http://dx.doi.org/10.5852/eit.2015.lll

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2015 • Martens K., Halse S. & Schon L

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Research article

urn:lsid:zoobank.org:pub:D405800A-4225-4A72-A541-DB122452352B

On the Bennelongia ttimala and B. triangulata lineages (Crustacea,

Ostracoda) in Western Australia, with the description of six new species

Koen MARTENS 1 - 2 - 5 , Stuart HALSE 3 - 6 & Isa SCHON 1 - 4 - 7

1 Royal Belgian Institute of Natural Sciences, Operational Directorate “Natural Environment”,

Freshwater Biology, Vautierstraat 29, B-1000 Brussels, Belgium.

Corresponding author: darwinula@gmail. com

2 University of Ghent, Department of Biology, K.L. Ledeganckstraat 35, B-9000 Gent, Belgium.

3 Bennelongia Environmental Consultants, 5 Bishop Street, Jolimont WA 6014, Australia.

4 University of Hasselt, Research Group Zoology, Agoralaan Building D,

B-3590 Diepenbeek, Belgium.

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

6 urn:lsid:zoobank.org: author: 676014BF-5979-49EC-BC8F-811214170111

7 um:lsid:zoobank.org:author:94232F10-7092-4E90-9071-64C4FDABE691

Abstract. The ostracod genus Bennelongia De Deckker & McKenzie, 1981 occurs in Australia and New

Zealand. We redescribe B. nimala from the Northern Territory and describe six new species from Western

Australia belonging to the B. nimala (five species) and B. triangulata sp. nov. (one species) lineages:

B. tirigie sp. nov., B. koendersae sp. nov., B. pinderi sp. nov., B. muggon sp. nov., B. shieli sp. nov. and

B. triangulata sp. nov. For six of these seven species, we could construct molecular phylogenies and

parsimonious networks based on COI sequences. We tested for specific status and for potential cryptic

diversity of clades with Birky’s 4 theta rule. The analyses support the existence of these six species and

the absence of cryptic species in these lineages. Bennelongia triangulata sp. nov. is a common species

in the turbid claypans of the Murchison/ Gascoyne region. Bennelongia nimala itself is thus far known

only from the Northern Territory. Bennelongia tirigie sp. nov., B. pinderi sp. nov. and B. muggon sp.

nov. occur in the Murchison/ Gascoyne region, whereas B. koendersae sp. nov. and B. shieli sp. nov.

are described from the Pilbara. With the six new species described here, the genus Bennelongia now

comprises 31 nominal species.

Keywords. Taxonomy, evolution, biodiversity. Western Australia, Pilbara

Martens K., Halse S. & Schon I. 2015. On the Bennelongia nimala and B. triangulata lineages (Crustacea,

Ostracoda) in Western Australia, with the description of six new species. European Journal of Taxonomy 111:

1-36. http://dx.doi.org/10.5852/eit.2015.111

Introduction

The genus Bennelongia was originally described by De Deckker & McKenzie (1981) from Queensland,

with Bennelongia harpago as type species. Subsequently, De Deckker (1981) redescribed and transferred

Chlamydotheca australis Brady, 1886 to Bennelongia and added a further three species to the genus:

European Journal of Taxonomy 111: 1-36 (2015)

Bennelongia barangaroo De Deckker, 1981, Bennelongia nimala De Deckker, 1981 and Bennelongia

pinpi De Deckker, 1981. De Deckker (1982) then described Bennelongia tunta De Deckker, 1982 from

Queensland. For about 30 years after those papers, nothing was added to the taxonomy of the genus

Bennelongia, but Halse (2002) highlighted its high frequency of occurrence in Western Australia (WA).

Then Martens et al. (2012, 2013) and Sheam et al. (2012) together added 19 new species, mostly from

WA. De Deckker & Martens (2013) illustrated how different the valve morphologies of adult and juvenile

Bennelongia species can be and how juvenile morphologies vary between the lineages within the genus.

Here, we redescribe Bennelongia nimala from the Northern Territory (NT) and describe six new species

in the B. nimala and Bennelongia triangulata lineages within the genus, thus bringing the total number

of species in the genus to 31 (see Discussion). This is the fifth contribution in the recent revision of the

genus Bennelongia.

