Zootaxa 5141 (6): 581–591 ISSN 1175-5326 (print edition)

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Copyright © 2022 Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)
https://doi.org/10.11646/zootaxa.5141.6.4
http://zoobank.org/urn:lsid:zoobank.org:pub:CC1937D3-94F8-4CC6-957D-48A496909916

New Early Cretaceous species of Zonocypris G. W. Müller, 1898

(Crustacea: Ostracoda) from the Crato Formation, Brazil
ENELISE KATIA PIOVESAN1,2*, GUSTAVO DIAS MELO1,5,
MARIA CRISTINA CABRAL3,4 & JULIANA GUZMÁN1,2
¹Laboratório de Micropaleontologia Aplicada/Laboratório de Geologia Sedimentar (LMA/LAGESE), LITPEG, Universidade Federal
de Pernambuco, Av. da Arquitetura, s/n, 50740-550, Recife, PE, Brazil.
²Universidade Federal de Pernambuco, Departamento de Geologia, Programa de Pós-Graduação em Geociências, PPGEOC, Av. da
Arquitetura, s/n, 50740-550, Recife, PE, Brazil.
� julitaguzmang@gmail.com; https://orcid.org/0000-0001-8717-4043
³Universidade de Lisboa, Faculdade de Ciências, Instituto Dom Luiz (IDL), Campo Grande, C1, 1749-016 Lisboa, Portugal.
� mccabral@fc.ul.pt; https://orcid.org/0000-0001-7898-049X
4
Universidade de Lisboa, Faculdade de Ciências, Departamento de Geologia, Campo Grande, C6, 4º, 1749-016 Lisboa, Portugal.
5�
gustavodiasmelo@gmail.com; https://orcid.org/0000-0002-3648-5383
*
Corresponding author. � enelise.katia@ufpe.br, https://orcid.org/0000-0002-0433-0395

Abstract

Two new species of Zonocypris G. W. Müller, 1898 are described from the late Aptian of the Araripe Basin, Northeast
Brazil: Zonocypris berthoui sp. nov. and Zonocypris dorsoconvexa sp. nov. Species of this genus occur from the Early
Cretaceous to the Holocene and are characterized by a small globular carapace, often ornamented over the whole surface
by concentric ridges with simple normal pore canals, showing a remarkable morphological stability during the evolution
of the genus. The first records of the genus point to an Atlantic origin. In Aptian deposits, species of Zonocypris are rare,
occurring in brackish/freshwater assemblages, with marine influence.

Key words: Ostracods, taxonomy, paleoecology, geographical distribution, Aptian

Introduction

The genus Zonocypris G. W. Müller, 1898 is recorded from Early Cretaceous to Holocene deposits, and was erected
for living specimens from freshwater environments in Madagascar. In the original description, the main diagnostic
characters of the genus are the ornamentation with concentric ribs, the adont hinge, the short and plump second an-
tenna and the presence of sexual dimorphism. Higuti & Martens (2012) pointed out the presence of a relatively well
developed postero-dorsal tooth in the Zonocypris s.s. hinge, either in the right or left valve. Fossil representatives
of this genus are very characteristic due to their small size, globular carapace and ornamentation pattern showing
prominent concentric ridges. According to Külköylüoðlu et al. (2021), 32 species of the genus Zonocypris have
been identified worldwide, including living and fossil species, with 16 species described in each group; while these
numbers are consistent for living species (cf. Meisch et al. 2019), they missed several fossil records, particularly
the oldest ones.
The oldest records of the genus are found in the Early Cretaceous; however, in these intervals the assignment to
the genus is doubtful and/or the species are in open nomenclature, most often due to their rarity in the samples. In
the Early Cretaceous, Zonocypris was recovered in the Pozo D-129 Formation, Barremian?–Aptian of the Gulf San
Jorge Basin, Patagonia, Argentina (Carignano et al. 2017); in the Cresmina Formation, top of the lower Aptian of
the Lusitanian Basin, Portugal (Cabral 1995; Pereira & Cabral 2005); and in the Crato Formation (upper Aptian) of
the Araripe Basin, Northeast Brazil (Berthou et al. 1994; Colin & Dépêche 1997; Do Carmo et al. 2004). A species
tentatively assigned to the genus was described in the Qiuncheg Formation (Aptian–Albian) in China (Tian & Zhao
1982). In the Late Cretaceous, Zonocypris became diversified, with several species described, occurring in South

