscopic growth

lines in the interspaces. Whorls about 514, distinctly

convex apically, next ones flatly convex and

lowest a little convex; base angularly- rounded.

Coiling is regular, the last whorl hardly descend-

ing in front. Suture weakly impressed apically,

shallow on the teleoconch with a faint white mar-

ginal line. Remnants of a pale cream periostracum

partially cover the last half whorl. Protoconch of

about 1 A whorls, bulbous; infrasutural band opaque

180

Jeff Parsons

cream, translucent pale flesh below. Apex (first

half whorl) obtusely pointed and a little exsert;

opaque cream (pale apical spot). Transition to the

teleoconch is weakly distinguished by a change in

whorl convexity and ground opacity. Teleoconch

pale flesh apically, following whorls dirty white

grading to white on the last.

Shell pattern is formed by a combination of

two independent spiral band networks, each with

three zones in alternate positions, and a circum-

umbilical band. A white subopaque stripe (mora)

about 1 mm wide, divides the bands clearly early

on the penultimate whorl; and ends in an opaque

greyish resting line, representing the former lip of

a resting stage (see Figs. 5, 6). Using Haniel’s

(1921) method of describing the pattern as seen on

the last half whorl, the band formula for this shell

is 023456.

Aperture oblique, semiovate and less than half

of the total shell length (0.45); very pale yellow

inside, clearly showing the external bands, but the

violet ones are stained brown. Parietal callus scarcely

perceptible, a thin colourless glaze; at the posterior

end of its margin, a very short thickened lump ad-

joins the termination of the outer lip (parieto-labral

tubercle), with nothing developed at the anterior

end. Outer lip (labrum) white, very thin, subre-

flexed and barely expanded, white margined within

the aperture. Columella white, narrow and thin-

walled; subvertical, angled away from the aperture

ventrally (abaperturally angled); twisted apically

before straightening to join with the basal margin

at an angle; weakly grooved at its root. Columellar

margin narrow, re volute, dilated above and taper-

ing to a narrow base, partially covering the narrow

umbilicus. Interior of umbilicus (umbilical inte-

rior) blocked in ventral view.

Animal and soft parts. Details unknown.

Distribution. Unknown.

Biology. Unknown.

Comparative notes. The A. (5.) contrarius

group from West Timor, which includes the nomi-

nate form and two accepted subspecies, share some

similarities with A. (A. ) laevus (s.s.). Each of the A.

(A) contrarius subspecies has pattern varieties that

are partially marked with continuous bands on the

spire or lower whorls, but usually the bands are in-

terrupted and indistinct or obsolescent.

Rarely some shells of A. ( S .) contrarius nikienesis

Rensch, 1931 develop banding like that of A. ( S .)

laevus (s.s.) (Fig. 18). Haniel (1921) indentified

such shells as being A. laevus ‘of the literature’.

These laevus-like shells have 3 to 6 purplish or

brown bands on the last whorl. The dark bands of

A. (A) laevus (s.s.) are reddish brown and violet

bands on the exterior surface, and brown on the

interior surface. Some shells of A. ( S .) contrarius

nikiensis have a similar disparity, with purple or

greenish external markings appearing brown inter-

nally (Figs. 16, 19). The overall shell shape, struc-

ture of the lip and columella, and lack of a dark

apex suggests A. ( S .) laevus (s.s.) has a close con-

nection to A. (A) contrarius nikiensis.

The shape and form of the columella is also

comparable to some species of the A. (A) incon-

stans group, e.g. A. (A) inconstans Fulton, 1898

and A. (A) wetaranus Haas, 1912. These species

have the root of the columella weakly impressed,

as does A. (5”.) laevus (s.s.). However, in A. (A) lae-

vus (s.s.) it is probably due to a lack of calcification

and not a distinctive feature of the species. A faint

yellow tint may occur on the columella and lip in

shells of A. (A) wetaranus. If the columella of A.

(A) laevus (s.s.) truly had a yellow tint when in-

spected by Muller, it has since faded to white.

Shells of A. (A) cf. laevus from Tutuala, Timor

L’este and laevus- like shells display another pat-

tern variation. Both commonly have solid dark

bands connected by alternating pale and dark blotches

(maculated zones) on the spire and solid or split

bands on the last whorl. In A. (A) cf. laevus these

maculated zones develop from bands with a paler

central zone (two-toned bands), which becomes in-

terrupted medially by pale coloured spots that fade-

away before the last whorl. In the laevus- like

shells, the maculated zones form by wide single

superior and peripheral bands becoming inter-

rupted medially by pale coloured spots, which are

later stained pinkish brown. The dark blotches

fade-away to leave a pinkish brown central zone

between each pair of purplish or brown bands,

creating two-coloured bands on the last whorl.

These maculated zones are absent in A. (A) laevus

(s.s.), which instead has a few faded blotches dis-

coloured to brown on the superior band; and the

two peripheral bands suffer pigment leakage and

become connected to form a two-toned band, pale

centrally. All three taxa show variation in the band

Lectotype designation and descriptions of two new subspecies of Amphidromus (S.) laevus (Pulmonata Camaenidae) 181

colour (tonal variation) creating intermittently

faded bands.

A. ( S .) laevus (s.s.) is here judged as a distinct

species. It can be separated from A. (S'.) contrarius

nikiensis by the following features: lip fused to the

body whorl; a minute parieto-labral tubercle at the

lip termination; continuous violet spiral bands; no

solid flammules; no vague flammules formed from

aligned dashes of interrupted bands; and nascent

reddish brown subsutural, supermedial and subme-

dial medial bands.

Remarks. The measurements of the shell as

positioned in the photos (Figs. 5, 6) give H 26.26

mm, D 14.69 mm and d 12.71 mm; and shell breadth

(B) is 13.42 mm (Fig. 5). Here H and B approxi-

mately fit Muller’s measurements of length 26.16

mm and breadth 13.08 mm, so it would seem

Muller measured this shell in a similar position to

that in figure 5. Original labels with the specimen

(Figs. 8, 9) refer to the first illustrations made of

it (Figs. 13, 14), with one label designating it as

the type shell (Sp.) and states Helix laeva a Mull.

(Fig. 9).

Muller described the colour of external surface

as lutescit, meaning coloured like white clay or

white with a muddy hue. Several times he described

the colour of the columella as lutea, assumed to

mean yellowish. The columella is now white with

some dirt inside, which blocks any view inside the

umbilicus to determine if it is white or tinged with

the colour of the circumumbilical band. Muller may

have been referring to this dirt giving the columella

a yellow tint at the time. He also described the shell

as having six brown bands, which must be referring

to the bands as seen inside the aperture. Figures 5

and 6 show a subreflexed lip, so it is probably a

subadult. The faint trace of a subsutural band is visible

behind the lip in the original digital images, but lost

in figure 6.

What is distinctive about this shell is the very

pale yellow palatal wall, which may have been

darker when collected in the 18th century. This

feature is absent in subadults of the other sub-

species of A. (S.) laevus. It is not peculiar to this

species and is discussed further below.

This shell was not figured by Muller and first fig-

ured by Martini in 1777 (Figs. 13), who wrongly

confused it with Helix inversa Muller, 1774 (A. (S.)

inversus). Chemnitz later figured it in 1786 but only

the dorsal view (Fig. 14). Three shells were located

in the ZMUC with a label identifies them as Or-

thostylus laevis (note “ laevis ” the typographical

error taken from Pilsbry, 1900). Another label ref-

erences the shell figured by Lister (1685) (Fig. 10),

thus suggesting they are Muller’s variation p. Pho-

tos taken by Tom Schiotte (ZMUC) show them to

be one adult and two juvenile shells indentified as

A. (S.) contrarius Muller, 1774. This is possibly a

case of the labels being mixed up with the wrong

specimens. This would mean that Muller’s variation

p is still to be found.

Muller noted a shell that Gualtieri figured in

1 742 (Fig. 1 1) is this species except for being dex-

tral. However, he did not mention the fact that in

the same volume he had named that particular

shell as Helix terebella (species number 319, p.

123). It is now accepted as Pyramidella terebel-

lum Muller, 1774 (Fig. 12), or as a synonym or

subspecies of P. dolabrata Linnaeus, 1758. Muller

(1774) placed the latter as species number 3 1 8 on

pages 121-122, accepting both as being terrestrial.

Muller’s comparison between the two species

must be due to the similarity in ground colour and

banding pattern. The obvious differences are P tere-

bellum has a columella with three folds, a simple

lip, a sharply attenuated spire and it is a marine

species, not terrestrial like it was assumed to be

at the time.

Chemnitz (1786) figured other shells that he

identified as A. ( S .) laevus (Tab. Ill, fig. 941-

948). His fig. 940 is Muller’s type shell (Fig. 14)

and shown marked with bluish (actually violet)

and brown bands, yet he repeats Muller’s descrip-

tion of it having six red brown bands. Yet for fig.

941 (Fig. 14), he clearly mentions that shell is

white with three reddish or bluish (possibly violet)

bands, which appears to mean reddish brown on

the spire and bluish on the last whorl. When com-

paring both shells, there is enough resemblance to

suggest they were possibly collected from the

same population. Unfortunately, this shell is yet to

be located and so was unavailable for this study.

Chemnitz’s fig. 949 is excluded because it is A.

(S.) furcillatus Mousson, 1849. It does appear that

subsequent authors (e.g. Reeve, 1849; von Martens,

1867 and 1877; and Pilsbry, 1900) have accepted

Chemnitz's account of A. (S.) laevus as accurate,

and have considered every shell similar in appear-

ance to those he figured and described as being the

same species.

182

Jeff Parsons

Amphidromus ( Syndromus ) laevus romaensis

Rolle, 1903

Type Material. Lectotype designated by Zilch,

1953 (Laidlaw & Solem, 1961; p. 654), currently in

the SM (Malakologie - SMF, 7574); ex. Sig. O. v.

Moellendorff collection (ex. H. Rolle); dimensions:

H 45.06 mm; D 22.95 mm; and H/D of 1.96; type

locality: Roma (Romang) Island, northeast of Kisar

and east of Wetar, Barat Daya Islands (Kepulauan

Barat Daya), Southwest Maluku Regency, Maluku

Province, Indonesia.

Description of the lectotype. Shell large

(45.06 mm high), sinistral, obliquely perforate and

very solid. Shape ovate-conic. Spire tall and regu-

larly tapered. Surface is glossy. Macrosculpture (un-

magnified): last whorl with occasional ridgelets,

more numerous towards the lip. Microsculpture (as

determined from digital images): protoconch

smooth; teleoconch with growth lines and very fine

growth threads. Whorls about 7, regularly increas-

ing in convexity; base rounded becoming angularly-

rounded behind the lip. Coiling is subregular with

the last whorl descending toward the lip. Suture

weakly impressed apically; shallow on the teleo-

conch with a thin white marginal line, more distinct

on the last whorl. Periostracum thin, pale tawny and

covers only the last whorl, thickening toward the

lip. Protoconch is opaque, dome-shaped, about VA

whorls; infrasutural band whitish; and pale yel-

lowish below fading away on the second whorl.

Apex opaque whitish (pale apical spot), obtusely

rounded and protruding. Transition to the teleo-

conch is distinguished by a change in whorl con-

vexity. Teleoconch pale yellowish apically,

following whorls white without gradation to a yel-

low last whorl.

Pattern combination as per the nominal sub-

species, but differs in band colouration and process

of modification. A single whitish mora about 1 mm

wide occurs close to the end of the penultimate

whorl, preceding a very thin and quite distinct

greyish resting line. The ground colour is rapidly

changed to yellow and banding modified after the

mora (post-mora modification) (see Fig. 21). The

banding pattern as seen on the last half whorl, gives

a formula of 023450.

Aperture oblique; strongly curved posteriorly,

semicircular and about 41% of the shell height; yel-

low deep inside and weakly showing the external

bands, white callused toward the lip. Parietal callus

very thin and colourless on the upper part; lower

third is white and flatly thickened toward a lump

beside the root of the columella (parieto-columellar

tubercle). Outer lip reflected, expanded and thick;

lower half with a vertically reflexed edge (rimmed).

Columella white, thick, rounded and wide; subver-

tical and abaperturally angled ventrally; oblique and

leaning outwards (proclined) laterally; forms a

distinct angle where it joins the basal margin (col-

umellar-labral junction); its base is gently curved

out (excurved) and projecting a little. Columellar

margin broad; a little dilated above and rolled over

the narrow umbilicus. Umbilical interior is indis-

cernible in the digital images.

Animal and soft parts. Details unknown.

Remarks. Laidlaw & Solem (1961: 654) con-

sidered it to be a variety of A. ( S .) laevus based on

the fact the distribution of the nominotypical form

was not known. However, it is regarded by the SMF

as being A. (S'.) laevus romaensis Rolle, 1903. The

shell is clearly marked as being the type shell indi-

cated by the letter “T.” inside the aperture along

with its catalogue number (Fig. 21 centre). The “a”

at the end of the catalogue number does indicate a

second specimen, however, Rolle (1903) mentioned

only one specimen and a search of the SMF data-

base (SeSam - Forschungsinstitut Senckenberg

2013) only located a single specimen. There is no

scale bar included in the digital images found on the

SMF database (Fig. 21) and no measurements given

either, so the measurements given by Rolle (1903)

are accepted as accurate. After taking measurements

from these digital images, it was concluded that

Rolle measured “D” parallel to the suture and “H”

perpendicular to that plane with the shell viewed

ventrally (Fig. 21 centre).

A number of field trips to Roma over the last

3 years have not confirmed the presence of this

subspecies (John Abbas, pers. comm. 2013).

Laidlaw & Solem (1961, p. 573; FMNH, specimen

CNHM 97362) mention another specimen; this

shell was unavailable for this study. A further search

on Roma is required to determine if it is still present

or ever was found on Roma, may be it was found

on a nearby satellite island instead of on Roma

itself. For now, due to its distinct appearance the

current status of this subspecies is accepted as

valid.

Lectotype designation and descriptions of two new subspecies of Amphidromus (S.) laevus (Pulmonata Camaenidae) 183

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Figures 5-7. Nominotypical subspecies A. (5.) laevus laevus : Figs. 5, 6. Lectotype, ZMUC-GAS-274 ex. Spengler Museum.

Figs. 8, 9. The original hand written tags with the lectotype, both citing the illustrations by Martini (1777) and Chemnitz

(1786), with (Fig. 9) identifying this specimen as the type shell, indicated by “Sp.”. Fig. 7. Closer view of theparieto-labral

node (circled) (Photos by Tom Schiotte, ZMUC). Fig. 10. Shell figured by Lister 1685 with text (t. 33, f. 31). Fig. 11. Shell

figured by Gualtieri (1742) with text (t. 4, f. M). Fig. 12. Pyramidella terebellum shown here for comparison (Maurice,

2013). Figs. 13, 14. Reproductions of original pre-1800 figures of Helix laeva. Fig. 13: lectotype by Martini (1777, Tab. I,

p. 416, figs. 8-9). Fig. 14: figures by Chemnitz (1786, Tab. Ill), (left) lectotype (Fig. 940) and (right) a shell very similar

to the lectotype, Chemnitz Collection (Fig. 941). Note: shells are not shown on the same scale.

184

Jeff Parsons

Distribution. Known only from Roma (Romang)

Island, northeast of Timor.

Biology. Unknown.

Amphidromus ( Syndromus ) laevus kissuensis

Rolle, 1903

Type Material. Lectotype designated by Zilch

1953 (Laidlaw & Solem, 1961; p. 633), currently in

the SM (Malalcologie - SMF, 7572), ex. Sig. O. v.

Moellendorff collection (ex. H. Rolle); dimensions

(see remarks): H 32.64 mm; D 17.77 mm and H/D

of 1.84; type locality: Kisar (Kissu) Island, north of

the eastern end of Timor Island, Barat Daya Islands

(Kepulauan Barat Daya), Southwest Maluku Re-

gency, Maluku Province, Indonesia.

Description of the lectotype. Shell medium

(32 mm high), sinistral, obliquely perforate and

quite solid. Shape distorted elliptic-pyramidal

with a moderately long and tapered spire. Surface

is shiny. Macrosculpture: lower teleoconch spo-

radically marked with growth threads. Microsculp-

ture (as determined from digital images):

protoconch almost smooth; teleoconch with nu-

merous growth lines and no discernible spiral mi-

crosculpture in the digital images. Whorls about

6 Vi, flatly to moderately convex on the spire; the

last is strongly convex opposite the aperture form-

ing a hump (laterally gibbose); base rounded to

somewhat flattened and sack-like behind the lip.

Coiling is irregular and distinctly distorted by the

last whorl’s gibbosity and descent toward the lip.

Suture shallow, somewhat impressed apically and

marked with a thin dull off-white edge. Perios-

tracum absent, worn off or removed (see remarks

below). Protoconch is dome-shaped with about

U /2 whorls; sub translucent pinkish brown. Apex

obtuse, slightly exsert; black (dark apical spot),

extending as an evanescent wedge and same-

coloured infrasutural band. Transition to the teleo-

conch weakly distinguished by a change in whorl

convexity and ground opacity. Teleoconch early

whorls stained pinkish brown between the brown

spiral bands and whitish above; the brown fades-

away on the fourth whorl; remaining whorls dirty

white.

Pattern combination as per the nominal sub-

species, except has different coloured bands. A pale

translucent grey weak mora occurs late on the an-

tepenultimate whorl; about 1 mm wide and divides

the bands cleanly (see Fig. 22). The band formula

for the lectotype as seen on the last half whorl is

12(34)56.

Aperture is oblique, semicircular and about

45% of the shell height. Palatal wall colourless and

pellucid, very clearly showing the external bands.

Parietal callus is pale straw yellow, very thin and

transparent; a small flattened parieto-columellar

tubercle present on the parietal callus margin be-

side the root of the columella. Outer lip white,

thickened, narrowly expanded and strongly re-

flected; external edge rimmed, distinctly so on the

basal margin; raised a little above the suture at its

termination (ascending termination). Columella

white, thickened and wide; straight and oblique,

abaperturally angled ventrally and proclined lat-

erally; angular at the columellar-labral junction

with its base projecting beyond it (extorted); weakly

impressed at its root. Columellar margin wide; di-

lated above and tapering toward its base; curled

over the narrow umbilicus, with its anterior edge

distinctly recurved above it. Umbilical interior

shows the ground colour of the last whorl, not

stained by the circumumbilical band and clearly

separate from it.

Animal and soft parts. Details unknown.

Variability. The spire is short to long with a

tapered to somewhat turreted (subturreted ) profile

and the surface may be dull. Protoconch is almost

smooth, occasionally with a few microscopic

growth threads (microthreads). Teleoconch has a

macrosculpture of growth threads throughout, or

admixed with ridgelets on the lower whorls; and a

microsculpture of numerous microscopic spiral

striae (microstriae) overlain by very fine growth

lines and microthreads apically, and coarser growth

lines elsewhere. Generally, only one mora is present

on the penultimate whorl, but commonly a second

one may be present on the antepenultimate or pre-

vious whorl; and rarely absent or more than two

present. The umbilicus is rounded (0.47—1.10 mm

wide) or rarely elongated (0.51 x 0.97 to 0.56 x

1.22 mm).

Commonly shells have distorted lower whorls

caused by irregular variation in coiling angle and

whorl expansion. Distortion either affects just one

part of the last whorl like in the lectotype (laterally

Lectotype designation and descriptions of two new subspecies of Amphidromus (S.) laevus (Pulmonata Camaenidae) 185

Figures 15-20. A. ( S .) contrarius nikiensis from the NikiNiki area, West Timor showing variation of ground colour, pattern,

interior, size and shape (JP). Fig. 15: brown flammulated form found closer to the coast and (Figs. 16 to 20) all found close

to NikiNiki. Fig. 18: Haniel’s A. ( S .) laevus 1 ' of the literature’. All shells except figure 17 have their periostracum still intact.

Note: natural size of shells. Figure 2 1 . Lectotype of A. ( S .) laevus romaensis ; SMF, Malalcologie - SMF, 7574a (a composite

of photos 19678- 19680, SeSam - Forschungsinstitut Senckenberg 2013). Figure 22. Lectotype of A. (5.) laevus kissuensis;

SMF, Malalcologie - SMF, 7572 (a composite of photos 19675-19677, SeSam - Forschungsinstitut Senckenberg 2013).

186

Jeff Parsons

gibbose), or affects both lower whorls (unequally

gibbose). Coiling may also be subregular with the

last whorl descending toward the lip and little or no

distortion of the whorls. The dark apical spot varies

as follows: apex to most of the first whorl darkly

stained with black fading to brown or reddish pur-

ple at its edge; extended as per the lectotype to form

a dark apical swirl, fading out toward or on the sec-

ond whorl, or not extended.

Two distinct colour morphs occur, pallibicinc-

tate and atrifasciate, with the appearance of the dark

apical spot and protoconch colour varying between

the morphs. Pallibicinctate refers to the two pale

coloured bands generally present, one each encircling

above and below the periphery. Since these shells

lack the dark bands, the band formula is 000000.

Atrifasciate refers to shells exhibiting dark coloured

bands.

The umbilical interior is clearly separate from

and never stained the circumumbilical band when

present, and shows only the shell’s ground colour.

Rarely the teleoconch has a few scattered translu-

cent grey flecks present. A. (S'.) laevus kissuensis

differs from the other subspecies by commonly

having a third overlying band network present con-

sisting of short to long segments of lime green lines

superimposed upon the other bands and their inter-

spaces. They appear in both colour morphs only on

the last whorl, variable in length, and occasionally

very slightly elevated above the shell’s surface as

spiral ribbons.

Distribution. Known only from Kisar, north of

Timor.

Biology. Unknown.

Remarks. According to Laidlaw & Solem

(1961, p. 633) the lectotype was first figured by

Zilch (1953, pi. 22, fig. 10). They did not state any

measurements of this specimen and incorrectly

referred to it as the holotype (p. 633). There is no a

scale bar included in the digital images (Fig. 22) in

the SMF database (SeSam - Forschungsinstitut

Senckenberg, 2013). The SMF classification for this

taxon is accepted as A. (S.) laevus kissuensis Rolle,

1903. The shell is clearly marked as being the “type

shell” indicated by the letter “T.” inside the aperture

along with its catalogue number (Fig. 22 centre).

The “a” at the end of the catalogue number indi-

cates a second specimen, and according to Rolle

(1903) there were two syntypes.

The lectotype is a stocky, primarily white shell

with brown bands and two very faded, subobsolete

yellow supermedial and submedial bands and the

subsutural band is absent. Yet Rolle mentioned

black, brown and yellow spiral bands, which sug-

gests the second shell had black bands with more

distinct yellow ones. Based on this and after taking

measurements from the digital image of the lecto-

type viewed ventrally (fig. 8 centre), this shell is

most likely Rolle’s larger syntype, since the calcu-

lated measurements closely match his: H 32 mm

and D 17 mm. Rolle’s second syntype has the di-

mensions of H 30 mm and D 16.2 mm.

Amphidromus ( Syndromus ) laevus janetabbasae

n. ssp.

Type Material. Number of shells examined:

total 25; adult 22 and 3 subadult (very thinly re-

flected lips); Holotype: AM C.483433; Paratypes:

LMA, (LMD/LOB 133653a-b) (2 shells); AM

C.483434 (5 shells); NHMUK 20120339 (4 shells);

JAC (2 shells); JP (11 shells); dimensions: H 33.91

mm; D 17.21 mm; and H/D of 1.97; type locality:

Moa Island, north-northeast of Timor Island, South-

west Maluku Regency, Malaku Province, Indonesia;

found in villagers’ plantations and the forest border-

ing them, on the eastern side of the road between

the foothills to the north of Wakarlili on the south-

west coast (Fig. 1).

Other material. JA: from the type locality; 49

adult shells.

Description of the holotype. Shell medium

(33.91 mm high), sinistral, obliquely perforate and

moderately solid. Shape ovate-conic with a moder-

ately long spire, profile somewhat flattened; H/D

ratio of 1.97. Surface is glossy; protoconch smooth-

ish; teleoconch macrosculpture of growth threads on

the lower whorls and occasional ridgelets on the last

whorl; and teleoconch microsculpture of numerous

spiral microstriae overlain by thin growth lines

throughout and occasional microthreads apically.

Whorls 6%, slightly or moderately convex; a small

section is a little swollen (subgibbose) just after the

mora; base angularly rounded. Coiling is subregular,

faintly distorting the lower whorls; last whorl de-

scending toward the lip. Suture impressed apically,

shallow below; bordered by a faint to bold white

Lectotype designation and descriptions of two new subspecies of Amphidromus (S.) laevus (Pulmonata Camaenidae) 187

marginal line. Periostracum removed in the holotype

(see remarks below). Protoconch is rotund with

about IV 2 whorls; opaque cream infrasutural band is

faint; ground subtranslucent pink; apex blunt, a little

exsert. Apical spot pale, not blackened; whole apex

opaque whitish. Transition to the teleoconch weakly

distinguished by several microthreads.

Teleoconch ground stained pink apically, fad-

ing-away on the third whorl; remaining whorls

whitish grading to yellow on the last; variously

marked with pale, medium or dark coloured spiral

bands, emerging on the second and third whorls form-

ing a pattern combination as per the nominal sub-

species, except more vividly coloured. A single

translucent pale grey mora is present roughly mid-

penultimate whorl; 1 mm wide, each side bordered

by an opaque greyish resting line; it marks a rapid

change in band colour and appearance (see Fig. 23).

The holotype has the band formula of 123456.

Aperture is oblique with a perch angle of 19°;

subovate and anteriorly subangular; AH/H ratio of

0.48. Umbilical interior white; moderately calcified

and translucent, clearly showing the dark external

bands. Parietal callus colourless; faintly calcified

and imperceptible; a minute parieto-columellar

tubercle present at the junction with the columella,

fading as a curved trace along the parietal callus

margin; and a minute parieto-labral tubercle adjoins

the outer lip’s termination. Outer lip has a white

face and faded pale yellow inner margin with faded

pale purple stains at both ends; thin, strongly re-

flected and narrowly expanded; face and external

edge flat; posterior termination slightly ascending.

Columella white; thickened, narrow and straight;

oblique and abaperturally angled (9°) ventrally;

subvertical and proclined laterally; angular at the

columellar-labral junction with a slightly excurved

base. Columellar margin white; curled over the um-

bilicus, partially covering it; face convex; cylindri-

cally dilated, its base tapered. Umbilicus is round

and narrow, 0.93 mm wide. Inner umbilicus tinged

by the circumumbilical band.

Animal and soft parts. Unknown as all speci-

mens were collected by locals for Mr. John Abbas

and received as empty shells.

Variability. Shells are variable in shape, com-

monly ovate-conic, but also oblong (like that of A.

( S .) latestrigatus Schepman, 1892) to elongate-

tapering when distorted.

This species is very variable in the morphology

of the aperture, palatal wall, outer lip, columella,

columellar margin and base. The umbilical interior

is very rarely yellow.

A well-developed parieto-columellar tubercle is

generally a trace line or absent; uncommonly occurs

as a thin curved line of callus in mature shells; and

rarely a short smudge at the root of the columella.

A parieto-labral tubercle adjoined to the lip

termination is rarely present, either a minute lump

(as in the holotype) or an elongate lump a few mil-

limetres long. The umbilicus is rounded and large

for the group (0.66-1 .84 mm wide) and rarely elon-

gated (0.92 x 1.14 to 0. 53 x 1.14 mm).

Faintly distorted subgibbose shells have less

pronounced swelling of the lower whorls than A.

(A) laevus kissuensis. However, the subregular

coiling in these shells causes a steeper descent angle

of the last whorl, creating an appearance of greater

distortion. Rarely true longitudinal distortion does

occur (2 shells) where irregular coiling elongates

the last whorl positioning it well below the pe-

riphery, thus exposing more of the previous whorl

than usual. Coiling may also be regular or subgib-

bose without distorted whorls. Atrifasciate shells

are the predominant form on Moa, with pallibicinc-

tate shells being rare. Both morphs have a similar

disparity in the colouration of the early whorls seen

in A. (A) laevus kissuensis.

In atrifasciate shells, the supermedial and sub-

medial bands may brown, blacken or redden to-

ward a mora and/or the lip, and the subsutural band

rarely develops a brown tinge. The most constant

band formulas are 020450 and 023450. All six

bands are commonly present and rarely are five or

all six partially absent on the last whorl, usually

ventrally. The less common band formula of

000450 is comparable with two variations of A. (A)

contrarius Muller, 1774. The pure form of 000450

with bands 1 to 3 absent on the spire is quite rare.

Commonly shells have other one or more bands on

the spire that are lost randomly or after a mora and

do not reappear on the last whorl. A. (5”.) contrarius

var. suspectus von Martens, 1864 displays pome-

granate supermedial and submedial bands and the

dark band formula of 000450, and rarely have

bands 1 to 3 present on the uppermost whorls. A.

(A) contrarius var. albolabiatus Fulton, 1896 has

only the pure form of 000450 and lacks the first

network bands.

188

Jeff Parsons

The external appearance of the lower whorls

shows a wider range of shell colours created by the

combination of periostracum, band and ground

colour. The colour of the protoconch and first net-

work bands is very variable. A variable dark apical

spot is present or absent. A mora either marks a hiatus

in the banding without modification of colour

and/or pattern, or marks a rapid change. Additional

shell pattern elements consist of dilution streaks

and/or shadowy deepening streaks, usually on the

last whorl.

Distribution. Based on current material avail-

able, this subspecies is currently known only from

type locality; it appears to be restricted to Moa, pos-

sibly found in the same vegetation type over the

whole of the island.

Biology. Found on the leaves, limbs and

branches of small trees and trunks of larger trees in

deciduous broadleaf forest and villagers’ plantations.

Etymology. Named in honour of Mr. John

Abbas’ daughter Janet.

Remarks. The holotype’s minute parieto-labral

tubercle is comparable to that of the nominal sub-

species. Its labral inner margin stains are due to pig-

ment leaked from the outer surface ground and

bands. Its lip is thinner than average, but equal in

thickness to a well-thickened lip seen in A. ( S .) con-

trarius (s.s.). The periostracum when present covers

only the lower whorls: thin, pellucid, dull; pale yel-

low to ochre; occasionally with darker or paler

streaks toward the lip. These streaks tend to appear

above the dilution or deepening streaks on the shell,

which dilute or deepen the ground and pattern colour

respectively. A. ( S .) kruijti R et F. Sarasin, 1899 has

similar periostracal streaks. The parietal callus is

colourless in fresh specimens, but becomes whitish

and deteriorates in older empty collected shells.

The pale purple inner labral stains are compara-

ble to those of other species. A. (S.) kuehni Moel-

lendorff, 1902 has ruddy anterior and posterior

stains (prominent externally, faded internally), and

any faint yellow staining is from an external

preapertural band showing through. In A. (S.) annae

von Martens, 1891 the entire outer margin is red-

dish purple (claret when dark, or magenta when

pale), across at least the lower lip face and some-

times faintly along the inner margin, with the deeper

tones shining through to the outside. Shells are

excessively variable in colouring and banding, but

commonly like that of the holotype. Compared to

A. (S'.) laevus kissuensis the supermedial and sub-

medial bands are never obsolescent on the spire, but

may appear faintly darker than the yellow or pale

saffron ground of the last whorl. Inner wall is the

preferred term to describe the exterior surface of the

previous whorl inside the aperture.

The translucent flecks are generally greyish, rarely

brownish, and sometimes have a same-coloured

shadow. These particular flecks are comparable to

a translucent brownish fleck and creamy shadow on

A. (S.) cf. laevus from Tutuala; and the comet like

markings of a black dot and yellow tail on A. (S.)

coeruleus Clench et Archer, 1932.

Amphidromus ( Syndromus ) laevus nusleti n. ssp.

Type Material. Number of shells examined:

total 15 (adult); Holotype: AM C. 483435;

Paratypes: LMA, LMD/LOB 133654a-b (2 shells);

AM C. 483436 (2 shells); NHMUK 20130070 (2

shells); JAC (2 shells); JP (6 shells); dimensions: H

32.33 mm; D 17.62 mm; and H/D of 1.83; type

locality: Leti Island, north-northeast of Timor Is-

land, Southwest Maluku Regency, Malaku Prov-

ince, Indonesia; found in forest on the hills to the

south of Sewaru village, which is on the northern

coastline of the island near Cape Tutukei (Fig. 1).

Other material. JP: 4 adult shells; 2 from the

type locality and 2 from forest behind the beach

near Cape Tutukei and Sewaru village. JA: from the

type locality; 1) 27 adult and 2 juvenile non-atrifas-

ciate shells; 2) 17 adult atrifasciate shells; 3) 6 adult

mid-banded shells

Description of the holotype. Shell medium

sized (32.33 mm high), sinistral, obliquely perforate

and heavily calcified. Shape ovate-conic; H/D ratio

of 1.83. Spire moderately long and subturreted. Sur-

face glossy; protoconch smoothish; teleoconch

macrosculpture of growth threads on the lower

whorls, and the last whorl also has numerous ridgelets

and a varix and teleoconch microsculpture of crow-

ded spiral microstriae crossed by growth lines and

microthreads. Whorls 6; convex and gradually ex-

panding; base angularly rounded becoming flat-

tened behind the lip. Coiling is subregular; last

whorl descends toward the varix and horizontal there-

Lectotype designation and descriptions of two new subspecies of Amphidromus (S.) laevus (Pulmonata Camaenidae) 189

after. Suture impressed apically and shallow on fol-

lowing whorls; bordered by a white marginal line.

Periostracum removed in the holotype (see remarks

below). Protoconch dome-shaped with steep sides,

1 1/2 whorls; ground translucent pink; apex subangu-

lar, obtusely pointed and a little exsert. Apical spot

dark; most of the first whorl stained black, be-

coming reddish purple marginally; extended along

the sutural margin as a thin dark apical swirl, fading

out on the second whorl. Transition to the teleo-

conch weakly demarcated by several microthreads.

Teleoconch ground stained pink apically; second

whorl whitened; fourth whorl stained very pale

straw yellow, grading to saffron toward the varix

and faded after it.

Pattern of spiral bands formed as per the nomi-

nal subspecies, but differently coloured and modi-

fied (see Fig. 24). The holotype has two incomplete

bands and so a formula of 0::450.

Aperture is oblique, subovate and posteriorly

rounded; perch angle 24° and AH/H ratio 0.49.

Aperture interior thickly calcified, subopaque;

whitish, a little glossy; external bands weakly vis-

ible, unmodified in colour. Parietal callus colour-

less; faintly calcified and inconspicuous;

parieto-columellar tubercle is a small lump adjoined

to the end of the columella margin’s flange; pari-

eto-labral tubercle is elongate, about 1 mm long.

Outer lip white; moderately thickened, strongly re-

flected and moderately expanded; face flat; lower

half of outer edge rimmed; posterior termination

ascending a little. Columella white, thick and

broad, a little twisted; subvertical and abapertu-

rally angled (5°) ventrally; oblique and proclined

laterally; angular at the columellar-labral junction,

its base extorted. Columellar margin is white; jut-

ted over the umbilicus, partially covering it; face

convex and well thickened, forming a flange ex-

tending to its insertion point; cylindrically dilated

and its base obliquely truncate with a subangular

jut. Umbilicus is round and narrow, 1 .08 mm wide.

Umbilical interior partially tinged by the circum-

umbilical band.

Variability. Whorls are flattened to convex;

sometimes the lower whorls are weakly to moder-

ately swollen (subgibbose to gibbose), or just the

last whorl is ventricose. Coiling is regular; subreg-

ular with the last whorl descending toward the lip;

or irregular with the lower whorls unequally gib-

bose and faintly to distinctly distorted. Spire short

to long with a somewhat flattened to subturreted

profile. Sculpture as per the holotype or a little

rough locally, with occasional ridges or growth

welts commonly on the last whorl, sometimes on

the penultimate whorl and rarely on the upper

whorls. The umbilicus is round, narrow to moder-

ately wide (0.54-1.34 mm) and rarely elongate to

rimate (0.65 x 1.18 to 0.35 x 1.10 mm). Umbilical

interior always tinged by the circumumbilical band.

Sometimes a single whorl forms the protoconch.

Dark apical spot is large as per the holotype; or

small with only the tip to whole apex stained black,

extending outward a little while fading to brown or

reddish purple at its edge.

Additional shell pattern elements consist of di-

lution or deepening streaks, sometimes the latter

are browned or blackened on the last half whorl;

and the teleoconch is rarely marked with translu-

cent grey or brownish flecks. Resting lines are gen-

erally a growth stria, but sometimes form a

ridgelet or swell up into a wide ridge formed from

a former lip (true varix). On Leti, pallibicinctate

shells are the predominant form, atrifasciate shells

are uncommon and mid-banded shells are rare.

There is a greater disparity in the colouration and

form of the pallibicinctate and atrifasciate shells

than that seen in A. (S.) laevus janetabbasae n.

ssp. and A. ( S .) laevus kissuensis. All three of

these colour morphs may have the first network

bands browned or blackened toward a mora

and/or the lip, rarely reddened beforehand. The

supermedial and submedial bands often appear

faintly darker than or fade into the yellow or saf-

fron ground of the last whorl.

A few shells have 6, 7 or 8 dark bands, but

extra bands inserted into the pattern created these

combinations. Only a single shell has the 000450

pattern with bands 2 and 3 on the upper whorls, a pat-

tern more commonly seen in A. (S.) laevus janetab-

basae n. ssp. Counting extra dark bands inserted

into the pattern, a maximum number of 12 and

minimum of 4 on the last whorl (subsutural, su-

permedial, submedial, and circumumbilical) bands

present.

Animal and soft parts. Unknown as all speci-

mens were collected by locals for Mr. John Abbas

and received as empty shells.

Distribution. Based on current material avail-

able, this subspecies appears to be restricted to Leti

190

Jeff Parsons

and probably found in the same vegetation type

over the whole of the island, including behind the

beaches.

Biology. Found on the leaves, limbs and branches

of small trees and trunks of larger trees in deciduous

broadleaf forest and vine thickets.

Etymology. The subspecies epithet is derived

from Nusleti, a historical Letinese name of Leti Is-

land (van Engelenhoven, 1997) and used here as a

noun in apposition.

Remarks. The holotype represents the average

conchological features of the subspecies, and the

average banding modification and pattern seen in

the atrifasciate shells. It also has a true varix that

occasionally develops in this subspecies. Its slate-

coloured dark bands are not present in the other

populations. Its parieto-labral tubercle is at full de-

velopment for the subspecies. It has the following

similarities with A. (5.) laevus janetabbasae n. ssp.:

columella and its margin are similarly formed;

parieto-labral tubercle is equal at full development;

and aperture is similar in shape. However, A. ( S .)

laevus nusleti n. ssp. tends to have an angular jut

at the base of the columellar margin; parieto-col-

umellar tubercle is a small lump or short ridge and

never fades as a curved trace or forms a long ridge;

and outer lip is opaque white with a flat or rimmed

edge. Periostracum covers only the lower whorls

when present: thin, pellucid, dull; and pale yellow

to citron-ochre; sometimes with darker periostracal

streaks towards the lip as in A. (S'.) laevus janetab-

basae n. ssp.

Pallibicinctate shells may have no translucent

grey morae present, or commonly have one or two

present, 0.5-3 mm marked by an opaque resting

line, and sometimes followed by a narrow whitish

or yellowish post-marginal band. Occasionally

wider multiple zones occur, composed of multiple

very thin morae. One non-type shell has several

deepening streaks stained with a reddish brown hue

on the last half whorl. Overall these more or less

yellow shells are comparable to A. (S.) contrarius

var. subconcolor von Martens 1867, a nearly uni-

form yellowish white shell with two very faint yel-

low bands encircling the last whorl. However, it

differs in the last whorl being darker yellow toward

the base and faded above; and it has the same pari-

etal tubercles and outer lip features as per A. (S.)

contrarius (s.s.).

This subspecies also shows a rare intermediate

form (6 shells), which shares features of both of the

other two forms thus proving they are only colour

morphs. These shells have a single very thin to thick

dark, brown or blackish band at the periphery

(mid-banded shells). The band is seen at the suture

on the spire or not (1 shell). The band is absent on

the last whorl in one shell, evanescent in another

and faint to bold in the other four shells.

Compared to A. (S.) laevus kissuensis and A.

(5.) laevus janetabbasae n. ssp. the band patterns

are extremely variable and more randomly devel-

oped; and partial bands are quite common, either

incomplete or evanescent. The atrifasciate shells

have less variation in band colour combinations

than those of A. ( S .) laevus janetabbasae n. ssp.,

but there is greater colour variation in the ground

and bands in its pallibicinctate shells. Compared

to all the other subspecies, on average, this sub-

species has a wider apex and a shorter spire for the

same spire width; therefore, it has a greater spire

angle. Partially interrupted bands caused by dilu-

tion streaks, tend to occur on the spire rather than

on the last whorl. Such interruptions occur behind

the lip on the only shell seen with all six bands

present. The formation of the superior band and

two-toned peripheral band is as follows: all three

bands are brown on the second whorl, changing to

slate on the third whorl; fade late on the penulti-

mate whorl; and then deepen again early on the

last whorl. On the fourth whorl, the space between

bands 3 and 4 becomes grey, thus forming a two-

toned band with a pale grey central zone, with the

whole band intermittently faded.

DISCUSSION

Compared to the A. ( S .) contrarius subspecies,

the A. (A.) laevus subspecies differ by the superme-

dial and submedial bands displaying hues other

than yellow; the submedial bands lack a same-

coloured zone suffused over them; and the lack of

axial or oblique flammules divided or not by a de-

coloured (i.e. of the ground colour, whitish) or yel-

low supermedial band. In addition, the

parieto-columellar tubercle is variably developed

among the subspecies, never shelf-like; the lip ter-

mination is fused to the body whorl and the parieto-

labral tubercle when present (minute or elongate

Lectotype designation and descriptions of two new subspecies of Amphidromus (S.) laevus (Pulmonata Camaenidae) 191

lump), so no gaps; and no canal beneath the suture

inside the aperture. They also lack a pinkish brown

medial zone between the bands of the superior and

peripheral band pairs seen in the maculated zones

of the laevus- like shells (Fig. 18). They may also

display post-mora modification of pattern ele-

ments, also seen in A. ( S .) contrarius hanieli

Rensch, 1931 (and in A. (S.) reflexilabris hanielanus

Rensch, 1931).

Even though some A. (S.) contrarius nikiensis

shells have a similar colouration and band pattern

(laevus-\i\.Q shells figure 18) to the lectotype of A.

( S .) laevus laevus , the latter has greater links to its

subspecies. The data in Table 1 clearly show this,

where all shell measurements are similar and no par-

ticular subspecies has anything outstanding dimen-

sional features. However, some distinctive features

can be noted: A. ( S .) laevus janetabbasae n. ssp.

5 mm

Figure 23. Holotype of Amphidromus (Syndrom us) laevus janetabbasae n. ssp.

Figure 24. Holotype of A. ( S .) laevus misled n. ssp.

192

Jeff Parsons

has a larger umbilicus; A. (A) laevus kissuensis has

a smaller aperture for the same sized shell; and on

average A. ( S .) laevus nusleti n. ssp. has the least

number of whorls, a lower H/D ratio and smaller

shells, with the largest shells coming from Moa,

Roma and Tutuala. All subspecies have a AH/H

ratio less than 0.5, plus similar columellar and

perch angles.

The Kisar, Roma and Tutuala populations all

have a rimmed lip, which is a distinctly reflected

edge. For A. ( S .) laevus nusleti n. ssp. the lip edge

is variably flat or rimmed, with the degree of re-

flection dependant on lip maturity. In A. ( S .) laevus

janetabbasae n. ssp., only a few gerontic shells

with greatly thickened lips have a weakly rimmed

edge. The shape, colouration and pattern similarities

of shells from Tutuala, Roma and Leti Islands (Leti,

Moa and Lakor) suggest they originated from a

common population. The A. (A) laevus romaensis

lectotype shows a great similarity in shape and pat-

tern modification to some shells of A. (A) laevus

janetabbasae n. ssp., and the lack of a dark apex.

But it has a stronger posterior lip curvature and a

wide flattened parieto-columellar tubercle com-

pared to A. (A) laevus janetabbasae n. ssp., which

has a thin curved line of callus (rarely a short

smudge) and a more angular aperture. The same tu-

bercle in A. (A) laevus nusleti n. ssp. is short and thick,

and a small-flattened lump in A. (A) laevus kissuensis.

A. (A) laevus kissuensis appears to have de-

veloped from a separate population to the other

subspecies, differing in several features. Most

shells have a whitish ground and the atrifasciate

shells rarely having a yellow last whorl. The sub-

sutural band when present is only ever yellow, of

the same hue as the supermedial and submedial

bands. The most distinctive feature is both colour

morphs may display a third band network of lime

green lines on the last whorl, something not seen

in the other populations studied. The closest ties

are between the subspecies from Leti and Moa,

isolated only by a narrow sea channel. However,

on Leti there is a predominance of the pallibicinc-

tate shells compared to a scarcity on Moa. This

suggests isolation was not only by a physical bar-

rier but also by niche preference. Pallibicinctate

shells prefer the drier and more open thorn forests

on Leti, while atrifasciate shells prefer more

shaded forests with a less open canopy on both

islands.

Pallibicinctate shells of the above three sub-

species are easily distinguishable. On Kisar, they

are uncommon white shells with two or three yel-

low bands, often obsolescent with some shells grad-

ing to almost pure white; and may have lime green

lines. Rare on Moa, shells with the last two whorls

pale yellow, plus two or three pale to dark yellow

bands, one or both may be partially tinged orange.

Common shells on Leti, with the last two or three

whorls of various yellow hues or grade to saffron;

plus two or three pale to dark yellow or orange

bands, often partially browned or blackened and

rarely partially reddened. Sometimes the subsutural

band is of different colour (rose or reddish orange)

to the other two bands; and rarely has a narrow rose

circumumbilical band (3 shells).

After a mora or growth flaw, atrifasciate shells

of A. (A) laevus janetabbasae n. ssp. commonly

show a patchy loss and colour change (pied forms),

or distinct and rapid colour change. The unmodified

dark bands tend to be black or rarely brown. In the

atrifasciate shells of A. (A) laevus nusleti n. ssp.

studied, unmodified dark bands are black, sooty,

blackish brown, brown or slate; and band modifi-

cation may occur before or after a mora, or occur

randomly and gradually without a mora present. A

single pied form from Leti has only the dark bands

partially absent, which return changed to pink

toward the lip. Both subspecies have a narrow to

wide circumumbilical band that rarely fails to enter

the umbilical interior, and the inner labral margin

may have coloured stains, also seen in shells from

Lakor. Either the purplish anterior stain is dominant

and fades-away as the lip thickens (Lakor and Leti),

or the longer yellow lateral stain is dominant and

weak at maturity (Moa). Similar yellow staining

seen in shells from Tutuala is in fact a preapertural

band showing through from the outside.

A. (A) laevus janetabbasae n. ssp. and A. (A)

laevus nusleti n. ssp. may have streaks that dilute

or deepen the band and ground pigment, often faint

or become distinct toward the lip. The Tutuala po-

pulation has these too, and dilution streaks some-

times faintly or distinctly interrupt the bands on the

spire. This happens in the other two subspecies too,

but only as short sections of affected bands. Ran-

dom partial dilution and deletion also forms inter-

rupted bands. Dilution streaks often disturb the

coloured suffusion on the last whorl, bleaching it

thus showing an undertone (discussed below).

Lectotype designation and descriptions of two new subspecies of Amphidromus (S.) laevus (Pulmonata Camaenidae) 193

The medial zone of two-toned bands may suffer

intermittent dilution, thus forming maculated zones

of pale and dark blotches with dark borders. These

maculated zones generally occur on the spire and

are either short with weak randomly spaced small

blotches (Leti and Moa), or long with wider alter-

nating blotches (Tutuala).

The first network bands may develop brown or

black staining toward a mora and/or the lip in both

colour morphs of A. ( S .) laevus nusleti n. ssp. In

atrifasciate shells of A. (S.) laevus janetabbasae n.

ssp., only the supermedial and submedial bands

have the same staining, commonly reddened too,

and it rarely affects the subsutural band. A. ( S .) lae-

vus (s.s.) shows a similar feature, except the same

bands are firstly invisible and then emerge on the

last whorl stained brown. Only in A. (S'.) laevus

janetabbasae n. ssp. are these bands completely

bleached to white after a mora and develop thin red

or blackish borders. These band modifications are

absent in the other populations. The subsutural band

when present is generally the same colour as the

other two bands in pallibicinctate shells, and com-

monly different in atrifasciate shells (magenta, red

or rose).

Distortion of the lower whorls is similar in

shells from Tutuala, Leti and Kisar, although the

latter have more gibbose whorls. A. (S.) laevus

janetabbasae n. ssp. tends to have elongated distor-

tion from a greater change in the coiling angle. A

weakly developed shoulder and/or an obsolete ly

subangular periphery on the last whorl may exag-

gerate distortion in A. (S.) laevus nusleti n. ssp.; a

dark apical spot is always present in A. (5.) laevus

kissuensis; very rarely absent in A. ( S .) laevus

nusleti n. sp. with only one shell known; present or

absent in shells from Tutuala, Moa and Lakor; and

absent in A. (S.) laevus (s.s.) and A. (S.) laevus

romaensis. Development of parietal tubercles varies

among the subspecies, often absent or at least very

weakly developed. A parieto-labral tubercle occa-

sionally occurs in some shells from three popula-

tions: a minute lump (A. (S.) laevus (s.s.), Leti and

Moa) or a small elongate lump perpendicular to the

lip (Leti and Moa), and the latter is seen in some

shells of A. (5.) sinistralis Reeve, 1849 and A. ( S .)

centrocelebensis Bollinger, 1918.

A parieto-colume liar tubercle develops into one

of four forms. The following is a comparison of A.

(S.) laevus subspecies (locality of each is in brack-

ets) with other species that also develop each one

of these. A. (S.) latestrigatus has a flattened lump

(Kisar, Roma, Moa and Leti), except it is reddish

purple. A. ( S .) beccarii Tapporone-Canefri, 1883

and A. (S.) annae have a faint trace or long thin line

(Moa). A long narrow ridge (Moa and Tutuala)

forms in A. ( S .) centrocelebensis and A. (S.) contrar-

ius (s.s.) (moderately developed). Lastly, a short

narrow ridge (Leti) develops in A. ( S .) maculatus

Fulton, 1896 and A. ( S .) kuehni. The same callosity

when well developed in A. ( S .) contrarius (s.s.) is

like a ledge jutting out from the inner wall. Even

this looks feeble compared to the ridge developed

in A. ( S .) sinistralis , which can be 1 to 2.5 mm thick

rising to the same plane as the outer lip and appear

as an extension of the columella.

Generally, A. (S . ) laevus kissuensis has a white

last whorl, rarely medium yellow. The other popu-

lations have various yellow tones on the last half to

the lowest three whorls. Atrifasciate shells from

Tutuala show the greatest variation of the last whorl

colouration, due to band pigment leakage suffusing

across the whorl or as localised spiral zones. This

suffusion is pink, magenta, purple, brown or greyish

green, with an undertone of cream or yellow. A. (S.)

laevus janetabbasae n. ssp. may have a pink suf-

fused last whorl and a yellow undertone, or as pink,

brown or pale reddish orange spiral or longitudinal

suffusion zones. A. (S.) laevus nusleti n. ssp. shows

only pink spiral suffusion zones.

According to Taylor (1914), an abrupt change

in pigment colour can be due to a change in diet.

Taylor also says if pigment-secreting cells are la-

tent or undeveloped during early shell develop-

ment, bands gradually or suddenly develop at the

commencement of a growth period, or they show

atrophy at the termination of a growth period that

causes the partial or complete loss of bands during

regrowth. This describes quite well some of the vari-

ation seen in the A. (, S .) laevus group. However, the

atrophy of the pigment-secreting cells may occa-

sionally be a gradual process with the bands slowly

fading-away. Taylor (1914) also discusses links for

banding variation in Theba pisana Muller, 1774.

He says in exposed habitats shells tend to have

delicate linear banding that tends to be irregularly

developed, while in less open and more shaded habi-

tats shells have more distinct and better developed

banding. This applies well with the banding of the

A. (S.) laevus group, with the addition of the palli-

194

Jeff Parsons

bicinctate shells being a form possibly adapted for

exposed habitats.

Addition pertains to extra bands inserted into

the pattern, not split from the main pattern bands,

and occur as either short or long segments (partial

bands). Division applies to main pattern bands that

split into narrower bands or lines (bandlets). Width

variation simply refers to different shells having

either thin or thick bands, or in combination. Fusion

ascribes adjacent bands gradually widening during

development and connect to form wider bands. If

fused bands dilate as well, then many bands may

connect behind the lip to form a partially very dark

shell (pseudomelanism). Spreading is band pig-

ment leaked into the interspaces as stains, short

streaks or connection of adjacent bands (two-toned

bands), and widespread or zoned suffusion. Pig-

mentation variation involves intermittent fading of

band and/or ground colour, occurring faintly along

growth lines or wider marks along bands that may

grade to maculated zones or even interrupted

bands. In comparison, dilution and deepening streaks

are pattern elements that modify the shell pigments

more strongly. Reduction refers to indistinctly or

irregularly developed bands, which are partially

faded and those that gradually fade (evanesce) or

gradually narrow before vanishing. Results of

studies on other banded snails suggest that the band-

ing pattern is probably under genetic control in this

species as well.

The A. ( S .) laevus group have another four

types of pigmentation variation. Firstly, incom-

plete xanthism relates to the pallibicinctate shells

where the yellow or orange bands are impercepti-

ble on a same-coloured last whorl (Leti); and sim-

ilarly coloured shells that have dark bands on the

upper whorls (Tutuala). Incomplete albinism ap-

plies to almost pure white shells with very faint or

obsolescent supermedial and submedial bands

(Kisar); and shells with dark banded upper whorls

and pure white lower whorls lacking first network

bands (Tutuala). Partial leucism refers to the pied

forms with short to long patch- like modification

of some to all bands after a mora (Moa and Roma).

It affects band colouration and presence, rarefy af-

fecting ground colour (Roma). No juvenile shells

were available to show the pied modification af-

fects bands on the base. Lastly, selective leucism

pertains to the discolouration or dilution of band

pigment without affecting the ground colour, for

at least part of the shell (Kisar, Leti, Moa and Tu-

tuala). For example, the bands change from brown-

ish to pinkish or from blackish to purplish, usually

abruptly or gradually after a mora, less often ran-

domly on the shell.

Overall the subspecies may share certain con-

chological features (e.g. weakly to distinctly abaper-

turally angled and/or proclined columella), but local

modifications of others (e.g. parietal tubercles and

sculpture) can be used to separate them in mature

shells.

CONCLUSIONS

In their isolation, each island population of A.

(S.) laevus has developed localised variations in

shell pattern and minor differences in shell charac-

ters not found in other populations. Differences in-

clude, but are not limited to: apex colouration (black

or dark apical spot, present or absent); coloura-

tion of the protoconch and lower whorls; thickness

and reflection of the lip and columella; surface sculp-

ture; and the development of parietal callosities.

Variation in the banding occurs via a number of mod-

ifications, in terms of: addition, division, width vari-

ation, fusion, spreading, pigmentation variation,

reduction, plus genetic control of the number of

bands present.

Collectively these variations have developed a

number of different subspecies, and the whole in-

tension of this study was to show that. All of the

above points show that the subspecies of A. (S.) lae-

vus are a group of snails that are highly polymorphic

in their shell colour and banding. This is especially

the case for the Tutuala population. Due to the lack

of study material, this study was unable to deter-

mine if shells like that of the A. ( S .) laevus laevus

lectotype occur on Lakor or at Tutuala. Further re-

search is required to determine if they are separate

subspecies. Knowledge of the conchological fea-

tures and distinctive phenotypes created by local

pattern variation, allows the subspecies to be distin-

guished in a mixed sample.

ACKNOWLEDGMENTS

I would like to thank John Abbas for the supply

of type and comparative material for this study, and

Lectotype designation and descriptions of two new subspecies of Amphidromus (S.) laevus (Pulmonata Camaenidae) 195

for photos of shells in his collection. I owe a debt

of gratitude to the following people: Danny Eibye-

Jacobsen, Assoc. Prof., Curator (of the mollusc col-

lection) and Tom Schiotte for photos (ZMUC); Dr.

Mandy Reid, Malacology Collection Manager for as-

sistance in depositing type material (AM); Jonathan

Ablett, Curator (of Non-Marine Mollusca and

Cephalopoda, Division of Invertebrates, Zoology

Department) for assistance in depositing type mate-

rial and their photographic unit for photos

(NHMUK); and Dr. Silke Stoll, Curator (of the

mollusc collection) for assistance in depositing type

material and photos (LMA).

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Schiff Gazelle gesammelten -Land- und Siiss Avasser-

Mollusken, Monatsberichte der Koniglichen Preussis-

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Martini F.H.W., 1777. Fortsetzung der vorlaufigen

Nachricht und Abbildung einiger linksgewundenen

Schnekken, Neue Mannigfaltigkeiten. 4. Jg: pp. 416-

418, Tab. I, figs. 8-9.

Maurice J., 2013. Pyramidella terebellum (Muller, 1774)

1035, PYRAMIDELLIDAE Gray, 1840, Fiche 1,

Mollusques de file de la Reunion. Available at:

http://vieoceane.free.fr/mollusques/intro_frame.htm

[Accessed 18 May 2013]

Muller O.F., 1774. Vermium terrestrium et fluviatilium,

seu, Animalium infusoriorum, helminthicorum et

testaceorum, non marinorum, succincta historia,

Volumen Alterum, Havniae et Lipsiae, apud Heineck

et Faber, ex officina Molleriana; Helix laeva : pp. 95-

96, No. 293 (not illustrated).

Pilsbry H.A., 1900. Manual of Conchology, Structural

and Systematic, with Illustrations of the Species, Se-

ries 2, Volume 13; Conchological Section, Academy

of Natural Sciences of Philadelphia, Philadelphia;

Amphidromus: pp. 127-234, pi. 46-71.

Reeve L., 1849. Conchologia iconica, or, Illustrations of

the shells of molluscous animals, Volume V, (Sep-

tember 1848); Reeve Brothers, London; Bulimus lae-

vus pi. 37, f. 216 b.

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Rolle H., 1903. Neue Amphidmmus- Formen, Nachrichts-

blatt der Deutschen Malakozoologischen Gesellschaft,

35 (No. 9, u. 10): Amphidmmus laevus var. romaensis

and A. /. var. kissuensis p. 157 (not illustrated).

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phidromus (, Syndromus ) laevus kissuensis Rolle 1903

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index.asp?objekt_id=562100&sprache kurz=en>

[Accessed: 24 March 2013]

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phidromus (, Syndromus ) laevus romaensis Rolle 1903

Available at: <http://sesam.senckenberg.de/page/index. asp?

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24 March 2013]

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

of Amphidmmus from Atauro Island, East Timor (Gas-

tropoda, Pulmonata, Camaenidae). Basteria 70: 23-28.

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Taylor J.W., 1914. Monograph of the land & freshwater

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Helicidae: pp. 1 99-446, pi. XX-XXXV (includes dis-

tribution maps).

Biodiversity Journal, 2014, 5 (2): 197-200

A new species of Agrilus Curtis, 1 825 from Brazil (Coleoptera

Buprestidae)

Gianfranco Curletti 1 & Letizia Migliore 2

'C/o Museo Civico di Storia Naturale, Parco Cascina Vigna, 10022 Carmagnola, Torino, Italy

2 Laboratorio de Ecologia Evolutiva de Insetos de Dossel, Departamento de Biodiversidade, Evolu?ao e Meio Ambiente, ICEB,

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

ABSTRACT A research of Museu de Entomologia da FEIS/UNESP, campus de Ilha Solteira, Sao Paulo

(SP) region, Brazil (MEFEIS) in order to monitoring the secondary xylophagous species,

showed the presence of a new species of Agrilus Curtis, 1825 (Coleoptera Buprestidae) that is

here described: Agrilus ( Agrilus ) flechtmanni n. sp.

KEY WORDS Brazil; Coleoptera; Buprestidae; Agrilus, new species.

Received 01.04.2014; accepted 15.05.2014; printed 30.06.2014

INTRODUCTION

A staff of the departement of Entomology of

FEIS/UNESP (Universitade Estadual Paulista,

Campus de Ilha Solteira, Brazil), made a research

for studying the biology of secondary xylophagous

insects.

Some species of Neotropical Coleoptera Ce-

rambycidae (Serville, 1835), manly belonging to

the genus Oncideres Serville, 1835 (Lamiinae) at-

tack tree species girdling the branches with the

aims of interrupting the lymphatics vessels and

killing the apical part where the larva will live

(Fig. 1). Some wood-boring secondary species de-

velop in those branches, taking advantage of the

particular habitat. Particularly favoured are ob-

viously small species that are able to colonize the

restant space made available by the primary host.

Oncideres species burrow large galleries in the

wood, leaving intact the bark only, and the species

specialized to colonize the remainder of the branch

are generally small and mainly sub-corticicolous.

Particularly favoured are some monovoltine species

of the genus Agrilus Curtis, 1825. The girdled

branches are weak and liable to breakage so they

are often found at the base of the trees, where they

were collected and placed in breeding containers,

waiting for the adults to emerge.

In the course of this research we obtained some

specimens of a new species of Agrilus (Curtis,

1825) that is described here.

MATERIAL AND METHODS

The study area is in Brazil, Sao Paulo (SP) re-

gion (Fig. 2). The specimens were provisionally

stored in formaldehyde, followed by dry prepara-

tion and glued on a card for the study and descrip-

tion and conservation.

The genitalia were placed on the same card.

The pictures were made with a Coolpix P6000

connected with a stereomicroscope Leica MZ6,

elaborated with Adobe Photoshop CS5 Extended

vers. 12.0 and stacked with Combine Z4 pro-

gram.

198

Gianfranco Curletti & Letizia Migliore

Figure 1. Branch girdled by Oncideres sp. (Coleoptera Ce-

rambycidae), French Guyana (photo S. Brule).

ABBREVIATIONS. MEFEIS = Museu de En-

tomologia da FEIS/UNESP, campus de Ilha

Solteira; Sao Paulo (SP), Brazil. MCCI = Museo

Civico di Storia Naturale di Carmagnola, Torino,

Italy.

Agrilus {Agrilus) flechtmanni n. sp.

Examined material (Figs. 3-5). Holotypus

male: BR[azil], SP, Ilha Solteira, UNESP campus,

20°25T1.65"S - 51°20'28.19"W, ex broken Anade-

nanthera falcata [now macrocarpa, Fabaceae

family], 3.III.2011, Flechtmann C.A.EI. legit (ME-

FEIS). Paratypes: 1 male and 3 females, idem,

respectively 18.III.2011, 26.IE2011, 19.III.2011,

10.IV.2011; 3 males and 5 females, Brazil, SP, Tres

Lagoas reforested degraded area, 20°44'55"S -

51°39'36"W, V, Nascimento leg., ex Cerambycidae-

girdled Anadenanthera macrocarpa branch on the

ground, 15.VII.2013 (MEFEIS and MCCI)

Description of holotypus. Length 4.4 mm.

Lengthened form; uniformly bronze-brown, less

brilliant, with yellow pale pubescence on elytra

forming three couples of pubescent spots. Vertex

slightly depressed, % width of anterior margin of

pronotum. Frons flat, green, glabrous, brilliant,

with sericeous sculpture. Clypeus without trans-

Figure 2. Study area: Brazil, Sao Paulo (SP) region, loca-

lity of the new species of Agrilus.

versal carina. Antennae short, green, serrate from

antennomere 4. Pronotum with lateral margins an-

teriorly arcuate and posterior angles less acute.

Premarginal carinula entire. Marginal carinae sub-

parallel, separated from base. Disc with a slight

depression before the scutellum. Sculpture trans-

verse and thickened. Anterior prostemal lobe cut

in the middle. Prosternal plate parallel-sided, bor-

dered. Scutellum carinate. Elytra with apices rounded

and denticulate. The first pair of elytral spot is on

the humeral callus, the second and third along the

suture before the middle and at the apical three-

quarters respectively; few thin hairs join the first

and second spots. Ventral side darker than the dor-

sum, uniformly covered by short thin pubescence.

Last visible ventrite rounded at the apex. Legs

with green reflections. Metatarsus shorter than

metatibia, basal metatarsomere shorter than the

sum of the following two (l<2+3). All claws bifid.

Aedeagus is shown in figure 5.

Variability. Length from 4.4 to 5.5 mm. In

some specimens the first and second couple of

elytral pubescence appear clearly separate without

the thin pubescence among them that characterize

the holotype. The females have frons copper and all

claws bifid as the males.

Etimology. After the name of the holotype col-

lector.

A new species of Agrilus Curtis, 1825 from Brazil (Coleoptera Buprestidae)

199

Figures 3, 4. Agrilus ( Agrilus ) flechtmanni n. sp., paratype female, lenght 5.5 mm. Figure 5. A. flechtmanni n. sp,

aedeagus, 1 .4 mm.

Remarks. For the shape, dorsal color and the

elytral spots, A. flechtmanni n. sp. is similar

and may be confused with A. aegrotus (Curletti

et Migliore, in press).

A. aegrotus differs for having the frons smooth

and not sericeous, the prostemal plate bordered and

enlarged at the top, sterna minus pubescent and

metatarsomeres more lengthened, with tarsal

formula 1 =2+3+4.

ACKNOWLEDGMENTS

We wish to thank Massimo Meregalli (Univer-

sity of Turin, Italy) for the suggestions, Stephane

Brule (Societe entomologique Antilles-Guyane) for

the picture of branch girdled by Oncideres sp. and

obviously Carlos Flechtmann (FEIS/UNESP, Uni-

versitade Estadual Paulista, Campus de Ilha

Solteira, Brazil) for the confidence in sending the

studied material.

REFERENCES

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

tis, 1829 nas colegoes do Museu de Zoologia da Uni-

versidade de Sao Paulo (Coleoptera Buprestidae).

Papeis Avulsos de Zoologia.

Curtis J., 1825. British Entomology; being illustrations

and descriptions of the genera of insects found in

Great Britain and Ireland: containing coloured figures

from nature of the most rare and beautiful species,

200

Gianfranco Curletti & Letizia Migliore

and in many instances of the plants upon which they Serville A., 1835. Nouvelle classification de la famille

are found. London, Printed for the author, volume 2, des Longicomes. Annales de la Societe entomologique

plates 51-98 with text, not paginated. de France, 4: 5-100.

