The bivalve Neithea from the Cretaceous of Brazil

Abstract

On the basis of new collections from the Sergipe and Camamu (Bahia) basins, revision of previously described material from the Pernambuco–Paraı́ba Basin and a reassessment of previous descriptions, five species of the pectinid bivalve Neithea are described from the Cretaceous of northeastern Brazil: N. (N.) alpina (d'Orbigny, 1847) from the Albian of the Camamu Basin; N. (N.) coquandi (Peron, 1877) from the Aptian–Cenomanian of the Sergipe Basin, the Albian of the Camamu Basin, broadly mid-Cretaceous beds of the Tucano Sul Basin (Bahia), and the Cenomanian of the São Luı́s Basin (Maranhão); N. (N.) hispanica (d'Orbigny, 1850) from the Albian–lower Turonian of the Sergipe Basin; N. (N.) bexarensis (Stephenson, 1941) from the Campanian of the Pernambuco–Paraı́ba Basin; N. (Neithella) notabilis (Münster in Goldfuss, 1833) from the Cenomanian of the Sergipe Basin. All species show a wide geographical distribution, in sharp contrast to previous studies that have indicated a highly endemic mollusc fauna in the Cretaceous of Brazil.

Full text

The bivalve Neithea from the Cretaceous of Brazil
Edilma de Jesus Andrade
a,
*, Jens Seeling
a
, Peter Bengtson
a
, Wagner Souza-Lima
b
a
Geologisch-Pala
¨ontologisches Institut der Universita
¨t Heidelberg, Im Neuenheimer Feld 234, DE-69120 Heidelberg, Germany
b
Fundac¸a
˜o Paleontolo
´gica Phoenix, Rua Geraldo Menezes de Carvalho, 218, Bairro Suissa, 49050-750 Aracaju-SE, Brazil
Abstract
On the basis of new collections from the Sergipe and Camamu (Bahia) basins, revision of previously described material from the
Pernambuco–Paraı
´ba Basin and a reassessment of previous descriptions, five species of the pectinid bivalve Neithea are described from the
Cretaceous of northeastern Brazil: N.(N.) alpina (d’Orbigny, 1847) from the Albian of the Camamu Basin; N. (N.)coquandi (Peron, 1877)
from the Aptian–Cenomanian of the Sergipe Basin, the Albian of the Camamu Basin, broadly mid-Cretaceous beds of the Tucano Sul Basin
(Bahia), and the Cenomanian of the Sa
˜o Luı
´s Basin (Maranha
˜o); N. (N.)hispanica (d’Orbigny, 1850) from the Albian –lower Turonian of the
Sergipe Basin; N. (N.)bexarensis (Stephenson, 1941) from the Campanian of the Pernambuco – Paraı
´ba Basin; N. (Neithella)notabilis
(Mu
¨nster in Goldfuss, 1833) from the Cenomanian of the Sergipe Basin. All species show a wide geographical distribution, in sharp contrast
to previous studies that have indicated a highly endemic mollusc fauna in the Cretaceous of Brazil.
q2004 Elsevier Ltd. All rights reserved.
Keywords: Aptian; Albian; Cenomanian; Turonian; Campanian; Cretaceous; Neithea; Pectinidae; Bivalvia; Sergipe; Camamu; Bahia; Pernambuco; Paraı
´ba;
Brazil; Taxonomy; Paleobiogeography
1. Introduction
The marine Cretaceous deposits exposed in the sedi-
mentary basins along the Brazilian continental margin
contain a rich macroinvertebrate fauna dominated by
molluscs. Ammonites and bivalves are generally the most
common and diverse groups. Among the bivalves, repre-
sentatives of the pectinid Neithea are abundant, commonly
well-preserved and dominate the fauna at certain levels, e.g.
in the Albian of the Sergipe and Camamu basins in
northeastern Brazil (Fig. 1). Systematic descriptions have
been carried out by White (1887), Maury (1925, 1937),
Oliveira (1939), Klein and Ferreira (1979), Brito (1967) and
Muniz (1993), but are now in need of revision. As is the case
with most of the earlier descriptions of fossil molluscs from
Brazil, many of these forms were originally described as
new species, subspecies or varieties, which gives the
impression of a highly endemic fauna. This reflects the
typological taxonomic concepts at the time of these studies
but is also, to some extent, the result of oversplitting. More
recent studies have shown that the Cretaceous of Brazil
contains few endemic mollusc species (e.g. Bengtson, 1996;
Seeling and Bengtson, 1999; this study). In our new material
and as a result of our reassessment of previously published
determinations, four species of the nominal subgenus and
one species of the subgenus Neithella are recognized in
Brazil, most of which have a wide geographical distribution.
White (1887) described Neithea quadricostata
(J. Sowerby, 1814) and Neithea sergipensis n. sp. from
Sergipe. In her revision of White’s (1887) material, Maury
(1937) maintained the latter species (here synonymized with
N. hispanica (d’Orbigny)) and split N. quadricostata into
the new subspecies N. quadricostata lastroensis and
N. quadricostata regoi, based primarily on new collections.
Duarte (1938) further reported N. quadricostata from the
same beds. Subsequently, Beurlen (1964a) referred these
forms to Neithea shawi Pervinquie
`re, 1912. We follow the
view of Dhondt (1973), who regarded N. shawi as a junior
synonym of Neithea (N.)coquandi (Peron). We also place
N. sergipensis, described from the Tucano Sul Basin in
Bahia (Oliveira, 1939), and N. (Neitheops)nana Ferreira
and Klein, 1979, from the Sa
˜oLuı
´s Basin in Maranha
˜o, in
synonymy with N. (N.)coquandi.
From the Camamu Basin in Bahia, Maury (1925)
described Pecten cf. quinquecostatus J. Sowerby, 1814.
Brito (1964) reported the genus Neithea, which he sub-
sequently described as N. bahiaensis n. sp. (Brito, 1967).
0895-9811/$ - see front matter q2004 Elsevier Ltd. All rights reserved.
doi:10.1016/j.jsames.2004.05.006
Journal of South American Earth Sciences 17 (2004) 25–38
www.elsevier.com/locate/jsames
*Corresponding author. Tel.: þ49-6221-546059; fax: þ49-6221-
545408.
E-mail addresses: edilma.andrade@urz.uni-hd.de (E. de Jesus
Andrade), jens.seeling@uni-hd.de (J. Seeling), bengtson@uni-hd.de
(P. Bengtson), wagnerl@hotmail.com (W. Souza-Lima).