Accepted by R. Smith: 13 May 2022; published: 31 May 2022 581

America, Neuquén Basin, Argentina (as Timiriasevia? sp. in Uliana & Musacchio 1978, pl. 2, fig. 30-31), North
America (Tibert et al. 2009), Europe, France (Babinot 2003), West Africa, Mali (Colin et al. 1996) and mostly in
India (e.g., Khosla et al. 2011; Kapur et al. 2018 and references therein).
In Brazil, fossil specimens of the genus Zonocypris were registered only in the Araripe Basin, where they are
rare (Berthou et al. 1994; Colin & Dépêche 1997; Do Carmo et al. 2004). The Brazilian Cretaceous non-marine os-
tracods have been studied since the nineteenth century (Jones 1860), particularly after 1960, especially due to their
biostratigraphic application (see Pinto & Sanguinetti 1958; Krömmelbein 1962, 1966; Viana 1966; Schaller 1969;
Krömmelbein & Weber 1971; Moura 1972; Coimbra et al. 2002; Poropat & Colin 2012). Despite several studies of
ostracods in Brazilian basins in this interval, the Zonocypris species have not been formally described so far, and
thus the descriptions of two species of the genus are the main objectives of the present study.

Geological setting

The analyzed material was collected from the well core 2-AR-SR-1A-CE (469 samples), Crato Municipality, and
from the outcrop 1BAr17 (eight samples), Três Irmãos Quarry, in Santana do Cariri Municipality, Araripe Basin,
Ceará State (Fig. 1). The studied samples belong to a succession included in the post-rift sequence of the Araripe
Basin, which corresponds stratigraphically to the Santana Group, comprising, from bottom to top, the Barbalha,
Crato, Ipubi and Romualdo formations (Assine et al. 2014), of Aptian age (Arai & Assine 2020; Melo et al. 2020).
The Crato Formation (c. 90 m thick), mainly characterized by interbedded limestones, shales and sandstones, has
been considered as a lacustrine system due to the apparent lack of true marine fossils (Coimbra et al. 2002; Assine
et al. 2014). However, recent discoveries of foraminifers, dinoflagellate cysts and typical marine ichnotaxa in the
Crato Formation (Varejão et al. 2021 and references therein) point to the presence of intermittent marine incursions
in this litho-stratigraphical unit. According to Varejão et al. (2021), the onset of the Crato Formation represents
a Transgressive Systems Tract followed by a Highstand Systems Tract. The transgressive surface and associated
deposits, mainly expressed by limestones, are succeeded by tide-dominated bay deposits with distinct subtidal,
intertidal and supratidal intervals, and after the Maximum Flooding Surface (a dark shale, below foreshore to shore-
face facies, marking the beginning of the Highstand Systems Tract), by evaporites. From Crato Formation strata,
abundant and diverse ostracods were recovered, including the very rare Zonocypris specimens presented here.

Material and methods

The samples were weighed (60 g of rock per sample) and mechanically crushed into fragments of approximately 1
cm. The fragmented sample was submerged in water for a period of 24 hours and then washed with running water
in a sieve set of 500, 250, 180 and 62 µm mesh, followed by oven drying at 60 ºC. Representatives of the genus
Zonocypris were only recovered in the fractions of 180 µm.
The specimens were imaged in a scanning electron microscope (SEM), at the Laboratório de Micropaleontolo-
gia Aplicada [Applied Micropaleontology Laboratory] of the Federal University of Pernambuco. The new species
are registered in Zoobank (the Official Registry of Zoological Nomenclature of the International Commission on
Zoological Nomenclature).
Higher classification of the Ostracoda follows Horne et al. (2002) and Meisch et al. (2019). Abbreviations: L =
length; H = height; W = width; C = carapace; RV = right valve; LV = left valve; very small carapace (L<0.40 mm);
small carapace (L = 0.40–0.50 mm); medium carapace (L = 0.51–0.70 mm).

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Figure 1. Map of the studied area showing the collected outcrop and well in the Araripe Basin, NE Brazil. Produced from
metadata provided by Geological Survey of Brazil - CPRM.