Biodiversity Journal, 2014, 5 (2): 201-208

A new species of rissoid of the genus Alvania Risso, 1 826 from

the E-Sicily: Alvania maximilicutiani n. sp. (Gastropoda Ris-

soidae)

Danilo Scuderi

Via Mauro de Mauro 15b, 95032 Belpasso, Catania, Italy; e-mail: danscu@tin.it

ABSTRACT Alvania maximilicutiani n. sp. is here described and figured as a new Mediterranean species

from the E-Sicily. The most similar species in morphological characters are A. clathrella

(Seguenza L., 1903), A. dalmatica Buzzurro et Prkic, 2007, A. dianiensis Oliverio, 1988, A.

dictyophora (Philippi, 1844), A. hallgassi Amati et Oliverio, 1985. All these species and other

similar Mediterranean and not Mediterranean congeners are here compared to the new species,

which differs by the very minute dimensions, being one of the smaller Alvania ever described,

the protoconch morphology and the colour pattern of the external soft parts. The Macaronesian

A. piersmai Moolenbeek et Hoenselaar, 1989, A. pouched Dautzenberg, 1889, A. spreta

(Watson, 1873) and other congeners are furthermore compared to A. maximilicutiani n.sp. The

new species could also resemble a dwarf form of A. lanciae, but to a more deepened exam of

the shell the latter species appears morphologically very different in both protoconch and

teleoconch characters. The type material of A. maximilicutiani n.sp. was collected in very shallow

waters in the rocky shores of the small village S. Giovanni Li Cuti (Catania, Italy).

KEY WORDS Gastropoda; Rissoidae; new species; taxonomy; Mediterranean Sea; Recent.

Received 13.04.2014; accepted 19.05.2014; printed 30.06.2014

INTRODUCTION

Rissoids of the genus Alvania Risso, 1826 un-

dergone a high adaptive radiation and are uniformly

distributed along the main marine biocenosis. Some

Authors debated whether they could be considered

as representative species of a separated family

(Golikov & Starobogatov, 1975), mainly based on

anatomical proofs which other Authors, through

further deepened studies, considered not as clear

and stable characters but only as anatomical schemes

variable among species (Ponder, 1984). The anatomy

of Alvania is thus comparable to that of Rissoa De-

smarest, 1814 being the animals rather similar in

structure and differences are due mainly to different

habitat preferences (Ponder, 1985). Similitudes are

close related as concerns the animal and shell be-

tween genera Alvania and Crisilla Monterosato,

1917 being the latter tentatively separated only as

a subgenus by Ponder (1985), but currently consid-

ered as a valid separated genus for the actual check-

lists (Clemam, 2013; MarBEF, 2013).

The Mediterranean represents an elective geo-

graphic area where rissoids exhibit wonderful pat-

terns of speciation (Bouchet in Giannuzzi-Savelli

et al., 1996), due to its environmental variability of

habitats. With 160 species of rissoids the Mediter-

ranean represents the most diverse site in the world

and plays an important role as one of the source of

rissoid-flow in the world (Avila et al., 2012). In the

202

Danilo Scuderi

Mediterranean sea and the adjacent Macaronesian

area (E- Atlantic) the genus Alvania is the most

abundant of rissoid species (Gofas, 1990; Van der

Linden, 1993; Hoenselaar, H.J. & Goud, J., 1998;

Avila, 2000; 2012), while lower number of species

are also present in the western Atlantic and Indo-

Pacific Ocean (Bouchet in Giannuzzi-Savelli et al.,

1997; Garilli & Parrinello, 2010), E- Africa (Gofas,

1999). In particular, species with non-planktotrophic

development seem to be limited to only the western

or the eastern part of the Mediterranean (Garilli &

Parrinello, 2010). Moreover, this basin is the site

with higher number of endemic species of Alvania

(Avila et al., 2012). Many species of Rissoidae are

reported as endemisms to restricted areas of Mediter-

ranean, particularly islands (Bogi et al., 1983; Oli-

verio, 1986; Amati & Oliverio, 1987; Oliverio,

1988; Giusti & Nofroni, 1989; Oliverio & Amati,

1990; Cecalupo & Quadri, 1995; Margelli, 2001;

Buzzurro, 2003; Micali et al., 2005; Buzzurro &

Landini, 2006; Buzzurro & Prkic, 2007; Oliver &

Templado, 2009). Numerous species of molluscs,

among which some of Alvania, are described as en-

demic to the E-Sicily, which represents a high hotspot

for the speciation of molluscs in the Mediterranean,

due probably to its variety of different environments

present in this area: A. dictyophora (Philippi, 1 844)

and A. clathrella (Seguenza L., 1903, ex Mon-

terosato ms) are an example.

Anew species of Alvania, named A. maximilicu-

tiani n.sp., was found in the E-Sicily coasts (Fig. 1)

and is here described and figured as new for science

and compared to the close similar congeners.

MATERIAL AND METHODS

Living materials of the new species were col-

lected by brushing the surface of little lava stones

inside a net of 0.5 mm mesh at a depth of 0.5 to

2.5 m. No empty material was found among the

shell grit collected handily with ARA. Drawings

of the external soft parts were obtained observing

the living animals in aquarium. Fossil materials of

similar species were studied and compared to the

new Alvania.

ACRONYMS. Museo del Dipartimento di Bio-

logia Animale, University of Catania, Italy

(MB AC); Alberto Villari malacological collection,

Messina, Italy (AVC); Bruno Amati malacological

collection, Roma, Italy (BAC); Danilo Scuderi ma-

lacological collection, Catania, Italy (DSC).

Alvania maximilicutiani n. sp.

Examined material. Holotypus, Catania, E-Si-

cily, Italy: S. Giovanni Li Cuti, in shallow water,

under lava stones. Paratypi, same data of holotypus,

11 living specimens.

Holotype in MBAC, n. MBLMC-CT-79; 1 paratype

in AVC; 1 paratype in BAC. Other paratypes in

DSC.

Description of holotypus. Shell (Fig. 6)

ovate-conic, stout, relatively strong, imperforate,

1.1 mm high x 0.65 mm broad. Teleoconch consti-

tuted by 1.9 whorls in adult specimens, separated

by marked sutures, bearing two spiral chords on the

first and rather on the majority of the last whorl:

only a trace of a third spiral chord appear almost at

the end-half of the last whorl. Spiral micro sculpture

is present over the surface of all the tele-whorls. The

axial ribs are angular, marked, opisthocline and in

number of 12 on the first tele- whorl, 14 on the body

whorls, never reaching the base of the shell. The

first adapical spiral chords are less marked than the

axial ribs and become stronger at the base. They

form a cancellate sculpture at the intersection with

the axial ribs, with not marked, almost rounded,

A new species of rissoid of the genus Alvania from the E-Sicily: Alvania maximilicutiani n. sp. (Gastropoda Rissoidae) 203

tubercles at the crossing points. At the base four

well marked spirals are present, except the last

which appears almost vanishing. Very thin spiral

threads cover the interspace and run over the axial

ribs too. The last whorl forms rather 76% of the

total shell height. Aperture ovate, drop shaped, with

thick uninterrupted peristome, which bears no den-

ticles and form a varix in the outer lip. Background

colour almost dark-brown in protoconch and first

tele-whorl, creamy in the body-whorl with two

small darker bands, the first sub-sutural, which

forms dark spots in correspondence of the tubercles

(one every three axial ribs) and the second, larger,

at the base.

Protoconch (Figs. 10, 11) low-rise, paucispiral

with direct development, constituted by 1.3 to 1.5

regularly convex whorls. Protoconch I slightly less

than 1 whorl, with 7-8 very thin spiral threads. Pro-

toconch II with 13-14 equally thin spiral threads.

The living animal (Fig. 12) is whitish as back-

ground colour and bears dark-brown to almost black

strips on head, anterior foot and opercular area. In

the head they are “V” shaped just before the eyes

and straight in the snout and foot, surrounding the

operculum and extending to the edge in the middle

of the foot: it is not visible on the sole of the foot,

where a whitish gland is visible in transparency in

the middle. The same gland is visible under the oper-

culum. Two whitish granular masses are also visible

on the snout just near the eyes. Few white stains are

present on both the cephalic tentacles. Only one

metapodial tentacle is visible. Operculum (Fig. 9)

thin, paucispiral, with eccentric nucleus.

Variability. All the fully developed adult spec-

imens collected seems not to differ in size (1-1.2

mm high x 0.65 mm wide) (Figs. 2-5; 6-8). The

only character which seem to vary is the colour of

the body whorl, entirely dark brown to creamy with

brown spots. Protoconch and first tele- whorl are al-

ways darker. The colour variability could be linked

to the shady habits of the animal. If so, the paler co-

loration could be considered typical and the darker

an adaptation.

Etymology. The specific name is in memory of

my father, Massimo, and also recalls the small vil-

lage, locus typicus of the present species, where he

spent his life as a fisherman and where he con-

ducted me to begin my marine biology career and

my malacological studies.

Biology and Distribution. The new species

seem confined exclusively to rocky very shallow

waters, under stones. Only living specimens were

hardly collected. It was never recorded among shell

grit. The new species was collected only from the

type locality in the Jonian sea, but, because of its

very small dimensions, it could be unnoticed and

its real distribution range could be wider.

Comparative notes. Because of the very small

dimensions of adult specimens, the peculiar charac-

ters of the teleoconch, particularly the number of

whorls, the sculpture and the colour pattern of the

shell, the protoconch sculpture and the external soft

parts of the living animal, the new species is unique

among all the Mediterranean and not Mediterranean

species of the same genus. The comparative notes

here following explore the possibility that a dwarf

form of a still known species is involved and

demonstrate the validity of the new species. So,

among Mediterranean species, A. maximilicutiani

n. sp. is morphologically similar to A. lanciae

(Calcara, 1845), A. aeoliae Palazzi, 1988, A.

datchaensis Amati et Oliverio, 1 987, A. fractospira

(Oberling, 1970) on one hand and to A. dictyophora,

A. hallgassi Amati et Oliverio, 1985, A. dianiensis

Oliverio, 1988, A. dalmatica Buzzurro et Prkic,

2007 on another hand. In the meantime the new

species is here compared to some Macaronesian

congeners: A. grancanariensis Segers,1999, A.

hoeksemai Hoenselaar et Goud, 1998, A. moniziana

(Watson, 1873), A. piersmai Moolenbeek et Hoense-

laar, 1989, A. pouched DautzQnbQYg, 1889, A. spreta

(Watson, 1873).

The possibility that the new species could be a

dwarf form of A. lanciae (Fig. 13), or of a close sim-

ilar species as A. datchaensis , A. fractospira , was

the first eventuality explored, but the different pro-

toconch, lacking the “orange skin” sculpture, colour

pattern of shell and soft parts make it easily recog-

nisable. Amati (2012) recently re-described and fi-

gured A. lanciae , A. arguta Locard et Caziot, 1900

and A. consociella Monterosato, 1884, considering

these two latter taxa, often distinguished for the

stouter shape and the larger dimensions, as syn-

onyms of the former. In the same paper he presented

an exhaustive comparison of A. lanciae with A.

datchaensis and. A. fractospira.

A. aeoliae has a more slender teleoconch, with

a colour pattern also similar, being almost darker in

the first whorls, paler with two darker spiral bands

204

Danilo Scuderi

in the body- whorl, but they do not form any stain

on the axial ribs. The protoconch has a different

shape, more elevated and with dense undulated spi-

ral sculpture like that of A. lineata, and the axial

ribs of teleoconch, which are opisthocline too, are

more numerous and with heavier spiral sculpture.

Two new species of Alvania were lately described

for the Mediterranean (Tisselli & Giunchi, 2013): A.

bozcaadensis Tisselli et Giunchi, 2013 and A. cam-

panii Tisselli et Giunchi, 2013. They are both quite

different from A. maximilicutiani n. sp. on account

of shell and protoconch differences. The former is a

species close similar to A. dorbignyi (Audouin, 1826)

as for the almost smooth protoconchandthe teleo-

conch, with characters of A. lineata mixed with those

of A. discors (Allan, 1818). The latter seems linked

to the A. datchaensis/A.fractospira group of species

as for both teleoconch characters and protoconch,

with “orange skin” sculpture.

Typical A dictyophora (Fig. 14) are usually 2.5—

2.8 mm, bear three spiral chords (excluding the 4

basal, present in the space between the upper attach-

ment of the external lip and the end of the base of

the shell) and 9-10 axial ribs in the bodywhorl. But

small specimens of this species (up to 1.4- 1.7 mm)

with only 2 spiral cHords are known (Fig. 15;

Palazzi & Villari, 2001, Fig. 27). These dwarf forms

of A. dictyophora resemble in shell morphology the

new species, which however is easily distinguish-

able by the smaller dimensions, the lower number

of whorls, the different protoconch and colour pat-

tern of the shell, the less marked spiral cords which

cross the more opisthocline axial ribs.

Shells of A. hallgassi and A. dianiensis are al-

most close similar, being the former only more del-

icate in teleoconch sculpture. The protoconch in

both species is constituted by 6 marked spiral

chords, on a smooth (A. hallgassi ) or granulated

with small dots background (A. dianiensis ). The

new species has the teleoconch different in dimen-

sions, shape, shell sculpture and colour pattern and

the protoconch with a different form and more con-

spicuous and delicate sculpture.

A. dalmatica is a species described few years

ago and shares with the above mentioned species a

bodywhorl with a similar sculpture, only slightly

more marked and with pointed tubercles at the in-

tersections. The new species differs for having a dif-

ferent shell colour pattern and sculpture of the last

whorl, while the similar protoconch differs for the

less dense and more marked spiral threads.

A. clarae (Nofroni et Pizzini, 1991) is an easter

Jonian species described for Zakinthos, Greece in

recent time, which shares with the new species a

similar protoconch sculpture and shell colour pat-

tern. Compared to the new species, it is higher and

with one more tele-whorl. The shape of teleoconch

is much more slender, with a different sculpture,

dense of marked tubercles some of which (those on

adapical first spiral chord) are oriented upward. The

subsutural zone is wider. The less wide base bears

one spiral chord less. The protoconch is shorter, less

wide, almost white and bears more marked spiral

sculpture.

A problematic taxon is A. peloritana (Aradas et

Benoit, 1870): Scuderi & Terlizzi (2012) exposed

their idea of this species, based on topotypical ma-

terials, which is different in both protoconch and

shell characters from the new species.

In a recent paper Garilli (2008) restored A. cin-

gulata (Philippi, 1836), which have a protoconch

similar to that of the new species as for general out-

line and sculpture, but the spiral threads are lower

in number and the teleoconch is veiy different in

general outline, more similar to a big A. tenera , and

sculpture, composed by thin and numerous axial

ribs crossed by spiral lines of the same consistence.

Among the numerous Macaronesian congeners

A. maximilicutiani n. sp. is here compared with

those of small dimensions and close similar mor-

phological characters. The new species resulted pe-

culiar in sculpture of teleoconch, characters and

dimensions of protoconch, resulting the smallest

species of Alvania in this area too, sculpture of

teleoconch and characteristics of protoconch. The

morphology of the external soft parts of many

species of this geographical zone are unknown: for

this reason comparisons between species of the liv-

ing animals were not possible.

A. grancanariensis is a species which could be

referred to the Mediterranean A. lanciae group, as

for general outline and sculpture of the teleoconch.

The darker colouration of the protoconch and first

tele-whorls make it similar to the new species,

which however could be easily distinguished for

the protoconch characteristics. Another congener

whose coloration is similar to this latter species is

A. hoeksemai, but the predominant spiral sculp-

ture, the heavier shell aspect and the different pro-

toconch make it easily differentiable from the new

species.

A new species of rissoid of the genus Alvania from the E-Sicily: Alvania maximilicutiani n. sp. (Gastropoda Rissoidae) 205

Figures 2-12. A. maximilicutiani n. sp. Figs. 2-5: paratype, S. Giovanni Li Cuti, H: 1 mm (D. Scuderi coll.). Fig. 5: detail

of protoconch, same data. Fig. 6: holotype, SEM photograph, H: 1.0 mm (MBAC). Figs. 7, 8: paratypes, same data of ho-

lotype, SEM photograph, both H: 1.1 mm (B. Amati coll, and A. Villari coll.). Fig. 9: upward view of operculum, H: 600

pm (same data of Fig. 7). Fig. 10: protoconch of holotype (257x328 pm), front view. Fig. 11: protoconch of paratype

(267x300 pm), lateral view. Fig. 12: external morphology of the soft parts. Figure 13. A. lanciae, S. Giovanni Li Cuti, H:

2.6 mm. Figures 14, 15. A. dictyophora, Salina, Eolie Is., H: 2.7 mm and 2.0 mm (B. Amati coll.).

206

Danilo Scuderi

The Madeiran A. moniziana shares with the new

species the general shape and number of tele-

whorls, but it is bigger, lacks axial sculpture and is

almost entirely whitish in colour, while the proto-

conch is different at all.

Apart the size, almost double, the different pat-

tern of shell colour, more light coloured with

marked white stains in the peripheral line of each

whorl, the sculpture, constituted by stronger axial

ribs and more numerous spiral chords, expecially

in the peripheral zone, and the different protoconch

distinguish A. piersmai, a small Canary recently de-

scribed species, from the new species.

A. poucheti is a small (2 mm high) species, dark

in colour, which resembles the new species for gen-

eral shape, but differs in dimensions, shell and pro-

toconch sculpture.

Juveniles specimens of A. spreta, a small dark

shell Madeiran species, resemble A. maximilicutiani

n. sp. as for the opisthocline shell sculpture and the

coloration. But the former is higher, the number of

whorls being equal, with more incised sutures and

more numerous spiral chords, which give to whorls

a less stouter aspect; the protoconch is different in

colour and sculpture and the external lip in adult

specimens is thicker and more rounded.

According to literature data (Dali, 1889, 1927;

De Jong & Coomans, 1988; Leal, 1989), among

Western Atlantic congeners few species properly

belonging to Alvania recall the new species in di-

mensions and general shell features. A. auberiana

(d'Orbigny, 1 842) has a similar colour pattern, but

is bigger, bears three spirals of the same thickness

of the axial ribs above the outer lip, of which the

uppermost is separated from suture, and has a dif-

ferent protoconch. As the latter species, A.faberi de

Jong et Coomans, 1988 bears a third spiral chord

on the last tele-whorl, while the uppermost colour

band is continuous and the protoconch has different

morphology. The general shape, dimensions and

shell sculpture, together with the different proto-

conch, discriminate A. nigrescens Bartsch et Reh-

der, 1939 from the new species. Another small

similar Caribbean congener is A. moolenbeeki De

Jong et Coomans, 1988 which shares with the new

species similar dimensions reaching 0.9 mm in

adult specimens, but it is almost entirely white and

has spiral sculpture predominant on the two tele-

whorls, a clear umbilicus is present and the proto-

conch has different dimensions and sculpture.

Two eastern Pacific congeners resemble the new

species in morphology, according to literature data

(Bartsch, 1912; Backer et al., 1930; Bartsch &

Rehder, 1939): both A. purpurea Dali, 1871 and A.

cosmia Bartsch, 191 1 are similar in teleoconch sculp-

ture, constituted by only two spirals above the outer

lip. Therefore, compared to the new species, they

are almost double in dimensions, they are consti-

tuted by three rather higher tele-whorls in adult spec-

imens and show a different shell colour pattern; the

protoconch is different. Three more species of the

same geographical area, A. almo Bartsch, 1911, A.

tumida Carpenter, 1857 and A. oldroydae Bartsch,

1911, share with the new species a similar general

outline, almost stubby and swollen, constituted by

only two or three tele-whorls. They all have a dif-

ferent sculpture, mainly constituted by spiral keels

and numerous less marked axial ribs, a different

shell colour pattern and protoconch.

Concerning the external morphology of the soft

parts the new species shows a peculiar character as

concerns the presence of only one single metapodial

tentacle. Usually species of Alvania are reported to

bear 3-7 metapodial tentacles (Ponder, 1985) and

previous observations of one single metapodial ten-

tacle by Jeffreys (1867) for A. punctura (Montagu,

1803), A. lactea (Michaud, 1832) and A. abyssicola

(Forbes, 1850) have been rejected on the basis of

Clark’s (1852) observations (Ponder, 1985). Now

the question on the number of metapodial tentacles

in species of this genus could be re-opened on the

basis of the present observations.

Among fossil species A. maximilicutiani n. sp.

shows some grossly resemblance with A. circum-

cincta SeguenzaG., 1873 and A. bicingidata Seguenza

L., 1903 and with species of Galeodinopsis Sacco,

1895 (Garilli, 2008), but show substantial differences

in dimensions, teleoconch sculpture and protoconch

shape and sculpture.

ACKNOWLEDGMENTS

I am indebted to Stefano Palazzi (Modena,

Italy), Bruno Amati (Roma, Italy), Pasquale Micali

(Fano, Italy) and Alberto Villari (Messina, Italy) for

bibliographic material and interesting comments

and suggestions. I am grateful to Fabio Liberto

(Cefalu) for the critical revision of the text. Frank

Swinnen (Lommel, Belgium) donated several

A new species of rissoid of the genus Alvania from the E-Sicily: Alvania maximilicutiani n. sp. (Gastropoda Rissoidae) 207

Macaronesian species to compare. Andrea Di Giu-

lio (Dipartimento di Biologia, Universita Roma Tre,

Roma, Italy) and Mauro Cavallaro (Istituto di Ve-

terinaria, Messina, Italy) allowed the realisation of

the SEM photographs.

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Biodiversity Journal, 2014, 5 (2): 209-212

Odostomia c rassa Jeffreys, 1 884 junior synonym of Tibersyrnola

unifasciata (Forbes, 1 844), new combination (Gastropoda

Pyramidellidae)

Pasquale Micali 1 , Italo Nofroni 2 & Carlo Smriglio 3

'Via Papiria 17, 61032 Fano, Pesaro-Urbino, Italy; e-mail: lino.micali@virgilio.it

2 Via B. Croce 97 00142 Rome, Italy; e-mail: italo.nofroni@uniromal.it

3 Via di Valle Aurelia 134/C, 00167 Rome, Italy; e-mail: csmriglio@alice.it

ABSTRACT Following the comparison with photos of type material of Odostomia crassa Jeffreys, 1884,

(Gastropoda Pyramidellidae) deposited at the British Museum of Natural History, and further

observations on specimens from whole Mediterranean, O. crassa is proved to be junior syn-

onym of Eulimella unifasciata (Forbes, 1844). The latter is here placed in genus Tibersyrnola

Laws, 1937 on the basis of the constant presence of flutings inside the whorls.

KEY WORDS Pyramidellidae; Tibersyrnola', new combinaton; recent; Mediterranean Sea.

Received 23.04.2014; accepted 15.05.2014; printed 30.06.2014

INTRODUCTION

The species Odostomia crassa Jeffreys, 1884

(Gastropoda Pyramidellidae) was described on par-

tially broken specimens and fragments collected in

north-eastern Atlantic and Mediterranean (Adventure

Bank, Sicily channel, 30-92 fms [corresponding to

50-150 m]). In the original description the new species

was compared only with Eulimella scillae (Scacchi,

1 835) and not with E. unifasciata, that is also reported

and drawn in the same work. We suppose that this is

due to the lack of complete specimens, and the

impossibility to see the shell profile, as well as the

lack of brown spiral band over the whorls, possibly

due to the bad preservation of studied fragments.

Original description (Jeffreys, 1884: 350):

"Shell cylindrical, remarkably thick and strong,

opaque, and glossy: sculpture none, except micro-

scopic lines of growth and the grooves with the

outer lip hereafter mentioned, as well as the pe- ri-

phery being slightly angulated: colour ivory-white:

spire long and finely tapering: whorls 5 only in the

fragments now described, although there would be

from 8 to 10 in perfect specimens; they gradually

increase in size and are flattened: suture slight,

rounded below: outer lip incrassated, furnished in-

side with 8 to 10 spiral striae or flutings, like those

in O. conoidea, O. tenuis, and O. conspicua, as also

in O. costaria and other Crag species: inner lip form-

ing an unusually thick and broad glaze on the pil-

lar: umbilicus none: tooth large, solid, prominent,

and winding round the pillar. Largest fragment L.

0.25 [about 6 mm], B 0.085 [about 2.1 mm]."

The species was figured by Jeffreys (1884: tav.

XXVI, figg. 7, 7a), but the original drawings, here

reported (Figs. 1, 2), are not clear and have not al-

lowed a sure recognition of this species by the later

Authors. Jeffreys compared the new species only

with E. scillae (Scacchi, 1835), stating that main

characters for separation are the “ strong tooth and

inside fluting of the outer lip ”. The fluting of the

outer lip seems to be the only character that separate

it from the other Lusitanic species.

210

P. Micali, I. Nofroni & C. Smriglio

Nordsieck (1972) found that the Jeffreys’s name

was pre-occupied by Odostomia crassa Thompson,

1 845 and proposed the new name Syrnola ( Tiber -

syrnola) wenzi Nordsieck, 1972.

Van Aartsen (1984), after examination of the

type material, concludes that this species is known

only for the original description and some frag-

ments at the British Museum of Natural History, no

one complete of protoconch.

This species was included in the recent lists of

species (Piani, 1980; Bruschi et al., 1985; Sabelli et

al., 1990; WORMS (World Register of Marine

Species, http://www.marinespecies.org/aphia.php?

p= taxdetaild&id=141048 searched on 21/03/2014)

with full validity, while in CLEMAM (Check List of

European Marine Mollusca Database, http://somali.asso.

fr/clemam/index.clemam.html searched on 11/01/2014)

it is considered doubtful.

MATERIAL AND METHODS

Thanks to the courtesy of Dr. Kathy Way and

Ms. Andreia Salvador (BMNHL) we obtained the

photos of the two broken specimens of the type se-

ries deposited at the BMNHL (Figs. 3, 4) with the

number 1885.11.5.1998. These fragments corre-

spond exactly with the original Jeffreys’s drawings

(1884, tav. XXVI, figg. 7, 7a). The larger one (Fig.

3) shows the sign of some fractures on the last

whorl, that may have caused the unusal

periphery /base profile. The two fragments are in

veiy poor conditions and it is quite surprising that

Jeffreys decided to describe a new species based on

a so poor material!

In addition we studied about 80 shells of Eulimella

unifasciata (Forbes, 1844) collected in various local-

ities, covering the whole Mediterranean, at depth rang-

ing from 120 and 500 m (coll. CSR, INR and PMF).

ABBREVIATIONS. British Museum of Natural

History, London = BMNHL; Carlo Smriglio collec-

tion (Rome, Italy) = CSR; Italo Nofroni collection

(Rome, Italy) = INR; Pasquale Micali collection

(Fano, Italy) = PMF.

RESULTS

Based on the original description/drawings and

the photos of type material, the main character,

which is also the unique one, useful for the specific

separation is the “ outer lip incrassated, furnished

inside with 8 to 10 spiral striae or flutings”. This

character has been re-evaluated in the more similar

mediterranean Eulimella species.

We were really surprised to find out that all the

examined specimens of E. unifasciata , as intended

by all European Authors, show the flutings inside

the outer lip, but these are well visible only in fresh

and transparent specimens (Figs. 8, 9), that are rarely

found; the flutings are observable both in fresh spec-

imens and in old ones but only in fresh and intact

specimens it is possible to observe the flutings in

transparency. In addition the external lip edge is

thin, sharp and smooth, and does not show any sign

of internal flutings (Figs. 5, 6), because these are

present up to a quarter of whorl from the aperture

and became visible only in specimens missing the

final portion of last whorl (Fig. 7). In transparent

specimens it is possible to observe that the flutings

appear just after the embryonic whorls, in number

of 3-4 and increase up to 6-8 on the last whorl.

Jeffreys (op. cit.) just after the description of O.

crassa , lists O. unifasciata , pointing out that a spec-

imen from the Gulf of Naples " shows also a grooved

or crenated mouth ".

Di Geronimo & Panetta (1973: 77) reported

Eulimella unifasciata (Forbes, 1844) for the Gulf

of Taranto, pointing out that " la superficie del lab-

hro esterno, che e rotto presenta dei solchi piuttosto

marcati [the surface of the external lip, that is bro-

ken, shows marked grooves]", but also Eulimella

crassa is reported in the same work as valid species,

although the figured specimen is of doubtful deter-

mination. Based on above observations we consider

that Odostomia crassa Jeffreys, 1884 should be

considered a junior synonym of Eulima unifasciata

Forbes, 1844.

As concern the genus where to include this species

in, almost all Authors placed it in genus Syrnola A.

Adams, 1860 with Syrnola gracillima A. Adams,

1860 as type species by monotypy. This genus was

described as follows (A. Adams, 1860: 405) “ Testa

subulata, recta, vitrea, polita; anfractibus plants;

suturis impressis. Apertura oblonga; labio in medio

plica obliqua instructo; labro simplici, acuto ”. Van

Aartsen (1994: 85), when dealing with the use of this

genus, states: “ It is my strong belief however, that a

fold on the columella is not enough to place species

in Syrnola”; Author also states that there is a possi-

ble type material of S. gracillima , at the Museum of

Victoria, consisting of the lower whorls only, which

Odostomia crassa Jeffreys, 1884 junior synonym of Tibersyrnola unifasciata (Forbes, 1844) (Gastropoda Pyramidellidae) 211

Figures 1-9. Tibersyrnola unifasciata (Forbes, 1844). Figs. 1,2. Original drawings. Figs. 3, 4. Type specimens used for ori-

ginal drawings. Fig. 5. Gorgona Island, LI, - 400 m (height 8.2 mm), the flutings inside the aperture are not visible in front

view. Fig. 6. Same specimen of figure 5, the flutings are visible in transparence from the back of aperture (see arrows). Fig.

7. Central Tyrrhenian sea, -380 m; flutings visible in a specimen having broken outer lip. Fig. 8. Anzio, RM, -400 m (4.4

mm), flutings not visible. Fig. 9. Same specimen of figure 8; flutings visible inside all whorls using greater magnification

and proper light angle.

212

P. Micali, I. Nofroni & C. Smriglio

does not correspond with original description due to

lack of columellar tooth. Really also E. unifasciata,

if not broken, has a smooth internal lip, therefore a

check of type material of S. gracillima could clarify

the presence of internal grooves.

Van Aartsen (1994) and van Aartsen et al. (2000)

have not evaluated the applicability of genus Tiber-

syrnola, as done by Nordsieck.

The subgenus Tibersyrnola used by Nordsieck

(1972) was proposed by Laws (1937: 303, 309)

(type species Syrnola semiconcava Marshall et

Murdoch, 1923 fossil from New Zealand) with the

following characters definition: “The shell for

which this name has been provided have all the char-

acters of Syrnola, but in addition the outer lip is

strongly lirate internally “. This taxon has been re-

cently accepted by Beu & Raine (2009) and later

on used with the same meaning by Robba (2013).

For the species dealt with in the present note, we

therefore propose the binomen Tibersyrnola unifas-

ciata (Forbes, 1844). Based on above conclusions

and the opinions of Dautzenberg & Fischer (1896)

and Penas & Micali (1999) the updated synonymy

shall be:

Tibersyrnola unifasciata (Forbes, 1844) ( Eulima )

= Odostomia crassa Jeffreys, 1884 not Thompson,

1845, nee O. pallida var. crassa O. G. Sars, 1878

= Syrnola ( Tibersyrnola ) wenzi Nordsieck, 1972

new name for Odostomia crassa Jeffreys, 1884

not Thompson, 1 845

= Eulimella smithi Verrill, 1881

In the Mediterranean this is the only species to

be placed in genus Tibersyrnola while some others

distributed along the West Africa coast, as Eulimella

endolamellata Schander, 1994, E. angeli Penas et

Rolan, 1997, E. vanhareni van Aartsen, Gittenber-

ger et Goud, 1998, E. boydae van Aartsen, Gitten-

berger et Goud, 2000, Turbonilla Candida de Folin,

1870 (= Odostomia lamothei Dautzenberg, 1912 =

O. etiennei Dautzenberg, 1912), could also be placed

in this genus.

ACKNOWLEDGMENTS

Image courtesy of Harry Taylor, NHMUK Pho-

tographic Unit. Thanks also to Dr. Andrea Di Giulio

(Dipartimento di Scienze, “Roma Tre” University,

Rome, Italy) for the SEM photos, executed at LIME

(Interdepartmental Laboratory of Electron Mi-

croscopy, “Roma Tre” University, Rome, Italy).

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talogo annotato dei molluschi marini del Mediterra-

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Bologna, 348 pp.

Biodiversity Journal, 2014, 5 (2): 213-216

A new record for the American Bullfrog, Lithobates cates-

beianus (Shaw, 1 802) (Amphibia Anura Ranidae), near Rome

(Latium, Italy)

Mauro Grano' & Cristina Cattaneo 2

'Via Valcenischia 24, 00141 Roma, Italy; e-mail: elaphe58@yahoo.it

2 Via Eleonora d’Arborea 12, 00162 Roma, Italy; e-mail: cristina.cattaneo76@libero.it

"■Corresponding author

ABSTRACT The American Bullfrog, Lithobates catesbeianus (Shaw, 1802) (Amphibia Anura Ranidae) is

native to North America. In Italy the introduction of this species dates back to the thirties of

last century and in few years the bullfrog expanded to the point that, at the end of the eighties

the species was known in more than 160 sites. In this paper, a new site of presence in Italy of

the L. catesbeianus is recorded in some ponds at Monterotondo Scalo, a locality near Rome.

KEY WORDS American bullfrog; Lithobates catesbeianus', invasive alien species; Latium; Rome.

Received 21.05.2014; accepted 14.06.2014; printed 30.06.2014

INTRODUCTION

Lithobates catesbeianus (Shaw, 1802) is an am-

phibian of family Ranidae, native to North Amer-

ica. It is a big frog because is able to exceed 30 cm

in length and 1600 g of weight. A very large and

obvious tympanic membrane is present in the tem-

poral area.

In Europe its presence is confirmed, as well as

for Italy, also for Belgium, France (Bordeaux), Ger-

many (Baden- Wurttemberg), United Kingdom (Sur-

rey), Greece, Holland (Breda) and Spain (Caceres).

In Italy the introduction of L. catesbeianus dates

back to the thirties of the last century and seems to

be related to food purposes. Indeed, it can be seen

from the literature that the first where this amphibian

was released was Mantova and, subsequently, it

would spread in short time in other territories thanks

to some peasants who would have used it for edible

purposes (Albertini & Lanza, 1987).

In few years the bullfrog expanded to the point

that, at the end of the eighties the species was

known in more than 160 sites (Scab, 2010).

Currently the presence of bullfrog in Italy is less

considerable, but reports remain for Lombardy

(Bergamo, Brescia, Cremona and Pavia), Veneto

(Verona and Rovigo), Piedmont (Asti and Torino),

Emilia-Romagna (Bologna, Modena, Ferrara,

Piacenza and Reggio Emilia), Tuscany (Firenze

and Pistoia) and Latium (in the province of Rome:

Maccarese, Torre in Pietra, Pomezia and Tor San

Lorenzo). In the year 2000 it was assumed that the

populations of Latium were extinct (Bagnoli, 2000),

but a subsequent paper confirmed its presence at

Maccarese (Pizzuti Piccoli & Cattaneo, 2008).

MATERIAL AND METHODS

The new site where the American Bullfrog was

detected, is located on the Via Salaria in locality

214

Mauro Grano & Cristina Cattaneo

Semblera, Monterotondo Scalo, about twenty kilo-

meters from Rome. It is a wetland of ten acres that

consists of four ponds adjacent to the left bank of

Tevere river (Fig. 1).

Originally these were clay quarries used by a

brick factory. At the end of the eighties these quar-

ries were abandoned, filled with waste materials

and subsequently with water. Initially, one of these

small lakes were used for sport fishing. Later were

no more used and this has allowed the rooting of

luxuriant vegetation (Fig. 2).

The hydrobiotope arisen is of some interest not

only for the existence of another species of amphi-

bian, Pelophylax klepton hispanicus (Bonaparte,

1839), but also for the significant presence of avi-

fauna that is stationed there.

RESULTS AND CONCLUSIONS

The sightings and detections of the croaking of

L. catesbeianus were carried out in all of the four

lakes. Geographic coordinates are the following:

42°03’52.91”N; 12°35’07.83 ,, E.

It has not been possible to quantify numerically

the presence of this frog (Figs. 3, 4), but it was as-

sumed, through visual and auditory detections, that

the population can be significant. The American

Bullfrog is an alien species considered damaging

for native amphibian populations, both for the large

size it can reach, and for its voracity (Scali, 2010).

L. catesbeianus besides is also a vehicle for the

spread of the dangerous fungus Batrachochytrium

dendrobatidis (Hanselmann et al., 2004; Gamer et

Figure 1. The study area (by Google earth). Figure 2. The study area. Figure 3. Lithobates catesbeianus from Montero-

tondo Scalo (courtesy by Alessandro Crea). Figure 4. L. catesbeianus from Monterotondo Scalo (courtesy by Alessandro

D’Alessio).

A new record for the American Bullfrog, Lithobates catesbeianus (Amphibia Anura Ranidae), near Rome (Italy) 215

al., 2006; Dejean et al., 2010; Ficetola & Scali,

2010), which is one of the most important causes

of rarefaction of numerous species of amphibians

in the world (Blaustein & Kiesecker, 2002; Kats &

Ferrer, 2003).

Recent studies (Ficetola et al., 2008) have

shown that the Italian populations of bullfrog have

originated from a considerably small strain, (less

than six females), thus highlighting the large

expansion capacity and rooting of this invasive

species (Scali, 2010).

Therefore, it is necessary to carry out regular

monitoring activities in order to record in time and

to avoid eventual invasive processes, especially in

the colonization of new sites also due to the strong

impact which may exert on ecology and community

structure of native amphibians (Andreone &

Marocco, 1999; Bologna et al., 2000). The greatest

danger is represented for native populations of

green frog, either as direct prey of L. catesbeianus ,

or as being subject to infection by the fungus B.

dendrobatidis.

Previously it was mentioned that the introduc-

tion of L. catesbeianus in Italy is related exclu-

sively to food purposes. On the contrary, the spread

of bullfrog in Latium is attributable exclusively to

the passive transport by humans. Individuals im-

ported at Maccarese and Torre in Pietra in 1974,

came from Castel d’ Ario (Mantova) place of origin

of the managers of three lakes used for sport fish-

ing in Latium (Bagnoli, 2000). It is surely know

that it was common practice to populate the lakes

used for sport fishing with a mixture of juvenile

fish coming from the areas around Mantova, where

the American bullfrog was certainly present with

well established populations, hence, together with

the fry there may have been many tadpoles of L.

catesbeianus (Andreone, 2005; Ferri, 2006).

This made it possible that this alien species

colonized new areas. Therefore, it could easily be

assumed that the presence of bullfrog in this site

should be related to the previous use of some of

these ponds for the practice of sport fishing.

ACKNOWLEDGMENTS

We are grateful to Augusto Cattaneo for his

invaluable help. We are also thankful to Alessio

Rivola for reporting and clear indications; to Alessan-

dro Crea and Alessandro D’ Alessio for the Ameri-

can bullfrog’s pictures; to anonymous referee.

Addendum

While the present contribution was in printing,

the authors have received a report of the presence

of the American Bullfr og, L. catesbeianus , in a new

site, still in the province of Rome. The new report

refers to some quarries currently filled with water,

located along the Via Flaminia in Civitella San

Paolo. Also these quarries are placed nearby the

Tevere river and are about twenty kilometers from

the old quarries of Monterotondo Scalo.

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In: Andreone F., Gromis di Trana C., Iussich E.,

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degli Anfibi e dei Rettili, Museo Regionale di Scienze

Naturali di Torino, Monografie XXVI, Torino, 192—

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Bagnoli C., 2000. Rana catesbeiana (Shaw, 1802). In:

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Blaustein A.R. & Kiesecker J.M., 2002. Complexity in

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amphibian populations. Ecology Letters, 5: 597-608.

Bologna M.A., Capula M. & Carpaneto G.M., 2000.

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Ficetola G.F., Bonin A. & Miaud C., 2008. Population

genetics reveals origin and number of founders in a

biological invasion. Molecular Ecology, 17: 773-

782.

216

Mauro Grano & Cristina Cattaneo

Ficetola G.F. & Scali S., 2010. Invasive Amphibians and

Reptiles in Italy. In: Di Tizio L., Di Cerbo A.R., Di

Francesco N., Cameli A., 2010. Atti VIII Congresso

Nazionale Societas Herpetologica Italica (Chieti, 22-

26 Settembre 2010), Ianieri Edizioni, Pescara, 335 pp.

Garner T.W.J., Perkins M.W., Govindarajulu P, Seglie

D., Walker S., Cunnigham A.A. & Fischer M.C.,

2006. The emerging amphibian pathogen Batra-

chochytrium dendrobatidis globally infects intro-

duced populations of the North American bullfrog,

Rana catesbeiana. Biology Letters, 2: 455-459.

Hanselmann R., Rodriguez A., Lampo M., Fajardo-

Ramos L., Aguirre A.A., Kilpatrick A.M., Rodriguez

J.P. & Daszak P., 2004. Presence of an emerging

pathogen of amphibians in introduced bullfrogs Rana

catesbeiana in Venezuela. Biological Conservation,

120: 115-119.

Kats L.B. & Ferrer R.P., 2003. Alien predators and

amphibian declines: review of two decades of science

and the transition to conservation. Diversity and

Distributions, 9: 99-110.

Pizzuti Piccoli A. & Cattaneo A., 2008. Rinvenimento di

un esemplare di rana toro, Lithobates catesbeianus

(Shaw, 1802) (Amphibia, Anura, Ranidae), in localita

Maccarese (Roma, Italia). Atti del Museo di Storia

Naturale della Maremma, 22: 119-122.

Scali S., 2010. Le specie alloctone in Italia: censimenti,

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Biodiversity Journal, 2014, 5 (2): 217-220

First record of Pempelia amoenella (Zeller, 1848) for Western

Europe (Lepidoptera Pyralidae)

Stefano Scalercio 1 *, Giuseppe Luzzi 2 & Marco Infusino 3

'Consiglio per la Ricerca e la sperimentazione in Agricoltura, Unita di Ricerca per la Selvicoltura in Ambiente Mediterraneo, c.da

Li Rocchi, 87036 Rende, Cosenza, Italy; e-mail: stefano.scalercio@entecra.it

2 Ente Parco Nazionale della Sila, Via Nazionale, 87055 Lorica San Giovanni in Fiore, Cosenza, Italy; e-mail: g.luzzi@parcosila.it

3 Associazione Palermoscienza, via Cirrincione 41, 90143 Palermo, Italy; e-mail: minfusino@unime.it

"■Corresponding author

ABSTRACT The first record of Pempelia amoenella (Zeller, 1848) (Lepidoptera Pyralidae) for Western

Europe is reported. The species was collected in Southern Italy, on the Ionian coast of Calabria,

where the vegetation is dominated by Tamarix, the known feeding plant of the larvae. Female

genitalia are figured for the first time.

KEY WORDS Mediterranean shrubland; Tamarix', diversity; Calabria.

Received 21.05.2014; accepted 09.06.2014; printed 30.06.2014

INTRODUCTION

Pempelia amoenella (Zeller, 1848) is a species

of Pyralidae belonging to the tribe of Phycitini, sub-

family Phycitinae. Described by Zeller (1848) as

Acrobasis amoenella , was sometimes mentioned as

Salebria amoenella and now is included in the

genus Pempelia (Hiibner, 1825). Mann (1867)

described Pempelia erberi for the island of Corfu,

Greece, but two years later the same author reported

P. erberi as a synonym of P. amoenella.

This species was rarely reported in literature and

very few data on its distribution are published.

Zeller (1848) indicated the European Turkey as the

locus typicus, without further details. Subsequently

in Europe it was collected in Croazia: Southern Dal-

matia (Mann, 1869) and Gravosa (Klimesch, 1942),

Montenegro: Cattaro (Caradja, 1910), Greece:

Corfu Island (Rebel, 1913), Macedonia (Klimesch,

1968), and Russia: Rostov-on-Don Province

(Poltavsky et al., 2009) and Astrakhan region,

Astrakhan Nature Reserve (coll. Tatyana A. Trofi-

mova). It was also generically reported for Albania

and Romania (Karsholt & Razosky, 1996). Outside

of Europe the species was collected inTurkey: Lille

Burgas (Rebel, 1913), Igdir province (Kogak &

Kemal, 2006) and Erzincan (coll. Zoological Mu-

seum, University of Copenhagen), Afghanistan

(Kogak & Kemal, 2012), Uzbekistan (Kogak &

Kemal, 20 12), Turkmenistan (coll. Siberian Zoolog-

ical Museum), Tadzhikistan (coll. Siberian Zoolog-

ical Museum), Kazakistan: Kyzyl-Orda region, Aral

lake, Karatup peninsula (coll. Tatyana A. Trofi-

mova), China: Kashgar, Xinjiang Province (Caradja,

1910) and Mongolia: Hovd Aimak, Mongolian

Altai, Uenchin-Gol Valley, 50 km N Uench (coll.

Tatyana A. Trofimova).

To date the corotype of P amoenella can be de-

fined as Centrasiatic-South East European.

Larvae feed on Tamarix L. (Tamaricaceae).

Mann (1867) observed larvae feeding on Tamarix

sp. where they live in silky structures built around

218

S. Scalercio, G. Luzzi & M. Infusino

the vegetative apex. The same behaviour was ob-

served by Klimesch (1942) who also found whitish

cocoon of this species on small tree branches.

Larvae pupate at the end of May. Adults were

observed to emerge from the end of June to the be-

ginning of July near the coastal line in southern

Croazia (Klimesch, 1942), whilst in Turkey were

collected later during the third week of July at

higher altitude (1200-1300 metres).

The habitat of P. amoenella is in coastal areas

and in salty and arid soils. It is more frequently

recorded at low altitude, especially in dune wood-

lands of coastal habitats, but, in Turkey, its altitudi-

nal range is extended up to 1300 metres.

MATERIAL AND METHODS

A light source was utilised to collect moths dur-

ing the night. Light source was a 160W mercury-

vapour lamp that reflected onto a white vertical

screen. Two operators were assigned to collect the

moths on the screen surface and on the ground

around the lamp.

The collecting site was located on the bed of the

Fiumara Trionto in the municipality of Crosia

(Cosenza), Southern Italy, at 90 metres of altitude

(lat.: 39°33’09”N; long.: 16 0 45’31”E) (Fig.l).

The so-called “fiumare” are streams with large

beds characterised by a torrential regime and devel-

opping primarily along a high altitude gradient, then

having a high erosive and transporting power. In

summertime the bed is usually dry and surface

water appears mainly from late September to late

June. The light source was positioned near to a

small riparian woodland dominated by Tamarix

africana Poir. and Nerium oleander L., mostly as-

sociated to Spartium junceum L., Asparagus acuti-

folius L., Rubus canes cens DC., Crataegus

oxyacantha L., Rosa sempervirens L., Verbascum

sinuatum L., Lagurus ovatus L., Vicia sativa L.,

Arum italicum Mill., Galactites tomentosa Moench,

Dracunculus vulgaris Schott, Artemisia vulgaris L.,

and Trifolium campestre Schreb. Around the small

woodlands and where the soil was stabile from

some years, grows a garrigue characterised by

Helichrysum italicum (Roth) G. Donand, Ephedra

distachya L., otherwise the soil is bare (Fig. 2).

Figure 1. Localisation of the collecting site of Pempelia

amoenella (Zeller, 1848).

Figure 2. Habitat of P amoenella in Italy.

Figure 3. Imago of Pempelia amoenella, Fiumara Trionto,

15.VI.2000, female, wingspan: 18 mm, coll. Unita di Ri-

cerca per la Selvicoltura in Ambiente Mediterraneo.

First record of Pempelia amoenella (Zeller, 1848) for Western Europe (Lepidoptera Pyralidae)

219

Figures 4-6. Female genitalia of P. amoenella. Fig. 4: general view (gen. praep. CRASAM-012). Fig. 5: details of papillae

anales. Fig. 6: details of comuti on the bursa copulatrix.

RESULTS AND DISCUSSION

One female of P. amoenella was collected at light

on 15 June 2000 (Fig. 3). This is the first record for

Italy and Western Europe of this interesting species.

During the night of sampling, mean temperature was

of 24°C, humidity rate of 75%, with no wind, the

40% of the moon surface was illuminated and the

sky was variably clouded. The sampling session

started at 9: 10 PM and lasted four hours.

Female genitalia are figured for the first time

(Figs. 4-6). The specimen and its genitalia (gen.

praep. CRASAM-012) are conserved in the collec-

tion of the Unita di Ricerca per la Selvicoltura in

AmbienteMediterraneo (CRA-SAM).

In Italy the habitat of P. amoenella is similar to

that of localities where the species was previously

collected on the eastern coastal areas of Adriatic

and Ionian seas, whilst phenology appears to be

anticipated due to the higher mean annual temper-

ature in the new discovered range. In fact, our

adult female is the earliest known capture for this

species.

220

S. Scalercio, G. Luzzi & M. Infusino

The presence of trans-ionic species in Southern

Italy is not a novelty. In fact, among macrolepi-

doptera at least 6 species of the Calabrian fauna

have a similar range, namely Oiketicoides lutea

(Staudinger, 1870) (Psychidae), Anthocaris damone

Boisduval, 1836 (Pieridae), Idaea determinate i

(Staudinger, 1876) (Geometridae), Aegle agatha

(Staudinger, 1861) (Noctuidae), Tiliacea cypreago

(Hampson, 1906) (Noctuidae) and Conistra ra-

gnsae (Failla-Tedaldi, 1890) (Noctuidae).

Further investigation along Ionian coastal areas

of Calabria can probably provide more detailed

information on the biology of P. amoenella.

ACKNOWLEDGMENTS

Many thanks to Tatyana A. Trofimova (Samara

State University, Russia) for providing distribu-

tional data from her private collection.

REFERENCES

Caradja A., 1910. Beitragzur Kenntnisuber die geographis-

che Verbreitung der Pyraliden des europaischen-

Faunen gebietesnebst Beschreibungeinigemeuer For-

men. Deutsche Entomologische Zeitschrift Iris, 24:

105-147.

Karsholt O. & Razowski J., 1996. The Lepidoptera of

Europe. Apollo Books, Stenstrupp, 380 pp.

Klimesch J., 1942. Uber Microlepidopteren-Ausbeuten

von Zatonbei Gravosa (Suddalmatien). Mitteilungen

Muenchener Entomologischen Gesellschaft, 32: 347-

399.

Klimesch J., 1968. Die Lepidopterenfauna Mazedoniens.

IV. Microlepidoptera. Posebno Izdanie Prirodonaucen

Muzej Skopje, 5: 1-201.

Kogak A.O. & Kemal M., 2006. Checklist of the Lepi-

doptera of Turkey. Priamus, suppl., 1: 1-196.

Kogak A.O. & Kemal M., 2012. Lepidoptera of Afghanistan.

Priamus, suppl., 26: 1-134.

Mann J., 1867. Zehnneue Schmetterlingsarten erhan-

dllungen der kaiserlich-kongiglichen zoologish-

botanischen Gesellschaft in Wien, 1867: 845-852.

Mann J., 1869. Lepidopteren, gesammeltwahrenddreier-

Reisennach Dalmatien in den Jahren 1 850, 1 862 und

1868. Verhandllungen der kaiserlich-kongiglichen

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371-388.

Poltavsky A.N., Artokhin K.S. & Silkin Y.A., 2009. To

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Province. Eversmannia, 17-18: 57-70.

Rebel H., 1913. Studien ber die Lepidopterenfaun der

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seums in Wien, 27: 281-334.

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(9) 641-691,(10) 721-754.

Biodiversity Journal, 2014, 5 (2): 221-224

An innovative, low-cost, small-scale rearing method for green

lacewings (Neuroptera Chrysopidae)

Laura Loru 1 , Xenia Fois 1 , Saminathan Vangily Ramasani 2 , Leonarda M. Fadda 1 & Roberto A. Pantaleoni 1,3,

'Istituto per lo Studio degli Ecosistemi, Consiglio Nazionale delle Ricerche, Sassari, Italy

2 Department of Crop Protection, AD AC & RI, Tamil Nadu Agricultural University, India

3 Dipartimento di Agraria, Universita degli Studi di Sassari, Italy

’Corresponding author, e-mail: pantaleo@uniss.it

ABSTRACT In this paper we describe an innovative, low-cost, small-scale green lacewing (Neuroptera

Chrysopidae) rearing method developed in our laboratories over a decade. The main simplifi-

cations of our method are represented by the replacement of a yeast-fructose liquid diet for

adults with bee pollen loads and by the use of Tenebrio molitor Linnaeus, 1758 larvae (Coleoptera

Tenebrionidae) as factitious prey for larvae. Moreover almost all the components of the rearing

cages derive from common cheap materials which can be easily assembled by anybody. Our

method proves to be adaptable from a small laboratory to a local farmer’s insectary and its

innovative aspects could be adopted in (and/or adapted to) mass rearing systems.

KEY WORDS bee pollen loads; mealworm beetle; factitious prey; biological control; beneficial insects.

Received 07.06.2014; accepted 18.06.2014; printed 30.06.2014

INTRODUCTION

Green lacewings (Neuroptera Chrysopidae)

have long been recognized as effective biological

control agents of a wide variety of arthropod pests,

but their use has been directed almost exclusively

toward the augmentative method for years (Canard

et al., 1984; McEwan et al., 2001). So a method of

mass-rearing was developed and a few species of

the genus Chrysoperla Steinmann, 1964 became

commercially available during the 1990s in the

USA, Europe and China (Wang & Nordlund, 1994;

Hunter, 1997). However, despite many promis-

ing laboratory tests and several instances of suc-

cessful field releases, failures often occurred prob-

ably due to the lack of integration between

research and commercial development (Tauber et

al., 2000).

In the last decade, the amount of research on

lacewing mass-rearing techniques has been station-

ary or even in decline and this is linked to the trend

of augmentative methods in biological control

(Warner & Getz, 2008; van Lenteren, 2012). So

reading the recent review of Pappas et al. (2011)

on the role of green lacewings in biological control,

we did not find any true innovations compared to

similar papers written more than 10 years ago

(Tauber et al., 2000).

Thirty five years ago, one of us (RAP) began to

rear green lacewings on a very small-scale for

taxonomic purposes (e. g. in order to obtain all the

stages of a given species or the adults from field

collected larvae). Working on the staff of Maria

Matilde Principi at the Alma Mater Studiorum, Uni-

versity of Bologna, Italy, his starting point was the

method described in Pasqualini (1975). This method

222

Laura Loru et alii

was subsequently modified in small steps in order

to make it less laborious, less expensive and to save

time. Recently, in the laboratory of the ISE-CNR

Sassari, Italy, other changes have been made so that

it can be used on a laboratory- sc ale for experimen-

tal purposes.

In this paper we want to describe our practices

in lacewing rearing because:

i) the well-known large-scale technology is

often not suitable when producing lacewings for

laboratory purposes and very few papers describe

small-scale rearing methods (McEwen et al., 1999);

ii) our method proves to be easily adaptable

from a small laboratory to a local farmer’s insectary

being a simple, low-cost technique;

iii) its innovative aspects could be adopted in

(and/or adapted to) mass rearing systems.

ADULT REARING

Over the years we have reared many Euro-

Mediterranean species with glyciphagous adults, scil-

icet pollen-and-nectar-feeders (Pantaleoni, 2014),

excluding the genus Chiysopa Leach in Brewster,

1815 which is predaceous. We have never had prob-

lems obtaining a new generation from gravid wild

females, very few species had difficulty in mating

and laying eggs. All of the Chrysoperla and almost

all of the Pseudomallada Tsukaguchi, 1 995 species

that we have dealt with have been reared continu-

ously for several generations, sometimes for years.

For laboratory purposes, specimens were main-

tained in a climatic chamber with 20 ± 2°C Tem-

perature, 65 ± 5% Relative Humidity, and a 16:8 h

Light: Dark Photoperiod. Among the rearing condi-

tions the key factor was the photoperiod, at Mediter-

ranean latitude almost all species enter in diapause

under short days.

The adult rearing unit is a cylinder, 1 00 mm in

height and 80 mm in diameter open at both ends,

obtained from a plastic water bottle. The inner part

of the cylinder was lined with type “Bristol” yellow

cardboard, by rolling up a 100 per 300mm rectangle

fixed by two clips. Both bases were closed with

square tulle mesh (1.4 mm openings), about 150

mm inside, secured with rubber bands. The cages

were kept vertically resting on a base. A water dis-

penser, containing cotton wool with a plastic cover

(half petri dish), was placed on the tulle netting at

the top of the container. The unit was put on a tray

covered with paper towels to absorb excrement.

Food consisted of honeybee pollen loads.

About fifty pollen loads, weighing more or less

300 mg, were put on the bottom of the cage. Water

was supplied by dampening the cotton in the dis-

penser, making sure that the water did not drip

down. Every unit hosted no more than four lacewing

pairs. The females laid eggs both on the paper and

on the tulle netting.

The maintenance of the adult rearing unit took

place twice a week. Eggs were isolated as necessary

in order to obtain larvae both for experimental pur-

poses and to renovate the laboratory colony. The

egg surplus had to be destroyed in order to prevent

hatching and subsequent attacks of larvae against

adults. With the same interval of time, twice a week,

the pollen was renewed and the cotton refilled with

water. The cotton, tulle netting and paper towels

were substituted at regular but longer intervals.

An experienced technician can easily manipu-

late the adults by transferring them into a glass tube

during rearing unit maintenance, otherwise, with

caution, it is possible to anaesthetise them (Loru et

al., 2010).

LARVAE REARING

Larvae were reared individually in order to

avoid cannibalism. We used transparent, plastic,

cylindrical containers both 25mm in height and in

diameter with a plastic lid. The eggs were isolated

by cutting the cardboard or tulle on which they had

been laid and were put singularly into the containers

using tweezers. As food we used mealworm beetle

larvae (the Coleoptera Tenebrionidae Tenebrio

molitor Linnaeus, 1758) previously killed with

ethyl acetate. Also a drop of fructose solution was

provided to the newly-hatched larvae (Pantaleoni,

2014). Different lacewing larvae instars were fed

on mealworms of the appropriate size, in particular

4-5 mm long and 0.5-1 mg in weight vs 1st instar,

6-8 mm long and 2-4 mg in weight vs 2nd instar,

9-12 mm long and 6-12 mg vs 3rd instar. Never-

theless an expert technician is perfectly able to sort

them by eye. Three mealworms were given to each

lacewing larva twice a week and every time residues

of the previous meal were eliminated. Initially a

An innovative, low-cost, small-scale rearing method for green lacewings (Neuroptera Chrysopidae)

223

little piece of paper towel was put into the container

in order to absorb exudates from dead mealworms

and offer a suitable support during cocooning.

When the cocoon was about one week old, the

residues of its last meal were eliminated. At the same

time a 30mm high piece of “Bristol” cardboard was

put into the container in order to provide support for

pharate adults and facilitate adult emergence. Adults

which had just emerged were sexed and put into the

container described above.

DISCUSSION

Our adult rearing unit (Fig. 1) can be easily as-

sembled by anybody. All the components derive

from common cheap materials and a few simple

tools such as scissors, a paper cutter, a ruler, a set

square and a pencil are needed. Only the cap of

the water dispenser is a specialized laboratory

item (half of a petri dish), but this is also easily

replaceable. Neither individual containers nor

possible communal cages for larvae have been de-

veloped although there are many opportunities for

improvement.

The replacement of a liquid meridic diet for

adults with a solid oligidic one, such as pollen

loads, is the main simplification of our method. Ac-

cording to our experience the management of a

liquid yeast-fructose diet is laborious. This diet

requires frequent cleaning or changing of the dis-

penser. It is both rapidly perishable and filthy, in-

sects often get themselves dirty becoming incapable

of flying or even moving. On the contrary pollen

loads are clean, easy to give and to remove and

more durable. In nature adult lacewings are essen-

tially pollen-feeders and pollen, as stated by

Nordlund et al. (2001), has good nutritional qualities.

Moreover adult lacewings harbour mutualistic

yeasts ( Turolopsis sp.) that synthesize essential

amino acids which are missing from their diet

(Hagen et al., 1970). The pollen loads contain these

yeasts, collected in the environment by bees (Gilliam,

1979). The slight disadvantage of using pollen is

the dramatic difference in quality of the various

kinds of pollen derived from different plants

(Nordlund et al., 2001).

Just as the pollen-feeder habit of adults drove us

towards our choice of pollen loads, the occasional

scavenger habits of lacewing larvae (repeatedly

Figure 1 . Adult rearing unit.

observed in nature by one of us (RAP), made us

choose the killed mealworms. The use of T. molitor

as factitious prey for lacewings was reported for the

first time at the IX International Symposium on

Neuropterology held in Ferrara, Italy, in 2005, but

it remained unpublished. Since then, only Pappas et

al. (2007) tested mealworms as a prey for lacewings,

but using “second instar mechanically injured”

instead of killed larvae of a larger size. Although

the aim of the paper was completely different, and

the use of T. molitor as factitious prey was not even

cited, Lorn et al. (2010) published the same kind

of data. Pappas et al. (2007), rearing a species of

Pseudomallada (then Dichochrysa Yang, 1991),

found a short lifespan and low female fertility in

specimens fed with mealworms. Lorn et al. (2010),

rearing a species of Chtysoperla, found a much longer

lifespan (max 120 days vs max 45 days) and a higher

fecundity (recalculated in order to compare the two

papers) (about 650 eggs/female vs 192).

The great advantages in the use of mealworms

as factitious prey are the minimal physical space re-

quired for their rearing, their high conversion effi-

ciency, their potential for massive production and

the chance to use organic waste materials as a food

source (Ramos -Elorduy et al., 2002).

The future of biological control in agriculture

will be played in two different arenas: the enhance-

ment of “big” technology (biotechnology, industry,

worldwide market) to apply to large scale produc-

224

Laura Loru et alii

tions, and the increase of “sustainable” technology

to apply to local development policies. In both

cases, the lacewings will be able to give a little help

as beneficial through their mass production or their

conservation by means of habitat management, but

in any case we should be able to rear them.

ACKNOWLEDGEMENTS

This work has been financed by Regione Au-

tonoma della Sardegna - L.R. 7 Agosto 2007, n. 7,

project COSMESAGRO “Progettazione, preparazione

e studio di inibitori ecosostenibili dell'attivita di

fenol- e polifenolossidasi sul controllo di melanine

di interesse nei settori cosmetologico e fitoiatrico”.

Grateful thanks to Marcella Fara for taking the photo.

REFERENCES

Canard M., Semeria Y. & New T.R., 1984. Biology of

Chrysopidae. Dr. W. Junk Publishers, The Hague,

294 pp.

Gilliam M., 1979. Microbiology of pollen and bee bread:

the yeasts. Apidologie, 10: 43-53.

Hagen K.S., Tassan R.L. & Sawall E.F., 1970. Some eco-

physiological relationships between certain Chysopa

(Neuroptera, Chrysopidae), honeydews and yeasts.

Bollettino del Laboratorio di Entomologia agraria

“Filippo Silvestri”, 28: 113-134.

Hunter C., 1997. Suppliers of Beneficial Organisms in

North America. California Environmental Protection

Agency, Department of Pesticide Regulation, Envi-

ronmental Monitoring and Pest Management Branch,

1997 Edition, 35 pp.

Loru L., Sassu A., Fois X. & Pantaleoni R.A., 2010.

Ethyl acetate: a possible alternative for anaesthetizing

insects. Annales de la Societe Entomologique de

France, 46: 422-424.

McEwen P., New T.R. & Whittington A.E., 2001. Lacewings

in the Crop Environment. Cambridge University

Press, Cambridge, 546 pp.

McEwen P., Kidd N.A.C., Bailey E. & Eccleston L.,

1999. Small-scale production of the common green

lacewing Chrysoperla carnea (Stephens) (Neuroptera,

Chrysopidae): minimizing costs and maximizing

output. Journal of Applied Entomology, 123: 303-306.

Nordlund D.A., Cohen A.C. & Smith R.A., 2001.

Mass-Rearing, Release Techniques and Augmenta-

tion. In: McEwen P.K, T.R New and A.E. Whitting-

ton 2001. Lacewings in the Crop Environment.

Cambridge University Press, Cambridge, pp. 303-319.

Pantaleoni R.A., 2014. Sweeten our crops. Sustain a high

diversity of beneficials through “sweet food” from

plants. Biodiversity Success Stories, 2: 18.

Pappas M.L., Broufas G.D. & Koveos D.S., 2007. Effects

of various prey species on development, survival and

reproduction of the predatory lacewing Dichochiysa

prasina (Neuroptera: Chrysopidae). Biological

Control, 43: 163-170.

Pappas M.L., Broufas G.D. & Koveos D.S., 2011.

Chrysopid Predators and their Role in Biological

Control. Journal of Entomology, 8: 301-326.

Pasqualini E., 1975. Prove di allevamento in ambiente

condizionato di Chrysoperla carnea Steph. (Neu-

roptera, Chrysopidae). Bollettino delflstituto di Ento-

mologia della Universita degli Studi di Bologna, 32:

291-304.

Ramos-Elorduy J., Gonzalez E.A, Hernandez A.R. & Pino

J.M., 2002. Use of Tenebrio molitor (Coleoptera:

Tenebrionidae) to Recycle Organic Wastes and as

Feed for Broiler Chickens. Journal of Economic En-

tomology, 95: 214-220.

Tauber M.J., Tauber C.A., Daane K.M. & Hagen K.S.,

2000. Commercialization of predators: recent lessons

from green lacewings (Neuroptera: Chrysopidae).

American Entomologist, 46: 26-37.

van Lenteren J.C., 2012. The state of commercial aug-

mentative biological control: plenty of natural ene-

mies, but a frustrating lack of uptake. Bio Control,

57: 1-20

Wang R. & Nordlund D.A., 1994. Use of Chrysoperla

spp. (Neuroptera: Chrysopidae) in augmentative

release programmes for control of arthropod pests.

Biocontrol News and Information, 15: 51N-57N.

Warner K.D. & Getz C., 2008. A socio-economic analysis

of the North American commercial natural enemy

industry and implications for augmentative biological

control. Biological Control, 45: 1-10

Biodiversity Journal, 2014, 5 (2): 225-228

Monograph

Preface

Studies on extant and fossils astridypeids

(Echinoidea Clypeasteroida)

Paolo Stara

Centro Studi di Storia Naturale del Mediterraneo, c/o Museo di Storia Naturale Aquilegia, Via Italia 63 Cagliari-Pirri and Geomuseo

Monte Arci, Masullas, Oristano, Sardinia, Italy; e-mail: paolostara@yahoo.it

Received 25.06.2013; accepted 30.05.2014; printed 30.06.2014

In: Paolo Stara (ed.). Studies on some astriclypeids (Echinoidea Clypeasteroida), pp. 225-358

This monographic volume is the result of the

need to clarify the diagnostic characters that really

distinguish species and genera belonging to this in-

teresting family of clypeasteroids.