Fig. 1. (a,b) Location of the Sergipe and Camamu basins in relation to other continental margin basins in northeastern and northern Brazil. Abbreviations of state names: AL, Alagoas; BA, Bahia; CE, Ceara
´; MA,
Maranha
˜o; PB, Paraı
´ba; PE, Pernambuco; PI, Piauı
´; RN, Rio Grande do Norte; SE, Sergipe. (c) Localities sampled in the Sergipe Basin. (d) Localities sampled in the Camamu Basin.
E. de Jesus Andrade et al. / Journal of South American Earth Sciences 17 (2004) 25–3826

Here, we refer most of the Neithea from the Camamu Basin to
N. (N.)alpina (d’Orbigny), with N.(N.) coquandi found at
only one locality.
Beurlen and Cobra (1960) reported N. sergipensis White
from Tamandare
´(near Estiva) and Gameleira (Ipojuca
district) in the Cabo Basin, south of the Pernambuco–
Paraı
´ba Basin. On the basis of this occurrence, the two
localities were correlated stratigraphically with the Albian
Riachuelo Formation in Sergipe. However, their species
assignment and correlation must be regarded as doubtful,
because: (1) Beurlen and Cobra (1960) did not provide
illustrations of the specimens; (2) Maury (1937) had
reported N. sergipensis not only from the Albian but also
from the ‘lower Turonian’ of Sergipe (probably uppermost
Cenomanian: Bengtson, unpublished results), as confirmed
by the stratigraphical range of the senior synonym N.(N.)
hispanica in our study; (3) the deposits at Gameleira were
subsequently dated as Turonian by Muniz and Almeida
(1988). From near the Estiva locality, on the banks of the
Ariquinda
´River, Beurlen (1964b) reported small specimens
of Neithea, without attempting any species assignment.
From Campanian beds near Alhandra (Paraı
´ba) in the
Pernambuco– Paraı
´ba Basin, Muniz (1993) described the
new species N. (N.)latericostata, which is here restudied
and synonymized with N. (N.)bexarensis (Stephenson,
1941).
The aim of our study is to describe and discuss the
occurrences of the bivalve Neithea in the Sergipe and
Camamu basins on the basis of new collections, restudy of
the material described by Muniz (1993) from the Pernam-
buco–Paraı
´ba Basin, and a reassessment of previous
descriptions and reports of the genus from Brazil. We also
discuss the stratigraphical range, paleobiogeographical
affinities, and ecological implications.
2. Geographical and geological setting
The Sergipe and Camamu basins, from where the new
material described here derives, are situated in the eastern,
coastal parts of Sergipe and Bahia in northeastern Brazil
(Fig. 1). They belong to the string of tensional rift basins
bordering the South Atlantic Ocean that formed as a result
of the late Mesozoic rifting and drifting apart of South
America and Africa. The sedimentary fill, which comprises
a Paleozoic– Lower Cretaceous non-marine and an Aptian –
Upper Neogene marine succession, is most complete and
extensively exposed in the Sergipe Basin. The geological
evolution and development of the marine Cretaceous along
the Brazilian continental margin, and the stratigraphy of the
Sergipe and Camamu basins in particular, have been
discussed by several authors. For more detailed information,
see Ojeda and Fugita (1976), Ojeda (1982), Bengtson
(1983), Chang et al. (1988), Koutsoukos et al. (1993), Feijo
´
(1995), Netto et al. (1995) and Berthou et al. (1996).
3. Material
The material for this study comprises 337 specimens
from 48 localities in the Sergipe Basin, 134 specimens from
eight localities in the Camamu Basin and 11 specimens from
one locality in the Pernambuco – Paraı
´ba Basin (Fig. 1). The
Aptian– Albian localities of Sergipe (Riachuelo Formation)
are described by Souza-Lima and Bengtson (in press), the
Cenomanian– Turonian localities (Cotinguiba Formation)
by Bengtson (1983, Appendix 1). A new locality in the
Cotinguiba Formation (Jardim 31) is described here
together with the eight localities collected in the Camamu
Basin (see Appendix A).
Previously studied material by White (1887) and Maury
(1937) from Sergipe, by Oliveira (1939) from the Tucano
Sul Basin, and by Ferreira and Klein (in Klein and Ferreira,
1979) from the Sa
˜o Luı
´s Basin is reassessed on the basis of
published descriptions and illustrations.
The new specimens are housed in the collections of the
Geologisch-Pala
¨ontologisches Institut, Universita
¨t Heidel-
berg, Germany (GPIH-BR) and the Fundac¸a
˜o Paleontolo
´-
gica Phoenix, Aracaju, Brazil (FPH). The specimens of
Oliveira (1939) are kept in the Departamento Nacional da
Produc¸a
˜o Mineral, those of Brito (1967) and Klein and
Ferreira (1979) in the Museu Nacional, both in Rio de
Janeiro, Brazil, and those of Muniz (1993) in the collections
of the Instituto de Geocie
ˆncias, Universidade Federal de
Pernambuco, Recife, Brazil (DG-CT-UFPE).
4. Stratigraphy
Koutsoukos and Bengtson (1993) established a
biostratigraphical scheme for the Aptian – Maastrichtian
of the Sergipe Basin on the basis of foraminifers and
ammonites. Their ammonite zonation provides the frame-
work for the stratigraphical distribution of the Neithea
species described from the basin. The oldest species is
N. (N.)coquandi, which first appears in the upper Aptian
Epicheloniceras–Diadochoceras –Eodouvilleiceras Zone
and extends into the Cenomanian. The occurrence of
N. (N.)coquandi [¼‘N. (Neitheops)nana’] in the Sa
˜o
Luı
´s Basin was dated by Klein and Ferreira (1979) as
broadly Cenomanian, whereas reliable dating of the
species from the Tucano Sul Basin [ ¼‘N. sergipensis’]
(Oliveira, 1939) must await more detailed field investi-
gations. N. (N.)hispanica is recorded from the upper
Albian to lower Turonian of Sergipe (local Mortoniceras
to Watinoceras amudariense –Kamerunoceras seitzi
zones). N. (Neithella)notabilis is restricted to the
Cenomanian of Sergipe, where it occurs from the lower
middle Cenomanian Acompsoceras spathi –Dunvegano-
ceras Zone to the lower upper Cenomanian Pseudocaly-
coceras harpax –Thomelites aff. sornayi Zone (Fig. 2). In
the Camamu Basin N.(N.) alpina and N.(N.) coquandi
occur in the Algodo
˜es Formation which, according to
E. de Jesus Andrade et al. / Journal of South American Earth Sciences 17 (2004) 25–38 27

