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Echinoids (Echinodermata) from the Neogene of Portugal mainland:

systematics review.

Conference Paper · July 2010

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 Echinoids (Echinodermata) from the Neogene
of Portugal mainland: systematics review
Equinóides (Echinodermata) do Neogénico de Portugal continental: revisão sistemática

P. Pereira1

1 Departamento de Ciências e Tecnologia, Universidade Aberta, Palácio Ceia, R. Escola Politécnica, 141-147. 1269-001 LISBOA. PORTUGAL.
Centro de Geologia da Universidade de Lisboa, Campo Grande, 1749-016 LISBOA. PORTUGAL. pecten@univ-ab.pt

Abstract: The Portuguese Neogene echinoid fauna was reviewed in order to establish a taxonomically
standardised echinoid database. 41 taxa from the Miocene of the Lower Tagus Basin and Algarve and two
taxa from the Pliocene of Mondego Basin were described. 25 taxa were reported for the first time from
Portugal mainland, while 22 of the previously reported Miocene species were excluded from the accepted
species list.

Key words: Cidaroida. Camarodonta. Cassiduloida. Clypeasteroida. Spatangoida.

Resumo: A presença de equinóides no Neogénico de Portugal é conhecida, pelo menos, desde há cerca
de 170 anos. Durante este período de tempo, foram publicados 36 trabalhos com referências à
ocorrência deste grupo de equinodermes em Portugal continental. No entanto, apenas seis apresentam
descrições sistemáticas detalhadas e, destes, apenas dois se referem a mais de uma espécie. No total, até
2003, tinham sido descritos 42 taxa do Miocénico e uma espécie do Pliocénico. Apresenta-se a revisão
sistemática da fauna de equinóides neogénicos, combinando a análise da literatura, a revisão das
colecções existentes em museus portugueses e estrangeiros, tendo em vista o estabelecimento de uma
base de dados taxonomicamente estandardizada dos equinóides neogénicos de Portugal continental.
Deste estudo resultou a descrição de 41 taxa, entre os quais um género e uma espécie novos para a
ciência, do Miocénico da Bacia do Baixo Tejo e Algarve e dois taxa do Pliocénico da Bacia do Mondego.
Das espécies miocénicas anteriormente referidas 22 foram excluídas da lista de espécies aceites.

Palavras-chave: Cidaroida. Camarodonta. Cassiduloida. Clypeasteroida. Spatangoida.