In particular, during the research on the Oligo-

Miocene species of Amphiope L. Agassiz, 1840

present in numerous outcrops of Sardinia (Stara et

al., 2012; Stara & Borghi, 2014) have been reported

several difficulties in specific distinction, when was

using only the set of morphological and morpho-

metric data normally used in the past (see Philippe,

1998).

As already observed by Durham (1955), almost

all of the authors prior to his monograph on

clypeasteroids were limited to the description of a

few morphological and/or morphometric data, such

as length, width and height of the test, petal-length

and distance stoma or periproct from the anterior

or posterior margin; all data disconnected from the

plate pattern of the shell and often described by

adjectives.

This practice, unfortunately, left many uncer-

tainties, as is clear from the discussion that lasted

for over a century (see Stara & D. Fois, 2014, and

references therein). In fact much has been dis-

cussed on several morphotypes belonging to the

genus Amphiope rather than Echinodiscus Leske,

1778, only on the basis of shape of their lunules.

The discussion, in fact, concerned about the use-

fulness of the shape of the two posterior lunules

(ellipsoidal elongated along the axis of the rear

ambulacra, or rounded to ellipsoidal transverse to

the rear ambulacra) in the diagnostic applied to the

systematic.

Philippe (1998), studying th q Amphiope popu-

lations from the Rhone Basin of South-Eastern

France, and highlighting the great variability of

shape and size of the lunules in the examined in-

dividuals, placed in synonymy with Amphiope

bioculata des Moulins, 1837, all nominal species

previously established in its and in other peri-

Mediterranean regions (except Amphiope boulei

Cottreau, 1914).

To make matters worse, at the current state of

historical research, several specimens used by the

authors as type-species, are nowhere to be found,

poorly defined and with stratigraphic data absent if

not conflicting (for Amphiope, for example, see des

Moulins, 1835-37; L. Agassiz, 1838-41; Cottreau,

1914, Philippe, 1998).

The generic distinction, however, is made easy

since Durham (1955) published the plate patterns

226

Paolo Stara

of two specimens that have been a type-function,

widely diffused by Smith & Kroh (2011) and used

by various authors, such as eg. Jansen & Mooi,

2011.

As admitted by Philippe (1998), given the many

uncertainties arose because of the supposed wide

variability of lunules in Amphiope , it would be

necessaiy to examine some sample of extant Echin-

odiscus, the genus closer to Amphiope. From this

correct observation we started to plan the work that

led to the publication of this monograph.

One of the main tools used in this study was

the examination of the plate pattern and internal

structure; to determine the usefulness and reliabil-

ity were examined more than 100 samples of Am-

phiope from different Sardinian's sites, including

more than 40 samples from a single locality (A.

lovisatoi Cotteau, 1895) (Stara et al., 2012; Stara

& Borghi, 2014) and more than 60 extant and fos-

sils "E chino discus" from many other locality

(Stara & Sanciu; Stara & M. Fois, 2014).

In particular, we have examined the plate pat-

tern by more than 30 samples of "Echinodiscus cf.

auritus" (Stara & M. Fois, 2014) from Mangili,

Province of Tulear (Madagascar). Of these, the

plates are numbered and have performed the neces-

sary checks of the stability of the encountered

characters.

The result of the research summarized in this

monographic volume has exceeded all expectations

and has allowed us to develop the tools to be used

for generic and specific distinction of echinoids

belonging to this family.

Meanwhile, it became clear that the variability

of the lunules was not the real problem, since Stara

& Borghi e Stara & Sanciu (2014) were able to dif-

ferentiate between different species (some of them

with a very high variability of lunules) of Sardinia

and many other locations.

Overcoming these issues is also fundamental to

achieve one of our main goals: to understand what

were the relationships that these populations have

had with the congeners of other regions of the Proto

Western Mediterranean (Stara & Rizzo, 2013; Stara

& Rizzo, 2014) .

Now we can propose, as a main tool for descrip-

tion of recent and fossils echinoids, analysis of the

plate pattern of the test and in particular those of the

oral interambulacrum 5 and oral / aboral ambulacra

I and IV.

In the case of fossils from different geological

epochs, with the same plate pattern, is also pro-

posed the analysis of the internal structure, since

that, as observed by Stara & Borghi (2014) with the

elapse of geological times, the structure shows

significant changes.

The trend shown by the sample of Sardinia

(over 100 specimens of Amphiope) indicates a

progressive reduction of the plates number and a

lightening of the structure of the internal supports

system.

The introduction of simple indices used for the

recognition of the shape (Shape Index) and the

size of lunules (Width Index) in Amphiope , as

done by Stara & Sanciu (2014), for example, al-

lowed to further differentiate groups of popula-

tions apparently similar. The use of other data

before overlooked such as the measure within the

ambitus of the interambulacrum 5 (Width at Am-

bitus) and the overall length of petalodium

(Petalodium Length) facilitated further discrimina-

tion between genera and species. Finally, when the

number of samples available makes it possible,

can not miss the statistical analysis, as is done by

Stara & Borghi (2014).

Other characters, such as the difference in the

shape and size of pedicellaria are certainly impor-

tant in supporting the distinction between species

and varieties, but never separately from the analysis

of the characters previously underlined. The use of

these tools has made possible the distinction of two

new genera and two new species within the family

Astriclypeidae Stefanini, 1912, and has allowed us

to lay the basis for the recognition of further differ-

entiation.

It was possible to achieve this work, thanks to

the availability of the web. The rapid access to

relevant documents, before traceable only in few

and far libraries; the ability to instantly contact

other researchers around the world and to get such

important information in real time; the possibility

of obtaining original photos of animals and places

in which examine the characteristics otherwise

unreachable and geographic data such as, eg.,

topography, vegetation type, type of coasts, alti-

tude of the mountains, has made it possible to

multiply a hundredfold the potential at our dis-

posal. And has certainly facilitated the realization

of this work.

Studies on extant and fossils astriclypeids (Echinoidea, Clypeasteroida)

227

Summary

Stara P. - Preface. Studies on extant and fossils

astriclypeids (Echinoidea Clypeasteroida): 225-228

Stara P. & Fois D. - Dispute about Echinodiscus

Leske, 1778 and Amphiope Agassiz, 1840 (Echinoi-

dea Astriclypeidae): 229-232

Stara P. & Rizzo R. - Paleogeography and dif-

fusion of astriclypeids (Echinoidea Clypeasteroida)

from Proto-Mediterranean basins: 233-244

Stara P. & Borghi E. - The echinoid genus Am-

phiope L. Agassiz, 1 840 (Echinoidea Astriclypei-

dae) in the Miocene of Sardinia: 245-268

Stara P. & Fois M. - Analysis on a sample of

Echinodiscus cf. auritus Leske, 1778 (Echinoidea,

Clypeasteroida): 269-290

Stara P. & Sanciu L. - Analysis of some astri-

clypeids (Echinoidea Clypeasteroida): 291-358

ACKNOWLEDGEMENTS

A heartfelt thanks to Enrico Borghi, of the

Societa di Scienze Naturali of Reggio Emilia, for

the support given to the overall success of this

volume. Thanks to the reviewers and in particular

to Andreas Kroh, of the Naturhistorisches Museum

of Vienna, for the advice dished out in compilation

of the work of Amphiope and for the patient revi-

sion that has allowed all of us to achieve unex-

pected results. I also thank on behalf of the entire

workgroup, institutions (museums and research

centers) that have allowed and encouraged this

research by allowing access to their fine collections

or use of your important data on their collections

or specimens: Museo di Paleontologia "D. Lo-

visato “, Dipartimento di Chimica e Geologia and

Dipartimento di Biologia animale ed Ecologia,

University of Cagliari; Museo Comunale di Storia

Naturale "G. Doria” in Genoa, and Dipartimento

del Territorio e delle sue Risorse, University of

Genoa; NHMUK, London; Natural History Mu-

seum of Denmark (Zoology), Copenhagen; PMBC

of Phuket (Thailand).

REFERENCES

Agassiz L., 1841. Monographic d'echinodermes

vivants et fossiles. Echinites. Famille des Clypeasteroides.

Seconde Monographic. Des Scutelles. Neuchatel: 149 pp.

Barbera C. & Tavernier A., 1989. II Miocene dei

dintomi di Baselice (Benevento) significato paleoecolo-

gico e paleogeografico. Atti 3° Simposio di Ecologia e

Paleoecologia delle comunita bentoniche. Taonnina, 12-

lb ottobre 1985. 1. de Geronimo (Ed.). 745-757.

Comaschi Caria I., 1972. Gli echinidi del Miocene della

Sardegna, Stabilimento Tipografico Edizioni Fossataro

S.p.A. Ed., Cagliari, 95 pp.

Cottreau J., 1914. Les echinides neogenes du Bassin

mediterraneen. Annales de l’lnstitut Oceanographique,

Monaco, 6: 1-193.

des Moulins C., 1837. Troisieme Memoire sur les

echinides. Synonymie general. Actes Societe Linneenne,

Bordeaux: 9: 45-364.

Durham J.W., 1955. Classification of clypeasteroid

echinoids. University of California Publications in Geo-

logical Sciences, 31: 73-198.

Jansen N & Mooi R., 2011. The Astriclypeidae: Phy-

logenetics of Indo-Pacific, super- flat, holey sand dollars.

Meeting abstract in: Society for Integrative and Compar-

ative Biology, 2011 Annual Meeting. Salt Lake City, UT,

USA.

Smith A.B. & Kroh A., 2011. The Echinoid Directory.

World Wide Web electronic publication.

http://www.nhm.ac.uk/scienceprojects/echinoids (ac-

cessed September 2013).

Stara P., Rizzo R., Sanciu L. & Fois D., 2012. Note

di geologia e paleoecologia relative ad alcuni siti ad Am-

phiope (Echinoidea: Clypeasteroidea) in Sardegna, Parva

Naturalia (2010-2011), 9: 121-171.

Stara P. & Rizzo R., 2013. Diffusion of Amphiope

Agassiz, 1 840 (Astriclypeidae, Clypeasteroida) from the

Western proto-Mediterranean Sea, towards the Eastern

Neotethys, XIII Giomate di Paleontologia. Perugia, May

23-25, 2013, Riassunti: 119-120, sessione poster.

Stara P. & Borghi E., 2014. The echinoid genus Am-

phiope L. Agassiz, 1840 (Echinoidea Astriclypeidae) in

the Oligo-Miocene of Sardinia (Italy). In: Paolo Stara

(ed.). Studies on some astriclypeids (Echinoidea Clypea-

steroida), pp. 225-358. Biodiversity Journal, 5: 245-268.

Stara P. & Fois D., 2014. Dispute about Echinodiscus

Leske, 1778 and Amphiope L. Agassiz, 1840 (Echinoidea

Astriclypeidae). In: Paolo Stara (ed.). Studies on some

astriclypeids (Echinoidea Clypeasteroida), pp. 225-358.

Biodiversity Journal, 5: 229-232.

Stara P. & Fois M. 2014. Analysis on a sample of

Echinodiscus cf. auritus Leske, 1778 (Echinoidea Cly-

peasteroida). In: Paolo Stara (ed.). Studies on some astri-

clypeids (Echinoidea Clypeasteroida), pp. 225-358.

Biodiversity Journal, 5: 269-290.

228

Paolo Stara

Stara P. & Rizzo R., 2014. Paleogeography and dif-

fusion of astriclypeids (Echinoidea Clypeasteroida) from

Proto-Mediterranean basins. In: Paolo Stara (ed.). Studies

on some astriclypeids (Echinoidea Clypeasteroida), pp.

225-358. Biodiversity Journal, 5: 233-244.

Philippe M., 1998. Les echinides miocenes du Bassin

du Rhone: revision systematique. Nouvelles Archives du

Museum d’HistoireNaturelle de Lyon, 36: 3-241, 249-441.

Biodiversity Journal, 2014, 5 (2): 229-232

Monograph

Dispute about Echinodiscus Leske, 1 778 and Amphiope L. Agas-

siz, 1840 (EchinoideaAstridypeidae)

Paolo Stara 1 & Daniele Fois 2

'Centro Studi di Storia Naturale del Mediterraneo, Museo di Storia Naturale Aquilegia, Via Italia 63, Cagliari-Pirri and Geomuseo

Monte Arci, Masullas, Oristano, Sardinia, Italy; e-mail: paolostara@yahoo.it

Museo di Storia Naturale Aquilegia, Via Italia 63, Cagliari-Pirri, Sardinia, Italy, e-mail: daniele.fois@tiscali.it

ABSTRACT Between the late 1800s and early 1900s, some European echinologists gave rise to a dispute

over belonging to the genus Amphiope Agassiz, 1 840, rather than Echinodiscus Leske 1778,

of some lunulate scutelliforms present in the Oligocene-Miocene deposits of France and

Italy. The problem has never been resolved, due to the fact that these echinologists consid-

ered the similarities or differences in shape, rather than structural ones. One of the nodes

of the dispute was the variability in shape and size of the lunules in Amphiope. Because of

all these problems, and also because of the impossibility to obtain and examine the struc-

tures of some type specimens of several species established in the past, the recognition of

new species is very complicated and research carried out so far, in many cases is doubtful

or controversial.

KEY WORDS Amphiope', Echinodiscus', lunules variability.

Received 25.06.2013; accepted 30.05.2014; printed 30.06.2014

In: Paolo Stara (ed.). Studies on some astriclypeids (Echinoidea Clypeasteroida), pp. 225-358

INTRODUCTION

The problems that have constituted the nodes of

the dispute which we summarize here, have been

the starting point for the studies carried out by dif-

ferent authors (see Stara, 2014). In particular, the

frequent lack of references relating to the structural

characteristics of Amphiope L. Agassiz, 1840 and

Echinodiscus Leske, 1778, so far established, and

the uncertainties due to the impossibility of com-

paring the type specimens of these species, have

greatly complicated these studies.

It should be said, however, that L. Agassiz

(1838-41), for example, had already meticulously

illustrated the complete plating of the two faces of

a specimen of Lobophora aurita ( Echinodiscus au-

ritus Leske, 1778) and that Loven (1872) published

an important work on the structure of echinoids.

With regards to the clypeasteroids, in particular,

Durham (1955) systematically had reproduced pat-

terns of the plates (plating) of a large number of

species, including those of Echinodiscus bisperfo-

ratus Leske (1887) and Amphiope bioculata (not

des Moulins 1837 type). Later, however, except in

rare cases (Kroh, 2005; Pereira, 2010) no one, to

our knowledge, reported platings of several other

nominal species belonging to the family Astriclypei-

dae Stefanini (1912).

To try to end the dispute which we summarize

here, other authors (Stara & Rizzo, 2014; Stara &

Fois M., 2014; Stara & Sanciu, 2014) proposed a

review of the main characters of some species be-

longing to this family (Astriclypeidae), using the

plate pattern of their tests, considering this the

main tool for the specific and generic diagnosis in

echinology.

230

Paolo Stara & Daniele Fois

THE DISPUTE

Shape of lunules, uncertainly of the generic

attribution and phylogenesis

Cottreau (1914), describing Amphiope boulei

Cottreau, 1914 from the Aquitanian of Carry

(Bouches-du-Rhone, France), stated that the axial

lunules are a primitive morphological character in

Amphiope. This primitive morphological character

is already present in some Oligocene species, such

as Amphiope agassizi des Moulins (1837) from the

Asterias-limestone of the Bordeaux Region, A.

pedemontana Airaghi, 1899, from Piedmont and

Liguria (Airaghi, 1899) and A. duff Gregory, 1911,

from Cyrenaica. According to this author, the axial

arrangement of the lunules persist in present-day

species of Tretodiscus [currently considered as syn-

onymous to Echinodiscus (Kroh, 2012)] bearing

elongated lunules or slits notches on the posterior

margin. These would be derived from Oligocene

species of Amphiope , and they were considered the

true Echinodiscus by Stefanini (1912).

According to Cottreau, also A. fuchsi Fourtau

(1901), from the Middle Miocene of Siwa (Siouah),

Egypt, was an Amphiope, as well as A boulei, while

"Amphiope" bearing elongated lunules was not the

typical form. As evidence of the kinship existing

between Amphiope and Echinodiscus, Cottreau ob-

served that juvenile individuals in Amphiope bioc-

ulata des Moulins, 1837 (the type species of this

genus) often bear pear-shaped elongated lunules

along the axis of the rear ambulacra. According to

this author, A. cherichirensis Gauthier, 1957, from

Tunisia and A truncata Fuchs (1883) from Middle

Miocene of Egypt, can be derived from the Oligocene

European "Amphiope" bearing axial lunules. These

would be derived from Middle Miocene Indian

echinoids, such as A. placenta Duncan, 1885, A.

desori Duncan et Sladen 1883, A. duncani Lambert,

1 907 and from Japanese ones, such as Echinodiscus

formosus (Yoshiwara, 1901). The latter one would

be the true ancestor of Tretodiscus ( Echinodiscus ),

which has slits and is typical of the Indian Ocean,

where it is represented by Tretodiscus elongatus

Duncan et Sladen, 1883 and E. bifissus Agassiz

( Lobophora ) 1840, the latter one corresponding to

the living Echinodiscus auritus Leslce, 1778 with

open slits on the back edge. Comparing the internal

structure of "Amphiope" bearing axial lunules with

that of the typical Amphiope (A. bioculata des

Moulins, 1835), Cottreau discovered that the two

structures were identical, but unfortunately, he did

not publish detailed descriptions of them.

Lambert (1915) wrote "Cottreau considered Am-

phiope agassizi as a very particular form, that joins

very closely Amphiope to Lobophora, (actually

Echinodiscus), and he proposed the suppression of

the latter genus. (...). The Amphiope (morpho) type

appeared in the Middle-Early Oligocene with A.

pedemontana, and it bearing elongated lunules in

the direction of the ambulacral axis, and retained

this character in a series of successive species: A.

agassizi in the Stampian (Middle Oligocene), A.

cherichirensis and A. baquiei in the Burdigalian and

A. truncata in the Early-Middle Miocene. Two

branches detached from this main trunk: the first

one in Aquitaine (France) during the Aquitanian,

with A. ovalifora and the series of closely related

Burdigalian and Serravallian species extinct in the

Tortonian with A. lorioli. The second one, devel-

oped in the Indo-Pacific region, firstly appeared dur-

ing the "Helvetian" with Tretodiscus elongatus, that

clearly represent an ancestral form of the present-

day T. laevis (A. Agassiz, 1872-74), T. biforis

(Gmelin, 1778) and T. rumphi Lambert et Thiery,

1914. Thus, the latter one does not descend from

the Miocene European Amphiope, but it directly

descended from the Oligocene T. elongatum

through a succession of intermediate Indian forms."

On the variability of lunules

Cottreau (1914) examined the variability of the

Amphiope' s lunules, using a sample from the Bur-

digalian of Saint-Cristol (Nissan, Herault, France).

He demonstrated their large variability in shape and

size, and thus considered these characters not as

diagnostic. He asserted that, despite the lunules are

rounded or broadly oval in transverse direction in

the adult specimens, A. bioculata could have elon-

gated lunules in the direction of the posterior am-

bulacra in the juvenile stages, as well as adults of

A. baquiei Lambert, 1907. Cottreau justified this

apparent anomaly by the replication of ancestral

characters in veiy young individuals.

More recently, Philippe (1998), based on the

hypothesis of a wide intraspecific variability of the

species of Amphiope from the Miocene of the

Rhone Basin (France), tried to order the systematics

Dispute about Echinodiscus Leske, 1778 and Amphiope L. Agassiz, 1840 (Echinoidea Astriclypeidae)

231

of this genus, synonymyzing a number of species,

and maintaining only two valid taxa: A. bioculata

and A. boulei.

DISCUSSION

Unfortunately, Philippe (1998) didn’t consider

important aspects, such as the internal test structure

and the test plating. Those features were partially

described by Durham (1955), Kroh (2005) and

Pereira (2010). However, the material studied by

Philippe did not come from the type locality indi-

cated by des Moulins (1837) (Souze-la-Rousse).

The Rhone Basin could even be considered the

typical area, but the stratigraphical range of the

sediments cropping out in this area is wide and the

age of the holotype of A. bioculata was not indi-

cated by des Moulins (1837). Additionally, the holo-

type of A bioculata seems to have been lost [given

that the type established by des Mulins (1837) be-

longed to his own collection (Meo Museum) and

being that des Moulins lived in Bordeaux, we

asked at the the local Natural History Museum if

in their collections there are the Des Molulins col-

lection. But we had no answer] and no description

or illustration of its internal structure, or plate

structures have ever been provided. Subsequent in-

terpretations are highly controversial (see Agassiz

L., 1838-40; Cottreau, 1914; Philippe 1998), thus

leading to an uncertainty in the systematics of the

genus. This problem and the need to assess the real

extent of the intraspecific variability of the species

of Amphiope comparing it with the living Echin-

odiscus species, are emphasized by Stara & Borghi

(2014) during the revision and characterization of

the Amphiope Sardinian’s species, and they're em-

phasized by Smith & Kroh (2011), who recom-

mended a systematic review of the entire genus.

Finally, this problem has been analyzed by Stara &

Fois M. (2014) on the bases of an “ Echinodiscus ”

cf. auritus sample.

On the other hand, with regard to other Echin-

odiscus species, illustrations and descriptions made

in the past (except Durham, 1955) concerned only

shape and basic test measures. Today, it is demon-

strated that the only basic measures, such as Test

Length, Test Width and Test Height, are not suffi-

cient to establish the real belonging to a species,

rather than another, since different species have

been grouped under a single morphotype who

answered to the name of Echinodiscus. For exam-

ple, let's take two cases: the description given by L.

Agassiz (in Agassiz & Desor, 1847) in the text

where he established the species Echinodiscus

tenuissimus from Waigiu (Western Papua, Indone-

sia) and the description of E. tenuissimus in Dollfus

& Roman (1981), in his publication on Red Sea

echinoids. In the first L. Agassiz says only that the

species has two small lunules back, but does not

contain any illustration concerning the test plating;

Dollfus & Roman, however, states only that “Za

van tenuissima ( Ag . & Desor, 1847) =E. laevisH/.

(Ag. 1873), consider ee par Mortens en (1948 d, p.

411-413) comme espece separee, n’existe pas en

mer Rouge (pi. 33, fig. 5-6). Elle differe d’ auritus

typique surtout par la position de l ’anus (qui est sur

la ligne joignant les milieux des lunules) et ses

lunules fermees , \

" The van tenuissima (...). Herself differs from

typical auritus for the position of the periproct

(which is on the line that joins the half of lunules)

and by closed lunules. ”

Regarding Echinodiscus bisperforatus var.

truncata, however, they show at least two morpho-

types (coming from diverse countries as Papua

New Guinea and Zanzibar) and a long synonymy,

based on the shape of the test and of the lunules

(short or long). This morphotype, in fact, had

already been well illustrated by L. Agassiz (1838—

40, pi. 11, figs. 11-16) as Lobophora truncata ,

(unknown origin) that differs from L. bisperforata

by shorter lunules.

It is evident that, in the absence of platings

description, regarding the specimens studied by sev-

eral authors mentioned in the synonymy, it is im-

possible to understand what the authors refer to,

when they talk about Echinodiscus tenuissimus

and/or about E. bisperforatus var. truncata.

From Dollfus & Roman (1981): analyzing the

beautiful images that illustrate the specimen from

New Caledonia, in which the plating is partially vis-

ible, it can be observed that the plating is not char-

acteristic of Echinodiscus , as will be illustrated best

in Stara & Sanciu (2014). Other specimens, such as

Echinodiscus bisperforatus var. truncata figured in

pi. 34, figs. 3-4, coming from New Britain (Papua

New Guinea) or as E. bisperforatus var. truncata

figured in pi. 35 figs. 1-2, coming from Zanzibar,

they differ in test shape and lunules length, but it is

not clear what is their plating.

232

Paolo Stara & Daniele Fois

CONCLUSIONS

It is evident that, in the absence of careful plating

analysis, it is not possible to determine the member-

ship of these specimens to one species/genus rather

than to another. We believe that the analysis of the

structure and in particular of the plating in echinoids

is the primary tool for diagnosing and that is very

difficult to confirm old descriptions based only on

morphology.

Therefore, to analyze the specimens of the fam-

ily Astriclypeidae, in this volume will be studied

most importantly their plating.

REFERENCES

Agassiz L., 1838-41. Monographic d’echinodermes

vivant et fossiles. Echinites. Famille des Clypeast-

eroides. Seconde Monographic. Des Scutelles. Neu-

chatel: 149 pp.

Agassiz L. & Desor E., 1847. Catalogue raisonnedel

especies, des genres, et des families d’Echinides.

Annales des sciences naturelles. Zoologie et biologie

animale, 3: 129-168, 5-35, 355-380.

Airaghi C., 1899. Echinidi del Bacino della Bormida-

Bollettino della Societa Geologica Italiana, 18: 140-

178.

Cottreau J., 1914. Les echinides neogenes du Bassin mediter-

raneen. Annales de Elnstitut Oceanographique, 6:

1-193.

des Moulins C., 1837. Troisieme Memoire sur les echinides.

Synonymie general. Actes de la Societe Linneenne

de Bordeaux, 9: 45-364.

Dollfus R. & Roman J., 1981. Les echinides de la Mer

Rouge, Monographic zoologique et paleontologique.

Ministere de EUniversites, Comite del Travaux His-

toriques et Scientifiques. Memories de la section des

Sciences. Bibliotheque Nationale, Paris, 1911, 143 pp.

Durham J.W., 1955. Classification of clypeasteroid

echinoids. University of California Publications in

Geological Sciences, 31: 73-198.

Kroh A., 2005. Catalogus Fossilium Austriae, Band 2,

Echinoidea neogenica. Verlag der Osterreichischen

Akademie der Wissenschaften, Wien, 210 pp.

Kroh A., 2012., Echinodiscus bisperforatus truncatus (L.

Agassiz, 1841). In: World Echinoidea Database.

Kroh A. & Mooi R. (Eds.). Accessed through: Kroh

A. & Mooi R. 2012 World Echinoidea Database at

http://www.marinespecies.org/echinoidea/aphia.php?

p=taxdetails&id=513717 on 2013-01-09

Lambert J., 1915. Revision des echinides fossiles du

Bordelais. II partie: Echinides de EOligocene. Actes

de la Societe Linneenne, 64: 13-59.

Loven S., 1872. On the structure of the Echinoidea. The

Annals and Magazine of Natural History, 4: 285-298,

376-385,427-444.

Pereira P, 2010. Echinoidea from the Neogene of Por-

tugal mainland. Palaeontos, Lisbon, 18: 154 pp.

Philippe M., 1998. Les echinides miocenes du Bassin du

Rhone: revision systematique. Nouvelles Archives

du Museum d’Histoire Naturelle de Lyon, 36: 3-241,

249-441.

Smith A.B. & Kroh A., 2011. The Echinoid Directory.

World Wide Web electronic publication.

http://www.nhm.ac.uk/scienceprojects/echinoids (ac-

cessed September 2013).

Stara P. & Borghi E. 2014. The echinoid genus Amphiope

L. Agassiz, 1 840 (Echinoidea Astriclypeidae) in the

Oligo-Miocene of Sardinia (Italy). In: Paolo Stara

(ed.). Studies on some astriclypeids (Echinoidea

Clypeasteroida), pp. 225-358. Biodiversity Journal, 5:

245-268.

Stara P. & Fois M., 2014. Analysis on a sample of Echi-

nodiscus cf. auritus Leske, 1778 (Echinoidea Clypeas-

teroida). In: Paolo Stara (ed.). Studies on some

astriclypeids (Echinoidea Clypeasteroida), pp. 225-

358. Biodiversity Journal, 5: 269-290.

Stara P. & Rizzo R., 2014. Paleogeography and diffusion

of astrilypeids (Echinoidea Clypeasteroida) from

Proto-Mediterranean basins. In: Paolo Stara (ed.).

Studies on some astriclypeids (Echinoidea Clypeas-

teroida), pp. 225-358. Biodiversity Journal, 5: 233-

244.

Stara P. & Sanciu L., 2014. Analysis of some as-

triclypeids (Echinoidea Clypeasteroida). In: Paolo

Stara (ed.). Studies on some astriclypeids (Echinoidea

Clypeasteroida), pp. 225-358. Biodiversity Journal, 5:

291-358.

Stefanini G., 1912. Osservazioni sulla distribuzione ge-

ografica, sulle origini e sulla filogenesi degli Scutel-

lidae. Bollettino della Societa Geologica Italiana, 30:

739-754.

Biodiversity Journal, 2014, 5 (2): 233-244

Monograph

Paleogeography and diffusion of astridypeids (Echinoidea

Clypeasteroida) from Proto-Mediterranean basins

Paolo Stara 1 * & Roberto Rizzo 2

'Centro Stndi di StoriaNaturale del Mediterraneo - Museo di Storia Naturale Aquilegia, Via Italia 63, Pirri-Cagliari and Geomuseo

Monte Arci, Masullas, Oristano, Sardegna, Italy; e-mail: paolostara@yahoo.it

2 Parco Geominerario, Storico e Ambientale della Sardegna, Via Monteverdi 16, Iglesias, Carbonia-Iglesias, Italy; e-mail:

robertorizzo@parcogeominerario.sardegna.it

* Corresponding author

ABSTRACT In this paper, the authors retrace the geological changes that during the Neogene have modified

the paleogeography of the Western Mediterranean up to its current set-up. It is assumed that

migration and probably also speciation of the involved astriclypeids (particularly Amphiope

L. Agassiz, 1840 and Echinodiscus Leske, 1778) are closely related to those changes.

KEY WORDS Paleogeography; Astriclypeidae; Oligo-Miocene; Mediterranean Sea.

Received 25.06.2013; accepted 30.05.2014; printed 30.06.2014

In: Paolo Stara (ed.). Studies on some astriclypeids (Echinoidea Clypeasteroida), pp. 225-358

INTRODUCTION

Currently we are dealing with investigations on

the Miocene echinoids of Sardinia and their relation-

ship with the echinological paleofaunas that during

the Cenozoic have migrated from, or towards, the

Proto-Mediterranean seas. In particular, great atten-

tion is given to the genus Amphiope Agassiz, 1841

(family Astriclypeidae Stefanini, 1912), which is

common in the Oligo-Miocene marine deposits of

Sardinia (Comaschi Caria, 1955; Stara et al, 2012;

Mancosu & Nebelsick, 2013; Stara & Borghi, 2014)

and its relationship both with congeners of peri-

Mediterranean regions and the phylogenetically

closest genera such as Echinodiscus Leske, 1778.

The clypeasteroids appeared at the end of the

Mesozoic or in the early Cenozoic. According to

Smith (2001), the oldest clypeasteroid genus is

Togocyamus Oppenheim, 1915, from the end of the

Paleocene of Senegal, Togo and Nigeria. It is as-

sumed that they evolved from the cassiduloids,

which were already present in the Maastrichtian, at

the end of the Cretaceous or in the early Paleocene,

and then spread and diversified through the world

oceans (Smith & Kroh, 2011). The large number of

fossil records from the Eocene of United States, Eu-

rope, Middle East, Taiwan, Japan and Africa, con-

firms this wide diffusion and diversification of

clypeasteroids, raising doubts as to whether all this

could have happened in a tens of millions of years

as assumed by Kier (1982). Wang (1984) argued

that Echinodiscus tiliensis was already present in

the late Paleocene or early Eocene in Taiwan,

although the remains of this species were poorly

preserved and their stratigraphic occurrence was

uncertain. Because of paucity of the fossil record

available for study, the discussion on the phyloge-

netic position of many of these fossils is still open.

Many genera of clypeasteroids lived in the Proto-

Mediterranean and/or peri-Mediterranean basins,

from Eocene to Miocene, as Sismondia Desor, 1857,

Clypeaster Lamarck, 1801, Scutclla Lamarck, 1816,

234

Paolo Stara & Roberto Rizzo

Parascutella Durham, 1953, Amphiope L. Agassiz,

1840 (Cottreau, 1914; Smith & Kroh, 2011); few of

these survived there until the Pliocene, such as

Clypeaster (Giannini, 1957; Cotteau et al., 1876-

1891). In Sardinia, in particular, Amphiope appeared

in the Chattian-Aquitanian and disappeared in the

Tortonian-Messinian age (Comaschi Caria, 1955,

1972; Stara et al., 2012a).

Nowadays, a number of clypeasteroid genera

inhabit wide areas that include environments rang-

ing from tropical to temperate, with some species

extending even further polewards, such as Echinar-

chnius Gray, 1825; they adapted to different eco-

logical niches, with preference for the inter-tropical

zone (Ghiold & Hoffmann, 1984, 1986).

Several members of the Astriclypeidae family,

found the ideal habitat in more or less limited geo-

graphic regions. Astriclypeus Verrill, 1867 has been

adapted from Oligocene to the present, in Japan,

China and Cambodia seas (Smith & Kroh, 2011).

Echinodiscus (herein assumed as a monophyletic

group) spread from the Oligo-Miocene throughout

the Indo-Pacific, as far as Australia and South

Africa, including the Red Sea and the Persian Gulf;

Amphiope and all other echinoids belonging to the

family Astriclypeidae, are absent from the present

Mediterranean Sea. Different scientists did not

agree on the generic attribution of astriclypeids with

two lunules aligned with the rear ambulacra, and

about the size and shape of the lunules variability

in Amphiope (Stara & D. Fois, 2014).

In the North-Western Mediterranean, Amphiope

(bearing transverse or rounded lunules) is recorded

from Chattian-Aquitanian to Tortonian-Messinian,

and it occurs in about thirty localities of the Rhone

Basin, south-eastern France (Philippe, 1998) and in

other thirty sites of Sardinia (Italy) (Stara et al.,

2012a; Stara & Borghi, 2014).

Furthermore, in the Tyrrhenian Basin Amphiopeis

reported in Corsica (Cotteau, 1 877) and in some Ital-

ian regions: Tuscany (Giannini, 1957), Campania

(Barbera & Tavernier, 1989), Calabria (Cottreau,

1914, Carone & Domning, 2007; our observations)

and Sicily (Garilli, 2010); further, to the West it is

found in some regions of Spain [Barcelona (Lambert,

1928a); Valencia and Alicante (our collections); Mal-

lorca and Menorca Islands (Llompart, 1983)] and Al-

geria (Pomel, 1887-1888; Cotteau et al., 1891).

Along the Atlantic-European coasts Amphiope is re-

ported in Portugal [Lisbon, etc. (De Loriol, 1896;

Pereira, 2010)] and in France [Aquitaine (Lambert,

1928b) and Touraine (our collections)]; along the

Atlantic- African coasts Amphiope is found in Angola

(De Loriol, 1905). To the East, Amphiope is reported

in both the Central Paratethys [Austria and Hungary

(Kroh, 2005)], in the eastern basins [Turkey (Nebel-

sick & Kroh, 2002)] and from the Middle East

regions [Egypt (Kroh & Nebelsick, 2003), Arabia,

Iraq (our observations) and Iran (Khaksar &

Moghadam, 2007)] to the Indian coasts (Mooi,

1989). Atypical forms of " Amphiope " with axial

lunules are mentioned, but they are less frequent and

mainly consist of Oligocene species found in France

[Aquitaine (Lambert, 1915)], Italy [Liguria-Pied-

mont (Airaghi, 1899, 1901), North Africa [Tunisia

(Gauthier, 1899), Libya and Egypt (Gregory, 1911;

Fourtau, 1899, 1904)] and in the Aquitanian of the

Rhone Basin, France (Cottreau, 1914; Philippe,

1998). In the Miocene of some regions of the Middle

East both forms are recorded (Kier, 1972) (Fig. 1).

NOTES ON THE EXAMINED ASTRICLY-

PEIDS ECOLOGY

The ecology and life styles of some clypeast-

eroids have been studied in the past: among others,

Merrill & Hobson (1970) observed Dendrasterex

centricus populations along the Pacific coast of

California and Mexico; Kang & Choi (2002) stud-

ied a population of Astriclypeus manni from the

Cheju island of South Korea, Nebelsick & Kampfe

(1994) examined, from a taphonomic point of view,

some populations of Echinodiscus auritus and

Clypeaster humilis in the Bay of Safaga, Red Sea,

Egypt. Kleitman (1941) observed that some clypeast-

eroids can live at temperatures ranging from 1 0°C

to 30°C, with best conditions between 24°C and

26°C; Nebelsick (1999) observed that most species

of astriclypeids lived in near-shore to infralittoral

sandy environments, with high to medium-high

wave energy and deep currents. The discovery of

Pliocene fossils of Echinarachnius at Lituya Bay

(North West Coast of Alaska) in the Arctic Circle,

corresponding to 59° north latitude (Merte, 1930)

and Late Miocene Amplaster and Monophoraster

along the Atlantic coast of the Province of Chubu

in Argentina, at 45°South (Martinez & Mooi,

2005), indicates that some clypeasteroids were and

are able to adapt to significant differences in tern-

Paleogeography and diffusion of astriclypeids (Echinoidea Clypeasteroida) from Proto-Mediterranean basins

235

Figure 1. Oligo-Miocene distribution of the the main morphotypes.

perature and salinity conditions. Stara et al. (2012),

comparing the sediments of 15 Sardinian sites of

Miocene Amphiope with those of 5 present beaches

observed that those populations live in environ-

ment characterized by sandy bottoms and shallow

water.

As summarized by Kroh & Nebelsick (2003),

Mellita , Encope, Leodia and Echinodiscus are all

shallow borrowers, whereas Dendrasterex centricus ,

that maintains a partially exposed vertical posi-

tion in the sediment is a suspension feeder (see

Merrill & Hobson, 1970).

In particular, with regard to the bathymetric

range of E. auritus, Dollfus & Roman (1981) ob-

served it at 1-2 meters in depth in the Red Sea, but

also dredged a number of specimens between 10

and 15 meters in depth; the samples studied from

Bohol (Philippines), were collected at about 50 me-

ters in depth and Mazzetti (1893) during the dredg-

ing session carried out in the Red Sea by the ship

"Scilla" in 1891-92, at Goubet Soghra, collected sev-

eral specimens between 40 to 1 00 meters in depth.

PALAEOGEOGRAPHY AND PHYLOGE-

NETIC RECONSTRUCTIONS

In order to better understand the relationships

between these echinoids, we need to reconstruct

their migration pathways. As noted by Stefanini

(1912), the "scutellidi" always spread in a rela-

tively limited geographical area. We suppose that

this fact depends on their lifestyles, linked to near-

shore sandy environments.

Probably, their larval dispersal was not very

wide and needed to find sandy bottoms near roosts.

This seems justified by the fact that their spread

seems to have proceeded along the coast or through

basins of limited depth.

In the paleo-biogeographic reconstruction,

however, one of the keystones is the completeness

of the knowledge of the paleofauna of the period

under study.

Unfortunately, as stated also by Harzhauser et

al. (2007), only some areas have been deeply inves-

tigated and therefore are well known.

236

Paolo Stara & Roberto Rizzo

DIFFUSION OF THE ASTRICLYPEIDS

FROM PROTO-WESTERN MEDITERRA-

NEAN BASINS

An interesting contribution on the temporal and

spatial distribution of “scutelliformes" was pub-

lished by Stefanini (1912), who assumed that the

North Ocean was a spreading center for these

groups of echinoids, where several species were

already present during the Eocene and Oligocene.

A further contribution came from Cottreau (1914),

who made a summary on the diffusion and evolu-

tion of echinoids (among others, also Amphiope) in

the context of the Mediterranean Neogene.

By using the latest knowledge of geology and

paleobiogeography, as we shall see later, it is pos-

sible to better define the temporal distribution of the

two basic morphotypes, that are the main object of

this study The first is " Amphiope " and “ Echinodiscus ”

with axial lunules (Figs. 2, 3), appeared during the

Rupelian in Italy (Piedmont and Liguria), Libya and

perhaps also in Tunisia, and subsequently diffused

in the Middle Oligocene (late Rupelian-Early

Chattian) of the Bay of Biscay (France). In the

Aquitanian, a similar morphotype is present in the

Basin of the Rhone and then in the Early Miocene

of Tunisia, Libya, (Burdigalian) Egypt. In the

Middle Miocene the diffusion area shifted deci-

sively towards the East. There are no citations of this

morphotype in the Western Proto-Mediterranean

basins, along the Atlanto-European coasts (from

the Bay of Biscay to down) and along the Atlanto-

African coasts (Fig. 1).

The second morphotype, Amphiope with round

or transverse lunules (Figs. 4, 5), appeared in the

Chattian- Aquitanian in Sardinia and in the Aqui-

tanian of France and Kabylies; it was widespread

during the Miocene in the Western Mediterranean

Basin, along the Atlanto-European and Atlanto-

African coasts, in the Paratethys, in the Middle

East, as for as India and perhaps to Japan (Fig. 1),

and went extinct during the Tortonian-Messinian in

Sardinia (Philippe, 1998; Smith & Kroh, 2011;

Stara et al., 2012). Another morphotype (Fig. 6),

characterized by small rounded lunules rather far

from the petaloid tips (Fig. 7), firstly appeared in

Libya during the Miocene; it showed some features

of both the previous main morphotypes.

Echinodiscus cf. auritus (Fig. 3) is already wide-

spread from the Gulf of Suez to the Indo-Pacific

coasts in the Plio-Pleistocene. This morphotype is

recorded in the Plio-Pleistocene of Suez (Fourtau,

1899), in the Isle of Kliarak (current Kliark Island)

of the Persian Gulf (Duncan & Sladen, 1883) and

of the Aru Islands in Indonesia (Currie, 1924), in

the late Pliocene and Pleistocene of Java (Jeannet

& Martin, 1937). Lastly, it appeared in Pleistocene-

Holocene sediments near Hurghada (Red Sea,

Egypt) accompanied by other forms of Echinodi-

scus. Lindley (2001) cited a similar morphotype

characterized by axial and medium-sized lunules,

in the Middle Miocene (Langimar beds) of the prov-

ince of Morobe (Papua New Guinea), but he as-

signed it by mistake to Echinodiscus bisperforatus.

Currently Echinodiscus cf. auritus seems to be

the astriclypeid with the widest spread surpassing

the lines of the two tropics, 30°North to 35°South.

Their presence is ascertained along the East African

coast of Mozambique and South Africa and along

the coasts of Madagascar. To the North it is ascer-

tained along the Red Sea, to the Gulfs of Suez and

Aqaba, (Dollfus & Roman, 1981) the Persian Gulf

and along the northern shores of the Indian Ocean

(Sakthivel & Fernand, 2014). Lastly, to the East, it

is widespread in the Malay Archipelago (Indone-

sia), Thailand (Putchakam & Sonchaeng, 2004),

Philippines, along the Gulf of Siam, China (Lane et

al., 2000) and Japan, reaching the Northern and

Western coasts of Australia and perhaps New Cale-

donia (Fig. 8).

Echinodiscus bisperforatus shows a similar

distribution: it was present in the Middle Miocene

of Makamby island, Northern Madagascar (Collignon

& Cottreau, 1927) and in the Pleistocene-Holocene

sediments of Hurghada in the Red Sea (our collec-

tions), but some morphotypes showing features

similar to those of E. bisperforatus ( E . formosus

Yoshiwara and E. yeliuensis Wang), were maybe

already present in the Middle Eocene and certainly

in Miocene of Taiwan.

Finally, the "E. tenuis simus" group seems to

have a limited distribution in northern latitudes of

the Indian Ocean to Oceania, but today it would be

absent from the eastern and southern coasts of

Africa (Fig. 8).

In the reconstruction proposed by Stara &

Rizzo (2013), the similarity between the echinoid

faunas of North Atlantic and Western Mediter-

ranean would have been facilitated by the opening

of the pre-Pyrenean Corridor, which took place

Paleogeography and diffusion of astriclypeids (Echinoidea Clypeasteroida) from Proto-Mediterranean basins

237

between Middle Eocene and Middle Oligocene

(Fig. 9), allowing direct exchanges between the

Atlantic faunas of the Bay of Biscay and those of

the Alpine Tethys or intra-AlKaPeCa basins (this

is an acronym used by Bouillin et al. (1986) to in-

dicate the micro-continent that moving away from

the European plate, would have given rise to differ-

ent regions of the actual Western Mediterranean).

After the closure of the pre-Pyrenean Corridor,

which probably has occurred during the Middle

Oligocene, the two faunas began to differentiate.

In addition, the almost complete separation between

the Alpine Tethys (from which the Proto-Western-

Mediterranean was bom) and the Western Neotethys

basins (according to the reconstructions of Stamp fli

et al. (2002), or basins resulting from detachment

of the AlCaPeKa micro-plates, according to

Carminati et al. (2012), also justifies a lot of the

differences observed between the Miocene faunas

of the Western Mediterranean and of the Eastern

Mediterranean (see Figs. 9-14). For example, in the

first area, "Amphiope" with axial lunules and

Figures 2-5. Moiphotypes based on the shape of rear ambulacral lunules/slits. First moiphotype, bearing axial lunules or

slits notching the posterior margin: Figure 2. “ Amphiope “ pedemontana, Oligocene, Val Bormida, Liguria and Piedmont,

Italy. Figure 3. “Echinodiscus”cf. auritus, Recent, Mangili, Tulear, Madagascar; Second morphotype, bearing rounded or

transverse lunules: Figure 4. Amphiope sp., Oligo-Miocene, Duidduru, Sardinia, Italy. Figure 5. A. nuragica, Oligo-Miocene,

Cuccuru Tuvullao, Sardinia.

Figures 6, 7. Moiphotype with small lunules far from the petal tips: 6 “ Amphiope ” boulei, Aquitanian, France (from Cot-

treau, 1914). Fig. 7. “ Amphiope ” sp., “Miocene”, Libya (NMHUK collections).

238

Paolo Stara & Roberto Rizzo

Scutella were absent, whereas in the second area

both these genera were widespread. Indeed, in the

Western Mediterranean, only Amphiope and Paras-

cutella are known (A. Kroh, personal communica-

tion, June 2012).

According to Stara & Rizzo (2013), from the

Sardinian- Provencal basins, derived from the frag-

mentation of the micro-continent AlKaPeCa, at

least three waves of migration of lunulate scutellids

may have originated: two from the East and one

from West. The first wave would have taken place

during the Oligocene through the corridor of the

Bormida Valley (Piedmont and Liguria) (Fig. 10),

the second at the beginning of the Miocene, through

the corridor of the Alpine Paratethys, the third was

a result of the fragmentation, the detachment and

their drift towards the south, of micro-plates, from

the continental margin of the Ibero-Provengal crust.

The second of these migration has been already

recognized by Kroh (2007), who stated that

the majority of the echinoid fauna of the Central

Paratethys is immigrant from the western Mediter-

ranean and partly shows similarities with that of the

Atlantic region. This migration took place in three

phases: the first wave would have started at the

beginning of the Miocene from the Rhone Basin

through the Alpine Tethys, the second and the third,

much later, according to Kroh (2007) took place

through the trans-Tethys Dinarids Corridor that led

to the Adriatic Neotethys. Some species wich

immigrated during the first phase had Atlanto-

Mediterranean affinities, those joining the second

and third phases were more closely related to the

faunas of the Eastern- African coasts.

The migration along the Val Bormida Corridor

has been hypothesized by Stara & Rizzo (2013),

based on the presence of a series of Scutella and

"Amphiope" pedemontana rich beds that crop out

in the Rupelian of Liguria and Piedmont. The hy-

pothesized migration is in accordance with the sim-

ilarity of some characters that these "Amphiope"

share with those of Rupelian from the coast of

Libya and those of the Middle Oligocene (Late

Rupelian-Early Chattian?) of the Gulf of Biscay.

In addition, this step is also traced by the spread of

Heterobrissus Manzoni et Mazzetti, 1878. This

Figure 8. Distribution of extant main morphptypes of “Echinodiscus” genus. Yellow dots: “Echinodiscus“ cf. auritus

group. Green dots, Echinodiscus bisperforatus group. Orange dots: “E chino discus” tenuissimus group.

Paleogeography and diffusion of astriclypeids (Echinoidea Clypeasteroida) from Proto-Mediterranean basins

239

Schematic palinspastic recontructions modified from: 1) Stampfli et al. (2002); 2-6) Rosembaum et al. (2002) and Carminati et al. (2012)

Legend

Morphotype

“Amphiope" and

Echinodiscus

Morphotype

Amphiope

Direction of

astriclypeids

migration

Emerged

Inland

Shelf

Oceanic

AlCaPeKa

crust basins basins basins microplates

Figures 9-14. Time scanning of Amphiope diffusion in the proto-Mediterranean basins. Figure 9. Middle Eocene-Lower

Oligocene connection between the Atlantic-Gulf of Biscay and the Provencal Basin. Figure 10. Morphotype 1 populations

begin their eastward Oligocenic migration through the Val Bormida Corridor. Figure 1 1 . Starting from a single distribution

center, located between the Biscay and the original intra-AlCaPeKa basin, morphotype 2 populations begin their Oligo-

Miocenic spread. Figure 12. Morphotype 2 is already widespread from Atlantic coasts to the far east; morphotype 1 is no

longer present in the western basins. Figure 13. Morphotype 2 reaches its peak in the Rhone Basin and in Sardinia;

morphotype 1 has spread from the Middle East to India. Figure 14. At the end of the Middle Miocene, morphotype 2 begin

to extinguish, whereas morphotype 1 has colonized the Indian Ocean and the Western Pacific Ocean. Based on the paleo-

geographic data from Stampfli et al., 2002; Rosenbaum et al. 2002; Carminati et al., 2012.

240

Paolo Stara & Roberto Rizzo

genus is present in the Oligocene of Caribbean Is-

lands (Jackson, 1922), in the Early Miocene of

Sardinia (Stara et al., 2012b), in the Middle

Miocene of Emilia and San Marino (Manzoni &

Mazzetti, 1 878), lastly in the Serravallian of Cyprus

(Currie, 1935; Smith & Gale, 2009), and today it is

widespread in the seas of China and South Eastern

Asia (Lane et al., 2000). So, the basins of the

Middle East suffered at least two waves of migrants

from N-NW, the first one through the Adriatic

Tethys during Oligocene and the second one

through the eastern Paratethys between the end of

the Early Miocene and the Middle Miocene.

In summary, from the Late Oligocene to the

Early Miocene, the Val Bormida Corridor had al-

ready closed as a result of Apennines orogeny,

while the Alpine Tethys Corridor shut at the end of

Burdigalian as a result of the Alpine orogeny. The

closing of these two corridors led to the isolation

or, at least, to a drastic reduction of the exchanges

between the eastern and western faunas of the Tethys

(or Proto-Mediterranean basins). This new situation

probably allowed the differentiation of the Oligo-

Miocene " Amphiope " with axial lunules from the

North- African and Middle-Eastern coasts. During

the Burdigalian, Amphiope with rounded or trans-

verse lunules was already present in the central

Paratethys and in Egypt. However, while it seems

clear that it arrived in the Paratethys crossing West

to East the canal north- Alpine, is not yet clear how

it arrived in Egypt. In fact, there is no evidence of

these echinoids, nor Parascutella, along the Miocene

Adriatic and Ionian seashores, favoring the conti-

nuity of their migration through the eastern basins,

already during the Middle-Early Miocene, to other

marine faunas. In any case, as a result of their mi-

gration, Amphiope went to Turkey, Egypt, Saudi

Arabia, Iran and finally to India and also in Iraq

(our observations). Finally, Harzhauser et al. (2007)

suggest that the complete disconnection between

the Proto-Mediterranean basin and the Indian

Ocean basin occurred at the end of the Burdigalian,

when the two faunas where already differentiated.

The apparent diachrony should be clarified

when the astriclypeids of the eastern regions faunas

will be studied. In fact, it is possible that faunas

from the West (as we assumed), but also from the

East, met in the Middle East area, since different

forms of “E chino discus” were already present

(doubtfully) in the Middle Eocene, but certainly in

the Lower Miocene, respectively, in the islands of

Taiwan and Japan.

Regarding the Mediterranean, according to Rogl

(1998), during the Miocene the two sides of the

Mediterranean were in full connection, while ac-

cording to Stampfli et al. (2002), these were com-

pletely separate. Much evidence is needed,

however, we argue in favor of this second hypothe-

sis. The reconstruction made by Stampfli et al.

(2002) suggests that the complete connection between

the Eastern and the Western Mediterranean would

have occurred much later, when the Calabrian mi-

croplate reached the Italian Apennines, at the end

of the Miocene or during the Pliocene. Although the

precise date of the disconnection between the basins

of the eastern Neotethys and the Indian Ocean is

still under discussion, Harzhauser et al. (2007)

agree with the development of different biota for

these two regions during the beginning of the

Middle Miocene.

THE SPREAD OF AMPHIOPE WITH

TRANSVERSE OR ROUNDED LUNULES

TO THE WEST-SOUTH-WEST

In the North-Western Mediterranean sedimen-

tary basin, Chattian-Aquitanian to Tortonian-

Messinian fossils of Amphiope with rounded or

transverse lunules have been found in many locali-

ties in the Rhone Basin (Philippe, 1998), and Sar-

dinia (Stara et al., 2012a; Stara & Borghi, 2014).

In detail, starting from density of Amphiope

deposits existing in a specific region, we can assume

that Amphiope appeared in a fairly restricted area

within the archipelago formed between the Basin

of the Rhone and Sardinia, from the end of the

Oligocene to the beginning of Miocene. According

to Rosenbaum et al. (2002) and Gattacceca et al.

(2007), in this period different microplates began

drifting towards the South forming that archipel-

ago (Figs. 11-14). The shift of these microplates

to the current position point lasted about 7 million

years, and during this time the fauna could (in

some cases) differ from the original giving rise to

new species, as it happened for example in Sar-

dinia, where 3 species [. Amphiope nuragica

(Comaschi Caria, 1955); Amphiope lovisatoi Cotteau,

1895, and Amphiope montezemoloi Lovisato,

1911] were confirmed and for the first time,

Paleogeography and diffusion of astriclypeids (Echinoidea Clypeasteroida) from Proto-Mediterranean basins

241

another two new ones have been described (Stara

& Borghi, 2014).

At the end of the Burdigalian the Sardinia-

Corsica microplate had completed its route after an

anticlockwise rotation, stopping more or less in its

current position; Calabria located in the East of Sar-

dinia, and it reached its current position only in the

Pliocene, the Kabylies had almost welded with

North Africa, the Betic-Rifian microplates were still

in the Alboran Sea, while the Balearic Islands were

more or less in the current position.

A part of Sardinia-Corsica and Balearic Islands

now detached itself, the other microplates, each

with its own specific fauna, to the contact with the

North African margin (for example the Kabylies) or

southern Europe (Iberia) were able to create further

migrations, which most likely occurred along the

sandy beaches adjacent to shallow depths (Ste-

fanini, 1912).

Pomel (1883, 1887-8) and Cotteau et al. (1876-

1891) reported the presence of Amphiope in the

Early Miocene of Cherchell and in the Middle

Miocene of Mleta, Oran, as well as in other places

of Kabylies (Algeria). Most likely, as suggested by

Stefanini (1912), starting from the Kabylies,

Amphiope populations reached the Atlantic Ocean

to continue towards South to colonize the area of

Bom Jesus (Angola, West Central Africa) during the

Middle Miocene. It is uncertain if the presence of

Amphiope in the region of Alicante and Valencia

during the Tortonian is due to a direct migration

from the North, since its presence is also reported

in the Middle Miocene in the region of Barcelona.

It seems logical that, starting from the South of the

Iberian Peninsula Amphiope has continued its coa-

stal migration as far as the Atlantic Ocean and back

along the coast of Portugal (Fig. 12). Pereira (2010)

reports: "The echinoid fauna of mainland Portugal

is closely related to that of the Mediterranean re-

gion. In fact, the biogeographic investigation of the

Portuguese echinoid fauna shows that a major part

of the Portuguese species is composed by Por-

tuguese immigrants from the Mediterranean area

(42.9% of the fauna in the Burdigalian and 60.9%

in the Middle Miocene). Endemism is low during

Miocene, with endemic species not exceeding 25%

of total Portuguese echinoid fauna".

Following its migration toward the North,

Amphiope reached the French coast until the Bay of

Biscay, where it has been reported in the Serravallian

deposits; its migration toward the North seems to

stop in the great inland sea that covered the Touraine,

where different sites related to Middle-Late Miocene

(Serravallian-Tortonian) are reported. However,

after the closure of the pre-Pyrenean Corridor, in

Aquitaine an endemic fauna probably developed

independently and directly from the original Aqui-

tanian Amphiope ovalifora Fallot, 1903.

In conclusion, along the Italian peninsula,

Amphiope was found in the Middle Miocene of

Tuscany (Giannini, 1957) and Campania (Barbera

& Tavernier, 1989); in the first case it is unclear

whether the migration is linked to the movement of

microplates along the Mediterranean, or if it oc-

curred directly from North along the peninsula coasts.

However, the presence of Amphiope in the Middle

Miocene of Campania and in the Tortonian of Sicily

(Garilli et al., 2010), can be connected with the ap-

proach of the Calabrian microplate (Fig. 14). Dur-

ing the Burdigalian this microplate moved to the

East bringing the original fauna, as stated by the find-

ings in the Tortonian deposits of Cessaniti near Vibo

Valentia (Cottreau, 1914).

CLIMATE CHANGE, LIMIT OF THE

DIFFUSION

To understand the diffusion of scutelliforms

living nearshore, we need to consider the trend of

climate change from the Cretaceous on to the Miocene,

and how it conditioned the life of organisms inhab-

iting the continents and oceans of the northern

hemisphere and Southern Africa. As summarized

by Harzhauser et al. (2007), the warm climate of the

Cretaceous continued into the Early Palaeogene,

with a distinct optimum that characterized the

Paleocene-Eocene transition. Starting in the Late

Eocene, a gradual decrease in temperature culmi-

nated around the Eocene-Oligocene boundary, lead-

ing to the formation of the first Antarctic ice cap.

From the late Oligocene times, the trend of increas-

ing temperature continued intermittently until the

Middle Miocene, when it reached its maximum

(Climate Optimum).

Around 14.2 Ma began the transition of the

Middle Miocene climate, characterized by the

cooling of surface waters and the expansion of the

East-Antarctic ice cap (Shevenell et al., 2004),

and during this time the extinction of Parascutella

242

Paolo Stara & Roberto Rizzo

and Amphiope , began, thus stopping their diffu-

sion to the south. Only for "Echinodiscus" migra-

tion will continue in the Indian Ocean and along

the coast of South Eastern Europe to settle in the

current positions.

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Monograph

The echinoid genus Amphiope L. Agassiz, 1 840 (Echinoidea

Astridypeidae) in the Oligo-Miocene of Sardinia (Italy)

Paolo Stara 1 & Enrico Borghi 2

'Centro Studi di Storia Naturale del Mediterraneo - Museo di Storia Naturale Aquilegia, Via Italia 63, Pirri-Cagliari and Geomuseo

Monte Arci, Masullas, Oristano, Sardinia, Italy; e-mail: paolostara@yahoo.it

2 Societa Reggiana di Scienze Naturali, Via Tosti 1, 42100 Reggio Emilia, Italy; e-mail: e.borghi@corghi.it

^Corresponding author

ABSTRACT The records of the genus Amphiope Agassiz, 1840 (Astridypeidae) from Sardinia are revised

on the basis of 110 specimens, collected from 15 localities of Oligo-Miocene age. Since the

morphological characters stated in the literature to distinguish the species of Amphiope de-

scribed in this region cannot provide a clear separation between them, analyses of the plate

patterns and of the internal test structure are introduced as taxonomic tools useful for

species-level taxonomy in this genus. Five different species of Amphiope are identified.

Three of the six species erected on the basis of fossil material from Sardinia are confirmed

as valid: Amphiope lovisatoi Cotteau, 1895, A. montezemoloi Lovisato, 1911 and A. nuragica

(Comaschi Caria, 1955). Two additional species are left in open nomenclature. The morpho-

logical descriptions and the stratigraphical distributions are updated and improved.

KEY WORDS Echinoidea; Amphiope', Oligo-Miocene; Sardinia; Mediterranean.

Received 25.06.2013; accepted 30.05.2014; printed 30.06.2014

Paolo Stara (ed.). Studies on some astriclypeids (Echinoidea Clypeasteroida), pp. 225-358

INTRODUCTION

The genus Amphiope L. Agassiz, 1 840 (Echinoi-

dea Astridypeidae) is known from the Oligocene

and Miocene of Central and Southern Europe, North-

ern Africa, Angola, Middle East, India (Smith &

Kroh, 2011).

It is well represented also in the Oligo-Miocene

of Sardinia (Fig. 1) since ten species of Amphiope

were recorded in the literature (Table 1), six of

which were erected as new taxa on the basis of

Oligo-Miocene fossils from this region: A. lovisatoi

Cotteau, 1895, A. dessii Lovisato in Cotteau, 1895,

A. montezemoloi Lovisato, 191 1, A. pallavicinoi Lo-

visato, 1914, A. calvii Lovisato, 1914 and A. nurag-

ica (Comaschi Caria, 1955). A. montezemoloi was

subsequently placed into synonymy with A. biocu-

lata (des Moulins, 1837), the type-species of the

genus, by Comaschi Caria (1955), Philippe (1998)

and Kroh (2005); the last author considered also A.

lovisatoi as synonymous with A. bioculata. Since

the type-specimens belonging to Lovisato’s collec-

tion, at that time housed at the “Regio Museo Mine-

ralogico e Geologico di Cagliari”, were lost in 1943

(Comaschi Caria, 1955), this work is based mainly

on new material collected from the respective type-

localities and on the holotype of A. nuragica, still

available to study at the “Dipartimento di Scienze

Chimiche e Geologiche, Universita di Cagliari”.

The genus Amphiope accounts for more than 40

species, most of which are nominal species in need

of revision, due to high intraspecific variation and

246

Paolo Stara & Enrico Borghi

Figure 1. Regions of Italy in which Amphiope has been

recorded in the literature (dark grey). Enlarged area (Sardinia):

finding localities examined in this study (black circlets), or

recorded in the literature (grey squares). Province of Sassari:

1 = Porto Torres; 2 = La Cracca; 3 = Bancali; 4 = San Giorgio;

5 = Sedini; 6 = Monte Oria Pizzinnu; 7 = Billiu and Monte

Sa Loca, near Chiaramonti; 8 = San Matteo, near Ploaghe; 9

= Ardara; 10 = Bessude; 1 1 = Bonnanaro; 12 = Monte Zarau,

near Torralba; 13 = Bonorva. Province of Oristano: 14 =

Nuraghe Caiu, near Villa Sant’ Antonio; 15 = Laconi; 16 =

Bruncu Muntravigu and Tanca Sierra, near Senis; 17 = Duid-

dura and Genoni (Nuoro); 18 = Monte Is Casteddus, near Isili;

2 1 = Santadi and Sa Lisporra, at Capo Frasca. Province of Ca-

gliari: 19 = Cuccuru Tuvullao; 20 = Strintu ‘e Melonis and

Nurri; 23 = Monte S. Michele; 24 = Capo S. Elia and Bonaria.

Medio Campidano province: 22 = Monte Arcuentu.

poor species definition (Smith & Kroh, 2011). The

complex taxonomy of this genus has been tradition-

ally based on the external morphological features,

mainly test outline, size and shape of lunules and

petals. Structural characters, largely used in the ta-

xonomy of other clypeastroids ( Durham, 1955;

Lohavanijaya, 1965; Mooi, 1989; Kroh, 2005; Jansen

6 Mooi, 2011), were almost overlooked in earlier

studies dealing with Amphiope and, although sev-

eral species of Amphiope have been described in the

literature, important features for species-level tax-

onomy, such as oral plating, were poorly illustrated

or omitted completely.

During a preliminary investigation we attempted

to find out whether the specimens from Sardinia

might be attributed to already known species, but a

particular difficulty was found in the use of the ex-

ternal characters alone, because comparison with

taxa whose structural characters are unknown re-

mained uncertain. This paper presents the results of

further studies based on a large and well preserved

fossil sample, to bring light into the problem of

classification of Amphiope. The main purpose is a

modem revision of Sardinian occurrences of Am-

phiope, using morphological and morphometric

analyses, with emphasis on the plate patterns and

the internal test support system.

Geological setting

The geology and palaeoecology of the Sardinian

Amphiope-bQdivmg localities cited in this study were

described by Stara et al. (2012) and Mancosu &

Nebelsick (2013). In the following a brief summary

is given of the type localities of the three species

revised from this region.

Central Sardinia (Marmilla). Three main ma-

rine sedimentary cycles have been recognized in

the Sardinian Basin, from late Oligocene to early

Messinian (Assorgia et al., 1997; Funedda et al.,

2000; Carmignani et al., 2001). The Amphiope-

bearing deposits in Central Sardinia belong to the

first cycle, extending from the late Oligocene to

the late Burdigalian. The Cenozoic sequence

starts with the late Oligocene-early Aquitanian

Ussana Formation (Pecorini & Cherchi, 1969),

consisting mainly of sediments of continental ori-

gin. The Ussana Formation is partly heteropic

with and is followed by the Nurallao formation,

late Chattian-early Burdigalian (Serrano et al.,

1997), which consists of the “Conglomerato di

Duidduru” member, made of coarse elastics from

transitional (deltaic) environments (Sowerbutts &

Underhill, 1998) and the littoral marine deposits

of the Arenarie di Serralonga member, late Oligocene-

early Aquitanian (Assorgia et al., 1997; Barca et

al., 2005), which yielded the Amphiope specimens

examined from this area. The Nurallao Formation

is partially heteropic with and followed by the

Calcari di Villagreca, dated to the late Oligocene-

early Burdigalian, and by the Marmilla Forma-

tion, dated to the Aquitanian (Cherchi et al.,

2008). The material studied from this area was

collected from eight localities: Villa Sant’Anto-

nio, Bruncu Muntravigu, Tanca Sierra, Duidduru,

The echinoid genus Amphiope L. Agassiz, 1840 (Echinoidea Astriclypeidae) in the Oligo-Miocene of Sardinia (Italy) 247

Recorded species

Cited by

Locality (Province)

A. bioculata (des Moulins, 1 837)

Lambert (1907)

Capo S.Elia (Cagliari)

Comaschi Caria (1955, 1972)

Cuccuru Tuvutlao (Cagliari)

Barca et al. (2000}

Thiesi (Sassari)

Spano et al. (2002)

Capo Frasca (Oristano)

A hollandei Cotteau, 1877

Cotteau (1895)

Castelsardo (Sassari), CapoS. ESia and Monte S,

Michele (Cagliari), San tad i (Oristano)

Lovisato (1911, 1914)

Capo Frasca (Oristano), Nurri (Nuoro), Toiralba

(Sassari)

A. Icvisatoi Cotteau, 1895 *

Cotteau (1895)

Billiu* near Chiaramonti (Sassari)

Comaschi Caria (1955, 1972)

Cuccuru Tuvutlao (Cagliari)

A. dessS Cotteau, 1 895 *

Cotteau (1895)

Nurri* (Nuoro)

Lovisato (191 4)

Bes sude (Sassari}

Comaschi Caria (1955, 1972)

Cuccuru Tuvullao (Cagliari)

A. mont ezemoloi Lovisato, 1911 *

Lovisato (191 1)

San Giorgio* near Sassari

A. deydieri Lambert, 1912

Gomaschi Caria (1955, 1972)

Cuccuru Tuvutlao (Cagliari)

A. transuersivora Lambert, 1912

Comaschi Caria (1955, 1972)

Cuccuru Tuvufiao (Cagliari)

A. calvii Lovisato. 1914 *

Lovisato (191 4)

San Matteo* near Ploaghe (Sassari)

A. pallavicinoi Lovisato. 1914*

Lovisato (191 4)

Monte Zarau* near Tonalba (Sassari)

Comaschi Caria (1955, 1972)

Cuccuru Tuvullao (Cagliari)

Amphiope sp

Lovisato (191 4)

Nulvi and: Bonorva (Sassari); Monte Arcuentu

(Oristano). Sant' Antonio Ruinas (actually Villa S.