Netto et al. (1995), is Albian to Turonian. Support for a
late Albian age of the beds in the Algodo
˜es area is
provided by the ammonite Elobiceras bahiaense (Maury,
1925)(Bengtson, 1996; Berthou et al., 1996). N. (N.)
bexarensis [¼‘N. (N.)latericostata’] from the Campa-
nian of the Pernambuco – Paraı
´ba Basin (Muniz, 1993)is
the youngest known occurrence of Neithea in Brazil.
5. Paleobiogeography and paleoecology
The Neithea faunas of Brazil are dominated by typically
Tethyan taxa such as N. (N.) hispanica,N. (N.) coquandi,
and N. (N.) alpina.N. (Neithella)notabilis is also common
in boreal regions of Europe and Japan and probably has a
cosmopolitan distribution. N. (N.) bexarensis is as yet only
known from the Campanian of the Pernambuco – Paraı
´ba
Basin and the Corsicana Formation of Texas (Maastrichtian;
Kennedy and Cobban, 1993). In general, the Brazilian taxa
are also characteristic in the upper Lower to Upper
Cretaceous of southern Europe (e.g. France, Austria,
Spain), the eastern Mediterranean (e.g. Jordan, Lebanon,
Israel, Turkey), northern and northwestern Africa (e.g.
Egypt, Libya, Algeria, Morocco), western Africa (Angola),
the North American Western Interior, the Caribbean and
northwestern South America (Peru) (Fig. 3). Close biogeo-
graphical relationships between Brazil and south Tethyan
regions during the Late Cretaceous, as shown by Neithea,
are confirmed by other benthic macrofossil groups, such as
Cenomanian oysters (Seeling and Bengtson, 1999) and
echinoids (Smith and Bengtson, 1991; Smith, 1992).
The conspicuous dominance of N. (N.)alpina in the
Camamu Basin, coupled with its absence from the nearby
Sergipe Basin poses a biogeographical problem. Collecting
failure can be ruled out, because the Albian beds of both
basins have been extensively collected. A ‘stratigraphical’
explanation, i.e. that corresponding beds are either absent or
not exposed, is also unlikely, because the Camamu material
appears to derive from various levels of the Albian. A purely
biogeographical explanation would imply a physical,
geographical, or ecological barrier separating the two basins
during the Albian.
Detailed study of the biogeographical patterns for
other fossil groups may cast light on this possibility.
Thus far, the only group that has been studied
extensively in both basins are the echinoids (Manso,
2003), which do not display any remarkable differences
in taxonomic composition. An anomalous situation is
noted, however, for the ammonites. In the Camamu
Basin ammonites are extremely rare, with only two
specimens found to date, the Elobiceras bahiaense of
Maury (1925) and a possible Yeharaites (Souza-Lima,
2003). By contrast, in the Sergipe basin, ammonites are
well represented, although of low diversity (Bengtson,
1996). This difference is difficult to explain for a group
as mobile as the ammonites unless a biogeographical
barrier is envisaged. With respect to the differences in
taxonomic composition of the Neithea faunas of the two
basins, a barrier may provide a plausible explanation,
although an alternative explanation might also be
sought in differing ecological conditions. For example,
N.(N.) alpina and N.(N.) coquandi may have been
adapted to the same or similar ecological niches, which
could have led to mutually exclusive competition that
favored N.(N.) alpina in Camamu and N.(N.) coquandi
in Sergipe.
In Brazil, Neithea is found mainly in carbonate rocks
representing shallow-marine environments. All species are
recliners that rested on soft substrates or with their convex
right valve partly buried, without being attached (‘iceberg
strategists’, e.g. Stanley, 1970; Jablonski and Bottjer,
1983). Some of the Neithea occurrences in Sergipe appear
to represent transported faunas. In most species, the right
valve is strongly convex and more durable than the flat to
slightly concave left valve, which is smaller, thinner, and
more fragile. A Neithea community buried in situ or
transported only a short distance would consist of
Fig. 2. Stratigraphical distribution of Neithea species in the Sergipe Basin.
Standard zones and local ammonite zones after Koutsoukos and Bengtson
(1993) and Walter and Bengtson (1998).
E. de Jesus Andrade et al. / Journal of South American Earth Sciences 17 (2004) 25–3828

approximately equal numbers of left and right valves.
Longer transport would remove or destroy the lighter, more
fragile left valves and deposit the heavier right valves at a
distance from the original site. The smaller number of left
valves found in some assemblages can be attributed to such
a process.
The degree of disarticulation, along with abrasion and
breakage, is a reliable indication of the degree of transport a
bivalve has undergone since its death (e.g. Boucot et al.,
1958). Most of the Brazilian Neithea shells are disarticu-
lated, although the degree of abrasion and breakage is
generally low. This condition suggests transport in a high-
energy regime, though of short duration.
6. Systematic paleontology
Family Pectinidae Rafinesque, 1815
Subfamily Neitheinae Sobetski, 1960
Genus Neithea Drouet, 1824
6.1. Subgenus Neithea Drouet, 1824
Diagnosis
Shell symmetrical, right valve with six primary ribs and a
variable number of secondary ribs in each interspace; left
valve nearly flat or slightly concave; auricles subequal or
unequal (revised after Hayami, 1965, and Dhondt, 1973).
6.1.1. Neithea (Neithea) coquandi (Peron, 1877)
(Fig. 4.1–4.5)
1862 Janira tricostata H. Coq.; Coquand, pp. 219, 220,
Pl. 13, Figs. 3 and 4.
1877 Janira coquandi Peron; Peron, pp. 501 – 509, Pl. 7,
Fig. 2.
1887 Neithea quadricostata Sowerby; White, pp. 37 – 39,
Pl. 4, Figs. 1 and 2.
1888 Janira welwitschi Choffat; Choffat, pp. 90, 91, Pl.
5, Fig. 11.
1912 Pecten (Neithea)shawi Pervinquie
`re; Pervinquie
`re,
pp. 136–138, Pl. 9, Figs. 1 – 6.
1925 Pecten cf. quinquecostatus Sowerby; Maury, pp.
558–561.
1937 Neithea quadricostata lastroensis, subsp. nov.;
Maury, pp. 138– 141, Pl. 5, Figs. 1, 16 – 18.
1937 Neithea quadricostata regoi, subsp. nov.; Maury,
pp. 140–143, Pl. 5, Fig. 14.
1939 Neithea sergipensis White; Oliveira, p. 80, Fig. 9.
1973 Neithea (Neithea)coquandi (Peron, 1877); Dhondt,
pp. 26–29, Pl. 3, Fig. 1 (with extensive synonymy).
Fig. 3. Geographical distribution of the Brazilian species of Neithea (after Dhondt, 1973, 1981, 1982, 1992; this paper). Map for the mid-Cretaceous based on
Barron (1987), Funnell (1990), and Scotese (2002).
E. de Jesus Andrade et al. / Journal of South American Earth Sciences 17 (2004) 25–38 29