Paleontological Systematic

Class OSTRACODA Latreille, 1802

Subclass PODOCOPA Sars, 1866

Order PODOCOPIDA Sars, 1866

Suborder CYPRIDOCOPINA Baird, 1845

Superfamily CYPRIDOIDEA Baird, 1845

Family CYPRIDIDAE Baird, 1845

Subfamily CYPRIDOPSINAE Kaufmann, 1899

Tribe ZONOCYPRIDINI Higuti & Martens, 2012

Genus Zonocypris G. W. Müller, 1898

Type-species: Zonocypris costata (Vávra, 1897)

Syn.: Zonocypris madagascarensis G. W. Müller, 1898

New Early Cretaceous species of Zonocypris Zootaxa 5141 (6) © 2022 Magnolia Press · 583
Zonocypris berthoui sp. nov.
(Fig. 2)

1994 Zonocypris sp. Berthou et al.: 541, pl. 1, figs 8–10.

1997 Zonocypris sp. Colin & Dépêche: 433, figs 2–14, 2–17.

ZooBank-link: urn:lsid:zoobank.org:act:879AA9BD-FBAA-4B02-945F-41844FAB3DE0
Etymology: In honor of the late Pierre-Yves Berthou, for his pioneering studies on ostracods of the Araripe
Basin, including the first record of the species described herein.
Type-locality: Nova Olinda Municipality, Ceará State, Northeast Brazil.
Type-horizon: Late Aptian. Crato Formation, Santana Group, Araripe Basin, Brazil. Outcrop sample 1BAr17a,
coordinates 7°06’52.3”S, 39°41’51.5”W.
Material: Four adult carapaces.
Holotype: LMA-00305, C, Adult female; L: 0.40 mm, H: 0.28 mm, W: 0.26 mm (Figs 2A–F). Outcrop sample
1BAr17a (Três Irmãos Quarry).
Paratypes: LMA-00331, C, Adult male, L: 0.42 mm, H: 0.28 mm. W: 0.27 mm (Figs 2G–H), 2-AR-SR-1A-CE
well core sample from depth interval 124.55–124.58 m; LMA-00332, Adult female L: 0.42 mm, H: 0.22 mm, W:
0.31 mm (Figs 2I–K), 2-AR-SR-1A-CE well core sample from depth interval 121.05–121.09 m.
Repository: Collection of the Laboratório de Micropaleontologia Aplicada [Applied Micropaleontology Lab-
oratory] (LMA) from UFPE, Recife, Brazil.
Diagnosis: A species of Zonocypris with a small, sub-ovate to sub-triangular carapace in lateral view, with the
typical ornamentation of thick, concentric ribs (except in the central area, where three sub-rectilinear ribs are pres-
ent) containing abundant normal pore canals; intercostal areas narrow and smooth.
Description: Small carapace, sub-ovate to sub-triangular in lateral view; strongly tumid, with biconvex dorsal
and ventral views. Valves slightly unequal, LV overlapping the RV on all sides. Greatest height located in antero-
median area, at anterior cardinal angle; height c. 2/3 of length in females. Greatest length below mid-height. Great-
est width slightly posterior to mid-length, larger than half of length in females, smaller in males. Dorsal margin
sub-rectilinear and short, ventral margin convex; anterior margin broadly rounded, posterior end infracurvate and
subacute. Sexual dimorphism present, with female carapace more inflated in dorsal view, and male carapace more
elongate and subtriangular laterally.
Surface entirely covered by coarse, sub-oval and regularly distributed concentric ribs, except in central area,
where three sub-rectilinear ribs are present. In dorsal and ventral views, ribs are subparallel to major axis. Intercostal
areas smooth and narrower than ribs. Abundant, round, simple normal pore canals (type A’ of Puri & Dickau 1969
and A1 of Danielopol et al. 2018) occur on ribs or adjacent to them throughout valve surface (Figs 2I and K). Some
best preserve pores with very thin lip around hole, seen at high-resolution (Figs 2F and J), possibly belonging to
the type A” of Puri & Dickau (1969) and A2 of Danielopol et al. (2018). Marginal pore canals also present, densely
arranged (Fig. 2E).
Presence of microdenticles in anteroventral area only observed when carapace is disarticulated or broken in this
zone (Fig. 2K). Other internal features not available.
Remarks: Normal pore canals and marginal pore canals are rarely referred to in the descriptions of Zonocypris
species, probably because they both require high magnifications of SEM images to be clearly observed; moreover,
many descriptions of Zonocypris species predated the widespread use of SEM, particularly those of recent species,
where these small structures are easier to see. However, in recent publications with SEM images, the normal pore
canals are often visible at high magnifications. We verified their presence in several fossil and extant species, always
located on the ribs or adjacent to them; in Whatley et al. (2002) the presence of normal pore canals on the ribs of
Zonocypris labyrinthicus Whatley, Bajpai & Srinivasan, 2002 was noted. The marginal pore canals are more dif-
ficult to observe. In Krstić (2006) there is a figure (Pl. LXVI, fig. 8) with the marginal pore canals of Zonocypris
membranae quadricella Stancheva, 1966; the author described them as “long, thin and densely arranged”.
The presence of microdenticles in the RV is common in extant and sub-fossil species of Zonocypris. In the
description of the extant Zonocypris cordata Sars, 1924, from South Africa, the author refers to a “row of minute
tubercles”, “both in front and behind” on the RV; microdenticles are also present in the anterior and posterior-ven-
tral parts of the selvage of the RV of a very well preserved Holocene species from Mozambique (MCC observa-
tion on material from M. J. Martins, ICArEHB, Universidade do Algarve, Portugal, Project InMoz -PTDC/HAR-