Antonio), Laconi, Genoni, Capo S. Elia and Monte S.

Michele fCsqliari).

A. nuragica (Comaschi Caria. 1955) *

Comaschi Caria (1956, 1972)

Cuccuru Tuvullao* (Cagliari)

Amphiope sp.

Stara et al., 2012

Cuccuru Tuvullao, Monte is Casteddus and Isili

(Cagliari); Nuraghe Caiu, Bruncu Montravigu and

Tanca Sierra (Oristano); Duidduru (Nuoro); Bancali;

La Crucca; Porto Torres; Ardara; Chiaramonti and

Bonnanaro (Sassari)

Table 1 . Records of Amphiope from Sardinia reported in the literature. An asterisk marks the species firstly described on the

basis of material collected from this region.

Genoni, Isili, Cuccuru Tuvullao and Capo Frasca

(see Fig. 1).

- Cuccuru Tuvullao (n. 19 in Fig. 1). The type-

locality of A. nuragica (Comaschi Caria, 1955) is

located 1.5 kinNE ofNuragus (Cagliari). Amphiope

occur in high numbers in a medium to coarse-

grained volcanoclastic sandstone (herein named C.

Tuvullao I), 2-2.5 m thick, corresponding to the

“Facies B” of Mancosu & Nebelsiclc, 2013; Fig. 2).

Amphiope is the most abundant taxon, followed by

Paras cutella sp., reworked balanids and bivalves.

Seven species of Amphiope were cited by Comaschi

Caria (1955) in this layer, including A. nuragica.

The echinoids are denuded, commonly fragmented

in the shape of pie-slices. Loosely packed or dis-

persed specimens are both in life position and up-

side down, their long axes being more or less paral-

lel to the stratal surfaces. This sand dollar deposit

is assigned to a shoreface environment and repre-

sents a multiple in situ reworking accumulation

(Mancosu & Nebelsick, 2013).

Fragments and rare complete specimens of Am-

phiope occur also at the base of the overlying richly

fossiliferous fine mudstone (herein named C.

Tuvullao II), corresponding to the “Facies C” of

Mancosu & Nebelsick (2013); Fig. 2). The fossil

content is dominated by gastropods, belonging to

the genus Turritella Lamarck, 1799, and by the bi-

valve Panopea Menard, 1807.

Northern Sardinia. Seven Amphiope-bQixnng lo-

calities were sampled in the Sassari province: Porto

248

Paolo Stara & Enrico Borghi

Torres, La Crucca, Bancali, San Giorgio, Chiara-

monti, Ardara and Bonnanaro. These deposits be-

long to the second Cenozoic sedimentary cycle, late

Bur digalian- early Langhian (Mancosu & Nebel-

sick, 2013). At the base, the lacustrine and fluvio-

deltaic sediments of the Oppia Nuova formation

(Funedda et al., 2000) are overlain by the Calcari

di Mores Formation, consisting of bioclastic lime-

stones and poorly cemented sands of shallow water

origin (Mazzei & Oggiano, 1990; Funedda et al.,

2000). The Calcari di Mores Formation is followed

by the Marne di Borutta Formation, which repre-

sents a deeper shelf facies.

- Chiaramonti (n. 7 in Fig. 1). Billiu, the type-

locality of A. lovisatoi, rests along the main road

from Ploaghe, close to Chiaramonti. The specimens

examined in this paper were collected from the sec-

tion studied by Stara et al. (2012) cropping out at

Monte Sa Loca, less than 1 km far from Billiu: it is

9 m thick and extends laterally for some 40 m. The

main Amphiope- bearing layer corresponds to the

“Facies C” of Mancosu & Nebelsick (2013; Fig. 4),

dated to the lower part (late Burdigalian) of the Cal-

cari di Mores Formation, and represents the lateral

extension of the strato-type of Billiu. It consists of

very coarse, poorly sorted, massive sandstone, about

1 m-thick, with carbonate cement, which, though dis-

continuous, can be traced over the entire length of

the outcrop. The dense accumulation of well-preser-

ved echinoids of Facies C is considered to represent

an autochthonous assemblage in a shoreface envi-

ronment (Mancosu & Nebelsick, 2013). Amphiope

prevails by far, followed in abundance by Agassizia

Yoshiyasu, 1987, Parascutella Durham, 1953, Echi-

nolampas Gray, 1825 and small bivalves, and is rep-

resented almost exclusively by complete tests (80%)

with subordinate fragmented tests.

- San Giorgio (n. 4 in Fig. 1). The type locality

of A. montezemoloi was never cited again since its

original description. It has been traced by one of the

authors (P.S.) following the indication of Lovisato

(1911; 1914), along the Sassari-Alghero railroad,

1.5 km far from the abandoned rail-station of San

Giorgio towards Olmedo. The Amphiope-bQSLvmg

outcrop consists of a coarse-grained sand-stone and

extends for a few square meters, only. Amphiope is

mainly represented by fragmented specimens in

chaotic position.

MATERIAL AND METHODS

The studied material consists of 1 1 0 specimens,

preserved as whole coronas deprived of the spines,

and several fragmented individuals, from 15 Miocene

Sardinian localities (Fig. 1). Most of them are

Figure 2. Amphiope : scheme of the biometric parameters measured in the studied specimens.

The echinoid genus Amphiope L Agassiz, 1840 (Echinoidea Astriclypeidae) in the Oligo-Miocene of Sardinia (Italy) 249

housed at the Museo di Storia Naturale “Aquilegia”

of Cagliari (MAC code): 95 specimens were col-

lected by one of the authors (P.S.), 12 specimens

were donated by private collectors. Three additional

specimens have been examined in the Dipartimento

di Scienze Chimiche e Geologiche, Universita di

Cagliari (UNICA); they include the holotype of A.

nuragica (code 9CC.8- 10504) and two specimens

classified as “A. calvii (code 3CC) and “A. dessii ”

(code 6CC- 10503) by Comaschi Caria (1955; pi. 1

and pi. 7, respectively). The other specimens from

Cuccuru Tuvullao described by Comaschi Caria

(1955) have not been traced at the UNICA. The

measurements of ten lost specimens, reported by

Lovisato (1911; 1914) and Comaschi Caria (1955),

were used in the statistical biometrical analyses;

since some data were lacking, they were taken from

the figures given by Comaschi Caria (1955: pi. 2,

figs. 1, 2, pi. 3, figs. 1, 2, pi. 4, figs. 1, 2, pi. 5, figs.

1, 2, pi. 6, figs. 1, 2, pi. 8, figs. 1, 2, pi. 9, figs. 1, 2,

pi. 13, fig. 1; 1972: pi. 44. figs. 3-4).

According to Lovisato (1911; 1914) the speci-

mens he provided for study to other echinologists,

including the type series used by Cotteau (1895) to

erect A. lovisatoi and A. dessii , returned back to his

collection at that time stored in the “Regio Museo

Mineralogico e Geologico di Cagliari”. All those

specimens, as well as the type material of the other

species based on fossils from Sardinia, with the ex-

ception of A. nuragica , were lost in 1943 during the

2nd World War (fide Comaschi Caria, 1955).

The internal structure was studied by sectioning

the test, and by X-ray; 5 specimens from Bancali, 3

from C. Tuvullao, 2 from Bonnanaro and 2 from

Chiaramonti were used for this purpose.

Morphological abbreviations (Fig. 2) -a = angle

between the axes of the two posterior petals; TL =

test lenght; TW = test width; TH = test height; Ll-

L2 = lunule length and width, respectively; L3 = dis-

tance posterior petal tip from the corresponding

lunule; L4 = distance apical system-posterior mar-

gin; L5-L6 = length and width of the frontal petal,

respectively; L7-L8 = length and width of the ante-

rior paired petal, respectively; L9-L10 = length and

width of the posterior petal, respectively; LI 1 = dis-

tance posterior border of periproct and of the test;

LI 2 = distance between the posterior border of the

peristome and of the periproct; LI 3 = antero-poste-

rior diameter of the basicoronal circlet. Measure-

ments of LI to LI 0 were taken from the left side of

the test, where possible. Systematic palaeontology

follows Kroh & Smith (2010).

Broken edges were indicated by dotted lines in

drawings, heavier lines indicate unbroken ambitus.

Plates were numbered according to Loven’s system

(CIT; compare Plate 1 Fig. 3b), interambulacral

plates were shaded in grey.

Geographic coordinates on the World Geodetic

System of 1984, WGS84.

Biometric analyses were carried out and data

analyzed using the software PAST -version 1.97

(2010) (Hammer & Harper, 2010; Hammer et al.,

2001), to help interpret the samples collected from

eight Sardinian localities. The original values of the

metric parameters were divided by TL to exclude

the effect of size, as suggested by Durham (1955),

Lohavanijaya (1965) and Pereira (2010). Principal

component analysis (PC A) is based on 71 speci-

mens [8 from Bancali, 4 from Bonnanaro, 34 from

Chiaramonti, and 25 from C.Tuvullao (22 from

layer I and 3 from layer II)]. The analysis utilizes

11 variables: TW/TL, TH/TL, Ll/TL, L2/TL,

L3/TL, L4/TL, L5/TL, L6/TL, L9/TL and L10/TL.

Univariate and bivariate analyses were based on a

data set taken from 79 specimens: 9 from Bancali,

38 from Chiaramonti and 28 from C.Tuvullao (25

from layer I and 3 from layer II) and 4 from Bon-

nanaro-San Giorgio.

RESULTS: MORPHOLOGICAL FEATU-

RES OF AMPHIOPE FROM SARDINIA

Lunules. Though a large variability is present in

the outline, lunules are commonly large-sized and

subcircular (Plate 4 Fig. 14c) to broad elliptical (Plate

4 Fig. 14b) at Bancali, Ardara, Villa S. Antonio,

Bonnanaro and San Giorgio. Similar shaped, but

smaller, lunules occur at Porto Torres and La Crucca,

whereas they are mainly broad elliptical (Plate 1

Figs. 1 , 2) at Chiaramonti and narrow-elliptical at C.

Tuvullao (Plate 2 Figs. 13b-d). Specimens from

different localities bearing similar-shaped lunules

often have different external test characters and/or

internal structure, so that test and lunules features are

not univocally linked. Lunules may be different even

in a single specimen (e.g. Plate 1 Fig. 3a).

Plate structure. A number of constant struc-

tural features are recognized in the studied material:

250

Paolo Stara & Enrico Borghi

- Interambulacral columns are always disjunct

adorally, plate 2b and sometime both plates 2a and

2b being separated from the basicoronals by en-

larged first ambulacral post-basicoronal plates.

- Oral interambulacra 1, 4, 5 are meridoplacous,

whereas interambulacra 2 and 3 may be either

amphiplacous or, more frequently, meridoplacous.

- The first post-basicoronal plates are the longest

of the series, both in ambulacral and interambu-

lacral columns. In interambulacrum 5, the plate 2b

is more elongate than 2a.

- There always is a higher number of plates abo-

rally than adorally in each column.

- There is almost the same number of plates in

each column of interambulacra 1, 4, 5 and ambula-

cra I and V: 14-15, seldom 16, plates in the speci-

mens of Bancali and Chiaramonti, whereas they are

more numerous (16-20) at C. Tuvullao. The same

condition occurs in interambulacra 2 and 3, and in

ambulacra II, III and IV: 14-16 plates in each col-

umn at C. Tuvullao I, 12-14 at Bancali and only 10-

13 at Chiaramonti.

- Lunules walls have vertical sutures (PI. 4 Fig.

8) corresponding to the Echinodiscus - type (“cross

linked” sutures, sensu Mooi, 1989). Adorally, lunules

begin to open in correspondence of the second pair

of post-basicoronal plates, whereas 1-2 couples of

plates separate lunules from the petal tips, aborally.

Plates encircling lunules are more numerous on the

aboral side (6-9) than adorally (3-5).

The composition of the oral interambulacrum 5

is characteristic (Table 2): at C. Tuvullao I the nor-

mal condition is 3 (33% of cases; Plate 2 Fig. lb)

to 4 plates (67%) in column 5b, whereas in all the

other samples there are only 2 post-basicoronal plates

in column 5a and 3 in 5b (Plate. 4 Figs. 11, 12).

Also the position of the periproct is highly char-

acteristic (Table 2): the specimens of C. Tuvullao II

have the periproct far from the posterior test margin

and bounded by the first pair of post-basicoronal

plates 2a/2b (Plate 3 Figs. 9b, 10b). At Bancali,

Bonnanaro and San Giorgio the periproct is invari-

ably associated with plates 2a/3b (Plate 3 Figs. 2a,

b; Plate 4 Figs. 11, 12), usually positioned halfway

along the suture. This is the most frequent condition

also at Chiaramonti, but at this locality the periproct

opens more distally, near the outer edge of plates

2a/3b towards plate 3a (Plate 1 Fig. 3b), or just at

the conjunction point 2a/3b/3a (9 cases of 27; Plate

1 Fig 4b), sometimes even between 3a/3b (3 cases).

At C. Tuvullao I the periproct is commonly found

(21 cases of 41 = 51%) at the conjunction point

2a/3a/3b (Plate 2 Figs, lb, 2b) or more distally

(22%; Plate 2 Fig. 3).

The interambulacral basicoronal circlet is very

variable in shape and size, even at the same locality.

The ambulacral circlet is more homogeneous: the

mean value of L13 is 11.3% TL at Bancali and

10.8% at C. Tuvullao I; it is smaller at Chiaramonti,

Bonnanaro and San Giorgio.

Locality

Position of the periproct

Plates present in the

ini era mb u Inc rum 5 adorally

2b fin

2b/2a/3b

2 a Jib

2a/3b/3a

3 b/3 a

3a/4l>

3b/3a/4b

3b/4a/4b

3a/4b

4a; 4b

4a/5b

5a/5b

Ardara

Bancali

Bonnanaro

Chiaramonti

C .Tuvullao

■*

C.TuvulIao

(ID

La Crucca

Porto

Torres

Senis

Table 2. Plates present in the interambulacrum 5 adorally and position of the periproct in the examined samples of Am-

phiope from Sardinia.

The echinoid genus Amphiope L Agassiz, 1840 (Echinoidea Astriclypeidae) in the Oligo-Miocene of Sardinia (Italy) 251

Internal test structure. It consists of a well

developed peripheral ballast system surrounding a

central cavity with domed ceiling. The extension of

the central cavity roughly corresponds to the peta-

loid area; adorally, it does not extend beyond the dis-

tal border of the first pair of post-basicoronal

ambulacral plates, in all samples. In the specimens

of Chiaramonti the central cavity has an almost flat

floor and is bordered by thin transversely elongate

straight walls delimiting a sub-pentagonal area (Plate

1 Fig. 8). At Bancali, Bonnanaro and San Giorgio

the first pillars of the radiating interambulacral but-

tresses (Plate 3 Fig. 5a, Plate 4 Fig. 7) are stronger

and get closer to the centre, thus forming an almost

“starring” outline (Plate 4 Fig. 15). At C. Tuvullao

I the floor of the interambulacra begin to thicken

close to the peristome and gradually rise towards

the first pillars of the radial supports (Plate 2 Fig. 11);

the central cavity has a rough sub-circular outline,

with floor and ceiling thicker than in the specimens

from Bancali and much thicker than those from

Chiaramonti, Bonnanaro and San Giorgio (Plate 3

Fig. 7). The convexity present on the external surface

of the petals contributes to strengthen the ceiling. In

the specimens from C. Tuvullao I the whole petal

surface is convex whereas only the interporiferous

areas are convex at the other localities.

The lantern muscle attachment structures always

consist of five fused interradial pegs (Plate 1 Fig.

8). The peripheral ballast system is made of a series

of sub-cylindrical pillars and walls extending from

the ceiling to the floor and crossed by cavities.

Towards the ambitus it becomes very dense, almost

massive, and crossed by micro-canals, in all the

examined samples.

On the whole, the support system is strongly

developed in the specimens of C. Tuvullao I and II,

whereas it is lighter and more complex at Bancali

and much lighter at Bonnanaro, San Giorgio and

Chiaramonti.

RESULTS OF BIOMETRIC ANALYSES

The PCA analysis resulted in three components

accounting for more than 70% of the total variance

in the data set. The first (PCI) explains 35.5% of

the variance; the ratios TW/TL and TH/TL enter

the heaviest loading into PCI. The second compo-

nent (PC2) is mainly controlled by L4/TL, the third

(PC3) by the lunule dimension variables. On the

whole, the PCA scatterplot (Fig. 4) shows large

overlaps in the distributions of specimens from the

different localities in multivariate space, delimited

by convex hulls. Only the samples from C. Tuvullao

I and Bonnanaro are clearly separate from that

of C. Tuvullao II. According to the PCA analysis

results, the specimens from C. Tuvullao I and II

present higher test and lunules with lower values of

LI and higher values of L2.

Descriptive statistics resulting from the univari-

ate analysis are reported in Table 3. The bivariate

plots confirm that the specimens from C. Tuvullao

II have much higher test (Fig. 3), the length of the

frontal ambulacrum and the distance of the periproct

from the posterior test margin have more elevate

values than those from layer I, the apical disc is

more centrally located. The sample from C. Tuvul-

lao I has higher tests and more transversely elon-

gated lunules than those from the other localities

(Fig. 5), with the exception of C. Tuvullao II. The

specimens from Chiaramonti and Bonnanaro-San

Giorgio have a very low test (Fig. 3). At Chiara-

monti the frontal petal has almost the same length

of the posterior petals (mean L9 = 95.1% L5),

whereas it is slightly longer at C. Tuvullao I, Ban-

cali, Bonnanaro (mean of L9 ranging from 86.6 to

88.8% L5), and much longer at C. Tuvullao II

(mean L9 = 79.1% L5, only).

On the whole, the statistical analyses indicate

the presence of five different morphotaxa in the stud-

ied material, corresponding to the samples from C.

Tuvullao I, C. Tuvullao II, Bonnanaro-San Giorgio,

Chiaramonti, Bancali.

DISCUSSION

The material under study indicates a large vari-

ability of the morphological characters stated to sep-

arate the species of Amphiope recorded from

Sardinia, mainly consisting of the external test fea-

tures: test outline, size and shape of lunules and

petals (Cotteau, 1877, 1895; Lovisato, 1911, 1914;

Cottreau, 1914; Comaschi Caria, 1955, 1972;

Philippe, 1998).

Using these “diagnostic” characters, Comaschi

Caria (1955) recognized seven species at C. Tuvullao I.

252

Paolo Stara & Enrico Borghi

Figure 3. Amphiope from five Sardinian localities: bivariate

plot of test height (values of H divided by TL) against test

length (TL, in mm). Legend: Ba = Bancali; Bon = Bonna-

naro; Ch = Chiaramonti; Ct I and Ct II = Cuccuru Tuvullao,

layer I and layer II, respectively.

Figure 4. Scatter diagram of PC A analysis based on speci-

mens of Amphiope from five Sardinian localities. The legend

is reported in figure 3.

Figure 5. Amphiope from five Sardinian localities: bivariate

plot of L1/L2 ratio against the test length (TL, in mm). The

legend is reported in figure 3.

However, intermediate cases are present in the

sample studied from this locality. On the other hand,

basing on the recent interpretation of A. bioculata

by Philippe (1998) as a taxon with large morpho-

logical variability and stratigraphical distribution,

the examined material from Sardinia should be at-

tributed to a single species. The specimens bearing

the “diagnostic” characters of A. lovisatoi, A. pallavi-

cinoi and A. calvii could be included in the vari-

ability range of “population B” of A. bioculata de-

scribed by Philippe (1998; fig. 15 b, e; pi. 16, fig.

6). The same case occurs with the A. transversivora-

A. deydieri-A. hollandei group, which is close to

“population A” (Philippe 1998; fig. 14 d-f; pi. 16,

fig. 4), and with A. montezemoloi, apparently cor-

responding to “population D” (Philippe 1998; fig.

17d, pi. 16, fig. 5).

The results of the biometric analyses clearly

show a number of significant differences and point

to the occurrence of at least five different morpho-

taxa in the studied sample.

To tackle the uncertainty, the analysis of the

structural features is introduced in the taxonomy of

Amphiope. The plate patterns and the inner test

structure have already been utilized in the classifi-

cation of other clypeasteroid genera (Durham,

1955; Lohavanijaya, 1965; Mooi, 1989; Jansen &

Mooi, 2011). The arrangement of the plates in echi-

noids is fixed early in ontogeny and the basic pat-

tern does not change during further growth in most

forms, especially in scutelliform echinoids in which

the plates forming the sharp ambitus are of special

form and prevent plate translocation from the aboral

to the oral side during later growth (Kroh, pers.

comm., 2012). The taxonomic potential of the plate

structure has been recently tested by Stara & Sanciu

(2014) in Echinodiscus Leske, 1778, a genus closely

related to Amphiope. The results indicate that the

position of the periproct, the shape and the number

of plates in the oral interambulacrum 5 clearly sep-

arate E. auritus Leske, 1778 from the other extant

species of Echinodiscus. These features are ex-

pected to provide a taxonomic potential also in Am-

phiope. Indeed, structural features are well pre-

served in the studied material from Sardinia and

show a low variability; additionally, the structural

differences identified between the Sardinian sam-

ples match with the diversities indicated by the

morphometric analyses.

The echinoid genus Amphiope L Agassiz, 1840 (Echinoidea Astriclypeidae) in the Oligo-Miocene of Sardinia (Italy) 253

Basing on the external test morphology, the

study of the structural characters and the statistic

biometric analyses, the specimens from Chiaramonti

(PL 1) are assigned to A. lovisatoi since they were

collected from the type locality of this species and

they are consistent with the original description and

illustration given by Cotteau (1895) and Lovisato

(1914): test very low with sharp margin and rather

broad, transversely elongated elliptical lunules. The

sample from this locality is clearly separated from

the others by the combination of structural charac-

ters: only two post-basicoronal plates present in the

oral interambulacral column 5a, periproct located in

the posterior part of the suture between plates 2a/3b,

internal structure very light with thin shell and sub-

pentagonal central cavity.

Well-preserved material recently collected from

San Giorgio, the type-locality of A. montezemoloi

Lovisato, 1911, and Bonnanaro, corresponds to the

original description of this species: large-sized and

anteriorly constricted test, with broad subcircular

lunules (Plate 3 Figs. 3a, b). A. montezemoloi differs

statistically from A. lovisatoi by a much larger test.

Additionally, the specimens from Chiaramonti have

smaller and more elongate lunules, the frontal petal

almost as long as the others, the internal structure

is lighter and the central cavity is subpentagonal not

starring as in A. montezemoloi.

The holotype of A. nuragica (Plate 2 Figs, la, b)

has four post-basicoronal plates in the oral interam-

bulacral column 5b, three in column 5a, and the

periproct is located at the conjunction of plates

2a/3a/3b. Comparison with the specimens collected

from the type-layer (C. Tuvullao I) with the holo-

type is entirely consistent. No significant statistical

differences were observed between the specimens

with the periproct close to the conjunction point 2a,

3 a, 3b and those with the periproct more posteriorly

located. The characteristic and strong internal test

structure is present in all the specimens from this

bed. Thus, the whole sample from C. Tuvullao I is

assigned to A. nuragica (Comaschi Caria, 1955).

The combination of the peculiar structural features

distinguishes A. nuragica from all the other Sar-

dinian examined samples.

The specimens from C. Tuvullao II stand apart

from all the others from Sardinia by much higher

test, apical system more centrally located, frontal petal

much longer than the posteriors and the periproct

bounded by the first pair of post-basicoronal plates.

Due to the scarcity of the available material these

specimens are assigned to Amphiope sp. 1, and left

in open nomenclature.

The specimens of Bancali (Plate 4 Figs. 1-3),

though rather similar to those from San Giorgio-

Bonnanaro, differ statistically from A. montezemo-

loi by more elevate test, larger ambulacral

basicoronal circlet and smaller lunules, and by a

stronger internal structure. The same differences sep-

arate them from A. lovisatoi; additionally, the spec-

imens from Chiaramonti are smaller and the

periproct, though bounded by plates 2a/3b as well

as in the sample of Bancali, is closer to the conjunc-

tion point 2a/3b/3a. Though the studied sample

looks like well differentiated, additional well-

preserved material is needed to corroborate the ob-

served differences and to confirm the occurrence of

a distinct species. Therefore, the specimens from

Bancali are assigned to Amphiope sp. 2.

Only scant and poorly preserved material is cur-

rently available from the type-localities of A. dessii

Lovisato in Cotteau, 1 895, A. pallavicinoi Lovisato,

1914 and A. calvii Lovisato, 1914. The oral plating

as well as the internal support arrangement of those

species were not reported in the original descriptions

and cannot be made out from illustrated specimens.

Cotteau (1895) attributed a test fragment from Sar-

dinia to A. hollandei Cotteau, 1877. Other Miocene

specimens from Sardinia were attributed to this

species (Lovisato, 1911; 1914). Since all that mate-

rial was lost (fide Comaschi Caria, 1955) the occur-

rence of A. hollandei in Sardinia is not confirmed.

Most of the Amphiope species described in the

literature lack primary data on the plating patterns

and the internal test structure and are therefore not

completely documented in terms of their morphol-

ogy. This is the case also for the type-species of Am-

phiope: the type locality and stratum of the

specimen “ variete 3 ” of Scutella bifora (see Lamarck,

1816) on which des Moulins (1837) based the diag-

nosis of Amphiope bioculata, are unknown; des

Moulins tentatively proposed “ terrain tertiaires ” of

Suze la Rousse in the Rhone Basin and Bordeaux

(France) as type-localities for that specimen. No

well preserved specimens from these localities have

been traced in public institutions (Philippe, 1998;

pers. comm. B. Martin Garin, March 2013) and the

plate patterns could not be taken from figures re-

254

Paolo Stara & Enrico Borghi

ported in the literature (Lambert, 1912). Philippe

(1998), when studying Amphiope from the Rhone

Basin, described also specimens from the Serraval-

lian of Suze la Rousse, but he could not describe

the plate patterns since they were not preserved.

As a consequence, the structural features of the

type-species, as well as of most of the earlier de-

scribed species of Amphiope, are still uncertain/

unknown thus preventing a reliable comparison with

the material under study, based on these important

characters.

SYSTEMATIC PALAEONTOLOGY

Family ASTRICLYPEIDAE Stefanini, 1912

Genus Amphiope L. Agassiz, 1840

Type species. Scutella bioculata des Moulins,

1837, by subsequent designation of Lambert (1907,

p. 49).

Emended diagnosis. (Partially modified from

Smith & Kroh, 2011). Test low with sharp margin.

Internal support well developed, consisting of

pillars and walls crossed by cavities. Towards am-

bitus, peripheral ballast system very dense, almost

massive and crossed by micro-canals. Apical disc

monobasal, sub-central or slightly anterior to cen-

tre, with four gonopores. Petals well developed;

short (about half radial length of test) and almost

closed distally. All five petals similar in length.

Ovate lunules or notches present in the posterior

ambulacra. Oral side flat or slightly concave. In-

terambulacra on the oral surface narrower than the

ambulacra, even at their widest point. Interambu-

lacra 1, 4 and 5 always meridoplacous adorally,

the interambulacral zones being separated by

enlarged first post-basicoronal ambulacral plates.

Interambulacra 2, 3 may be either amphiplacous

or meridoplacous adorally. Basicoronal circlet

pentastellate with interambulacral plates forming

the points. Peristome small, subcentral or slightly

anteriorly located. Periproct circular, small, open-

ing between the first, the second or the third pair

of post-basicoronal interambulacral plates. Two to

five post-basicoronal plates present in the interam-

bulacrum 5 adorally. Food grooves well devel-

oped, bifurcating at the edge of the basicoronal

plate; they do not reach the margin.

Posterior pair of food grooves running around

the lunules; finer distal branches well developed.

Ambulacra a little wider than interambulacra at am-

bitus. Tuberculation dense, made of very small, per-

forate and crenulate tubercles, larger on the oral

face than aborally.

Distribution. Oligocene and Miocene. Central

and Southern Europe, North Africa, Middle East,

India, Angola (Smith & Kroh, 2011).

Sardinian species included:

• Amphiope lovisatoi Cotteau, 1895. Late Burdi-

galian.

• Amphiope montezemoloi Lovisato, 1911. Late

Burdigalian- early Langhian.

• Amphiope nuragica (Comaschi Caria, 1955). Late

Chattian-early Aquitanian.

• Amphiope sp. 1. Late Chattian-early Aquitanian.

• Amphiope sp. 2. Late Burdigalian- early Langhian.

Remarks. Both Amphiope and Echinodiscus

Leske, 1778 show two lunules in the posterior am-

bulacra. Echinodiscus differs in having axially elon-

gated, slit-like lunules or notches and posterior

petals shorter than the others.

Amphiope lovisatoi Cotteau, 1895

Plate 1 Figs. 1-8; Plate 3 Fig. 7b

1895 Amphiope Lovisatoi Cotteau - Cotteau p. 16

pi. 3, fig. 15

1914 Amphiope Lovisatoi Cotteau - Lovisato, p.

118, pi. 2, figs. 6a-b

Non 1955 Amphiope lovisatoi Cotteau - Comaschi

Caria, p. 9, pis. 9, 11, 12.

Type-locality and horizon. Chiaramonti (Sas-

sari). The type-layer described by Lovisato (1914)

corresponds to the “Facies C” of Mancosu &

Nebelsick (2013; Fig. 4), attributed to the lower part

of the Calcari di Mores Formation, dated to the late

Burdigalian.

Type material. Cotteau (1895) did not detail

the composition of the type-series provided by

Lovisato, nor designated a holotype. All those

specimens, as well as the others from Chiaramonti

belonging to the Lovisato’s collection, were lost

in 1943 (fide Comaschi Caria, 1955). They could

not be traced by the authors at UNICA. A neotype

is proposed herein, to clarify the diagnostic char-

The echinoid genus Amphiope L Agassiz, 1840 (Echinoidea Astriclypeidae) in the Oligo-Miocene of Sardinia (Italy) 255

acters of this nominal taxon which cannot be ex-

tracted from published descriptions and illustra-

tions of the lost type. Additionally, A. lovisatoi

has been synonymised with other species of Am-

phiope, and the structure features firstly described

in this paper are not visible in the illustration pro-

vided by Cotteau (1895) and Comaschi Caria

(1955).

Neotype: MAC.PL1706 (PI. 1 Figs.la-c and

5a-b), a specimen with both faces well preserved

(TL = 73, TW = 77.5, TH = 8.5 mm). It was recov-

ered at Monte Sa Loca (40°44’55.15” N,

8°49’59.20” E), from the main Amphiope-bearing

bed (Facies C of Mancosu & Nebelsick, 2013),

which represents the lateral extension of the type-

layer cropping out at Billiu (Stara et al., 2012).

Billiu is less thanl km far from Monte Sa Loca,

both are located in the suburbs of Chiaramonti.

The road-cut of Billiu currently yields only scarce

and poorly preserved fossil material, whereas at

Sa Loca abundant and well preserved material is

available to study.

Examined material. The studied sample from

the type-locality consists of 61 complete tests

(MAC code: PL1301-3, PL1317, PL1413, PL1418-

20, PL1422-4, PL1427, PL1429, PL1567-70,

PL1572-80, PL1583, PL1585-7, PL1692-99,

PL1700-1707, PL1709-1714, PL1715-18, PL1720-

23, PL 1726) and 7 test fragments.

Revised diagnosis. Middle-sized species of

Amphiope with low test, sharp margin and broad,

transversely elongated elliptical lunules. A low

number of plates is present in each ambulacral

and interambulacral columns, only two post-basi-

coronal plates occur in the interambulacral col-

umn 5a adorally. Periproct bounded by plates

2a/3b, rather close to the posterior test edge. In-

ternal structure very light, with thin shell and sub-

pentagonal central cavity.

Description. Middle sized test (mean TL= 76

mm). Outline sub-circular to anteriorly con-

stricted, usually slightly transversely elongated,

posteriorly rounded (Plate 1 Fig. 1) or subtruncate

(Plate 1 Fig. 3a). Maximal width located subcen-

trally. Test very low (mean TH = 10.2% TL), with

maximum height positioned anteriorly. Marginal

indentations well developed in ambulacra II and

IV (Plate 1 Figs. 1, 2). Shallow notches may occur

also in ambulacra I, III, V and interambulacra 1

and 4. Test edge sharp.

Ambulacra . Frontal and posterior petals similar

in length. Poriferous zones depressed; interporifer-

ous zones slightly raised. Interporiferous area larger

than the corresponding poriferous one. Lunules

transversely elongate and broad elliptical (Plate 1

Figs. 1, 2 and 7d); the shape-variability includes

also rare small subcircular (Plate 1 Fig. 7c) and nar-

row elliptical (Plate 1 Fig. 7a) lunules. Ambulacra

slightly depressed adorally, along their central suture.

Ambulacral basicoronal circlet small.

Interambulacra . Only 14-16 plates in each col-

umn of interambulacra 1 , 4, 5 and ambulacra I and

V; 10-13 plates in interambulacra 2 and 3, as well as

in ambulacra II, III and IV. Adorally, only two, some-

times a small part of the third, post-basicoronal plates

in the interambulacral column 5 a adorally, three of

them in column 5b (Plate 1 Figs. 3b, 4b, 5b).

Periproct . Commonly found in the distal half of

the suture 2a/3b (Plate 1 Figs. 3b, 5b).

Internal structure . Reduced, with spaces be-

tween elements larger than the calcite elements

comprising the buttress system (Plate 1 Fig. 6).

Central cavity bordered by five transversely elon-

gate straight walls, delimiting a sub-pentagonal area

(Plate 1 Fig. 8). Peripheral ballast system very

dense, almost massive towards the ambitus.

Other features as for the genus . See Table 3 for

descriptive statistics.

Remarks. Most of the “diagnostic” features

stated for A. lovisatoi by Cotteau (1895) and Lo-

visato (1914) are very variable in the studied sam-

ple from Chiaramonti and cannot provide a clear

separation from the other Sardinian species. Some

specimens do not correspond to “posteriorly rounded

test” (e.g. Plate 2 Fig. 3a), or to “deep notches in

the margin” (e.g. Plate 2 Fig. 4a), as stated for this

species. Only “test middle-sized and depressed” and

“sharp margin” are confirmed as valid distinctive

characters by this study.

A. lovisatoi was synonymised with A. bioculata

by Kroh (2005), however, the figured specimen

from Austria differs from the Sardinian fossils by

much higher test (TH = 20% TL) and by the frontal

petal, which is clearly longer than the posterior

paired petals (L9/L5 = 0.80 in the Austrian speci-

men, against 0.95 in the sample of Chiaramonti).

Comaschi Caria (1955) cited this species at C.

Tuvullao. However all the specimens from that

locality bearing external features similar to those of

A. lovisatoi had very different plate patterns and

256

Paolo Stara & Enrico Borghi

much stronger internal structure, corresponding to

that of A. nuragica.

Occurrence in Sardinia. Chiaramonti, Calcari

di Mores Formation, late Burdigalian.

Amphiope montezemoloi Lovisato, 1911

Plate 3 Figs. 1-6, 7a

1911. Amphiope Montezemoloi Lovisato - Lovisato,

p. 43, pi. 6, figs, la, b

1928. Amphiope montezemoloi Lovisato - Lambert

[36], p. 23, pi. 8, fig. 4

1955. Echinodiscus ( Amphiope ) bioculata var. mon-

tezemoloi Comaschi Caria - Comaschi Caria, p.

184, pis. 14, 15

1972. Amphiope bioculata des Moulins - Comaschi

Caria, p. 42.

Type-locality and horizon. Near the aban-

doned railway station of San Giorgio (Sassari) to-

wards Olmedo. The Amphiope- bearing outcrop,

belongs to the Calcari di Mores Formation, late

Burdigalian- early Langhian (Carmignani et al.,

2001 ).

Type material. Lovisato (1911) did not des-

ignated a holotype nor detailed the composition of

the type-series. According to Comaschi Caria

(1955) all those specimens were lost; they were not

traced by the authors at the UNICA. The validity of

A. montezemoloi is still debated, since it has been

synonymised with other Amphiope species, and the

plate patterns were not described. Thus, a neotype

is herein designated.

Neotype: MAC code PL1827 (Plate 3 Fig. la, b).

It consists of a large complete specimen with oral

structure partially visible (TL =116, TW = 133, TFI

= 10, Lll = 14 mm). It comes from Stazione di San

Giorgio, Sassari, the type locality of this species

(40°41 , 13.68 ,, N, 8°27’03.41”E), from the Calcari

di Mores Formation.

Examined material. The studied material

includes also 2 large test fragments from the type-

locality (PL 1828-9) and 3 specimens (PL 1674-6)

from Bonnanaro (Sassari).

Revised diagnosis. Large sized species with

low lateral profile and very large, subcircular to

slightly transversely elongate, lunules. Frontal petal

longer than the others. In the oral interambulacrum

5, plate 2b very elongate and only 2 post-basicoro-

nal plates present in column 5a. Ambulacral basi-

coronal circlet very small. Internal structure

reduced, central cavity with starring outline.

Description. Large sized test with transversely

elongate and anteriorly restricted outline. Test veiy

low (mean TH = 9.8% TL), almost flattened adapi-

cally. Margin posteriorly sharp, anteriorly more

rounded and 3-3.5 mm thick.

Ambulacra . Frontal petal longer than the poste-

riors. Poriferous zones depressed; interporiferous

zones slightly raised. Lunules very large, subcircu-

lar to broad elliptical with moderately transversely

elongate outline. Ambulacra slightly depressed ado-

rally along their central suture.

Interambulacra . The first post-basicoronal plate

(2b) in the oral interambulacrum 5 is very elongate

(Plate 3 Figs. 2a, b) and plate 2a is close to the pos-

terior margin. Three post-basicoronal plates are

present in column 5b, only two plates in column 5a.

Periproct . Small (mean diameter = 1.7% TL),

rather close to the posterior test margin and located

along the suture 2a-3b.

Internal structure . Rather complex but reduced,

with thin shell and large spaces between the calcite

elements comprising the buttress system (Plate 3

Figs. 5a, c). Central cavity large (Plate 3 Figs. 4, 7b)

with flat and thin floor. On the ceiling, the inter-

poriferous areas of the petals are convex, the porif-

erous areas slightly concave. Peripheral ballast sys-

tem dense towards the ambitus.

Other features as for the genus . See Table 3 for

descriptive statistics.

Remarks. Large-sized test and broad subcircu-

lar to elliptical lunules (Lovisato, 1911) are confir-

med by this study as distinctive features of A.

montezemoloi. The other “diagnostic” characters

stated in the original description cannot provide a

clear separation from the other examined species.

In particular, the “irregularities on the aboral sur-

face” described by Lovisato (1911) were not ob-

served in any specimen. The apical disc is “eccentric

towards the anterior test edge”, but the measures of

the two specimens reported by Lovisato (1911), as

well as the mean of L4 in the examined sample of A.

montezemoloi , almost correspond to those of A. lo-

visatoi and Amphiope sp. 2 (see Table 3).

The echinoid genus Amphiope L Agassiz, 1840 (Echinoidea Astriclypeidae) in the Oligo-Miocene of Sardinia (Italy) 257

A. montezemoloi was placed into synonymy

with A. bioculata by Comaschi Caria (1955),

Philippe (1998) and Kroh (2005). However, since

at present a comparison with A. bioculata based on

the structural characters is not possible, the separa-

tion between the two is herein maintained.

Occurrence in Sardinia. San Giorgio and

Bonnanaro (Sassari), Calcari di Mores Formation,

late Burdigalian- early Langhian.

Amphiope nuragica (Comaschi Caria, 1955)

Plate 2 Figs. 1-13

1955. Echinodiscus ( Amphiope ) nuragica Co-

maschi Caria - Comaschi Caria, p. 186, pi. 1

1955. Echinodiscus (Amphiope) deydieri Lambert -

Comaschi Caria, p. 186, pis. 2-3

1955. Echinodiscus ( Amphiope ) transversivora

Lambert - Comaschi Caria, p. 187, pi. 4

1955. Echinodiscus (Amphiope) pallavicinoi Lo-

visato - Comaschi Caria, p. 188, pis. 5-6, pi. 13,

fig. 2

1955. Echinodiscus (Amphiope) calvii Lovisato -

Comaschi Caria, p. 188, pis. 7-8, pi. 13, fig. 1

1955. Echinodiscus (Amphiope) lovisatoi Cotteau -

Comaschi Caria, p. 189, pi. 9; PI. 11, fig. 2, pi.

1955. Echinodiscus (Amphiope) dessii Lovisato -

Comaschi Caria, p. 190, pi. 10, pi. 11, fig. 1.

Type material. Holotype (UNICA code 9CC.8-

10504; Plate 2 Figs, la, b and 9). TL = 106, TW =

107, TH =16 mm. It was the sole specimen attrib-

uted to this species by Comaschi Caria (1955).

Type-locality and horizon. Layer I of Cuc-

curu Tuvullao (Cagliari; 39°47’ 18.88” N,

9°26’55.54” E), corresponding to facies “B” of

Mancosu & Nebelsick (2013), Nurallao formation,

Arenarie di Serralonga member, late Chattian-early

Aquitanian.

Examined material. Two additional whole tests

from C. Tuvullao I (UNICA, code 3CC and 6CC-

10503). Nineteen specimens and 66 large test frag-

ments from the same layer are housed at the MAC

(PL1590-1, PL1678-80, PL1684, PL1727, PL1820,

PE1829; PL1835-1844).

Revised diagnosis. Test high, with narrow and

transversely elongate lunules. Petal surface convex,

including the poriferous areas. High number of

plates in ambulacral and interambulacral columns.

Three to four post-basicoronal plates in each

column of the oral interambulacrum 5. Periproct

close to the conjunction of plates 2a/2b/3a, or more

posteriorly located. Internal support system strongly

developed, with thick shell and roundish outline of

the central cavity.

Description. Medium to large sized test. Out-

line subcircular in the holotype, but more frequently

transversely elongate and restricted anteriorly (PI.

2 Figs. 5-7). Maximal width located subcentrally.

Test high (mean TH = 14.2% TL). Marginal inden-

tations slightly developed in ambulacra II and IV,

shallow notches may occur also in ambulacra I, III,

V and interambulacra 1 and 4. A shallow but

distinct anal notch may be present. Posterior edge

thin, the anterior margin is thicker.

Ambulacra . Petals almost closed distally (Plate

2 Fig. 12), the frontal one slightly longer than the

others. Maximal petal width about one half to two-

thirds of petal length. External petal surface slightly

raised and convex, including the interporiferous

zones. Angle between the axis of posterior petals

large: mean a = 76°, against a mean of 71° at the

other localities. A transversely elongate lunule is

present in each posterior ambulacrum. Shape and

size of lunules variable (PI. 2, Figs. 13a-d). They

are commonly narrow (Plate 2 Fig. 13b) to rather

broad elliptical (Plate 2 Fig. 13a), seldom sub-

polygonal (Plate 2 Fig. 7). A small protuberance is

present along the internal margin of the lunules in

the holotype and in another specimens (Plate 2

Figs. 1, 3). Ambulacra slightly depressed adorally

along their central suture. Ambulacral basicoronal

circlet rather large.

Interambulacra . There are 1 6-20 plates in each

column of interambulacra 1, 4, 5, as well as in am-

bulacra I and V; 14-16 plates in interambulacra 2

and 3 and in ambulacra II, III and IV. Adorally, at

least 3, frequently also 4, post-basicoronal plates

present in each column of the interambulacral

column 5a adorally (Plate 2 Fig. lb, 3, 4).

Peristome . Small (mean diameter = 3.3% TL).

Periproct . It opens along the distal half of the

suture 2a/3b, close to the conjunction point

2a/3b/3a (as in the holotype, compare Plate 2 Fig.

258

Paolo Stara & Enrico Borghi

lb, 2b) or more posteriorly located (Plate 2 Fig.

3). It is close to the posterior test edge (mean LI 1 =

9.1 % TL).

Internal structure . Well developed, with thick

shell and spaces between elements narrower than the

elements comprising the buttress system (Plate 2

Fig. 11). The floor of the central cavity in the inter-

ambulacra begins to thicken close to the peristome

and gradually rises, extending radially towards the

first pillars of the radial supports (Plate 3 Fig. 7c).

Central cavity with a rough sub-circular outline.

Other features as for the genus . See Table 3 for

descriptive statistics.

Remarks. The noticeable test height (TH =

14.2% TL), the periproct close to the posterior

edge (Lll = 9.1% TL) and the characteristic

shape and proportion of lunules, stated by Co-

maschi Caria (1955) for A. nuragica, are confirmed

by this study as valid distinctive characters. The

other “diagnostic” features are variable in the stud-

ied sample and cannot provide a clear separation

from the other examined species. In particular, ir-

regularities in the lunules outline, as the promi-

nences described by Comaschi Caria (1955),

occur randomly in other species (Plate 1 Fig. 7e;

Plate 2 Figs. 1, 3) and have no taxonomic value.

The morphology and the plate patterns of two spec-

imens assigned by Comaschi Caria (1955) to A.

dessii and A. calvii (UNICA, code 6CC-10503

and 3CC, respectively), correspond to those of A.

nuragica and are therefore assigned to this species.

The same case occurs with specimens bearing

external features corresponding to A. deydieri, A.

transversivora , A. pallavicinoi, A. calvii and A.

lovisatoi, all species recorded from this locality

by Comaschi Caria (1955); additionally they are

not statistically separable from the others from C.

Tuvullao I.

Occurrence in Sardinia. Cuccuru Tuvullao

(Cagliari), Nurallao formation, Arenarie di Serra-

longa member, late Chattian-early Aquitanian.

Amphiope sp. 1

Plate 3 Figs. 8-10

Examined material and horizon. 3 specimens

(MAC code PL1681, PL1685 and PL1834) and 1

test fragment (PL 1684), from layer II of C.Tuvullao

(Cagliari, 39°47’18.88”N, 9°26’55.54”E), corre-

sponding to facies “C” of Mancosu & Nebelsick

(2013), Nurallao formation, Arenarie di Serralonga

member, late Chattian-early Aquitanian.

Description. Middle to large sized test with

thick margin. Outline transversely elongate and

slightly anteriorly restricted. Maximal width located

subcentrally. Test very high (mean TH = 20.7%

TL), with maximal height slightly anterior of the

apical disc (Plate 3 Fig. 8c). Marginal indentations

slightly developed in ambulacra II, III and IV. Faint

notches may occur also along the posterior test

edge, in ambulacra I and V and interambulacra 1

and 4.

Ambulacra . Frontal petal distinctly longer

than the posteriors (mean L9 = 79.1% L5).

Maximal petal width about one half to two-thirds

of petal length. Poriferous zones depressed; in-

terporiferous zones slightly raised. Lunules tran-

sversely elongate, narrow elliptical and close to

the tips of the posterior petals (mean L3 = 4.6%

TL). Ambulacra slightly depressed adorally along

their central suture. Ambulacral basicoronal cir-

clet small.

Interambulacra . Adorally, only two, maximum

three, post-basicoronal plates are present in each col-

umn of interambulacrum 5 (Plate 3 Figs. 9b, 10b).

Periproct. Bounded by the first pair of post-

basicoronal plates 2a/2b (Plate 3 Figs. 9b, 10b) and

far from the posterior test margin (mean Lll =

13.8% TL).

Other features as for the genus . See Table 3 for

descriptive statistics.

Remarks. The specimens from C. Tuvullao II

stand apart from all the others examined from Sar-

dinia by their much higher test, more centrally lo-

cated apical disc, frontal ambulacrum much longer

than the posterior ones and lunules closer to the tips

of the posterior petals.

In addition, it differs from A. nuragica by the

presence of only three post-basicoronal plates in

each column of interambulacrum 5 adorally and the

periproct bounded by the first pair of post-basi-

coronal plates.

The plate patterns in oral interambulacrum 5 cor-

respond to the drawings of A. bioculata reported by

Durham (1955), Kroh (2005) and Pereira (2010), but

the attribution of those schemes to the type-species

The echinoid genus Amphiope L Agassiz, 1840 (Echinoidea Astriclypeidae) in the Oligo-Miocene of Sardinia (Italy) 259

is doubtful since they were not based on topo-typic

material. Besides, the specimens of C. Tuvullao II

differ by more transversely elongate lunules.

Occurrence in Sardinia. Cuccuru Tuvullao

(Cagliari), Nurallao formation, Arenarie di Serra-

longa member, late Chattian-early Aquitanian.

Amphiope sp. 2

Plate 4 Figs. 1-15

Examined material. Ten complete specimens

(MAC PL343, PL547-553, PL1665, PL1836) and

ten fragments from Bancali (Sassari, 40°43 , 55.66 ,, N,

8 0 26’55.54” E), collected in a bioclastic sandstone

at “outcrop 2” described by Stara et al. (2012), attrib-

uted to the Calcari di Mores Formation, late Burdi-

galian-early Langhian.

Description. Middle to very large sized test,

with rather thin margin. The test width reached up

to 133 mm in complete specimens, but some frag-

ments points to complete test with length up to

about 170 mm. Outline slightly transversely elon-

gate and restricted anteriorly (Plate 4 Figs. 1-3).

Maximal width located subcentrally. Maximum

test height slightly anterior of the apical disc (Plate

4 Figs. 4-6). Marginal indentations well developed

in ambulacra II and IV, faint indentations may

occur also in ambulacra I, III and V and interam-

bulacra 1 and 4. Oral side flat or slightly concave.

Food groves well developed (Plate 4 Figs. 2, 3), bi-

furcating at the edge of the basicoronal circle. Pos-

terior pair of food grooves running around the

lunules, not reaching the margin; finer distal branches

well developed.

Apical system . Distinctly anterior to centre

(mean L4 = 59.7% TL).

Ambulacra . Frontal petal slightly longer than the

others. Maximal petal width about one half to two-

thirds of petal length. Poriferous zones depressed;

surface of the interporiferous zones slightly convex.

A broad and slightly transversely elongate lunule is

present in each posterior ambulacrum. Lunules

commonly subcircular (Plate 4 Figs. 14c, d) to

broad elliptical (Plate 4 Fig. 14e). Ambulacra

slightly depressed adorally along their central suture.

Ambulacral basicoronal circlet rather large (mean

L13 = 11.3% TL).

Interambulacra . There are 14-16 plates in each

column of interambulacra 1, 4, 5 and ambulacra I

and V, 12-14 plates in interambulacra 2 and 3 and

in ambulacra II, III and IV. Only 3-4 post-basicoro-

nal plates present in each column of oral interam-

bulacrum 5 (Plate 4 Figs. 11, 12).

Peristome . Small (mean diameter = 2.8 % TL).

Periproct . Small (mean diameter = 2% TL), lo-

cated between the second pair of post-basicoronal

plates (Plate 4 Figs. 11, 12) and rather far from the

posterior test margin.

Internal structure . Well developed, with rather

thick shell wall and spaces between elements nar-

rower than the elements comprising the buttress

system. Central cavity with almost starring outline

and interambulacral radial supports extending

towards the centre.

Other features as for the genus . See Table 3 for

descriptive statistics.

Remarks. Amphiope sp. 2 differs statistically

from A. montezemoloi by more elevate test, larger

ambulacral basicoronal circlet and smaller lunules;

it has also a stronger internal structure. The same

differences separate Amphiope sp. 2 from A. lo-

visatoi; additionally, the specimens from Chiara-

monti are smaller and the periproct, though bounded

by plates 2a/3b as well as in the sample of Bancali,

is closer to the conjunction point 2a/3b/3a.

The closely related specimens from Ardara (Sas-

sari) likely belong to this species, though a larger

sample from this locality is needed to confirm this

hypothesis.

Occurrence. Bancali, probably also Ardara,

(Sassari), Calcari di Mores Formation, Late Burdi-

galian-early Langhian.

STRATIGRAPHICAL DISTRIBUTION AND

EVOLUTIVE TRENDS

The genus Amphiope is represented in Sardinia

by forms with rounded or transversely elongate

lunules, only. The first records of Amphiope with

axial lunules are dated to the middle Oligocene of

the Gulf of Biscay, Val Bormida (Piedmont, Italy)

and Libya, whereas Amphiope with transverse

lunules appeared in the late Oligocene-early Miocene

in the area between the Gulf of Biscay and the intra-

AlCaPeKA Basin (Stara & Rizzo, 2013). At that

260

Paolo Stara & Enrico Borghi

Plate 1 .Amphiope lovisatoi Cotteau, 1895, Calcari di Mores Formation, late Burdigalian, Chiaramonti (Sassari). Figs, la-

c Neotype (PL 1706, TL = 73 mm), aboral (a), oral (b) and antero (to the left)-posterior (to the right) lateral view (c). Fig. 2.

Aboral view of PL1317 (MAC code), TL = 82 mm. Figs. 3a, b. Plate patterns of PL1704 (TL = 82 mm); a) aboral side, b)

oral side. Figs. 4a, b. Plate structure of PL 1574 (TL = 80 mm); a) aboral side, b) oral side. Figs. 5a, b. Plate patterns of

PL1706 (TL= 73 mm); a) aboral side, b ) oral side. Fig. 6. Internal structure: cross-section along the radial axis of ambulacrum

I. Figs. 7a-e. Variation range of test outline and lunules shape. All specimens in aboral view, if not otherwise specified, a)

PL1308 (oral view; TL = 78 mm), b) PL1312 (TL = 76.2 mm), c) PL1311 (oral view; TL = 85 mm), d) PL1306 (TL = 89

mm), e) PL1303 (TL = 83.5 mm). Fig. 8. Internal view of oral surface, with almost flat floor and straight walls (E) delimiting

the subpentagonal outline of the central cavity. In Figs. 6 and 8: A= central cavity, B = first peripheral elements (pillars or

walls), C = radial cavity of the interambulacram 5 leading to the periproct, D = lantern supports, E = straight walls at the

periphery of the central cavity, F = massive peripheral support system, G = small cavities. Scale bar equals 1 cm.

The echinoid genus Amphiope L. Agassiz, 1840 (Echinoidea Astriclypeidae) in the Oligo-Miocene of Sardinia (Italy) 261

Plate 2. Amphiope nuragica (Comaschi Caria, 1955), Nurallao formation, late Chattian-early Aquitanian, Cuccuru Tuvullao

(Cagliari). Figs. 1-4, 10 Plate structure drawings with interambulacral plates shaded grey. 1) Holotype (UNICAcode 9CC.8;

TL = 100 mm), aboral (la) and oral (lb) views. 2) PL 1680 (TL = 100 mm), aboral (2a) and oral (2b) views. 3) PL 1591 (TL

= 81 mm), oral view. 4) PL1684 (TL = 91 mm), oral view. 10) PL1835 (TL = 89 mm), aboral view. Fig. 5. PL1836 (TL =

93 mm), aboral view. Fig. 6. PL1835 (TL = 89 mm), aboral view. Fig. 7. PL 1837 (TL = 88.5 mm), aboral view. Fig. 8.

PL1838 (TL = 82 mm), aboral view. Fig. 9. Antero (to the right)-posterior (to the left) lateral view of PL1820 (TL = 98.2

mm). Fig. 11. Close up view of a cross-section through the radial axis of the ambulacrum I. B = first peripheral elements

(pillars), C = radial cavity of the interambulacrum 5 leading to the periproct, F = massive peripheral support system, G =

small cavities, H = floor of the central cavity. Fig. 12. Close up of the petals (PL1 835, TL = 89 mm). Figs. 13a-d. Variability

of test outline and lunules shape. All specimens in aboral view, a) PL 1839 (TL = 59.4 mm), b) PL 183 8 (TL = 82 mm), c)

PL 1840 (TL = 83.6 mm), d) PL1841 (TL = 98.2 mm). Scale bar equals 1 cm.

262

Paolo Stara & Enrico Borghi

Plate 3. A. montezemoloi, San Giorgio and Bonnanaro (Sassari). Figs. la-b. Neotype (PL1827): aboral (a) and adoral (b)

views , TL=121 mm. San Giorgio. Figs 2a-b. Oral plating structure of PL1675 (a) and PL1676 (b). Bonnanaro. Figs. 3a, b.

Specimen PL1676 (TL=90 mm): aboral (a) and oral (b) views. Bonnanaro. Fig. 4. Aboral view of PL1828; margins of the

central cavity marked as dotted lines. San Giorgio. Figs. 5 a- c. Test fragment (PL 1830): adoral (b) and internal views taken

from points x (a) and y (c), respectively. Bonnanaro. Figs. 6a-c. Aboral (a), oral (b) and antero (to the left)-posterior (to the

right) lateral (c) views of PL1675 (TL=1 1 2 mm). Bonnanaro. Figs. 7a-d. Scheme of the internal structure in an antero (to

the right)-posterior (to the left) axial section of specimens from: a) A. montezemoloi , San Giorgio, b) A. lovisatoi , Chiara-

monti, c) A. nuragica, C.Tuvullao, d) Amphiope sp. 2, Bancali. Amphiope sp. 1, Nurallao formation, late Chattian-early

Aquitanian, of Cuccuru Tuvullao (Cagliari). Figs. 8a-c. PL 1834 (TL=95.5 mm); aboral (a), adoral (b) and antero (to the

right)-posterior (to the left) lateral (c) views, respectively. Figs. 9a, b. Plating patterns of PL1681 (TL=90 mm); a) aboral

side, b) oral side. Figs. 10a, b. Plating patterns of PL 1685 (TL=101 mm); a) aboral side, b) oral side. In Figs. 5a-c: A =

central cavity, B=first peripheral elements (pillars or straight walls), C=radial cavity of the interambulacrum 5 leading to

the periproct, F=massive peripheral support system, G=small cavities. Scale bar equals 1 cm.

The echinoid genus Amphiope L. Agassiz, 1840 (Echinoidea Astriclypeidae) in the Oligo-Miocene of Sardinia (Italy) 263

Plate 4. Amphiope sp. 2, Calcari di Mores Formation, late Burdigalian-early Langhian, Bancali (Sassari). Figs. 1-3. 1) PL343,

TL = 73 mm). 2) PL553, adoral view (TL=55.8 mm). 3) PL552, adoral view (TL=73.4 mm). Figs. 4-6. Antero (to the right)-

posterior (to the left) lateral views of PL343 (4), TL=73 mm, PL550 (5), TL=79.5 mm and PL551 (6), TL=88.5 mm. Fig. 7.

Close up view of the internal structure: cross-section of a test fragment along the radial axis of ambulacrum I. B = first pillars

in the radiating buttresses of interambulacrum 5, C=radial cavity of the interambulacrum 5 leading to the periproct, F=massive

peripheral support system, G=small cavities. Figs. 8-9. PL550 (TL=79.5 mm). 8) close up view of the petals; 9) plating

scheme of a lunule (oblique view), with cross-linked wall. Fig. 10. Close up view of the tuberculation in the oral interambu-

lacrum 1; fg=food groove. Figs. 11-13. Plate diagrams, with interambulacral plates shaded grey. Oral side: 11) PL551Ba,

TL=88.5 mm; 12) PL552, TL=73.4 mm. Aboral side: 13) PL553, TL=55.8 mm. Figs. 14a-e. Variation range of test outline

and lunules shape. All specimens in aboral view, if not otherwise specified; a) PL548 (TL=93 mm), b) PL552 (oral view;

TL=73.4 mm), c) PL550 (TL=79.5 mm), d) PL553Ba (oral view; TL=55.8 mm), e) PL1281 (TL=81.7 mm). Fig. 15. X-ray

photograph (PL549), showing the starring outline of the central cavity. Scale bars equal 1 cm.

264

Paolo Stara & Enrico Borghi

time Sardinia begun to separate from Europe and

the progressive marine ingression transformed it

into an archipelago surrounded by an epicontinental

sea (Gattaceca et ah, 2007). A. nuragica and Am-

phiope sp. 1, from the late Chattian-early Aquitanian

Nurallao formation, inhabited shallow water envi-

ronments of that sea and represent the earliest

record of this genus in Sardinia and one of the

oldest record of Amphiope with transverse lunules

(Stara et al., 2012). The resemblance, based on

comparison of test and lunules features alone, with

the almost coeval population “A” of A. bioculata

described by Philippe (1998) from the Aquitanian

of the Rhone Basin, suggests that at that time

closely related, may be the same, species of Am-

phiope inhabited Sardinian and Provencal shallow

environments.

In the middle Miocene Sardinia was encircled

by a deep sea. During that period Amphiope devel-

oped into new species (Fig. 6): A. lovisatoi, A. mon-

tezemoloi and Amphiope sp. 2 of the Burdigalian-

early Langhian of the Calcari di Mores Formation,

and Amphiope sp. of the Early Serravallian Capo

Frasca Sud Formation (Funedda et al., 2000); Spano

et al., 2002). Fovisato (1914) cited Amphiope also

from a “ sottile banco di calcare breccioso compat-

tissimo, sotto al calcare argilloso ” outcropping at

Monte S. Michele (Cagliari, n. 24 in Fig. 1). This

sediment corresponds to the “Pietra Forte” belong-

ing to the Calcari di Cagliari Formation, dated to

the late Tortonian- early Messinian Cherchi et al.,

1978), thus representing, so far, the most recent pop-

ulation of the genus Amphiope.

A major evolutionary change observed in the

studied sample points to the progressive reduction

and the increasing complexity of the internal sup-

port system. Thus, the dense and strong internal but-

tress and the thick shell of A. nuragica , should be

regarded as primitive characters in Amphiope with

transverse lunules. A decreasing number of plates

in the ambulacral and interambulacral columns and

the progressive migration of the periproct towards

the peristome, from plates 3b/3a in the Aquitanian

A. nuragica , to the distal part of the suture 2a/3b in

the Burdigalian A. lovisatoi, to the proximal part of

2a/3b in the early Fanghian A. montezemoloi and

Amphiope sp. 2, are also observed (Fig. 6). The only

exception to the last two trends is represented by

the “ancient” Amphiope sp.l (Fig. 6), with the pe-

riproct bounded by the first post-basicoronal plates

(2a/2b) and a low number of plates in the oral inte-

rambulacrum 5.

Paleoecological data reported for the Amphiope-

bearing Sardinian localities (Stara et al., 2012) in-

dicate that Amphiope was a deposit feeder, living in

shallow sandy settings, with middle to high water

energy and tropical climate.

CONCLUSIONS

The examined material shows that previously

described criteria used to distinguish between the

fossil species of Amphiope cited in Sardinia are not

sufficiently diagnostic, mainly due to the marked

intra-specific variation of the external test features.

The well preserved specimens available to

study from this region enable to describe the plate

patterns and the internal test support system. Based

on these features and the results of the morphome-

tric analyses, three different species are recognized

in the examined material: A. nuragica, late

Chattian-early Aquitanian of Cuccuru Tuvullao

(Cagliari), A. lovisatoi, late Burdigalian of Chiara-

monti (Sassari), A. montezemoloi, late Burdigalian-

early Fanghian of San Giorgio and Bonnanaro

(Sassari). Two groups of specimens from the late

Chattian-early Aquitanian of C. Tuvullao (layer II)

and the late Burdigalian-early Fanghian of Bancali,

though well differentiated, are assigned to Am-

phiope sp. 1 and Amphiope sp. 2 respectively, and

left in open nomenclature due to the scarcity of the

available material.

The stratigraphical distribution of Amphiope in

Sardinia ranges from the late Chattian-early Aqui-

tanian of Cuccuru Tuvullao, which represents one

of the earliest records of Amphiope with transverse

lunules in the Mediterranean, to the late Tortonian-

early Messinian of Monte San Michele (Cagliari),

the last being the most recent record of this genus.

The main evolutionary trends observed in the

studied sample from Sardinia are the progressive

reduction and the increasing complexity of the in-

ternal support system of the test. Also a decreasing

number of plates in the ambulacral and interambu-

lacral columns and the approaching of the periproct

towards the peristome are observed, though they

need confirmation based on a larger fossil sample.

The results of this study highlight the validity of

the structural characters as taxonomic tools at the

The echinoid genus Amphiope L Agassiz, 1840 (Echinoidea Astriclypeidae) in the Oligo-Miocene of Sardinia (Italy) 265

Specks

M ca n

(mm)

Range

( nun)

S.E

A. lovisatoi

76.0

52-100

1.64

A. montezetmtloi

1 09.0

90-121

6.65

.4. nuragica

92.7

81-100

1.33

Amphiope sp. 1

95.5

90-101

3.17

Amphiope sp. 2

95.9

55.8-137

8 .71

S pe tics

Mean

% Did

Range

% Dan

S.E

A. lovisatoi

105.4

94-113

0.69

A. montezemoloi

1 05.7

100-11Q

230

A. nuragica

1 06.3

101-116

0.77

Amphiope sp. 1

1 09.3

102-114

3.71

Amphiope sp. 2

104.7

102-110

0,87

A. lovisatoi

10.2

6. 8-15.1

0.29

A. rnttnieremofn i

9,8

8.3-12,2

0,99

A. nuragica

14.2

10.5-17.2

0.38

Amphiope sp. 1

20,7

19.8-22.2

0,77

Amphiope sp. 2

1 7 3

8.8-14.6

0.58

A. lovisatoi

1 1.2

8.3-15.1

0.26

A . mo nlezemolo i

14.4

11.6-16.7

132

A . nuragica

8,2

6.0-JI.2

0.37

Amphiope sp. 1

9.5

8.2-11.9

1.22

Amphiope sp. 2

1 t.7

10.8-13.9

0.44

1,2

A . lovisatoi

17.1

1 1-22

0.42

A. montezemoloi

16,9

142-18,3

0.93

A . nuragica

18.0

14,1 -23.5

0.61

Amphiope sp. 1

18.9

17.8-20,8

0.95

Amphiope sp. 2

15.0

11.1-18.2

0.78

A. lovisatoi

5.7

2,8-85

0.21

A- mo niezenwloi

5.0

3. 9-6,6

0,60

A. nuragica

5.3

2 2-8.0

0,35

Amphiope sp. 1

4.6

3, 3-5. 4

0.64

Amphiope sp, 2

4.8

3, 3-6,8

0,49

A. lovisatoi

59.4

5 1 ,9-65.2

9.51

A. montezemoloi

59,3

55.1-62.2

1 .52 j

A. nuragica

57.4

50.0-63.0

0.68

Amphiope sp. 1

53.3

49 5 -57.6

2.35

Atnph f ope sp. 2

59.7

53-66

1.42

A. lovisatoi

24.1

20.8-28.5

0.34

A , montezemoloi

24.7

22.8-26.6

0.98

A. nuragica

25.2

23.0-27.2

0,30

Amphiope sp. 1

? i

27.7

26,7-28.7

0,58

Amphiope sp. 2

24.7

2 1 -9-29.0

0 74

1,6

A. lovisatoi

14,8

11 .8-17.3

0.20

,4. montezemoloi

14,7

132-15,6

0,53

.4, nuragica

14.1

U .1-16.5

0.27

Amphiope sp. 1

15.3

14.4-16,8

0.75

Amphiope sp. 2

15.1

1 2.8 -t 6.3

0.43

A. lovisatoi

22.9

193-26,0

0.30

,4. montezemoloi

2 1.4

20.0-22.7

0,59

A. nuragica

22.4

18.4-26,0

0.47

Atnph tope sp. 1

2 1.9

IK, 8 -24,4

1.64

Amphiope sp, 2

21.7

17.9-24.2

0,80

1,10

A. lovisatoi

! 5.2

I2.5-I9.lt

0,25

A. montezemoloi

13.8

13.4-14.2

0.27

.4. nuragica

14.3

1 1.0-16.2

0.03

Amphiope sp. 1

13.9

11 .9-15.6

1.07

Amphiope sp, 2

14.4

U ,6- 15.8

0,42

LI t

A. lovisatoi

12.0

9.8-14.0

0.91

,4. montezemoloi

! 2.0

1225-14.9

0.63

A. nuragica

9.1

6.3-12.8

0.38

Amphiope sp, 1

13,8

122-15,8

1,06

Amphiope sp. 2

13.0

10.0-15.8

0.69

Table 3. Descriptive statistics of Amphiope from Sardinia.