Fig. 4. (4.1–4.5) Neithea (Neithea)coquandi (Peron): (4.1) right valve, (4.2) lateral view of right valve, FPH 1059-I, (4.3) left valve, FPH 1063-I, from
Riachuelo 1, lower Albian, (4.4) right valve, FPH 1325-I, from Sa
˜o Joaquim 4, lower Albian, (4.5) right valve, FPH 1390-I, from Porto dos Barcos 2, upper
Aptian. (4.6–4.12) Neithea (Neithea)alpina (d’Orbigny): (4.6) right valve, FPH 1391-I, from Boipeba 2, Albian, (4.7) right valve, FPH 1329-I, (4.8) right
valve, (4.9) lateral view of right valve, FPH 1332-I, (4.10) right valve, FPH 1341-I, from Cangaı
´ba 2, Albian, (4.11) right valve, FPH 1392-I, from Cangaı
´ba 1,
Albian, (4.12) right valve, FPH 1243-I, from Boipeba 3, Albian, (4.13 –4.17) Neithea (Neithea)hispanica (d’Orbigny): (4.13) right valve, GPIH-BR C507.N2,
from Japaratuba 11, lower Turonian, (4.14) right valve, GPIH-BR C618.10b, from Japaratuba 14, upper Cenomanian, (4.15) right valve, GPIH-BR C385.4,
from Jerico
´4, middle or upper Cenomanian, (4.16) left valve, GPIH-BR C375.51, from Cruzes 9, middle or upper Cenomanian, (4.17) right valve, GPIH-BR
C618.10a, from Japaratuba 14, upper Cenomanian. (4.18– 4.20) Neithea (Neithea)bexarensis Stephenson, Campanian: (4.18) right valve, DG-CT-UFPE 2841
(holotype of Neithea (Neithea)latericostata Muniz, 1993), (4.19) right valve, DG-CT-UFPE 2842, (4.20) right valve, DG-CT-UFPE 2853, all from near
Alhandra, Paraı
´ba. (4.21– 4.26) Neithea (Neithella)notabilis (Mu
¨nster in Goldfuss): (4.21) right valve, GPIH-BR C513.2, from Porto da Cruz 1, middle or
upper Cenomanian, (4.22) right valve, GPIH-BR C514.2, from Tibu
´rcio 1, upper (?) Cenomanian, (4.23) right valve, GPIH-BR C239.20, from Mosquito 1,
middle Cenomanian, (4.24) right valve, (4.25) left valve, (4.26) lateral view of right valve, GPIH-BR C514.6, from Tibu
´rcio 1, upper (?) Cenomanian. All
specimens £1.5, except 4.8, 4.9 £2, and 4.7, 4.11, 4.22, 4.23 £3.
E. de Jesus Andrade et al. / Journal of South American Earth Sciences 17 (2004) 25–3830

1979 Neithea (Neitheops)nana Ferreira et Klein sp. nov;
Klein and Ferreira, p. 526, Fig. 4.
Material
Four specimens from one locality in the Camamu Basin:
Algodo
˜es 3 (FPH 1375-I, 1381-I, 1382-I, 1384-I). 296
specimens from 27 localities in the Sergipe Basin: Boa Hora
2 (FPH 1189-I-91-I); Catete Velho 1 (FPH 893-I-894-I);
Coqueiro 1 (FPH 895-I-898-I); Engenho Lira 1 (FPH 1194-
I-1201-I), 2 (FPH 899-I-925-I, 927-I-929-I, 931-I-934-I);
Jerico
´2 (GPIH-BR C382.1a, 1b, 2, 3), 7 (GPIH-BR C383.1-
2, 3a, 3b, 4), 8 (GPIH-BR C387.1); Jorda
˜o 12 (FPH 939-I);
Limeira 3 (FPH 940-I-944-I), 4 (FPH 945-I), 5 (FPH 946-I-
950-I, 1326-I), 6 (FPH 981-I-986-I), 7 (GPIH-BR R34.1);
Maruim 1 (FPH 951-I-952-I); Pedra Branca 20 (FPH 953-I-
954-I), 21 (GPIH-BR R51.14); Porto dos Barcos 1 (FPH
955-I-965-I), 2 (FPH 960-I-971-I, 1206-I-1208-I, 1390-I);
Praia 9 (GPIH-BR C339.22, C339b.4, 8, 22), 13 (FPH 1187-
I-1188-I); Riachuelo 1 (FPH 972-I – 977-I, 933-I –934-I,
996-I–1005-I, 1008-I, 1010-I, 1012-I – 1090-I, 1092-I,
1095-I–1108, 1110-I–1115-I, 1117-I, 1121-I, 1122-I,
1124-I, 1125-I, 1126-I, 1129-I – 1133-I, 1136-I – 1141-I,
1143-I–1146-I, 1148-I – 1174-I), 2 (FPH 1182-I – 1185-I);
Sa
˜o Joaquim 4 (FPH 1325-I); Sı
´tio Novo 4 (FPH 1203-I–
1205-I); Usina Central 12 (FPH 1211-I– 1212-I); Timbo
´9
(GPIH-BR R114.2).
Description and discussion
Most specimens are medium-sized (height 20 – 40 mm,
length 19–30 mm). The largest specimen (FPH-1188-I) has
a height of 53 mm and a length of 40 mm. There are also a
number of smaller, juvenile individuals (Table 1). The
species is characterized by having three unequal secondary
ribs intercalated between each pair of principal ribs
(Fig. 4.1), with the middle secondary rib more strongly
developed (Fig. 4.5). This arrangement can be seen in the
majority of our specimens; however, in some specimens
there are only two and in others as many as four secondary
ribs between any two principal ribs. The distribution of
secondary ribs in these specimens then appears as follows:
either (i) four secondary ribs between the first two principals,
(ii) four secondary ribs between the last two, (iii) four
secondary ribs between the first two and the last two
principals, or (iv) only two secondary ribs between any two
principal ribs (Fig. 4.4). However, the variation usually
affects only a single pair of principal ribs; that is, in these
specimens the number of secondary ribs between two
principal ribs is normally three.
Zucon et al. (1998) examined the variability of ribbing
in specimens collected from the locality Riachuelo 1 and
distinguished 10 morphotypes. The dominant morphotype
(two-thirds of all specimens) carries three secondary ribs
between each pair of principal ribs, and the second most
common morphotype (one-sixth of all specimens) has
four secondary ribs in one of the interspaces. The other
eight morphotypes are represented by a few specimens
only. Our material from Riachuelo 1 displays similar
proportions among the different morphotypes.
Some of our specimens differ from typical North
African representatives of N. (N.) coquandi in having
larger auricles (A.V. Dhondt, Brussels, personal com-
munication, September 2002). However, in all other
respects these specimens are typical N. (N.) coquandi.
Beurlen (1964a) assigned the specimens from Sergipe
described by White (1887) and Maury (1937) as
N. quadricostata to N. shawi Pervinquie
`re, 1912, and
mentioned their similarity to N. welwitschi (Choffat,
1888), probably on the basis of their strongly developed
middle secondary ribs.
Three specimens of Neithea from two localities (Santa
Cruz and near Inhambupe) in the Tucano Sul Basin, Bahia
collected by Melo Junior (1939) were described by Oliveira
(1939) as N. sergipensis. However, judging from his
description and illustration these specimens are not refer-
able to N.(N.) hispanica, as are the types of N.sergipensis
from Sergipe, but agree in all respects with N.(N.)
coquandi.
Neithea (Neitheops)nana Ferreira and Klein (in Klein
and Ferreira, 1979) from the Sa
˜o Luı
´s Basin, was based on
few, poorly preserved specimens. The description and
illustration fit the concept of N. (N.) coquandi and the two
Table 1
Dimensions of selected, representative specimens of N. (N.) coquandi
Specimen number Height (mm) Length (mm)
FPH 896-I 24 21
FPH 897-I 21 14
FPH 898-I 13 9
FPH 901-I 35 31
FPH 902-I 32 27
FPH 905-I 16 13
FPH 906-I 18 14
FPH 931-I (left valve) 9 8
FPH 937-I 18 17
FPH 941-I 26 23
FPH 956-I 11 9
FPH 973-I 21 17
FPH 976-I 33 30
FPH 993-I 41 36
FPH 996-I 24 19
FPH 997-I 22 18
FPH 1005-I 9 8
FPH 1016-I (left valve) 15 14
FPH 1124-I 6 4
FPH 1174-I 25 21
FPH 1182-I 15 14
FPH 1188-I 53 40
FPH 1194-I 24 19
FPH 1197-I 11 9
FPH 1198-I 9 7
FPH 1211-I 28 20
FPH 1325-I 19 16
FPH 1375-I (left valve) 10 10
FPH 1381-I 12 10
FPH 1384-I 13 11
E. de Jesus Andrade et al. / Journal of South American Earth Sciences 17 (2004) 25–38 31

