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http://dx.doi.org/10.5852/ejt.2012.33
WWW. europeanj oumaloftaxonomy. eu
2012 • Higuti J. & Martens K.
This work is licensed under a Creative Commons Attribution 3.0 License.
Research article
um:lsid:zoobank.org:pub:85998904-3543-461B-A741-CC6959AA48DE
Description of a new genus and species of Candonopsini
(Crustacea, Ostracoda, Candoninae) from the alluvial valley
of the Upper Parana River (Brazil, South America)
Janet HIGUTI' & Koen MARTENS^ *
^ Universidade Estadual de Maringa, Nupelia, PEA, Av. Colombo 5790, CEP 87020-900,
Maringa, PR, Brazil. E-mail:
[email protected]
Royal Belgian Institute of Natural Seienees, Freshwater Biology, Vautierstraat 29, B-1000 Brussels,
Belgium and University of Ghent, Dept of Biology, K.E. Eedeganekstraat 35, B-9000 Ghent, Belgium.
* Corresponding author:
[email protected]
' um:lsid:zoobank.org:author:3A5CEE33-280B-4312-BF6B-50287397A6F8
" um:lsid:zoobank.org:author:9272757B-A9E5-4C94-B28D-F5EFF32AADC7
Abstract. The genus Candobrasilopsis gen. nov. is here deseribed, with C. rochai gen. nov. sp. nov.
as type speeies, from the alluvial valley of the Upper Parana River. The enigmatie Candonopsis
brasiliensis Sars, 1901 is here redeseribed and transferred to this new genus, the new eombination being
CandobrasUopsis brasiliensis (Sars, 1901). The new eandonid genus belongs to the tribe Candonopsini,
beeause of the absenee of the proximal seta on the eaudal ramus. It is elosely related to Latinopsis
Karanovie & Datry, 2009, beeause of the relatively short terminal segment of the mandibular palp
(length less than 1.5 times the basal width, while this segment is longer than three times the basal width
in Candonopsis) and the large and stout b-seta on the Tl. However, it differs markedly from Latinopsis
in the size and shape of the ealeified inner lamellae of both valves and in the type of hemipenis. We
also diseuss the doubtful alloeation of several other genera to the Candonopsini, raise Abcandonopsis
Karanovie, 2004 to generie status and reassess the uneertain position of Candonopsis anisitsi Daday,
1905 within Latinopsis.
Key words. Candobrasilopsis rochai gen. nov. sp. nov., eaudal ramus, parallel evolution, taxonomy.
Higuti J. & Martens K. 2012. Deseription of a new genus and speeies of Candonopsini (Crustaeea,
Ostraeoda, Candoninae) from the alluvial valley of the Upper Parana River (Brazil, South Ameriea).
European Journal of Taxonomy 33: 1-31. http://dx.doi.org/10.5852/ejt.2012.33
Introduction
There are approximately 2000 deseribed speeies and ca. 200 deseribed genera of Reeent, free-living,
non-marine ostraeods in the world (Martens et al. 2008; Martens & Savatenalinton 2011). About 1000
speeies and 100 genera of these belong to the family Cyprididae Baird, 1845, and ca. 500 speeies
to the family Candonidae Kaufmann, 1900. The latter family was thought to be the most speeiose in
the northern hemisphere and to be of relatively reeent origin (Danielopol 1978), but several reeent
papers doeument the presenee of (sometimes speeiose) lineages in the (sub-)tropies and in the southern
hemisphere (Broodbakker 1983; Martens 1992, and several papers on Australian eandonids summarised
1
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European Journal of Taxonomy 1-31 (2012)
in Karanovic 2007). Especially the diversity and diversification of the subterranean ostracods in the
Pilbara area (NW Australia) is truly astonishing.
Close to 300 species have been described and reported from South America and more than 100 species
are known from Brazil (Martens & Behen 1994; Higuti & Martens 2012). Recent work on the alluvial
valley of the Upper Parana River (Higuti et al. 2007, 2009a, b, 2010) has contributed significantly to
the general knowledge of ecology and biogeography of Brazilian freshwater ostracods, especially those
occurring in the pleuston of fioating plants. Those same collections also served for taxonomic revisions
and several new species and genera have meanwhile been described from lakes, river beds and channels
of the Upper Parana fioodplain (Higuti et al. 2009c, in press; Higuti & Martens 2012). Most of these new
taxa belong to the family Cyprididae; the Candonidae were thus far not treated in any detail.
Here, we describe a new genus and a new species, Candobrasilopsis rochai gen. nov. sp. nov. and re¬
describe an old enigmatic species, Candonopsis brasiliensis Sars, 1901, which was labelled incertae
sedis by Karanovic & Marmonier (2002), confirmed by Karanovic & Datry (2009). This species is here
also transferred to Candobrasilopsis gen. nov. We discuss the taxonomy of the tribe Candonopsini as
well as the position of several species and genera.
Material and Methods
Study area
The alluvial valley formed by the Upper Parana River incorporates various fiuvial systems; each of
these has a variety of enviro nm ent types, ranging from the river itself, over connecting channels l inkin g
with open lakes and, finally, closed lakes. The Upper Parana River consists of a large braided channel,
with an extensive fioodplain (maximum width of 20 km) and high sediment accumulation in its bed,
creating sand bars and islands of diverse sizes (from some hundreds of meters to several kilometers in
length) (Agostinho & Zalewski 1996; Agostinho et al. 2004). The alluvial valley of the Upper Parana
River, 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) (Figure 1).
Material
The material for the present paper was collected in March, July and November of 2004 and in February
of 2011, 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 (Eichhornia crassipes, Pistia stratiotes, Hydrocotyle rannunculoides
and Salvinia spp.) 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-YSl, YSl 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.
All specimens with OC numbers are stored in the Ostracod Collection of the Royal Belgian Institute of
Natural Sciences, Brussels (Belgium). The material with MZUSP numbers is stored in the Museu de
Zoologia da Universidade de Sao Paulo, Sao Paulo (Brazil).
2
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HIGUTI J. & MARTENS K., New genus and speeies of Candonopsini from Brazil
Morphological analyses
Ostracods were dissected with valves stored dry in micropalaeontological slides and soft part in glycerine
in sealed slides. Drawings of soft parts were made with camera lucida with a compound microscope
(WILD HEERBRUGG). Valves were illustrated and measured using scanning electron microscopy
(Philips XE30 SEM at RBINS, Brussels).