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ARQ/28148/2017, ongoing studies); in the extant Zonocypris mardinensis Külköylüoðlu et al., 2021, from Turkey,
the presence of a row of fine denticles is referred to, but only in the posterior part of the calcified inner lamella of the
RV (in the selvage, as we can see in fig. 4F, of Külköylüoðlu et al. 2021). The lack of descriptions of these structures
in fossil species is probably due to the rarity of loose valves where they can be observed.

Figure 2. Zonocypris berthoui sp. nov. (A–F) LMA-00305, holotype, female carapace, (A) general right view, (B) general
left view, (C) general dorsal view; (D) general ventral view, (E) detail of D, showing normal and marginal pore canals, (F)
detail of E, simple normal pore canals (type A” of Puri & Dickau 1969 and A2 of Danielopol et al. 2018); (G–H) LMA-00331,
paratype 1, male carapace, (G) general dorsal view, (H) general left view; (I–K) LMA-00332, paratype 2, female carapace, (I)
dorsal view, detail of anterior part, simple normal pore canals (type A’ of Puri & Dickau 1969 and A1 of Danielopol et al. 2018),
(J) detail of I, simple normal pore canals (type A” of Puri & Dickau 1969 and A2 of Danielopol et al. 2018), (K) left view, detail
of anteroventral denticulation.

New Early Cretaceous species of Zonocypris Zootaxa 5141 (6) © 2022 Magnolia Press · 585
 Zonocypris berthoui sp. nov. differs from other Early Cretaceous species of the genus mainly by its sub-tri-
angular to ovoid outline and the ribs distribution pattern. Compared to Zonocypris? expansa Tian & Zhao, 1982,
from the Aptian-Albian of NE China, it is significantly smaller, with a very different lateral outline and thicker ribs.
Zonocypris sp. recorded by Carignano et al. (2017) in the Barremian?–Aptian interval of Argentina has a rounded
sub-trapezoidal outline in lateral view, a greatest width clearly in the posterior third and more asymmetrical arrange-
ment of the concentric ribs in the center of the valve.
Some species from the Late Cretaceous resemble our proposed species. In India, Zonocypris labyrinthicus
Whatley, Bajpai & Srinivasan, 2002 from the Maastrichtian of Mohagaonkala, and Zonocypris gujaratensis Bhan-
dari & Colin, 1999, from the Upper Maastrichtian-Base of Paleocene? of Anjar, Kachchh, Gujarat State, also show
sexual dimorphism, but have a very small carapace with a more rounded lateral outline than Z. berthoui sp. nov.;
moreover Z. labyrinthicus has RV>LV and lacks the ribs in the central part of the valves, and in Z. gujaratensis the
greatest height is almost in the middle of the length. Zonocypris spirula Whatley & Bajpai, 2000, from the Late
Cretaceous and Paleogene of India, differs from the present species in having a very small carapace with spirally
arranged ornament comprising a single helicoidal rib coiled on the valve surface. The Campanian Zonocypris digi-
talis Babinot, 2003, from south-east France, presents a similar rib pattern, but differs in the truncated anterior and
posterior margins, and in size, which is very small. The specimen of Zonocypris? sp. from the Campanian–Early
Maastrichtian? of Mali, figured in Colin et al. (1996), very likely belongs to the genus and shows similarities with
Z. berthoui sp. nov., however it has an almost round outline and the intercostal areas are pitted.
Occurrence: Late Aptian, Crato Formation, Araripe Basin, Barbalha, Crato and Nova Olinda municipalities,
Ceará State, Brazil (Berthou et al. 1994; Colin & Dépêche 1997; this study).