INTRODUCTION PREVIOUS STUDIES

The occurrence of the class Echinoidea in the
Neogene of Portugal has been known at least since the
1830’s. Since then, 36 papers have mentioned the
occurrence of this echinoderm group in Portugal
mainland (Pereira, 2008). However, no more than six of
these papers present detailed systematic descriptions
(Agassiz, 1841; Michelin, 1861; de Loriol, 1896;
Zbyszewski, 1953; Ferreira, 1962; and Kotchetoff et al.,
1975) and only two of them describe (and illustrate)
more than one species: de Loriol (1896) and Kotchetoff
et al. (1975).
The systematics review of the Neogene echinoids
from Portugal mainland presented here is particularly
significant not only because of this rather small amount
of published papers concerning echinoid descriptions
but also because almost all echinoid species from the
Portuguese Neogene have been described basing on one
to three specimens and no systematics review has ever
been done.
This systematics review also provides taxonomic
standardization that is essential for large-scale
palaeobiogeographic studies.
The first palaeontological study with reference to an
echinoid genus from the Portuguese Neogene was
published by Smith (1847) in a study “on the age of the
Tertiary beds of the Tagus” where he reported the
occurrence of the genera Clypeaster, Scutella, Micraster
and Echinus.
The first reference to an echinoid species was by
Agassiz (1841) in his monograph about the living and
fossil echinoderms where he described and illustrated
the species Scutella smithiana based on an incomplete
specimen collected by James Smith from the Miocene
of Lisbon.
Twenty years later, Michelin (1861) in his
monograph about the fossil Clypeaster described and
illutrated the species Clypeaster olissiponensis based on
a single specimen collected in Miocene sediments from
the surroundings of Lisbon.
The first Portuguese publication with references to
Neogene echinoids was produced by Cotter (1879): in a
list of fossils from the Portuguese Tertiary basins he
reported the occurrence of the genera Spatangus,
Echinolampas, Clypeaster and Scutella from the Lower
Tagus Basin.
Near the end of the 19th century, de Loriol (1896) in
a monograph about the Portuguese (Lower Tagus Basin
and Algarve) Tertiary echinoderms described twenty-
nine species or varieties of Miocene echinoids, nine of
them new to the science (Arbacina mutellaensis,
Scutella lusitanica, S. roquetei, Clypeaster
palencaensis, C. mutellensis, C. delgadoi,
Echinolampas hemisphaericus var. maxima, Brissopsis
lusitanicus and Opissaster cotteri) and 20 that were
previously known in other places (Cidaris avenionensis,
Rhabdocidaris sismondai, Psammechinus dubius,
Scutella subrotunda, S. faujasii, Amphiope palpebrata,
Clypeaster crassicostatus, C. olissiponensis, C. acclivis,
C. altus, C. tauricus, C. michelotti, C. latirostris, C.
laganoides, C. marginatus, Heteroclypeus semiglobus,
Echinolampas hemisphaericus, Echinanthus
aremoricus, Schizaster scillae and Spatangus corsicus).
This monograph included a table made by Berkeley
Cotter with the lithostratigraphic distribution of the
echinoids from the Lower Tagus Basin.
From 1940 onwards, mainly due to the work of
George Zbyszewski and Veiga Ferreira, many studies
contributed a lot to the better knowledge of the diversity
and stratigraphic and palaeogeographic distribution of
the Portuguese Neogene faunas, including echinoids.
Ferreira (1961) published a record of the echinoids
from the Neogene of Portugal mainland and Azores and
Madeira Archipelagos, with a minor revision,
stratigraphic distribution and provenance of the material
of all the species cited in all previous studies. This
author reported the occurrence of 33 species in Portugal
mainland: Cyathocidaris avenionensis, Prionocidaris
sismondai, Psammechinus dubius, Arbacina
mutellensis, Echinocyamus pusillus (the first echinoid
species reported to occur in the Pliocene of Portugal),
Clypeaster crassicostatus, C. palencaensis, C. acclivis,
C. mutellensis, C. olissiponensis, C. altus, C. tauricus,
C. delgadoi, C. michelotti, C. latirostris, C. laganoides,
