http://dx.doi.org/10.5852/eit.2Q14.106

www. europeani ournaloftaxonomv. eu

2014 • Higuti J. & Martens K.

This work is licensed under a Creative Commons Attribution 3.0 License.

Research article

urn:lsid:zoobank.org:pub:3C2078C7-B4B0-4976-8CED-CD2AA97E9E8F

Five new species of Candoninae (Crustacea, Ostracoda) from the

alluvial valley of the Upper Parana River (Brazil, South America)

Janet HIGUTI 1 & Koen MARTENS 2 *

1 Universidade Estadual de Maringa, Nupelia, PEA, Av. Colombo 5790, CEP 87020-900,

Maringa, PR, Brazil, E-mail: [email protected]

2 Royal Belgian Institute of Natural Sciences, Freshwater Biology, Vautierstraat 29, B-1000 Brussels,

Belgium and University of Ghent, Biology, K.L. Ledeganckstraat 35, B-9000 Ghent, Belgium,

E-mail: [email protected]

^corresponding author e-mail: [email protected]

1 urn:lsid:zoobank.org:author:3A5CEE33-280B-4312-BF6B-50287397A6F8

2 urn:lsid:zoobank.org:author:9272757B-A9E5-4C94-B28D-F5EFF32AADC7

Abstract. The present paper describes five new species of candonid ostracods in two genera:

Pseudocandona agostinhoi sp. nov., P. cillisi sp. nov., P. claudinae sp. nov., Candobrasilopsis elongata

sp. nov. and C. acutis sp. nov. The three species of Pseudocandona belong to the caribbeana- group

in this genus. With the two new species of Candobrasilopsis , this genus now comprises four species.

Candobrasilopsis elongata sp. nov. is the most common of the five new species described here, while C.

acutis sp. nov. and P. claudinae sp. nov. are known from one locality only, which is furthermore the same

for both species: a small streamlet entering the Parana River. With the description of the present five

species, the number of species known from the Parana River alluvial valley, including the Taquaru 9 u

lakes, now amounts to 49.

Key words. Pseudocandona, Candobrasilopsis , streams, taxonomy

Higuti J. & Martens K. 2014. Five new species of Candoninae (Crustacea, Ostracoda) from the alluvial valley

of the Upper Parana River (Brazil, South America). European Journal of Taxonomy 106: 1-36. http://dx.doi.

org/10.5852/eit.2014.106

Introduction

There are about 2000 Recent non-marine ostracod species in the world (Martens et al. 2008). Close to

300 of these species have been described and reported from South America and more than 100 species

are now known from Brazil (Martens & Behen 1994; Higuti et al. 2009a, 2013; Higuti & Martens

2012a, 2012b). Recent work on the alluvial valley of the Upper Parana River (Higuti et al. 2007, 2009b,

2009c, 2010; Mormul et al. 2010) has contributed significantly to the general knowledge of ecology and

biogeography of Brazilian freshwater ostracods, especially those occurring in the pleuston of floating

plants. Floating plants such as species in the genera Eichhornia , Pistia and Salvinia are typical of South

American (floodplain) lakes, and pleuston, the animal communities living in the root systems of such

plants, are thus typical of such lakes.

European Journal of Taxonomy 106: 1-36 (2014)

The ecological realm of pleuston, next to the plankton and benthos (see Por 1995), is especially important

in floodplain lakes. Sediments in floodplain lakes have low oxygen levels most of the time and sudden

water level rise can render sediments anoxic virtually overnight. Thus, floating plants become important

substrates for colonization by ostracods, since these macrophytes will simply follow rising water levels

and local variables such as oxygen will change minimally on the root systems of the floating vegetation.

Many otherwise benthic organisms can thus be found in the pleuston in South American floodplain

lakes.

The collections made to study the ecology of the ostracods in the pleuston in the alluvial valley of

the Upper Parana River (Higuti et al. 2007, 2009b, 2009c, 2010; Mormul et al. 2010) also served for

taxonomic revisions and several new species and genera of ostracods have meanwhile been described

from lakes, river beds and channels of this floodplain (Higuti et al. 2009a, 2013; Higuti & Martens

2012a, 2012b). Most of these new taxa belong to the family Cyprididae; only one paper thus far treated

two species of Candonidae from this floodplain in detail (Higuti & Martens 2012b), describing a new

genus and species, Candobrasilopsis rochai Higuti & Martens, 2012, and redescribing and generically

transferring another species, Candobrasilopsis brasiliensis (Sars, 1901).

Porto

Primavera

BRAZIL

SAO

PAULO

STATE

Taquaruqu

0 400 800km

MATO GROSSO

CunA-baChamd

PARANA

STATE

0 2 4 6 8 10km

Fig. 1 . Map of the study area, indicating the localities of three new species of Pseudocandona and two new

species of Candobrasilopsis.

HIGUTI J. & MARTENS K., Five new Candoninae species from Brazil

Here, we describe five new species of candonids from this floodplain in southern Brazil (South America);

three new species belong to the genus Pseudocandona Kaufmann, 1900 and two new species to the genus

Candobrasilopsis Higuti & Martens, 2012. Several new species from this and other Brazilian floodplains

remain to be described, e.g., in the genera Physocypria (Candonidae), Cypretta and Strandesia as well

as in several genera of the Cypridopsinae (Cyprididae).

Material and Methods

Study area

The Upper Parana River floodplain is located between the Porto Primavera Reservoir and the Itaipu

Reservoir, extending over about 230 km. In this area, three conservation units were created: “Area

de Prote 9 ao Ambiental das Ilhas e Varzeas do Rio Parana” (100,310 ha; an Environmental Protection

Area), the “Parque Nacional de Ilha Grande” (78,800 ha; a National Park), and the “Parque Estadual

do Ivinheima” (70,000 ha; a State Park) (Agostinho & Zalewski 1996; Agostinho et al. 2004). The

floodplain, apart from the main channel of the Parana River, also includes the Ivinhema and Baia Rivers

and associated with it, the isolated lakes of the Taquaru 9 u area (Souza Filho & Stevaux 2004) (Fig. 1).

Also several streams enter into the system, and these remain largely understudied.

Material

The samples were collected during the expeditions of 2004, 2011 and 2012 in the alluvial valley of the

Upper Parana River. Ostracods were sampled using a rectangular net (28 cm x 14 cm, mesh size ca. 160

pm) hauled close to the sediment-water interface for littoral collections. Floating vegetation was hand-

collected, and roots were thoroughly washed in a bucket. The residues were washed in the same hand net.

Water temperature and dissolved oxygen (Oxymeter-YSI, YSI Incorporated World Headquarters, Yellow

Springs, Ohio, U.S.A.), pH (pHmeter-Digimed, Digimed, Sao Paulo, Brazil) and electrical conductivity

(conductivimeter-Digimed, Digimed, Sao Paulo, Brazil) were measured close to the surface of the water

(Table 1).

All specimens with OC numbers are stored in the Ostracod Collection of the Royal Belgian Institute of

Natural Sciences, Brussels. The material with MZUSP numbers are stored in the Museu de Zoologia da

Universidade de Sao Paulo, Sao Paulo (Table 2).

Morphological analyses

Ostracods were dissected with valves stored dry in micropalaeontological slides and soft parts

in glycerine in sealed slides. Drawings of soft parts were made with a camera lucida attached to a

compound microscope (Olympus, BX51). Valves were illustrated and measured using scanning electron

microscopy (Philips XF30 SEM at RBINS, Brussels).

European Journal of Taxonomy 106: 1-36 (2014)

Table 1. Localities (and some of their characteristics) from which the five new species of candonids were collected. Locality names of 1 to 9 are from

the Ivinhema River System, 10 to 18 from the Baia River System, 19 to 22 from the Parana River System and locality 23 is from the Taquarugu System.

Bold = type localities of the new species. WT = water temperature (°C), EC = electrical conductivity (pS cm 1 ), DO = dissolved oxygen (mg L 1 ), Pa =

Pseudocandona agostinhoi sp. nov., Pc = P. cillisi sp. nov., Pel = P. claudinae sp. nov., Ce = Candobrasilopsis elongata sp. nov., Ca = C. acutis sp. nov.

HIGUTI J. & MARTENS K., Five new Candoninae species from Brazil

European Journal of Taxonomy 106: 1-36 (2014)

Table 2. Locality data of specimens used for illustrations in the present paper.

Pseudocandona agostinhoi sp. nov.

All specimens were sampled on 16 Jul. 2012, Peroba Lake (PAR 710), Eichhornia crassipes.

Valves (male): MZUSP 32655, MZUSP 32660, MZUSP 32661, OC. 3304

Valves (female): OC. 3303, MZUSP 32667, MZUSP 32668, 0C. 3305

Soft parts (male): MZUSP 32655, MZUSP 32653, MZUSP 32656

Soft parts (female): MZUSP 32654

Pseudocandona cillisi sp. nov.

All specimens were sampled on 7 Nov. 2004, Aurelio Lake (PAR 150), floating plants. The specimen

MZUSP 32681 was sampled on 10 Nov. 2004, Baia River (PAR 195), Eichhornia crassipes.

Valves (male): MZUSP 32681, MZUSP 32671, MZUSP 32677, OC. 3362

Valves (female): MZUSP 32672, MZUSP 32678, MZUSP 32680, 0C. 3364

Soft parts (male): MZUSP 32673

Pseudocandona claudinae sp. nov.