Zonocypris dorsoconvexa sp. nov.

(Fig. 3)

ZooBank-link: urn:lsid:zoobank.org:act:F2A2EDDA-1D45-4290-8FB4-89C3D10B00C2
Etymology: The specific name is derived from Latin convexus (meaning convex) and refers to the dorsal outline.
Type-locality: Crato Municipality, Ceará State, Northeast Brazil.
Type-horizon: Late Aptian. Crato Formation, Santana Group, Araripe Basin, Brazil. 2-AR-SR-1A-CE well
core (coordinates: 7°17’29.88” S and 39°23’34.84” W) sample, depth interval 134.00–134.05 m.
Material: three carapaces (two adults; one juvenile).
Holotype: LMA-00333, adult male; L: 0.42 mm, H: 0.24 mm, W: 0.27 mm (Figs 3A–D). 2-AR-SR-1A-CE well
core sample, depth interval 134.00–134.05 m.
Paratypes: LMA-00306, female, L: 0.41 mm, H: 0.27 mm, W: 0.27 mm (Fig. 3E), 2-AR-SR-1A-CE well core
sample, depth interval 134.00–134.05 m. LMA-00334, C, juvenile, L: 0.33 mm, H: 0.21 mm. W: 0.22 mm (Figs
3F–H), 2-AR-SR-1A-CE well core sample, depth interval 124.55–124.58 m.
Repository: Collection from Laboratório de Micropaleontologia Aplicada [Applied Micropaleontology Labo-
ratory] (LMA) from UFPE, Recife, Brazil.
Diagnosis: A species of Zonocypris with a small, sub-rectangular to sub-ovate carapace in lateral view, with the
typical ornamentation of concentric ribs almost horizontal in the central and ventral areas; intercostal areas smooth
and slightly thicker than the ribs.
Description: Small carapace, sub-rectangular to sub-ovate in lateral view. Carapace ovoid, in dorsal and ventral
views, posteriorly broadened, anteriorly acuminate. LV larger than RV, uniformly overlapping it along all margins,
except in the ventral margin. Greatest height at mid-length in females, in front of middle in males. Greatest length
below mid-height. Greatest width in posterior third. Dorsal margin convex, ventral margin almost straight; anterior
margin sub-rounded, posterior margin truncated and projected downwards. Sexual dimorphism pronounced, with
females higher, less elongated, more inflated and dorsally more convex than males.
Concentric ribs parallel to margins on carapace periphery. In central region of carapace, ribs more triangular
shape, being almost horizontal in ventral zone. Intercostal areas smooth and slightly thicker than ribs. On ribs, or
adjacent to them, simple normal pore canals (type A’ of Puri & Dickau 1969 and A1 of Danielopol et al. 2018) are
abundant, some of which seemingly with marginal lip (type A” of Puri & Dickau 1969 and A2 of Danielopol et al.
2018), bad preservation of specimens not allowing for more detailed description (Fig. 3H). Marginal pore canals
also present, more densely arranged (Fig. 3D).

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 Internal features not seen.
Remarks: This species differs from the previous Zonocypris berthoui sp. nov. in general outline, mainly the
dorsal one, less robust ribs and the posterior position of greatest width. Zonocypris dorsoconvexa sp. nov. resembles
Zonocypris sp. 1 from the Early Aptian of Portugal (Cabral 1995), but the latter has a straighter ventral margin and
a more marked posterior end; the ornamentation is also slightly different with the ribs more inclined in the ventral
area. The species Zonocypris sp., recorded from the Early Cretaceous from Argentina (Carignano et al. 2017), dif-
fers from Zonocypris dorsoconvexa sp. nov. in the more rounded anterior margin, and position of the greatest width
in the posterior quarter of the carapace.
Occurrence: Late Aptian, Crato Formation, Araripe Basin, Crato Municipality, Ceará State, Brazil (this
study).