Aragatuba
0 2 4 6 8 10km
53 ° 40 ’
53 ° 30 ’
53 ° 20 ’
53 ° 10 ’
TaquaruQU
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^ Mato Grosso
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Parana
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Fig. 1. Map of the area, indicating localities of Candobrasilopsis rochai gen. nov. sp. nov. and C. bra-
siliensis (Sars, 1901) comb. nov. were collected. Detailed occurrences are indicated in Table 1. Eocality
13 (samples PARI93 and PARI95) is the type locality of C. rochai gen. nov. sp. nov.
3
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Table 1. Localities where Candobrasilopsis rochai gen. nov. sp. nov. and Candobrasilopsis brasiliensis (Sars, 1901) eomb. nov. were eolleeted. IVI =
Ivinhema River System, BAI = Baia River System, PAR = Parana River System, TAQ = Taquaru 9 u System, floating = mixed floating plants, WT= water
temperature (®C), EC = eleetrieal eonduetivity (pS em'^), DO = dissolved oxygen (mg L'^). Number 13 refers to the type loeality of C. rochai gen. nov.
sp. nov. (Table eontinued on page 5).
European Journal of Taxonomy 1-31 (2012)
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5
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European Journal of Taxonomy 1-31 (2012)
Abbreviations used in text and figures
Al
Antennula
A2 =
Antenna
Cp =
Carapaee
H
Height of valves
L
Length of valves
Lpp =
Left prehensile palp
LV =
Left valve
Is =
Lateral shield of hemipenis
Md =
Mandibula
ms =
Medial shield of hemipenis
Mxl =
Maxillula
Rpp =
Right prehensile palp
RV =
Right valve
T1
First thoraeopod
T2
Seeond thoraeopod
T3
Third thoraeopod
W
Width
Chaetotaxy of the limbs follows the model proposed by Broodbakker & Danielopol (1982), revised for
the A2 by Martens (1987) and for the T3 by Meiseh (2000). Higher taxonomy of the Ostraeoda follows
the synopsis by Home et al. (2002).
Results
Taxonomic descriptions
Class Ostraeoda Latreille, 1806
Subelass Podoeopa G.W. Muller, 1894
Order Podoeopida Sars, 1866
Suborder Cypridoeopina Baird, 1845
Superfamily Cypridoidea Baird, 1845
Family Candonidae Kaufinann, 1900
Subfamily Candoninae Kaufinann, 1900
Tribes
Candonini Kaufinann, 1900; Candonopsini Karanovie, 2004; Cryptoeandonini Karanovie, 2007;
Danieloeandonini Karanovie, 2007; Humphreyseandonini Karanovie, 2005, Namibeypridini Martens,
1992; Terrestrieypridini Pinto et al, 2005; Trapezieandonini Karanovie, 2007.
Tribe Candonopsini Karanovie, 2004
Diagnosis (new, only partly following the original diagnosis)
Candonid speeies, with earapaee elongated in lateral view and narrow in dorsal and ventral views.
Caudal ramus well-developed, distally with two elaws and a distal seta, but with proximal seta missing.
Genera
Candonopsis Vavra, 1891, Latinopsis Karanovie & Datry, 2009, Candobrasilopsis gen. nov.
Not aeeepted here is the alloeation of Abcandonopsis Karanovie, 2004 (here raised to generie rank),
Cubacandona Broodbakker, 1983, Caribecandona Broodbakker, 19H3, MarococandonaMarmonier et
al, 2005 andPioneercandonopsis Karanovie, 2005 (see diseussion below).
6
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HIGUTI J. & MARTENS K., New genus and speeies of Candonopsini from Brazil
Genus Candobrasilopsis gen. nov.
um:lsid:zoobank.org:act:6DA166F4-39Dl-4AB3-8EC6-8C5C53ADB198
Diagnosis
Anterior calcified inner lamella broad, anterior inner 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 infiated, distal finger long, hook-like.
Hemipenis of triangular type, without protmding lobes a and b. Caudal ramus without proximal seta,
distal seta reduced to a small spine.
Etymology
Named after the country of Brazil (Brasil in Portuguese), with ''Cando-' and ''-opsis’" denoting
relationship to Candonopsis.
Type species
Candobrasdopsis rochai sp. nov. (by original designation).
Other species
Candonopsis brasiliensis Sars, 1901; 1 Candonopsis anisitsi Daday, 1905.
Differential diagnosis
This new genus is closely related to Candonopsis Vavra, 1891 (with type species C. kingsleyi (Brady
& Norman, 1870)) and to Latinopsis Karanovic & Datry, 2009 (type species L. patagonica Karanovic
& Datry, 2009). It differs from Candonopsis s.s. mainly in the presence of a short terminal Md-palp
segment, by the fact that the prehensile palps have 2 distal setae (1 in Candonopsis) and by the relatively
simple attachment of the caudal ramus (1 lateral branch in Latinopsis and in Candobrasilopsis gen. nov.,
3 in Candonopsis s.s.).
The new genus differs from Latinopsis (only L. patagonica) mainly in the presence of a broad anterior
calcified inner lamella with sinuous inner margin (narrow and with evenly rounded inner margin in
Latinopsis) and by the absence of protruding lobes ‘a’ and ‘b’ in the hemipenis (present in Latinopsis).
Remarks
We propose to tentatively transfer C. anisitsi Daday, 1905 to Candobrasilopsis gen. nov. based on the
morphology of the hemipenis and the prehensile palps (see discussion below).
Candobrasilopsis rochai sp. nov.
Figs 2-6, 7A-B
um:lsid:zoobank.org:act:152E5E0F-0FF9-4A83-B8EA-6424D6A07858
Candonopsis annae (Mehes, 1914) - Higuti et al. 2009b: 664. — Higuti et al. 2010: 267.
Diagnosis
A species of the tribe Candonini, with elongated valves, greatest height well behind the middle; carapace
in lateral view with weakly rounded dorsal margin, not straight. Anterior calcified inner lamella in both
valves broad; almost 1/4 of total length and with inner margin sinuous, not parallel to valve margin. Base
of right prehensile palp broad, almost triangular, distal part curved, distal tip slightly expanded, set with
two subequal lateral setae. Eeft prehensile palp larger and more elongated, with hook-like distal part.
7
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European Journal of Taxonomy 1-31 (2012)
distal tip slightly swollen. Hemipenis with triangular Is, distinctly pointed, proximo-dorsal expansion
small, ms rounded and without ventral, wart-like expansion.