All specimens were sampled on 4 Oct. 2012, Matagal Stream (PAR 755), littoral.

Valves (female): MZUSP 32683, MZUSP 32684

Soft parts (female): MZUSP 32682

Candobrasilopsis elongata sp. nov.

All specimens were sampled on 2 Feb. 2011, Patos Lake (PAR 461), Eichhornia crassipes.

Valves (male): MZUSP 32687, MZUSP 32690, MZUSP 32691, MZUSP 32692

Valves (female): MZUSP 32686, MZUSP 32699, MZUSP 32696, MZUSP 32697

Soft parts (male): MZUSP 32685

Candobrasilopsis acutis sp. nov.

All specimens were sampled on 4 Oct. 2012, Matagal Stream (PAR 755), littoral.

Valves (male): MZUSP 32700, MZUSP 32704, OC. 3358

Valves (female): MZUSP 32701, MZUSP 32706, MZUSP 32707, OC. 3360

Soft parts (male): MZUSP 32703

Abbreviations used in text and figures

CpD

CpRl

CpV

CRa

Lpp

LVi

Ante nn ula

Ante nn a

Carapace

Carapace dorsal

Carapace right lateral

Carapace ventral

Caudal ramus

Attachment of caudal ramus

Dorsal branch of caudal ramus

Height of valves

Janet Higuti

Length of valves

Left prehensile palp

Lateral shield of hemipenis

Left valve

Letf valve interior

Mandibula

Medial shield

HIGUTI J. & MARTENS K., Five new Candoninae species from Brazil

Mxl

Rio

Rpp

RBINS

RVi

Maxillula

Rake-lilce organ

Right prehensile palp

Royal Belgian Institute of Natural Sciences, Brussels, Belgium

Right valve interior

First thoracopod, sometimes called Maxilla = Mx2

Second thoracopod

Third thoracopod

Valve

Ventral branch of caudal ramus

Width of carapace

The nomenclature of the limb chaetotaxy mostly follows Broodbakker & Danielopol (1982), for the

second antenna we follow the revised model proposed by Martens (1987), while for the second and third

thoracopods, Meisch’s nomenclature (2000) is followed. Higher taxonomy of the Ostracoda follows the

synopsis by Horne et al. (2002).

Results

Taxonomic descriptions

Class Ostracoda Fatreille, 1806

Subclass Podocopa G.W. Muller, 1894

Order Podocopida G.O. Sars, 1866

Suborder Cypridocopina Baird, 1845

Superfamily Cypridoidea Baird, 1845

Family Candonidae Kaufmann, 1900

Subfamily Candoninae Kaufmann, 1900

Genus Pseudocandona Kaufmann, 1900

Type species

Candona insculpta G.W. Mueller, 1900 (N.B. Candona pubescens Koch, 1837 is an unused senior

synonym of this species).

Other congeneric species in South and Central America (Neotropical)

Pseudocandona antilliana Broodbakker, 1983; Pseudocandona caribbeana Broodbakker, 1983;

Pseudocandona cubensis Broodbakker, 1983; Pseudocandona geratsi Broodbakker, 1983;

Pseudocandona annae (Mehes, 1914) following Broodbakker (1983) and Karanovic & Datry (2009).

Diagnosis

Carapace variously shaped, usually relatively short and stout, rarely elongated or triangular in lateral

view. Surface of adults valves smooth or pitted, usually with long, stiff and perpendicularly attached

setae. FV overlaps RV. Setal group of the second segment of the mandibular palp with 3 to 5 setae (plus

beta-seta). Basal segment of T3 with 3 setae (dl, d2 and dp), medial seta of penultimate segment always

missing, terminal segment with one short (hi) and two long setae (h2 and h3). Hemipenis with at least

three distal lobes (a, b, h).

European Journal of Taxonomy 106 : 1-36 ( 2014 )

Remarks

1. All three new species described here have a group of 4 ventral setae (+ 1 (3-seta) on the second

segment of the Md-palp. In addition, claws G1 and GM of the A2 (in females) and claws G2, GM

and zl (in males) are long. The combination of these characters places the three new species in the

car ib be ana-group of the genus according to the scheme of Namiotko & Danielopol (2004). See the

general discussion of the present paper for further discussion on this species group.

2. The genus Pseudocandona is speciose, and most species occur in the Palaearctic and the Nearctic,

including a large radiation in the ancient Lake Baikal (Siberia). In the differential diagnoses in the

present paper we will only compare our new species to those already described from the Neotropical

region, although of course their morphology was also checked against the Holarctic species of which

good descriptions and illustrations are available.

3. Pseudocandona agostinhoi sp. nov. is described in full, the descriptions of the other two new species

in this genus are abbreviated.

Pseudocandona agostinhoi sp. nov.

urn:lsid:zoobank.org:act:479455Al-91AD-4866-AADB-B8D6EB3892Bl

Figs 2-5

“Candona ” sp. 4 nov. sp. Higuti et al, 2007: 1935.

“Candona” sp. 2 n. sp. Higuti et al ., 2009c: 664.

LL Candona ,, sp. 2 n. sp. Higuti et al., 2010: 267.

Diagnosis

Carapace rather elongated in lateral view, and with LV overlapping RV on all sides, especially at

anterior and posterior extremities. Anterior calcified inner lamella in both valves relatively broad. LV

with anterior and posterior inner list, situated well away from the valve margin. Second segment of Md

palp ventrally with 4 setae (+ 1 (3-seta). Prehensile palps distally hoolc-like and set with two thin, sub-

apical setae. Penultimate segment of T3 not divided; seta hi slightly more than half the length of seta

h2. Hemipenis with lobe a small and sub-quadrate, with rounded distal margin; lobes b and h largely

overlapping, except on ventro-apical side, asymmetrically rounded and pointed towards the ventral side.

Etymology

The new species is named after Prof. Dr Angelo Antonio Agostinho (Nupelia, Universidade Estadual de

Maringa, Maringa, Brazil), in recognition of his vast contributions to the knowledge on the biodiversity

and the conservation of Brazilian freshwater environments, ma inl y in ichthyology, and also to

acknowledge him for being one of the founders of Nupelia (Research Group in Limnology, Ichthyology

and Aquaculture).

Type material

All type material was collected on 16 Jul. 2012 by JH and students, by washing roots of Eichhornia

crassipes (PAR 710) in a bucket and by filtering the material in a hand net with mesh size 160 pm. See

Table 1 for measurements of water chemistry at time of collecting.

Holotype

$, with soft parts dissected in glycerine in a sealed slide, and with valves stored dry in a

micropalaeontological slide (MZUSP 32653).

Allotype

5, dissected and stored like the holotype (MZUSP 32654).

HIGUTI J. & MARTENS K., Five new Candoninae species from Brazil

Paratypes

8 33 dissected and stored like the holotype (MZUSP 32655, MZUSP 32656, MZUSP 32657,

MZUSP 32658, MZUSP 32659, OC.3302, OC.3306, OC.3307); three 3 carapaces stored dry in

micropalaeontological slides (MZUSP 32660, MZUSP 32661, OC.3304); eight $$ dissected and stored

like the holotype (MZUSP 32662, MZUSP 32663, MZUSP 32664, MZUSP 32665, MZUSP 32666,

OC.3303, OC.3308, OC.3309); three $ carapaces stored dry in micropalaeontological slides (MZUSP

32667, MZUSP 32668, OC.3305).

Other material investigated

A 3 (MZUSP 32669) and a $ (MZUSP 32670) carapace stored dry in micropalaeontological slides from

Ventura Lake (PAR 1) and Pintado Lake (PAR 213), respectively, are also deposited in the Museu de

Zoologia da Universidade de Sao Paulo, Sao Paulo.

Type locality

BRAZIL: Peroba Lake in Upper Parana River floodplain in roots of the floating plant species Eichhornia

crassipes. Coordinates: 22°54 , 39.4” S, 53°38 , 34.5” W.

Differential diagnosis

Pseudocandona agostinhoi sp. nov. can be distinguished from P. geratsi Broodbakker, 1983 by the

shape of the valves (dorsal margin straight and oblique in the middle in Pa., rounded in P.g.), of the

hemipenis (lobe a with distal margin rounded in P.a., straight in P.g.) and of the prehensile palps (Rpp

evenly rounded in P.a., with blunt angles in P.g.; Lpp more narrow and position of setae more distal in

P.a.). Pseudocandona antilliana Broodbakkker, 1983 also has the middle part of the dorsal margin of the

valves straight in the female, but running parallel to the ventral margin, not sloping as in P. agostinhoi

sp. nov., while also in this species the distal margin of lobe a of the hemipenis is straight (rounded in

P. agostinhoi sp. nov.). Pseudocandona caribbeana Broodbakker, 1983 and P. cubensis Broodbakkker,

1983 both have short and high carapaces, with straight dorsal margin parallel to the ventral margin, and

with at least P. caribbeana with an anterior rostrum in dorsal view. Both of the latter species are known

from females only.