N = number of specimens, S.E. = standard error.

Amphiope sp.

(M.S. Michele, Cagliari)

Amphiope sp.

(Capo Frasca)

Amphiope sp. 2

Amphiope sp.1

A, nuragica

Figure 6. Scheme of the major evolutionary trends observed

in the studied sample of Amphiope from Sardinia, with the

only exception of Amphiope sp. 1 . Over time the number of

plates in the oral interambulacrum 5 decreases and the

periproct migrates toward the peristome, in relation to post

basicoronal plates number.

266

Paolo Stara & Enrico Borghi

M orphologicul

characters

A. lovisatoi

Cotteau, 1895

A. m ontezemotoi

Lovfeato, 1 91 1

A. nuragica

(Contuse hi Cana,

1955)

Am phi oiTe sp. 1

Amphiope sp.2

Test size (TL)

small

(mean 76 mm)

large

(mean 1 09 mm)

middle sized

( mean 92,7 mm)

middle sized

(mean 95.5 mm)

very large

(> 1 30 mm)

Test height

low

(H=l 0.2% Dap)

very low

(H=9,8% Dap)

middle height

(H=I4.2% Dap)

very high

<11=20.7% Dap)

middle height

(H=12,3% Dap)

Test margin

thin

middle thickness

rather thick

middle thickness

Lunules

size and shape

small

slightly elongate

very targe

almost subcircular

large

narrow elliptical

large

narrow elliptical

large

subcircular-

elliptical

Location of the

periproct

distal half of the

suture 2a/3b

halfway along the

suture 2a/3b

close to the

conjunction of

plates 2 a/3 a/ 3b

bounded by plates

2a/2b

halfway along the

suture 2a/3b

Number of plates in

intcrambulacra 2, 3

and ambulacra II -IV

10-13

14-16

12-14

Number of post-

basicoronals in oral

interambulacrum 5a

Number of post-

basicoronals in oral

ii 1 1 er ambu lac ru m 5 b

Central cavity

subpentagonal

starring

subcircular

starring

Internal structure

light and complex

rather light,

complex

simple and strong

rather strong

Other characteristics

- Shell very thin

- Shell thin

- Plate 2b of oral

imerambulacrum 5

very elongate

- Shell thick

- Petal surface

convex, including

the poriferous areas

- Shell very thick

- Lunules very close

to posterior petals

- Apical disc more

centrally located

than the others

- frontal petal much

longer than the

posteriors

Table 4. Specific characters in the studied species.

specific level in Amphiope and indicate that a re-

view, based on these features, of the earlier de-

scribed species of Amphiope is needed to improve

the poorly resolved taxonomy of this genus and to

bring light into the diffusion of Amphiope in the

Mediterranean and in the eastern Atlantic.

ACKNOWLEDGEMENTS

We are grateful to Andreas Kroh (Natural

History Museum of Vienna), Stefano Dominici

(Museo di Storia Naturale, Sezione di Geologia e

Paleontologia, Universita di Firenze) and Carlo

Corradini (Dipartimento di Scienze Chimiche e

Geologiche, Universita di Cagliari) for valuable ad-

vice and support during the preparation of the man-

uscript. We thank Gianluigi Pillola (Dipartimento

di Scienze Chimiche e Geologiche, Universita di

Cagliari), for allowing one of us (P.S.) to take pho-

tographs of the holotype of A. nuragica, and Andrea

Mancosu (Dipartimento di Scienze Chimiche e

Geologiche, Universita di Cagliari), for information

about the geologic setting of Ardara. We warmly

thank Roberto Rizzo (Parco Geominerario Storico

Ambientale della Sardegna) for information about

the geology of the Sardinian localities, Bertrand

Martin Garin (Centre de Paleontologie, Universite

de Provence, Marseille, France) for information

about the Amphiope specimens housed in that Insti-

tution, and Carlo Cabiddu (Villanovaforru), Sergio

Caschili (Cagliari), Massimo Scanu (Sanluri), Vin-

cenzo Incani (Masullas), Mario Doria (Sassari) and

Giampaolo Troncia (Pau), for the loan of fossil spe-

cimens. Pedro Pereira (Departamento de Ciencias

e Tecnologia - Universidade Aberta de Lisboa, Por-

The echinoid genus Amphiope L Agassiz, 1840 (Echinoidea Astriclypeidae) in the Oligo-Miocene of Sardinia (Italy) 267

tugal) provided us with a first version of the biome-

trical statistical analyses. We are grateful also to Da-

vide Serra, for allowing access to the fossilifer-ous

site of Cuccura Tuvullao, and Mario Lai (3S, La-

boratori Analisi Immagini, Capoterra), for provid-

ing the radiographs utilized in this work.

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Biodiversity Journal, 2014, 5 (2): 269-290

Monograph

Analysis on a sample of Echinodiscus cf. auritus Leske, 1 778

(Echinoidea Clypeasteroida)

Paolo Stara 1 & Maurizio Fois 2

'Centro Studi di Storia Naturale del Mediterraneo - Museo di Storia Naturale Aquilegia, Via Italia 63, Pirri-Cagliari and Geomuseo

Monte Arci, Masullas, Oristano, Sardinia, Italy; e-mail: paolostara@yahoo.it

2 Dipartimento di Scienze della Vita e dell'Ambiente, Universita degli Studi di Cagliari, Via Ing. Tommaso Fiorelli 1, Cagliari, Italy;

e-mail: prionace85@hotmail.it

^Corresponding author

ABSTRACT In order to ascertain the extent of the natural intraspecific variability of living and fossil

echinoids belonging to the family Astriclypeidae Stefanini, 1912, morphometric and struc-

tural aspects were examined in a number of specimens of extant Echinodiscus cf. auritus

Leske, 1778, from Madagascar and Philippines. The data obtained will be compared, in a

following work, with those of other echinoids belonging to the same family. The analysis of

the results indicates, for the sample studied, a great variability in the length of the posterior

ambulacral notches, in the petaloid length and in the position of the periproct respect to the

posterior margin, while the study of the complete scheme of the plates has clarified the sta-

bility and constancy of some parts of this scheme and the variability of other. On the basis

of these observations, it has been claimed that the variability of these measures is not so

extensive as to affect or determine specific distinctions, if used without careful analysis of

the plating pattern in particular in the interambulacrum 5 and in the ambulacra I and II. The

results of these analyses, finally, suggests that these echinoids belong to a different genus,

than Echinodiscus Leske, 1778.

KEY WORDS Astriclypeidae; Echinodiscus cf. auritus', morphometric variability; Recent.

Received 25.06.2013; accepted 30.05.2014; printed 30.06.2014

Paolo Stara (ed.). Studies on some astriclypeids (Echinoidea Clypeasteroida), pp. 225-358

INTRODUCTION

The variability in shape of some parts of the test

in echinoids belonging to family Astriclypeidae Ste-

fanini, 1912, has been a problem discussed for a long

time (Stara & Fois D., 2014). Considering the wide

variability in the extinct genus Amphiope L. Agassiz,

1840, Philippe (1998), in the absence of comparative

studies on the intraspecific variability in living echi-

noids belonging to the same family, poses in syn-

onymy many Amphiope species present in the

Miocene levels of the Rhone Basin (Southern

France). In fact, given the large variability in size and

shape of the lunules of these echinoids, he concluded

that most nominal species previously established

should be in synonymy with the type species: Am-

phiope bioculata des Moulins, 1837. The same dif-

ficulties are also experienced by Stara & Borghi

(2014) during the study of Sardinian’s Amphiope

species, some put in synonymy by Philippe (1998)

and, later, also by other authors. In this case, howe-

ver, the study of the species from Sardinia was ad-

dressed by analyzing both the variability of lunules,

both the internal structure and the scheme of the

270

Paolo Stara & Maurizio Fois

plates that compose the test, as already done by

other authors for this family of echinoids (Durham,

1955; Kroh, 2005; Pereira, 2010).

Therefore, the goal of this work is to verify the

extent of the morphometric variability in some

echinoids belonging to the genus Echinodiscus, in

order to compare it with that observed by Stara &

Borghi (this volume) on a sample of at least a hun-

dred of specimens of different species of Amphiope

collected in a number of Oligo-Miocene Sardinian

outcrops.

MATERIAL AND METHODS

The samples examined are housed in the collec-

tions of the Museo di Storia Naturale Aquilegia of

Cagliari (acronym MAC). Being irregular echinoids

in bilateral symmetry, with the symmetry plane pass-

ing along the line stoma - procto, we realize that it

was the distance measured along the test length

(TL) could provide the major reference point, ac-

cording by Kier (1972) and Stara & Borghi (2014).

Therefore, specimens were ordered on the basis of

the TL, as done previously by Lohavanijaya &

Swan (1965), Alexander & Ghiold (1980) and

Stelmle (1990).

We detected the structure of the test, showing the

relative platings of the plates, as was done by Durham

(1955), but including both faces of specimens. The

internal structure was studied by sectioning the test,

and by X-ray; 3 specimens from Mangili, 2 from

Philippines were used for this purpose.

Because of the similarities between the two gen-

era object of the parallel study, to reach homo-

geneity we use on Echinodiscus the same set of

measurements used to collect data on Amphiope by

Stara & Borghi (this volume) with the addition

of some further measures, such as the attached

drawings.

Morphological abbreviations (Fig. 1) B angle be-

tween major axis of the two lunules; TL = Test

Length; TW = Test Width; TH = Test Height; WA=

ambulacral and interambulacral width at ambitus;

L1-L2 = lunule length and width, respectively; L3 =

distance between petal tip and corresponding lunule,

Figure 1. Biometric parameters measured in the studied samples.

Analysis on a sample of Echinodiscus cf. auritus Leske, 1778 (Echinoidea Clypeasteroida)

271

L4 = distance apical system-posterior margin, L5-L6

= length and width of the frontal petal, respectively;

L7-L8 = length and width of the anterior paired petal,

respectively; L9-L10 = length and width of the

posterior petal, respectively; LI 1 = distance between

periproct and the posterior border of the test; LI 2 =

distance between periproct and peristome, LI 3 =

front-rear diameter of the basicoronal circlet. PL =

Petalodium Length; o pc = periproct diameter; o ps

= peristome diameter; E = summation. To reduce the

lunules shape and dimension into a numeric value,

we introduced a Shape Index (SI) corresponding to

the ratio L2/L1 and a Width (wideness) Index (WI)

corresponding to the formula (LI + L2 ) / 2. In the

chapters of the analysis/descriptions of samples/spec-

imens, the interambulacrum 5 was abbreviated as

inter 5.

Measurements were taken with caliper and/or

metallic decimeter (with division to the millimeter)

measures of TL and thickness of the anterior and

posterior margins were taken in millimeter; all other

linear measurements were reported as percentage

TL; while the angular divergence between the axis

of the notches, were detected in degrees, using a

computer graphics program. The use of the percent-

ages was adopted to decrease the influence of the

variations due to the growth and to the difference

in size. Finally, the width of the different ambu-

lacral and interambulacral areas at margin (WA),

was performed measuring the distance between the

interradial sutures of each area.

Plates were numbered according to Loven

(1872) system and interambulacral areas were

shaded in grey. The systematic classification used,

follows Kroh & Smith (2010).

Note

Since it is veiy rare that minute spines, pedicel-

laria, parts of the lantern or even the soft parts of

the intestines conserve, the information which we

consider useful for the classification of fossils will

be searched in the structure of the test and, where

possible, in the internal support. Moreover, even

many tests of living clypeasteroids, because of the

fast taphonomic processes they are subject, they

come to us devoid of spines. In order to observe and

compare the internal structures of some specimens

examined, we also used specimens dissections and

numerous x-rays in supero-inferior position.

Critical value of the Spearman rank correlation

ro (a, n), with a = 0.05 and n = number of samples,

are extract from the table of critical values obtained

through the software SuppDist (Wheeler, 2005) im-

plemented in R and based on the method of Kendall

& Smith (1939).

OBSERVATIONS ON SAMPLES

Sample 1

Echinodiscus cf. auritus Leske, 1778

Recent, Philippines (Plate 1 Figs. 1-8; Plate 2

Figs. 1-9; Figs. 2 a-d; Tables 1, 2).

Examined material. 5 specimens from Talibon

in the north of the Bohol island, dredged at 50 m

depth, code MAC.IVM 206 - 210, TL. 131 - 154

mm; 5 specimens from Oslob, Cebu Island,

MAC.IVM 211-215, TL 125 - 173 mm; 2 speci-

mens of generic provenance "Philippines"

MAC.IVM81, 133; TL 150 and 152 mm.

General. In detail, the line of the ambitus is

almost always polygonal, with almost rectilinear

sections in correspondence of the different ambu-

lacral and interambulacral areas. The line of the

rear edge, with the two ambulacral notches, forms

a "dovetail", which is always very irregular and

sometimes is asymmetrical (Plate 1 Figs. 1, 2). In

some cases, the "tail" protrudes from the ideal line

of the ambitus, in other cases it does not arrive

there because of its shortness; sometimes it is not

straight and draws an arch that seems to continue

the roundness of the rest of the ambitus. For this

reason, it often happens that the two sides of each

notch show a different length. The irregularities

that occur along the margins, or even inside the

notches, sometimes seem to be caused by attacks

of predators, or by deformities present at birth, or

by growth malformations. The test is very de-

pressed (mean = 8% TL); the highest part of the

test is located just before the apical disc which is

eccentric forward, in a range that should be 54-

58% TL. From a side-view, the profile is not uni-

formly tapered but, starting from the posterior

margin, it goes up with a slight slope along the

length of the notches, then it increases rapidly until

just beyond the peak, from where it descends

rapidly towards the anterior margin. The ambital

272

Paolo Stara & Maurizio Fois

margin is thin and sharp rear (0.8 to 1.5 mm), and

it becomes thicker more and more, as it gets closer

to the front where the thickness varies from 2.5 to

3.5 mm. The adoral surface is flat or slightly plano-

concave, with the inner point near the peristome;

the last one is opened in a central position.

Test structure. In summary, the structure can

be divided into two main parts: the central cavity,

which houses most of the viscera, and the periph-

eral system that mechanically "supports" the whole

test (Plate 2 Fig. 1). It is noted that the distribution

of the interior supports (pillars, buttresses, “trabec-

ulae”) is not linked to the plates distribution, so

they do not constitute the inner extension of their

structure.

Visceral hollow. This central cavity contains

both the gut and the main vital parts (inner part of

the respiratory organs, reproductive organs, etc.). In

plan view, the shape is roughly rounded-polygonal,

with the base line lying between the two rear am-

bulacra and located at right angled of the inter. 5.

From the rear wall to the front, the length of the hol-

low corresponds to an average of ~ 43% TL. From

this cavity opens some pocket containing several

vital organs, and two elongated cavities: one of

these opens along the ambulacmm II and contains

the caecum, while the other one, longer, contains

the terminal intestines along the inter. 5 and leads

into the periproct. L. Agassiz (1838-41) described

in detail the path of the intestine compared to vis-

ceral hollow. From the aboral view the visceral cav-

ity of these echinoids coincide with the width of the

petalodium, whose PL ~ 43% TL (Table 2). From

the adoral side, instead, the visceral hollow is

roughly contained into the limits of the first distal

ambulacral post-basicoronal plates that are particu-

larly extended, as you can see from the relative plat-

ing. The visceral hollow ceiling is supported by a

system of 5 trabeculae that, from the lateral sup-

ports of the cavity (pillars and cellular tissue) run

along the interambulacra, between the petals, until

they join the reinforcement ring that supports the

apical disc and the components connected to it

(Plate 2 Fig. 2). The poriferous areas are flat, or in

some cases slightly sunken. The Aristotle's lantern

rests around the peristome, which opens at the cen-

ter of the floor and has a sub-pentagonal outline.

The auricles which support the lantern, flanked dis-

tally from navicular pits, stand on the basicoronal

interambulacral plates (Plate 2 Figs. 6d, 7a). The

plank is formed mainly by post-basicoronal ambu-

lacral plates; these ones are thin, measuring less

than 1 mm thick, but they are reinforced by a "net-

work“ mesh structure more and more thick and

dense as it gets closer to the side walls of the vis-

ceral hollow. Looking down the hollows, it is noted

that the lantern covers most of the floor because it's

relatively large in this population of echinoids. The

lantern is "balanced" on the auricles, and it is

moved by a system of ligaments attached to the per-

radial ambulacral sutures.

Apical disc. The apical disc is star- shaped with

the points turned towards the interambulacra and is

formed by a small madreporite (mean width = 5%

TL), in which the 4 genital pores open. The genital

and the ocular pores can be observed only on naked

specimens (Plate 2 Fig. 3).

Peripheral ballast system. This system is

formed by a sandwich composed by the oral and

aboral parts of the test, each with its own combi-

nation of plates, and from an internal "cushion"

formed by pillars or by buttresses of tissue with a

cellular structure, crossed by more or less wide

cavities. This "cushion" (with a probable coelomic

origin) is formed by emi-structures that extend

from the plates and are joined together in a plane

more or less median (Plate 2 Figs. 4a, 6a). You note

that the structure formed by radial ribs and micro-

canal that constitutes each plate, and which is evi-

dent after erosion, is the evidence of the direction

of the growth of the same, which follows the dis-

tribution of the microchannels of the aquifer

system and that isn't related to the direction of

growth of the underlying internal structures. This

complicated "sandwich" system constituted from

plates, buttresses and trabeculae, reaches the am-

bitus where, in general, the last plate is "refolded"

on itself in a peculiar way, forming precisely the

margin. Counting the plates, in fact, you will

sometimes consider that the last plate of a face is

also the first one of the other face; only in some

cases the two plates that delimit the upper and

lower face appear to join up to the margin.

To illustrate this peripheral system of reinforce-

ment, we also used simple radiological techniques,

which show well the pillars, buttresses support and

internal cavities distribution, and that is becoming

a potential diagnostic tool (Plate 2 Fig. 1).

Analysis on a sample of Echinodiscus cf. auritus Leske, 1778 (Echinoidea Clypeasteroida)

273

Ambulacral areas. Along these areas opens

the petals and the upper part of rear notches. The

petals are closed, long twice their width and are

sub-equal to each other (mean L5 = 23% TL, L7 =

19% and L9 = 20.5%). In 7 out of 12 specimens of

this group, only one pair of post-basicoronal ambu-

lacral plates occlude those of inter. 5; in the remain-

ing specimens (MAC.IVM206; 207, 208, 209, 211,

212; 81) the inter. 5 is occluded by two pairs of

post-basicoronal plates (Figs. 2b, c). The doubling

of the post-basicoronal ambulacral plates in the am-

bulacra I and V, sometimes leads to a shifting of the

position of the periproct towards the rear margin.

In the ambulacra I and V, the width at the ambitus

WA, is on average the 23% TL and the plates per

column are 14-15; in the ambulacra II, III, IV, the

WA are on average 37% TL but the plates are 12-

13 in the ambulacra II and IV, while they are 13-14

on the ambulacrum III.

Notches. The notches extend along the perra-

dial sutures of the rear ambulacrum and they are

opened in the posterior margin. Differences in

shape and asymmetries observed in these notches,

are common and therefore we have considered

them as normal. The length of the notches (LI) in

this sample varies from 18 to 27% TL (mean

23%). Because of the constant presence of defor-

mations and/or malformations along the notches,

the detection of L2 is not significant from a statis-

tical standpoint. The notches are surrounded by 4

and 4 plates per column on the oral face and by 4-

5 plates per column on the aboral one, approach-

ing, therefore, to a 1/1 ratio (Fig. 2a-c). Between

the petal tip and the corresponding notch there are

3-4 couples of plates, while, on the oral face the

notches opens between the plates 4a/5b or 5a/4a.

The B angle is small and on average is almost 57°

(Table 2).

Interambulacral areas. The WA in interam-

bulacra 1 and 4, is 40% TL and the plates per co-

lumn are 14-15; in the interambulacra 2, 3, 5,

however, the WA is on average 33% TL, while the

plates per column are 15-16. The WA remains rela-

tively constant, with the areas of the interambulacra

1 and 4 always larger than the other. The number of

post basicoronal plates in inter. 5 is always 3 in

column “a” and 3-4 in column “b”.

Periproct. The periproct is round (range 1.5-2,

average 1.8% TL) and far from the rear margin (LI 1

= 14.5 -s- 24.5% TL, average = 20%). Except a clearly

abnormal specimen, the periproct always opens

along the suture between the post-basicoronal plates

2b and 2a, that are partially paired (Figs. 2b, c).

Food grooves. The adoral face is crossed by

very branched food grooves. Every single column

in the ambulacral plates is traversed in the center

by a main groove, which are joined to other numer-

ous secondary thinner and less deep grooves (Fig.

2d). Other large grooves are joined to the main one

at various levels: on the ambulacra I, II, IV and V

these converge to the greater one in the middle of

the corresponding ambulacral column; on the III,

however, in several cases the secondary grooves

converge closer to the stoma, within the post basi-

coronal area. Finally, the two main grooves of each

column are joined exactly at the height of the tip of

the basicoronal ambulacral plates, that then lead to

the stoma. In the rear ambulacra they run along both

sides of the notches. The ambulacral plates crossed

by food grooves have a greater deflection near of

the stoma.

Peristome and Aristotle’s lantern. The peri-

stome varies from rounded to sub-pentagonal

shape and is relatively small (mean = 3% TL). The

basicoronal circlet is small (mean LI 3 = 11% TL).

The basicoronal interambulacral plates are quite

regular and always are disjointed from the corre-

sponding post-basicoronal ones. The Aristotle's

lantern is very flat and especially developed in

width, and it is formed by five jaws delta- wing

shaped, whose bisector is a groove within where

the teeth run horizontally. The teeth are knife

shaped, with the sharp end facing toward the stoma

(Plate 2, Figs. 5, 8); Always along the bisector near

the stoma, a small fork and the corresponding

navicular dimples that fit in the underlying auricle

are located in each jaw. The jaws are held together

by ligaments attached to perradial ambulacral

sutures. We verified if there was a direct relation-

ship between the size of the lantern and the size of

its basicoronal circlet. In the specimen of Bohol,

MAC.IVM209 (TL = 152 mm) the size of the

basicoronal circlet (LI 3) is 11.5% TL; the height

of the pentagon formed by the Aristotle's lantern,

as in the plating, measures ~ 40 mm, equal to

26.3% TL; the distance between the support of the

inter. 5 and the edge of the stoma is of ~ 6.5 mm

and the diameter of the stoma exceeds 3.5 mm.

274

Paolo Stara & Maurizio Fois

Fig. 2: “ Echinodiscus ” cf. auritus, Recent, Philippines: a, b) respectively, aboral and oral plate structure of MAC.IVM

210; c) oral plate structure of MAC.IVM 211: d: food grooves.

Specimen TL TW TH L) L2 L3 L4 L5 L6 L7 LK L9 L10 Ml L12 L13 o?c oSro

IVM215

! 73

103

7.5

21.5

13.5

1 1.5

9.5

21.5

IVM208

131

104

7.4

16.5

9.5

1.5

2.H

1VM210

141

104

7.4

1.5

17.5

8.5

22,5

28,5

10.5

1.8

2.5

I VM 206

148

103.5

7.4

11.5

18.5

24.5

9.5

1,5

2.5

IVM209

! 52

106.5

9.2

22.5

2.5

15.5

24.5

18.5

1 1.5

2.8

IVMQ81

152

102

7.5

12.5

22.5

14.5

n.5

1.6

1 VM 213

152

1 06

8.5

12.5

56.5

24.5

12.5

1 1,5

21.5

10.5

30.5

12.5

3.5

IVM207

154

101

2.5

22.5

20.5

10.5

17.5

11.5

1.8

2,5

[VM214

156

108

13.5

12.5

31.5

11.5

3.2

I V M 2 1 2

157

103

22.5

3.5

13.5

12.5

20.5

31,5

3.5

IVM21 1

1 59

105

4.5 ,

11.5

9.5

16.5

il.5

1,6

3.5

101

7,4

1.5

10,0

9.0

16.5

9.0

17.0

8.0

16.5

28.0

9.5

1.5

2.5

Range

108

9.2

4.5

17.0

13.5

22.5

12.5

23,0

12.5

24.5

35.0

12.5

3.5

Mean

104

2.5

10.5

20.5

31,5

1.8

Variance

4,5

3.5

4.5

0,5

Range- V

4.7

120

31.5

Table 1. Morphometric data of “Echinodiscus” cf. auritus (Philippines, Recent). TL in mm, other data in % TL.

Analysis on a sample of Echinodiscus cf. auritus Leske, 1778 (Echinoidea Clypeasteroida)

275

Plate 1: Echinodiscus cf. auritus from Philippines (Recent): external features. Figures 1, 2, 3. Aboral, adoral and lateral

views of MAC.IVM 212. Figure 4. Close up view of the pointed spines on the inner slits surface. Figure 5. Primary and se-

condary aboral spines. Figure 6. Claviform primary and secondary aboral spines. Figure 7. Tuberculation and food grooves

of the peristomial area. Figure 8. Periproct, tuberculation and food grooves on posterior areas.

276

Paolo Stara & Maurizio Fois

Plate 2. Echinodiscus cf. auritus from Philippines (Recent): internal features. Figure 1 . Radiograph of specimen MAC.IVM

215 (Dap =173 mm): a) central hollow; b) teeth on wing of disarticulated lantern; c) caecum cavity; d) terminal intestine

cavity. Figure 2. Internal view of ceiling of central hollow: a) interporiferous areas; b) poriferous areas; c) apical disk rein-

forcement. Figure 3. Corresponding external view of apical disc: a) genital pores; b) madreporite. Figure 4. Fragment of the

test with: a) separation surface of internal structures; b) peripheral pillar and buttresses; c) walls delimiting the central

hollow. Figure 5. Aristotle's lantern of MAC.IVM 209, upper view: a) jaw; b) teeth. Figure 6. MAC.IVM 209: antero-po-

sterior section (from the left to the right); a) separation surface of internal structures; b) peripheral pillar and buttresses; c)

walls delimiting the central hollow; d) floor of the central hollow; e) lantern supports. Figure 7. a) lantern support; b) rib in

a network of floor reinforcement; c) peristoma. Figure 8. MAC.IVM 209: Aristotle’s lantern in position.

Analysis on a sample of Echinodiscus cf. auritus Leske, 1778 (Echinoidea Clypeasteroida)

277

In the specimens MAC.IVM81, with the same

length as the previous one, L13 is ~ 12% TL; the

height of the pentagon of Aristotle's lantern mea-

sures 38 mm, equal to 25 % TL; the distance be-

tween the support of the interambulacmm 5 and the

stoma is 7 mm, equal to 4.6% TL, while the di-

ameter of the stoma is 3 mm. In the specimen

MAC.IVM206 (TL =148 mm), the lantern mea-

sures 30 mm, which corresponds to ~ 20% of the

TL. So the size of Aristotle's lantern varies, in the

specimens tested, around 20-26% TL and does not

seem related to the width of the basicoronal circlet

(LI 3). The basicoronal interambulacral plates are

flat and studded with large tubercles crenulated and

perforated. The spines that surround the stoma,

post-mortem, are inclined to cover it completely.

Tuberculation and spines. In the oral face the

tuberculation is dense, consisting of primary tuber-

cles medium sized, poorly differentiated and ex-

tended over the face, and secondary tubercles that

form a granulation background, scarcely distinguish-

able. The tubercles are larger around the periproct

and the smaller sized ones are found particularly

along the main food groves (Plate 1 Figs. 7, 8). In

the aboral face the tubercolation is undifferentiated,

thick and petite, evenly distributed over the entire

surface.

The spines are well differentiated according to

the areas in which they are located, the larger are

surrounded by numerous small ones, that are also

much thinner. Both types are more or less cylin-

drical, slightly thinner distally, and traversed by

numerous thin longitudinal ribs, and both end

with a rounded tip (Plate 1 Figs. 4-6). Spines of

medium and large size are found around the

stoma; other ones with a medium-length cover

ambulacral center areas, including the entire

length of the notches, even on the inner surfaces;

other spines, much shorter and thinner and with a

hinted club-shaped, however, cover the areas

around the food grooves. Along the ambitus and

along the notches there are often curved and sharp

spines. Also, together spines of medium length,

cylindrical and with pointed termination, there are

stubby, short and clearly club-shaped spines. Even

all the aboral face is covered from thickly spines,

generally smaller than those already described.

Apart the surfaces of the petals, where the spines

are cylindrical and pointed, and near of the notches,

there are transitions forms of spines between

Specimen

Apical

disc

[VM206

148

4.5

IVM207

154

4.5

IVM2GS

131

IVM209

152

5.5

I VM21 0

141

1VM2M

159

IVM212

157

IVM213

152

IVM2I4

156

i VM215

173

4.5

Mean jLjyg-gJ

4.6

43.4

33,4

iLiiicc jSpHnj

4-5.5

41-48

30-38

54-61

Table 2. Morphometric data of apical disc, PL, WA and B,

in “ Echinodiscus ” cf. auritus (Philippines, Recent). TL in

mm, B in degree, other data in % TL.

those of the adoral face and those of the aboral

one, while all the other spines have club-shaped

with the ends decidedly thickest of the rest of the

stem (Plate 1 Fig. 6).

Sample 2

Echinodiscus cf. auritus Leske, 1778

Recent, Madagascar (Plate 3 Figs. 1-7; Plate 4

Figs. 1-7; Figs. 3a-c; Tables 3-7)

Examined material. 32 specimens from Man-

gili, Province of Tulear, caught in back-barrier

lagoon, ~ 5 to 8 m deep, MAC.IVM82 - 113, TL =

74 -s- 140 mm.

General. The ambital line is almost polygo-

nal; the posterior margin line, sited between the

two notches (the tail), is always irregular and often

very asymmetric. Although in smaller individuals

seem to prevail lines with a rounded shape, the

larger individuals present clearly truncate lines.

278

Paolo Stara & Maurizio Fois

As in the specimens coming from Philippines,

in some cases the "tail" comes out from the ideal

rounded line in the ambitus (Plate 3 Figs. 1, 2).

The test is medium - large (max TL =140 mm),

with a consistent variability of the relationship

TL/TW, and very depressed (mean TH = 8% TL).

The highest part of the test is located at the apical

disc that is forward eccentric (mean L4 = 55%

TL). The side profile is slightly and almost uni-

formly sub-conical. Starting from the posterior

margin, it rises almost uniformly and then de-

creases as the inclination, finding the maximum in

an area that has the apical disc in the center. This

trend is repeated to the anterior margin, just a little

steeper (Plate 3 Fig. 3). Only on some larger spec-

imens the line of the rear profile rises more gently

towards the top and remained low over the "tail"

of the echinoid. On the specimen MAC.IVM111

(TL =140 mm), the ambitus is thin and sharp pos-

teriorly (0.8 mm), but it thickens more and more

as it gets closer to the front, where the margin is

often ~ 2.5 mm and rounded. The adoral surface

is flat or slightly plano-concave, with the inner

point near the peristome; the last one opens in a

central position (L11+L12 mean = 50% TL). The

periproct is far from the posterior margin (Lll =

17 to 23.5 % TL). Especially in larger specimens

there are showy deformations, due to both growth

malformations, or to damage from predations.

Many of these damages from predation are self-

repaired in life, since they are covered with new

tubercles.

Test structure. The structure of the test of

these echinoids is not very different from that of

the specimens coming from the Philippines; this

structure can then be divided into two main parts:

the central one, which contains visceral cavity,

and the peripheral one, which contains the pillars

and buttresses made of cellular structure (Plate 4

Figs. 1, 4).

Visceral hollow. On the specimen

MAC.IVM111, this cavity has a rounded to sub-

pentagonal plan, with the base placed between the

rear ambulacra and right-angled to the interambu-

lacrum 5; the distance between the extreme rear and

the front of the hollow is approximately 38%

TL. In general, from an upper view the echinoid

present the visceral hollow circumscribed into the

petalodium (mean PL = 40), while the floor ends

before the distal limit of the first post-basicoronal

ambulacral plates. The ceiling structure of the

visceral hollow is comparable to that of the speci-

mens coming from Philippines; the surface of the

interporiferous areas are raised, while that the porif-

erous ones are flat or very slightly concave. Unlike

the specimens coming from the Philippines, the

relationship between basicoronal and post-basi-

coronal plates is very changeable even in the same

specimen itself. This population shows a high vari-

ability in the development of the basicoronal inter-

ambulacral and the post-basicoronal ambulacral,

while the development of the other plates is normal

(Plate 5 Figs. 3-5).

Apical disc, the small apical disc (about 5.5%

TL) is formed by the star shaped madreporite, and

presents 4 genital and 5 ocular pores (Plate 3 Fig.

4). All the specimens are adult and present all the 4

genital pores open.

Peripheral ballast system. Also for the

structure of the visceral hollow, there are no fun-

damental differences compared to those of the spec-

imens coming from Philippines. The ribs, the

pillars and the reinforcing buttresses seem more

delicate than the ones of the Philippine specimens

(Plate 4 Figs. 4-5).

Ambulacral areas. The petals are sub-equal,

long about twice of their width and always closed,

but the overall shape varies also conspicuously

(mean L5 = 21%; L7 = 19%; L9 = 18.5% TL).

There are interporiferous areas wide 1.2 to 2 times

than the poriferous ones.

Given the numerical strength and the size range

of this group of echinoids, we detected changes in

petals size in relation to the size of the test (see

Table 4). In all the specimens of this group, only a

pair of post-basicoronal ambulacral plates occlude

those of the interambulacrum 5 in the oral face.

Unlike the Philippine sample, in this one the rela-

tionship between the basicoronal and post-basi-

coronal interambulacral plates is highly variable,

with also several joint plates in the same specimen

(table 5). In the ambulacra I and V, the WA is on

average the 23% of TL and the plates per column

are 14-15; in the ambulacra II, III and IV, WA is

on average the 37% TL, while the plates per

column are 12-13 in the II and IV, and 13-14 in the

ambulacrum III.

Analysis on a sample of Echinodiscus cf. auritus Leske, 1778 (Echinoidea Clypeasteroida)

279

Rear notches. There is a high frequency of

malformations and deviations in the population, due

either of delayed or unfinished processes of disjunc-

tion, occurred perhaps in the course of the individ-

ual development. In specimens MAC.IVM108, 111

and 112, the two long edges of the notches have not

completed the disjunction, in other cases the

notches are very malformed. Often they are also

very asymmetric, so the two notches are quite dif-

ferent in the same individual. In this group of echi-

noids the disjunction of lunules interests 4-5

couples of plates on the oral face and 4-5 on the

aboral ones, with a 1/1 ratio. Between the petal tips

and the beginning of the notches there are 4 or 5

plates per column, in the oral face the notches opens

between post-basicoronal plates 3a/3b. B range from

49° to 62° for an average of 55° (Table 6).

Interambulacral areas. The WA of the inter-

ambulacra 2, 3, 5 measures on average the 32% TL;

on the interambulacra 1 , 4, however, measures on

average the 40% TL. The number of plates in the

ambulacra 1 and 4 are 14-16, such as in the inter-

ambulacra 2 and 3, while on the inter. 5 they are 13-

14. In the oral face the inter. 5 is always occluded

and it has always 4 and 3 post- basicoronal plates

per column.

Periproct. The periproct is small, because in

young specimens it is the 2.6% while is about the

2% TL in the larger ones. Lll is less variable than

in the sample coming from the Philippines, but the

average of Lll is always ~ 21% TL. In almost all

the specimens (31 of 33) the periproct opens

between the plates 2/2b in the inter. 5; only in two

specimens it is at the end of said suture, in position

2a/2b/3a (Table 5).

Food grooves. The food grooves in this sample

do not differ much from those coming from the

Philippines, only in some specimens, the two grooves

of each ambulacral column join together a little be-

fore the tip of the basicoronal plates (Fig. 3c).

Peristoma, Aristotle’s lantern and basico-

ronal circlet. In this group of echinoids the peri-

stoma is circular, with a diameter ranging from 4

mm on smaller samples, up to 3 mm of the larger

ones (overall average 3.5%TL). The basicoronal

circlet is small (mean L13 = 11.5% TL). The basi-

coronal interambulacral plates are completely irreg-

ular, with considerable variations in length and

shape, which can diversify from triangular to

lanceolate and very elongated. Often, even in the

same individual, there are some post-basicoronal

plates in contact and other disjointed; also, the

second interambulacral plates, disjointed or not, can

be found both in meridoplacous or amphiplacous

condition (see Table 5). The Aristotle's lantern sup-

ports stand, on the floor of the visceral hollow of

the basicoronal interambulacral plates (Plate 4 Figs.

4, 6) ; given that the length of the plates varies

greatly, we observed that the relative position of the

supports is independent. In the specimen MAC.

IVM112 (TL =140 mm) the pentagon formed by

the lantern measures 15.3% TL, absolutely far from

the 20.2% of the TL detected in the sample

MAC.IVM206 (TL =148 mm) coming from the

Philippines. In another sample of TL = 1 16 mm, the

lantern measures about 15% TL.

From what has been observed, it is clear that the

abnormal length of the basicoronal interambulacral

plates does not seem to lead to consequent struc-

tural changes.

Tuberculation and spines. On the oral face the

tuberculation is dense, consisting of medium sized

tubercles, poorly differentiated and extended over

the entire surface. The tubercles are larger around

the periproct (Plate 3, Figs. 5-7) and the smaller

ones are found particularly along the main food

grooves. On the aboral face the tubercolation is

undifferentiated, thick and petite, evenly distributed

over the entire surface.

The echinoids of this sample are devoid of spi-

nes, except in some restricted areas, such as the

edges of the stoma and the inner surfaces of the

notches. The spines that we have observed are not

dissimilar to those observed in the Philippine

sample.

DISCUSSION

One of the problems that has arisen in the paral-

lel work done by Stara & Borghi (2014), was the

evidence of a marked decrease in the number of

plates and a reduction in the internal structure, during

the geological time.

It is shown, therefore, the need to count all the

plates of echinoids we examined (Table 7) subject-

ing the obtained data to a statistical examination.

280

Paolo Stara & Maurizio Fois

Plate 3. Echinodiscus cf. auritus from Mangili, Madagascar (recent): external features. Figures 1, 2, 3. Aboral, oral and

lateral views of MAC.IVM 81. Figure 4. View of the apical disc with madreporite (a), genital (b) and ocular (c) pores.

Figure 5. Stoma surrounded by medium sized tubercles and small spines. Figure 6. Large (a) and small (b) primary tubercles;

scarcely differentiated tuberculation in MAC.IVM8 1 .

Analysis on a sample of Echinodiscus cf. auritus Leske, 1778 (Echinoidea Clypeasteroida)

281

Plate 4. Echinodiscus cf. auritus from Mangili, Madagascar (Recent), internal features. Figure 1. Radiograph of MAC.IVM

109 (Dap = 125 mm). Figure 2. Aristotle's lantern, oral view. Figure 3. Close up of the support system and teeth. Figure 4.

Fragment and antero-posterior test section of MAC.IVM 103 with: a) peripheral pillar and buttresses; b) walls delimiting

the central hollow. Figure 5. Close-up of supports into the last aboral and adoral plates. Figure 6. Floor of the central hollow

with ribs in a network of reinforcement. Figure 7. Small Aristotle’s lantern in position.

282

Paolo Stara & Maurizio Fois

Plate 5. Echinodiscus cf. auritus. Instability of the disjunction in basicoronal interambulacral plates in“ Echinodiscus” speci-

mens. Figures 1, 2. E. bisperforatus (South Africa, Recent): 1) interambulacra 2, 3 continuous; 1, 4 and 5 disjunct; 2) all

plates disjunct. Figures 3, 4, 5. E. cf. auritus (Mangili, Madagascar, Recent), MAC.IVM110: interambulacra 1, 2, 3, 4

continuous; 5 disjunct; MAC.IVM115 interambulacra 1, 2, 3, 5 disjunct, 4 continuous; MAC.IVM84 interambulacra 1, 2

continuous; 3, 4, 5 disjunct. Figure 6. E. cf. auritus (Philippines, Recent), MAC.IVM206: interambulacra 1, 2, 3, 4 disjunct.

Analysis on a sample of Echinodiscus cf. auritus Leske, 1778 (Echinoidea Clypeasteroida)

283

Plate 6. Comparison of Aristotle’s lantern of “Echinodiscus” cf auritus in specimens from a) Mangili; b) Philippines; c) jaws.

This has documented, as expected, given the

preliminary data (Table 3) that in our sample of

adult individuals, while doubling the size, the num-

ber of plates per column in some interambulacra

and ambulacra does not vary significantly (see

Figure 4 and conclusions).

This confirms the fact that the reduction of plate

numbers observed in geologically younger speci-

mens is credible. Moreover, even Durham (1955)

cites a case like this, with a decrease of plates

observed in Echinocardium from subsequent geo-

logical periods.

With regard to the statistical analysis, however,

the samples analyzed show a distribution of length

(TL) that goes from a minimum of 54 mm to a

maximum of 15 1 .5 mm (mean 104.5); standard de-

viation 20.7; range 77.5; median 103.3 (see Fig. 4,

Length Frequency Distribution).

For each sample were also collected the counts

related to the number of plates present in the various

ambulacral and interambulacral column. Analyzed

here are those related to oral and aboral face of the

interambulacrum 5 (defined respectively 5 - oral

and 5 - aboral) and those of the single-sided aboral

(3-aboral) of the III ambulacrum. To assess the in-

dependence of these measures in relation to TL, we

test by "Spearman ps" the null hypothesis HO: ps =

0 ("no correlation") for each of the following pairs

of variables, considered separately: TL - 5 - oral;

TL - 5 - aboral, TL - III aboral. The level of signif-

icance chosen is a <0.05.

Scatter plots obtained for each pair of variables

do not seem to show any linear relation between the

variables represented. This seems also suggested by

the test of Spearman (ps), which does not allow us

to reject the null hypothesis ("no correlation") with

a 95% confidence level, as reported in Table 8.

We have not considered necessary, because evi-

dent, to verify statistically the variation in the size

of the back notches in this sample, which is rela-

tively low. In fact, in a lot of sizes ranging from 21

to 28% TL (mean 24%), we observed a variation of

29% on average value).

Another goal of this work was to try to under-

stand the real extent of variability in LI, PL, Lll,

WAandB.

The variability of LI is significant and similar

in both samples (20-28%, mean 23% TL, against

LI 21-28%, mean 24% TL), but does not seem to

affect the possibility of specific distinction.

284

Paolo Stara & Maurizio Fois

Specimen

L lo

111

■

L13

tiPc

t>Slo

IVM82

101

5.5

22.5

3,5

54,5

20,5

8,5

18.5

8,5

2,5

IVM83

100

2.5

54.5

19.5

17.5

8.5

17.5

7.5

27.5

11.5

2.5

[VM84

102

7.5

24.5

3,5

12,5

53,5

19.5

8,5

8.5

11.5

IVM85

21,5

17,5

8,5

17,5

10,5

3,5

IYM86

102

12,5

53,5

21.5

18,5

8.5

21,5

27.5

11.5

IVM87

102

3.5

8.5

17,5

26.5

11.5

2.5

3.6

IVM88

103

6.5

19.5

7,5

16.5

7,5

22.5

27.5

I1,S ,

1VM89

8,5

18.5

8.5

28.5

11.5

2.5

IVM90

103

7.5

20,5

8,5

18,5

8,5

8.5

30.5

12.5

2.5

3.5

IVM9I

21,5

14.5

21.5

9.5

18.5

18,5

18.5

12.5

IVM92

97.5

8.5

23,5

2.5

12.5

22.5

10,5

19.5

7.5

22,5

2,2

IVM93

23.5

2,5

13.5

22.5

8.5

28.5

10.5

2.5

IVM94

101

24.5

2.5

21.5

18,5

17,5

7.5

27.5

2.5

3.5

J VMM

21,5

7.5

17.5

21.5

2.5

3.5

IVM96

urn

7.5

27.5

21.5

26.5

1 1-5

2.2

3.5

tVM97

103

8.5

2,5

22.5

18,5

IVM98

104

102

21.5

18.5

17.5

9.5

11.5

2.2

3.5

[VMM

108

3,5

20.5

17,5

9.5

11,5

■J

3.5

1VMIOO

108

102

9.5

115

20.5

22.5

27.5

11.5 |

2,5

3,3

JVM 101

III

95.5

1.5

20.5

9.5

20.5

2,2

3.2

IVMJ02

1.2

!00

2,5

12.5

21.5

9.5

18,5

8,5

8.5

30.5

12.5

2.5

3.5

IVMI03

2.5

12.5

10.5

19.5

1 1.5

2.2

3.5

JVM 104

IIS

100

15.5

20.5

9.5

18.5

8.5

19.5

29.5

11.5

1.8

3.5

1VMI05

119

102

13.5

55,5

18.5

9.5

30.5

1.8

3.5

JVM 106

120

16,5

56,5

8.5

16,5

8.5

17.5

11.5

3.5

IVMJ07

121

!00

24,5

8.5

18.5

23.5

28.2

11.5

JVM 108

121

(02

1.5

8,5

18.5

28.3

11.5

2.2

3.2

1VMI09

125

9.5

12.5

18.5

8.5

11.5

JVM 110

135

106

25.5

11.5

8,5

21.5

9,5

18,5

31.5

60.5

JVM 1 1 1

132

1O0

27.5

0.5

12.5

8.5

17.5

22-2

30.2

11.8

2.5

JVM 112

140

105

12,5

1.5

23,5

9.5

18,5

11.5

2.2

2.5

IVMII3

126

101

1.8

11.5

22.5

9,5

19.5

9.5

19.5

95,5-

5,5-

21-

11.5-

53.5-

19.5-

7.5-

16.5-

17-

7.5-

IS-

26.5-

IQ-

1.8-

2,5-

Range

106

12.5

1-5

16.5

56,5

23,5

10,5

3.5

Mean

100.3

8.3

2,1

13.2

21.18

8,8

!8.6

9.1

19.5

20.9

29.2

11.5

2.3

3.5

Variance

10.5

4,5

1.5

2.5

6.5

1.7

1.5

Range - V

10.4

84.3

29,1

190.4

37,8

5,4

18.8

24.1

25,6

28,7

222

73,9

42.8

Table 3. Morphometric data in “Echino discus” cf. auritus, Recent, from Mangili, Tulear, Madagascar.

Analysis on a sample of Echinodiscus cf. auritus Leske, 1778 (Echinoidea Clypeasteroida)

285

Figure 3. Echinodiscus ” cf. auritus from Mangili, Tulear, Madagascar (Recent): a, b) respectively, aboral and oral plate

structure of MAC.IVM 87; c) food grooves.

f. auri f us ( n= 3 2 i

Length Frequenev Distribution

»!.s »,S N.S 104.3 ltl.S ISIS l«S.S IM.S

Scatter plot (oral face}

in n? r ,j lac rji p late s 5 1 n = 27)

O J 1 <—

0 5D 100 ISO

100

LIKIN lott'Hl Inn)

test lenjjlh (moi)

Scatter plot (aboral face} Scatter plot (aboral face)

interambulacraJplaies 5 (nni7j a inbLilaeral plates 3 (n»27)

a J

100

ISO

200

10 0

test length (own)

Test length (mm)

Figure 4. Ratio between number of plates and echinoids size, in a sample from Mangili, Tulear, Madagascar.

Figure 5. Ratio, by increasing dimension, between odd petals and corresponding number of pores.

286

Specimen

Dap

coupl

e of

pores

IVM 82

5.3

13.06

13.55

1 V M 83

75.6

14.3

1 2.65

12.8

IVM 84

78.7

15.7

14.45

13.85

1 VM85

79.3

17.1

14,3

14.3

IVMSG

80.2

17.8

15.2

14.3

1 VM

81.3

14.5

14.3

1 VM8S

82.2

13.5

1 VMS9

18.2

15.6

15.4

1 VM90

88.3

18.4

15.5

15.4

1 VM91

88.6

18.8

16.8

16.5

1 VM92

92.9

20.3

18.2

17.8

J VM93

93.7

20.8

17,8

17,45

1 VM94

95.6

20.6

17.3

16.2

1 VM95

96.5

19.7

17,85

16,35

1 VM96

97.9

19.5

16.65

17,45

I VM97

102.5

21.8

18.5

19.7

1 VM 98

104.2

22.4

17.95

1 VM99

108.2

21.8

18.65

18.25

rvMioo

1 10.8

23.4

20.3

19.55

I VM 10 1

111.4

23.4

20.55

( 11.8

23.8

19.6

20.5

24.8

20.6

20.3

1 VM 1 04

1 17.8

25.4

21 ,7

21.25

I VM 105

t 1 9.2

26.4

23.3

21.75

1 VM 1 06

119.7

23,1

19,65

1 VM 1 07

1 20.7

21.05

21.35

IVM 1 08

120.8

25.2

20.9

21.6

1 VM 109

i 24.9

23.35

IVM 1 10

1 34.9

32.1

28.8

27.8

Table 4 (upper). Data of petals dimension and corresponding

number of pores in ambulacral areas of “Echinodiscus” cf.

auritus, Recent, from Mangili.

Table 5 (right). Data of interambulacra disjunction variabi-

lity in “ Echinodiscus“ cf. auritus, from Mangili: C = Conti-

nues; D = disjoint; A = Amphiplacous; M = Meridoplacous.

Specimen

lal

Ia2

Ia4

fa5

p lutes

pOfi, Pc

IV MS 2

D-A

D-M

D-A

4a-5b

2a-2b

IVMK3

C-A

D-A

4a-4b

2a-2b

IVM 8 4

D-M

D-A

C-A

C-M

D-A

4a-4b

2a-2b

IV MS 5

D-M

C-A

D-M

D-A

4a- 5b

2a-2b-3b

IVM86

C-M

C-A

D-A

4a-5b

2a-2b

FVMS7

C-A

C-M

D-A

4a- 5b

2a -2 b

IVM8S

D-M

C-A

D-M

D-A

4a-4b

2a-2b

IV MS 9

D-A

D-M

D-A

4a- 5b

2a-2b

(VM90

D-M

D-A

C-A

D-M

D-A

4a- 4b

2a-2b

IVM 1 )!

D-M

C-A

D-A

D-M

4a-4b

2a.2b-3b

1VM92

D-M

D-A

D-M

D-A

4a-5b

2u-2b

IVM93

CVM

C-A

D-A

D-M

D-A

4a-4b

2a 2 b

IVM94

D-M

D-A

D-M

D-A

4a- 4b

2a-2b

IVM 1 ) 5

C-A

D-M

D-A

4a -4b

2a-2b

IVM96

D-M

C-A

D-M

D-A

4a-5b

2a-2b

IVM97

D-A

C-M

D-A

5a-5b

2a-2b

1VM98

C-M

C-A

C-M

D-A

4b-5a

2a-2b

IV MW

D-M

D-A

C-M

D-A

4a-5b

2a-2b

IVM 1 00

C-M

D-A

C-A

D-A

5a-5b

2a-2b

IVM! 01

D-M

D-A

C-A

D-M

D-A

4a- 5b

2a-2b

IVM 10S

C-A

C-M

D-A

4a -4b

2a-2h

IVM 103

C-A

C-M

D-A

4a- 5b

2a-2b

IVM 104

4a-4b

2a-2b

IVM 105

C-M

D-A

D-M

D-A

4a- 5b

2a -2 b

LVM106

C-M

C-A

D-A

C-A

D-A

4a-5b

2a-2b

T VM ] 07

C-M

D-A

C-A

D-A

4a-5b

2a-2b

IVM 1 0S

C-M

C-A

C-M

D-A

4a-5b

2a-2b

1 VM 1 (19

C-A

D-A

4a-4b

2a-2b

IVM HO

D-A

4a-4b

2a-2b

IVM 1 1 1

C-A

D-A

4a-5h

2a-2b

IVM 112

D-M

D-A

D-M

D-A

4a-4b

2a-2b

IVM] 13

D-M

D-A

D-M

D-A

-i.!-4V

2a-2b

IVM 081

D-A

D-M

D-A

D-M

D-A

4A4b

2a-2b

IVM 206

D-M

D-A

D-M

5A-5B

2a 2 b

IVM 207

D-M

D-A

D-M

D-A

4A4B

2a-2b

IVM20S

D-M

D-A

D-M

4A-5B

IVM 2 10

D-M

D-A !

D-A

D-M

4a-5b

2a-2b

IVM209

D-M

D-A

D-M

4a-4h

2a-2b

IVM2II

D-M

D-A

D-M

4a-4b

2a-2b

IVM212

D-M

d-a ;

D-A

D-M

4a -5b

2a-2b

IVM213

D-M

D-A

D-M

4a-4b

2a-2b

IVM214

D-M

D-A

D-M

4a-4b

2a-2b

IVM2I5

D-M

D-A

D-M

4a -4b

2a-2b

Analysis on a sample of Echinodiscus cf. auritus Leske, 1778 (Echinoidea Clypeasteroida)

287

Specimen

Apical

di sc

PL WA

1VM82

4.5

IVM83

t V M84

4.5

1VM85

4.5

1VMS6

4.5

[VMS 7

4.5

IV MSS

4.5

1VM89

4.5

IVM90

4.5

1VM9I

IVM92

5.5

1VM93

5.5

1VM94

5,5

IVM95

1VM96

1VM97

5.5

1VM98

5.5

IVM99

5.5

JVM 100

1 V M HI 1

IVMI02

5.5

IV Ml 03

[ V Ml 04

IV M 1 05

1 V M 1 06

5.5

1 V M 1 07

1 V M 1 08

4.5

IV Ml 09

IVM1 10

4.5

IVM 11 I

4.5

l V M 11 2

mean

5.6

39.8

32.3

55.4

Range

4 - 6

36-47

29 -37

48 -

Table 6. Data of apical disc, PL, WA at ambitus on interamb.

5, and angle B, in “ Echinodiscus ” cf. auritus, from Mangili.

The same can be said about the variation of PL,

which is higher (37-48% TL) in the sample of

Mangili compared to that of the Philippines (40-

48% TL) but with an average much lower (41

versus 43% TL). Also the variation of Lll is high

enough, but similar in the two samples (21% TL).

Even the measurement of WA to the interambula-

crum 5 and of the 13 angle, shows normal variability,

but, already at a first comparison with data from

other species of “Echinodiscus " , it seems to pro-

vide significant results.

In fact, the low average of WA and the low

grade of B, together with the still stretched shape,

makes these echinoids very characteristic and sepa-

rates them significantly from other groups of the

same family.

This will be useful to make direct comparisons

between samples of "Echinodiscus" mdAmphiope,

as done by Stara & Sanciu (2014).

CONCLUSIONS

Tests and observations obtained allow us to

several considerations. The variability of the

notches, and in particular of the LI is quite high,

while that of L2 is not verifiable because of

frequent malformations, however, these variability

does not seem to change the appearance of these

echinoids. The same consideration should be made

for the variability of the periproct position in

respect to the posterior margin. This is always very

high when compared with that detected in other

specimens of Echinodiscus seen in the literature

(see Stara & Sanciu, this volume). Other mea-

surements taken do not appear to show significant

levels of variability. In reference to the plating

(pattern plate), there is a great variability in the

disjunction of the post basicoronals in interambu-

lacra 1, 2, 3 and 4. An analysis of this unexpected

appearance is shown in Stara & Sanciu (2014).

Very important results relate to other fundamental

aspects for all those who will compete in this

family's systematic. Specifically, it was possible

to detect a marked stability in the development of

the scheme of interambulacrum 5 and in that of the

adjacent ambulacra I and V.

In fact, the periproct position and the plate shape

in interambulacrum 5, are very stable in both stud-

ied samples. Also the plates number per column

288

Paolo Stara & Maurizio Fois

I V

lla

ab ,

lib

Ilia

—

lib

nib

IVMS2

IVM83

1VM84

IV MSS

!VM86

13 !

1VM87

1VM8G

1VM89

1VM90

1VM91

! VM92

IVM93

IVM94

!VM95

FVM96

IVM97

13 |

IVM9S

IVM99

IVM100

JVW101

14 '

13 1

IVM102

1VM103

IVM104

IVM105

IVM106

1VM107

! VIM 108

14 |

IVM109

15 ;

IVM110

—

112

IVM111

1VM112

1VM113

Table 7. Numbers of plates (post basicoronal only) in some ambulacra and interambulacra in the sample of “Echinodiscus”

cf. auritus from Mangili. or = oral side; ab = aboral side; ZL = summa of oral + aboral number of plates.

Analysis on a sample of Echinodiscus cf. auritus Leske, 1778 (Echinoidea Clypeasteroida)

289

TL -5 oral

TL - 5 aboral

TL- 3 aboral

(a) r,

0.267

0.173

-0.218

(bk (0.05; 27)*

0.382

0,382

If a < b then

H a : p s = 0

TRUE

Table 8. * Critical value of the Spearman rank correlation rs

(a, n), with a = 0.05 and n = number of samples. This value

is extract from the table of critical values obtained through

the software SuppDist (Wheeler, 2005) implemented in R

and based on the method of Kendall & Smith (1939).

relative to interambulacrum 5 and to the two adja-

cent ambulacra seem to be very stable for all size

classes considered (see diagram in Fig. 4). On the

contrary, it was observed a good linear relationship

between the petals length and sample size. It is

considered, for each sample, the petaloid III with

the relative increase of the number of pairs of pores

(Fig. 5).

Is interesting to note that, despite the significant

variability of Lll, the position of the periproct,

compared to the scheme of the plates of these echi-

noids does not vary.

Also the size of PL, WA and B seems to charac-

terize these echinoids, particularly since, already at

a first glance, it distinguishes them very well, from

other groups of the same family, as you can see better

in Stara & Sanciu (2014). Even the observation of

the structure of the floor of the central hollow,

which seems to differ from that of other genera like,

for example, Amphiope, it may be helpful for future

comparisons between species of genera apparently

neighbours, through the work of those correlations.

Is very important, finally, the difference in

Aristotle's lantern size for the same TL between

Madagascar and Philippines specimens (Plate 6).

This suggests that there may be two distinct

species, but this will be subject of another work.

ACKNOWLEDGEMENTS

We warmly thanks Enrico Borghi of the Societa

di Scienze Naturali of Reggio Emilia, for critical

reading of the manuscript; and Mario Lai (3S,

Laboratories images, Capoterra) for scoring the

radiographs to the examined specimens.

REFERENCES

Agassiz L., 1838-1841. Monographic d'echinodermes

vivants et fossiles. Echinites. Famille des Clypeas-

teroides. Seconde Monographic. Des Scutelles. Neu-

chatel, 149 pp.

Alexander D.E. & Ghiold J., 1980. The functional signif-

icance of the lunules in the sand dollar, Mellita quin-

quiesperforata. Biological Bulletin, 159: 561-570.

Durham J.W., 1955. Classification of clypeasteroid

echinoids. University of California Publications in

Geological Sciences, 31: 73-198.

Kendall, M.G. & Babington Smith B., 1939. "The

Problem of m Rankings". The Annals of Mathemat-

ical Statistics, 10: 275-287.

Kier P.M., 1972. Upper Miocene Echinoids from the

Yorktown Formation of Virginia and their environ-

mental significance. Smithsonian contributions to

paleobiology, 13: 1-40.

Kroh A., 2005. Catalogus Fossilium Austriae, Band 2,

Echinoidea neogenica. Verlag der Osterreichischen

Akademie der Wissenschaften, 56: 1-210.

Kroh A. & Smith A. B., 2010. The phylogeny and classi-

fication of post-Palaeozoic echinoids, Journal of

Systematic Palaeontology, 8: 147-212.

Lohavanijaya P. & Swan E. F., 1965. The separation of

post-basicoronal areas from the basicoronal plates in

the interambulacra of the sand dollar, Echinarachnius

parma (Lamarck). Marine Biological Laboratory The

Biological Bulletin, 129: 167-180.

Loven S., 1872. On the structure of the Echinoidea.

Annals and Magazine of Natural History, 4: 285-298,

376-385, 427M44.

Pereira P., 2010. Echinoidea from the Neogene of Por-

tugal mainland. Palaeontos, 18: 1-154.

Philippe M., 1998. Les echinides miocenes du Bassin du

Rhone: revision systematique. Nouvelles Archives du

Museum d’Histoire Naturelle de Lyon, 36: 3-241,

249-441.

Smith A.B. & Kroh A., 2011. The Echinoid Directory.

World Wide Web electronic publication.

http://www.nhm.ac.uk/scienceprojects/echinoids (ac-

cessed September 2013).

Stara P. & Borghi E., 2014. The echinoid genus Am-

phiope L. Agassiz, 1840 (Astriclypeidae) in the

Oligo-Miocene of Sardinia (Italy). In: Paolo Stara

(ed.). Studies on some astriclypeids (Echinoidea Cly-

peasteroida), pp. 225-358. Biodiversity Journal, 5:

245-268.

Stara P. & Fois D., 2014. Dispute about Echinodiscus

Leske, 1778 and Amphiope L. Agassiz, 1840 (Echi-

290

Paolo Stara & Maurizio Fois

noidea Clypeasteroida Astriclypeidae). In: Paolo

Stara (ed.)- Studies on some astriclypeids (Echi-

noidea Clypeasteroida), pp. 225-358. Biodiversity

Journal, 5: 229-232.

Stara P. & Sanciu L., 2014. Analysis of some as-

triclypeids (Echinoidea Clypeasteroidea). In: Paolo

Stara (ed.). Studies on some astriclypeids (Echi-

noidea Clypeasteroida), pp. 225-358. Biodiversity

Journal, 5: 291-358.

Stelmle F.W. 1990. Population dynamics growth, and

production estimates for the sand dollars Echi-

narachnius parma. Fishery bulletin U.S., 88: 179—

189.

Wheeler, B. 2005. The SuppDist package, version 1.0-

13. Gnu Public License version 2. http://cran.r-pro-

ject.org/web/packages/SuppDists/index.html

Biodiversity Journal, 2014, 5 (2): 291-358

Analysis of some astriclypeids (Echinoidea Clypeast-

eroida)

Paolo Stara 1 * & Luigi Sanciu 2

'Centro Stndi di Storia Naturale del Mediterraneo - Museo di Storia Naturale Aquilegia, Via Italia 63, Pirri-Cagliari and Geomuseo

Monte Arci, Masullas, Oristano, Sardinia, Italy; e-mail: paolostara@yahoo.it

* Corresponding author

The systematic position of some astriclypeid species assigned through times to the genera

ABSTRACT Amphiope L. Agassiz, 1840 and Echinodiscus Leske, 1778 is reviewed based on the plating

pattern characteristics of these two genera universally accepted, and on the results of new

studies. A partial re-arrangement of the family Astriclypeidae Stefanini, 1912 is herein pro-

posed, with the institution of Sculpsitechinus n. g. and Pciraamphiope n. g., both of them char-

acterized by a peculiar plating-structure of the interambulacrum 5 and of the ambulacra I and

V. Some species previously attributed to Amphiope and Echinodiscus are transferred into

these two new genera. Two new species of Astriclypeidae are established: Echinodiscus

andamanensis n. sp. and Pciraamphiope raimondii n. sp. Neotypes are proposed for Echin-

odiscus tenuissimus L. Agassiz, 1840 and E. auritus Leske, 1778, since these species were

still poorly defined, due to the loss of the holotypes and, for E. auritus , also to the unclear

geographical/stratigraphical information about the type-locality. A number of additional nom-

inal fossil and extant species of " Echinodiscus " needs revision based on the same method.

KEY WORDS Astriclypeidae; Amphiope ; Paraamphiope; Echinodiscus', Sculpsitechinus; Oligo-Miocene.

Received 28.02.2014; accepted 14.03.2014; printed 30.06.2014

Paolo Stara (ed.). Studies on some astriclypeids (Echinoidea Clypeasteroida), pp. 225-358.

INTRODUCTION

The classification of the astriclypeid echinoids

Amphiope L. Agassiz, 1840 and Echinodiscus

Leske, 1778 have been traditionally based on the

external morphological features, mainly test

outline, size and shape of lunules and petals (see

Durham, 1955). Structural characters, largely used

in the taxonomy of other clypeastroids, were prac-

tically ignored in earlier studies dealing with these

genera, and although several species have been

described in the literature, important features for

species-level taxonomy, such as oral plating, were

poorly illustrated or omitted completely.

MATERIAL AND METHODS

The studied specimens are housed in the fol-

lowing public institutions: MAC (Museo di Storia

Naturale Aquilegia) and UNICA (Department of

Animal Biology and Ecology, University of

Cagliari) Cagliari, Italy; MSNDG (Museo di Storia

Naturale Doria, Genoa) and UNIGE.SM (Dip.Te.

Ris, University of Genoa), Genoa, Italy; NHMUK

(National History Museum of United Kingdom)

London, England; ZM (Zoological Museum of

Denmark, University of Copenhagen) Copenhagen,

Denmark; PMBC (Phuket Marine Biological

Centre), Phuket District, Thailand.