C. marginatus, Scutella faujasii, S. lusitanica, S.
roquetei, S. subrotunda, Amphiope palpebrata,
Echinolampas hemisphaericus, E. barcinensis, E. aff.
lovisatoi, Heteroclypeus semiglobus, Echinanthus
aremoricus, Schizaster scillae, Schizaster sp., Brissopsis
lusitanicus, Opissaster cotteri, Pericosmus latus,
Agassizia (Anisaster) mossomi and Spatangus corsicus.
The last important contribution to the knowledge of
the Neogene echinoid palaeodiversity of Portugal
mainland dates from 35 years ago. Kotchetoff et al.
(1975) collected and described several echinoids from
the Miocene of Foz da Fonte – North Penedo (Lower
Tagus Basin) that they assigned to Echinocardium
olissiponensis sp. nov., Clypeaster papilionensis sp.
nov., C. intermedius, Clypeaster sp., Prionocidaris
sismondai, Spatangus ocellatus, Pericosmus latus,
Pericosmus sp., Plagiobrissus sp. and a “grosse espèce
à test mince”.
SYSTEMATICS REVIEW
The systematics review of the Neogene echinoid
fauna included the revision of echinoid material existing
in Portuguese and foreign museum collections and the
searching and analysis of specific literature (about 400
papers and monographs).
The studied material, about 1200 specimens, belongs
to the following museums (or other institutions): Museu
Geológico (MG), Portugal; Departamento de Geologia
da Faculdade de Ciências da Universidade de Lisboa,
Portugal; Laboratório de História Natural da Batalha,
Portugal; Museu de Arqueologia e Etnografia do
Distrito de Setúbal, Portugal; Naturhistorisches Museum
Wien, Austria; The Natural History Museum, England;
Musée des Confluences (former Musée d’Histoire
Naturelle de Lyon), France; Museo Municipal
Paleontológico de Estepona, Spain.
Initial classification of the specimens was based on
character analysis. Whenever possible (depending on
the availability of suitable numbers of specimens and
their preservation) these classifications were
subsequently tested by morphometric analyses.
Potentially powerful multivariate analysis (e.g. principal
component analysis, landmark methods, or canonical
discriminant analysis) could not be employed due to the
high number of missing values in the data sets. Instead,
bivariate statistics was favoured.
The species accepted were the smallest
morphologically diagnosable groups. During this
process, several previously established species turned
out to form gradational series rather than distinct
groups. Previously employed species concepts (e.g., de
Loriol, 1896) led to the differentiation of closely related
forms based on subtle differences in size and shape.
This went so far as to recognize nearly each individual
as a separate species. An example of this is the genus
Clypeaster, with 14 species reported from the Lower
Tagus Basin.
The list of Miocene accepted species includes three
cidarids [Eucidaris zeamays (Sismonda, 1842),
Prionocidaris avenionensis (des Moulins, 1837) e P.
sismondai (Mayer, 1864)], five camarodonts
[Genocidaris catenata (Desor in Agassiz & Desor,
1846), Genocidaris sp., Monilechinus portucallensis
nov. gen. nov. sp., Psammechinus dubius dubius
(Agassiz, 1840) e Schizechinus sp.), six cassiduloids
(Pliolampas elegantula ? (Cotteau, 1883), Studeria sp.,
Hypsoclypus subpentagonalis (Gregory, 1891),
Echinolampas (E.) barcinensis Lambert, 1906,
Echinolampas (E.) hemisphaerica (Lamarck, 1816) e
Echinolampas (E.) schultzi Kroh, 2005], eleven
clypeasteroids [Clypeaster altus (Leske, 1778), C.
campanulatus (Schlotheim, 1820), C. latirostris
Michelin, 1861, C. marginatus Lamarck, 1816, C.
olisiponensis Michelin, 1861, Echinocyamus sp. 1,
Echinocyamus sp. 2, Parascutella lusitanica (de Loriol,
1896), P. smithiana (Agassiz, 1841), Parmulechinus sp.
e Amphiope bioculata (des Moulins, 1837)], and sixteen
spatangoids [Brissopsis crescentica Wright, 1855, B.
ottnangensis Hoernes, 1875, Schizobrissus sp.,
Brissidae indet., Spatangus delphinus Defrance, 1827,
Echinocardium olisiponensis Kotchetoff et al., 1975,
Hemipatagus ocellatus (Defrance, 1827), Lovenia sp.,
Pericosmus latus (Desor in Agassiz & Desor, 1847),
Pericosmus sp. 1, Opissaster cotteri de Loriol, 1896,