Pseudocandona agostinhoi sp. nov. differs from P. annae (Mehes, 1914) in several aspects of its

morphology. The latter species is more elongated with a longer straight section of the dorsal margin. In

P. agostinhoi sp. nov., the LV overlaps the RV on all sides, and especially strongly along the anterior

and posterior margins, whereas the overlap appears to be minimal according to the drawings by Mehes

(1914). The Rpp has a fully rounded dorsal-distal part in P. agostinhoi sp. nov., whereas this has a

straight margin in P. annae. Finally, lobes a, b and h of the hemipenis in P. agostinhoi sp. nov. are almost

equally long, whereas lobe a is much shorter in P. annae.

Description of male

Carapace in right lateral view (Fig. 2C) rather elongated, showing LV overlapping RV on all sides, with

larger overlap at both anterior and posterior extremities, external surface of valves smooth. Cp in dorsal

and ventral views (Fig. 2D-E) lancet-shaped, sharply pointed anteriorly, more broadly so posteriorly,

greatest width situated slightly behind the middle.

Valves (Fig. 2A-B) with greatest height situated behind the middle (and with a blunt angle there),

posteriorly rather broadly rounded, anteriorly less so. RV and LV of similar shape; anterior calcified

inn er lamella more broadly rounded than posterior calcified inner lamella. LV with anterior and posterior

inner lists, situated closely to valve margin, yet clearly separated from it.

European Journal of Taxonomy 106: 1-36 (2014)

Fig. 2. Valves of Pseudocandona agostinhoi sp. nov. A-E S and F-J $. A, F. LV, internal view (A =

MZUSP 32655, F = OC. 3303). B, G. RV, internal view (B = MZUSP 32655, G = OC. 3303). C, H. Cp,

right lateral view (C = MZUSP 32660, H = MZUSP 32667). D, I Cp, dorsal view (D =MZUSP 32661,1 =

MZUSP 32668). E, J. Cp, ventral view (E = OC. 3304, J = OC. 3305). Scale bars: A-J = 500 pm.

HIGUTI J. & MARTENS K., Five new Candoninae species from Brazil

Fig. 3. Limbs of Pseudocandona agostinhoi sp. nov. (c?). A. A2 (MZUSP 32656). B. A2, detail of the

last segment (MZUSP 32656). C. A1 (OC. 3302). D. Mxl, respiratory plate (OC. 3302). E. T3 (MZUSP

32655). F. Md, coxal plate (MZUSP 32653). G. T2 (MZUSP 32653). H. T1 (MZUSP 32653). Scale bars:

A-H = 50 pm.

European Journal of Taxonomy 106: 1-36 (2014)

Fig. 4. Limbs of Pseudocandona agostinhoi sp. nov. (c?)- A. Mxl (MZUSP 32653). B. Md palp (MZUSP

32655). C. Zenker organ (MZUSP 32653). D. Right prehensile palp (MZUSP 32653). E. Left prehensile

palp (MZUSP 32653). F. Caudal ramus (MZUSP 32653). G. Attachment of the caudal ramus (MZUSP

32653). H. Hemipenis (MZUSP 32653). Scale bars: A-H = 50 pm.

HIGUTI J. & MARTENS K., Five new Candoninae species from Brazil

Fig. 5. Limbs of Pseudocandona agostinhoi sp. nov. ($). A. A2, detail of the last segments (MZUSP

32654). B. Caudal ramus (MZUSP 32654). C. T1 (MZUSP 32654). Scale bars: A-C = 50 pm.

European Journal of Taxonomy 106: 1-36 (2014)

A1 (Fig. 3B) with basal segment with 2 relatively short dorsal setae, one at ca. mid-length, the other

sub-apical, and two dorsal, sub-apical setae: one long, the other slightly longer than half the length of

the first. Second segment with one dorso-apical seta (reaching beyond basis of fourth segment), no

ventro-apical seta present. Third segment with one sub-apical dorsal set, reaching beyond basis of fourth

segment, no ventral seta present. Fourth and fifth segments with two longer dorso-apical and 1 shorter

ventro-apical setae. Sixth and seventh segments most elongate of all; sixth segment with two longer

dorso-apical and one shorter ventro-apical setae. Seventh (terminal segment) slightly sinuous, bearing

one short and two longer setae and one aesthetasc Ya, the latter shorter than the shorter seta.

A2 (Fig. 3A) with basal segment broad, wide and relatively long, basally with 2 unequal setae, one

relatively long, the other ca. half as long as the first; one long ventro-apical seta also present. Remnant of

exopod consisting of a short plate, with one long and two unequal shorter setae. Endopod consisting of

4 segments (penultimate segment divided). First endopodal segment long, carrying one long aesthetasc

Y at the base of the ventral side, and one long and one short ventro-apical setae. Second endopodal

segment shorter and smaller, rectangular with one ventral aesthetasc yl inserted about mid length, 3

t-setae, with tl almost same length as t2, t2 and t3 modified into male-bristles; dorso-apically with 1

seta. Third endopodal segment with apical chaetotaxy sexually dimorphic, with zl and z3 being short

setae, z2 being modified into a long claw; G2 a long claw, G1 a long seta and G3 a short seta, aesthetasc

y2 short and ventro-apically inserted. Terminal (fourth) segment small (Fig. 3B), distally with claws GM

(long) and Gm (short), aesthetasc y3 with its companion seta, fused at the basis and of unequal length,

and seta g, longer than y3.

Md with coxa (Fig. 3F) relatively slender, distally set with a series of strong teeth.

Md-palp (Fig. 4B) consisting of 4 segments. Basal segment dorsally carrying the respiratory plate (not

fully shown), ventro-apically with 2 long setae, 1 hirsute si seta, the other seta smooth, the minute alpha

seta, and seta s2 short, stout and hirsute. Second segment dorso-apically with 2 setae of unequal length,

ventro-apically with a group of 5 setae: 3 long, one shorter, and the short beta-seta. Third segment with

a group of 3 smooth dorso-subapical setae (2 long and one shorter), a central group of two setae, the

ventral-most one being the gamma seta, and ventro-apically with 2 smooth setae. Terminal segment

sub-quadrate, apically set with two large claw-like setae, one distally hirsute and three shorter setae of

unequal length.

Mxl (Fig. 4A) with a basal (basipodite) part carrying a large respiratory plate (exopodite), 3 endites and

a two-segmented palp (endopodite). Respiratory plate elongated, carrying more than 10 long respiratory

rays (Fig. 3D). Palp with first segment carrying 3+1 apical setae; terminal segment short and broad,

carrying 2 longer claws and 4 short setae (one shorter than the other). Chaetotaxy of three endites

impossible to determine. Sideways directed bristles near first endite stout, one long and another short.

Tl (Fig. 3H) consisting of basal part (basipodite), carrying respiratory plate (not shown), a palp

(endopodite, modified to prehensile palp in males) and an endite distally set with ca. 10 setae of different

morphology and length. Basal plate set with one long and stout ‘b+seta, a long and more slender ‘d’-seta

and one ‘a’-seta. Prehensile palps (Figs 4D-E) distally hook-like and set with two, thin sub-apical setae;

palps slightly asymmetrical, left prehensile palp (Fig. 4E) margin tightly rounded.

T2 (walking limb. Fig. 3G) with 4-segmented endopodite (penultimate segment divided) and elongated.

First segment with short seta dl. Knee-segment devoid of seta d2. First segment of endopod elongated,

with short ventro-apical seta. Second segment also with one short ventro-apical seta. Third segment with

two ventro-apical setae, one short, one slightly longer. Terminal segment with one short apical and one

short sub-apical seta and a long apical claw.

HIGUTI J. & MARTENS K., Five new Candoninae species from Brazil

Table 3. Measurements of specimens of the three new species of Pseudocandona and two new species of Candobrasilopsis described in the present

paper.

European Journal of Taxonomy 106: 1-36 (2014)

T3 (cleaning limb. Fig. 3E) as typical of the family. First segment with three setae, one short medial (dl),

one short subapical (d2) and one long apical (dp). Second segment without seta. Penultimate segment

with one long subapical seta (g). Terminal segment carrying three setae: 2 long, one of which hirsute (h2

and h3), and one short and hirsute (hi).

Caudal ramus (= fiirca. Fig. 4F) with stout ramus and two stout apical claws. Long proximal setae, distal

seta a small spine. Attachhment to caudal ramus (Fig. 4G) long and stout, distally bifurcated and with

additional lateral branch at ca. mid-length. Length ratio ramus / largest claw = 1.92.

Hemipenis (Fig. 4H) relatively small, with lobe a short, and with distal margin rounded, lobes b and h

largely overlapping, reaching about as far as lobe a, but asymmetrially pointing towards the ventral side.

Labyrinth with first (ascending) branch long, thin and slender; post-labyrinthal spermiduct S-shaped and

simple.

Zenker organ (Fig. 4C) short and broad, with at least 5 spinous whorls.

Description of female

Valves (Fig. 2F-G) similar to those in the male; Cp (Fig. 2H) in lateral view with bigger overlap at

the anterior extremity; Cp in D and V views (Figs 2I-J) equally narrow and lancet-shaped, in V view

highlighting the overlap.

Soft parts (Fig. 5) largely as in the male, but with sexually dimorphic A2 and Tl.

A2 (Fig. 5A) with setae 11-4 setae like, not transformed; zland z3 short and slender setae, z2 longer and

slender. Claws Gl, G3 and GM all reaching to about the same point and claw G2 short.