292

Paolo Stara & Luigi Sanciu

Figure 1. Biometric parameters measured in the studied samples.

Some other specimens, used for comparison, are

kept in private collections, as cited.

43 specimens attributed to Echinodiscus and 29

to Amphiope were examined. 8 Amphiope fossils

from Touraine, France; 1 Echinodiscus fossil from

Hurgada, Egypt; 1 ‘Amphiope“ fossil from Liguria,

Italy and, 1 “ Echinodiscus ” Recent from Lembeh,

North Sulawesi, Indonesia; 2 Echinodiscus Recent

from Nosy Be, Madagascar; 33 “ Echinodiscus

auritus ” Recent from Mangili, Madagascar; 10

“ Echinodiscus ” Recent from Philippines; 1 “ Echin-

odiscus ” Recent from Indonesia (Borneo) are

housed in MAC; after study, some specimens will

be deposited at the UNICA; 3 “ Amphiope ” fossils

from Liguria, housed to the MSNDG and 2 fossil

specimens of “Amphiope” at the UNIGE.SM (Dip.

Te. Ris); 21 Echinodicus and 10 “ Amphiope ” at the

NHMUK; 1 “ Echinodiscus ” at the ZM; 6 Echin-

odiscus at the PMBC.

Three specimens of “ Echinodiscus ” used for

comparison belong to private collections; some

plating patterns were taken from illustrations re-

ported in the literature.

Measures taken as in figure 1 . The plating pat-

tern follows Durham (1955 ) and, when possible,

includes both sides of the specimen. To highlight

the sutures, humidification by denatured ethyl alco-

hol has been used for extant species, a mixture of

water and hydrochloric acid (ca. 2%) for some fos-

sil specimens. The internal structure was studied by

sectioning the test, and in some cases by X-ray.

Morphological abbreviations as in figure 2: B =

angle between major axis of the two lunules; TL =

test length; TW= test width; TH = test height; Ll-

L2 = lunule length and width, respectively; L3 = di-

stance between posterior petal-tip and lunule, L4 =

distance between apical system-posterior margin,

L5-L6 = length and width of the frontal petal,

respectively; L7-L8 = length and width of the ante-

rior paired petal, respectively; L9-L10 = length and

width of the posterior petal, respectively; LI 1 = di-

stance between periproct-posterior margin test; L12

= distance between the posterior border of the peri-

stome and of the periproct, LI 3 = front-rear diame-

ter of the ambulacral basicoronal circlet. PL =

petalodium lenght; WA= ambulacral and interam-

bulacral width at ambitus; o pc = periproct diame-

ter; o ps = peristome diameter ; E = summation. To

describe the lunules shape and dimension into a

numeric value, we introduced a Shape Index (SI)

corresponding to the ratio L2/L1 and a Width Index

(WI) = (LI + L2) / 2.

Species with doubtful taxonomic attribution are

marked by quotation marks.

Analysis of some astriclypeids (Echinoidea Clypeasteroida)

293

I. DESCRIPTION OF THE SAMPLES

EXAMINED

“ Amphiope ” sp.

Plates 1, 2; Tables 1, 2

Examined material. Eight specimens from

Channay-sur-Lathan, Touraine, France, Late Ser-

ravallian-Early Tortonian, TE 47 - 73 mm.

Description. Small to medium sized echinoid

with low test and small rounded lunules. The lunules

show a low variability range: SI ranges from 1 to

1.6 (lunules roundish to slightly transversely elon-

gated), WI ranges from 9 to 10.5 (small lunules). In

this sample, the lunules variability equals on the

average 34% LI and 27% L2.

In the oral interambulacrum 5 there are only the

post-basicoronal plates 2a, 3a and 2b, 3b and, in

some cases, a small portion of 4b. The plate 2a is

long and staggered with respect to 2b; the periproct

opens between 2a/3b. Aborally, the tips of the pos-

terior petals are separated from the lunules by 1 or

2 couples of plate.

“Amphiope” pedemontana Airaghi, 1901

Examined material. Rupelian, Piedmont and

Liguria, Italy. The holotype was housed in the Civic

Museum of Natural Histoiy of Milan; it was lost dur-

ing the Second World War. The specimens in the

Genoa museum indicate the occurrence of two dif-

ferent morphotypes under this name: one of them

corresponds to the description of " Amphiope " pede-

montana Airaghi, 1901, the other seems different.

First morphotype

Figures 2a, b; Table 3

Examined material. Three specimens: MSNDG.

N25 from Pareto, MSNDG. N12 14 from Cairo Mon-

tenotte and MSNDG.N1218 from unknown locality,

TL 53 ^ 61 mm; two illustrations given by Airaghi

(1899 and 1901) of a specimen from Dego and an-

other one from Santa Giustina; one specimen from

Merana: MAC.PL2014, TL, 71 mm, TH 7 mm.

Description. Small to medium sized form, with

small axial lunules, small and open petals, very de-

pressed test and drop-shaped periproct. In the oral

interambulacrum 5 there are only the post-basi-

coronal plates 2a, 3a and 2b, 3b, all of them large

and paired; the periproct opens between plates

2a/3b (Figs. 2a, b).

Second morphotype

Fig. 2c; Table 3

Examined material. Two whole specimens

(UNIGE.SM-VI-P-(5)-DN and UNIGE.SM-VI-

DR) and 2 test fragments, from Pareto.

Table 1. Moiphometric data of Amphiope sp. 3. TL in mm, other measures in % TL.

294

Paolo Stara & Luigi Sanciu

Figure 2. “Amphiope” pedemontana (Oligocene, Liguria and Piedmont, Italy): a, b: respectively, aboral and oral plate

structure of MSNDG. 12 1 8; c: “ Amphiope'' sp. 1 (Oligocene, Liguria and Piedmont, Italy), oral plate structure of UNIGE.SM-

VI-P5-DN; d: “ Amphiope ” arcuata (Miocene, Libya), oral plate structure.

Amphiope sp. 3

Apx

PL 1668

52°

PL 1669

64°

PL1821

54°

PL 1822

7 1 °

PL 1823

PL 1824

67°

PL 1825

64°

PL 1826

53°

mean

60.7°

range

50-

29*

52-71

Table 2. Apx, PL, WA and « data of Amphiope sp. 3.

TL in mm, « in degree, other measures in % TL.

Description. Small sized form with closed

petals and a notch along the posterior margin, close

to the periproct. The periproct is rounded and opens

between plates 2a/3b. In the interambulacram 5

there are only the post-basicoronal plates 2a, 3a and

2b, 3b, that are large and paired.

"Amphiope" arcuata Fuchs, 1882

Fig. 2d; Table 4

Examined material. Five specimens from the

“Miocene” of the Libyan desert (locality not speci-

fied), housed in the NHMUK (code El 67 1-2,

E1674-6), TL 35-79 mm.

Description. Small to medium sized echinoid,

with very low test and thin ambitus; test outline

rounded or sub-trapezoidal. In the interambulacrum

5 there are two plates per column: 2a, 3a and 2b 3b;

the plates 2a/2b are staggered. These specimens are

characterized by small ovoid axial lunules, distant

from the corresponding petal tips. Lunules show a

low variability since SI ranges from 0.45 to 0.76

(axially elongated lunules) and WI ranges from 8

to 10.5 (small lunules). On average the lunules

variability equals 34% of LI and 27% of L2. PL

ranges from 42 to 46% TL.

"Amphiope" duffi Gregory, 1911

Plate 3 Figs. 1-6

Examined material. Rupelian, Libya. Two syn-

types housed in the NHMUK: CY66/E1 1350, from

Sidi Rof Diasiasia, Cyrenaica, TL = 37 mm and

Cy264/E11349, from Ain Sciahat, Cyrenaica, TL =

39 mm.

Description. Small and low test with thin

ambitus; test outline sub-rounded. Adoral face

unknown. Width of the interambulacrum 5 at the

ambitus about 23% TL. In Cy66 the petals are dis-

tally open. Petals are sub-equally sized; PL = 47%

Analysis of some astriclypeids (Echinoidea Clypeasteroida)

295

TL. Lunules are apparently open (notches?), but it

is not clear whether this is due to damage; in C66

they are very small and ellipsoidal. B measures 65°.

On the aboral side each lunule is separated from the

petal tip by 3-4 couples of plates and is surrounded

by 3 - 4 couples of plates. The apical disc is small

(~ 8% of TL) and star-shaped. The internal structure

is unknown. Number of plates per column only par-

tially visible (see Table 7).

“ Echinodiscus tenuissimus ” L. Agassiz, 1 847

Examined material. Recent, Indian Ocean, In-

donesian Archipelago, Oceania and China Sea.

Remarks. The holotype was established by L.

Agassiz (1847) in Agassiz & Desor (1847) on the

basis of a specimen with small axial lunules, from

Waigiu (New Britannia, Western Papua, Indonesia)

and housed at the Museum of Natural History, Paris.

Actually the holotype is wanting (personal commu-

niation by Sylvain Charbonnier, June. 03. 20 14). The

group of Recent specimens under study indicate the

occurrence of three different morphotypes: one of

them shows some characteristics of the genus type

E. bisperforatus Leske, 1778, the others seem dif-

ferent.

Figure 3. Plate structures of adoral side of the “ Echinodiscus tenuissimus ” morphotypes; postbasicoronal plates of

interambulacrum 5 colored - a, first morphotype; b, second morphotype; c. third moiphotype.

Figure 4. “ Echinodiscus bisperforatus truncatus“, plate pattern of oral (a) and aboral sides (b).

296

Paolo Stara & Luigi Sanciu

Table 3. M morphometric data of Echinodiscus pedemontanus (former Amp hi ope pedemontana)

and Echinodiscus sp. 1. TL in mm, other measures in % TL.

NHMUK.E76I61

NHMUK.E76162

NHMUK.E76166

NHMUK.E76165

NHMUK.E76164

Mean

Table 4. Morphometric data of Paraamphiope arcuata (former Amphiope arcuata ). TL in mm, other measures in % TL.

Analysis of some astriclypeids (Echinoidea Clypeasteroida)

297

First morphotype

Fig. 3a

Examined material. One specimen from Lem-

beh Channel, North Sulawesi (Indonesia), MAC.

IVM 207, TL = 50 mm; one specimen from New

Caledonia, NHMUK. 198 1.1 1.2.25, TL = 112 mm;

one specimen from Palau, Micronesia,

NHMUK.59.7.1.14, TL= 120 mm; two specimens

from Lembeh Channel, North Sulawesi (Indonesia)

TL 50-65 mm from the M. Fantin collection and one

from Noumea, Baie des Citrons, New Caledonia, TL

68 mm, from the F. Hattemberger collection.

Description. Middle size test echinoids with

small slit like axial lunules and small petals. Very

flat test and thin ambitus, with an elongated and

more rounded anteriorly outline. In the oral inter-

ambulacrum 5 there are two-three plates in column

a(2a, 3a, 4a) and three in column b (2b, 3b, 4b); in

which the plates 2a and 2b are more or less stag-

gered and the periproct opens between plates 2a/2b.

The B angle is low (65-70°, mean 67°) and WA at

interambulacrum 5 is small (mean 32% TL).

Second morphotype

Fig. 3b

Examined material. Two specimens from Pak

Meng Beach, Trang Province, Thailand; PMBC.

26346, 2842, TL = 81 and 66 mm; two specimens

from Noparat Tara Beach, Krabi Province,

PMBC.2843, 2830, TL = 66,2 and 54,6 mm; one

specimen from PMBC Jetty South, Phuket,

PMBC2844, TL = 66,2 mm; one specimen from West

side of Ko Yao Yai, Phuket, housed in the NHMD.Z

n° ZMUC-ECH-1001, TL 37 mm (see also Waren &

Crossland, 1991: figs. 10a, c); one specimen from

“Thailand”, Recent (based on a illustration in “www.

Echinoids NL” by Bas van der Steld, Netherlands).

Description. Small size tests echinoids, with

ovoidal axial lunules, very flat test and thin ambitus,

sub-rounded in shape. In the oral interambulacrum

5 there are two postbasicoronal plates per column

(2a, 3a and 2b, 3b), paired and wide. The B angle is

small (75,5°). The WA at interambulacrum 5 is

about 38% TL. Since only a small sample is avail-

able to study, it is not possible to verify the vari-

ability of the lunules.

Third morphotype

Fig. 3c

Examined material. One specimen from Indone-

sia (Borneo), Recent, MAC.IVM206, TL =53 mm.

Description. Small sized echinoid, with a flat test

and slit-like axially elongated lunules. Petals small,

closed distally. In the oral interambulacmm 5 there

are 2 postbasicoronal plates per column (2a, 3a - 2b,

3b), with the first two staggered. The 2b is in am-

phiplacous contact with the first postbasicoronal

plates of ambulacra I and V. Between the petal tips

and the notches there are 3 couples of plates, and the

periproct opens between plates 2a/3b. The WA at the

interambulacrum 5 is 38% TL; the B angle is 80°.

Echinodiscus bisperforatus truncatus

(L. Agassiz, 1841)

Examined material. Some Recent specimens

examined in the Fantin collection (Venice, Italy),

labeled E. truncatus (Fig. 4a, b), recently dredged

near Singapore, allowed to observe the plate struc-

ture and other characteristics. It differs from the

previous “second morphotype” by some noticeable

features. The echinoids collected in Singapore have

the plate pattern that match with those of the second

morphotype, but have the peristome smaller, more

branched food grooves, the apex much further for-

ward, lunules longer and slit-lilce and the ambital

outline with the posterior margin truncated. To com-

pare other characters we took some pictures of E.

truncatus in situ, from www.wildsingapore.com

(Mega Marine Survey of Singapore) and we have

established new differences, as we will see in the

discussion ad in the systematics chapters.

Echinodiscus bisperforatus Leske, 1778

Plates 4, 5; Table 5

Examined material. Recent, Red Sea and

Indian Ocean. Seven specimens from South Africa

(locality not specified) NHMUK: NHMUK 2013.7-

13, TL = 26 - 62 mm, eleven specimens from

Wakiro, Massawa, Eritrea, Red Sea, NHMUK.

1 965 .1.11 -20, TL = 46 = 69 mm, one specimen from

Pangani, Tanga, Tanzania, NHMUK. 1957. 5. 2 1.3.

TL = 84 mm; two specimens from Nosy Be Island,

Northern Madagascar, TL = 25 and 45 mm, the

smaller one housed at the (MAC.IVM208), and the

large one from a private collection.

298

Paolo Stara & Luigi Sanciu

Sped men

L 1 1

NHMUK. 1965-6-1-1!

9.8

115

12.5

101

NHMUK. 1965-6-1 -12

67.5

120

2.5-5

8.5

109

NHMUK. 1965-6-1 -13

119

31.5

2.5-5

103

NHMUK. 1965-6-1-14

8.5

122

2.5-5

102

NHMUK. 1965-6-1-15

113

NHMUK. 1965-6-1 -16

115

NHMUK. 1965-6-1 -17

119

9.5

NHMUK. 1965-6-1-18

7.5

112

8.5

107

NHMUK. 1965 -6- 1-19

113

107

NHMUK. 1965-6-1-20

6.5

113

104

NHMUK. 1965-6-1-2!

6.5

109

mean

8.6

115.4

30.2

9.2

41.2

105

range

6.5-1 1

109-120

28-34

2.5-5

7-12.5

46- 54

38-47

101-109

Specimen

MAC.IVM 208

113

3.5

Specimens

Dap

LI 1

NHMUK 301 3.7

9.5

115

3-3-4

5.5

103

NHMUK 30 13-8

115

33.5

3 -4-4 ,5

5.5

105

NHMUK 30 13.9

116

3-3-3. 5

106

NHMUK. 301 3. 10

53.5

114

33.5

34-5

106

NHMUK.30l3.il

115

34-5

108

NHMUK .301332

122

444

117

NHMUK .3013.13

6-6-6

! 02

mean

7.9

1 15.7

—

4.5

107

range

7-9.5

113-122

31-38

—

3-5.5

45-53

102-117

Speci men

Ll 1

NHMUK, 1957.5.21 -3

111

3.5-3 .5-3

lit

Table 5. Simplified morphometric data of Echinodiscus bisperforatus from different localities;

TL in mm, B in degree, other measures in % TL.

Analysis of some astriclypeids (Echinoidea Clypeasteroida)

299

Figure 5. “ Echinodiscus

sp. 2“ (Pleistocene-

Holocene, Hurgada,

Red Sea, Egypt), plate

pattern of oral (a) and

aboral sides (b).

Figure 6. “ Lobophora

aurita”, plate pattern,

taken from the illustra-

tion by L. Agassiz

(1838-41: table 14,

figs. 1, 2): adoral (a)

and aboral view (b).

Description. Middle size tests echinoids, with

flat test with rounded to sub-trapezoidal outline. In

the interambulacrum 5 there are two plates per

column, 2a, 3a and 2b, 3b, paired and wide (see

Plate 5 Fig. 2). The B angle is about 110°; the

lunules are very long and show a low variability.

The WA is high (47-50% TL).

Echinodiscus sp.

Fig. 5a, b

Examined material. One specimen from the

Pleistocene of Hurghada, Red Sea, Egypt, MAC.PL

1850, TL = 21 mm.

Description. Small sized echinoid, with flat test,

thin ambitus and test outline rounded. In the oral in-

terambulacrum 5 there are two plates per column

(2a, 3a, 2b, 3b), paired and wide, and the periproct

opens between plates 2a/3b. The B angle is 80°.

Echinodiscus desori Duncan et Sladen, 1883

Plate 6 Figs. 1-6

Examined material. Four specimens from the

Miocene of the Gujarat State, northern India:

NHMUK.E7 8129, TL 49 mm; NHMUK.E724b,

TL 39 mm; NHMUK.E78128a (TL 47 mm) and b

(TL 47.5 mm).

Description. Small size and very depressed test

(TH = 7-5-11% TL). The ambitus is thin and with

sub-rounded outline. The oral surface is exposed

only in specimen NFIMUK.E78128a, with the plat-

300

Paolo Stara & Luigi Sanciu

ing pattern only partially visible. The WA at the in-

terambulacrum 5 is about 30 ^ 35% TL in

NHMUK.E724b. The petals are sub-equal in size,

they are distally open or tend to opening (e.g. in

NHMUK.E724b). The axial lunules are medium

sized, ellipsoidal shaped. In NHMUK.E78129 they

are both incomplete. The B angle is low (68° to 74°).

“ Amphiope bioculata“ des Moulins, 1835

Plate 7 Figs. 1-11

Examined material. Based on eleven specimens

illustrated by Cottreau (1914), from the Helvetian

(Burdigalian in Philippe, 1998) of Saint-Cristol,

Nissan, Herault; pi. VI, figs. 1-11, TL 43 -s- 67 mm.

Description. Small sized echinoids, slightly

wider than long (TW = 103 110, the mean mea-

sure is 106% TL). Test depressed, however the

measure of the height is unknown. The ambitus

outline is sub-rounded. Inflections occur in the

ambitus in correspondence with the ambulacra II,

III and IV. The adoral surface is flat or slightly

concave; plating not detectable. The petals are

closed and sub-equal; PL ranges between 42 to

55% TL (mean 49.5% TL). The lunules are very

distorted, relatively small and rounded and close

to the tips of the corresponding petal. It is clear

that the size variability of the lunules is high

(Table 6; Figs. 11, 12), with a variation range of

LI which exceeding 50% on the average value and

that of L2 which exceeds 45% on the average

value. However, the SI varies from 0.95 to 1:47

Speci-

mens

Fig. 1

9.5

0.95

9.75

Fig. 2

1.25

Fig. 3

1 .08

12.5

Fig. 4

1.08

12.5

Fig. 5

8.5

10.5

1.23

9,5

Fig. 6

7.5

1.33

8.75

Fig. 7

7.5

1.33

8.75

Fig. 8

1.12

8.5

Fig. 9

1.47

Fig. 10

1.33

10.5

Fig. 11

1,37

9.5

mean

9.13

1.22

10.20

range

7,5-12

9-14

0.95-1.47

9-12.5

variance

49.2

45.4

Table 6. Variability data of the lunules in Amphiope

bioculata in Cottreau's sample.

Figure 7. Aboral plate structures of “Echino discus” from Taiwan and Japan; a: E. formosus, ?Eocene-Miocene, Taiwan

(from an illustration in Tokunaga, 1901, pi. 1, figs. 1, 2); b: E. yeliuensis, Miocene, Taiwan (from Wang, 1984, pi. 1, fig. 2a,

b); c: E. cikuzenensis, Oligo-Miocene, Japan (from Takano et al., 2007, pi. 1, fig. 12); d: transiens, Miocene, Japan

(from Nisiyama, 1966, pi. 17, fig. 1).

Analysis of some astriclypeids (Echinoidea Clypeasteroida)

301

while the WI ranges from 9 to 12.5, indicating that

the lunules are always rounded, while varying

especially in amplitude.

“ Echinodiscus auritus ” Leske, 1778

Fig. 6a, b

Examined material. Based on the illustration of

Lobophora aurita by L. Agassiz (1841), as L.

aurita, pi. 14, fig. 1,2; TL 110 mm, TH 11% of

TL; Recent, Red Sea, Egypt.

Description. Medium to large-sized echinoid

with polygonal ambitus outline and two long poste-

rior notches. The adoral surface is flat or slightly

plano-concave. In the oral interambulacrum 5 there

are four postbasicoronal plates per column (2a, 3a,

4a, 5a and 2b, 3b, 4b, 5b); the periproct opens far

from the posterior margin (LI 1 = 19% TL) between

plates 3b/3a. Aborally, there are 5-6 couples of plates

between the notches and the posterior petal tips.

Remarks. The illustration given by L. Agassiz

(1841) was the first to highlight the plate structure

of this echinoid. These data are not even reported

in the recent works, for which we detect the com-

plete plating, which is veiy different from those of

Mangili and from the Philippines, examined by

Stara & Fois M. (2014).

Echinodiscus /brjwtfsws Yoshiwara, 1901

Fig. 7a

Examined material. Based on the illustration by

Tokunaga (1901-3: pis. 14 and I, fig. 2); TL 100 mm.

1 specimen from Middle Eocene? to Miocene, Hatto,

Kelung, Taiwan. Plating of the aboral side taken from

fig. 2, pi. I; inclination of the lunules as in pi. II fig. 2.

Description. Medium to large sized echinoid

(max TL = 140 mm), with depressed test and with

sub-ellipsoidal ambitus outline. The estimated WA

of the interambulacrum 5 obtained by measuring

the half visible, seems to be the 50% TL. The petals

are closed, sub-equal in size; Tokunaga affirms that

the anterior odd petal is 25% and the other are

22.5% TL, but the illustration indicates that they all

measure the 20% TL. The lunules are large and

ellipsoidally shaped. The B angle is large (111°).

The lunules are surrounded by 5 pair of plates on

the aboral face. The partial number of plates per

column is shown in Table 7.

Echinodiscus yeliuensis Wang, 1982

Fig. 7b

Based on illustration in Wang (1984), from the

Taliao Formation (Aquitanian), of Yeliu, Taiwan,

pi. I, fig. 2a, b; topotype n° NTUG - [E] - 81.42; TL

= 112 mm; TW =131 mm.

Description. Medium to large sized echinoid

with depressed test (TH = 10% TL). The ambitus

outline is sub-trapezoidal, wider near the rear.

Only a part of the plating of the apical surface is

detectable. The estimated measure of WA at the

interambulacrum 5 is 34% TL. The petals are sub-

equal in size; the petalodium is wide (PL = 52%

of TL). The lunules are long and narrow, broader

anteriorly, lanceolate shaped and their axis devi-

ates substantially from the corresponding petals

(B angle about 1 14°). There are 2 couples of plates

between the petals tips and the corresponding

lunules.

Echinodiscus cikuzenensis Nagao, 1928

Fig. 7c

Examined material. Based on illustration in

Takano etal. (2007) pi. 1, fig. 11; 1 specimen of Edu-

cation Kawai, Oligocene-Miocene in age, from

Chugoku and Kyushu Province, Japan. TL unknown.

Description. The test outline in sub-rounded.

The aboral face is incomplete. The petals are sub-

equal in size; the PL is small (44% TL). The lunules

are large, sub-ellipsoidal shaped and deformed and

very close to the corresponding petal tip. The right

lunule is surrounded by 6 couples of plates on the

aboral side. The B angle is 73°.

Echinodiscus transiens Nisiyama, 1968

Fig. 7d

Examined material. Based on the specimen il-

lustrated in Nisiyama (1966), pi. 17, fig. 1; IGPS

collection, No. 37773, from the Yamaga Formation,

Miocene, Yamaguchi Prefecture, Japan. TL 102 mm.

Description. Medium sized, with a depressed

test; TH unknown. Test outline sub-rounded. The

aboral face is incomplete. The petals are sub-

equal and the petalodium is wide (52% TL). The

unique visible lunule is large and sub-rounded

shaped.

302

Paolo Stara & Luigi Sanciu

Specimens

ta 5

amb

atnb

la 1

1a4

amb

Amb

Ja2

Ia3

atnb

III

Echinodiscus formosus

x+6

x+5

Middle Eocene

x+6

x+5

Echinodiscus pederttaniaitus

x+9

X+8

...

Rupeliaai

x+8

X+7

Echinodiscus tiheenensis

x+6

x+10

x+4

x+8

X+5

Qligo - Miocene

x+6

x+9

x+5

x+R

Echinodiscus wihiaais

x+10

k+7

X+R

x+II

X+10

Eariy Miocene

x+10

x+8

X+8+

x+ll

X+10

Anifihiofte mmigica

4+9

7+9

8+9+

4+11

4+10

5+4

4+6+

4+S

5+5

MA C PL 1684, Oi igo-M \ oeone

5+10

R+10

7+9

3+10

x+l t

5+4

4+4

5+6+

4+7

6+5

AmpMofie nuragjca

4+15

7+10

8+1 1

4+12

x+13

x+5

4+10

5+6

MAC. PL 1680, 01 igo-M iocene

5+14

7+11

7+10

x+12

4+13

5+5

x+6

4+11

X+10

6+5

Amfthinpe sp, 2

MAC PI I aie

3+1 1

7+8

7+7

4+7

x+R

6+5+

x+4

5+10

4+ II

5+5

4+11

7+7

6+7

x+8+

4+7

6+5+

x+5

4+11

5+10

5+6

Btodigahan

Amphiope sp. 2

3+1!

8+8+

8+7

4+10

3+-

6+4

6*4+

4+9

5+9

6+^

MAC.PL552. Late

Burdiaalian

5+10

8+9

8+8

3+10

4+«

6+5

6+6+

4+9

6+4

.1 mf>h .sp. 3 MAC:. PL 1 669

3+10

6+7

7+7

4+1!

3+10

5+4+

5+5

4+9

4+10

6+4

Sen-avail ian ■ Tortonian

4+1!

7+7

6+7

3+ 1 1

4+9

5+4

6+4

4+!0

3+9

5+4

Sht ipsitechimts temt issimtts

3+6

7+8

7+6

5+8

13+

4+9

5+6

6+ (-)

4+H

S-K-)

6+^

Recent

4+6

7+7

7+6

4+6

10+

5+10

6+()

6+H

5+H

4+M

6-K

Sat Ipsilechinus a teilus

—

4+10

7+7

5+11

4+10

6+6

6+6+

5+9

5+10

6+6

M A C, IV MR 7, Recent

5+10

7+8

6+7

4+1 1

5+!0

7+5+

6+6

5+9

4+10

5+7

Sculpsilechimts sp. I

4+1!

7+7

7+8

5+11

4+10

6+6

6+7

4+10

5+11

6+6

MAC IV M2 10, Recent

4+1!

8+7

7+6

4+11

5+11

6+7

6+6

5+10

5+ II

7+6

Echimclisctts andumunciviis

3+9

5+6

5+5

4+8

4+R

5+5

5+9

4+6

6+5

Thai la India, Recent

3+9

5+5

5+8

4+8

6+4

4+9

4+5

6+6

Echi nodi sews tmnea Ms

3+10

6+6

6+4

4+10

3+9

5+6

5+5

4+8

5+5

Singapore. Rcoeni

3+9

6+5

5+6

4+9

5+5

4+R

5+5

Echinodiscus Uspafomtus

3+9

7+6

8+6

5+1!

4+9

6+5

6+4

4+10

5+!0

7+4

4+9

8+5

7+6

4+10

6+5

6+4

4+10

5+10

7+4

1 1

Table 7. Number of plates in some Astriclypeids species. Ia = interambulacram; Amb = ambulacrum;

£ = summation of oral and aboral plates per column.

II. MAIN CHARACTERISTICS DISTINGUI-

SHING AMPHIOPE FROM ECHINODISCUS

Durham (1955: 154, fig. a, b) and Smith &

Kroh (2011) indicated some characteristics of the

oral plating which distinguish Amphiope from

Echinodiscus. In species belonging to Amphiope

in the oral interambulacrum 5 there are two postba-

sicoronal plates in column a (2a, 3a) and three in

column b (2b, 3b, 4b), with the plate 2b more elon-

gated and staggered than the 2a and in amphipla-

cous contact with the first two adjacent

postbasicoronals ambulacral plates (Fig. 8a). In

species belonging to Echinodiscus , in the oral in-

terambulacrum 5 there are two postbasicoronal

plates per column in wich the first two plates 2b/2a

are similar-sized and paired (Fig. 8b).

From Kroh (2005), Pereira (2010) and Stara &

Borghi (2014), we have taken other characteristics

that distinguish Amphiope from others genus. Since

the shape of the lunules in E. bisperforatus is hardly

distinctive, we found the other characters of this

genus by a number of samples stored in a museum,

how above documented.

In the aboral face of Amphiope, there are always

one-two pairs of plates between the petal tips and

the corresponding lunules and the plates surround-

ing the lunules are arranged in a radial manner

(Plate 8 Figs. 1, 2; Plate 9 Figs. 1, 2; Plate 10 Figs.

1, 2). In the aboral face of E. bisperforatus, there

Analysis of some astriclypeids (Echinoidea Clypeasteroida)

303

Figure 8. Plate patterns of adoral sides of Amphiope “bioculata“ and Echinodiscus bisperforatus from Durham

(1955: p. 154, figs, a, b) and Smith & Kroh (2011); in red the oral postbasicoronal interambulacral structures.

Figure 9. Sculpsitechinus auritus from Mangili (Recent, Tulear, Madagascar); a, b: respectively,

aboral and oral plate structure of MAC.IVM 87; c: food grooves scheme.

are similarly one-two pairs of plates between the

petal tip and the corresponding lunules, but the

plates surrounding the lunules are arranged in a lin-

ear manner (Plate 9 Fig. 5). Finally, Smith & Kroh

(2011), state that Amphiope have roundish-ovoid

transverse lunules, while Echinodiscus have ovoidal

axial lunules or notches.

Based on these characters, and others that have

already been published in the pages of Amphiope

and Echinodiscus genus in Smith & Kroh (2011),

we have included in these genera the morphotypes

whose the plate patterns of interambulacrum 5,

coincided with those described by Durham (1955)

and Smith & Kroh (2011).

According to logic, the forms that do not match

with any of the two types have been treated by us

and characterized as belonging to several new

genera. Then, in the following we will use the terms

Amphiope and Echinodiscus to indicate any form

of astriclypeids corresponding to the description

summarized above.

304

Paolo Stara & Luigi Sanciu

DISCUSSION ON SYSTEMATICS ASPECT

The specimens of " Echinodiscus auritus ” de-

scribed by Stara & Fois (2014) do not match with

the above reported concepts of Echinodiscus and

Amphiope. The first two postbasicoronal plates 2b

and 2a in the oral interambulacrum 5 are partially

staggered and the total number of plates per column

is higher (4 in column b and 3-4 in column a) than

those of Echinodiscus and Amphiope (see Plate 1 0

by comparison). Also the couples of plates present

between the tip of each petal and the corresponding

notch is higher than those of Echinodiscus and

Amphiope (see Plate 9).

Furthermore, the size of the petalodium (PL) is

significantly lower than that of Amphiope , where as

WA is lower than that of both genera. Even the com-

plicate and dense food grooves branching is peculiar.

Based on these characters this form is attributed to a

new genus, Sculpsitechinus genus novum, The defi-

nition of the corresponding type species is reported

in the following systematics chapter (see Figs. 9a-c).

The oral structure of the sample of "Amphiope"

sp. from Channay-sur-Lathan (Plate 2 Fig. 6)

matches the schemes published by Durham (1955),

Pereira (2010), Smith & Kroh (2011) and Stara &

Borghi (2014). The plate 2b of the interambulacrum

5 is longer and staggered with respect to the 2a; the

lunules are more or less roundish and surrounded

by plates arranged in a radial manner. The size of

the lunules do not correspond to the samples from

Sardinia (Stara & Borghi, 2014). It is clear that this

foim belongs to the genus Amphiope. However it is

left in open nomenclature, Amphiope sp. 3, since

the type species of A. bioculata Des Moulins, 1837

still needs definition.

Both the morphotypes recognised within the

sample of ""Amphiope” pedemontana Airaghi, 1901

have an oral plate structure corresponding to the

plating pattern of Echinodiscus given by Durham

(1955) and Smith & Kroh (2011). Also the axially

elongate lunules and the plate arrangement around

them indicate that they belong to the genus Echin-

odiscus (Figs. 2a-c).

The first morphotype corresponds to the original

description and is herein assigned to E. pedemon-

tanus (Airaghi, 1901). The other form is left in open

nomenclature, Echinodiscus sp. 1, since only two

are available to study and they are poorly pre-

served.

The specimens of ""Amphiope duffy" Gregory,

1911, from Cyrenaica (Libya) (Plate 3 Figs. 1-6),

show a plate structure in the aboral side which is

quite different from the plate patterns of Amphiope

and Echinodiscus. The arrangement of the plates

surrounding the lunules is linear as in Echinodi-

scus but there are 3-4 couples of plates between

the lunules and the corresponding petal tips. It is

noticeable that one of the petals is open distally,

as in E. pedemontanus . Since the oral plating of

these echinoids is not visible, better preserved ma-

terial is needed to clear the systematic position of

this form.

* 15 " ^ ^ pn

| 14 -

13 - — ■ — —

12 -

11 * —

10 J

ABC D £ F

Figure 10. Numbers of post-basicoronal plates comparison

in Amphiope from different geological age. A, B: A. nurag-

ica, Oligo-Miocene, respectively inter. 5 and ambulacrum

III. C, D: Amphiope sp. 2, Late Burdigalian, respectively

inter. 5 and amb. III. E, F: Amphiope sp. 3, Late Serraval-

lian, Tortonian, respectively, inter. 5 and amb. III.

The oral and aboral structures of the large sample

of E. bisperforatus from various localities of the In-

dian Oceans and the Red Sea mach with the plating

schemes given by Durham (1955), Jansen & Mooi

(2011), Smith & Kroh (2011). It seems likely that

different species may be present within the studied

sample (Plate 4 Figs. 1-8; Plate 5 Figs. 1-3), howe-

ver further studies are needed to clear the question.

Samples attributed to ""E. tenuissimus"" L. Agas-

siz, 1847. The first morphotype (Fig. 3a) has both

the oral and aboral plate structure that does not

match those of E. bisperforatus Leske, 1778, and

is closer to "E. auritus ” studied by Stara & M. Fois

Analysis of some astriclypeids (Echinoidea Clypeasteroida)

305

(2014). The plates arrangement of the oral interam-

bulacrum 5 and the number of couples of plates be-

tween the petal tips and the corresponding lunules

is high and matches those of “E. auritus ” (Fig. 9a,

b). Also the particular branching of the food grooves

matches with that of “E. auritus ”. Based on these

observations this morphotype is placed into Sculp-

sitehinus n. gen.

Since the holotype of E. tenuissimus, from Wai-

giou, eastern Indonesia is wanting, a specimen from

Lembeh, North Sulawesi (eastern Indonesia) is pro-

posed as neotype of Sculpsitechinus tenuissimus (L.

Agassiz, 1847) (Plate 11 Figs. 5, 6).

The second morphotype from Andaman Sea of

Thailand, has the plating structures of oral and

aboral faces that match with Echinodiscus, since in

the oral interambulacrum 5 the postbasicoronal

plates 2b and 2a are paired as well as 3b and 3a

(Fig. 3b). Also the axial lunules and the structure

of the plates surrounding them matches with those

of E. bisperforatus.

This morphotype which is a true Echinodiscus

and, given the differences between it and the other

species of this genus, as we shall see in the chapter of

the systematic, is here named E. andamanensis n. sp.

The third morfotype from Indonesia (Borneo),

has the plating structure of the oral interambula-

crum 5 with the two first postbasicoronal plates

staggered as in Amphiope (Fig. 3 c), but it has the

plate arrangement that encircling the lunules as in

Echinodiscus.

It is evident, now, that this form belongs to a

new genus. Therefore, we introduce Paraamphiope

n. g., as it has some similarities with Amphiope.

This morphotype is named Paraamphiope rai-

mondii n. sp., after the collector who donated the

specimen to our museum.

The specimens labeled Echinodiscus truncatus

from Singapore, has a plate structure of the oral inter-

ambulacrum 5 that matches with that of Echinodi-

scus (see Fig. 4), but they differ from other species

of Echinodiscus in many features, that make us con-

sider this a tme E. truncatus (L. Agassiz. 1841).

Also the specimen collected from Hurgada, Red

Sea, shows the plate structure of the oral inter-am-

bulacrum 5 corresponding with that of Echin-

odiscus (Figs. 5a, b). They differs from E.

andamanensis n. sp. and E. truncatus by the posi-

tion of the periproct, that opens more rearmost, be-

tween the plates 2a/3b/3a. This is likely a different

Figure 11. Comparison of lunules size variability in Am-

phiope, Echinodiscus and Sculpsitechinus species. A-B: re-

spectively, LI and L2 variability in A. lovisatoi. D, E:

respectively, LI and L2 variability in “A. bioculata ii in Cot-

treau (1914). G, H: respectively, LI and L2 variability in A.

nuragica. J, K: respectively, LI and L2 variability in E. trun-

catus. M, LI variability in E. bisperforatus. N, O: respecti-

vely, LI and L2 variability in S. auritus. R, S: respectively,

LI and L2 variability in E. andamanensis n. sp. U, V: re-

spectively, LI and L2 variability in S. tenuissimus.

Figure 12. Comparison of differences and variability in

lunules shape utilizing SI. C: Amphiope lovisatoi. F: “A.

bioculata ” in Cottreau (1914). I: A. nuragica. L: Echinodis-

cus truncatus. P: Sculpsitechinus auritus. T: E. andamanen-

sis n. sp. W: S. tenuissimus .

species but, since the sole specimen available to

study is poorly preserved, it is left in open nomen-

clature: Echinodiscus sp. 2.

The examined specimens of " Amphiope "

arcuata Fuchs, 1882, from the ’’Miocene" of

306

Paolo Stara & Luigi Sanciu

Libya, has the oral plate structure on interambu-

lacrum 5 (Fig. 2d) that matches with Paraam-

phiope raimondii n. sp. It differs by P. raimondii

by greater distance between petal tips and lunules

and by longer lunules. Based on these characters

this form is attributed to Paraamphiope genus

novum and assigned to Paraamphiope arcuata

(Fuchs, 1882).

The specimens labeled as Echinodiscus desori

Duncan et Sladen, 1883 are incomplete and the sole

oral face visible is only partially legible. Using the

available data, these echinoids probably belong to

Echinodiscus, by the shape of the lunules and by

the arrangement of the plates surrounding them

(Plate 6 Figs. 1-6). Also the petals, clearly open

distally, connect them to E. pedemontanus.

The sample of “Amphiope bioculata ,, des Mou-

lins described by Cottreau (1914) (Plate 7 Figs.

1-11) likey belongs to the genus Amphiope L.

Agassiz, 1840, by the large petalodium, that in

some specimens gets up to 60% TL, the roundish

lunules with a SI value of about 1.5, and the

distance of the lunules from the tips of the petals

which is very short. However, it is not possible to

attribute these specimens to A. hioculata, since their

plate structure was not reported by Cottreau (1914)

and, on the other hand, the type species of Am-

phiope still needs defining.

The “Lobophora aurita ” illustrated by L. Agas-

siz 1840 (Fig. 6), clearly belongs to the Sculps itech-

inus n. gen., by the plate structures, close to that of

S. auritus (Leske, 1778) and S. tenuissimus (L.

Agassiz, 1847) (see Plate 11 by comparison). The

oral interambulacrum 5 has 4 couples of postbasi-

comal plates, with 2b and 2a partially staggered and

low WA value. There are six couples of plates be-

tween the petal tips and the corresponding notches

and the PL is very small. It differs from the above

mentioned species by the periproct that opens be-

tween plates 3a and 3b.

Since the original specimen is wanting, this spe-

cies is left in open nomenclature: Sculp sit echinus

sp. The species E. formosus, E. yeliuensis, E. cike-

zenensis and E. transiens were based on specimens

with the oral face covered by sediments. Lacking

the important characters of the oral face, such as the

interambulacral plating, a comparison with the type

species E. bisperforatus is unreliable.

For any other consideration see the conclusions

chapter.

DISCUSSION ON MORPHOMETRIC AND

MORPHOLOGICAL ASPECTS

In the following some relevant characters and

morphometric values highlighted by the studies of

Stara & Fois (2014) and Stara & Borghi (2014) are

compared with the results of this study. This can be

useful for further studies to improve the knowledge

of this interesting family of echinoids.

The sample of Amphiope examined by Stara &

Borghi (2014) and in this work, represents a time

span that ranges from the Chattian-Aquitanian to

the Serravallian-Tortonian (about 13-14 Ma).

Furthermore, this sample confirms what has been

observed by Stara & Borghi (2014): in the echi-

noids belonging to this family, during the geologic

time, there was a downward trend with a decreasing

total number of plates. The sample examined in this

study also includes other genera of astriclypeids like

Echinodiscus, which are present from Rupelian to

Recent, Paraamphiope, which runs from the middle

Miocene to the present and Sculp sit echinus that

may have been present in the Miocene and is very

wide spread in the Recent.

Durham (1955) noted that the number of plates

on the oral face is fixed at the end of metamorpho-

sis, whereas some new plates are formed in the abo-

ral face during the early stages of growth (e.g. from

2-3 mm to 10-15 mm TL). No significant variation

in the total number of plates per column was ob-

served by Durham on both oral and aboral faces of

the examined adult individuals (TL = 50 to 62 mm).

The same result emerges also from the available

sample of Sculp sitechinus auritus (former E. auri-

tus) from Mangili, consisting of about thirty speci-

mens with TL ranging from about 70 mm to 150

mm (see Stara & Fois M., 2014). Smith (2005) con-

firmed that the number of plates in adult clypeast-

eroids remains almost unchanged during the stages

of growth in this group of echinoids (see Fig. 10).

In samples of Amphiope, on the other hand, we

can see one particularity: there is a decrease in the

overall plate number as a consequence of the "geo-

logical age”. Kier (1982) noted that there was a

trend within cassiduloids for a decrease in the

number of plates through time, and this may reflect

a general trend towards fewer and earlier formed

plates (from Smith, 2005).

We can deduce that the Amphiope with greater

number of plates is more archaic than that with

Analysis of some astriclypeids (Echinoidea Clypeasteroida)

307

lower number of plates. It would be logical that this

should also be observed in "Echinodiscus" , but it is

not obvious, in part due to the heterogeneity of the

treated samples, which look more like a paraphyletic

group than a monophyletic one, and in part due to

the presence of too many gaps in the fossil records.

On the variability in size and shape of the lu-

nule/notches

From the comparison between the samples

examined, we observed that lunules are variable in

shape and in dimension, and that the greater vari-

ability seems to affect only some species and some

populations.

The sample utilized by Cottreau (1914) seems

to show the largest variability range of the lunules

(variance), which is respectively 49% on LI av-

erage and the 45% on L2 average; the other samples

decidedly show a lower variability, which ranges

between 22% in E. bisperforatus and 41% in

Paraamphiope arcuata.

Moreover, the finding of distorted lunules has

been the normality, as noted by Stara & Borghi

(2014) on over 100 complete specimens of Am-

phiope from Sardinia and many fragments with

lunules belonging to different species and localities,

and this can often make worthless the measures.

Comparing in a graph the size of LI and L2,

detected in a larger sample [40 specimens of A.

lovisatoi (data from Stara & Borghi, 2014); 11 "A.

bioculata" in Cottreau; 25 A. nuragica] however,

we see that the sample of Cottreau's "Amphiope" is

not the more variable, but that the more variable is

A. lovisatoi from Sardinia. The graph (Fig. 11)

shows that, despite the significant variability in the

size of lunules, remain clear the specific differences

(see in particular the difference between "A. biocu-

lata' ' in Cottreau and A. nuragica ).

Using the SI and WI data, in “A. bioculata“ in

Cottreau, SI range from 0.95 to 1.47 (mean 1.22)

(Table 6); in A. nuragica the SI range from 2 to 3

(mean 2.4). As demonstrated (see Table 8), this

system highlights the real differences very well.

Now, if we compare the SI of the various samples

utilized in the first graph (Fig. 11), the specific dif-

ferences between A. lovisatoi, "A. bioculata" and A.

nuragica become very evident, (Fig. 12 C, F and I).

About the samples of the other genera ( Echinodiscus

and Sculpsitechinus ), instead, it is seen that the vari-

Figure 13. Lll comparison, respectively with Sculpsitechi-

nus species (A), Echinodiscus bisperforatus group (B) and

others Echinodiscus (C).

ability of lunules is much lower and the specific dif-

ference is highlighted much more through the mea-

surement of LI and L2 (see Fig. 11, samples J-V).

Finally, the lunules variability exists, but this

does not make difficult to specific distinction.

Indeed, it is demonstrated that the shape of the lunes

(measured with the SI) becomes really distinctive

between species.

On the plates arrangement encircling lunu-

les/notches

The position of the lunules along the ambu-

lacrum has visibly changed during time, but we can

evaluate this change in the oldest species only par-

tially, given that in most of the literature only the

aboral face is shown and is sometimes incomplete.

Now, there are at least two possibilities: dif-

ferent starting point or finishing point of the lunules,

and different number of couples of plates surround-

ing the lunula in the oral and aboral side.

As noted by Stara & Borghi (2014) the number

of couples of plates that surround the lunules can

vary greatly from species to species, and in partic-

ular look different between geologically younger

species from geologically older ones.

In Amphiope from Chattian-Early Aquitanian

from Cuccuru Tuvullao, the couples of plates that

encircle aborally the lunules are 8-9, while in the

oral face are 4-4 (about half); in the specimens

308

Paolo Stara & Luigi Sanciu

Figure 14. B angle comparison, respectively with: S culp-

sitechinus species (A), Echinodiscus andamanensis n. sp.,

E. truncatus, Echinodiscus sp. 1, Echinodiscus sp. 2 (B),

and E. bisperforatus group (C).

Figure 15. Wa comparison, respectively, with: Sculpsitechi-

nus species (A), Echinodiscus and Paraamphiope species

(B), and Echinodiscus bisperforatus group (C).

Figure 16. PL comparison, respectively, with: Amphiope

species (A), Echinodiscus and Paraamphiope species (B),

and Sculp sitechinus species (C).

from Channay-sur-Lathan the situation is 6-6

against 3-4, with an aboral/adoral ratio sharply

decreasing.

In E. pedemontanus the couples of plates on

aboral side ranges among 4-5 and 5-6; in the adoral

side, however, they range from 2-2 to 3-3. It seems

clear that the more archaic characteristics (greater

number of plates) are located in A. nuragica (Co-

maschi-Caria, 1955), so this last one can not

descend from E. pedemontanus , but could derive

from a more archaic ancestor.

In other Echinodiscus fossils, we can observe

the following: in E. cikuzenensis, on aboral side, the

couples of plates are 6-6, the highest number be-

tween the fossils of their genus, but we do not know

the number of the corresponding adoral ones (as-

suming that they are 8, the highest number known,

the total number will be 14 plates per column, very

far from the 16-20 of A. nuragica ).

On the other hand, E. bisperforatus shows more

plates than E. andamanensis n. sp., which has the

lowest number of plates of all (see Table 6).

In all samples of Amphiope, between the lunules

and the corresponding petal tips there are one or

two pairs of plates (not occluded); while on the oral

side the lunules begin constantly from the second

pair of the post-basicoronal plates, (see Plate 9 Figs.

1,2; Plate 10 Figs. 1,2)

In E. pedemontanus there are 3-4 couples of

plates between lunules and the corresponding petals

tip, as in Paraamphiope arcuata.

Summarizing, in other forms of Echinodiscus ,

excluding E. bisperforatus, between the petal tip

and the corresponding lunula/notch there are

two-three pairs of plates (Plate 9 Fig. 4), and

these are arranged in a linear manner, as in E.

bisperforatus. Therefore these characteristics seem

constant and diagnostic and in the future it will be

necessary to take them into account. For more

information see the respective plates and the plate

patterns reported in plates 9, 10.

Migration of the periproct

The migration of the periproct is one of the main

evolutionary processes of irregular echinoids

(Durham, 1955; Kier, 1982); from the apical disc the

periproct migrates towards the peristoma, viz, from

the aboral surface shifts to the oral one. Echinoids

of this family always have the periproct in the oral

Analysis of some astriclypeids (Echinoidea Clypeasteroida)

309

face, between the peristome and the ambitus and it

is clear that the periproct does not change its position

with respect to individual plates after those plates

have started to form, at the end of the metamorpho-

sis, except for occasional cases (Durham, 1955).

But with the passing of geological time, its posi-

tion is not fixed and immutable, nor is the plating

that surrounds it. Thus, the "migration" of the

periproct occurs simultaneously with changes in the

plating and width of the plates, and in the internal

structure, too. We noticed plate pattern modifica-

tions, that we believe to be diagnostic, as well as an

increase or decrease in number, or breadth and shape

changes, of the post-basicoronal interambulacral

plates on inter. 5 and on the ambulacra I and V.

Considering the A. nuragica, Amphiope sp. 2 (in

Stara & Borghi, 2014) and Amphiope sp. 3 series,

which covers a Chatham Aquitanian to Serravallian-

Tortonian time-span, the position of the periproct to

the relative plates seems to be indicative to the

effect of time and evolution; indeed, the distance

from the posterior margin has even decreased (L 1 1

varies from a minimum of 10% of TL in A. nuragica

to 4% TL in Amphiope sp. 3) simultaneously to the

decreasing in size of the last echinoids.

In A. nuragica sample the periproct position

relative to the plates is veiy variable; and this last

one may open both between the plates closest to the

rear edge (3a/4b), both between the most anterior

ones (2a/3b), never between 2b/2a. We also noted

that in this archaic form, the periproct position with

respect to the related plates and their number, ap-

pear very inconstant, unlike the most recent forms.

In the sample of Amphiope sp. 2 the position-

plates ratio is more steady, so the periproct always

opens between plates 2a/3b, as in Amphiope sp. 3.

However, in Cuccuru Tuvullao outcrop one can also

find the morphotype with predominantly transverse

lunules, Amphiope sp. 1 (see Stara & Borghi, 2014),

with the periproct opened along the suture between

the first two plates 2b/2a. This could indicate a con-

vergent evolution of two close species, evolved at

different speeds and in different environments, and

probably found themselves in the same locality only

by accident (in a slightly different times).

This last situation may depend on the well dif-

ferentiated Oligo-Miocene faunas, and on the in-

complete scene of the previous evolutionary steps,

due to the fossil record gaps. Perhaps it will be

Sample

range

mean SI

range

mean

Variance

A m ph lope hiocu la ta .

tn Cottreau. 1914

0.95-1.47

1.22

9+ 12.5

10.20

49.2

45.4

Amphiope nut ugica

2 + 3

2.4

1 1,5+ 15

13.5

Amphiope sp. 2

1.2+ 1.5

1.3

11 + 16

Amphiope sp. 3

1+1.6

1.26

9+10.5

9.8

Echinodiscas

pedemonuimis

0.26+0.54

0.37

9.2+1 1.2

10.4

—

Pa i a amp h io pe arcu at a

0.45+0.76

0.63

8+10.5

9.3

Sad pan echinus sp. 1

—

Sculps it edrimis auritus

—

Echinodiscus

hisperforatus

—

Table 8. Comparison of index and data variation, between Amphiope samples and other astriclypeids genera.

310

Paolo Stara & Luigi Sanciu

possible to answer to this question by furthering

studies on the structure of the samples from

Provence and the Bay of Biscay ones.

In samples of Sculpsitechinus from Mangili and

from Philippines the periproct always opens along

the suture between the post-basicoronal plates

2b/2a, while in the specimen from the Red Sea,

illustrated by L. Agassiz (1840: pi. 14 fig. 2) and

reproduced here in figure 6, it opens in the rearmost

position, between the plates 3b/3a.

In the group of living Echinodiscus andama-

nensis n. sp., E. truncatus and E. bisperforatus, the

periproct opens along the suture between the

plates 2a/2b or at the junction 2a/2b/3b. In any

case, these forms differ from the Sculpsitechinus

“group” in which the periproct-posterior margin

distance and the plates number on the inter. 5 is

higher. In fact, within this group of living echi-

noids, the periproct distance (LI 1) varies from 11

to 25% TL, within a plating with more plates per

column in interambulacrum 5 (2-3 in column “a”

and 4-4 in column “b”).

Plates number and shape on the interambu-

lacrum 5 and periproct position

In plate 1 0 are summarized the results of our ob-

servations about these characteristics. Highlighting

the diversity in the genera there are three platings.

The distance between the periproct from the rear

edge, its position along the perradial suture in the

inter. 5, the shape and the relationship established

between the various plates that form in particular

the inter. 5 and the ambulacra I and V, seem highly

diagnostic at the level of genus and species.

The distance of the periproct from the posterior

margin is a characteristic which is considered to be

very important by ancient authors. As we have

already seen, its position is partially related to the

arrangement, shape and size of the plates of the

inter. 5. However, for the same plating, the distance

may be diagnostic for the species, if it is confirmed

by statistically significant numbers. Here we

simply report what, in general, has been detected

in the small samples which we examined (Fig. 13).

Angle p and WA

Given that rounded lunules can not show

angles with respect to the corresponding petal, the

problem could be solved only studying the mor-

photypes with elongated lunules and in particular

those elongated axially. In particular, the B angle

seem very significant; important data are drawn

by the quantification of this peculiar situation in

tables 9. These data highlight different groups,

corresponding to different species and / or genera,

and in particular highlight Sculpsitechinus (B =

55° to 67°), Echinodiscus and Paraamphiope (B

= 70° to 85°) and E. bisperforatus (B = 105° to

111 °).

A characteristic which, up until now has been

underestimated, is the size at the ambitus of the

various ambulacral and interambulacral sectors. In

particular, the WA at interambulacrum 5 appears to

be very important, seeing it differentiates two of the

genus studied by us: Echinodiscus and Sculp-

sitechinus. Furthermore, the E. bisperforatus group

differentiates itself from the other.

Petalodium

One of the important aspects in these echinoids

is the petalodium length (PL), which can be very

different from group to group. In this comparison

we considered a total of 54 specimens of Am-

phiope, according to the table 10. On the speci-

mens from 1 1 different Sardinian localities (see

also Stara & Borghi, 2014), the PL size ranges

from 47 to 57% TL (mean 52 N42). In the totality

of the specimens from Italy, Spain, France and

Iran, the dimensions range from 45 to 60% TL (see

Plate 8 Fig. 1), with the majority between 48 and

53%. In the sample of Sculps it ehinus auritus from

Mangili in Stara & Fois (2014) PL is 34-45% TL,

as in the sample of S. tenuissimus (29-45% TL)

(see Plate 9 Fig. 8).

In E. bisperforatus PL is very variable, reach-

ing from 40 to 52% TL. In this species, the front

odd petal is always longer than the rear ones,

which are always decidedly shorter. The size of the

remaining " Echinodiscus " is very diversified and

difficult to interpret, given the scarcity of the

material available.

Another interesting feature is the presence of

open petals in different species. In particular in E.

pedemontanus the petals are all open. Some open

petals are visible, however, also in E. desori,

"Amphiope" duffi and, occasionally, even in E.

bisperforatus (Fig. 16).

Analysis of some astriclypeids (Echinoidea Clypeasteroida)

311

Sample

WA- range

B - range

A nip hi ope mtragica

49-53

38 -46

—

A. sp. 2

50-54

37-40

—

A. sp. 3

50-53

29-36

—

A. h to ( ul at a in C ot t re a u, 1914

48-54

—

Sculpsiiechi/tus sp, 1

41 -48

30-38

57°

54 -59°

Sculp sitechi mis tulearei is is

36-47

29-37

55°

48 - 62°

Snip si tech in us aunt us

28-34

51 -59°

Sculp siiee hi nits tenuissimus

30-45

31 -35

67°

65 - 70°

Paraamph tope arena fa

42-46

33 -35

85°

75-100°

Pa/ a an t ph lop e > 'ai mo nil it

80°

Ech i no i Use t ts ties art ( In d ia )

46-48

30-35

71°

68 - 74°

"Am phi ope" Ditffi

65°

Ech i no disc us pedemonta tuts

40-44

35 -37

85°

73 -93°

Ech i nodiscus sp. 1

44-46

69°

70-87°

Echi nodiscus sp. 2

—

80°

E. andamanensis

45.5-53

37-38

75.5°

68 - 85°

E. cikuzenensis

—

73°

E. hisperforatus (Red Sea)

38-47

46-54

105°

102 - 110°

E. hisperforatus (S. Africa)

43 -50

45 - 53

107°

i 02 - 117°

E. h isperfo ra tus (Tanzania)

111°

E.Jelt ttetisis

1 52 ?

114°

E. transiens

—

Table 9. Comparison of PL, WA and B range data in a large sample of astriclypeids. B in degree, other measures in % TL.

312

Paolo Stara & Luigi Sanciu

Amphiope

(Sardinia. Italy)

47-57

52 N42

Amphiope (Alicante,

Spagna)

48-51

49 N2

A mp hi ope (Charm ay-s n r-

Lathan, France)

51-53

53 N10

Amphiope biocuiata in

Cottrcau, 1914

48-55

50 Nil

A mp hi ope sera a in i (France)

A mp hi ope d eydieti ( France)

A mp hi ope b aq uiei (France)

A mp hi ope el lipti ca ( F ran ce)

A mp hi ope h ol la nde i

(Corsica)

Amphiope cf biocuiata

(Iran)

Table 10. Comparison of PL data in a large sample

of Amphiope. Data in % TL.

Variability of the disjunction/contact be-

tween basicovonal and post-basicoronal

plates

Random disjunctions between the basicoronal

interambulacral plates and the related post basi-

coronal ones can be observed in many samples. For

example, Sculpsitechinus tenuissimus from New

Caledonia, S. auritus from the Red Sea and from

Tulear, Madagascar, E. bisperforatus from Eastern

Africa (see Jansen & Mooi, 2011) have high vari-

ability. The problem has already been studied by

Durham (1955), who pointed out that more archaic

scutellids show the basicoronal plates in contact

with the following postbasicoronals, and that the

separation is observable only in the most recent

genera. He also noted that in Dendraster excentricus

(Eschscholtz, 1831) from the Pacific coast of United

States, juvenile individuals shown the basicoronal

plates in full contact with the following ones.

Furthermore, during growth, the second plate of

each ambulacra grew faster than the others until its

separation from the second interambulacral ones, as

indeed is observed in most representatives of the

Astriclypeidae family.

Of all the species studied by Durham where this

variation occurred, Echinarachnius showed the

largest variability. Lohavanjiaya & Swan (1965)

also studied this problem in more detail on some

populations of Echinarachnius parma (Lamarck,

1816) from the coasts of New Hampshire (USA).

These authors noted that there was a wide variabil-

ity in the loss of contact between the basicoronal in-

terambulacral plates and the corresponding post-

basicoronal ones for each column, but it also varied

the amphiplacous or meridoplacous conditions of

the contact, when it was present. It demonstrated

that the variation in the number of plates involved

in the phenomenon followed individuals growth

(size increase), and conceived that the phenomenon

was caused by a selective response to genetically-

induced modifications, at least partially, by different

environmental factors for the different places where

the tested samples lived. As for Durham's observa-

tions, we believe as normal (not diagnostic) the

presence of basicoronal interambulacral plates in

contact with the following post-basicoronal ones.

From the results obtained in particular from

Stara & Fois M. (2014) on the sample of Sculp-

sitechinus auritus (Former Echinodiscus cf. auritus)

from Mangili it is clear that the disjunction between

the basicoronal and post-basicoronal plates in Inter.

5 is constant, but also that there is no constancy in

disjunctions between the corresponding plates in

other interambulacra (see Plate 5 Figs. 3-5).

Moreover, from what emerges from the analysis of

our sample, but especially from the sample (about

100 specimens) observed by Stara & Borghi (2014)

were not basicoronal interambulacral plates in con-

tact joint in Amphiope.

Differences in internal structure

As we have seen in the tested sample, while the

morphology of these two groups of astriclypeids

may be similar, the difference in the internal struc-

tures can be substantial. All groups have a single

central visceral hollow with peripheral walls and

pillars, but the floor reinforcement systems of the

central cavity are profoundly different.

In Sculpsitechinus the floor is supported by a

dense network of thin trabeculae or ribs (see Stara

Analysis of some astriclypeids (Echinoidea Clypeasteroida)

313

& Fois M., 2014: pi. 2 fig. 7; pi. 4 figs. 6, 7),

whereas in Amphiope the floor is supported by a

system of thick masses, with appearance of callos-

ity, modeled in different shapes depending on the

species (see Plate 2 Figs. 2-4). Also the floor of the

central cavity of Parascutella (personal observa-

tion) (but also of Astriclypeus ) seems to have the

same floor support structure that characterize Am-

phiope species.

Instead, the ballast system of all the astriclypeids

is crossed by a different number of cavities more or

less large, and by several micro-channels, which are

distributed differently. A characteristic that does not

appear to have been recognized enough so far is the

size of Aristotle's lantern.

In percentage, the Aristotle's lantern is much

larger in Sculpsitechinus sample from the Philippines

than in the sample from Mangili, Madagascar (see

pi. 6 in Stara & Fois, 2014).

However, some results obtained so far are very

interesting. For example, the size of the Aristotle’s

lantern in Parciamphiope raimondii n. sp. is very

large [27% of TL in a central hollow that measure

46% TL(Plate 19 Fig. 6) if compared to the 15-18%

TL that characterize the Aristotle’s lantern of S.

auritus from Mangili (see Plate 23 Fig. 4) or the

15% TL of the Aristotle’s lantern of Sculpsitechi-

nus tenuissimus from Lembeh.

Other peculiarities

In Amphiope , the food grooves are always sim-

ple (Plate 1 Fig. 7), while they are always more or

less branched in Echinodiscus and largely branched

in Sculpsitechinus (Plate 22 Fig. 4).

INFLUENCE OF PALEOGEOGRAPHY

DURING EOCENE-MIOCENE

Stara & Rizzo (2014), hypothesized that the

Oligocene closure of the pre-Pyrenean corridor

caused a separation (or the exchanges decreasing)

between the North-Western Atlantic faunas and the

Mediterranean ones. To understand the conse-

quences of this, we need to study the evolutionary

course of these faunas, in particular on the basis of

the structural aspects.

From initial observations it appears that already

in the late Rupelian-Early Chattian the scutellids

faunas of the Bay of Biscay were well differenti-

ated. Even the " Amphiope " bearing axial lunules

from Rupelian of Val Bormida had at least two

morphotypes (Stara & Rizzo, 2013; 2014). In Early

Miocene, the Rhone Basin was inhabited by " Am-

phiope " boulei Cottreau, 1914, a particular mor-

photype with small ellipsoidal axial lunules

positioned far from their petals (Plate 14 Fig. 1);

also during the middle Miocene, in Libya a similar

morphotype appeared characterized by smaller and

rounded lunules positioned far from the petals tip

(Plate 14 Fig. 2). At the same time in India, Echin-

odiscus desori lived together E. placenta Duncan

et Sladen, 1883, a form characterized by ellipsoidal

axial lunules far away from the corresponding

petals tip (Plate 14 Fig. 3). During Middle Miocene

in Papua New Guinea lived another similar form,

with long and narrow lunules (comparable with

those of Sculpsitechinus tenuissimus (Plate 14 Fig.

4). Other morphotypes not appear so clear, as the

“ Echino discus" sp. from Miocene of Libya (Plate

14 Fig. 6), which has lunules open posteriorly,

resembling the Recent Sculpsitechinus auritus.

Even Amphiope with rounded or transverse

lunules was already well-differentiated, and wide-

spread: this morphotype is found in the Bay of

Biscay, in the Rhone basin, in central Sardinia and

in the Kabylies.

Stara & Borghi (2014) found two different

species of Amphiope with transverse lunules, both

originating from Cuccuru Tuvullao, Sardinia,

Chattian-Aquitanian in age: Amphiope nuragica,

and Amphiope sp. 1. Not far from this locality

(both from the spatial and temporal point of view),

in the localities of Duidduru, Bruncu Montravigu

Nuraghe Caiu and Tanca Sierra, also a form char-

acterized by rounded lunules (Stara et al., 2012)

was present.

To complete our knowledge of the differentia-

tions occurred between the Biscay faunas and those

of the Proto-Western Mediterranean, it will be

necessary to know the structure of " Amphiope "

agassizi from Middle Oligocene, and A. ovalifora

from the Aquitanian of the Atlantic coast, and

furthermore "A." boulei from the Rhone basin.

Given the wide temporal and the spatial distribution

of Amphiope, as previously described, it is probable

that numerous speciation events occurred even in

different French regions, as occurred in Sardinia.

314

Paolo Stara & Luigi Sanciu

EVOLUTIONARY TRENDS

Now, as proposed by Cottreau (1914), Amphiope

would be descended from some Atlantic-European

"Echinodiscus" , deriving also from E. formosus,

because this last might be geologically the oldest.

But the situation seems more complex and the cur-

rent phylogenetic tree needs to be reviewed.

Obviously, this requires a careful study of cladistics,

and so for now we will only formulate hypotheses

based on observations arising from this work.

Echinodiscus formosus from ?Middle Eocene

and E. yeliuensis from Early Miocene of Taiwan,

already had some features comparable to those of

the living E. bisperforatus (e.g. a similar B angle).

This character and the lack of similar forms in the

Oligo-Miocene peri-Mediterranean basins, suggest

that this morphotype is derived from ancient faunas

of the China Sea. But that does not seem true for

other forms of Echinodiscus.

On the other hand, other common features such

as the lunules shape and their distance from the

corresponding petals, seem to connect " Amphiope "

boulei, " Echinodiscus" placenta, ecc. (see Plate 14)

to the group of Sculpsitechinus.

To clarify the relationship between the four

groups which have emerged from this study (Am-

phiope, Echinodiscus, Paraamphiope and Sculp-

sitechinus, it is necessary to study more the internal

structure of the various fossil forms of the far east

and those that linked the north American faunas to

the European ones.