Etymology
The new species is named after Prof Dr Carlos Eduardo Falavigna da Rocha (USP, Sao Paulo, Brazil), in
recognition of his vast contributions to the taxonomy of Brazilian Copepoda, but also in acknowledgement
of his initiative to start research on terrestrial Ostracoda in South America.
Type material
All type material was collected on 10 Nov. 2004 by the authors, by washing roots of Eichhornia cras-
sipes (PAR 195) ?indPistia stratiotes (PAR 193) over a hand net with mesh size 160 pm. See Table 1 for
measurements of water chemistry at time of collecting.
Holotype
1 (f, with soft parts dissected in glycerine in a sealed slide, and with valves stored dry (MZUSP.28102).
Allotype
1 $, dissected and stored as the male (MZUSP.28103).
Paratypes
2 (f(f dissected and stored as the holotype (MZUSP.28104, MZUSP.28105); 4 3 carapaces stored dry
in micropalaeontological slides (OC.3291, MZUSP.28106, MZUSP.28107, MZUSP.28108); 4 $$
dissected and stored as the holotype (OC.3292, MZUSP.28109, MZUSP.28111, MZUSP.28110); 3 ?
carapaces stored dry in micropalaeontological slides (OC.3293, MZUSP.28112, OC.3294).
Type locality
Baia River in Upper Parana River Floodplain, in floating plants (E. crassipes, P. stratiotes), coordinates:
22°40’37.5” S, 53°12’29” W (locality number 13 with samples PAR193 and PAR195 on Figure 1).
Other material investigated
See Table 1 for a list of localities where the present species was collected. Specimens MZUSP.28113,
MZUSP.28114, MZUSP.28115 {SS) and MZUSP.28116 ($) are here used for illustrations and are
also deposited in the Museu de Zoologia da Universidade de Sao Paulo, Sao Paulo (Brazil). Illustrated
specimens are listed in Table 2.
Differential diagnosis
The new species differs from the congeneric species (C. brasiliensis), and indeed from all other
Candonopsini, by the shape of the valves, by the peculiar shape of the prehensile palp (with distal part
swollen in Rpp) and by the size and shape of the hemipenis.
Description of male
Valves (Fig. 2A, B) elongated, with greatest height situated behind the middle (and bluntly pointed
there) anteriorly rather narrowly 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; inner margin of anterior calcified inner lamella slightly
sinuous in both valves.
Carapace in right lateral view (Fig. 2C) showing FV overlapping RV slightly on all sides; external
surface of valves smooth.
8
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HIGUTI J. & MARTENS K., New genus and speeies of Candonopsini from Brazil
Fig. 2. Candobrasilopsis rochai gen. nov. sp. nov., valves (A-F S and G-K 5 )- A, G. LV, internal view
(A: MZUSP.28113, G: MZUSP.28103). B, H. RV, internal view (B: MZUSP28113, H: MZUSP28103).
C, I. Cp, right lateral view (C: OC.3291, I: OC.3293). D, J. Cp, dorsal view (D: MZUSP28106,
J: MZUSP28112). E, F, K. Cp, ventral view (E: MZUSP28108, F: MZUSP28107, K: OC.3294). Scale
bars: A-D = 200 pm; E, F = 250 pm; G, H = 400 pm, I-K = 300 pm.
9
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European Journal of Taxonomy 1-31 (2012)
Fig. 3. Candobrasilopsis rochai gen. nov. sp. nov., limbs ( 6 '). A. A1 (MZUSP.28102). B. A2 (MZUSP.
28114). C. Md palp (MZUSP.28114). D. Md, coxal plate (MZUSP.28114). E. Mxl (Chaetotaxy
incomplete) (MZUSP.28105). F. Mxl, respiratory plate (MZUSP.28114). G. Attachment of the caudal
ramus (MZUSP.28104). Scale bars: A-G = 50 pm.
10
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HIGUTI J. & MARTENS K., New genus and speeies of Candonopsini from Brazil
Fig. 4. Candobrasilopsis rochai gen. nov. sp. nov., limbs (cJ). A. Tl, left clasping organ (MZUSP.28114).
B. Right clasping organ (MZUSP.28114). C. T2 (MZUSP.28114). D. T3 (MZUSP.28104). E. Caudal ramus
(MZUSP.28104). F. Detail of caudal ramus (MZUSP.28115). Scale bars: A-F = 50 pm.
11
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European Journal of Taxonomy 1-31 (2012)
Fig. 5. Candobrasilopsis rochai gen. nov. sp. nov., limbs ($). A. A2 (MZUSP.28110). B. Mdpalp (OC.3292).
C. A1 (OC.3292). D. Md, coxal plate (MZUSP.28109). E. Mxl, respiratory plate (MZUSP.28116). Scale
bars: A-E = 50 pm.
12
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HIGUTI J. & MARTENS K., New genus and speeies of Candonopsini from Brazil
Fig. 6. Candobrasilopsis rochai gen. nov. sp. nov., limbs ($). A. T3 (OC.3292). B. Attachment of the
caudal ramus (MZUSP.28110). C. T2 (MZUSP.28109). D. Caudal ramus (OC.3292). E. Mxl (Chaetotaxy
incomplete) (OC.3292). F. T1 (MZUSP.28111). Scale bars: A-F = 50 pm.
13
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European Journal of Taxonomy 1-31 (2012)
In dorsal view (Fig. 2D), carapace lancet-shaped, sharply pointed in the anteriorly, more broadly so
posteriorly, greatest width situated well behind the middle.
Carapace also in ventral view (Fig. 2E, F) lancet-shaped, with anterior side more acutely pointed than
posterior side.
A1 (Fig. 3A) with 5 terminal segments relatively elongated (L = at least 1.5 x basal width). Basal part
of A1 (= undivided protopodite) with 2 long sub-apical, ventral setae and two shorter setae on the dorsal
side, one at ca. mid-length, the other sub-apical. Next (second) segment with one long (reaching beyond
basis of penultimate segment) dorso-apical seta, no ventro-apical seta present. Third segment with one
small sub-apical seta on both ventral and dorsal sides. Fourth and fifth segments with two longer dorso-
apical and 1 shorter ventro-apical setae. Sixth segment with three longer dorso-apical and one shorter
ventro-apical setae. Seventh (terminal segment) most elongate of all, slightly sinuous, bearing one short
and two longer setae and one aesthetasc Ya, the latter shorter than the shorter seta.
A2 (Fig. 3B) with basal segment very broad, wide and relatively long, basally with 2 unequal setae, one
relatively long and slender, the other very short and broad, both hirsute; apically with a ventral seta.