Tl (Fig. 5C) with basal part as in the male and an endite distally set with ca. 13 setae of different

morphology and length. Endopod a broad palp, with three unequal setae.

Caudal ramus (Fig. 5B) with stout ramus and two stout apical claws. Long proximal setae, distal seta a

small spine. Length ratio ramus / largest claw = 1.79.

Measurements

See Table 3.

Pseudocandonci cillisi sp. nov.

urn:lsid:zoobank.org:act:DC713AC3-23AA-4056-AE25-020DD6A1257C

Figs 6-7

“Candor” sp. 1 n. sp. Higuti et al., 2009c: 664.

“Candona ” sp. 1 n. sp. Higuti et al., 2010: 267.

Diagnosis

Valves small, high and short in lateral view. Anterior calcified inner lamella in both valves relatively

broad; posterior calcified inner lamella very narrow. Second segment of Md palp with 4 setae. Prehensile

palps distally hoolc-like, relatively short and set with one sub-apical spine and one sub-apical setae.

Penultimate segment of T3 not divided; seta hi ca. Vs of the length of seta h2. Hemipenis with lobe

a short and broad, distal margin only slightly rounded, lobes b and h almost completely overlapping,

asymmetrically rounded and pointed towards the ventral side.

HIGUTI J. & MARTENS K., Five new Candoninae species from Brazil

Fig. 6. Valves of Pseudocandona cillisi sp. nov. A-E $ and F-J $. A, F. LV, internal view (A = MZUSP

32681, F = MZUSP 32672). B, G. RV, internal view (B = MZUSP 32681, G = MZUSP 32672). C, H.

Cp, right lateral view (C = MZUSP 32671, H = MZUSP 32678). D, I. Cp, dorsal view (D = OC. 3362,

I = OC.3364). E, J. Cp, ventral view (E = MZUSP 32677, J = MZUSP 32680). Scale bars: A-B = 200

pm; C-E, H-J = 500 pm; F-G = 300 pm.

European Journal of Taxonomy 106: 1-36 (2014)

Fig. 7. Limbs of Pseudocandona cillisi sp. nov. ((?). A. T3 (MZUSP 32673). B. T2 (MZUSP 32673).

C. Md palp (MZUSP 32673). D. Caudal ramus (MZUSP 32673). E. Attachment of the caudal ramus

(MZUSP 32673). F. Left prehensile palp (MZUSP 32673). G. Hemipenis (MZUSP 32673). H. Right

prehensile palp (MZUSP 32673). Scale bars: A-H = 50 pm.

HIGUTI J. & MARTENS K., Five new Candoninae species from Brazil

Etymology

The new species is named in honour of Julien Cillis (RBINS), in recognition of his continuous technical

assistance with the scanning electron microscopy (SEM) images of ostracod valves at RBINS for more

than 2 decades.

Type material

All type material was collected on 7 Nov. 2004 by the authors, by washing roots of floating plants (PAR

150) in a bucket and the material was filtered in a hand net with mesh size 160 pm. See Table 1 for

measurements of water chemistry at time of collecting.

Holotype

S carapace stored dry in a micropalaeontological slide (MZUSP 32671).

Allotype

5, with soft parts dissected in glycerine in a sealed slide, and with valves stored dry in a

micropalaeontological slide (MZUSP 32672).

Paratypes

A S dissected,with LV stored dry in a micropalaeontological slide (OC. 3363); four SS dissected and

with the valves lost (MZUSP 32673, MZUSP 32674, MZUSP 32675, MZUSP 32676); two S carapaces

stored dry in micropalaeontological slides (MZUSP 32677, OC. 3362); four § carapaces stored dry in

micropalaeontological slides (MZUSP 32678, MZUSP 32679, MZUSP 32680, OC. 3364).

Other material investigated

A S dissected, with soft parts lost and with valves stored dry in a micropalaeontological slide (MZUSP

32681) from Baia River (PAR 195) is also deposited in the Museu de Zoologia da Universidade de Sao

Paulo, Sao Paulo.

Type locality

BRAZIL: Aurelio Lake in Upper Parana River floodplain in a mix of floating macrophyte species.

Coordinates: 22 0 41 , 36.5 ,, S, 53°13 , 52” W.

Differential diagnosis

Pseudocandona cillisi sp. nov. has valves which are much shorter and higher than those of P. geratsi

and P. antilliana , while the (straight) dorsal margin is much more sloping than in P. caribbeana and P.

cubensis. The new species also differs from P. annae Mehes, 1914 in the shape of the valves, which are

more elongated in the latter species.

Description of male

Valves (Fig. 6A-B) small, high and short in lateral view, with greatest height situated behind the middle

(and bluntly pointed there), anteriorly and posteriorly rather broadly rounded, RV and LV of highly

similar shape; anterior calcified inner lamella broadly rounded, posterior calcified inner lamella narrow

and narrower towards the dorsal side.

Carapace in right lateral view (Fig. 6C) showing LV overlapping RV slightly on all sides; anteriorly and

posteriorly widely beyond the RV, external surface of valves smooth.

In dorsal view (Fig. 6D), carapace lancet-shaped, sharply pointed in the anteriorly, more broadly so

posteriorly, greatest width situated well behind the middle.

European Journal of Taxonomy 106: 1-36 (2014)

Carapace also in ventral view (Fig. 6E) lancet-shaped, with anterior side more acutely pointed than

posterior side, in V view with slight rostrum.

Md-palp (Fig. 1C) consisting of 4 segments. Basal segment dorsally carrying the respiratory plate (not

shown), ventro-apically with 2 long setae (only one hirsute = si), the alpha seta and a short, stout and

hirsute seta s2. Second segment dorso-apically with 2 setae of unequal length, ventro-basally with a

short, stout and hirsute seta and ventro-apically with a group of 5 setae: 2 long setae, 2 shorter (one

hirsute), and the short beta-seta. Third segment with a group of 3 smooth dorso-subapical setae of

unequal length, a central group of two setae, one of which being the gamma seta, and a ventro-apical

group of 2 smooth setae. Terminal segment subquadrate, chaetotaxy as in P. agostinhoi sp. nov.

Tl: Prehensile palps (Fig. 7F, H) distally hook-like and set with two unequal sub-apical setae; palps

slightly asymmetrical, robust left prehensile palp (Fig. 7F) margin tightly rounded.

T2 (walking limb - Fig. 7B) with 4-segmented endopodite (penultimate segment divided) and elongated.

First segment with short seta dl. Knee-segment devoid of seta d2. First segment of endopod elongated,

with short ventro-apical seta. Second segment also with one short ventro-apical seta. Third segment with

two ventro-apical setae, one short, one slightly longer. Terminal segment with one short apical and one

short sub-apical seta and a long apical claw.

T3 (cleaning limb, Fig. 7A) as typical of the family. First segment with three setae, one short medial

(dl), one short subapical (d2) and one long apical (dp). Second segment without seta. Penultimate

segment with one long subapical seta (g). Terminal segment carrying three setae: 2 long (h2 and h3) and

one very short (hi).

Caudal ramus (furca. Fig. 7D) with stout ramus and two stout apical claws. Tong proximal setae, distal

seta a small spine. Attachment to caudal ramus (Fig. 7E) long and stout, distally bifurcated and with 2

additional lateral branches. Length ratio ramus / largest claw = 2.0.

Hemipenis (Fig. 7G) relatively small, with lobe a short and broad, distal margin only slightly rounded,

lobes b and h almost completely overlapping, asymmetrically rounded and pointed towards the ventral

side. Labyrinth with first (ascending) branch long, thin and slender; post-labyrinthal spermiduct S-shaped

and simple.

Description of female

Valves (Fig. 6F-G): Cp in lateral view (Fig. 6H) and in D and V views (Fig. 6I-J) similar to those in the

male.

Measurements

See Table 3.

Pseudocandona claudinae sp. nov.

um:lsid:zoobank.org:act:198F7F5F-49D0-4883-845A-386E0CAA4970

Figs 8-9

Diagnosis

Valves in lateral view elongated and with dorsal margin rounded, greatest height situated well behind

the middle; anterior calcified inner lamella in both valves broader than very narrow posterior calcified

inner lamella. Carapace in dorsal and ventral views relatively narrow; LV overlapping RV on all sides.

HIGUTI J. & MARTENS K., Five new Candoninae species from Brazil

Seta hi less than half the length (< ca . A) of seta h2. Caudal ramus with distal seta clearly developed, not

spine-like.

Etymology

The new species is named in honour of Claudine Behen (RBINS), in recognition of her continued

technical assistance with the line drawings of ostracod limbs over many years at RBINS.

Type material

All type material was collected on 4 Oct. 2012 by the authors and students, by hand net with mesh size

160 pm. See Table 1 for measurements of water chemistry at time of collecting.

Holotype

5, with soft parts dissected in glycerine in a sealed slide, and with valves stored dry in a micro-

palaeontological slide (MZUSP 32682).

Fig. 8. Valves of Pseudocandona claudinae sp. nov. ($). A. LV, internal view (MZUSP 32683). B. RV,

internal view (MZUSP 32683). C. Cp, right lateral view (MZUSP 32684). D. Cp, dorsal view (MZUSP

32684). E. Cp, ventral view (MZUSP 32684). Scale bars: A-E = 500 pm.