It seems clear, however, that this trend has led

to the current situation, in which we can see that,

while S. auritus is spread throughout the Indian

Ocean to the islands of the Western Pacific, the

form S. tenuissimus seems confined to the Western

Pacific (see Fig. 17). In these two forms, however,

may also be included various species which only by

new studies, based on more consistent sampling and

analysis of pedicellaria and / or DNA, can be distin-

guished.

Among others, the most widespread form of

Echinodiscus remains E. bisperforatus, while other

forms seem very localized in restricted areas (see E.

andamanensis n. sp. in the Andaman Sea and E.

tmncatus in the Singapore coasts). Even in this case,

new studies, in part based on the analysis of the

structure but also (for the living species) on other

analysis, may better clarify their distribution areas.

For now, in figure 17 you can see the distribu-

tion areas of living forms so far recognized in this

work.

OLD AND NEW PHYLOGENETIC HYPO-

THESES

From the phylogenetic point of view, although

several aspects still remain unclear, today we can

say with reasonable certainty that in the dispute be-

tween Stefanini (1912) and Cottreau (1914) both

had a share of reason. In fact, the thesis supported

by Stefanini (1912) (he thought astriclypeids bear-

ing axial lunules were real Echinodiscus and not

Amphiope) is here confirmed for E. pedemontanus

(former A. pedemontana).

However, as argued by Cottreau, the ancestor of

"A." boulei could also be the "Amphiope" with

small axial lunules positioned far from their petals

tips [such as "Amphiope" sp., from the Libyan

desert (Plate 14 Fig. 2) and such E. placenta from

India (Plate 14 Fig. 3]. We partially agree with him

when he states that the living Sculpsitechinus auri-

tus (former Echinodiscus cf. auritus ) that colonized

the entire Indo-Pacific area, could be derived from

these echinoids. In fact, if we compare morphology

and distance lunules -petals in these echinoids, with

the one detectable in "E. bisperforatus" from Papua

New Guinea (Lindley, 2001) (Plate 14 Fig. 4) and

S. tenuissimus from Lembeh North Sulawesi and

other East Pacific locality, (Plates 21, 22), we will

see that they are relatively overlapping. We do not

agree with Cottreau, however, when we examine

the oldest E. formosus and E. yeliuensis from Tai-

wan. In fact, B angle and distance between lunules

and respective petals tips, seem to suggest them as

being the ancestors of the living E. bisperforatus.

The fact that some features (shape of the plates

in inter. 5 and the periproct position, etc...) accost

them very closely to the E. pedemontanus, let us as-

sume at least two possibilities. First presumes that

already during the Middle Eocene these as-

triclypeids were very diversified and spread along

the shores from the Atlantic to the China Sea;

second, however, one presumes that from a single

common ancestor who lived in the northern basins

of the Atlantic Ocean during the Eocene, two forms

detached. These last ones migrated then in opposite

directions: one towards the inland basins of the

Analysis of some astriclypeids (Echinoidea Clypeasteroida)

315

Figure 17. Distribution of extant main “Echin .odiscus” morphotype, Sculps itechinus and Paraamphiope species. From Mooi

(2014) the GBIF portal; Ashley (personal communication, Dec. 2013); Hattenberger (personal communication, Dec. 2013),

Mooi (1989). Other data by cited authors. Yellow dots: Sculpsitechinus auritus group. Blue dots: S. tenuissimus. Orange

dots: Echinodiscus andamanensis n. sp. Grey dots: E. truncatus. White dots: “ Echinodiscus ” cf. tenuissimus. Green dots:

E. bisperforatus group. Light blue dots: Paraamphiope raimondii n. sp.

Proto-Mediterranean sea and then to the Middle

East; the other one towards the Western Pacific

basins to the north of the Eurasian continent.

Figure 18 shows the phylogenetic hypothesis

emerged from this research.

TAXONOMIC CONCLUSIONS

In the samples we analyzed there are clear inter-

nal structural differences between Echinodiscus and

Amphiope. Important differences can also be ob-

served at the specific level, especially in the mor-

phology of the central hollow floor.

To understand some of undetermined features in

the fossil individuals, the use of living species spec-

imens for the comparison allowed us to solve sev-

eral problems that were unsolved for a long time.

Despite the great variability in shape and

lunules/notches size shown by some groups of as-

triclypeids, the shape of the lunule remains an impor-

tant data for the specific distinction. We have seen that

other characters are also useful to specific and generic

distinction; particularly, we should consider the plat-

ings of the two test faces, and the differences in the

internal structure, where possible. The comparison of

pedicellaria and spines, not always considered in this

work, need further studies, particularly in the living

populations and can help us in the determination of

the variety and/or species, also by molecular examen.

The detection of the test plating allowed com-

parisons based on reliable data; the use of appro-

priate indicators in the statistical comparison, as

operated by Stara & Borghi (2014) can provide,

moreover, a further diagnostic tool.

Among the astriclypeids examined in this work,

the specimens of the Chattian-Aquitanian from

Cuccuru Tuvullao have the highest number of plates

in the inter. 5, the backward position of the periproct

(with respect to the post-basicoronal plates on inter.

5) and also the highest number of couples of plates

surrounded the lunules. These characters and the

massive and strong structure make them apparently

the most archaic of all the taxa included in this com-

316

Paolo Stara & Luigi Sanciu

parison group. It follows that this Amphiope does

not descend from E. formosus, but from an older

common ancestor. This is also true for E. pedemon-

tanus, “ Amphiope ” duffi and E. desori, which can

not be the ancestor of the said Amphiope.

Also the comparison of the internal structures be-

tween Amphiope and Sculps itechinus makes it clear

that these two groups are not as similar as they seem,

but their common origin moves further back in time.

As a result of these observations, it is clear that

all forms of Amphiope bearing round or transverse

lunules, today grouped under the specific name of A.

bioculata, as proposed by Philippe (1998) need to be

revised, since their distinctive characters have not yet

been published up to now or have been underesti-

mated. It is obvious, moreover, that among the species

of Amphiope of Sardinia and those of the group of

“Echinodiscus” there is no direct connection.

In this group of comparison, the knowledge of

the arrangement, number and size of the post-basi-

coronal plates that characterizes the oral face of the

inter. 5 and the aboral one, shows a different subdi-

vision of the genera and a species distinction previ-

ously unrecognized. Referring to the data obtained with

our present work, we can say that the petalodium

size has a diagnostic importance at generic level: it is

small in echinoids of the new group Sculpsitechinus

(30-47%) and wider in Amphiope (45-60%). In any

case, this characteristic must be always used together

with others, since in some groups, such as the E. bi-

sperforatus, it is very variable.

Given the different morphologies and morpho-

metric diversity observed between the samples of

“E. bisperforatus” group here examined (Plate 4

Figs. 1-8), we believe that there is also the basis for

looking for the presence of different species, but this

will be the subject of future research. Jansen & Mooi

(20 11) propose the examen of the pedicellaria of liv-

ing echinoids to differentiate species. In paleontol-

ogy, unfortunately, this possibility is almost always

precluded, since the soft parts or the minute parts

hardly preserve in the sediment. However, careful

observation of the skeletal parts, such as test, inter-

nal structures, Aristotle’s lantern, can partially allow

the distinction between genera and also between

species, acting as a support of the soft parts study.

Despite the lack of available data, we can already

say that the genus of French ' Amphiope " bearing

axial lunules, such as A. agassizi and A. boulei, and of

the East regions, such as E. placenta , from Miocene

of India, are not real Amphiope or Echino discus',

these species should be re-studied and assigned to

different genera. However, the morphological and

morphometric comparison of Miocene astriclypeids

as ‘ Amphiope boulei”, “ Amphiope ” sp. from Libya,

"E. bisperforatus" from Papua New Guinea, with

those that characterize the new genus Sculpsitechi-

nus, allow us to assume that the first ones may be

the ancestors of the latter one and all are detached

from Amphiope and Echinodiscus.

Even the series from ?Eocene to Miocene, Echin-

odiscus formosus-E. yeliuensis and living E. bisper-

foratus could be consistent. In fact, all these echinoids

share some distinctive characters such as the angle B

and the distance between lunules and petals tips,

which places themselves in a close phylogenetic

relationship, and detaches them from both Amphiope,

Sculpsitechinus and Paraamphiope. Strictly speaking,

even the Echinodiscus formosus, E. jeliuensis and E.

bisperforatus series should be moved in a separate

genus, but also in this case it is needed to restudy the

specimens of Taiwan and deeply study also the dif-

ferent forms of living "E. bisperforatus".

It is also evident the diversity of E. transiens

from all other supposed congeners, in particular by

the dimension of the sole visible lunule, and by the

petalodium size, that would fit it between the real

Amphiope. Even in these cases, however, nothing

certain can be defined, until we know the oral face

plating of the specimens in object.

With regards to the astriclypeids present in the

Middle East Miocene, despite the different works

published (among others, see Kier, 1972), the illustra-

tions and platings published are insufficient to deter-

mine with any certainty the belonging to a genus

rather than another. From bibliographical data we be-

lieve it could be Echinodiscus or Paraamphiope, but

only a new study will clarify the real systematic po-

sition of these echinoids. We add only that, according

to a specimen present in NHMUK London, Am-

phiope was also present in the Miocene of Mosul

(Iraq) and in the Gulf of Aqaba (Arabian peninsula).

Finally, although we have observed that the con-

dition of open petals is quite common in the as-

triclypeids and perhaps also in other scutellids of

neighboring families, this important character, men-

tioned by Airaghi (1 899; 1901), joins Echinodiscus

pedemontanus to "Amphiope" duffi and E. desori,

and allows us to reconfirm the existing link be-

tween their regions.

Analysis of some astriclypeids (Echinoidea Clypeasteroida)

317

Figure 18. Hypothesis of phylogenetic relationships between Amphiope, Echinodiscus,

Paraamphiope and Sculp sit echinus genera.

318

Paolo Stara & Luigi Sanciu

All three of these echinoids have some petals

which are open or partially open, an uncommon (or

ignored) character in echinoids of this family.

Indeed, we observed in E. bisperforatus that even

one or more petals are open or tend to be open.

Based on these conclusions, in figure 18 is proposed

a new phylogenetic hypothesis, waiting appropriate

cladistic new studies on the genera and species of

this family.

In conclusion, according to the results high-

lighted, four clusters emerge at the generic level of

species hitherto treated and a new distribution of

living species studied as in figure 17.

1 . Amphiope, including: A. nuragica; Amphiope

sp. 1; Amphiope sp. 2 from Bancali; Amphiope sp.

3 from Channay-sur-Lathan and all other species

from Sardinia treated by Stara & Borghi (2014)

2. Paraamphiope, including “ Amphiope "

arcuata from Libya and “ Echinodiscus tenuis-

simus ” from Indonesia, here renamed Paraam-

phiope arcuata and P raimondii n. sp.

3. Echinodiscus , including: E. bisperforatus ; E.

andamanensis n. sp. and E. truncatus; E.formosus,

E. cikuzenensis, E. jeliuensis, E. desori; finally, E.

pedemontanus and Echinodiscus sp. 1 from Italy

and Echinodiscus sp.2 from Red Sea.

4. Sculpsitechinus, including all the “ Echinodis-

cus cf. auritus"', “ Echinodiscus tenuis simus" of

New Caledonia, Micronesia and some “ E . tenuis-

simus' > '’ from Indonesia; " Echinodiscus bisperfora-

tus " of Papua New Guinea, which would be a new

species. All renamed here as follows: Sculpsitechi-

nus auritus from Mangili; S. tenuissismus, Sculp-

sitechinus sp. 1 and Sculpsitechinus sp. 2.

All other nominal species of “ Amphiope^ and

“ Echinodiscus ” discussed herein and not included

in these four groups will have to be reviewed, given

the few characteristics known at the present.

Finally, a clear zonation of living Sculpsitechi-

nus and Echinodiscus in the Indo-Pacific Seas is

highlighted, as a prelude to further investigations

about the old bibliographic citations on the presence

of " Echinodiscus cf. tenuis simus n in the Oceania

and in the Andaman Sea (see Fig. 17).

In order to facilitate the understanding of the

main characteristics that differentiate the species

and genera treated here, we have summarized the

main differences in Tables 13 and 14.

Speci-

mens

Apx

IVM 82

4.5

JVM 83

1 VM84

4,5

JVM 85

4.5

JVM 86

4.5

1 VM 87

4.5

1 VMKS

4.5

JVM 89

4.5

J VM90

4,5

1VM91

JVM 92

5.5

JVM93

5.5

JVM 94

5.5

1 VM95

JVM 96

JVM97

5.5

JVM 98

5.5

J VM99

5,5

I VM 100

JVM 101

IVM 102

5.5

JVM 103

I VM 1 04

I VM ! 05

IVM 106

5.5

IVM 107

IVM 108

4.5

IVM 109

I VM 1 10

4.5

IVM 1 I 1

4.5

IVM 1 12

mean

5.6

39.8

32.3

55.4

Range

4-6

36-47

29-37

48-62

Table 1 1 . Apx, PL, WA andB data in the sample of Sculp-

sitechinus auritus. B in degree, other data in % TL.

Analysis of some astriclypeids (Echinoidea Clypeasteroida)

319

Table 12 (right).

Specimen

™ 1

1.4

LI 1

Simplified mor-

/: ch i/ifjdfjlEWt iiiuhjftwfifnsLK PMBC

9.1

11 J

5.6

0.49

phometric data of

26346

Echinodiscus ,

Fa hintxbsi ax ait d timei n fits 1 *M 1 2(42

17,6

6.R

0,38

Paraamphiope

EchinattiteruA un dtnnts th'f l.six PM IK’ 264 3

111

5,2

0.43

7j6

and Sculpsitechi-

/; ch triad i sett* undiimiimrjisis PM 1 K 2644

15,5

4.9

0.31

9j6

nus samples from

F ch inadixeus undiimammxui PM I K ’ 2X30

54.6

9.1

14.9

5.2

0.35

4.1

different locali-

ties; TL in mm, fi

EchimMiimLi urtdanwttwisis PM IK’ 2(4 3. 1

5.1

0,39

in degree, other

imJijmiHi 'd.iA 2ML'('-f it' H - i 00 !

6,7

0.44

45.5

measures in %

F. ti fi da mam NL

1X3

4,1

0.30

6.R

TL.

Mean E tmdankincttsis

9.5R

14.3

5.4

0.44

6 A

75.5

MAC.PL 1850

0.33

MAC.1VM 206.

13.5

6.6

0.33

E truncates . S in gmp*irc S 1 3 7a

14,6

4.2

0.28

F'. livneatex, Suifsiporc SI 37

15.3

3.9

0.25

51.5

F ■ truncates, Siii(Sip(src SI

19.6

4.2

0.21

10,8

E. truncates. Sm^iporc S2

17.6

4.2

0.21

Table 13 (down).

Comparison be-

E. intricates. Singapore S3

17.6

3.4

0.19

11.5

Mean E truncates

16.9

3.98

0.22

10.4

tween different

old and new

Si'ulps itcelitwis temassirnus

TH 1

1.4

Li 1

astriclypeid gen-

K Col, in Dollies & Rorruin

121

115

0.32

era: contact bet-

NHMUK. 59,7.1.14

112

0,4

ween interambu-

lacral plate 2b

NHMUK.mi.il 2,25

120

125

3.5

0.28

ANU 6B549

0,28

-■

and the adjacent

MAC IV M 207

0,30

ambulacral plates.

A= amphiplacous;

Lcmbcb 1

0,38

—

M=meridopla-

Lcmbdi 2

0,30

t6.5

cous. B in degree,

N Cnl, I (Hattcmb. cal!)

4,5

0.28

other data in %

TL.

Mean Scutpsitischinus tenuis simus

11.7

4,2

0,28

13,2

32,5

37.5

665

Genus

Main characters o

f inter. 5

other

Asiridypeus

5 ambulacral lu miles; floor of the central hollow

reinforced by massive thickening; highly branched

food grooves

Amphiope

45-61

29-46

4-13

Transverse or rounded posterior lumtles; Moor of the

central hollow reinforced by massive thickening;

simply food grooves

Ptiraumph io pe

42-46

31-38

75-

7100

3,5-12

Axial posterior kinules; Moor of llic central hollow

reinforced by ribs; highly branched posteriorly food

grooves

Echinodiscus

"tenuissimus group’’

40-50

35-38

70-81

5.5-8

Axial posterior lunulcs; floor of the central hollow

reinforced by ribs; simply food grooves; sometime

branched posteriorly food grooves

Echinodiscus

1 "h ispetfi jra ( us

group"

38-50

45-54

100-

117

3.5-12

Axial posterior lunulcs; floor of the central hollow

reinforced by ribs; sometime branched posteriorly

food grooves

Sett Ip.si fee hi nus

30-47

30-33

48-70

11-24

Axial posterior lunulcs; floor of the central hollow

reinforced by itelwork of ribs or trabeculae; highly

branched food grooves

320

Paolo Stara & Luigi Sanciu

Plate 1 . Amphiope sp. 3 from Charmay-sur-Lathan, France (late Serravallian-early Tortonian): external features. Figs. 1-3. Aboral,

adoral and antero (to the left)-posterior (to the right) lateral view of MAC.PL1 823; Fig. 4. Apical disk with (a) madreporite (b)

genital pores; the other pores at the tips of the petals are ocular pores; Fig. 5. Stoma, basicoronals circlet with tuberculation

and food grooves; Fig. 6. Aboral view with (a) undifferentiated tuberculation; Fig. 7. Pattern of very simple food grooves.

Analysis of some astriclypeids (Echinoidea Clypeasteroida)

321

Plate 2. Amphiope sp. 3 from Channay-sur-Lathan, France (late Serravallian-early Tortonian): internal features and plating.

Fig. 1 . radiography of MAC.PL1668; a: central hollow; b: caecum cavity; c: terminal intestine cavity; d: small disarticulated

Aristotle's lantern. Fig. 2. Test fragment showing the internal structures of the central hollow. Fig. 3. Cross antero (to the

right)-posterior (to the left) section of the test; a: central hollow; b: wings of the Aristotle's lantern c: middle conjunction

plan of the reinforcement structures. Fig. 4. Antero-posterior cross-section of the echinid; a: lantern supports; b: section of

ceiling; c: massive floor reinforcement; d: pillars and buttresses of the peripheral reinforcement system. Fig. 5. Plating of

aboral face of MAC.PL1668. Fig. 6. Plating of adoral face of MAC.PL1668.

322

Paolo Stara & Luigi Sanciu

Plate 3. “ Amphiope ” dujfi from Sidi Rof Diasiasia, Cyrenaica, Libya (early Oligocene). Figs. 1, 2. Aboral view of NHMUK:

CY66/E11350 and corresponding aboral plating. Figs. 3, 4. Open posterior right petal and aboral tuberculation of

NHMUK.CY66. Figs. 5, 6. Ambulacrum II with open tip and aboral tuberculation on NHMUK.CY264. In particular, from

Figures 3 and 5 it is noted that in the tip of the petals no plate is occluded.

Analysis of some astriclypeids (Echinoidea Clypeasteroida)

323

Plate 4. Echinodiscus bisperforatus from different localities (Recent). Figs. 1-3. Aboral, adoral and antero (to the left)-pos-

terior (to the right) lateral view of NHMUK.2013.7 from South Africa. Figs. 4, 5. Aboral and antero (to the left)-posterior

(to the right) lateral view of NHMUK.2013.3, from Eritrea. Figs. 6, 7. Aboral and antero (to the left)-posterior (to the right)

lateral view of NHMUK.1957.5.21.3, from Tanzania. Fig. 8. Aboral face of juvenile IVM.206 from north Madagascar.

324

Paolo Stara & Luigi Sanciu

Plate 5. Echinodiscus bisperforatus, platings and peculiarities in specimens from different localities (Recent). Figs. 1, 2.

Plating of aboral and adoral face in two specimen from South Africa. Fig. 3. Plating of aboral face in MAC.IVM.206,

juvenile from Madagascar. Fig. 4. Scheme of food grooves in a specimen from South Africa. Fig. 5. Open anterior odd petal

in NHMUK.2013.7 from South Africa. Fig. 6. Peri-oral tuberculation in specimen NHMUK.2013.7 from South Africa.

Analysis of some astriclypeids (Echinoidea Clypeasteroida)

325

Plate 6. Echinodiscus desori from India (Miocene). Fig. 1. Aboral face of NHMUK.E78128a. Fig. 2. Aboral face of

NHMUK.E78129. Fig. 3. Aboral tuberculation, NHMUK.E78129. Fig. 4. Ambulacrum IV with open tip, NHMUK .E78129.

Fig. 5. Plating of aboral face of NHMUK.E78128a. Fig. 6. Plating of aboral face of NHMUK.E78129. Is noticeable that

these samples have in common with those of Cyrenaica some petals open.

326

Paolo Stara & Luigi Sanciu

Plate 16

L ISvJ II.' I IS . ,KA| IHQtTl

tCMINrUfcS DU NtOGENE M£DiTELRRANLE^

Ajtjphiope

Plate 7. In this fine example of morphological variability of a small portion of Amphiope population, Cottreau (1914) shows

visually what the morphometric data has confirmed. But inadvertently he also highlights that none of these forms can match

with those of other species, such as, for example, A. nuragica. However, looking closely at the lunules, one can also see the

normal deformations and growth differences between the two lunules of the same specimen.

Analysis of some astriclypeids (Echinoidea Clypeasteroida)

327

Plate 8. Examined features colored in reddish; interambulacra shaded gray. Figs. 1,2. Amphiope montezemoloi, arrangement of

plates surrounding lunules in oral and aboral face. Figs. 3, 4. Echinodiscus sp. 2, plate arrangement of interambulacum 5 on

oral and aboral faces; numbering according to Foven’s System. Fig. 5. Sculpsitechinus auritus, plates between notches and petal

tips; measure of ambulacral and interambulacral areas at ambitus. Fig. 6. S. auritus, plates between basicoronals and notches.

328

Paolo Stara & Luigi Sanciu

Plate 9. Comparison of number of plate couples between lunules and petal tips on aboral faces-examined features colored

in reddish, interambulacral columns shaded gray. Fig. 1 .Amphiope nuragica. Fig. 2. Amphiope sp. 2 fromBancali, Sardinia.

Fig. 3. Paraamphiope arcuata. Fig. 4. Echinodiscus sp. 2. Fig. 5. Echinodiscus bisperforatus. Fig. 6. Sculpsitechinus auritus.

Fig. 7. Sculpsitechinus sp. Fig. 8. Sculpsitechinus tenuissimus.

Analysis of some astriclypeids (Echinoidea Clypeasteroida)

329

Plate 10. Comparison of number, shape and disposition of plates on oral interambulacrum 5 - examined features colored in

reddish, other interambulacral columns shaded gray. Fig. 1. Amphiope nuragica. Fig. 2. Amphiope sp. 2 from Bancali,

Sardinia. Fig. 3. Paraamphiope arcuata. Fig. 4. Paraamphiope raimondii. Fig. 5. Echinodiscus sp. 2. Fig. 6. Echinodiscus

bisperforatus . Fig. 7. Sculp sit echinus tenuissimus. Fig. 8. Sculp sit echinus auritus.

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Paolo Stara & Luigi Sanciu

Plate 11. Plating features comparison in Sculpsitechinus species, interambulacral columns shaded gray. Fig. 1, 2. Sculp-

sitechinus auritus. Fig. 3, 4. Sculpsitechinus sp. 1 from the Philippines. Fig. 5, 6. Sculpsitechinus tenuissimus from Lembeh,

Indonesia. Fig. 7. Sculpsitechinus sp. (in Agassiz, 1841). We can observe some common features: large number of plates

between lunules/notches and the petal tips; high number of plates in the oral interambulacrum 5.

Analysis of some astriclypeids (Echinoidea Clypeasteroida)

331

Plate 12. Petals open and petals closed in living and fossils species. Figs. 1. Sculpsistechinus auritus from Mangili: closed

petal tip with occluded plates. Figs. 2. Amphiope nuragica : closed petal tip with occluded plates. Figs. 3, 5, 6. Echinodiscus

pedemontanus : anterior odd petal open;3 and 5 with gradual tip plate downsizing. Fig. 4. “ Echinodiscus ” duffi : posterior

right petal tip open.

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Paolo Stara & Luigi Sanciu

Plate 13. Random contact between basicoronal and relate postbasicoronal interambulacral plates in Echinodiscus and Sculp-

sitechinus. Figs. 1,2. E. bisperforatus (South Africa). 1: interambulacra 2, 3 in contact; 1, 4, 5 disjoint; 2: disjoint. Figs. 3-5.

S. auritus (Mangili): MAC.IVM1 10, interambulacra 1, 2, 3, 4 in contact; 5 disjoint; MAC.IVM1 15 4 in contact; MAC.IVM84,

interambulacra 2, 3 in contact; 3, 4, 5 disjoint. Fig. 6. S. sp. 1 (Philippines) MAC.IVM218: interambulacra all disjoint.

Analysis of some astriclypeids (Echinoidea Clypeasteroida)

333

Plate 14. Other astriclypeids morphotypes. Fig. 1. Aboral face of “Amphiope” boulei (pi. 5, fig. 9). Fig. 2. Aboral face of “A. cf.

bioculata” from Libya, specimen NHMUKE5788. Fig. 3. Aboral face of “Echinodiscus” placenta (Duncan & Sladen, 1883: pi.

52 fig. 1). Fig. 4. Aboral face of Sculpsitechinus sp. 2, in Lindley, 2001 (Fig. 7d). Fig. 5. Adoral face of “Amphiope” sp. from

Libya, Miocene, NHMUK E79772. Fig. 6. Adoral plating pattern of “Amphiope” sp. from Libya, Miocene, NHMUK E79772.

334

Paolo Stara & Luigi Sanciu

Species

posit.

Lunules

or notches

Space (1,3) and plates between

limulesand (a) petals or (b)

basi coronals

other

Pa ramp hi ope a ana to

2a/3b

11.5

3-4

Lunules ellipsoidal in shape

Pa ra mphi op e ra imondii

2aOb

1-2

2-3

Lunules slit-like

Ech mod iscus pedemontaims

2a/3b/3

3h/3a

3-4

Petals open

Ech in o d iscus andaman easts

2 b/ 2a

5.4

2-4

Petals dosed: 13 = 75,5°;

WA = 38% TL

Ech in od. iscus I ru ncafes

2b/2a/3

Petals closed; mean S =67°;

WA = 38% TL

Ech in od iscus h isperfot a tus

2b/2a

2h'2a/3

30-

1-2

Virtually closed petals; long slit-

like lunules

mean (J - 100°; WA = 48-50 %TL

Sculpsitechinus sp.

(in L. Agassiz )

3b/3a

6-6

3-4

Posterior notches

Sculps it ech inns nil ea reus is

2a/2b

3-3

Posterior notches

Sculps it a h imts t en uissi mus

2a/2b

4-6

2-3

posterior lunu les ; L 1 1 = 1 1 -26;

13^66,5°; WA-32,5 TL

Table 14. Comparison between different species here studied. Data LI -3 in % TL.

III. SYSTEMATICS

In this chapter we will discuss the species and

taxonomic groups that have been modified or pro-

posed as a consequence of our observations. Others,

such as Amphiope and Astriclypeus remain unal-

tered and are not considered / modified by us.

Family Astriclypeidae include the genera: Astri-

clypeus Verrill, 1867 ; Amphiope L. Agassiz, 1840;

Echinodiscus Leske, 1778; Paraamphiope genus

novum; Sculpsitechinus n. g.

Class ECHINOIDEA Leske, 1778

Subclass EUECHINOIDEA Bronn, 1860

IRREGULARIA Latreille, 1825

MICROSTOMATA Smith, 1984

NEOGNATHOSTOMATA Smith, 1981

Order CLYPEASTEROIDA L. Agassiz, 1835

Suborder SCUTELLINA Haekel, 1896

Infraorder SCUTELLIFORMES Haekel, 1896

Superfamily SCUTEELIDEA Gray, 1825

Family ASTRICLYPEIDAE Stefanini, 1912

Main characters of the family ASTRICLYPEIDAE

1 . Main visceral central hollow, with floor rein-

forced by a network of thin trabeculae or by solid

calcitic masses in apparently calloused form; pe-

ripheral buttressing developed as dense honey-

combed meshwork of cellular structure;

2. Central or sub-central apical system with 4

gonopores;

3. Width of ambulacral and interambulacral

zone at ambitus depends on the species or genera

4. Petals well developed and closed or nearly

closed distally, sometimes open;

5. Small basicoronal circlet, with the interam-

bulacral elements usually pointed, but not strongly

Analysis of some astriclypeids (Echinoidea Clypeasteroida)

335

projected distally; all interambulacra disjointed or

virtually disjointed on the oral surface;

6. Posterior interambulacral area always dis-

jointed adorally and separated by enlarged first pair

of post-basicoronal ambulacral plates;

7. Periproct oral open, along the perradial suture

of the post-basicoronal plates in inter. 5;

8. Perradial lunules or notches in some or all

ambulacra;

9. Food grooves well developed, bifurcated at the

edge of the basicoronal circlet and branched distally.

From Smith & Kroh (2011, accessed September

2013), as emended.

Genus Echinodiscus Leske, 1778

=Echinoglycus Leske, 1778, p. 197 (nomen nudum)

= Lobophora Agassiz, 1841, p. 64, not Curtis, 1825

in Smith & Kroh (2011).

=Tretodiscus Pomel, 1883, p. 71 (objective)

=Tetrodiscus Lambert & Thiery, 1921, p. 323

(nomen vanum) from Smith & Kroh (2011).

Species included

E. formosus Yoshiwara, 1901, Middle ?Eocene,

Miocene, Taiwan

E. pedemontanus (Airaghi, 1899), Rupelian, Italy

E. chikuzenensis Nagao, 1928, Oligo-Miocene,

Japan

E. yeliuensis Wang, 1982, Early Miocene, Taiwan

E. bisperforatus Leske, 1778, Pleistocene-Recent,

Indo-Pacific

E. andamanensis n. sp., Recent, Indo-Pacific;

E. truncatus (L. Agassiz, 1841), Recent, Indo-

Pacific

E. desori Duncan et Sladen, 1883, Miocene, India

Echinodiscus sp. 1, Rupelian, Italy

Echinodiscus sp. 2, Pleistocene-Holocene, Egypt

Other species attributed to this genus, that need

to be revised

Echinodiscus placenta Duncan et Sladen, 1883,

Miocene, India

Echinodiscus ellipticus Duncan et Sladen, 1883,

Miocene, India

Echinodiscus ginauensis Clegg, 1933, Saudi Arabia

and the Persian Gulf

Diagnostic features

1 . Test sometimes slightly indented laterally in

ambulacra II and IV; thin and sharp margin;

2. Main visceral central hollow with floor rein-

forced by a network of thin trabeculae;

3. Petals sometimes open; the posterior pair

shorter than the rest, the anterior odd sometime

being the longest;

4. Posterior ambulacra with axial ellipsoidal

lunules, long slit-like lunules or notches;

5. Periproct open next to the rear margin on

inter. 5;

6. Food grooves branched distally;

7. Angle between the lunules from 70 to 110°;

8. Width at ambitus of interambulacrum 5 from

36 to 53% TL;

9. Tube-feet extending into interambulacral zones;

10. Post-basicoronal plates 2a/2b, 3a/3b on inter.

5 large and paired, forming an obtuse triangle;

1 1 . Only 2-4 plates present between the lunules

and the tips of respective petals.

From Smith & Kroh (2011 , accessed September

2013), as emended.

Echinodiscus is distinguishable from the other

genera, by the first two couple of post-basicoronal

plates in inter. 5 large and paired, whereas in Am-

phiope and in Paraamphiope n. g. they are staggered,

with the first one elongated and in Sculps itechinus

they are smaller and only partially paired; further-

more, the contact of the first post-basicoronal plates

in inter. 5 and the related ambulacral is meridopla-

cous in Echinodiscus , while in Amphiope and

Paraamphiope these is amphiplacous. Echinodiscus

is distinguished from Sculpsitechinus as having only

2^4 couples of plates between the lunules and the tips

of respective petals, instead of 3-6, and by the

periproct position, which is very close to the pos-

terior margin (2,5-13% TL) while it is more distant

in Sculpsitechinus (11-26% TL).

Echinodiscus andamanensis n. sp.

Plate 15 Figs. 1-5, Table 12.

Synonymy. The synonymy includes only the

citations that certainty belong to this species.

1971, Echinodiscus tenuissimus L. Agassiz,

1847, Clark A.M. & Rowe F.W.E, pp. 144-145

336

Paolo Stara & Luigi Sanciu

1987, Echinodiscus tenuissimus L. Agassiz, 1847,

Bussarawit S. & Hansen B. (n.v.)

1991, Echinodiscus tenuissimus L. Agassiz, 1847,

WarenA. & Crossland M.R., p. 106

2004, Echinodiscus tenuissimus L. Agassiz, 1847.

Putchakam S. & Sonchaeng P., p. 424, pi. 1

2005, Echinodiscus tenuissimus L. Agassiz, 1847.

Van der Steld b., Electronic Web Publ., accessed

sept. 2013

Examined material. Holotype: specimen from

Pak Meng Beach, Trang Province, Thailand, inven-

tory n° PMBC 26346.1 TL 81 mm. Other speci-

mens from Andaman coast of Thailand housed in

the PMBC, Phuket, Thailand: PMBC.2842, TL =

66 mm, from Pak Meng Beach; PMBC. 2843, TL =

66.2 mm and PMBC. 2830, TL = 54.6 mm, from

Noparat Tara Beach, Krabi Province; PMBC. 2 844,

TL = 66.2 mm, from PMBC Jetty-South, Phuket

Province. The series fromPak Meng Beachinc ludes

5 specimens, inventory numbers 2842.1-5, TL

65.8-79.2 mm. 1 specimen from West side of Ko

Yao Yai, Phuket, housed in the NHMD.Z n°

ZMUC-ECH- 1001, TL 37 mm (See also Waren &

Crossland, 1991 figs. 10a, c); 1 specimen from

“Thailand”, Recent (based on a illustration in

“www. Echinoids NL”). In the latest specimen the

TL is unknown, but the platings are well legible.

The PMBC material was collected by S, Bussarawit

and C. Nielsen, in 1975-1980, on sandy mud, at low

tide and (PMBC jetty- South) on reef flat, sand.

Description. Small size, depressed test. Ambitus

outline sub-rounded (TW ~ 105 -s- 110% TL). Oral

surface flat, peristome sub central. The inter. 5 has 2

post-basicoronal plates per column, the first two

large and triangular, the second one larger, forming

the complex a broad-based triangle; the width at the

ambitus is ~ 38% TL. The periproct is veiy close to

the rear edge (Lll = 6.6 % TL) and small (2-^3%

TL), and it opens along the suture between the first

two post-basicoronal 2b/2a plates or between

2b/2a/3a, in the samples examined (Plate 15 Figs. 1-

5). The peristome is round and large size (almost 5%

of the TL); the basicoronal ambulacral circlet is

small (LI 3 = 10% TL). The petals are closed, the odd

petal is longer than the other; petalodium size 49 %

of TL. The lunules are short and axial (LI = 14 %

TL), narrow (L2 = 5.4 % TL) and with a B of 75.5°.

Each lunule is separated from the corresponding

petal tip by 2-3 couples of plates and surrounded by

3-5 couples of plates per column on the aboral face,

and by 3-4 on the oral one. The apical disc is star-

shaped and small in size (~ 8-10% TL). The internal

structure and the size of the Aristotle's lantern were

not detected. However, the complete plating was de-

tected (Plate 15 Figs. 3, 4). The number of plates per

column is shown in tab. 6. The food grooves are sim-

ple (Plate 15 Fig. 5); the main food grooves run

through the center of each ambulacral column, start-

ing from long stretches parallel to the ambitus. The

distribution of tuberculation is linked to the shape

and distribution of the food grooves. Large tubercles

can be found in the basicoronal interambulacral

plates and along the sutures that lead to the post

basicoronal plates. Large tubercles also cover the pe-

riphery of the post basicoronal interambulacral plates,

moving up the ambitus where the tubercles are

smaller. Medium sized tubercles also cover a band

along the perradial ambulacra sutures and close to

the lunules toward the stoma, and the periproct; the

tuberculation is apparently absent along the main

food grooves. On the aboral face the tuberculation

is undifferentiated, evenly distributed, dense and

small, all over the surface in all the specimens.

Etimology. From Andamane coasts of Thai-

land, locus typicus.

Distribution. Recent, Thailand coast of An-

damane Sea, Noparat Tara Beach, Krabi Province;

Pak Meng Beach, Trang Province; PMBC jetty-

South, Phuket Province. The type locality is Trak

Meng Beach, Trang Province, Thailand 7°29’57.69”

N, 98°49 , 08.51” E, on sandy mud, low tide.

Comparative notes. E. andamanensis n. sp.

differs from E. pedemontanus in that all of its petal

are closed, the periproct is rounded in shape, instead

of drop-shaped, and in that the periproct opens be-

tween plates 2a/2b, against 3b/3a, in oral interam-

bulacrum 5; moreover, E. andamanensis n. sp.

differs from E. bisperforatus due to the shape of the

lunules, that are longer and twisted in the last one

and due to the B angle that is 75.5° against 110°. E.

andamanensis n. sp. have the WA at inter. 5 equal

to 38% TL against 50% of E. bisperforatus. E. an-

damanensis n. sp. differs from Echinodiscus sp. 1

by the size of the stoma that is large (=>4 % TL)

while in Echinodiscus sp. 1 is very small (<2.5%

TL) and by the position of the periproct, which lies

between 2a/2b in inter. 5, instead between 3a/3b.

Furthermore, E. andamanensis has 5-7 aboral

Analysis of some astriclypeids (Echinoidea Clypeasteroida)

337

Plate 15. Echinodiscus andamanensis n. sp., Recent, Andaman coasts of Thailand. Figs. 1, 2. Aboral and adoral face of

holotype PMBC.26346, from Pak Meng Beach. Figs. 3, 4. Aboral and oral plating pattern of specimen ZMUC-ECH-1001.

Fig. 5. Food grooves scheme of specimen in Van Der Bas. Fig. 6. Oral face of PMBC.2643.1, from Noparat Tara Beach, in

which are well visible the long coronal spines

338

Paolo Stara & Luigi Sanciu

couples of plates in the ambulacra I and V against

9-9. E. andamanensis n. sp. differs from E. trunca-

tus in having the stoma much wider, spines much

denser and longer (4.9 % TL against 3% TL) and

simpler food grooves. Finally, E. andamanensis dif-

fers from Echino discus sp. 2 in having the periproct

that opens between plates 2a/2b instead 2a/3b/3a.

Echinodiscus pedemontanus (Airaghi, 1899; pi.

XXII, IV, fig. 4)

Plate 16 Figs. 1-8; Figs. 2a, b; Tables 3, 6, 8, 9

1899, Amphiope pedemontana Airaghi, p. 17, pi.

VI, fig. 4a, b.

1901, Amphiope pedemontana Airaghi, p. 188, pi.

XXII (IV), fig. 4.

Type specimens. The whole type-series, located

at the Natural History Museum of Milan, was lost

in the bombing during the last World War. The sam-

ples we studied are housed at the Museo di Storia

Naturale “G. Doria” of Genoa (MSNDG) and one

at the MAC, Cagliari. The sample inventoried with

the number MSNDG. 12 18 is assigned as Neotype.

Examined material. Three specimens:

MSNDG. 25 from Pareto, MSNDG.1214 from

Cairo Montenotte and MSNDG. 12 18 lacking indi-

cation of the locality; one specimen MAC.PL2014

from Merana (Alessandria Province). Illustrations

of the samples described by Airaghi (1899 and

1901) were also examined.

Emended diagnosis. Species of medium-small

size, depressed lateral profile, narrow and elongated

axially lunules on the posterior ambulacra. Frontal

odd petal slightly longer than the others and always

open. In the oral face on inter. 5 there are only two

pairs of post-basicoronal plates, with the first two

large and paired.

Description. Small-medium sized, with more

or less axially elongated lunules on posterior am-

bulacra. Depressed test (mean TH = 12% TL) with

the highest point anterior to the apical disc. The

margin is thin and uniform; ambitus rounded in out-

line and wider posteriorly. The frontal odd petal is

slightly longer than the others and is open or almost

open; the other one tends to be open, and the two

rear ones are the shortest. Interporiferal and poriferal

areas raised; sometime the poriferal zone is slightly

sunken, with the first ones 1 to 1.5 times larger than

the others. The lunules are small (mean WI = 10.4),

more elongated along the axis of the posterior am-

bulacra and narrow (mean SI = 0.37).

Only two post-basicoronal plates are present

in each column on the oral inter. 5, with the first

two plates being large and paired. The WA of

inter. 5 at ambitus is, on average, 35% TL; on

MAC.PL2014, the only one not deformed; B is

93°. The periproct is small, elongated and drop-

shaped (wide 2.2% and long 3.5% of TL), close

to the posterior test margin and open between

plates 2a/3b/3a or 3b/3a. Internal structure and

other features as for the genus.

Distribution. Type locality and horizon. Val

Bormida, Liguria and Piedmont. Molare Formation,

Rupelian. Occurrence in Italy: Dego, Mioglia,

Pareto, Squaneto, Santa Giustina, Giusvalla, Cairo

Montenotte, Merana.

Comparative notes. E. pedemontanus differs

from E. bisperforatus, E. andamanensis and E.

truncatus in the shape of the front odd petal, which

is always open, and the periproct position that opens

more posteriorly, between the second two postba-

sicoronal plates; it also differ from Echinodiscus sp.

2 from Hurgada (Egypt) in the petals shape. E. pede-

montanus differ from E. bisperforatus in the shape

and length of the lunules; on the B angle that is 86°

against 110°. Finally, E. pedemontanus have sub-

equal petals and simpler food grooves, while E. bis-

perforatus have the front odd petal longer and the

posterior petals always much shorter than the others

and much complex food grooves.

Echinodiscus sp. 1

Plate 17 Figs. 1-6; Fig. 2c; Tables 3, 6, 8, 9

Examined material. Two specimens: UNIGE.

SM-VI-P-(5)-DN and UNIGE. SM-VI-DR and two

large fragments: UNIGE.SM-DS and UNIGE.SM-

VI-VI-DP.

Diagnosis. Small-medium sized species, with a

depressed lateral profile and axially elongated

lunules in the posterior ambulacra. Petals sub-equal,

large and closed, the frontal one a little longer than

the others. In the oral face on the inter. 5, there are

only two post-basicoronal plates per column, large

and paired. In the rear margin there is a clear notch

that arrives close to the periproct.

Analysis of some astriclypeids (Echinoidea Clypeasteroida)

339

Plate 16. Echinodiscus pedemontanus from Liguria and Piedmont, Italy (Oligocene). Figs. 1-3. Aboral, adoral and antero

(to the left)-posterior (to the right) lateral view of MSNDG 1218. Fig. 4. Antero (to the left)-posterior (to the right) lateral

view of MAC.PL2014. Figs. 5, 6. Aboral and adoral view of MAC.PL2014. Fig. 7. Apical disc close-up of MAC.PL2014.

Fig. 8. Magnification of open frontal odd petal in MSNDG 1218.

340

Paolo Stara & Luigi Sanciu

Description. Small-medium sized, with more

or less axially elongated lunules in posterior am-

bulacra. Depressed test (TH = 10.5% TL) with the

highest point anterior to the apical disc. Thin

margin, anteriorly a little thicker than the rear. The

ambulacra have sub-equal petals, closed, with

poriferous and interporiferous zone similar in

width. Small and narrow lunules, elongated along

the axis of the posterior ambulacra. There are only

two post-basicoronal plates per column in inter. 5 ;

the first two are large and paired. The periproct is

small (width = 2.5% of the TL) and round; close to

the posterior margin and open along the suture be-

tween 3b-3a. The internal structure and other fea-

tures as for the genus.

Distribution. Locality and horizon: Val Bormida,

Liguria and Piedmont, Molare Formation, Ru-

pelian. Occurrence: Pareto e Santa Giustina (Lig-

uria) Italy.

Comparative notes. Echinodiscus sp. 1 differs

from E. pedemontanus in having all the petal closed,

by the periproct shape, sub-rounded instead of drop-

shaped, and by the characteristic indentation on the

posterior margin of the interambulacrum 5, absent

in all other known Echinodiscus. Echinodiscus sp.

1 differs from E. bisperforatus in shape, length and

angle of the lunules. In Echinodiscus sp. 1 B is 93°

against 1 1 0° of E. bisperforatus and the WA at inter.

5 is only 35%, against 50% TL. Echinodiscus sp. 1

differs from E. andamanensis because in the last one

the stoma is very large (> 5% of the TL) and by the

position of the periproct, which lies between the

plates 2a/2b in the inter. 5. Furthermore, E. andama-

nensis has 5-7 aboral couples of plates in the ambu-

lacra I and V against 9-9. Echinodiscus sp. 1 differs

from E. truncatus in having the periproct that open

between the plates 3b-3a, while in E. truncatus it

opens between plates 2b/2a/3b.

Remarks. The specimen is inventoried as

UNIGE.SM VI (P5) DN, and consists of a small-

medium sized specimen (TL = 76 mm, TW = 104%

TL, TFI 8 mm), with both faces visible.

Echinodiscus sp. 2

Platel8 Figs. 1-3, 6; Plate 8 Figs. 3, 4; Tables 9, 12

Examined material. 1 specimen, MAC.PL

1850, TL = 21 mm.

Description. Small size echinoid, very flat test

and thin ambitus, with rounded to sub-rounded out-

line. In the inter. 5 there are two plates per column,

paired and wide. The B angle is 80°, the axial

lunules are narrow; the periproct opens between

plates 2a/3a/3b. The anterior odd petal are the long-

est and the two posterior pair are shorter. Internal

structure not detected.

Distribution. Locality and horizon: Pleistocene-

Holocene from Hurghada, Red Sea, Egypt.

Remarks. Echinodiscus sp. 2 differs from E.

pedemontanus in that all of its petal are closed, the

periproct is rounded in shape, instead of drop-shaped;

Echinodiscus sp. 2 differs from E. bisperforatus by

the shape and size of the lunules, that are longer and

twisted in the last one and due to the B angle that is

80° against 110°. Echinodiscus sp. 2 differ from

Echinodiscus sp. 1 by the size of the stoma that is

very large (> 5% TL) while in Echinodiscus sp. 1

is very small (<2.5% TL) and by the position of the

periproct, which lies between 2a/2b on oral interam-

bulacrum 5, instead between 3a/3b. Echinodiscus

sp. 2 differs from E. truncatus in having the stoma

much wider and simpler food grooves.

Echinodiscus truncatus (L. Agassiz, 1841)

p. 66; pi. 11, figs. 11-16

Platel8 Figs. 4-6; Figs. 4a, b, Figs. 3, 4; Tables 9, 12

1841, Lobophora truncata L. Agassiz, pp. 66-67,

pi., 11, fig, 11-16

1855, Echinodiscus truncata Gray, p. 20 (n.v),

1872, Echinodiscus truncatus Gray, Gray, p. 122 (n.v)

1921, Amphiope ( Tetrodiscus ) laevis Klein (Mel-

lita), Lambert J. & Thiery R, p. 323

1948, Echinodiscus bisperforatus truncatus (L.

Agassiz), Mortensen, pp. 410-411, pi. 71, figs.,

6, 18

1914, Echinodiscus bisperforatus var. truncatus

Clark H. L., p. 42

1925, Echinodiscus bisperforatus var. truncata (L.

Agassiz, 1841) Clark H.L., p, 171

1981, Echinodiscus bisperforatus truncatus (L.

Agassiz, 1841) Dollfus R. & Roman J., p. 102,

data (n.v. 1855-1872 taken from Kroh, A., 2012)

Examined material. 2 specimens from Changi

East coast, Singapore, in the Fantin collection: 137,

Analysis of some astriclypeids (Echinoidea Clypeasteroida)

341

Plate 17. Echinodiscus sp. 1 from Liguria Italy (Oligocene). Figs. 1-3. Aboral, adoral and antero (to the left)-posterior (to

the right) lateral view of UNIGE.SM VI (P5) DN. Figs. 4, 5. Aboral and adoral plate pattern of UNIGE.SM VI (P5). Fig. 6.

Aboral view of UNIGE.SM- VI-DR. Figs. 2, 5. Despite the large deformations and distortions of lunules and shell, the

generic characters remain intact and legible in the plate pattern of oral interambulacrum 5.

342

Paolo Stara & Luigi Sanciu

137A, TL = 51 and 60 mm (Plate 18 Figs. 4-6);

three specimens from Kampong Pasir Ris, North

East, Singapore; based on pictures from Ria Tan

(web site www.wildsingapore.com, 2014); named

S.l-3, TL unknown, but complete of spines.

Description. Small-medium size echinoid, flat

test and thin ambitus with sub-rounded outline

truncated at the posterior edge. The apical disc is

eccentric forward (L4 = 57% TL; L 13 small (mean

= 14% TL). In the inter. 5 there are two plates per

column, paired and wide. B is 67° on average; the

WA vary from 37 to 40% TL. The lunules are axial

and slit - like in shape; SI vary from 0.19 to 0.28

(mean = 0.22) and WI is 8.68 on average. The

periproct is small and opens between plates

2b/2a/3b,with LI 1 on average 10% TL. The petals

are sub-equal in size and PL is about 50% TL long.

The food grooves are finely branched in all the am-

bulacra (Plate 4 Fig. 4). The primary spines are

short (about 3% TL) and sparse; the tubercolation

is visible in Plate 18 Figs. 3, 5.

Distribution. Locality and horizon: Recent,

Singapore.

Comparative notes. E. truncatus differs from

E. andamanensis n. sp. in having the stoma smaller,

spines much sparse and shorter (3% TL against 4.9

% TL) and much branched food grooves; E. trun-

catus differs from E. pedemontanus in that all of its

petal are close and the periproct is rounded in shape,

instead of drop-shaped; E. truncatus differs from E.

bisperforatus by the shape of the lunules, that are

longer and twisted in the last one and due to the B

angle that is about 67° against 110°; E. truncatus

differs from Echinodiscus sp. 1 by the size of the

stoma that is very small (<2.5% TL) while in Echin-

odiscus sp. 1 is very large (> 5% TL) and by the

position of the periproct, which lies between

2b/2a/3b in oral interambulacrum 5, instead be-

tween 3a/3b. E. truncatus differs from Echinodiscus

sp. 2 in having the periproct that opens between

plates 2b/2a/3b instead 2a/3b/3a.

Remarks. Agassiz L. (1841: 66), named these

species Lobophora truncata because the ambital out-

line truncated at the posterior margin; among other

features this species showed well food grooves, more

branched than in E. bisperforatus. In addition, the

lunules are shorter and a bit larger than in E. bisper-

foratus (formerly Lobophora bifora). The specimen

described by L. Agassiz was deposited at the “Mus-

eum of Paris” but where it came from is unknown.

Clark H.L. (1914) cites seven specimens from

New Caledonia and two from Penang (Malaysia),

but we believe that the New Caledonia's specimens

belong to E. tenuissimus group, In fact, the infor-

mation in our possession, says that in New Caledo-

nia there are not E. bispeiforatus and E. truncatus,

but only echinoids belonging from the Sculpsitehi-

nus tenuissimus group (formerly E. tenuissimus ),

As distinctive features, Clark H.L. (1914) mentions

short lunules and short petals.

The same author (p. 171) confirms that he has

observed several specimens from New Caledonia

and from Penang (Malaysia), but he “doubts” that

these correspond to “E. tenuissimus ”, and says that

these specimens "would look like" to E. truncatus.

Dollfus & Roman (1981) consider E. truncatus

a variety of E. bisperforatus and cite (p. 102) Sin-

gapore as locality where the described specimens

came from. All this shows how much uncertainty

there is in the recognition of certain species without

the use of structural characteristics.

Paraamphiope n. gen.

Type species. Paraamphiope raimondii n. sp.;

the holotype is housed in the Department of Animal

Biology and Ecology, University of Cagliari

(UNICA).

Description. Diagnostic features:

1 . Sub pentagonal visceral hollow width almost

47% TL;

2. Main visceral central hollow with wall rein-

forced by a network of thin trabeculae;

3. Petalodium small in size (from 42 to 47%

TL); petals well developed and almost closed

distally;

4. Posterior axial ambulacral lunules ellipsoidal

or narrow slits;

5. 3 to 4 pairs of plates between petals and

lunules;

6. Periproct open less than 13% TL from the

posterior margin;

7. The first two plates in inter. 5 must be stag-

gered with the 2b in amphiplacous contact with the

post basicoronal ambulacral plates;

8. Food grooves very branched distally near the

rear edge;

Analysis of some astriclypeids (Echinoidea Clypeasteroida)

343

Plate 18. Fossils and living Echinodiscus species. Figs. 1,2. Echinodiscus sp. 2, aboral and antero (to the left)-posterior (to the

right) lateral view ofMAC.PL1850, from Flurgada, Egypt. Fig. 3. Echinodiscus truncatus , aboral face of malformed individual

(specimen 137A from Fantin collection), Recent, Singapore. Fig. 4. Food groovers scheme of E. truncatus from Singapore.

Figs. 5, 6. E. truncatus, oral and aboral view of specimen 137, from Fantin collection, Recent, Singapore.

344

Paolo Stara & Luigi Sanciu

9. width at ambitus at inter. 5 zone, measuring

almost 36% TL;

10. B angle about 88°;

1 1 . Tube-feet extending into interambulacral zones.

Etimology. Para = affinity with the relate genus

Amphiope

Distribution. From Libya and Indonesia, Miocene

to Recent.

Remarks. Paraamphiope n. g. differs from Echin-

odiscus in having the first two post-basicoronal

plates of inter. 5 which are staggered whereas they

are always large and paired in the second; moreover,

in Paraamphiope n. g. the contact by post-basicoro-

nal ambulacral plates in inter. 5 is amphiplacous, as

in Amphiope , while this is meridoplacous in Echin-

odiscus. Paraamphiope n. g. differs from Amphiope

in that has axial lunules separated by 3 to 4 couples

of plates from the respective posterior petals, in the

latter they are rounded or transverse and separated

from respective petals tip by only 1 -2 couples of

plates; Paraamphiope n. g. has very branched food

grooves in the posterior part of the test, in Amphiope

they are veiy simple and in Sculpsitechinus these are

highly branched and developed on the entire adoral

surface. Paraamphiope have a petalodium long 42-

46% TL, against 45-60% of Amphiope and 30-45%

TL of Sculpsitechinus. Paraamphiope n. g. differs

from Sculpsitechinus by the position of the periproct

that is close to the rear margin (2.5-13% TL) against

11-26% TL.

This genus includes the following species:

P. raimondii n. sp., Recent, Indonesia (Borneo)

P. arcuata (Fuchs, 1882), Miocene, Egypt and

Libya

Paraamphiope raimondii n. sp.

Plate 19 Figs. 1-7; Tables 9, 12

Examined material. Holotype, MAC.IVM

206, TL 53 mm housed in the Department of

Animal Biology and Ecology (UNICA), Cagliari,

Italy.

Diagnosis. Small-medium sized species, with a

low side profile and slit-like lunules axially elonga-

ted in the posterior ambulacra. Petals closed

distally, with the front odd longer than the other and

the posterior ones slight shorter. In the oral inter. 5

there are 2 post-basicoronal plates in column "a"

and 2 in column "b", with the first two ones stag-

gered; between the petals and the notches there are

3 or 4 couples of plates, and the periproct opens

along the suture between plates 2a/3b.

Description. Small-medium sized echinoid (TL

53 mm), with depressed test (TH = 11% TL) with

the higher point anterior to the apical disc and a thin

margin, rounded in outline. Petaloid medium size

(42% TL); petals closed, with the frontal odd longer

than the other. Poriferous zone flat, interporiferous

ones slightly raised, with interporiferous size rang-

ing from 1.5 to 2 those poriferous. Lunules axially,

more long than large (LI =20 mm; L2 = 6.6 mm)

and surrounded by 4 couple of plates on the aboral

side and by 3 in the oral one. In the inter. 5 there are

2 couples of post-basicoronal plates, the first two

2b and 2a staggered and the 2b in amphiplacous

contact with the first ambulacral postbasicoronal

plates; in this interambulacrum the WA is 36% TL.

The periproct is small (3% TL) and sharply

rounded, close the posterior margin (7% of TL) and

open along the suture 2a/3b. Main visceral central

hollow with wall reinforced by a network of thin

trabeculae; peripheral buttressing developed as

dense honeycombed meshwork of cellular struc-

ture; Aristotle's lantern width almost 27% of TL and

large but short caecum cavity. The food grooves are

very branched posteriorly; tubercolation well differ-

entiated adorally, dense and poorly differentiated

aborally (see Plate 19). Other data in relate Tables.

Etimology. From the name of S. Raimondi,

the collector that have donated the specimen to the

museum.

Distribution. Type locality and horizon: Re-

cent, Indonesia (Borneo). Occurrence: Recent, In-

donesia (Borneo).

Comparative notes. P. raimondii n. sp. differs

from P. arcuata in the shape of the lunules and in

the shorter distance of the lunules from the petals

tip. Moreover, in the oral side of P. arcuata , the

lunules open after two couples of post-basicoronal

plates in ambulacra I and V, while in P raimondii

lunules open after only one couples of plates.

Finally, the tubercles are absent or scarce along the

perradial sutures in P. raimondii and are always

present in P. arcuata.

Analysis of some astriclypeids (Echinoidea Clypeasteroida)

345

Plate 19. Paraamphiope raimondii n. sp., Recent, Indonesia. Figs. 1-3. Aboral, adoral and antero (to the left)-posterior (to

the right) lateral view of MAC.IVM206. Fig. 4. Close-up of spines and tubercolation in the oral area. Fig. 5. Scheme of

food grooves, more branched posteriorly. Fig. 6. Radiograph in supero-inferior projection with well visible support system

and a large Aristotle’s lantern. Fig. 7. Oral plate pattern.

346

Paolo Stara & Luigi Sanciu

Paraamphiope arcuata (Fuchs, 1882)

Plate 20 Figs. 1-6; Tables 6, 9

1882, Amphiope arcuata Fuchs, p. 31

1899, Amphiope arcuata Fuchs, Fourtau R., p. 698

1911, Amphiope arcuata Fuchs, Gregory, p. 667

1914, Amphiope truncata Fuchs, 1882, Cottreau,

p. 55

1920, Amphiope arcuata Fuchs, Furtau, p. 40

1920, Amphiope arcuata Fuchs, Migliorini, p. 153

Examined material. The material studied by us

is labeled as Miocene, Libyan desert (locality not

specified), housed in the NHMUK in London, with

code E1671-2, E1674-6, TL 35 - 79 mm.

Diagnosis. Small-medium sized species, with a

low side profile and ovoid lunules axially elongated

in the posterior ambulacra. Petals closed distally,

sub-equal in size. In the oral face of the inter. 5 there

are 2 post-basicoronal plates in column "a" and 2

(sometimes also a small part of the fourth plate is

visible) in column "b", with the first two staggered;

between the petals tip and the lunules there are 3 or

4 couples of plates, and the periproct opens between

plates 2a/3b.

Decription. Size small-medium (in our sample

max TL =79 mm) as wide as long. Test depressed

(TH = 6+ 12% TL). The highest part of the test lies

on the apical disc, which is sub-central. The am-

bitus outline is subrounded to subtrapezoidal; the

adoral surface is flat or slightly plano-concave with

the inner point near the peristome, which is sub-

central. There are 2 post-basicoronal plates in

column “a” and 2-3 in column “b”, in which the

2b is wide and elongated (like as in Amphiope ) and

are in amphiplacous contact with the relate post-

basicoronal ambulacrals. The periproct is small (2-

3% TL) and opens between the post- basicoronal

plates 2a/3b on inter 5; Lll varies from 4 to 14 %

TL. The peristome is round and measure from

3.5% to 5.5 TL.

The petals are just closed in larger specimens,

but the frontal odd seems open in the smaller spec-

imen. The Petalodium is of medium size (42 -s- 47

% TL). The lunules are very small and ellipsoidal

shaped. The B angles ranges between 88° to 96°.

Each lunule is separated from the corresponding

petal by 4-5 couples of plates and surrounded by

4-4 couples of plates on the aboral side, against

3-4 couples on the adoral one.

Apical disc with a small (~ 6% TL) star-shaped

madreporite, with 4 genital pores, all open also in

the smaller individuals. Internal structure and size

of Aristotle's lantern were not detected.

The main food grooves are simple and run

through the center of each column in the ambulacra,

starting from small branches parallel to the ambitus

or from the ambitus itself (in E76164). Short sec-

ondary branches grow along the grooves on the pos-

terior ambulacra and near the lunules and the

periproct. Tubercolation is poorly differentiated on

the oral face; tubercles are large on the basicoronal

interambulacral plates and on the post-basicoronal

ambulacral ones. In the interambulacra the tubercles

diminished in size farther from the center; large

tubercles surround the periproct. The tubercolation

covers with small tubercles also major food grooves.

On the aboral face the tubercolation is undifferen-

tiated, evenly distributed, dense and petite, over the

entire surface.

Distribution. Libya, Miocene. Locality and

horizon: Syouah, Gebel Ndefer, Egypt (the Holo-

type is housed in the Naturhistorisches Museum of

Vienna) and Libyan desert (Tobruc area), Middle

Miocene.

Comparative notes. Morphologically, P. ar-

cuata differs from “ Amphiope ” truncata Fuchs,

1882, in its smaller size, smaller petalodium, lunules

outline much more ovoids and the food grooves less

branched distally; P. arcuata differs from P. rai-

mondii n. sp. by the shape of the lunules and by the

longest distance of the lunules from the petal tips.

Moreover, the tubercles are always present along

the perradial sutures while in P. raimondii are ab-

sent or scarcely.

Remarks. Under careful observation of the spec-

imen E76164, the rear part of the test seems incom-

plete and for this reason the measurements and

plating are biased in this way. It is unclear whether

this anomaly occurred before or during the process

of fossilization.

The illustrations of the type species provided by

Fuchs (1882: 31, pi. XI, figs. 4-6) correspond, from

a morphological point of view, to the specimens stud-

ied by us. Cottreau (1914) puts this species in

synonymy with A. truncata Fuchs, 1882 and A. fuchsi

Fourtau, from the Middle Miocene of Egypt, then,

Analysis of some astriclypeids (Echinoidea Clypeasteroida)

347

Plate 20. Paracimphiope arcuata from Libya (Miocene). Fig. 1. Aboral view of NHMUK.E76162. Figs. 2, 3. Antero (to the

left)-posterior (to the right) lateral and adoral view of NHMUK.E76161. Fig. 4, 5. Plate pattern of aboral and adoral face of

NHMUK.E76164. Fig. 6. Food grooves scheme of NHMUK.E76161. In this example it is evident the great outline vari-

ability, in particular in the rear of the ambitus, due (Figs. 4, 5), perhaps, to defects in fossilisation.

348

Paolo Stara & Luigi Sanciu

in figure 23 he illustrates A. arcuata as type. How-

ever, the scheme of the plates of the other nominal

species has never been published, and therefore we

believe that they should remain a separate species.

Sculp sit echinus n. gen.

Type species. Sculpsitechinus auritus (Leske,

1778)= Echinodiscus aunt us Leske, 1778.

As Neotype are assigned specimen MAC.

IVM109 and is housed in the Department of Ani-

mal Biology and Ecology, University of Cagliari

(UNICA).

Description. Diagnostic features:

1 . Subcircular or vaguely polygonal visceral hol-

low, with the floor reinforced by a network of ribs;

2. Petalodium small, PL about 30-48% of TL;

petals always closed distally;

3. Pentastellate basicoronal circlet, with the in-

terambulacral plates that can be elongated distally,

usually separated from the post-basicoronal ones;

4. Posterior ambulacra with axial notches or

lunules; the lunules shape may vaiy from ellipsoidal

to narrow slits like, which may be open to the

ambitus;

5. 3 to 6 couples of plates are present between

petals tip and lunules/notches;

6. Periproct far from the rear margin almost 13

- 26% of TL;

7. 3 or 4 post-basicoronal plates per column in

inter. 5, with the first two partially paired and nor-

mally in meridoplacous contact with the relate am-

bulacrals;

8. Pood grooves very branched and spread over

all the oral surface;

9. Width at inter. 5 zone at ambitus about 30-

38% of TL;

10. B angle within 48° to 70°;

11. Tube-feet extending into interambulacral

zones.

Etimology. Sculpsit = carved: the name derives

from the rear notches that characterized the species

Echinodiscus auritus Leske, 1778, transferred here

to Sculpsitechinus auritus (Leske, 1778).

Distribution. Prom Indian Ocean, Red Sea,

Persian Gulf to West-Pacific. Time span: from

Middle Miocene to Recent.

Comparative notes. Sculpsitechinus n. gen. dif-

fers from Amphiope and Echinodiscus in that it has

3 to 6 pairs of plates between the posterior petals

tip and the respective lunules, whereas there are

only 1-2 in Amphiope and 2-4 in Echinodiscus ;

also the first two post-basicoronal plates in inter. 5

are relatively small and only partially coupled, in

Amphiope and Paraamphiope they are always stag-

gered, with the first one longer, and in Echinodiscus

they are always large and paired. Sculpsitechinus n.

gen. differs from the other genera also by the posi-

tion of the periproct that is far from the rear margin

(13-26% TL), while in the other ones this distance

ranges from 2.5 to 13% of the TL. furthermore,

Sculpsitechinus n. g. differs from Amphiope also in

having a smaller petalodium (30-47% against 45-

60%). Sculpsitechinus n. g. differs from Echinodis-

cus by the smaller width of inter. 5 at the ambitus

(30-38 against 35-54), and the lower angle be-

tween the lunules (B = 54°-70° against 70°-117°).

finally, Sculpsitechinus n. g. differs from Amphiope

and Echinodiscus by the food grooves highly

branched on the whole adoral surface.

Remarks. This genus includes the following

species:

S. auritus (L. Agassiz, 1838); Recent; Tulear, Mada-

gascar, Red Sea, Indian Ocean and West Pacific.

S. tenuissimus (L. Agassiz, 1847) Recent; Lem-

beh, North Sulawesi and Waigeo, West Papua (In-

donesia); New Caledonia, Papua New Guinea and

Palau, Micronesia.

Sculpsitechinus sp. 1 ; Recent; Bohol and Oslob

islands, Philippines.

Sculpsitechinus sp. 2, Middle Miocene; Papua

New Guinea.

Sculpsitechinus auritus (Leske, 1778)

Plate 21 Tigs. 1-7; Tables 6, 9, 11, 12 (see also

Table 3 in Stara & Lois D., 2014)

1778, Echinodiscus auritus , Leske N.G., p. 138

1778, Echinodiscus inauritus Leske N.G., p. 138

1816, Scutella bifissa Lamarck J.B.P.A., p. 10

1817, Scutella bifissa Savigny, pi. 7 fig. 3 (n.v)

1826, Scutella bifissa Lamarck, Auduin, p. 210

(n.v)

1841, Lobophora aurita L. Agassiz, pp. 70-71, pi.