Remnant of exopod consisting of a short plate, one long and two very short setae. Endopod consisting of
4 segments (penultimate segment divided). First endopodal segment long, carrying one long aesthetasc
Y on the ventro-basal side, and one long and one short ventro-apical setae. Second endopodal segment
shorter and smaller, but still rectangular with one ventral aesthetasc yl inserted about mid length, 3
t-setae, with tl a long, hirsute setae, t2 and t3 modified in the male-bristles. Dorso-apically with 3 short
setae of unequal length. 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 setae, aesthetasc yl short and ventro-apically inserted. Terminal (4^*^) segment small, distally with
claws GM (long) and Gm (short), aesthetasc y3 with its companion seta, fused at the basis and of c equal
length, and seta g, slightly longer than y3.
Md with coxa (Fig. 3D) relatively elongated, distally set with a series of strong teeth. Md-palp (Fig. 3C)
consisting of 4 segments. Basal segment dorsally carrying the respiratory plate (not shown), ventro-
apically with 2 long hirsute setae (s 1 & s2) and the alpha seta, the latter proximally with a broad base and
a fiagellum-like, distal part. Second segment dorso-apically with 2 seta of unequal length, ventro-basally
with a short, stout and hirsute seta and ventro-apically with a group of 5 setae: 3 long and hirsute setae
(similar to the two s-setae of the previous segment), a shorter, less hirsute seta and the short and hirsute
beta-seta. Third segment with a group of 3 dorso-subapical setae, the latter smooth, a central group of
two setae, one of which being the gamma seta and ventro-apical group of 4 unequal setae. Terminal
segment rectangular, apically set with two large claws and an uncertain number of smaller setae.
Mxl (Fig. 3E) 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 20 respiratory
rays, some quite short, others long (Fig. 3F). Palp with first segment carrying 3+1 apical setae; terminal
segment short and broad, carrying 2 longer claws and c 3 short setae. Chaetotaxy of three endites
impossible to determine. Sideways directed bristles near first endite long and stout.
Tl (sometimes called Mx2 - Fig. 4A) consisting of basal part (basipodite), carrying respiratory plates
(not shown), a palp (modified to prehensile palp in males) and an exopodite (?) distally set with 14-15
setae of different morphology and length. Basal plate set with one long and stout ‘b’-seta, a shorter and
more slender ‘d’-seta and two ‘a’-setae of unequal length. Prehensile palps (Fig. 4A, B) one-segmented,
distally hook-like and set with two unequal, sub-apical setae; palps slightly asymmetrical, right prehensile
palp (Fig. 4B) basally slightly more swollen and with distal hook like expansion apically swollen.
14
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HIGUTI J. & MARTENS K., New genus and speeies of Candonopsini from Brazil
T2 (walking limb - Fig. 4C) with 4-segmented endopodite (penultimate segment divided) and elongated.
First segment with long seta dl. Knee-segment devoid of seta d2. First segment of endopod especially
elongated, with short ventro-apical seta. Second segment also with one short ventro-apical seta. Third
segment with two ventro-apical seta, one short, one slightly longer. Terminal segment with one short
apical and one short sub-apical seta and a long apical claw.
Fig. 7. — A, B. Candobrasilopsis rochai gen. nov. sp. nov. A. Hemipenis (MZUSP.28104). B. Zenker
organ (MZUSP.28115). — C, D. C. brasiliensis (Sars, 1901) comb. nov. C. Hemipenis (OC.3296). D.
Zenker organ (OC.3296). Scale bars: A, C, D = 50 pm, B = 100 pm.
15
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European Journal of Taxonomy 1-31 (2012)
T3 (cleaning limb - Fig. 4D) as typical of the family. First segment with three long setae, one medial
(dl), one subapical (d2) and one apical (dp). Second segment with one subapical seta (e). Third segment
with one long subapical seta (f). Penultimate segment with one short and curved subapical seta (g).
Terminal segment well-separated from penultimate segment and carrying three setae: one long and
reflexed (h3) and two side-ways directed (hi & h2), the latter subequal.
Caudal ramus (fiirca - Fig. 4E) with stout ramus and two stout apical claws. Proximal setae missing,
distal seta a small spine. Attachment to caudal ramus (Fig. 3G) long and stout, distally bifurcated and
with additional lateral branch at c mid-length. Length ratio ramus/largest claw = 1.51.
Hemipenis (Fig. 7A) relatively small, with rounded ms and bluntly pointed, triangular Is, the latter
furthermore with small proximo-dorsal expansion; labyrinth short and stout, postlabyrinthal spermiductus
narrow and straight, without additional coils, but with a weak bent at the most distal part.
Zenker organ (Fig. 7B) short and broad, with at least 5 spinous whirls.
Description of female
Valves (Fig. 2G, H) and Cp (Fig. 21) in lateral view similar to those in the male, RV in inner view dorsally
more rounded and less bluntly pointed than in the male. Cp in D and V views (Fig. 2J, K) equally narrow
and lancet-shaped, in D view with slight rostrum, in ventral view without the pronounced (anterior and
posterior) rostral keels, typical of the male.
Soft parts (Figs 5B-E, 6A-E) largely as in the male, but with sexually dimorphic A2 and Tl.
A2 (Fig. 5A) with setae tl-4 setae like, not transformed; zl and z2 short and slender, seta z3 short and
stout. Claws Gl, G2, G3 and GM all reaching to about the same point.
Tl (Fig. 6F) with basal part as in the male. Endopod a broad palp, with two short and one long distal
setae.
Caudal ramus: length ratio ramus/largest claw = 1.40.
Measurements
See Table 3.
Ecology
See below under C. brasiliensis.
Remarks
This species was originally identifled by us as Candonopsis annae (Mehes, 1914), but subsequent
detailed analysis of the valves and soft parts showed that this was not so, and that the populations from
the alluvial valley of the Upper Parana River needed be described as a new species.
Some specimens (Fig. 2F) give the impression that they have an anterior rostrum and a postero-ventral
keel. This was observed in 2 male specimens, after their carapaces were dried for SEM. So either these 2
specimens belong to a different species, which is not very likely, or the weak calciflcation of the valves
(typical of Candonopsis-Mks species) has made those parts of the valve implode while drying in air.
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HIGUTI J. & MARTENS K., New genus and speeies of Candonopsini from Brazil
Table 2. Locality data of Candobrasilopsis rochai gen. nov. sp. nov. and Candobrasilopsis brasiliensis
(Sars, 1901) comb, nov., material used here.
Candobrasilopsis rochai gen. nov. sp. nov.