European Journal of Taxonomy 106: 1-36 (2014)

Paratypes

3 $$ dissected and stored like the holotype (MZUSP 32683, OC3365, OC3366); a $ carapace stored

dry in micropalaeontological slides (MZUSP 32684).

Type locality

BRAZIL: Matagal Stream in Upper Parana River floodplain in sediment amongst grasses and other

terrestrial vegetation in very shallow water (named littoral in Table 1). Coordinates: 22°54 , 26.4” S,

53°32’27.7”W.

Differential diagnosis

Pseudocandona claudinae sp. nov. can be distinguished from P antilliana, P. caribbeana and P. cubensis

by the elongated shape of the valves and the rounded dorsal margin. Pseudocandona claudinae sp. nov.

appears to be closely related to P. geratsi, but the female valves of P. claudinae sp. nov. resemble those

of the males in P. geratsi , while the female valves in this latter species are shorter and higher than those

of the male. As no males of P. claudinae sp. nov. are known, the copulatory appendages cannot be

Fig. 9. Limbs of Pseudocandona claudinae sp. nov. (§). A. Md palp (MZUSP 32682). B. Attac hm ent

of the caudal ramus (MZUSP 32682). C. T2 (MZUSP 32682). D. Caudal ramus (MZUSP 32682). E. T3

(MZUSP 32682). Scale bars: A-E = 50 pm.

HIGUTI J. & MARTENS K., Five new Candoninae species from Brazil

compared. In addition, the attachment of the caudal ramus in P. geratsi has two lateral branches, whereas

there is only one in P. claudinae sp. nov.

Pseudocandona claudinae sp. nov. differs from P. annae by the more elongated valves, with a dorsal

margin which is fully rounded (with a straight part in P. annae).

Description of female

Valves (Fig 8A-B) small and short, with greatest height situated behind the middle. Anteriorly and

posteriorly rather broadly rounded, RV and FV of highly similar shape; anterior calcified inner lamella

broadly rounded, posterior calcified inner lamella very narrow and almost disappearing towards the

dorsal side.

Carapace in right lateral view (Fig. 8C) showing FV overlapping RV slightly on all sides; external

surface of valves smooth. In lateral view dorsally more rounded.

In dorsal view (Fig. 8D), carapace lancet-shaped, sharply pointed anteriorly, more broadly so posteriorly.

Carapace in ventral view (Fig. 8E) lancet-shaped, with anterior side more acutely pointed than posterior

side.

Md-palp (Fig. 9A) consisting of 4 segments. Basal segment dorsally carrying the respiratory plate (not

shown), ventro-apically with 2 long setae (only one hirsute (= si), the thin and smooth alpha seta and a

short and stout hirsute seta (= s2). Second segment dorso-apically with 2 setae of unequal length, ventro-

basally with a short, stout and hirsute seta and ventro-apically with a group of 5 setae: 2 long setae, 2

shorter, and the short and beta-seta. Third segment with a group of 3 dorso-subapical setae of unequal

length, all smooth, a central group of two setae, one of which being the gamma seta, and a ventro-apical

group of 2 setae, one long and one short, both smooth. Terminal segment subquadrate, chaetotaxy as in

P. ago s tin hoi sp. nov.

T2 (walking limb. Fig. 9C) with 4-segmented endopodite (penultimate segment divided) and elongated.

First segment with short seta dl. Knee-segment devoid of seta d2. First segment of endopod elongated,

with short ventro-apical seta. Second segment also with one short ventro-apical seta. Third segment with

two ventro-apical setae, one short, one very long. Terminal segment with one short apical and one short

sub-apical seta and a long apical claw.

T3 (cleaning limb. Fig. 9E) as typical of the family. First segment with three setae, one short medial (dl),

one short subapical (d2) and one long apical (dp). Second segment without seta. Penultimate segment

with one long subapical seta (g). Terminal segment carrying three setae: 2 long (h2 and h3) and one very

short (hi).

Caudal ramus (furca. Fig. 9D) with stout ramus and two stout apical claws. Fong proximal setae, distal

seta a small spine. Attachment to caudal ramus (Fig. 9B) long and stout, distally bifurcated and with

additional lateral branch at ca. mid-length.

Male unknown

Measurements

See Table 3.

European Journal of Taxonomy 106: 1-36 (2014)

Genus Candobrcisilopsis Higuti & Martens, 2012

Type species

Candobrasilopsis rochai Higuti & Martens, 2012

Other species

Candobrasilopsis acutis sp. nov.; C. brasiliensis (Sars, 1901); C. elongata sp. nov.

Diagnosis

Anterior calcified inner lamella broad, anterior inn er margin sinuous, posterior calcified inner lamella

narrow, posterior inner margin running parallel to valve margin. Terminal Md-palp segment short

(L < 1.5 x basal width). Prehensile palps one-segmented, base inflated, distal finger long, hook-like.

Hemipenis of triangular type, without protruding lobes a and b. Caudal ramus without proximal seta,

distal seta reduced to a small spine.

Remarks

As the genus was characterised by a full description of its type species, C. rochai , by Higuti & Martens

(2012b), the descriptions of the following two new species are shortened.

Candobrasilopsis elongata sp. nov.

urn:lsid:zoobank.org:act:197FB8BE-2703-4FA6-97CA-2D40A8342325

Figs 10-11

Candonopsis sp. 3 n. sp. in Higuti et al., 2009c: 664.

Candonopsis sp. 3 n. sp. in Higuti et al., 2010: 267.

Diagnosis

A species with very elongated valves, greatest height well behind the middle; in lateral view with

dorsal margin straight for more than half the total length. Anterior calcified inn er lamella in both valves

broad, almost % of total length and with inner margin sinuous, not parallel to valve margin. Both valves

externally set with elongated micro-ridges.

Prehensile palps one-segmented. Right prehensile palp set with two subequal and robust lateral setae.

Left prehensile palp larger and more elongated, with hook-like distal part, distal tip slightly swollen.

Hemipenis atypical of the genus, with leaf-like lobe Is, distally rounded and proximally narrowing, lobe

ms broad, distally with almost straight margin, ventro-distally bluntly pointed.

Etymology

The present species is named after the very elongated carapace.

Type material

All type material was collected on 2 Feb. 2011 by JH and students, by washing roots of Eichhornia

crassipes (PAR 461) in a bucket and by filtering the material in a hand net with mesh size 160 pm. See

Table 1 for measurements of water chemistry at time of collecting.

Holotype

$, with soft parts dissected in glycerine in a sealed slide, and with valves stored dry in a micro-

palaeontological slide (MZUSP 32685).

HIGUTI J. & MARTENS K., Five new Candoninae species from Brazil

Fig. 10. Valves of Candobrasilopsis elongata sp. nov. A-F <$ and G-F $. A, G. FV, internal view (A =

MZUSP 32687, G = MZUSP 32686). B, H. RV, internal view (B = MZUSP 32687, H = MZUSP 32686).

C, I. Cp, right lateral view (C = MZUSP 32690,1 = MZUSP 32699). D, J. Cp, right lateral view detail

(D = MZUSP 32690, J = MZUSP 32699). E, K. Cp, dorsal view (E = MZUSP 32692, K = MZUSP

32696). F, L Cp, ventral view (F = MZUSP 32691, F = MZUSP 32697). Scale bars: A-C, E-I, K-F =

500 pm; D, J = 20 pm.

European Journal of Taxonomy 106: 1-36 (2014)

Allotype

A §, dissected and stored like the holotype (MZUSP 32686).

Paratypes

Five S3 dissected and stored like the holotype (MZUSP 32687, MZUSP 32688, MZUSP 32689, OC.

3353, OC. 3354); three S carapaces stored dry in micropalaeontological slides (MZUSP 32690, MZUSP

32691, MZUSP 32692); five $ $ dissected and stored like the holotype (MZUSP 32693, MZUSP 32694,

OC. 3355, OC. 3356, OC. 3357) and a $ dissected with valves lost (MZUSP 32695); four $ carapaces

Fig. 11. Limbs of Candobrasilopsis elongata sp. nov. (f). A. Left prehensile palp (MZUSP 32685). B.

Right prehensile palp (MZUSP 32685). C. Md palp (MZUSP 32685). D. Attachment of the caudal ramus

(MZUSP 32685). E. Caudal ramus (MZUSP 32685). F. Hemipenis (MZUSP 32685). Scale bars: A-E = 50

pm; F = 100 pm .

HIGUTI J. & MARTENS K., Five new Candoninae species from Brazil

stored dry in micropalaeontological slides (MZUSP 32696, MZUSP 32697, MZUSP 32698, MZUSP

32699).

Type locality

BRAZIL: Patos Lake in Upper Parana River floodplain in floating plants (E. crassipes). Coordinates:

22°49 , 48.2” S, 53°33’20.3” W.

Differential diagnosis

Candobrasilopsis elongata sp. nov. differs from C. rochai and C. brasiliensis by the more elongated

shape of the valves, and from C. acutis sp. nov., which also has elongated valves, by the presence of

external micro-ridges on the valves. Candobrasilopsis elongata sp. nov. can be distinguished from its

three congeners by the atypical hemipenis, which has a lobe Is with rounded (not pointed) distal margin,

and a lobe ml with almost straight distal margin (not rounded).