Material and methods

Collections

Ostracods were collected from pans and lakes with a hand net of mesh size of 250 pm during several

field trips (see below) (Fig. 1). Material for morphological analyses originated from both these ‘new’

collections and from earlier samples from all over WA, mostly collected by the research group of one

of us (SH) and preserved in a collection housed at the Department of Parks and Wildlife, Perth. The

molecular analyses were successful only with newly collected material, using either living specimens or

specimens sorted directly in the field and preserved in 100% ethanol. Consequently, molecular analyses

were limited to six of the seven species (re-)described here. Tocations of populations used for the present

paper are indicated on the map in Fig. 1. Type material of the new species is deposited in the Western

Australian Museum, Perth, Australia (WAMC numbers) and in the Ostracod Collection of the Royal

Belgian Institute of Natural Sciences, Brussels, Belgium (OC numbers) (see Table 1).

Morphological analyses

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

glycerine in sealed slides, or with soft parts used for molecular analyses. Drawings of soft parts were

made with a camera lucida on a compound microscope (Leica, DM 2500 at Bennelongia Environmental

Consultants, Perth). Valves were illustrated and measured using scanning electron microscopy (Philips

XT30 SEM at the Royal Belgian Institute of Natural Sciences, Brussels).

Molecular analysis

We used the Qiagen Blood and Tissue extraction kit following the manufacturer’s protocol to extract

DNA from 54 ostracods representing six species of the Bennelongia nimala and B. triangulata lineages.

The universal PCR primers of Folmer et al. (1994) for amplifying part of the mitochondrial COI region

were applied with the following conditions: 25 pi volumes of the HotStar Master Mix (Qiagen; 1.5 mM

MgCfy 200 pM dNTP, Tris Cl, KC1, (NHfySCfy 1.25 U Taq) and 0.1 pM of each primer were applied.

In a T personal Thermoblock (Biometra), we conducted PCRs with 15 min at 95°C, 40 cycles with 1 min

at 95°C, 1 min at 44° C, 1 min at 72° C, followed by a final extension step for 10 min at 72° C. Success

of PCR amplifications was verified by agarose gel electrophoresis and staining of gels with Gelred™.

We cleaned PCR products with the GFX™ PCR DNA and gel band purification kit (GE Healthcare)

and sequenced them in both directions with the universal primers and the Big Dye kit (ABI) on an ABI

3130X following the manufacturer’s protocol.

No fresh (living) material of B. shieli sp. nov. was obtained and this species is not represented in the

molecular phylogenetic tree and networks.

MARTENS K., HALSE S. & SCHON I., Bennelongia lineages in Western Australia

Analyses of sequence data

We used BioEdit (Hall 2007) to visualize sequence chromatograms. Sequence editing included

alignments of the forward and reverse sequence of each individual with ClustalX (Larkin et al. 2007),

followed by manual checking and correcting of ambiguities and trimming the final alignment to equal

length. We confirmed identity of the obtained sequences by BLAST searches (Altschul et al. 1990) in

Genbank. The optimal model of molecular COI evolution was assessed with 88 or 24 models and the

AICc criterion in jModeltest 2.1.1 (Darriba et al. 2012). We reconstructed phylogenies with two different

methods, Bayesian Inference (BI) in Mr Bayes 3.2 (Ronquist et al. 2011; with 5 million generations,

sampling every 100 th generation, a burn-in of 25% and the parameters identified by jModeltest for 24

different models) and the Maximum-Likelihood method in PhyML (Guindon & Gascuel 2003; with 1000

Fig. 1 . Map of Western Australia with localities of Bennelongia species described in the present paper.

European Journal of Taxonomy 111 : 1-36 ( 2015 )

Table 1 . Individual measurements of specimens used for the present descriptions. All measurements

were done using SEM (see Material and methods). If a molecular sequence was available for the same

specimen, the GenBanlc registration number is also given. However, some specimens were used as

whole animals for DNA sequencing, and thus no measurements are available. The present table therefore

does not list all 54 specimens for which sequences are available. Specimens in bold are holotypes.

Abbreviations: see Material and methods.

Museum nr

KMWA/OS

Genbank

Bennelongia

species

Locality

67 ?