species are here synonymized. N. (N.) coquandi differs from
N. (N.) alpina from the Albian of the Camamu Basin in
having three, commonly unequal secondary ribs, with the
middle one most strongly developed.
There has been nomenclatorial confusion about N. (N.)
coquandi, starting from the first description as Janira
tricostata by Coquand (1862). For a discussion of the
problem, see Dhondt (1973, p. 28).
Occurrence
Widely distributed in the Albian – Santonian of southern
Europe, the Middle East and northern and western Africa
(e.g. Austria, France, Spain, Algeria, Egypt, Iraq, Israel,
Lebanon, Somalia, Angola); upper Aptian – Cenomanian of
the Sergipe, Camamu, and Sa
˜o Luı
´s basins and indetermi-
nate middle Cretaceous (?) of the Tucano Sul Basin, Brazil.
6.1.2. Neithea (Neithea) alpina (d’Orbigny, 1847)
(Fig. 4.6–4.12)
1847 Janira alpina d’Orbigny, 1846 [sic]; d’Orbigny, pp.
643, 644, Pl. 446, Figs. 4–8.
1855 Neithea occidentalis; Conrad, p. 269.
1852 Pecten quadricostatus var.; Roemer, p. 64, Pl. 8,
Fig. 4a–c.
1888 Janira ficalhoi, Choffat; Choffat, pp. 89, 90, Pl. 5,
Figs. 8–10.
1928 Pecten (Neithea)irregularis Bo
¨se 1910; Adkins,
p. 126, Pl. 17, Fig. 8.
1928 Pecten (Neithea)occidentalis (Conrad); Adkins,
p. 126.
1947 Pecten (Neithea)occidentalis (Conrad); Stanton,
pp. 43, 44, Pl. 27, Figs. 1–4.
1960 Pecten (Neithea)occidentalis (Conrad); Perkins,
pp. 65, 66, Pl. 17, Figs. 22 and 23.
1965 Neithea (Neithea)ficalhoi (Choffat); Hayami,
pp. 302–305, Pl. 42, Figs. 5 and 16.
1967 Neithea bahiaensis n. sp.; Brito, pp. 5 – 7, Pl. 1,
Figs. 1–6.
1973 Neithea (Neithea)alpina (A. d’Orbigny, 1847);
Dhondt, pp. 15– 20 (with extensive synonymy).
Material
130 specimens from 7 localities in the Camamu Basin:
Boipeba 1 (FPH 1238-I – 1240-I), 2 (FPH 1280-I, 1391-I), 3
(FPH 1241-I– 1244-I, 1283-I – 1285-I); Cangaı
´ba 1 (FPH
1245-I– 1279-I, 1286-I – 1315-I, 1317-I – 1324-I, 1388-I,
1392-I), 2 (FPH 1328-I – 1342-I, 1345-I – 1366-I); Taipus 3
(FPH 1327-I, 1389-I, 1393-I); Tatus 1 (FPH 1281-I –1382-I).
Description and discussion
Most specimens are small (height 11–25 mm, length
8– 21 mm) with only a few medium-sized individuals
(Table 2). The right, convex valve is characterized by
having two equal secondary ribs between six principal ribs.
The principal ribs are subdivided, with weak subordinate
ribs on both sides, which gives the principal ribs a tripartite
aspect (e.g. Figs. 4.8– 4.11). The lateral areas carry a
variable number of radial riblets (Fig. 4.8). In most
specimens the ribs are relatively sharp, particularly on
poorly preserved internal moulds. Where the shell is
preserved, the ribs have a more rounded outline
(Fig. 4.12). The auricles are subequal and relatively small.
In most specimens the auricles are poorly preserved and
their ornamentation hardly visible.
Brito (1967) discussed the resemblance of his new
species N. bahiaensis to N. ficalhoi (Choffat, 1888), the
latter now a synonym of N. (N.)alpina, and differentiated
them on the basis of the stronger secondary ribs in
N. bahiaensis. In that his material is conspecific with our
specimens, N. bahiaensis is here considered a junior
synonym of N. (N.)alpina.
Hayami (1965, p. 289) discussed the group of N. ficalhoi,
including Janira alpina d’Orbigny, 1847,J. ficalhoi
Choffat, 1888,N. occidentalis Conrad, 1855,N. george-
townensis Kniker, 1919, Vola irregularis Bo
¨se, 1910,
V. subalpina Bo
¨se, 1910, Pecten texanus Roemer, 1852,
and P. tricostatus Bayle, 1849. All these species are very
similar and most probably conspecific (Hayami, 1965). In
her revision of Neithea,Dhondt (1973) discussed Hayami’s
(1965) observations and concluded that there is no reason to
Table 2
Dimensions of selected, representative specimens of N. (N.) alpina
Specimen number Height (mm) Length (mm)
FPH 1245-I 13 10
FPH 1253-I 20 16
FPH 1254-I 26 25
FPH 1255-I 13 15
FPH 1257-I (left valve) 17 18
FPH 1258-I (left valve) 13 14
FPH 1259-I (left valve) 12 10
FPH 1260-I 23 20
FPH 1263-I 17 13
FPH 1264-I 19 18
FPH 1265-I (left valve) 15 16
FPH 1268-I (left valve) 20 19
FPH 1273-I 17 14
FPH 1283-I (left valve) 25 27
FPH 1293-I (left valve) 34 37
FPH 1294-I (left valve) 13 13
FPH 1295-I 18 17
FPH 1297-I 15 13
FPH 1300-I 30 26
FPH 1302-I 19 16
FPH 1313-I 12 10
FPH 1329-I 11 7
FPH 1348-I 26 23
FPH 1349-I 21 18
FPH 1350-I 19 13
FPH 1357-I 17 12
FPH 1360-I (left valve) 13 14
FPH 1361-I (left valve) 27 25
FPH 1366-I 11 8
FPH 1388-I 15 10
E. de Jesus Andrade et al. / Journal of South American Earth Sciences 17 (2004) 25–3832