Description of male

Carapace in right lateral view (Fig. IOC) showing LV overlapping RV slightly on all sides. Carapace,

in dorsal (Fig. 10E) and ventral views (Fig. 10F), narrow and lancet-shaped, with anterior and posterior

side pointed, greatest width situated well behind the middle.

Both valves (Fig. 10A-B) very elongated, with greatest height situated behind the middle (and bluntly

pointed there), anteriorly rather broadly and posteriorly rather narrowly rounded, RV and LV of highly

similar shape; anterior calcified inner lamella broadly rounded, posterior calcified inner lamella very

narrow and almost disappearing towards the dorsal side; inner margin of anterior calcified inner lamella

slightly sinuous in both valves. External surface of both valves striated (Fig. 10D).

Al, A2, Mxl, Tl-3 as typical of the genus and not illustrated.

Md-palp (Fig. 11C) consisting of 4 segments, terminal segment short (L < 1.5 x basal width).

Prehensile palps (Fig. 11A-B) one-segmented, distally hook-like and set with two unequal and robust,

sub-apical setae; both palps slightly asymmetrical, left prehensile palp (Fig. 11 A) basally slightly more

swollen.

Caudal ramus (Fig. 11E) with stout ramus and two stout apical claws. Proximal setae missing, distal

seta a small spine. Attachment to caudal ramus (Fig. 11D) long and stout, distally bifurcated and with

additional lateral branch at ca. mid-length. Length ratio ramus / largest claw = 1.85.

Hemipenis (Fig. 1 IF) atypical of the genus, with lobe Is leaf-like, distally rounded and proximally

narrowing, lobe ms broad, distally with almost straight margin, ventro-distally bluntly pointed; an

additional small ventral lobe present proximal of lobe ms. Internal labyrinth short and stout, postlabyrinthal

spermiduct narrow and straight; internal copulatory process stout, beak-like and sharply pointed.

Description of female

Valves (Fig. 10G-H) and Cp (Fig. 101—J) in lateral view similar to those in the male. Cp in D and V

views (Fig. 10K-L) equally narrow and lancet-shaped, with anterior and posterior side pointed.

Soft parts as typical of the genus and, except for sexually dimorphic ones, similar to those of the male.

Measurements

See Table 3.

European Journal of Taxonomy 106: 1-36 (2014)

Candobrasilopsis acutis sp. nov.

urn:lsid:zoobank.org:act:2894DAF9-9CF7-4A60-B25F-6ED4BFEF8E09

Figs 12-13

Diagnosis

A species with very elongated valves, greatest height well behind the middle; carapace in lateral view

with weakly rounded dorsal margin, nearly straight. Anterior calcified inner lamella in both valves

broad, almost Vs of total length and with inner margin sinuous, not parallel to valve margin. External

valve surface smooth, devoid of ridges.

Both prehensile palps one-segmented. Right prehensile palp with distal part curved, distal tip slightly

expanded, set with two subequal lateral setae. Left prehensile palp narrower and more elongated, with

hoolc-like distal part, distal tip slightly swollen. Hemipenis with triangular and sharply pointed lobe Is.

Etymology

Named after “pointed” (Latin = acutis). The shape of lobe Is of the hemipenis is triangular and distally

very pointed.

Type material

All type material was collected on 4 Oct. 2012 by the authors and students, by hand net with mesh size

160 pm. See Table 1 for measurements of water chemistry at time of collecting.

Holotype

S carapace stored dry in a micropalaeontological slide (MZUSP 32700).

Allotype

5, with soft parts dissected in glycerine in a sealed slide, and with valves stored dry in a

micropalaeontological slide (MZUSP 32701).

Paratypes

A S with soft parts dissected in glycerine in a sealed slide and with LV stored dry in a micropalaeon¬

tological slide (OC. 3359); two SS dissected and with the valves lost (MZUSP 32702, MZUSP 32703);

two S carapaces stored dry in micropalaeontological slides (MZUSP 32704, OC. 3358); a $ with soft

parts dissected in glycerine in a sealed slide and with LV stored dry in a micropalaeontological slide

(MZUSP 32705); a $ with soft parts dissected in glycerine in a sealed slide and with the valves lost (OC.

3361); three $ carapaces stored dry in micropalaeontological slides (MZUSP 32706, MZUSP 32707,

OC. 3360).

Type locality

BRAZIL: Matagal Stream in Upper Parana River floodplain in sediment amongst grasses and other

terrestrial vegetation in very shallow water (named littoral in Table 1). Coordinates: 22 0 54’26.4” S,

53°32 , 27.7” W.

Differential diagnosis

Candobrasilopsis acutis sp. nov. is characterized by the shape of both prehensile palps and especially by

the very pointed lateral shield of the hemipenis, by which it can easily be distinguished from the other

three species in the genus. The shape of the valves of C. acutis sp. nov. is quite similar to those of C.

elongata sp. nov., but C. acutis sp. nov. lacks the clear external striation of the latter species (compare

Figs 10D, J with 12D, H).

HIGUTI J. & MARTENS K., Five new Candoninae species from Brazil

Fig. 12. Valves of Candobrasilopsis acutis sp. nov. C-F S and A-B, G-J 5. A. FV, internal view. (MZUSP

32701) B. RV, internal view (MZUSP 32701). C, G. Cp, right lateral view (C = MZUSP 32700, G =

MZUSP 32706). D, H. Cp, right lateral view, detail (D = MZUSP 32700, H = MZUSP 32706). E, L Cp,

dorsal view (E = MZUSP 32704,1 = MZUSP 32707). F, J. Cp, ventral view (F = OC. 3358, J = OC. 3360).

Scale bars: A-C, E-G, I-J = 500 pm; D, H = 20 pm.

European Journal of Taxonomy 106: 1-36 (2014)

Description of male

Carapace very elongated in right lateral view (Fig. 12C), with greatest height situated behind the middle,

anteriorly and posteriorly broadly rounded; LV overlapping RV slightly on all sides; external surface of

valves smooth (Fig. 12D). In dorsal view (Fig. 12E) and ventral view (Fig. 12F) carapace lancet-shaped,

sharply pointed anteriorly, more broadly so posteriorly.

Internal valve anatomy as in the female (see below).

Al, A2, Mxl, Tl-3 as typical of the genus and not illustrated.

Fig. 13. Limbs of Candobrasilopsis acutis sp. nov. (c?)- A. Right prehensile palp (MZUSP 32703).

B. Left prehensile palp (MZUSP 32703). C. Hemipenis (MZUSP 32703). D. Caudal ramus (MZUSP

32703). E. Attachment of the caudal ramus (MZUSP 32703). F. Md palp (MZUSP 32703). Scale bars:

A-F = 50 pm.

HIGUTI J. & MARTENS K., Five new Candoninae species from Brazil

Md-palp (Fig. 13F) consisting of 4 segments, terminal segment short (F < 1.5 x basal width).

Prehensile palps (Fig. 13A-B) both segmented, distally hook-like and set with two unequal, sub-apical

setae; palps slightly asymmetrical, right prehensile palp (Fig. 13A) basally slightly more swollen and

with distal hook-like expansion apically swollen.

Caudal ramus (furca. Fig. 13D) with stout ramus and two stout apical claws. Proximal setae missing,

distal seta a small (hirsute) spine. Attachment to caudal ramus (Fig. 13E) long and stout, distally

bifurcated and with additional lateral branch at ca. mid-length. Fength ratio ramus / largest claw = 1.84.

Hemipenis (Fig. 13C) typical of the genus, with small and rounded ms and very pointed, triangular Is.

Description of female

Valves (Fig. 12A-B) very elongated, with greatest height situated behind the middle, anteriorly and

posteriorly broadly rounded; RV and FV of highly similar shape; anterior calcified inner lamella broadly

rounded, posterior calcified inner lamella very narrow and almost disappearing towards the dorsal side;

inner margin of anterior calcified inn er lamella slightly sinuous in both valves.

Carapace in right lateral (Fig. 12G), dorsal (Fig. 121) and ventral views (Fig. 12J) similar to those in the

male.

Soft parts as typical of the genus and, except for sexually dimorphic ones, similar to those of the male.

Measurements

See Table 3.

Discussion

Taxonomy of Pseudocandona and Typhlocypris

Vejdovsky (1882) erected Typhlocypris as a subgenus of Cypris to accommodate the distinct species

Cypris eremita Vejdovsky, 1880, while Kaufmann (1900) established the genus Pseudocandona for

the species Candona pubescens Koch, 1837. For historical reasons, the species Cypris eremita was

transferred to Pseudocandona and Typhlocypris became an unused senior synonym (Danielopol 1978;

Meisch 2000). Karanovic (2005) synonymised the whole of Pseudocandona with the older taxon

Typhlocypris , simply because the older name has priority over the more recent name following the

ICZN. Whereas this was technically correct, the name Pseudocandona was so well-used at that stage

that this move was not in the interest of nomenclatorial stability. For some time, this created an uneasy

and confusing situation.

Meisch (2000) foreshadowed a solution, by suggesting that the eremita- group of Pseudocandona has a

clearly distinct morphology, sufficient to consider it a genus separate from Pseudocandona.