Figure 3. Zonocypris dorsoconvexa sp. nov. (A–D), LMA-00333, holotype, male carapace, (A) general ventral view, (B)
general dorsal view, (C) general right view, (D) ventral view, anterior part, marginal pore canals; (E) LMA-00306, paratype 1,
female carapace, general left view; (F–H) LMA-00334, paratype 2, juvenile carapace, (F) general right view, (G) general left
view, (H) ventral view, anterior part, simple normal pore canals (types A’ and A”? of Puri & Dickau 1969 and A1 and A2? of
Danielopol et al. 2018).

New Early Cretaceous species of Zonocypris Zootaxa 5141 (6) © 2022 Magnolia Press · 587
Discussion

Taxonomic and paleogeographic remarks

The species of Zonocypris presented here are the first formally described from Early Cretaceous deposits. Most
of their morphological characteristics, such as the small dimensions, globose shape, ornamentation consisting of
concentric ribs, simple normal pore canals, probably of two types (A’ and A” of Puri & Dickau 1969; A1 and A2
of Danielopol et al. 2018), located on the ribs or adjacent to them on the entire lateral surface, LV > RV, marginal
pore canals abundant, presence of antero-ventral microdenticles in the RV, are typical features of the genus over
time (from the Early Cretaceous to today). This may be easily verified in the literature concerning descriptions
and illustrations of other species of Zonocypris, both from the Mesozoic (e.g. Bhandari & Colin 1999; Whatley
& Bajpai 2000; Whatley et al. 2002; Babinot 2003) and Cenozoic (e.g. Sars 1924; Mostafawi 1994; Krstić 2006;
Külköylüoðlu et al. 2021), which show remarkable morphological stability in the genus. The main differences seem
to be the increase in size (from very small and small to medium), the more elongated lateral outline, presence of
posterior “extensions” and intercostal spaces frequently reticulated/pitted in the Cenozoic species. Some species
from the Late Cretaceous are more elongate in lateral view, when compared with the Early Cretaceous Zonocypris
species (Whatley et al. 2002), or more inflated in the median region (Khosla et al. 2011), and generally present very
strong ribs (e.g. Whatley & Bajpai 2000; Whatley et al. 2002, Babinot 2003). The widely distributed Zonocypris
membranae Livental, in Suzin 1956, occurs from the Miocene to late Pleistocene (see Külköylüoðlu et al. 2021) and
clearly shows an increase in size relative to the Cretaceous species, as well as a more elongate carapace with thin-
ner ribs and reticulated intercostal spaces (e.g. several images of Z. membranae quadricella Stancheva, in Krstić,
2006). These trends are also observed in extant species, for instance Z. costata (Vávra), in Cabral et al. 2004 and Z.
mardinensis Külköylüoðlu et al. 2021.
The oldest known representatives of Zonocypris have an Aptian age, occurring in South America (Brazil, Ar-
gentina), Europe (Portugal) and most doubtfully China (Aptian–Albian). The Chinese species Z.? expansa Tian,
1982 (in Tian & Zhao 1982) has a different lateral outline, with a very convex ventral margin, suggesting the de-
velopment of a subtle ala, not observed in the other species of the genus. The dispersion center of the genus is still
unclear, but the known occurrences strongly suggest an Atlantic origin, linked to the opening of this ocean, when the
marine connection between the Central and South Atlantic, and therefore Tethys and Atlantic oceans, was already
established (Melo et al. 2020; Varejão et al. 2021 and references therein).