Valves (6')
MZUSP.28113 - Piranha Lake (PAR 115), littoral, coll. 05 Jul. 2004.
OC.3291, MZUSP.28106, MZUSP.28107, MZUSP.28108 - Baia River (PAR 195), Eichhornia
crassipes, coll. 10 Nov. 2004.
Valves ($)
MZUSP.28103, OC.3293, MZUSP.28112, OC.3294 - Baia River (PAR 195), Eichhornia crassipes,
coll. 10 Nov. 2004.
Soft parts (3)
MZUSP.28102, MZUSP.28104, MZUSP.28105 - Baia River (PAR 195), Eichhornia crassipes, coll.
10 Nov. 2004.
MZUSP.28114 - Cortado Channel (PAR 234), Pistia stratiotes, coll. 11 Nov. 2004.
MZUSP.28115 - Ivinheima River (PAR 222), Eichhornia crassipes, coll. 10 Nov. 2004.
Soft parts ($)
OC.3292, MZUSP.28109, MZUSP.28110 - Baia River (PAR 195), Eichhornia crassipes, coll. 10
Nov. 2004.
MZUSP.28116 - Cortado Channel (PAR 234), Pistia stratiotes, coll. 11 Nov. 2004.
MZUSP.28111 - Baia River (PAR 193), Pistia stratiotes, coll. 10 Nov. 2004.
Candobrasilopsis brasiliensis (Sars, 1901) comb. nov.
Valves (cJ)
MZUSP.28118 - Samambaia 1 Lake (PAR 102), floating, coll. 04 Jul. 2004.
MZUSP.28119, OC.3297, OC.3298 - Manezinho Backwater (S2), Eichhornia crassipes, coll. 11
Nov. 2004.
Valves ($)
MZUSP.28126, OC.3301 - Samambaia 1 Lake (PAR 102), floating, coll. 04 Jul. 2004.
MZUSP.28127, MZUSP.28125 - Manezinho Backwater (S2), Eichhornia crassipes, coll. 11 Nov. 2004.
Soft parts (3)
MZUSP.28117 - Manezinho Backwater (S2), Eichhornia crassipes, coll. 11 Nov. 2004.
OC.3295, OC.3296 - Pontal Lake (PAR 41), littoral, coll. 15 Mar. 2004.
Soft parts ($)
MZUSP.28125, OC.3299, OC.3300 - Manezinho Backwater (S2), Eichhornia crassipes, coll. 11 Nov.
2004.
Hemipenis
MZUSP.28117 - Manezinho Backwater (S2), Eichhornia crassipes, coll. 11 Nov. 2004.
OC.3295, OC.3296, MZUSP.28120 - Pontal Lake (PAR 41), littoral, coll. 15 Mar. 2004.
MZUSP.28121, MZUSP.28122, MZUSP.28123, MZUSP.28124 - Mane Cotia Lake (PAR 422),
Eichhornia crassipes, coll. 1 Feb. 2011.
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European Journal of Taxonomy 1-31 ( 2012 )
Table 3. Measurements of illustrated speeimens of Candobrasilopsis rochai gen. nov. sp. nov. and
Candobrasilopsis brasUiensis (Sars, 1901) eomb. nov.
Species
Sex
Code
V/Cp
L (mm)
H (mm)
W (mm)
Candobrasilopsis rochai gen. nov. sp. nov.
s
MZUSP.28113
LVi
748
353
RVi
739
356
OC.3291
CpRl
744
350
MZUSP.28106
CpD
766
218
MZUSP.28107
CpV
806
224
MZUSP.28108
CpV
773
217
?
MZUSP.28103
LVi
783
361
RVi
770
357
OC.3293
CpRl
753
342
MZUSP.28112
CpD
739
211
OC.3294
CpV
756
221
Candobrasilopsis brasUiensis (Sars,
s
MZUSP.28118
LVi
977
493
1901) eomb. nov.
RVi
976
475
MZUSP.28119
CpRl
931
463
OC.3297
CpD
902
300
OC.3298
CpV
917
301
?
MZUSP.28125
LVi
928
451
RVi
924
446
MZUSP.28126
CpRl
962
480
OC.3301
CpD
978
320
MZUSP.28127
CpV
912
288
Candobrasilopsis brasUiensis (Sars, 1901) eomb. nov.
Figs 7C-D, 8-13
Candonopsis brasUiensis Sars, 1901: 45-46.
Candonopsis brasUiensis 1984: 439-447. — Martens & Behen 1994:14. — Higuti et al. 2007:
1935. — Higuti et al. 2009b: 664. — Higuti et al. 2010: 161. — Mormul et al. 2010: 189.
Candonopsis brasUiensis incertae sedis - Karanovie & Datry 2009: 5.
Diagnosis
A speeies of the tribe Candonini, with elongated valves, greatest height well behind the middle; earapaee
in lateral view with dorsal margin straight over 2/3 of the length. Anterior ealeified inner lamella in both
valves broad; almost 1/5 of total length and with inner margin sinuous, not parallel to the valve margin.
Base of right prehensile palp narrow and rounded, distal part eurved, distal tip slightly expanded, set
with two subequal lateral setae. Left prehensile palp larger and more elongated, with hook-like distal
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HIGUTI J. & MARTENS K., New genus and speeies of Candonopsini from Brazil
part, distal tip not swollen. Hemipenis with triangular Is, bluntly pointed, bearing large and rounded
proximo-dorsal expansion, and with rounded ms bearing a ventral, wart-like expansion.
Type material and type locality
Sars (1901: 46) wrote: “Only 2 female specimens of this form were secured. They were found in one of
my aquaria prepared with mud from Sao Paulo”. No detailed locality information is provided. Given de
expansion of the city of Sao Paulo over the past 100 years, the actual locality from which the mud was
collected has meanwhile almost certainly been destroyed.
Remark
Sars (1901) based his description and illustration (only a dorsal and left lateral view of a carapace) on
2 female specimens only. As females in this genus do not have the most specific characters which are
in the hemipenis and prehensile palps of the male, we have decided not to rely on the type material to
redescribe this species. Rather, we use new material from the Upper Parana River, about 700 km WNW
from Sao Paulo City.
Material used for the present redescription
See Table 1 for an overview of localities in the alluvial valley of the Upper Parana River where this
species was found.