Namiotko et al. (2014) meanwhile have indeed reinstated Typhlocypris as a separate genus for the

eremita- lineage, while Danielopol et al. (2012) have created the genus Marmocandona for the zschokkei-

species group in Pseudocandona. We follow this position and maintain all other species groups for

the time being within Pseudocandona , although, as foreshadowed by Danielopol (1978), at least the

Pseudocandona species of the Siberian Fake Baikal will eventually have to be transferred to a new

genus.

For a full discussion on the taxonomic and nomenclatorial history of Typhlocypris and Pseudocandona ,

see Namiotko et al. (2014).

European Journal of Taxonomy 106: 1-36 (2014)

Taxonomic position of Candona annae Mehes, 1914

Broodbakker (1983) described four species in the genus Pseudocandona from the West Indies and

also transferred the Columbian species Candona annae Mehes, 1914 to this genus, albeit implicitly.

Mehes (1914, fig. lOd) had indeed illustrated four long setae (small beta setae not drawn) on the second

segment of the Md-palp, which falls within the diagnosis of this genus. Furtos (1936) provided excellent

illustrations of a single male specimen from Florida that she referred to this species, but which had a

fairly different lobe “a” on the hemipenis and with the second segment of the Rpp being a lm ost evenly

rounded. Furtos (1935) described P. annae var septentrionalis from Massachusetts, mainly differing

from the Florida specimen by the presence of a distal seta on the caudal ramus and the absence of a fused

zone in the valves. Valve shape, and shape of hemipenis and of the prehensile palps are similar in both

forms described by Furtos.

According to Broodbakker (1983), the specimen reported by Furtos (1936) from Florida was in fact

not P. annae , but a different species, possibly closely related to either his P. geratisi , or more likely P.

antilliana. Later on in the same paper, he wrote that the Florida male of Furtos (1936) could very well

belong to P. antilliana. Broodbakker further mentioned that also P. elliptica (Furtos, 1933) from Ohio is

closely related to P. antilliana. Broodbakker did not comment on the position of P. annae septentrionalis

(Furtos, 1935).

Karanovic (2006) transferred P. annae to the re-instated genus Typhlocypris Vejdovsky, 1882. Karanovic

(2005) explicitly transferred P. geratsi Broodbakker (1983) to Typhlocypris , while Karanovic & Datry

(2009) also moved P. caribbeana Broodbakker, 1983 to Typhlocypris , but did not mention the position

of the other Pseudocandona species described by Broodbakker (1983) in their list of South and Central

American candonids. As outlined above, Typhlocypris now only comprises the old ‘ P. eremita ’ group, so

none of the South and Central American species belong to this genus.

Karanovic (2006) provided a re description of P. a. septentrionalis , based on one of the original dissections

of a male by Furtos, and synonymised it with P. annae s.s. At the same time, Karanovic (loc.cit.) also

sank P. antilliana Broodbakker, 1983 into the synonymy of P. annae. Strangely enough, the caudal

ramus, originally illustrated by Furtos (1935) with a clear distal seta, one of the major characters of this

form, lacks this seta in the illustration by Karanovic (2006).

Finally, Karanovic (2005) illustrated P. cf. geratsi (in Typhlocypris , sic) from interstitial waters near

Perth, Western Australia, but also illustrated some aspects of the morphology of the holotype of P. geratsi

Broodbakker, 1983, including a hemipenis in erection, not illustrated by Broodbakker (loc.cit.). Such

a wide distribution for a candonid species would be highly unusual, though of course not impossible.

In conclusion, we here maintain P. annae and P. antillana as separate species, with the male from Florida

most likely belonging to the latter species. The positions of P. annae septentrionalis (Furtos, 1935) and

of the Australian species P. cf. geratsi remain uncertain.

Taxonomic position of the presently described Pseudocandona species

With the eremita- group now back in Typhlocypris and the zschokkei- group allocated to Marmocandona,

Pseudocandona now comprises species from the compressa-, rostrata-, prespica- and the caribbeana-

groups (according to Namiotko & Danielopol 2004), together with the Lake Baikal species flock.

Namiotko & Danielopol (2004) allocated P. caribbeana and P. cubensis to a separate species group (the

caribbeana-group ), but P. antilliana and P. geratsi to the rostrata- group within the genus. Nevertheless,

both P. geratsi and P. antilliana have 4 ventral setae (plus beta-seta) on the second segment of the Md-

palp, and thus belong with P. caribbeana and P. cubensis to the same species group, which is not the

rostrata- group of which the species have 3 setae plus the beta seta there. Namiotko & Danielopol (2004,

HIGUTI J. & MARTENS K., Five new Candoninae species from Brazil

Table 4. Re-appraisal of the homology of claws and setae on two species in the P. caribbeana group,

based on Broodbakker (1983) and Martens (1987), NA = not applicable.

Female

Male

Species

Modern

nomenclature

Morphology

Nomenclature in

Broodbakker 1983

Morphology

P. agostinhoi sp. nov.

short claw

long claw

short claw

long claw

short claw

long claw

short seta

long claw

medium seta

short seta

P. antilliana

short claw

longest claw

long claw

short claw

long claw

short claw

long claw

short claw?

long claw

medium seta

short seta

medium seta

table 2) provided an alternative set of characters to distinguish between the different species groups in

Pseudocandona , namely by looking at the length of claws on the A2, relative to the combined lengths

of the second and third segments of the endopod. Also here, the new species fall well into the range of

the species of the caribbeana- group (including P. geratsi and P. antiliana). For example, the ratio G1 /

E2+3 in females is 1.6-1.9 for the caribbeana- group, and 1.68 for P. agostinhoi sp. nov.; the ratio GM/

E2+3 in females is 1.3-1.5 for the caribbeana- group and 1.38 for P. agostinhoi sp. nov.

One potential pitfall in interpreting the illustrations of Broodbakker (1983) is the use of different

nomenclature for the claws and setae on the A2. Martens (1987) demonstrated that, in the sexual

dimorphism in A2-chaetotaxy, form and shape of the claws and setae can change dramatically, but

relatively position does not. The work by Broodbakker (1983) predates this finding, and thus setae and

claws were named based on appearance, not on relative position. Table 4 corrects this situation and

homologizes Broodbakker’s nomenclature with the modern one. In doing so, the close relationship

between our and Broodbakker’s species becomes even more clear.

There are some small differences that need further investigation. For example, the reduced seta s2 on the

Md-palp is ca. twice as long in P. antilliana as in P. agostinhoi sp. nov., while the seta zl in the female is

illustrated as a short claw in P. antilliana (Broodbakker 1983, fig. 4D), which would be unusual in females.

In all, however, the three new species here allocated to Pseudocandona appear to belong in one lineage

with the four species described by Broodbakker (1983), which, following Namiotko & Danielopol

(2004), we here continue to name the caribbeana- group.

European Journal of Taxonomy 106: 1-36 (2014)

From a zoogeographical point of view, uniting these seven species into one lineage, and maybe including

also others like P. annae , seems logical.

Ecology

The five new species of candonids were found in 23 localities in the alluvial valley of the Upper Parana

River. The most common species of the five newly described here is Candobrasilopsis elongata sp. nov.,

which was recorded from 16 localities; Pseudocandona agostinhoi sp. nov. was found in nine localities,

P. cillisi sp. nov. in six, and C. acutis sp. nov. and P. claudinae sp. nov. were only recorded in one, and

the same, locality.

The latter two new species were found in sediments (named littoral in Table 1) of the Matagal stream,

with pH 6.2, a low value of dissolved oxygen (1.73 mg L 1 ) and a high value of electrical conductivity

(179.9 pS cm 1 ). At the time of collecting, the surface water available was less than lm 2 , and only a few

mm of standing water hidden by many grasses could be sampled.

Candobrasilopsis elongata sp. nov., was recorded in localities with pH values ranging between 4.7 and

7.2, electrical conductivity between 25.8 and 49.6 pS cm' 1 and dissolved oxygen between 0.14 and 8.3

mg L' 1 . The predominant type of substrate in which this species was found was Eichhornia crassipes.

Pseudocandona agostinhoi sp. nov. occurred in both types of substrate, namely littoral and Eichhornia

crassipes , with pH values ranging between 5.4 and 6.9, electrical conductivity between 28.1 and 64.1

pS cm 1 and dissolved oxygen between 0.9 and 7.2 mg LA Finally, P. cillisi sp. nov. was found in

environments with pH values ranging between 4.9 and 6.1, electrical conductivity between 27.0 and

41.8 pS cm -1 and low values of dissolved oxygen, between 1.0 and 4.5 mg LA Also this species was

predominant in Eichhornia crassipes.

Acknowledgements

Julian Cillis and Claudine Behen (RBINS) provided technical assistance with the SEM and the line

drawings, respectively. Jaime Luiz Lopes Pereira (Maringa, Brazil) offered help with the map. JH

received postdoctoral grants for short to more extended stays at the Royal Belgian Institute of Natural

Sciences (Brussels, Belgium) from CAPES (Process: BEX1452/08-8), from The Global Taxonomic

Initiative (GTI) and from the Belgian Science Policy (Belspo). This research fonns part of projects

supported by CNPq (Process: 472434/03-9, 478487/2010-0, 476130/2010-7 and 558118/2009-7 (Long-

Term Ecological Research - LTER)), SETI/Funda 9 ao Araucaria/MCT/CNPq (n. 232/10), SISBIOTA

(MCT/CNPq/Funda 9 ao Araucaria) and Nupelia. Dr David Horne (London) and Dr Dan Danielopol

(Graz) made important comments on an earlier draft of the manuscript.