separate N. ficalhoi and N. alpina.Dhondt (1973, p. 19) also
mentioned N.texana (Roemer), N.subalpina (Bo
¨se) and
N.irregularis (Bo
¨se) from North America as similar to
N. alpina. However, the former two differ from the alpina–
ficalhoi group by their lack of tripartite principal ribs. These
species probably fall within the variability of N. alpina.
However, for a definite decision about the synonymy of
these species, a revision of the North American faunas is
needed.
Dhondt (1973) did not mention N. occidentalis Conrad, a
species that also belongs to this group of North American
taxa. In his original description of that species, Conrad
(1855) emphasized the ‘trifid character’ of the principal ribs,
which is produced by a raised line or fine rib on the flanks of
the principal ribs. In this respect Conrad’s species falls
within the concept of N. alpina, as noted by Hayami (1965),
who placed it in his N. ficalhoi group. According to Dhondt
(personal communication), N. alpina and N. occidentalis are
probably distinct species, and the latter a synonym of
N. syriaca Conrad, 1852. However, judging from the
original descriptions and the discussion by Dhondt (1973),
N.syriaca differs from N. alpina by the absence of tripartite
principal ribs, its four unequal intercalary ribs, and the
smoothness of its areas and auricles.
Occurrence
Albian– Maastrichtian of Europe (e.g. France, Austria),
northern Africa and the Near East (e.g. Morocco, Algeria,
Egypt), Asia (e.g. India, Japan), and North America and the
Caribbean (USA, Jamaica); Albian of the Camamu Basin,
Brazil.
6.1.3. Neithea (Neithea) hispanica (d’Orbigny, 1850)
(Fig. 4.13–4.17)
1850 Janira hispanica d’Orbigny, 1850; d’Orbigny,
p. 170.
1887 Neithea sergipensis (sp. nov.); White, pp. 39 – 40,
Pl. 3, Figs. 6 and 7.
1912 Pecten (Neithea)aequicostata (Lamarck); Pervin-
quie
`re, pp. 135, 136.
1937 Neithea sergipensis White; Maury, pp. 142, 143, Pl.
5, Figs. 6 and 7.
non-1939 Neithea sergipensis White; Oliveira, p. 80, fig.
9[¼N.(N.) coquandi (Peron, 1877)].
1957 Neithea hispanica d’Orbigny; Dartevelle et al., Pl.
9, Fig. 12.
1973 Neithea (Neithea)hispanica (A. d’Orbigny, 1850);
Dhondt, pp. 14, 15 (with extensive synonymy).
Material
36 specimens from 18 localities in the Sergipe Basin:
Biriba 1 (GPIH-BR C517.7); Cruzes 4 (GPIH-BR C365.6),
6 (GPIH-BR C371.50), 8 (GPIH-BR C358.18, 119, 123,
126, 360.28), 9 (GPIH-BR C375.51), 16 (GPIH-BR
C380.4); Fazenda Nova 1 (FPH 1228-I); Japaratuba 4
(GPIH-BR C109.9), 11 (GPIH-BR C507.N1, N2), 14
(GPIH-BR C618.5, 10a, 10b, 10c, 15, 17, 20, 46); Jardim
19 (GPIH-BR C116.N1), 31 (GPIH-BR C658.26); Jerico
´4
(GPIH-BR C385.4); Machado 3 (GPIH-BR C247.5); Praia
10 (FPH 1236-I), 12 (FPH 1237-I); Pedra Branca 18 (FPH
1229-I–1235-I); Timbo
´7 (GPIH-BR C450.8).
Description and discussion
Relatively small; few specimens exceeding a height of
10 mm, the largest specimen reaching approximately
25 mm (Table 3). The right, convex valve, where well
preserved, shows 21 equal primary and secondary ribs
separated by narrow interspaces. The left valve is flat.
Lateral areas are smooth; auricles are not preserved.
With its overall equal ribs N. (N.)hispanica is easily
distinguished from most other Neithea species. However, it is
very similar to N. (N.) aequicostata (Lamarck, 1819) and has
often been referred to that species. N. (N.)hispanica differs
from N. (N.)aequicostata in its smaller number of ribs,
18–21 in N. (N.)hispanica and 23 – 36 in N. (N.)aequico-
stata. The geographical and stratigraphical distribution of the
two species is also different, N. (N.)hispanica being a typical
Tethyan form, which, according to Dhondt (1973), is never
found with the boreal N. (N.)aequicostata. All equal-ribbed
Brazilian Neithea are here referred to N. (N.)hispanica.
The similarities of N.sergipensis White, 1887 to N. (N.)
aequicostata were mentioned already by Newton (1916).
According to White (1887) and Maury (1937),
N. sergipensis is characterized by its small size; however,
in all other respects their specimens fit the concept of N. (N.)
hispanica and therefore may represent juvenile individuals.
Our material includes varying growth stages, from very
small (Figs. 4.13 and 4.15) to medium-sized (Figs. 4.14 and
4.17) specimens. Rennie (1945) described N. moutai and
N. salinasensis from Angola, which may be synonymous
with N. (N.)hispanica (Dhondt, 1973).
Occurrence
Albian of Angola (Freneix, 1966), Cenomanian of
northern Africa and south-western Europe (Dhondt, 1973)
Table 3
Dimensions of selected, representative specimens of N. (N.) hispanica
Specimen number Height (mm) Length (mm)
FPH 1228-I 9 9
FPH 1229-I 5 4
FPH 1230-I 8 7
FPH 1232-I 5 4
FPH 1234-I 6 5
FPH 1235-I 5 5
FPH 1236-I 8 6
FPH 1237-I 10 8
GPIH-BR C385.4 13 11
GPIH-BR C507.N2 16 13
GPIH-BR C618.10a 25 19
GPIH-BR C618.10b 23 20
E. de Jesus Andrade et al. / Journal of South American Earth Sciences 17 (2004) 25–38 33