Specimens are here used for illustrations and are also deposited in the museums indicated above. Eight
SS, with soft parts dissected in glycerine in a sealed slide and with valves stored dry (MZUSP.28117,
OC.3295, OC.3296, MZUSP.28120, MZUSP.28121, MZUSP.28122, MZUSP.28123, MZUSP.28124);
four S carapaces stored dry in micropalaeontological slides (MZUSP.28119, OC.3297, OC.3298, and
MZUSP.28118). Three $ $, with soft parts dissected in glycerine in a sealed slide and with valves stored
dry (MZUSP.28125, OC.3299, OC.3300); three $ carapaces stored dry in micropalaeontological slides
(MZUSP.28127, MZUSP.28126, OC.3301).
Differential diagnosis
This species differs from the congeneric species (C. rochai gen. nov. sp. now), by the shape of the
valves, by rounded basal part of the right prehensile palp and by the large hemipenis with blunt tip and
the ms with a ventral wart-like expansion.
Redescription of male
Valves (Fig. 8A, B) elongated, with greatest height situated well behind the middle (and bluntly pointed
there), middle 2/3 of the dorsal margin straight and sloping towards the anterior margin; anterior margin
rather narrowly and posterior rather broadly 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.
Carapace in right lateral view (Fig. 8C) showing FV overlapping RV slightly on postero-dorsal and
ventral sides; external surface of valves smooth.
In dorsal (Fig. 8D) and ventral (Fig. 8E) views, carapace lancet-shaped, bluntly pointed anteriorly, more
broadly so posteriorly, greatest width situated well behind the middle.
A1 (Fig. 9C) with 5 terminal segments relatively elongated (F = at least 1.5 x basal width). Basal
segment of A1 (= undivided protopodite) with 2 long sub-apical, ventral setae and two shorter setae on
the dorsal side, of the latter one inserted at c mid-length, the other sub-apical. Second segment of basal
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European Journal of Taxonomy 1-31 ( 2012 )
Fig. 8. Candobrasilopsis brasiliensis (Sars, 1901) comb, nov., valves (A-E 3, F-J $). A, F. LV,
internal view (A: MZUSP.28118, F: MZUSP.28125). B, G. RV, internal view (B: MZUSP28118, G:
MZUSP28125). C, H. Cp, right lateral view (C: MZUSP28119, H: MZUSP28126). D, L Cp, dorsal
view (D: OC.3297,1: OC.3301). E, J. Cp, ventral view. (E: OC.3298, J: MZUSP.28127). Seale bars: A,
B, H, I = 200 pm; C-G, J = 400 pm.
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HIGUTI J. & MARTENS K., New genus and speeies of Candonopsini from Brazil
Fig. 9. Candobrasilopsis brasiliensis (Sars, 1901) comb, nov., limbs (cJ). A. A2 (MZUSP.28117). B. A2,
detail of the last segment (MZUSP.28117). C. A1 (MZUSP.28117). D. Mdpalp (MZUSP.28117). E. Mxl
(Chaetotaxy incomplete) (OC.3296). F. Md, coxal plate (MZUSP.28117). Scale bars: A-F = 50 pm.
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European Journal of Taxonomy 1-31 ( 2012 )
Fig. 10. Candobrasilopsis brasiliensis (Sars, 1901) comb, nov., limbs ((?). A. T1 (palp not shown)
(MZUSP.28117). B. Attachment of the caudal ramus (OC.3295). C. T3 (MZUSP.28117). D. Right clasping
organ (MZUSP28117). E. Left clasping organ (MZUSP28117). F. T2 (MZUSP28117). G. Mxl, respiratory
plate (OC.3296). H. Caudal ramus (OC.3295). Scale bars: A-H = 50 pm.
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HIGUTI J. & MARTENS K., New genus and speeies of Candonopsini from Brazil
Fig. 11. Candobrasilopsis brasiliensis (Sars, 1901) comb, nov., limbs ($). A. A2 (MZUSP.28125). B. A2,
detail of the last segment (MZUSP.28125). C. Mdpalp (MZUSP.28125> D. A1 (MZUSP.28125). E. Mxl
(Chaetotaxy incomplete) (OC.3299). F. Md, coxal plate (MZUSP.28125). Scale bars: A-F = 50 pm.
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European Journal of Taxonomy 1-31 ( 2012 )
Fig. 12. Candobrasilopsis brasiliensis (Sars, 1901) comb, nov., limbs ($). A. Caudal ramus (MZUSP.28125).
B. Attachment of the eaudal ramus (MZUSP.28125). C. T1 (OC.3300). D. T3 (OC.3300). E. T2
(MZUSP.28125). F. Mxl, respiratory plate (OC.3299). Seale bars: A-F = 50 pm.
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HIGUTI J. & MARTENS K., New genus and speeies of Candonopsini from Brazil
Fig. 13. Candobrasilopsis brasiliensis (Sars, 1901) comb, nov., hemipenis. A. MZUSP.28117.
B. OC.3295. C. OC.3296. D. MZUSP.28120. E. MZUSP28121. F. MZUSP28122. G. MZUSP28123.
H. MZUSP28124. Scale bars: A-F = 50 pm.
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European Journal of Taxonomy 1-31 (2012)
part with one long (reaching beyond basis of fifth segment) dorso-subapical seta, no ventro-apical seta
present. Third segment with one small sub-apical dorsal and ventral setae. Fourth and fifth segments
with two longer dorso-apical and 1 shorter ventro-apical setae. Sixth segment with three longer dorso-
apical and one shorter ventro-apical setae. Seventh (terminal segment) most elongate of all, slightly
sinuous, bearing one short and two longer setae and one aesthetasc Ya, the latter shorter than the shorter
seta.
A2 (Fig. 9A, B) with basal segment broad, wide and relatively long, basally with 2 unequal setae, one
relatively long and slender, the other very short and broad, both hirsute; apically with a ventral seta.
Remnant of exopod consisting of a short plate, one long and two very short setae. Endopod consisting of
4 segments (penultimate segment divided). First endopodal segment long, carrying one long aesthetasc
Y on the ventro-basal side, and one long and one short ventro-apical setae. Second endopodal segment
shorter and smaller, but still rectangular with one ventral aesthetasc yl inserted about mid length,
3 t-setae, with tl a long, hirsute setae, t2 and t3 modified in the male-bristles; this segment dorso-
apically with 3 short setae of unequal length. 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 setae, aesthetasc yl short and ventro-apically inserted. Terminal (4^*^) segment
(Fig. 9B) small, distally with claws GM (long) and Gm (short), aesthetasc y3 with its companion seta,
fused at the basis and with seta almost twice as long as aesthetasc, and seta g, clearly longer than y3.