References

Agostinho A. A. & Zalewski M. 1996. Aplanlcie alagdvel do alto rio Parana: importancia epreservagao.

{Upper Parana River Floodplain: Importance and Preservation). Maringa, EDUEM, 100.

Agostinho A.A., Gomes L.C., Thomaz S.M & Hahn N.S. 2004. The Upper Parana River and its

floodplain: Main characteristics and perspectives for management and conservation. In: Thomaz S.M.,

Agostinho A.A. & Hahn N.S. (eds) The Upper Parana River and its Floodplain: Physical Aspects,

Ecology and Conservation: 381-393. Backhuys Publishers, Leiden.

Broodbakker N.W. 1983. The subfamily Candoninae (Crustacea, Ostracoda) in the West Indies. Bijdragen

tot de Dierkunde. 53: 287-326.

Broodbakker N.W. & Danielopol D.L. 1982. The chaetotaxy of Cypridacea (Crustacea, Ostracoda) limbs:

proposals for a descriptive model. Bijdragen tot de Dierkunde 52: 103-120.

HIGUTI J. & MARTENS K., Five new Candoninae species from Brazil

Danielopol D.L. 1978. Uber Herkunft und Morphologie der Siisswasser-hypogaischen Candoninae

(Crustacea, Ostracoda). Sitzungsberichten der Osterreichischen Akademie der Wissenschaften.

Mathematisch-Naturwissenschaftliche Klasse, Abteilungl. 187: 1-162.

Danielopol D.L., Namiotko T. &. Meisch C. 2012. Marmocandona nov. gen. (Ostracoda, Candoninae),

with comments on the contribution of stygobitic organisms to micropalaeontological studies. Kolner

Forum fur Geologie und Palaontologie 21: 13-16.

Furtos N.C. 1935. Fresh-Water Ostracoda from Massachusetts. Journal of the Washington Academy of

Science 25 (12): 530-544.

Furtos N.C. 1936. Freshwater Ostracoda from Florida and North Carolina. The American Midland

Naturalist 17 (2): 491-522.

Higuti J. & Martens K. 2012a. On a new cypridopsine genus (Crustacea, Ostracoda, Cyprididae) from

the Upper Parana River Floodplain (Brazil). Zootaxa 3391: 23-38.

Higuti J. & Martens K. 2012b. Description of a new genus and species of Candonopsini (Crustacea,

Ostracoda, Candoninae) from the alluvial valley of the Upper Parana River (Brazil, South America).

European Journal of Taxonomy 33: 1—31. http://dx.doi.org/10.5852/ejt.2012.33

Higuti J., Velho F.F.M., Fansac-Toha F.A. & Martens K. 2007. Pleuston communities are buffered

from regional flood pulses: the example of ostracods in the Parana River floodplain, Brazil. Freshwater

Biology 52: 1930-1943. http://dx.doi.org/ 10.1111/i.l365-2427.2007.01821.x

Higuti J., Meisch C. & Martens K. 2009a. On Paranacypris samambaiensis gen. nov., sp. nov. (Crustacea,

Ostracoda), the first South American psychrodromid from the alluvial valley of the Upper Parana River,

Brazil. Journal of Natural History 43: 769-783. http://dx.doi.org/10.1080/002229308027025Q6

Higuti J., Fansac-Toha F.A., Velho F.F.M., Pinto R.F., Vieira F.C.G. & Martens K. 2009b. Composition

and distribution of Darwinulidae (Crustacea, Ostracoda) in the alluvial valley of the upper Parana River,

Brazil. Brazilian Journal of Biology 69: 253-262. http://dx.doi.org/10.1590/S1519-698420090002000Q4

Higuti J.,Velho F.F.M., Fansac Toha F.A. & Martens K. 2009c. Biodiversity of non-marine ostracods

(Crustacea, Ostracoda) in the alluvial valley of the upper Parana River, Brazil. Brazilian Journal of Biology

69 (suppl): 661-668. http://dx.doi.org/10.1590/S1519-6984200900030002Q

Higuti J., Declerck S.A.J., Fansac-Toha F.A., Velho F.F.M. & Martens K. 2010. Variation in ostracod

(Crustacea, Ostracoda) communities in the alluvial valley of the upper Parana River (Brazil) in relation

to substrate. Hydrobiologia 644: 261-278. http://dx.doi.org/10.1007/sl0750-010-Q122-l

Higuti J., Schon I., Audenaert F. & Martens K. 2013. On the Strandesia obtusata/elliptica lineage

(Ostracoda, Cyprididae) in the alluvial valley of the Upper Parana River (Brazil), with the description of

three new species. Crustaceana 86 (2): 182-211. http://dx.doi.org/10.1163/15685403-00003160

Horne D.J., Cohen A. & Martens K. 2002. Taxonomy, morphology and biology of Quaternary and living

Ostracoda. In\ Holmes J.A. & Chivas A.R. (eds) Biology, Taxonomy and Identification Techniques'.

5-36. American Geophysical Union, Washington DC.

Karanovic I. 2005. On the genus Typhlocypris Vejdovsky, 1882 (Crustacea, Ostracoda, Candoninae) with

the description of two new species. Systematics and Biodiversity 3: 375-406. http:// dx. doi.org/10.1017/

S1477200005001738

Karanovic I. 2006. Recent Candoninae (Crustacea, Ostracoda) of North America. Records of the Western

Australian Museum, Suppl. 71: 1-75.

Karanovic I. & Datry T. 2009. Overview of the Candoninae (Crustacea, Ostracoda) of South America and

the West Indies, with the description of two new species and one new genus. Zootaxa 2267: 1-25.

European Journal of Taxonomy 106: 1-36 (2014)

Kaufmann A. 1900. Cypriden und Darwinuliden der Schweiz. Revue Suisse de Zoologie 8: 209^123.

Martens K. 1987. Homology and functional morphology of the sexual dimorphism in the antenna of

Sclerocypris Sars, 1924 (Crustacea, Ostracoda, Megalocypridinae). Bijdragen tot de Dierkunde 57: 183—

190.

Martens K. & Behen F. 1994. A checklist of the non-marine ostracods (Crustacea, Ostracoda) from

South-American inland waters and adjacent islands. Travaux scientifiques du Musee d’Histoire naturelle

de Luxembourg 22: 1-81.

Martens K., Schon I, Meisch C. & Horne D.J. 2008. Global diversity of ostracods (Ostracoda, Crustacea)

in freshwater. In: Balian E. et al. (eds.) Freshwater Animal Diversity Assessment. Hydrobiologia 595:

185-193. http://dx.doi.org/10.1007/slQ750-007-9245-4

Mehes G. 1914. Siisswasser-Ostracoden aus Columbien und Argentinien. Memoires de la Societe naturelle

de Neuchatel 5: 639-663.

Meisch C. 2000. Freshwater Ostracoda of Western and Central Europe. Spektrum Akademischer Verlag

GmbH, Heidelberg, Berlin.

Mormul R.P., Thomaz S.M., Higuti J. & Martens K. 2010. Ostracod (Crustacea) colonization of a native

and a non-native macrophyte species of Hydrocharitaceae in the Upper Parana floodplain (Brazil): an

experimental evaluation. Hydrobiologia 644: 185-193. http://dx.doi.org/10.1007/sl0750-010-Q112-3

Namiotko T. & Danielopol D.L. 2004. Review of the eremita species-group of the Pseudocandona

Kaufmann (Ostracoda, Crustacea), with the description of a new species. Revista Espahola de

Micropaleontologia 36: 117-134.

Namiotko T., Danielopol D.L., Meisch C., Gross M. & Mori N. 2014. Redefinition of the genus

Typhlocypris Vejdovsky, 1882 (Ostracoda, Candonidae). Crustaceana 87: 952-984 + S1-S4. http://

dx.doi.org/10,1163/15685403-00003338

Por F.D. 1995. The Pantanal of Mato Grosso (Brazil), World’s Largest Wetlands. Kluwer Academic

Publishers, Dordrecht.

Souza Filho E.E. & Stevaux J.C. 2004. Geology and geomorphology of the Baia-Curutuba-Ivinheima

River complex. In: Thomaz S.M., Agostinho A.A. & Hahn N.S. (eds) The Upper Parana River and its

Floodplain: Physical Aspects, Ecology and Conservation: 1-29. Backhuys Publishers, Leiden.

Vejdovsky F. 1882. Thierische Organismen der Brunnenwasser von Prag\ 1-70 (Ostracoda: 64-65,

table 7).

Manuscript received: 25 May 2014

Manuscript accepted: 30 September 2014

Published on: 16 December 2014

Topic editor: Rudy Jocque

Desk editor: Kristiaan Hoedemakers

Printed versions of all papers are also deposited in the libraries of the institutes that are members of

the EJT consortium: Museum National d’Histoire Naturelle, Paris, France; Botanic Garden Meise,

Belgium; Royal Museum for Central Africa, Tervuren, Belgium; Natural History Museum, London,

United Kingdom; Royal Belgian Institute of Natural Sciences, Brussels, Belgium; Natural History

Museum of Denmark, Copenhagen, De nm ark