and Peru (Dhondt and Jaillard, 1997); upper Albian – lower
Turonian of the Sergipe Basin, Brazil.
6.1.4. Neithea (Neithea) bexarensis (Stephenson, 1941)
(Fig. 4.18–4.20)
1941 Pecten (Neithea)Bexarensis Stephenson, n. sp.;
Stephenson, pp. 136, 137, Pl. 22, Figs. 7 and 8.
1993 Neithea (Neithea)latericostata sp. nov.; Muniz,
pp. 68–70, Pl. 4, Figs. 2– 4.
Material
Eleven specimens from ‘Locality 2’ of Muniz (1993),
near Alhandra, Paraı
´ba: DG-CT-UFPE 2841 (holotype of
N. (N.)latericostata Muniz), 2842–2843, 2848,
2850–2853, 2856, 2858, 2861.
Description and discussion
Small to medium-sized (Table 4). The convex right valve
carries two secondary ribs between each pair of principal
ribs. Each principal and secondary rib has three to five radial
striae. The lateral areas and the subequal auricles also are
ornamented by fine radial striae.
Muniz (1993) placed his new species N. (N.) latericos-
tata in the same group as N.(N.) bexarensis, characterized
by having two secondary ribs. He separated the two species
on the basis of smooth lateral areas in the latter. However,
this conclusion apparently was based on a misinterpretation,
because in the original description of N.(N.) bexarensis
(Stephenson, 1941, p. 136) the lateral areas and auricles
were said to be ornamented with riblets. In Stephenson’s
and Muniz’s material, the interspaces between the primary
and secondary ribs are wider than those between the
secondaries, a feature that, according to Muniz (1993),is
more strongly developed in the material from Paraı
´ba. In our
view, the two species cannot be separated on the basis of this
slight difference. The specimens from Paraı
´ba restudied
here lie well within the morphological concept of N.(N.)
bexarensis as defined by Stephenson (1941).
N.(N.) bexarensis resembles N. (N.) alpina (d’Orbigny),
but the latter has more prominent, angular, tripartite
principal ribs, as well as a more acute ventral margin. The
principal morphological character for separating the two
species is the presence of fine striae on the ribs of N.(N.)
bexarensis (Figs. 4.18 and 4.19), a feature never observed in
N.(N.) alpina.
In its general outline N. (N.) bexarensis is similar to
N. (N.) hispanica (d’Orbigny), but the latter has equal and
more numerous ribs that lack the radial striation seen in
N. (N.) bexarensis.
Occurrence
Maastrichtian of Texas, USA; Campanian of the
Pernambuco– Paraı
´ba Basin, Brazil.
6.2. Subgenus Neithella Hayami, 1965
Diagnosis
Shell inequilateral, asymmetrical, relatively small; right
valve with four to six broad ribs with fine striae in the
interspaces; anterior auricle usually pointed and much larger
than the posterior one (after Hayami, 1965; Dhondt, 1973).
6.2.1. Neithea (Neithella) notabilis (Mu
¨nster in Goldfuss,
1833)
(Fig. 4.21–4.26)
1833 Pecten notabilis Mu
¨nster; Goldfuss, p. 56, Pl. 93,
Fig.3a–c.
1847 Janira longicauda d’Orbigny, 1846; d’Orbigny,
pp. 639, 640, Pl. 445, Figs. 9–14.
1847 Janira cometa d’Orbigny, 1846; d’Orbigny,
pp. 640, 641, Pl. 445, Figs. 15–19.
1847 Janira digitalis d’Orbigny, 1846; d’Orbigny,
pp. 642, 643, Pl. 446, Figs. 1–3.
1939 Pecten (Neithea)Hourcqi nov. sp.; Collignon, p. 13,
Pl. 1, Figs. 16 and 17.
1973 Neithea (Neithella)notabilis (G. von Mu
¨nster in
Goldfuss, 1833); Dhondt, pp. 66 – 70, Pl. 4, Fig. 2 (with
extensive synonymy).
Material
Five specimens from three localities in the Sergipe
Basin: Mosquito 1 (GPIH-BR C239.20), Porto da Cruz 1
(GPIH-BR C513.2) and Tibu
´rcio 1 (GPIH-BR C514.2, 5, 6).
Description and discussion
Small to medium-sized (Table 5). The valves carry six
prominent principal ribs, whose interspaces are covered
with four fine, unequally developed secondary ribs. In four
of the five specimens studied the primary ribs on the right
valve are sharp and narrow (Figs. 4.21 – 24), whereas those
on the left valve are narrow, with flattened crests (Fig. 4.25).
The lateral area is smooth. The asymmetrical, inequilateral
outline and the very large anterior auricle (Figs. 4.21 and
4.23) contribute to the clear differentiation of N. (Neithella)
notabilis from all other species of Neithea from Brazil.
Table 4
Dimensions of N. (N.) bexarensis
Specimen number Height (mm) Length (mm)
DG-CT-UFPE 2841 25 24
DG-CT-UFPE 2842 34 28
DG-CT-UFPE 2843 (left valve) 25 26
DG-CT-UFPE 2848 16 13
DG-CT-UFPE 2850 16 13
DG-CT-UFPE 2851 18 16
DG-CT-UFPE 2852 23 20
DG-CT-UFPE 2853 20 11
DG-CT-UFPE 2856 10 9
DG-CT-UFPE 2858 34 Incomplete
DG-CT-UFPE 2861 23 18
E. de Jesus Andrade et al. / Journal of South American Earth Sciences 17 (2004) 25–3834