Schizaster eurynotus Sismonda, 1841, Schizaster ? sp.
1, Ova karreri (Laube, 1869), Agassizia algarbiensis
Ferreira, 1962 e A. zitteli Fuchs, 1883] (see Fig. 1). The
list of Pliocene taxa includes only camarodonts
(Camarodonta indet.) and one clypeasteroid
[Echinocyamus pusillus ? (Müller, 1776)] (see Fig. 1).
The list of rejected echinoid species from the
Neogene of Portugal comprises 22 species, which were
considered misidentifications or junior synonyms of
other species.
CONCLUSIONS
The study of the Portuguese Neogene echinoids has
been neglected for many years. In fact, the last (and
only) serious systematics study of the Portuguese
Neogene echinoid fauna dated from late nineteenth
century (de Loriol, 1896).
After more than one century, the author (Pereira,
2008) presents a taxonomically standardised echinoid
database: 41 taxa from the Miocene of the Lower Tagus
Basin and Algarve and two taxa from the Pliocene of
Mondego Basin. Among these 43 taxa, it was described
one new genus and one new species and 24 (23 Miocene
and one Pliocene) other taxa were reported for the first
time from the Neogene of Portugal mainland. From the
previously reported Miocene species, 22 were excluded
from the accepted species list.
ACKNOWLEDGEMENTS
The author wishes to acknowledge the Portuguese
and foreign museum keepers that provided access to the
collections under their care. The first visits to the
Natural History museums of London and Vienna were
financed by grants of the Calouste Gulbenkian
Foundation and SYNTHESYS, respectively.
REFERENCES
Agassiz, L. (1841). Monographie d'echinodermes
vivants et fossiles. Échinites. Famillie des
Clypéasteroides. Seconde Monographie. Des
Scutelles. Neuchatel, 149 pp., 32 pls.
Cotter, J.C.B. (1879). Contribuições para o
conhecimento da Fauna terciária de Portugal –
Fósseis das Bacias terciárias marinas do Tejo, do
Sado e do Algarve. Jornal de Sciencias
Mathematicas, Physicas e Naturaes, 26, 11 pp.
Ferreira, O.V. (1961). Equinídeos do Miocénico de
Portugal continental e ilhas adjacentes.
Comunicações dos Serviços Geológicos de Portugal,
45: 529-564. 17 pls.
Ferreira, O.V. (1962). Nota sobre a presença do género
“Agassizia” no Miocénico do Algarve.
Comunicações dos Serviços Geológicos de Portugal,
46: 293-295, 1 pl.
Kotchetoff, B; Kotchetoff, Y. and Ferreira, O.V. (1975).
Contribution à la connaissance des gisement
fossilifères miocènes au Nord du Cap d’Espichel.
Comunicações dos Serviços Geológicos de Portugal,
59: 59-106, 8 pls.
de Loriol, P. (1896). Description des Echinodermes
tertiaires du Portugal [Accompagnée d’un tableau
stratigraphique par J. C. Berkeley Cotter]. Mémoires
de la Direction des Traveaux Géologiques du
Portugal, 50 pp., 13 pls.
Michelin, M.H. (1861). Monographie de Clypéastres
fossiles. Mémoires de la Société Géologique de
France, Paris, Série 2, 7(1): 101-147, 36 pls.
Pereira, P. (2008). Echinoids from the Neogene of
Portugal mainland: Systematics, Palaeoecology,
Palaeobiogeography. Ph.D. thesis, Univ. of Lisbon,
200 pp., 50 pls.
Smith, J. (1847). On the age of the Tertiary beds of the
Tagus, with a catalogue of the fossils. The Quaterly
Journal of the Geological Society of London,
London, 3: 410-423, 6 pls.
Zbyszewski, G. (1953). Notícia explicativa da Folha 31-
A (Santarém) da Carta Geológica de Portugal na
escala 1:50.000. Serviços Geológicos de Portugal,
16 pp.
 FIGURA 1. Characteristic echinoids from the Miocene of Portugal. a-c: Psammechinus dubius dubius (Agassiz, 1840), MG 3383c; d: Echinolampas
(Echinolampas) hemisphaericus (Lamarck, 1816), MG 3807; e: C. olisiponensis Michelin, 1864, MG 3409; f: Clypeaster marginatus Lamarck, 1816,
MG 3404; g: Parascutella smithiana (Agassiz, 1841), MG 3416; h: P. lusitanica (de Loriol, 1896), MG 3768; i: Amphiope bioculata (des Moulins,
1837), MG 3417; j: Schizaster eurynotus Sismonda, 1841, MG 3472; k-l: Agassizia algarbiensis Ferreira, 1962, MG 3793a; m-n: Echinocardium
olisiponensis Kotchetoff et al., 1975, MG 3795. Each white (or grey) bar of the scale bars equals 5 mm.

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