Md with coxa (Fig. 9F) relatively elongated, distally set with a series of strong teeth. Md-palp (Fig. 9D)
consisting of 4 segments. Basal segment dorsally carrying the respiratory plate (not shown), ventro-
apically with 2 long hirsute setae (s 1 & s2) and the alpha seta, the latter proximally with a broad base
and a fiagellum-like, distal part. 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: 3 long and hirsute
setae (similar to the two s-setae of the previous segment), a shorter, less hirsute seta and the short and
hirsute beta-seta. Third segment with a group of 3 dorso-subapical setae, the latter smooth, a central
apical group of two setae, one of which being the gamma seta and ventro-apical group of 4 unequal
setae. Terminal segment rectangular, apically set with two large claws and an uncertain number of
smaller setae.
Mxl (Fig. 9E) with a basal (basipodite) part carrying a large respiratory plate (exopodite), 3 endites
and a two-segmented palp (endopodite). Respiratory plate (Fig. lOG) elongated, carrying more than
20 respiratory rays, some quite short, others long. Palp with first segment carrying 3+1 apical setae;
terminal segment short and broad, carrying 2 longer claws and c 3 short setae. Chaetotaxy of three
endites impossible to determine. Sideways directed bristles near first endite long and stout.
Tl (sometimes called Mx2 - Fig. lOA) consisting of basal part (basipodite), carrying respiratory plates
(not shown), a palp (modified to prehensile palp in males) and an exopodite (?) distally set with 14-15
setae of different morphology and length. Basal plate set with one long and stout ‘d’-seta, a shorter
and more slender ‘b’-seta and two ‘a’-setae of unequal length. Prehensile palps (Fig. lOD, E) one-
segmented, distally hook-like and set with two unequal, sub-apical setae; palps slightly asymmetrical,
Rpp (Fig. lOD) slightly smaller than Epp, distal part of apical hook slightly swollen in Epp, not at all in
Rpp.
T2 (walking limb - Fig. 1 OF) with 4-segmented endopodite (penultimate segment divided) and elongated.
First segment with long seta dl. Knee-segment devoid of seta d2. First segment of endopod especially
elongated, with short ventro-apical seta (e). Second segment also with one short ventro-apical seta (f).
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.
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HIGUTI J. & MARTENS K., New genus and speeies of Candonopsini from Brazil
T3 (cleaning limb - Fig. IOC) as typical of the family. First segment with three long setae, one medial
(dl), one subapical (d2) and one apical (dp). Second segment with one subapical seta (e). Third segment
with one long subapical seta (f), Penultimate segment with one short and curved subapical seta (g).
Terminal segment well-separated from penultimate segment and carrying three setae: one long and
reflexed (h3) and two side-ways directed (hi & h2), the latter subequal.
Caudal ramus (fiirca - Fig. lOH) with stout ramus and two stout apical claws. Proximal setae missing,
distal seta a small spine. Attachment of caudal ramus (Fig. lOB) long and stout, distally bifurcated and
with additional lateral branch before mid-length. Length ratio ramus/largest claw = 1.64.
Hemipenis (Fig. 7C) large, with medial shield rounded and with additional ventral protuberance, lateral
shield bluntly pointed, triangular and with rounded, proximo-dorsal expansion; labyrinth short and
stout, postlabyrinthal spermiductus narrow and straight, without additional coils, but with a weak bent
at the most distal part.
Zenker organ (Fig. 7D) short and broad, with ca. 5 spinous whirls.
Redescription of female
Valves (Fig. 8F, G) and Cp (Fig. 8H) in lateral view similar to those in the male; valves in inner view
slightly more elongated and less high than in the male. Cp in D and V views (Fig. 81, J) slightly more
slender than in the male.
Soft parts (Figs IIC-F, 12A, B, D-F) largely as in the male, but with sexually dimorphic A2 and Tl.
A2 (Fig. IIA, B) with setae tl-4 setae like, not transformed; zl-3 short and slender setae. Claws Gl, G2,
G3 and GM all reaching to about the same point.
Tl (Fig. 12C) with basal part as in the male. Endopod a broad palp, with two short and one long distal
setae.
Caudal ramus: length ratio ramus/largest claw = 1.61.
Measurements
See Table 3.
Ecology
Together, both species have been found in 29 localities, in a total of 48 localities sampled in the alluvial
valley of the Upper Parana River, they occurred sympatrically in only 11 localities.
Candobrasilopsis rochai gen. nov. sp. nov. occurred in 17 localities in the alluvial valley. The pH ranged
between 5.1 and 6.6, electrical conductivity between 13.2 and 66.9 pS cm'^ and dissolved oxygen
between 1.2 and 8.6 mg L f
Candobrasilopsis brasiliensis comb. nov. was found in 26 localities, with pH values ranging between
4.7 and 6.5, electrical conductivity between 23.6 and 114.9 pS cm'^ and dissolved oxygen between 0.6
and 13.3 mg L f
These species were predominant in several types of substrates as sediment (named littoral in Table 1)
and different species of floating macrophytes {Eichhornia crassipes, Pistia stratiotes, Hydrocotyle
ranunculoides and Salvinia spp.) in different habitats (lakes, channels and rivers) (Table 1).
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European Journal of Taxonomy 1-31 ( 2012 )
Remarks
As some small differences in hemipenis morphology between two different populations of this species
were detected, we dissected a longer series of males from 3 populations and illustrated both hemipenes
of each individual (Fig. 13). It is clear that indeed some variability exists, i.e. in the size of the proximo-
dorsal expansion of the Is, as well as in the ventral expansion of the ms. Some of this variability might
be a result of different positions of these organs between cover slip and glass slide. Therefore, we deem
all of these populations to be conspecific.
Discussion
Candobrasilopsis versus Latinopsis
Karanovic & Datry (2009) erected the genus Latinopsis, with the type species L. patagonica Karanovic
& Datry, 2009 from Chile, based mainly on the fact that the length of the terminal segment of the Md-
palp is much shorter in this South American genus than in Candonopsis s.s. We agree that this is a
significant character that merits the distinction of a separate lineage within the Candonopsini.