14, figs. 3, 7

Analysis of some astriclypeids (Echinoidea Clypeasteroida)

349

1892, Echinodiscus auritus Leske, Boutan L., p.

29, 46, 47

1893, Echinodiscus auritus Leske, de Loriol P, p.

375

1894, Echinodiscus auritus Leske, Mazzetti G, pp.

222, 225-226 (n.v)

1 899, Echinodiscus auritus Leske, Fourtau R, pag.

540

1904, Tetrodiscus auritus Fourtau, pag. 425, 444

(n.v)

1914, Amphiope ( Tetrodiscus ) aurita Leske, Four-

tau R, p. 88 (n.v.)

1948, Echinodiscus auritus Leske, Mortensen T.,

pp. 400-406

1955, Echinodiscus auritus Leske, Tortonese E.,

p. 38

1971, Echinodiscus auritus Leske, Clark A.M. &

Rowe F.W.E., p. 144

1971, Echinodiscus auritus Leske, James D.B. &

Pearce J.S., p. 99

1981, Echinodiscus auritus Leske, Dollfiis R. &

Roman J., pp. 97-99

2014, Echinodiscus auritus Leske, Stara & Fois M.

Examined material. Neotype: MAC.IVM.109,

TL 125 mm, Recent, from Mangili, Tulear Prov-

ince, Madagascar. Housed in the Department of

Animal Biology and Ecology (UNICA), Cagliari,

Italy. 31 specimens from Mangili, Tulear Province,

Madagascar, caught in back-barrier lagoon, ~ 5 to

8 m in deep, in sandy-mud, MAC.IVM82 - 113-1

TL = 74 140 mm, housed in the Museo di Storia

Naturale Aquilegia, Cagliari and in the Department

of Animal Biology and Ecology (UNICA), Cagliari,

Italy.

Diagnosis. Medium sized species, with a low

side profile and axially elongated notches in the pos-

terior ambulacra. Petals closed distally, in a small

petalodium. In the oral interambulacrum 5 there are

3 post-basicoronal plates in column “a” and 4 in

column “b”, with the first two partially coupled;

between the petals tips and the notches there are 4

- 5 couples of plates, and the periproct opens along

the suture between plates 2b/2a in interambula-

crum 5.

Description. Medium-sized echinoid with a

almost polygonal ambital outline; the posterior mar-

gin line, sited between the two notches (like a tail),

is always irregular and often very asymmetric.

Although in smaller individuals a rounded outline

seems to prevail, the larger individuals present

clearly truncate lines; however the ambitus outline

can vary greatly. The adoral face is flat or slightly

plano-concave. The periproct is small and far from

the posterior margin (Lll = 18-24% TL). The

plating structure is reported in Plate 23.

In this samples the Aristotle’s lantern measures

about 15% TL.The petalodium is medium size (PL

= 35-40% TL) and the petals are sub-equal, twice

as long as the width and always closed (L5 = 18%;

L7 and L9 = 17% TL); the poriferous areas are 1.2

to 1 .5 times wider than the poriferous ones. The api-

cal disc measures 6% TL. Only one pair of post-

basicoronal ambulacral plates occlude the interam-

bulacrum 5. The notches are surrounded by 4-5 cou-

ples of plates on the oral face and by 4-5 on the

aboral one. Between the petal tip and the beginning

of the notch there are 6 couples of plates per column.

B is approximately 55° and WA at interambulacrum

5 is on average 32% TL. In the oral side of the in-

terambulacrum the periproct opens between post-

basicoronal plates 2b-2a. The stoma is pentagonal,

with a diameter of 4% TL; LI 3 = 11% TL. The ba-

sicoronal interambulacral plates are all irregular,

with some in contact and others disjointed. The

tuberculation is dense, made up of medium sized

tubercles, poorly differentiated and extended over

the entire aboral surface. The tubercles are larger

around the periproct and the smaller ones are found

particularly along the main food grooves. On the

aboral face the tubercolation is undifferentiated,

thick and petite, evenly distributed over the entire

surface. The food grooves are very branched out

over the entire oral surface.

Distribution. Tulear, Madagascar; Indian

Ocean, Red Sea, Persian Gulf, Oceania West-

Pacific Ocean. Recent.

Comparative notes. S. auritus differs from S.

tenuissimus and from Sculp sit echinus sp. 2, by

having notches against lunules. S. auritus differs

from Sculp sit echinus sp. 1, in having smaller

Aristotle’s lantern and greater size.

Remarks. To establish this species, Leske

(1778: 202), did not mention the locality where the

specimens studied come from and neither the mus-

eum in which these specimen has been deposited.

The author, however, has not even provided an

illustration of the sample that he described.

350

Paolo Stara & Luigi Sanciu

Plate 21. Sculpsitechinus auritus. Recent, Mangili, Madagascar. Figs. 1-3. Aboral, adoral and antero (to the left)-posterior

(to the right) lateral view of MAC.1VM109. Fig. 4. Radiograph taken in super- inferior position of MAC.IVM109, in which

is visible the small Aristotle’s lantern. Figs. 5, 6. Plate pattern of aboral and adoral faces of MAC.1VM110. Fig. 7. Well

branched food grooves scheme.

Analysis of some astriclypeids (Echinoidea Clypeasteroida)

351

Despite our historical research, followed by a

information request to the Natural History Museum

in Leipzig, the city where Leske has worked for

long time, we could not find the type specimens.

Kindly, R. Schiller from the Museum of Natural

History in Leipzig, communicated us (June, 17.

2014) that are not found in their museum specimens

of the Leske’s collection. Thanks to R. Schiller, R.

Wolf, from the Zoologic Collection of the Univer-

sity of Leipzig has informed us that some of his old

collections were dispersed in several other mu-

seums in Germany but they do not possess these

samples. Our historical research will be continued

but, being necessary to know some features that are

not described by the first author, as the test plate

structure, we felt it opportune to name as Neotype

a specimen of Sculps itechinus auritus (former

Echinodiscus cf. auritus in Stara & Fois M., 2014)

from Mangili, Madagascar.

Sculp sit echinus tenuissimus (L. Agassiz, 1847)

Plates 22, 23; Figs. 9a-c; Tables 6, 9, 12

1847, Echinodiscus tenuissima (L. Agassiz & Desor

E.,)

1847, Lobophora tenuissima L. Agassiz & Desor

E., p. 78

1861, Lobophora deplanchei Michelin (n.v.)

1863, Lobophora texta A. Agassiz, p. 359

1872-74, Echinodiscus laevis A. Agassiz (n.v.)

1881, Echinodiscus biforis Pfeffer (n.v.)

1948, Echinodiscus bisperforatus var. truncatus,

Mortensen T., pp. 409, 411,413

1971, E. tenuissimus Agassiz, 1847, Clark A.M. &

Rowe F.W.E, p. 148

1986, E. tenuissimus Agassiz, 1847 De Ridder C.

(n.v.)

Data (n.v. 1861-1881 taken from Kroh, A., 2014)

Examined material. The Type material is not

traceable in the Museum of Natural History of Paris

where it was housed (see Agassiz & Desor, 1 847).

For these reasons we had to establish a Neotype.

Neotype: one specimen from Lembeh Chan-

nel, North Sulawesi, Indonesia, MAC.IVM207,

TL 50 mm (Plate 21 Figs. 1, 2, housed in the De-

partment of Animal Biology and Ecology

(UNICA), Cagliari, Italy.

Two specimens from Lembeh Channel, North

Sulawesi, Indonesia, MAC.IVM207-208, and two

from Fantin Collection TL = 50-60 mm; one speci-

men from New Caledonia, NHMUK. 1981.11 .2.25,

TL 1 12 mm, TH 10.5 mm, one specimen from Palau,

Micronesia, NHMUK.59.7. 1.14, TL 120 mm, TH 11

mm. We have also considered a specimen from

“New Caledonia” by literature in: Dollfuss &

Roman (1981, table 33 figs. 5, 6), TL 121 mm; and

one by personal communications and photos by F.

Hattemberger collection, TL 68 mm, collected at

the depth of 2 meter in a sandy beach from

Noumea, Baie des Citrons, New Caledonia.

Diagnosis. Medium sized species, elongated (in

the specimens of New Caledonia and Palau) with a

maximum width very rear of the center (mean TW

= 97% TL), profile low, small and ellipsoidal-

shaped to slit- like lunules. Petalodium highly vari-

able (30 to 38% TL) smaller in the samples of New

Caledonia and in those of North Sulawesi, the

greater one in the specimen from Palau. In the oral

face of the inter. 5 there are 2-3 post-basicoronal

plates in column "a" and 3 in column "b", with 2a

and 2b partially paired; between the petals and the

lunules there are 4-5 couples of plates.

Description. Medium sized; small, narrow and

elongated ambulacral lunules (LI = 12% and L2 =

4% TL, with WI = 7.7 and SI = 0.33) and with a

narrower angle B (67 °). Depressed test (~ 9% TL),

with the most highly point anterior to the apical

disc. Thin margin, more thick anteriorly. Sub-equal

petals, closed distally, with the front one slightly

longer than the others; interporiferal zone wider

1—1.5 times of the poriferous ones. In the inter. 5

there are 2-3 post-basicoronal plates in column “a”

and 3 in column “b”, with the first two partially

coupled. The WA at the inter. 5 is 31% TL, one of

the lowest among Sculp sit echinus. The periproct is

small (2.5% TL), round shaped, far from the pos-

terior margin (11 to 18% TL) and open between

plates 2b/2a or 2b/3a/3b in inter. 5. The peristome

is sub-pentagonal and small (3.5% TL), sub-central.

Other features as for the genus. For any other data

see the relate tables and plates.

Distribution. Lembeh Channel, North Su-

lawesi; Waigeo, West Papua (Indonesia); New

Caledonia, Papua New Guinea and Palau Sibuan,

(Micronesia and Melanesia) and perhaps Japan.

Recent.

352

Paolo Stara & Luigi Sanciu

Comparative notes. S. tenuissimus differs

from S. auritus and from Sculps itechinus sp. 1 in

that it has lunules instead notches. S. tenuissimus

differ from Sculpsitechinus sp. 2 by the SI index

that is greater (0.33) against 0.18 of Sculpsitechi-

nus sp. 2. The data, however, is not sufficient to

separate with certainty this species, because the

oral plate structure of Sculpsitechinus sp. 1 is still

unknown.

Remarks. As already mentioned in the para-

graph on Echinodiscus andamanensis n. sp., not

having been published the plating of the type

species Echinodiscus tenuissimus , under this name

are included several morphotypes based on the test

and lunules shape, coming also from very different

geographical areas. However, the description of

the type species made by Agassiz & Desor (1847:

78) is really laconic "species very flat, with small

lunules, corresponding to the ambulacra pair pos-

terior” and the sample deposited at the time in the

"Museum of Paris", is now wanting (pers. comm.

Sylvain Charbonnier, June 03.2014). Agassiz &

Desor (1847) mentions the geographical origin

(Waigiou), which corresponds to New Britannia

(Indonesia). In the zonation resulted from our

observations, in this area only Sculpsitechinus

species are present. It seems clear, therefore, the

need to appoint a neotype from the closest geo-

graphical area.

Sculpsitechinus sp. 1

Plates 1, 2; Table 1 in Stara & Sanciu (2014)

Examined material. 12 specimens, Recent,

MAC IVM 81; MAC.IVM206 - MAC.IVM215;

MAC.IVM233, housed in the MAC, Cagliari, Italy.

5 specimens from Oslob (TL 131 ^ 154 mm), 5

specimens from Cebu (TL 152 to 173 mm); two

examples of generic origin "Philippines" (TL 121

and 152 mm).

Diagnosis. Large sized specie (up to 173 mm),

with low side profile (mean TH = 12% TL), narrow

and elongated ambulacral notches open on the pos-

terior margin. Sub-equal petals, closed distally, with

the frontal odd petal sometimes slightly longer than

the other ones. In the oral face of the inter. 5 there

are 3 post- basicoronal plates in column "a" and 4

in column "b", with the first two partially coupled

and the periproct that opens between the plates

2a/2b. Between the petals tip and the beginning of

the notches there are 4 or 5 couples of plates.

Description. Large in size (in the studied sam-

ple TL 121 4- 173 mm), with more or less axially

elongated ambulacral notches open on the posterior

margin. Depressed test with the highest point ante-

rior to the apical disc. Thin margin, more thick an-

teriorly. The petals are closed, sub-equal, with the

frontal odd one a little longer than the other; porif-

erous zone flat or slightly sunken, interporiferous

ones slightly raised, with interporiferous areas 1.5

to 2 larger than the poriferous ones. Notches vary-

ing in length from 18 to 27% of TL and are more or

less narrow. The WA at inter. 5 is on average 33%

TL and the B angle is on average 57°. The periproct

is small (2.5% TL), rounded in shape, far from the

posterior margin (16-24% TL) and always open

along the suture 2b/2a. The internal structure con-

sists of a central visceral hollow and a peripheral

support structure. The hollow is sub-rounded to

polygonal shaped and its size corresponds to the

petalodium length (PL ~ 42% TL), the floor is thin

and reinforced by a structure made by a network of

thin trabeculae. The system of pillars and buttresses

is similar to the S. auritus. the Aristotle's lantern is

very large (20-24% of TL in specimen 140 mm

long). Lor descriptive statistics see Stara & Lois M.

(2014).

Distribution. Island of Talibon (Bohol) and

Island of Oslob (Cebu), Philippines. Recent.

Comparative notes. Sculpsitechinus sp. 1 dif-

fers from S. auritus by a larger Aristotle's lantern

that measures 20-24% TL against 15-18% TL in a

specimens 140 mm long. Sculpsitechinus sp. 1 dif-

fers from S. tenuissimus by a less number of couples

of plates between petals tip and lunules/notches,

which are 3-4 against 5-6 and have notches instead

lunules.

Sculpsitechinus sp. 2

Plate 14 Pig. 4

Based on the illustration in Lindley (2001:128,

fig. 7d.

2001, Echinodiscus bisp erf or atus Leske, 1778.

Lindley, p. 128.

Analysis of some astriclypeids (Echinoidea Clypeasteroida)

353

Plate 22. Sculpsitechinus tenuissimus, Recent, Lembeh (Indonesia) and New Caledonia. Figs. 1, 2. Aboral and adoral view

of MAC.IVM207 from Lembeh, North Sulawesi, Indonesia. Figs. 3, 4. Plate pattern of aboral and adoral faces of

MAC.IVM207. Fig. 5. Radiograph taken in supero-inferior position of MAC.IVM207; are visible the parts of the small

Aristotle’s lantern. Fig. 6. Aboral view of well preserved specimen from New Caledonia (F. Hattemberg collection).

354

Paolo Stara & Luigi Sanciu

Plate 23. Sculpsitechinus tenuissimus, other example from New Caledonia. Figs. 1-3. Aboral, adoral and antero (to the left)-

posterior (to the right) lateral view of NHMUK. 1981.11.2.25. Fig. 4. Food grooves very branched scheme of

NHMUK. 198 1 . 1 1 .2.25. Fig. 5. Plate pattern of aboral face of NHMUK. 198 1.1 1.2. 25. Figs. 6, 7. Aboral and adoral plate

pattern from figs. 5-6, PI. 33 in Dolfuss & Roman (1981).

Analysis of some astriclypeids (Echinoidea Clypeasteroida)

355

Examined material. One specimen TL = 70

mm; TW = 65 mm, housed in the Departement of

Geology, Australian National University of Cam-

berra; code ANU 60549.

Description. Test depressed of medium size,

discoidal with anterior semi-circular outline, trun-

cated posteriorly. Central apical system not well

legible. Petals straight, narrow, distinctly closed

distally; PL medium (43% TL); poriferous areas

slightly less than the width of interporiferous ones.

Axial, narrow Lunules in the posterior ambulacra.

Tubercolation not legible.

Given its general shape, its large distance sepa-

rating the lunule from the respective petals tips, and

given the small B angle, this echinoid appears very

near to S. tenuissimus.

Lindley (2001) accost this form to E. truncatus

“ lunule length relative to test radius or petal length

as a diagnositic character. The possession of closed

posterior lunules of a similar length to petals

clearly indicates assignment of the Aseki specimen

to Echinodiscus bisperforatus Leske, 1778. E. te-

nuissimus (L. Agassiz, 1847), a similar species in

many respects, possesses lunules shorter than

petals. Although Mortens en (1948: 409) observed

that the length of lunules varies very considerably

within this species, it is useful to note that the lunules

of the Aseki specimen are at most about as long the

petals, a diagnostic character of var. truncatus (L.

Agassiz, 1841)”.

Besides, the size of the lunules (LI = L2 =

15% TL and 2.8% TL and SI = 0.18) is different

from that of S. tenuissimus, in which LI = 12%

TL and and L2 = 4% TL, with an SI = 0, 33. The

data, however, is not sufficient to separate with

certainty this species from S. tenuissimus. In fact,

the features of the oral face and of the plate struc-

ture are unknown.

Distribution. Langimar Beds, Middle Miocene,

Aseki Village (Morobe Province), Papua New

Guinea.

Remarks. Based on its geological age, we be-

lieve that it is an ancestral species of S. tenuissimus

and S. auritus currently living in the same regions.

However, in the absence of further details such

species is left in open nomenclature: Sculpsitechi-

nus sp. 2.

ACKNOWLEDGEMENTS

We warmly thanks Enrico Borghi of the

Societa di Scienze Naturali of Reggio Emilia, for

critical reading of the manuscript; Claudia Puddu

for the careful translation of the manuscript and

for helpful comments on geologic parts; Gianluigi

Pillola and Carlo Corradini, responsible for the

Museo di Paleontologia "D. Lovisato " at the

Dipartimento di Chimica e Geologia, Universita

di Cagliari; Maria Tavano of the Museo Comu-

nale di Storia Naturale "G. Doria” in Genoa, and

Maria Cristina Bonci of Dip.Te.Ris, Universita di

Genoa; Timothy Ewin, Curator of Invertebrate

Paleontology of the Department of Earth Sci-

ences, and Consuelo Sendino who helped us in

the work, as well as Andrew Cabrinovic, Curator

of the Division of Aquatic Invertebrates of the

Life Sciences Department, the Natural History

Museum London; Tom Schotte, collection man-

ager, Echinodermata, of the Natural History Mus-

eum of Denmark (Zoology), Copenhagen, and

Charatsee Aungtonya Reference Collection of the

Phuket Marine Biological Center, Phuket, Thai-

land, to make available numerous specimens pre-

served in the collections of museum and their

respective departments; Cristian Biagioni of the

Dipartimento di Scienze della Terra, Universita di

Pisa, for the Amphiope fragments verification on

electron microscope; Roberto Rizzo of the Parco

Geominerario Storico e Ambientale della Sardegna,

for his support in the preparation of the geological-

paleogeographic parts; Mario Lai (3S, Laboratori

immagini, Capoterra) for scoring numerous radio-

graphs of the examined specimens; Luca Rai-

mondi (Radiologia Novi Ligure) for producing

some other X-ray radiograph. We are grateful,

also, to David Serra for allowing us to access on

outcrops in Cuccuru Tuvullao and Luciano Con-

cas (Arbus), Sergio Raimondi (Genoa) and Marco

Pantin (Venice) for providing us with some inter-

esting specimens of living scutellids. We thank

also Heidi Lriedhoff (Norderstedt) for information

on the Pleistocene deposits of Hurghada, Sergio

Caschili (Cagliari) for the loan some specimens

of A. nuragica; Ashley Miskelly (Sydney) and

Lranck Hattenberger (Nova Caledonia), to give us

information and/or photos of specimens of “Echin-

odiscus” locality.

356

Paolo Stara & Luigi Sanciu

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Biodiversity Journal, 2014, 5 (2): 359-366

Taxonomic notes on the genus Pseudoapterogyna Escalera,

1914 (Coleoptera Scarabaeoidea Melolonthidae) in Sicily

Ignazio Sparacio

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

ABSTRACT All Sicilian records of the genus Pseudoapterogyna Escalera, 1914 (Coleoptera Scarabaeoidea

Melolonthidae) are revised. As a result four species are reported to occur in Sicily: P euphytus

lamantiai n. ssp. (for the populations of Pantelleria Island, previously attributed to P euphytus

s.l.), P vorax (Marseul, 1878) from Lampedusa Island, P. pellegrinensis (Brenske in Ragusa,

1893) from Western Sicily (to which all P euphytus records from Sicily need to be attributed),

and P michaelis n. sp. from Western Sicily.

KEY WORDS Melolonthidae; Pseudoapterogyna', Sicily; taxonomy.

Received 18.05.2014; accepted 22.06.2014; printed 30.06.2014

INTRODUCTION

The Sicilian Pseudoapterogyna Escalera, 1914

(Coleoptera Scarabaeoidea Melolonthidae) have

been attributed thus far to P. euphytus (Buquet,

1 840), a species deemed to occur through Algeria,

Tunisia and Sicily (Rottenberg, 1871; Ragusa,

1873; 1874; 1893; Bertolini, 1899; Heyden et al.,

1883; Luigioni, 1929; Porta, 1932; Baraud, 1977;

1985; 1992; Arnone et al., 1995; Carpaneto &

Piattella, 1995; Sparacio, 1995; Smetana & Krai,

2006; Arnone, 2010; Ballerio et al., 2010).

Examination of type material of P. euphytus plus

a large quantity of specimens from Sicily and North-

ern Africa allowed me to re-evaluate the taxonomic

status of the Sicilian populations of Pseudoapterog-

yna. As a result P euphytus euphytus proved not

to occur in Sicily, being restricted to Algeria and

Tunisia, while the Sicilian populations of Pseu-

doapterogyma proved to represent three distinct

taxa: P. euphytus lamantiai n. ssp. from Pantelleria

Island (Sicily Channel), P pellegrinensis (Ragusa,

1892) and P. michaelis n. sp., both endemics of

North-Western Sicily.

I consider Pseudoapterogyna Escalera, 1914 as

a separate genus (according with Baraud, 1985;

1992), and not as a synonym of Geotrogus Guerin-

Meneville, 1842, as recently proposed by Coca-

Abia (2003, see also Smetana & Krai, 2006).

ACRONYMS AND ABBREVIATIONS. V.

Aliquo collection, Palermo, Italy (CVA); M.

Arnone collection, Palermo, Italy (CMA); M.

Bellavista collection (CMB); Armando Monastra

collection, Palermo, Italy (CAM); M. Romano col-

lection, Capaci, Palermo, Italy (CMR); I. Sparacio

collection, Palermo, Italy (CIS); A. Tetamo collec-

tion, Palermo, Italy (CAT); Collection of Diparti-

mento di Biologia Animale University of Catania,

Italy (CMC); Collection of Museo Civico di

Storia Naturale "Giacomo Doria", Genova, Italy

(MCSNG); Collection of Museum National d'His-

toire Naturelle, Paris, France (MNHN); m = male/s;

f = female/s. Unless otherwise stated, the collector of

the beetles in the field is the owner of the collection.

360

Ignazio Sparacio

Pseudoapterogyna euphytus euphytus Buquet,

1840

Examined material. 1 ex labelled: Ex Musaeo

Mniszech / a Euphytus Bqt alg. type / Museum

Paris - ex coll. R. Oberthur / SYNTYPE - Rhizotrogus

euphytus Buquet, 1840 / SYNTYPE / SYNTYPE -

Pseudoapterogyna euphyta (Buquet, 1840) /

MHNHN EC4183. 1 ex labelled: Ex Musaeo

Mniszech / Museum Paris - ex coll. R. Oberthur /

SYNTYPE - Rhizotrogus euphytus Buquet, 1 840 /

SYNTYPE / SYNTYPE - Pseudoapterogyna eu-

phyta (Buquet, 1840) / MHNHN EC4184. Algeria,

Costantine, Bonvouloir, 1 m (MCSNG); Algeria,

Costantine, Henon, 3 m and 1 f (MCSNG); Algeria,

Batna, 27 marzo 1952, ex coll. G. Fiori, 1 m

(MCSNG); Algeria, Bone, 1874, Puton, 2 m and 2

f (MCSNG); Tunisia, Tunisi, Aut.-Inv. 1981-2, G.

e L. Doria, 3 m (MCSNG); Tunisia, Tunisi dint.,

1881, G. e L. Doria, 1 m

Biology and Distribution. Aduls often found

under stones. Algeria and Tunisia.

Remarks. P. euphytus was described from Con-

stantine in Algeria (Buquet, 1840). The synonyms

of this species are the following (Baraud, 1985;

1992; Smetana & Krai, 2006): maculicollis Fair-

maire nee Villa, 1860; biskrensis Marseul, 1878

locus typicus: Biskra, Algeria; tuniseus Fairmaire,

1884 locus typicus: Tunis, Tunisia; dilutus Fair-

maire, 1860 locus typicus: Tunis, Tunisia.

The study of P. euphytus type material (Figs. 1,

2), and of several specimens from the type locality

and other places in Algeria and Tunisia allows to

summarize the diagnosis of this species (see also

Baraud, 1985) as follows:

Males with fully developed metathoracic wings,

females are flightless. Length 12-16 mm.; yellow-

ish-brown, disc of pronotum sometimes darker;

dorsal surface sub-opaque; antennae 1 0-segmented,

scape elongate and dilated distally, club developed,

shorter than fiinicle (club/funicle = 0.64); pronotum

glabrous, only a few sparse short erect setae on an-

terior margin, surface micro-reticulated, punctation

with large and dense punctures (the distance be-

tween the punctures being subequal or inferior to

their diameter); posterior angles well marked, in

most cases distinctly projecting backwards, pre-

ceded by a slight sinuosity at lateral sides; base of

pronotum marked by a trasversal row of coarse punc-

tures; elytral surface densely microreticulated, with

shallow and poorly defined puncturation. Anterior

tibiae tridentate on external margin, basal tooth very

short, sometimes absent. Posterior tibiae without Ca-

rina on dorsal side or, at least, proximally. 1° metatar-

somere short in both sexes. Claws toothed at base

and with a spine well developed. Pygidium finely

wrinkled and micro-reticulated with puncturation

made of shallow scattered large punctures mixed to

other much smaller and denser punctures. Posterior

coxae of males far from median coxae. Aedeagus

with parameres, in lateral view, narrowed distally

with very elongate and pointed apex.

Females have a more convex dorsum, a smaller

antennal club, shorter tarsi, posterior angles of

pronotum often pointed, and larger and coarser ely-

tral punctation.

Pseudoapterogyna euphytus lamantiai n. ssp.

Examined material. Holotypus male (CIS):

Pantelleria (Sicily, Italy), Sesi, 10.V.1991, 1 m;

Paratypes: ibidem, 30.IV. 1995, 2 m and 4 f (CIS).

Pantelleria, 11.1906, S. Sommier, Rhizotrogus eu-

phytus Buq., det. Sabatinelli, 4 m (MCSNG). 1

male labelled: “Pantelleria” and 1 male without

label, likely from the same locality as above (see

below) E. Ragusa collection (CMC).

Description of holotypus. Length 12 mm.

Yellowish-brown, disc of pronotum sometimes

dark brown; antennae, palpi, legs, pubescence, and

ventral surface yellowish; dorsal surface sub-

opaque. Head with deep and dense punctation and

a thin, slightly raised transverse carina; clypeus

slightly emarginate at middle of anterior margin.

Antennae 1 0-segmented, club 3 -segmented, about

half the length of funicle (club/funicle = 0.52);

scape dilated distally, almost as long as the 2°, 3°

and 4° segments together. Pronotum with maximum

width just before middle, sides crenulated, subrec-

tilinear with a little sinuation before posterior pro-

truding angles; anterior margin subrectilinear; basal

bead complete, flattened and punctate; pronotal

sculpturing made of middle-sized deep sparse punc-

tures (the distance between the punctures being

more than double their diameter) clearly visible on

the micro-reticulated surface; the anterior margin

and sides of pronotum bear short and sparse fine

setae. Scutellum triangular, bearing some short

Taxonomic notes on the genus Pseudoapterogyna Escalera, 1914 (Coleoptera Scarabaeoidea Melolonthidae) in Sicily 361

setae near base, with big shallow punctures. Elytra

subparallel, slightly dilated at apical third, with

micro-reticulated surface bearing some deep small

punctures; humeral callus visible. Anterior tibiae

tridentate, with a very weak hardly visible basal

tooth. Posterior tibiae without carina on dorsal side.

Tarsi about twice longer than tibiae. First posterior

tarsomere shorter than the 3° tarsomere. Claws toothed

at the base and with a spine well developed. Pygid-

ium finely wrinkled, micro-reticulated, with punc-

tures small. Metathoracic wings fully developed.

Aedeagus with parameres strong, narrowed distally,

with slightly elongate apex.

Variability. Length 11.5-13 mm; the disc of

pronotum may be yellowish in colour, like the rest

of the body. The little carina of dorsal side of pos-

terior tibiae may be absent or very reduced. Females

are flightless, have a more convex dorsum and ely-

tra more dilated backward, with dorsal punctation

sparse and shorter tarsi; posterior tarsi little longer

than posterior tibiae.

Etimology. Latin noun in the genitive case.

After Tommaso La Mantia, University of Palermo,

Italy, in acknowledgement of his oustanding exper-

tise on the Sicily channel islands.

Biology and Distribution. Adults found

under stones, active during Spring. Records from

other seasons (see Arnone et al., 1995) are likely

due to findings of dead specimens under stones,

often found in good condition due to the dry envi-

ronment. Endemic to Pantelleria Island.

Comparative notes. P. euphytus lamantiai n.

ssp. differs from P. euphytus euphytus from Algeria

and Tunisia by the following characters: smaller

size, males with dorsum less convex, more parallel

body (especially the elytra), punctation (in particu-

lar that of pronotum) sparser and less deep, antennal

club longer, sculpturing of pygidium without big

punctures, and shape of parameres with stouter and

shorter apex.

Remarks. All previous records from Pantelleria

Island, such as Ragusa (1875: sub Rhizotrogus

Gerardi Buq.), Bertolini (1899: sub Rhizotrogus

Gerardi Buq.), Luigioni, 1929 (sub Rhizotrogus eu-

phytus Buquet), Liebmann (1962: sub Rhizotrogus

KuiauB Porn*

ex Cckl 1 •

ft.Oberthvir

SYNTYPE

Rhizotrogus r

suphytus Buquef 1840

SYNTYPE

Pseudoapterogyna

auphyta (Buquet. 1840)

MNHN

EC4184

5 mm

Mnrswchl

SYNTYPE

Rhizotrogus .

auphytus Buquet. 1840

SYNTYPE

Psaudoaptarogyna

auphyta I Buquet 1840)

MNHN

EC4183

Figures 1, 2. Syntypes of Pseudoapterogyna euphytus (MHNHN).

Figure 3. Aedeagus of the P. euphytus from Costantine, Algeria, length 6.3 mm (MCSNG).

362

Ignazio Sparacio

euphytus), Ratti, 1987 (sub P. euphytus ) and finally

Amone et al. , 1995 (sub P. euphytus ), actually refer

to P. euphytus lamantiai n. ssp.

Pseudoapterogyna pellegrinensis (Brenske in

Ragusa, 1893)

Examined material. Lectotypus male: Falde

3. [Monte Pellegrino leg. E. Ragusa], E. Ragusa

collection (CMC); paralectotypes, idem 8 males

and 3 females, without label, likely from the same

locality as above (see below) E. Ragusa collection

(CMC); Palermo, Addaura, 17.I.1970-10.II.1970

(CVA); Monte Cuccio (Palermo), 4.IV.1970-

27. III. 1971 (CVA); Cinisi (Palermo), 7.1.1973-

1 l.II. 1973-7 and 19.III.1973 (CVA); Capaci

(Palermo), 11.11.1973 (CVA); Mazara del Vallo

(Trapani), 16.IV.1976, 9.1.1983, 15.11.1983,

2. IV. 1983, 13.V.1983, 6.XII.1986, 11.IV.1987

(CVA); Foce Fiume Belice (Trapani) 13. IV. 1984

(CVA). Cinisi (Palermo), 11.11.1975 (CAM); Mazara

del Vallo (Trapani), 6.II.1984 (CAM); Torretta (Pa-

lermo), 23.11.1977 (CAM). Capaci (Palermo)

9.1.1973, 8. II. 1973, 6.III.1973, 27.1.1980 (CMR);

Carini (Palermo), 7.III.1973 (CMR); Cinisi (Pa-

lermo) 19. III. 1973 (CMR); Castelluzzo litorale

(Trapani), 10.IV.2011 (CMR); Capaci (Palermo),

27.1.1980, 3. II. 1980, 17.11.1980 (CMA); Cinisi

(Palermo), 2 l.III. 1986 (CMA); Palermo, Addaura,

23.XII.1972, leg. A. Carapezza (CMA); Campo-

bello di Mazara: Cave di Cusa (Trapani), 7.IV.1985

(CMA); Castelluzzo litorale (Trapani), 14.III.1999,

17.11.2002, 10.IV.2011 (CMA); Isole Egadi, Favi-

gnana (Trapani), 30. IV. 1969 leg. B. Massa (CMA);

Mazara del Vallo (Trapani), 9.1.1983, 30.IV.1983

CMA). Palermo, Tommaso Natale, 23.III.1978, 1

m (CIS); Capaci (Palermo), 17.11.1980, 4 m and 3

f; idem, 16.1.1983, 1 m 10.V.1991, 1 m(CIS); Sfer-

racavallo, LaConza, 18.1.1981, 2 f; idem 10.V.1991,

1 m (CIS); Mazara del Vallo (Trapani), l.II. 1981, 1

m (CIS); Isola delle Femmine (Palermo), 25.11.1983,

1 m 10.V.1991, 1 m (CIS); Cinisi (Palermo),

25.11.1983, 1 m 10.V.1991, 1 m (CIS); Terrasini (Pa-

lermo), 12. IV. 1983, 1 m; idem, 24.IV.1993, 1 f;

idem, 2.IV.2002, 1 f 10.V.1991, 1 m (CIS); Monte

Cofano (Trapani), 14.IV. 1991, 1 m (CIS).

Biology and Distribution. A Mediterranean

maquis dweller, usually found in open disturbed

maquis, with stony grounds. Adults of P pellegri-

nensis are found under stones, or walking, or (males

only) flying during Spring months. Larvae are ri-

zophagous and found underground. Endemic to

North-Western Sicily, in strong rarefaction near Pa-

lermo where it has disappeared from many locali-

ties in the last thirty years; very localized in the area

of Trapani.

Comparative notes. Compared to P. euphytus ,

P. pellegrinensis is bigger and wider (length 15-18

mm), shiny and has a less convex dorsum. Diagno-

sis: reddish-brown. Male: club/funicle = 0.52, prono-

tum wider just before anterior half, with punctures

smaller, spaced and shallow, irregularly distributed,

thinner at the sides; sides less curved, slightly

sinuated just before posterior angles, posterior

angles well-marked but not protruding. Elytra

wrinkled, with irregular longitudinal striae, with

hardly visible micro-retic-ulation and punctures big

and deep. Posterior tibiae distinctly keeled on the

dorsal surface. Ventrites shiny, the first three with

a longitudinal depression. Pygidium much wider at

base, coarsely wrinkled, micro-reticulated and with

punctures small and very dense; apical margin

slightly emarginate. Aedeagus with parameres

stouter, slightly elongate distally.

Female: females are wingless, with a more con-

vex dorsum and elytra more dilated backward; pos-

terior tibiae distinctly keeled on the dorsal surface;

posterior tarsi little longer than posterior tibiae.

Remarks. In Sicily, P euphytus was reported

for the first time by Rottemberg (1871: “Am Fuss

des M. Pellegrino”) and mentioned several times by

Ragusa (1873; 1874; 1893).

Ragusa (1874), however, was not convinced of

the identity of this beetle “... il Barone di Rottem-

berg ... lo crede il R. Euphytus Buq... Quest’ ins etto

merita d’essere attentamente studiato ...”. [“Rot-

temberg ... he believes it is R. euphytus Buq. ... this

insect deserves to be carefully studied”].

In his " Catalogo ragionato dei Coleotteri di Si-

cilia", Ragusa (1893) provided a long comment on

P euphytus in a footnote in which he translates a

letter that he received from Brenske (to whom he

had “ comunicato tutte le specie di Rhizotrogus di

Sicilia ” ["sent all the species of Rhizotrogus of

Sicily”]). Brenske, after examining the bibliography

on P euphytus available at that time, indicates that

the Sicilian populations of the species should be dis-

tinguished from those of North Africa: “( 1 ) Var. pel-

Taxonomic notes on the genus Pseudoapterogyna Escalera, 1914 (Coleoptera Scarabaeoidea Melolonthidae) in Sicily 363

legrinensis Brenske var. nov. ... esse si devono se-

parate, se non come due specie distinte, una come

varieta dell’altra . ... La differenza piii evidente sta

nella punteggiatura del pygidio, che nell ’ insetto di

Sicilia non e lucido, ed e leggermente aggrinzita,

mentre l ’algerino e oltre di cid ricoperto di grossi

e forti punti. Per la specie di Sicilia io scelgo il

nome di pellegrinensis, per indicare la localita dove

questa specie e stata fin ’ora trovata .”

[“(1) Var. pellegrinensis Brenske var. nov. ...

they must represent two distinct species or at least

two distinct varieties. The most obvious difference

is in the punctation of pygidium, which in the insect

from Sicily is not shining, and slightly wrinkled,

while in the Algerian specimens, in addition to this

character, is covered by big and strong punctures.

For the species of Sicily I choose the name pelle-

grinensis to indicate the location where this species

has been found so far.”]

It is quite clear from the footnote of Ragusa's

1893 paper that the name "pellegrinensis" and the

description come from Brenske. I think therefore

that Brenske is alone responsible both for the name

and for satisfying the criteria of availability other

than publication, hence, according to art. 50.1.1 of

the Code, Brenske is the author of the name ” 'pelle-

grinensis'’ which needs to be quoted as Bresnke in

Ragusa. Ragusa provided only the translation of

Brenske's letter, but this circumstance, in my opin-

ion, is not sufficient to change the sole responsi-

bility of Brenske.

The populations of Sicily, so far reported as

P. euphytus, show clear differences from P. euphy-

tus and therefore must be attributed to P. pelle-

grinensis. Reports of P. euphytus from the

Lampedusa Island (Failla Tedaldi, 1887; Heyden

et al., 1891; Goggi, 2004) are likely records of P.

vorax (see below).

In Ragusa's collection there are 14 specimens

of Pseudoapterogyna from Sicily (see also Amone,

2010); only two of them bear a locality label. One

of them comes from Pantelleria (together with

other two specimens of the series of 14 speci-

mens) and therefore belongs to P. euphytus laman-

tiai (see above), while the other bears the

following label:: "Falde, 3, E. Ragusa. [Monte

Figure 4. Pseudoapterogyna euphytus lamantiai n. ssp. Fig. 5. Idem, aedeagus, length 5.6 mm. Fig. 6. P. pellegrinensis.

Fig. 7. Idem, aedeagus, length 6 mm. Fig. 8. P. michaelis n. sp. Fig. 9. Idem, aedeagus, length 6 mm. (photos M. Romano).

364

Ignazio Sparacio

Pellegrino leg. E. Ragusa]" (foothills, March, E.

Ragusa. [Monte Pellegrino leg. E. Ragusa]". I

hereby designate this latter specimen as the lec-

totypus of P. pellegrinensis. The following red

handwritten label has been added to it: Lectotypus

- Pseudoapterogyna pellegrinensis Brenske in

Ragusa, 1893, 1. Sparacio des. 2014. Ragusa used

to add the locality label only to the first specimen

of a series (Amone, 2010), therefore the remaining

ten specimens of the series, all belonging to P. pel-

legrinensis , have to be consdiered as paralectotypi.

Pseudoapterogyna vorax (Marseul, 1878)

Examined material. Sicily (Agrigento), Lampe-

dusa, 4.VI.1975, 1 m {Pseudoapterogyna vorax

Mars. J. Baraud det.), legit B. Massa (CM A); idem,

1 f, 5.VI.1975 (CMA); Sicilia (Agrigento), Isola di

Lampedusa, 15.V.1983, 1 f (CIS).

Biology and Distribution. Adults active in

May and June. Records from other months (Amone

et al., 1995; Ballerio et al., 2010) are likely findings

of dead specimens, often found under stones.

P vorax is widespread from Morocco to Libia

(Baraud, 1985). Baraud (1977) was the first to report

the occurrence of this species in Lampedusa (Sicily

Channel), later Smetana & Krai (2006) added a

record from Lampione (an islet next to Lampedusa).

Remarks. A comparison between the holotype

of P. vorax (MNHN, locus typicus: Algeria, Batna)

and a few specimens from Lampedusa Island did

not reveal any relevant difference between the two

populations.

Pseudoapterogyna michaelis n. sp.

Examined material. Holotypus male: Monte

Cofano (Trapani, Sicily, Italy), 20.XI.2011 (CIS).

Paratypes: idem, 2 m (CIS); idem, 17.XI.20 13, 31

m (CIS); idem, 20 m, legit A. Tetamo (CMB); idem,

8 m (CAT); idem, 1 m, legit I. Sparacio (MNHN);

idem, 1 m, legit I. Sparacio (MCSNG).

Description of holotypus. Length 14.5 mm.

Shiny. Reddish-brown, with darker pronotum;

palpi, antennae, tarsi, lateral margins of pronotum,

and underside yellowish-brown; long yellowish-

brown erect setae are present around the body, denser

in the central part of both the anterior and posterior

margins of pronotum, the latter having also a very

dense tuft of long fine recumbent setae. Sternum

with dense yellowish setation. Antennae 10-seg-

mented; scape elongate and distally dilatate, almost

as long as 2°-3° and 4° segments together; 2° seg-

ment very short; club shorter than the 7 previous

segments together (club/fiinicle = 0.60). Clypeus

with anterior margin emarginate at middle.

Head covered with big deep dense punctures; a

small transverse carina slightly raised and not reach-

ing the sides is present on clypeus. Pronotum trans-

verse, 2.3 times wider than long, sides slightly

curved at basal, norrowed distally, maximum width

at basal half; posterior angles obtuse; anterior mar-

gin slightly curved forward, posterior margin pro-

jecting backward in the middle with basal bead thin

and with sparse little punctures; a smooth longitu-

dinal line is present in the middle of pronotum;

pronotal surface smooth (not microreticulated),

sculpture formed by medium sized deep dense punc-

tures, regularly distributed over the entire surface

(the distance between the punctures sub equal to

twice their diameter). Scutellum wide, subtriangu-

lar, with curved sides, micro-reticulated, with deep

dense punctation, concentrated mainly at sides,

covered by the dense recumbent long fine setation.

Humeral callus present. Elytra broad, dorsally

flattened, wider at distal third, coarsely striated,

with punctures large, dense and deep at the base

then densely wrinkled at apical third; elytral apex

slightly divergent. Pygidium with poorly defined

shallow sparse punctures on densely micro-reticu-

lated surface. Tarsal claws toothed at base. Anterior

tarsi elongate, 1.75 longer than the corresponding

tibiae. Anterior tibiae tridentate on external margin.

Posterior tarsi elongate, 1.75 longer than the corre-

sponding tibiae, 1 ° tarsomere distinctly shorter than

the 3°. Posterior tibiae without carina on dorsal side.

Metathoracic wings fully developed.

Aedeagus, in lateral view, with sub-parallel

parameres, distally slightly wider, with apex short,

sharp and slightly curved.

Lemale unknown.

Variability. Body length 13-18 mm; colour of

the dorsal surface is sometimes much darker, with

pronotum almost completely dark. In some speci-

mens there is a short and weak carina on the top of

the dorsal face of posterior tibiae.

Taxonomic notes on the genus Pseudoapterogyna Escalera, 1914 (Coleoptera Scarabaeoidea Melolonthidae) in Sicily 365

Figure 10. Distribution of the Sicilian members of the genus

Pseudoapterogyna and of P. euphytus euphytus in North

Africa (rhombus). P. euphytus lamantiai : triangle; P pelle-

grinensis : circles; P. michaelis : star; P. vorax (also in Mo-

rocco): squares.

Etimology. Latin noun in the genitive case.

This new species is dedicated to my friend Michele

Bellavista (Palermo, Italy).

Biology and Distribution. All specimens col-

lected were found still alive in potholes with water,

during an Autumn sunny day without wind. P.

michaelis n. sp. seems to be an Autumn species and

is known only from the type locality in North-West-

ern Sicily, where it occurs in sintopy with P. pelle-

grinensis, the latter being however a Spring species.

The collecting locality is characterized by a stony

landscape, with sparse small trees in a disturbed

Mediterranean maquis.

Comparative notes. P. michaelis n. sp. differs

from North African Pseudoapterogyna for the fol-

lowing characters (see also Baraud, 1985, pp. 413—

415, in particular points 9 and 16): antennal club

slightly shorter than funicle, shape and setation of

pronotum (long fine setae along basal and fore mar-

gins), thin basal bead of pronotum finely and

sparsely punctate, obtuse posterior angles of prono-

tum without sinuation, posterior tibiae without

carina on dorsal side, shape of parameres.

Remarks. Distribution of the Sicilian Pseu-

doapterogyna is showed in figure 10.

They can be easily identified using the fol-

lowing key:

1. Tarsal claws with a short basal tooth

only P. michaelis

- Tarsal claws with a distinct sharp tooth just

above basal tooth 2

2. Sides of pronotum without sinuation just before

basal angle, which is obtuse and rounded. Prono-

tum without distinctly crenulate lateral margins.

Aedeagus with parameres not narrowed distally

and apex very short (lateral view) P vorax

- Sides of pronotum with a sinuation just before

basal angle, which is acute or at a right angle.

Pronotum with distinctly crenulate lateral mar-

gins. Aedaegus with parameres distally narrowed

(lateral view) 3

3. Dorsal surface shiny. Posterior tibiae with upper

face longitudinally carinate. Pygidium coarsely

wrinkled, micro-reticulated and with punctures

small and very dense; apical margin slightly

emarginate P. pellegrinensis

- Dorsal surface sub-opaque. Posterior tibiae

without carina on upper face, or with a slight

carina limited to the proximal part. Pygidium

with different surface sculpturing 4

4. Pigidial punctation made of mixed dense small

punctures and sparser larger ones on a micro-retic-

ulate background. Parameres with veiy long and

sharp apex (lateral view) P. euphytus euphytus

- Pigidial punctation made of small deep punc-

tures on a coarsely wrinkled background. Aedea-

gus with parameres with apex short (lateral

view) P. euphytus lamantiai

ACKNOWLEDGEMENTS

I am grateful to R. Poggi (Museo Civico di Storia

Naturale, Genoa, Italy), O. Montreuil and A. Man-

tilleri (Museum National d'Histoire Naturelle, Paris,

France), G. Sabella (Dipartimento di Biologia Ani-

male University of Catania, Italy), E. Piattella (Uni-

versity “Sapienza” of Rome, Italy), A. Rey (Genoa,

Italy), V. Aliquo (Palermo, Italy), Antonella Monas-

tra (Palermo, Italy). I am also grateful to M.S.

Colomba (University of Urbino, Italy), F. Liberto

366

Ignazio Sparacio

(Cefalu, Italy), M. Bellavista (Palermo, Italy), and,

particularly, to A. Ballerio (Brescia, Italy), M.

Amone (Palermo, Italy), and M. Romano (Capaci,

Italy) for support in the field and during the prepa-

ration of this paper.

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Biodiversity Journal, 2014, 5 (2): 367-370

On the distribution of some Mediterranean Cerithiopsis

Forbes et Hanley, 1 850 (Caenogastropoda Cerithiopsidae)

Luigi Romani 1 & Stefano Bartolini 2

'Via delle ville 79, 55013 Lammari, Lucca, Italy; e-mail: luigiromani78@gmail.com

2 Via E. Zacconi 16, 50137 Firenze, Italy; e-mail: stefmaria.bartolini@alice.it

’Corresponding author

ABSTRACT New records extend the distribution range of some Cerithiopsis Forbes et Hanley, 1850

(Caenogastropoda Cerithiopsidae): C. ladae Prkic et Buzzurro, 2007, C. pulchresculpta

Cachia, Mifsud et Sammut, 2004 and C. iudithae Reitano et Buzzurro, 2006.

KEY WORDS Cerithiopsis', Cerithiopsidae; distribution; Mediterranean Sea; new records.

Received 26.03.2014; accepted 19.05.2014; printed 30.06.2014

INTRODUCTION

The genus Cerithiopsis Forbes et Hanley, 1850

(Caenogastropoda Cerithiopsidae) gathers a large

pool of species consistently associated with sponges

(Marshall, 1978), it appears heterogeneous and

possibly polyphyletic (Cecalupo & Robba, 2010;

Prkic & Mariottini, 2010; Modica et al., 2013).

Currently the genus is intended conservatively

and only recently some species have been assigned

to the new genus Nanopsis Cecalupo et Robba,

2010 relying on subtle differences of the proto-

conch (Cecalupo & Robba 2010). Otherwise Scud-

eri & Criscione (2011) stated that the description

of a new genus could not depend on protoconch

features only.

In this respect more than 20 species of Cerithiop-

sis occurs in the Mediterranean Basin (Campani

2014, pers. comm.; Gofas, 2013; Gofas & Le Re-

nard, 2014), many of which have been described

during the past few years with apparently narrow

ranges. Here we provide new data on distribution

of three Cerithiopsis species recently described in

order to give a little contribution to the knowledge

of this complex group. A thorough global revision

taking into account not only shell morphology but

also anatomical and genetical features will be

necessary to reveal the actual relationships among

them.

MATERIAL AND METHODS

Cerithiopsis ladae Prkic et Buzzurro, 2007 from

lie Rousse and Livorno were collected by algal

washing. C. pulchresculpta Cachia, Mifsud et Sam-

mut, 2004 from Argentario by brushing Posidonia

rhizomes. All other specimens were picked up from

shell grit samples collected by SCUBA diving.

ACRONYMS. CA = collection P.G. Albano

(Bologna, Italy); CB = collection S. Bartolini

(Firenze, Italy); CP = collection A. Pagli (Lari,

Pisa, Italy); CRL = collection L. Romani (Lam-

mari, Lucca, Italy); CRA= collection A. Raveggi

(Firenze, Italy); CSC = collection C. Sbrana

(Livorno, Italy); CSF = collection F. Siragusa

(Livorno, Italy); sh = empty shell(s); sp: speci-

men^) collected alive.

368

Luigi Romani & Stefano Bartolini

RESULTS AND DISCUSSION

Cerithiopsis ladae Prkic et Buzzurro, 2007

Examined material. lie Rousse (Corsica, France),

30 m, 1 sp (CB); Livorno (Tuscany, Italy), 0.5 m,

1 sh (CRA); Calafuria (Livorno, Italy), 31 m, 1 sp

(CRA); Elba island (Livorno, Italy), 40 m, 1 sh

(CB); Punta Ala (Grosseto, Italy), 0.5 m, 1 sh (CB);

Palinuro (Salerno, Italy), 05-2006, 30 m, 1 sh

(CSF); Lampedusa island (Agrigento, Italy), 60 m,

1 sh (CRA); Malta, 60 m, 1 sh (CSC).

C. ladae “ribbed form”: Cannizzaro (Catania, Italy),

40 m,l sh (CB); Getares (Algeciras, Spain), 15 m,

1 sh (CB).

Remarks. The present species (Figs. 2-5) has a

dark brown, pupoid shells with a blunt cylindrical

protoconch, smooth and white. It was described

from Dalmatian coast (Prkic & Buzzurro, 2007) and

subsequently recorded from Spain (Penas et al.,

2006; Oliver, 2007; Gofas et al., 2011; Oliver et al.,

2012), Brindisi, E-Apulia (Scuderi & Terlizzi,

2012) and Eastern Sicily (Scuderi & Criscione,

2011). Our records extend its range to the Tyrrhe-

nian Sea and Strait of Sicily.

We would like to take the opportunity of signal-

ing some specimens from Cannizzaro (Sicily) and

Getares (S-Spain) with axial riblets on the lower

whorl of the protoconch (Figs. 4, 5). This feature

wasn’t reported in the original description but it was

already noticed (Prkic 2008, pers. comm.). It con-

firms that the protoconch variability in C. ladae is

wider than previously recorded, both in sculpture

and in whorls morphology.

Cerithiopsis pulchresculpta Cachia, Mifsud et

Sammut, 2004

Examined material. La Herradura (Granada,

Spain), 30 m, lsh (CRA); lie Rousse (Corsica,

France) 40 m, 2 shs (CB); Castelsardo (Sassari,

Italy), 45 m, 2 shs (CB); Calafuria (Livorno, Italy),

31 m, 3 shs (CB); Capraia island (Livorno, Italy),

70 m, 1 sh (CP); Capraia island (Livorno, Italy),

31 m, 1 sh (CRL); Giannutri island (Grosseto,

Italy), 55 m, 1 sh (CRA); Secca delle Murelle

(Viterbo, Italy), 27 m, 1 sh (CSC); Punta Cam-

panella (Naples, Italy), 50 m, 1 sh (CB), 1 sh

(CRA); Palinuro (Salerno, Italy), 50 m, 1 sh (CB);

Palinuro (Salerno, Italy), 30 m, 1 sh (CSF); Prvic

island (Krk, Croatia), 40 m, 2 shs (CB); Corfu is-

land (Greece), 15 m, 1 sh (CB); Corfu island

(Greece), 58 m, 1 sh (CRA).

Remarks. The species (Figs. 6, 7, 11, 12) has a

brown conical-pupoid shell with a cylindrical bica-

rinated protoconch, crossed by close thin and dense

axial riblets, hazelnut brown- hazelnut in colour.

The present species was described from Malta (Ca-

chia et al., 2004) and then reported from Calabria

(Vazzana, 2010) and Sicily (Scuderi & Criscione,

2011). Our records widely extend its range to the

north-central Tyrrhenian Sea, Corsica, Sardinia, NE

Adriatic, NE Ionian and S Spain.

Cerithiopsis iudithae Reitano et Buzzurro, 2006

Examined material. Calafuria (Livorno, Italy),

31 m, 1 sh (CRA); Elba island (Livorno, Italy), 40

m, 1 sh (CB); Argentario (Grosseto, Italy), 15 m,l

sh (CA); Secca delle Murelle (Viterbo, Italy), 27

m, 1 sh (CSF); Punta Campanella (Naples, Italy),

50 m, 1 sh (CB); Prvic island (Krk, Croatia), 40 m,

1 sh (CB).

Remarks. The species (Figs. 9, 10) has a brown

conical-pupoid shell with a white protoconch sculp-

tured by two spiral chords crossed by thin and

undulated axial riblets. It was described from east-

ern Sicily (Reitano & Buzzurro, 2006) and then re-

ported from Apulia (Trono & Maori, 2013). Our

records extend its range to the Tyrrhenian Sea and

NE Adriatic.

The difficulty to correctly recognise many dif-

ferent species, which share a close similar teleo-

conch and protoconch morphology, is often due to

the general status of the collected specimens. If

they are not found in perfect conditions, important

characters, as the protoconch morphology, could

not allow the identification of the materials col-

lected. Moreover the particular habitat and the

small size of all the species of this family of gas-

tropods has probably contributed to misidentifica-

tions and lack of data of many species. Putting

together all these facts, in our opinion new exami-

nations of specimens of collectors and further col-

lecting materials could lead researchers to

re-evaluate the distribution range of many species

and their real diffusion status.

On the distribution of some Mediterranean Cerithiopsis Forbes et Hanley, 1850 (Caenogastropoda Cerithiopsidae)

369

Fig. 1 . Sampling localities of Cerithiopsis iudithae (black stars), C. ladae (black circles), C. ladae “ribbed form” (black diamond),

C. pulchresculpta (black triangles). Figs. 2, 3 . C. ladae and protoconch, lie Rousse, Corsica, France, 2 mm. Figs. 4, 5. C. ladae

“ribbed fonn” and protoconch, Getares, Spain, 1.9 mm. Figs. 6, 7. C. pulchresculpta and protoconch, Calafuria, Tuscany, Italy,

2.8 mm. Figs. 8, 9. C. iudithae and protoconch, Punta Campanella, Naples, Italy, 2.9 mm. Fig. 10. C. iudithae, Prvic, Croatia,

2.8 mm. Fig. 11. C. pulchresculpta, LaHerradura, Spain, 3.2 mm. Fig. 12. C. pulchresculpta (Prvic, Croatia), 2.8 mm.

370

Luigi Romani & Stefano Bartolini

ACKNOWLEDGMENTS

We are grateful to G.P. Albano (Bologna, Italy),

A. Pagli (Lari, Pisa, Italy), S. Raveggi (Firenze,

Italy), C. Sbrana (Livorno, Italy), F. Siragusa

(Livorno, Italy), who loaned biological material,

and to T. Manousis (Epanomi, Greece), J. Prkic

(Split, Croatia), E. Quaggiotto (Vicenza, Italy) who

kindly provided us informations. We are also grate-

ful to C. Bogi (Livorno, Italy) for useful suggestions

and to E. Campani (Livorno, Italy) for reading and

improving the manuscript.

REFERENCES

Cachia C., Mifsud C. & Sammut P.M., 2004. The marine

shelled mollusca of the Maltese Islands. Part 4: The

classes Caudofoveata, Solenogastres, Bivalvia,

Scaphopoda & Cephalopoda. Leiden, Backhuys Pub-

lishers vi + 270 pp., 25 pi.

Cecalupo A. & Robba E., 2010. The identity of Murex

tubercularis Montagu, 1803 and description of one

new genus and two new species of the Cerithiopsidae

(Gastropoda: Triphoroidea). Bollettino Malacologico,

46: 45-64.

Gofas S., 2011. Familia Cerithiopsidae. In: Gofas S.,

Moreno D. & Salas C. (Eds.), Moluscos marinos de

Andalucia. Volume 1 . Malaga: Servicio de Publica-

ciones e Intercambio Cientifico, Universidad de

Malaga, pp. 152-161.

Gofas, S., 2013. Cerithiopsis Forbes & Hanley, 1850.

Accessed through: World Register of Marine Species

at http://www.marinespecies.org/aphia.php7pMaxde-

tails&id=137764 on 2014-03-12.

Gofas S. & Le Renard J. (Eds.), 2014. Cerithiopsis

Forbes & Hanley, 1850. Accessed through: CLE-

MAM: Check List of European Marine Mollusca at

http://www.somali.asso.fr/clemam/

index.clemam.html on 2014-03-12.

Marshall B.A., 1978. Cerithiopsidae (Mollusca: Gas-

tropoda) of New Zealand, and a provisional classifi-

cation of the family. New Zealand Journal of Zo-

ology, 5: 47-120.

Modica M.V., Mariottini P., Prkic J. & Oliverio M., 2013.

DNA-barcoding of sympatric species of ectoparasitic

gastropods of the genus Cerithiopsis (Mollusca: Gas-

tropoda: Cerithiopsidae) from Croatia. Journal of the

Marine Biological Association of the United King-

dom, 93: 1059-1065.

Oliver Baldovi D., 2007. Catalogo de los Gasteropodos

testaceos marinos de la parte Sur del Golfo de Valen-

cia (Espana). Iberus, 25: 29-61.

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ropodos marinos de las islas Columbretes (Mediter-

raneo occidental). Iberus, 30: 49-87.

Penas, A., Rolan, E., Luque, A. A., Templado, J., Moreno,

D., Rubio, F., Salas, C., Sierra, A., Gofas, S., 2006.

Moluscos marinos de la isla de Alboran. Iberus, 24:

25-151.

Prkic J. & Buzzurro G., 2007. Anew species of Cerithiop-

sis (Gastropoda Cerithiopsidae) from Croatian coasts.

Triton, 15: 1-4.

Prkic J. & Mariottini P., 2010. Description of two new

Cerithiopsis from the Croatian coast, with comments

on the Cerithiopsis tubercularis complex (Gas-

tropoda: Cerithiopsidae). Aldrovandia, 5: 3-27.

Reitano A. & Buzzurro G., 2006. Descrizione di una

nuova specie di Cerithiopsidae per le coste della

Sicilia orientale (Mollusca Triphoroidea). II Natu-

ralista Siciliano, 30: 549-554.

Scuderi D. & Criscione F., 2011. New ecological and ta-

xonomical data on some Ptenoglossa (Mollusca,

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Sea). Biodiversity Journal, 2: 35-48.

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dell' Alto Jonio. Grifo Ed., 188 pp.

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(Trono, 2006): corrigenda e addenda. Bollettino

Malacologico, 49: 26-48.

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Biodiversity Journal, 2014, 5 (2): 371-373

Rediscovery of the critically endangered cyprinid fish Epalze-

orhynchos bicolor (Smith, 1 93 1) from West Thailand (Cyprini-

formes Cyprinidae)

Sitthi Kulabtong 1,2 *, Siriwan Suksri 3 , Chirachai Nonpayom 4 &Yananan Soonthornkit 2

'Save wild life volunteer Thailand, Wangnoi District, Ayuttaya Province 13170, Thailand

fisheries Program, Faculty of Agro-Industrial Technology, Rajamangala University of Technology Tawan-ok Chantaburi Campus,

Chantaburi, Thailand

Reference Collection Room, Inland Fisheries Resources Research and Development Institute, Department of Fisheries, Thailand

4 5 34/26 Soi Phaholyothin 58, Phaholyothin Rd. Sai Mai, Bangkok, Thailand

’Corresponding author, email: kulabtong2011@hotmail.com

ABSTRACT In the present paper, we report on the critically endangered cyprinid fish, Epalzeorhynchos

bicolor (Smith, 1931) “rediscovered” in Maeklong Basin, West Thailand. Moreover, distribu-

tion data and biological observations of this species are also provided.

KEY WORDS Epalzeorhynchos bicolor; Cyprinidae; Maeklong Basin; Thailand.

Received 01.04.2014; accepted 18.05.2014; printed 30.06.2014

INTRODUCTION

The freshwater cyprinid fish genus Epalze-

orhynchos Bleeker, 1855, order Cypriniformes

Bleeker, 1859 and family Cyprinidae Cuvier, 1817,

has been reported for Southeast Asia only (Kottelat

& Whitten, 1996; Doi, 1997; Monkolprasit et al.,

1997; Yang & Winterbottom, 1998).

According to the current taxonomic status of

this genus, it comprises 4 valid species:

E. frenatum (Fowler, 1937) from Chao Phraya

Basin, Maeklong Basin in Thailand and Mekong

Basin in Indochina;

E. kalopterum (Bleeker, 1850) from South Thai-

land to Indonesia;

E. munense (Smith, 1934) from Mekong Basin

in Indochina;

E. bicolor (Smith, 1931) from Central and West

Thailand (Smith, 1931; Vidthayanon et al. 1997;

Kottelat, 2013).

RESULTS

The cyprinid fish Epalzeorhynchos bicolor

(Fig. 1) is an endemic fish of Thailand. The distri-

bution of this species is reported only for Lower

Chao Phraya Basin, Bangpakong Basin and

Lower Maeklong Basin (Smith, 1931; Vidthayanon

et al., 1997; Vidthayanon, 2005, 2011). It has been

threatened by mass collecting for aquarium trade,

pollution of many sources and habitats destruction

(Vidthayanon, 2011) and, according to the IUCN

Red List of Threatened Species (Vidthayanon,

2005, 2011), this fish is a threatened species. In

1996, it was even thought to be extinct in the wild

since there was no documented evidence of it the

last more than 50 years (Kottelat & Whitten,

1996).

Currently, the status of the species is poorly

known. In 201 1 , Dr. Chavalit Vidthayanon assessed

that the species is still extant in the Chao Phraya

372

SlTTHI K.ULABTONG ET ALII

Figure 1. Epalzeorhynchos bicolor from Maeklong Basin, West Thailand, standard length 66 mm.

Basin but strictly localized, nevertheless, its loca-

tion is still unclear (personal comment). On the con-

trary, in the same year, the population of E. bicolor

was reported to be extirpated in Maeklong Basin

and Bangpakong Basin (Vidthayanon, 2011).

In a survey project of the first author at Lower

Maeklong Basin, West Thailand (carried out dur-

ing February 2013) the author found only one speci-

men of E. bicolor in the rocky dam around the

mainstream of Maeklong River near the water gate

of Maeklong Dam, Muang District, Kanchanaburi

Province, Lower Maeklong Basin, West Thailand

(Fig. 2).

This fish lives in gaps between the rocks and

its habitat is characterized by large rocks and a

sandy bottom. This area is fast flown by tides and

the depth of water is more than 1 meter. In the

same area, we found many other fish species,

including:

CLUPEIF ORME S CLUPEIDAE

Clupeichthys goniognathus Bleeker, 1855

OSTEOGLOSSIFORMES NOTOPTERIDAE

Notopterus notopterus (Pallas, 1769)

C YPRINIF ORME S CYPR1NIDAE

Rasbora aurotaenia Tirant, 1885

Barbonymus schwanenfeldii (Bleeker, 1854)

Cirrhinus molitorella (Valenciennes, 1 844)

Opsarius koratensis (Smith, 1931)

Mystacoleucus marginatus (Valenciennes, 1 842),

Osteochilus vittatus (Valenciennes, 1842)

Osteochilus microcephalus (Valenciennes, 1842)

C YPRINIF ORME S BALITOLIDAE

Nemacheilus masyae Smith, 1933

Homaloptera smithi Hora, 1932

CYPRINIFORMES COBITIDAE

Acanthopsoides gracilentus (Smith, 1945)

Pangio oblonga (Valenciennes, 1 846)

SILURIF ORME S BAGRIDAE

Pseudomystus siamensis (Regan, 1913)

BELONIFORMES HEMIRAMPHIDAE

Dermogenys siamensis Fowler, 1934

SYNBRANCHIFORMES MASTACEMBELIDAE

Mastacembelus favus Hora, 1924

Rediscovery of the critically endangered cyprinid fish Epalzeorhynchos bicolor from West Thailand

373

PERCIFORMES NANDIDAE

Pristolepis fasciata (Bleelcer, 1851)

PERCIFORMES AMBASSIDAE

Parambassis siamensis (Fowler, 1937)

CONCLUSION

In conclusion, at present, the occurrence of E.

bicolor in the wild is certainly confirmed in Lower

Maeklong Basin, Kanchanaburi Province, West

Thailand, whereas it is still unclear in the Chao

Phraya Basin due to the lack, to date, of docu-

mented evidence.

ACKNOWLEDGMENTS

We wish to thank the anonymous reviewers for

their invaluable editorial advice. A very special

thank to Dr. Chavalit Vidthayanon and Mr.

Anuratana tejavej for providing available data for

this species; and to Mr. Adisorn Nonpayom and

Mr. Varin Pornrojnangkool for helping us during

the field survey.

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