The interspaces between the primary ribs are also wider and
shallower than those in most specimens of the species
illustrated from Europe. The ribbing in N. (Neithella)
notabilis is, however, variable. Petho
¨(1906, p. 227, Pl. 15,
Figs. 10 and 11) described the very similar Neithea
almusana from the ‘Upper Senonian’ of Yugoslavia,
which has a more curved umbo. However, Petho
¨’s species
is based on only two specimens and may be conspecific with
N. (Neithella)notabilis. A related species is N. (Neithella)
valangiensis (Pictet and Campiche, 1870), which differs in
its well-developed striae on all ribs, less angular principal
ribs, stronger secondary ribs, and a narrower beak (Dhondt,
1973).
Occurrence
Widely distributed from the Aptian to the Cenomanian –
Turonian boundary of Europe (Dhondt, 1973); Aptian of
Japan and Cenomanian – Turonian of Madagascar; middle
and upper (?) Cenomanian of the Sergipe Basin, Brazil.
Acknowledgements
We thank biology students A.F. Costa, R.M. Farias and
M.F.S. Pinto (Universidade Federal de Sergipe, Brazil) for
preparing and measuring some of the specimens. S. Viana
(Universidade Federal de Pernambuco, Brazil) provided
photographs and specimens of N.bexarensis. J.S. thanks
S. Walter, S. Schneider (Universita
¨tHeidelberg,
Germany), and A. Herrmann (Penn State University,
USA), and P.B. thanks S.I. Bengtson (Universita
¨t Heidel-
berg) for field assistance. K. Will (Universita
¨t Heidelberg)
is acknowledged for photographic work. Part of the field-
work was financed by the German Academic Exchange
Service (DAAD) (E.J.A., J.S.) and the Swedish Natural
Science Foundation (NFR) (P.B.). J.S. received financial
support from the German Research Council (DFG), Grant
No. Be 1382/9. A.V. Dhondt (Koninklijk Belgisch
Instituut voor Natuurwetenschappen, Brussels) and
I. Dieni (Universita
`degli Studi di Padova, Padua)
provided careful and constructive reviews of the
manuscript.
Appendix A. Locality descriptions
Kcsp ¼Cotinguiba Formation, Sapucari Member
Kagg ¼Algodo
˜es Formation, Germa
ˆnia Member
Kagq ¼Algodo
˜es Formation, Quiepe Member
Section ¼outcrop height over 0.5 m.
Each locality designation is followed by a code in
brackets, which is provided as a shorthand for use on maps,
charts and specimens. The code consists of three letters
taken from the locality name, followed by the locality
number. For descriptions of localities not included here, see
Bengtson (1983) and Souza-Lima and Bengtson (in press).
A.1. Sergipe Basin
A.1.1. Jardim 31 (JAR-31)
UTM 8 823 610N/727 620E. Topographical map sheet:
SC.24-Z-B-V Japaratuba. Geological map sheet: SC.24-Z-
B-V-1 Carmo
´polis.
Section on hillside facing S at well-site CP-1260.
Altitude ca. 35 m.
Kcsp: Hard, nodular, light yellowish to beige limestone.
A.2. Camamu Basin
A.2.1. Algodo
˜es
References to the area: ‘… um calcareo marinho
fossilifero, exposto perto de Algodo
˜es, a norde
´ste da villa
de Marahu
´’(Maury, 1925). ‘Na margem direita do rio
Algodo
˜es, seis kilometros a nordeste da cidade de
Marahu
´…’ (Maury, 1930). Localidade 35-IT-3 (Coordena-
c¸a
˜o da Produc¸a
˜o Mineral, 1974). ‘…pequeno povoado Porto
dos Algodo
˜es, situado a meio caminho de Marau
´a
Campinho’ (Lima, 1981).
A.2.1.1. Algodo
˜es 3 (ALG-03). UTM 8 444 400N/503 800E.
Topographical map sheet: SD-24-Z-A-I-1-NO Marau
´.
Section and debris on the north river bank, near
mangrove margin. Altitude ca. 1 m.
Kagq: Friable light grey to light pink calcareous
siltstones and claystones, moderately bioclastic, with local
dolomitization.
A.2.2. Boipeba
References to the area: ‘… provenientes de afloramentos
de calca
´reos do litoral nordeste da Ilha Boipeba, situada na
a
´rea de Camamu…’ (Brito, 1964). ‘… coletados por J. I.
Fonseca, na Ilha Boipeba, em 1962,…’ (Brito, 1967).
‘Litoral nordeste da Ilha Boipeba, a
´rea de Camamu…’
(Brito, 1981).
A.2.2.1. Boipeba 1 (BPB-01). UTM 8 498 950N/508 550E.
Topographical map sheet: SD-24-X-C-IV-1-NO Velha
Boipeba.
Large boulders and debris along northeastern shore of
Boipeba island near Velha Boipeba village. Altitude 0 – 2 m.
Table 5
Dimensions of N. (Neithella)notabilis
Specimen number Height (mm) Length (mm)
GPIH-BR C239.20 10 7
GPIH-BR C513.2 11 9
GPIH-BR C514.2 23 20
GPIH-BR C514.5 [30] 27
GPIH-BR C514.6 36 32
Measurement in brackets is inferred.
E. de Jesus Andrade et al. / Journal of South American Earth Sciences 17 (2004) 25–38 35

Kagg: Dolomitized, locally saccharoidal and vuggy
cream limestones.
A.2.2.2. Boipeba 2 (BPB-02). UTM 8 498 950N/508 600E.
Topographical map sheet: SD-24-X-C-IV-1-NO Velha
Boipeba.
Section on hillside facing NE and large boulders and
debris along northeastern shore of Boipeba island near
Velha Boipeba village. Altitude 0 – 4 m.
Kagg: Dolomitized, locally saccharoidal and vuggy
cream limestones.
A.2.2.3. Boipeba 3 (BPB-03). UTM 8 499 000N/508 600E.
Topographical map sheet: SD-24-X-C-IV-1-NO Velha
Boipeba.
Section on hillside facing NE and large boulders and
debris along northeastern shore of Boipeba island near
Velha Boipeba village. Altitude 0 – 3 m.
Kagg: Dolomitized, locally saccharoidal and vuggy
cream limestones.
A.2.3. Cangaı
´ba
References to the area: ‘… na ilha Cangaı
´ba, situada
pro
´ximo a
`ilha Grande, na baı
´a de Camamu, municı
´pio de
Camamu…’ (Santos, 1958). ‘… coletados por A. Galea
˜o,
em 1926, na Ilha Gangaiba…’ (Brito, 1967).
A.2.3.1. Cangaı
´ba 1 (CGB-01). UTM 8 462 150N/498
100E. Topographical map sheet: SD-24-V-D-VI-4-SE
Camamu.
Debris along northwestern shore of Cangaı
´ba island on
tidal flat. Altitude 0 m.
Kagg: Dolomitized, locally saccharoidal, vuggy and
fossiliferous cream limestones.
A.2.3.2. Cangaı
´ba 2 (CGB-02). UTM 8 462 150N/498
250E. Topographical map sheet: SD-24-V-D-VI-4-SE
Camamu.
Section on hillside facing NE and debris along north-
eastern shore of Cangaı
´ba island on tidal flat. Altitude 0 m.
Kagg: Dolomitized, locally saccharoidal, vuggy and
fossiliferous cream limestones.
A.2.4. Taipus
A.2.4.1. Taipus 3 (TPU-03)
UTM 8 458 350N/507 400E. Topographical map sheet:
SD-24-X-C-IV-3-SO Barra Grande.
Section near shore, surrounding ‘Taipus hill’. Altitude
1–20 m.
Kagg: Dolomitized, locally saccharoidal, vuggy and
fossiliferous cream limestones.
A.2.5. Tatus
A.2.5.1. Tatus 1 (TAT-01)
UTM 8 451 000N/500 100E. Topographical map sheet:
SD-24-Y-B-III-2-NE Barcelos do Sul.
Section on hillside facing SW and debris and boulders on
tidal flat along southwestern margin of Tatus island.
Altitude 0 m.
Kagg: Dolomitized, locally saccharoidal, vuggy and
fossiliferous light brown limestones.
References:‘… e principalmente nos ricos jazigos da ilha
do Tatu…’ (Lima, 1981).
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