Here, we describe a new genus with the same character, the short terminal segment on the mandibular
palp, but which differs markedly from Latinopsis in a number of characters. Firstly, in the size and shape
of the anterior calcified inner lamella, which is narrow and of which the inner margin is evenly rounded,
running parallel to the valve margin in Latinopsis patagonica, but which is ca. 3 times broader in
Candobrasilopsis and which has a sinuous inner margin in both valves, not parallel to the anterior valve
margin. In fact, the anterior calcified inner lamella in Latinopsis species have a juvenile appearance,
and if the illustrations in Karanovic & Datry (2009: figs lA, 3A) are correct, one would suspect a
heterochronic valve development in this genus. Secondly, the hemipenis of Latinopsis patagonica is
not at all of the type that is typical of 'Candonopsis'’ species: it has two distinct lobes (named a and b in
Karanovic & Datry, loc.cit.) and as such is more of the 'Candona' ^./.-type (see extensive descriptions
in Danielopol 1978). The hemipenis in 'Candonopsis' s.l, as in Candobrasilopsis gen. nov. and in
C. anisitsi, has a large, triangular and pointed lateral shield (named lobe a in Karanovic & Datry, 2009)
and an almost non-existant medial shield (Fig. 7A, C). The Candonopsini thus have at least 2 separate
lineages within the Neotropics.
The position of Candonopsis anisitsi Daday, 1905
Karanovic & Datry (2009) also included several (older, and ill described) species into their new genus,
including Candonopsis anisitsi Daday, 1905 from Paraguay. Of this latter species, they re-described type
material and illustrated several of the limbs (not the valves, which were apparently missing in the type
material). It is clear from this re-description that there are several differences between L. patagonica
and C. anisitsi, for example (1) in the type of hemipenis, which has a 'Candona' - like appearance with
lobes a and b in Latinopsis patagonica, but has the real 'Candonopsis'’ - type with triangular lateral
shield (Is) and reduced medial shield (ms) in Candonopsis anisitsi (see above); (2) the morphology
of the prehensile palps (almost symmetrical in Latinopsis patagonica and with distinct asymmetry in
C. anisitsi and (3) in the size, shape and position of the seta ‘b’ on the endopod of the T1 (seta b large
and stout and placed distinctly away from seta d in Latinopsis and Candobrasilopsis gen. nov.; small
and positioned closely to seta d in C. anisitsi). There also appear to be differences in other aspects of the
chaetotaxy between L. patagonica and C. anisitsi, but these are more difficult to interpret. For example,
the rest of the chaetotaxy of the mandibular palp appears to be different between both species.
We agree that both species belong to related lineages within Candonopsini, and differ from Cando¬
nopsis S.S., but do not necessarily belong in the same genus.
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HIGUTI J. & MARTENS K., New genus and speeies of Candonopsini from Brazil
New material of this species needs to be re-examined to determine the exact shape and position of seta
b on T1 and of the shape and size of the anterior calcified inner lamella (and of the inner margin) in this
species. The other species transferred to Latinopsis by Karanovic & Datry (2009), namely Candona
columbiensis Mehes, 1914 and Candonopsis falclandica Vavra, 1898, need to be re-examined in detail
to see in which genus they belong.
Taxonomy of the Candonopsini
The taxonomic identity of the Candonopsini relied first and foremost on the absence of the proximal seta
of the caudal ramus. This feature is still the first delimiting factor to distinguish between ‘real’ candonids
on the one hand and candonopsines on the other hand. There are at present three genera with a well-
developed caudal ramus (i.e. a strong ramus, with 2 well-developed claws) in which the proximal seta
is missing: Candonopsis Vavra, 1891, Latinopsis Karanovic & Datry, 2009 and Candobmsilopsis gen.
nov. These genera most likely belong to the same phyletic clade and can thus be united in a tribe, the
Candonopsini. However, meanwhile several species with further reductions in the chaetotaxy of the caudal
ramus have been described within Candonopsis s.s. (e.g. C. westaustraliensis Karanovic & Marmonier,
2002), while Marococandona Marmonier et al, 2005 comprises one species with fully developed caudal
ramus (M. danielopoli) and one species in which the caudal ramus consists of only a ramus and one apical
claw (M nicolae). Moreover, several other genera with strongly reduced caudal rami have been lodged in
the Candonopsini by Karanovic & Datry (2009) and Karanovic (2012): Abcandonopsis Karanovic, 2004
(here raised to generic rank), Cubacandona Broodbakker, 1983, Caribecandona Broodbakker, 1983,
Marococandona Marmonier et al., 2005 and Pioneercandonopsis Karanovic, 2005.
We have issues with this taxonomic situation, as it is almost certain that not all of these species and genera
effectively belong to the same phyletic lineage. A reduction in size and chaetotaxy of the cypridoid
caudal ramus has occurred many times in different lineages and is in fact one of the most common
examples of parallel evolution (homeomorphy) in non-marine Ostracoda (see also McKenzie 1982).
The polyphyletic nature of the Candonopsini as defined by Karanovic (2012) is further demonstrated
by the fact that the different genera have several other characters in which they differ from each other:
“Carapace shape various (sic) (reniform, elongated, trapezoidal) (...), Al 7-segmented (...) or the
number of segments reduced. Male sexual bristles on A2 present (...) or absent.” (Karanovic 2012:
258). Other differences occur in hemipenis morphology.
A revision of the Candonopsini is urgently needed so that the different lineages which are presently
grouped within it, can be recognised and can be properly classified.
Acknowledgements
JH acknowledges Thijs Van der Meeren (Brussels, Belgium) for help with the drawings. Julien Cillis and
Claudine Behen (Brussels, Belgium) provided technical assistance with the SEM and the line drawings,
respectively. Kristiaan Hoedemakers has assisted greatly at various stages in the production of this paper.
Jaime Luiz Lopes Pereira (Maringa, Brazil) offered help with the map. JH received grants for short
to more extended visits to the Royal Belgian Institute of natural Sciences (Brussels, Belgium) from
CAPES (Process: BEX 1452/08-8), from The Global Taxonomic Initiative (GTI-CBD/ 2010 and GTI-
CBD/ 2012), from the Belgian Science Policy (Belspo) and from International Mobility Program (n®
003/2011) of the Universidade Estadual de Maringa. This research forms 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)), Funda 9 ao Araucaria/CNPq (n. 232/10) and Nupelia.
29
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European Journal of Taxonomy 1-31 ( 2012 )
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Manuscript received: 30 October 2012
Manuscript accepted: 12 December 2012
Published on: 17 December 2012
Topic editor: Rudy Jocque
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; National Botanic Garden
of Belgium, Meise, Belgium; Royal Museum for Central Africa, Tervuren, Belgium; Natural History
Museum, London, United Kingdom; Royal Belgian Institute of Natural Sciences, Bmssels, Belgium;
Natural History Museum of Denmark, Copenhagen, Denmark.
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