Paleoecological aspects
The living species of Zonocypris mainly occur in Africa, with also rare records in South-Central America, Australia
and Turkey (Karanovic 2012; Külköylüoðlu et al. 2021). They are considered detritivores, inhabiting warm, slightly
alkaline, shallow permanent continental freshwater environments, such as lakes (e.g. Mazzini 2011; Karanovic
2012; Mazzini et al. 2013). Külköylüoðlu et al. (2021), reported that the two living species of the genus in Tur-
key inhabit warm (15–30ºC), slightly acidic to alkaline (pH 6.81–8.44), low to well oxygenated (3.05–18.8 mg/l)
waters, tolerating salinity (electrical conductivity 103-1910 µS/cm; salinity 0.05–0.39 ppt) values within a lim-
ited elevational range (336–991 m). This freshwater feature of extant Zonocypris is common to Cenozoic species,
mainly Miocene to Pleistocene, and to older species of the genus. Mazzini et al. (2013) state that the Zonocypris
membranae quadricella Stancheva (Late Miocene–Early Pleistocene), is considered the only species of the genus
adapted to brackish environments, with all the remainder being freshwater. Most of the Late Cretaceous Zonocypris
species were found in the Deccan “intertrappean beds” of India (species list in Külköylüoðlu et al. 2021), known to
yield non-marine microfossil assemblages; the French Late Cretaceous Z. digitalis Babinot also occurs in a typical
freshwater-oligohaline assemblage (Babinot 2003).
Early Cretaceous Zonocypris species occur in freshwater assemblages, together with species of brackish ostra-
cod genera. In our material, Zonocypris berthoui sp. nov. and Z. dorsoconvexa sp. nov., always rare, occur together
with abundant specimens of Pattersoncypris Bate, Damonella Anderson, Theriosynoecum Branson and Brasacypris
Krömmelbein and, less abundantly, Cypridea Bosquet, Alicenula Rossetti & Martens, Neuquenocypris Musacchio
and Ilyocypris Brady & Norman representatives; despite belonging to the non-marine ostracod group, most of these
genera tolerated a slight increase in salinity (e.g. Carbonel et al. 1988 and references therein; Horne 2002). The
same ostracod association was demonstrated in previous works on the Crato Formation from the Araripe Basin (Ber-
thou et al. 1994; Colin & Dépêche 1997). The lithology of the strata from which the Zonocypris berthoui sp. nov.
and Z. dorsoconvexa sp. nov. were recovered corresponds to limestone (outcrop 1BAr17), shale and heterolithic

588 · Zootaxa 5141 (6) © 2022 Magnolia Press PIOVESAN et al.

facies in flaser and wavy bedding (2-AR-SR-1A-CE well core, depth interval 121.05–134.05 m) from the lower
part of the Crato Formation. From the same strata of the well core, besides ostracods, benthic foraminifers were
also recovered. The foraminifera specimens are currently being identified for future studies, although their pres-
ence indicates marine influence, suggested as well by the heterolithic bedding, mostly formed in tidally influenced
environments of brackish water. Our preliminary facies analyses, and benthic foraminifera recognition are in accor-
dance with Varejão et al. (2021), who interpreted the Crato Formation deposition occurring in a broad non-marine
(coastal/marginal—tide-dominated bay) environment with short-lived marine incursions.
In deposits of the Lower Cretaceous from Argentina, Zonocypris sp. was associated with Charophyta species
and non-marine ostracods of the families Limnocytheridae and Darwinulidae. In the Upper Cretaceous Argentinian
rocks, one species from the genus Zonocypris was recorded (as Timiriasevia? sp.) by Uliana & Musacchio (1978),
mainly associated with several Ilyocypris species. In the Portuguese ostracod assemblage from the top of lower Ap-
tian of Rio de Mouro, Lusitanian Basin, well discussed in Cabral & Colin (1998), the single specimen of Zonocypris
sp. 1 (in Cabral 1995) occurs together with abundant charophytes and freshwater-oligohaline ostracods belonging to
the genera Cypridea and Rosacythere Colin (Limnocytheridae, Timiriaseviinae), and the marine euryhaline genus
Cytheropterina Mandelstam.
Despite its rarity, the presence of Zonocypris, in similar assemblages of brackish environments (with intermit-
tent marine influence), in the Early Cretaceous, suggests that the ancient forms of the genus were more tolerant to
environment parameters, particularly salinity, than the recent ones.

Acknowledgements

We thank the Laboratório de Micropaleontologia Aplicada [Applied Micropaleontology Laboratory] (LMA- Feder-
al University of Pernambuco) team for their help with this study. We also thank the reviewers J. C. Coimbra (Brazil)
and A. P. Carignano (Argentina) for their useful suggestions and comments. Editorial guidance by Robin Smith was
much appreciated. This work was supported by Brazilian National Petroleum Agency (ANP) and PETROBRAS
through the projects ARTUNJA: Correlações bioestratigráficas dos sistemas flúvio-lacustres das fases rifte e pós-
rifte das bacias do Araripe, Jatobá e Tucano Norte, NE do Brasil / nº 2017/00263-2” and “Implantação da infraestru-
tura do Laboratório de Micropaleontologia Aplicada da Universidade Federal de Pernambuco / nº 2018/00320-9”.
M. C. Cabral was supported by Fundação para a Ciência e Tecnologia (FCT), I.P./MCTES, National Funds (PID-
DAC)—UIDB/50019/2020.

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