CpRL

CPD/V

WAMC55564

KMWA.1361

triangulata

SIKE07

1850

1158

1923

1188

WAMC55565

KMWA.885

triangulata

SIKE07

2192

1308

2286

1350

WAMC55567

KMWA.886

triangulata

S1KE07

1892

1161

1950

1158

OC3368

KMWA.1364

triangulata

S1KE07

2190

1338

WAMC55569

KMWA.1365

triangulata

SIKE07

2138

1122

WAMC55570

KMWA.1366

triangulata

SIKE07

2168

1220

OC3369

KMWA.1367

triangulata

SIKE07

1818

1100

WAMC55571

KMWA.1368

triangulata

S1KE07

1917

1038

WAMC55572

KMWA.1369

triangulata

S1KE07

1933

1065

OC3367

KMWA.244

triangulata

CB54

1931

1072

2147

1122

WAMC55574

OS.254

triangulata

CB54

1888

996

1990

1006

WAMC55575

OS.122

triangulata

CB75a

1800

1075

WAMC55576

KMWA.884

triangulata

SIKE11

1961

1022

2069

1061

WAMC55577

KMWA.885

triangulata

S1KE11

2339

1300

2486

1297

WAMC55578

KMWA.701

triangulata

SIKE21

2173

1360

2238

1377

WAMC55579

KMWA.702

triangulata

S1KE21

2178

1331

WAMC55580

KMWA.703

triangulata

SIKE21

2161

1350

no nr

KMWA.1189

KP006594

triangulata

ESK101

1963

1230

2035

1255

no nr

KMWA.229

triangulata

OSTR10B

2030

1100

no nr

KMWA.230

triangulata

OSTR10B

2010

1030

no nr

KMWA.231

triangulata

OSTR10B

2080

1160

no nr

KMWA.232

triangulata

OSTRIOC

2260

1250

no nr

KMWA.234

triangulata

OSTRIOC

2160

1360

2270

1410

OC3371

KMWA.1037

KP006566

nimala

NT/12/01

1500

861

1640

962

WAMC55585

KMWA.1038

KP006565

nimala

NT/12/01

1520

872

1700

958

OC3372

KMWA.1039

nimala

NT/12/01

1730

1020

WAMC55586

KMWA.1040

nimala

NT/12/01

1680

1710

WAMC55587

KMWA.1043

KP006569

nimala

NT/12/03

1510

863

1630

954

WAMC55588

KMWA.1044

KP006570

nimala

NT/12/03

1480

853

1630

946

WAMC55589

KMWA.1047

nimala

NT/12/01

1546

902

1700

994

WAMC55590

KMWA.1048

KP006568

nimala

NT/12/08

1554

906

1748

1012

WAMC55591

KMWA.llll

nimala

CNJR-109

1383

781

1479

867

WAMC55592

KMWA.1112

nimala

CNJR-109

1462

852

1567

904

WAMC55593

KMWA.1113

nimala

CNJR-109

1450

840

1585

942

WAMC55594

KMWA.1114

nimala

CORN 9/v/09

1481

850

1594

937

OC3373

KMWA.1116

nimala

CORN 9/v/09

1627

1010

WAMC55595

KMWA.164

tirigie

SIEK4

1080

603

1190

650

WAMC55596

KMWA.169

tirigie

SIEK4

1180

692

1300

737

MARTENS K., HALSE S. & SCHON L, Bennelongia lineages in Western Australia

WAMC55597

KMWA.168

tirigie

SIEK4

1090

603

1100

659

WAMC55598

KMWA.165

tirigie

SIEK4

1290

800

WAMC55599

KMWA.166

tirigie

S1EK4

1300

755

WAMC55600

KMWA.167

tirigie

SIEK4

1280

724

WAMC55605

KMWA.1169

KP006532

tirigie

ESKI06

1029

593

WAMC55606

KMWA.1170

tirigie

ESKI06

1196

740

WAMC55607

KMWA.1171

tirigie

ESKI06

1136

638

WAMC55608

KMWA.1172

tirigie

ESKI06

1063

603

WAMC55609

KMWA.1173

tirigie

ESKI06

1042

597

WAMC55610

KMWA.1176

KP006539

tirigie

ESKI08

1129

660

OC3374

KMWA.1178

tirigie

ESKI08

1178

663

OC3375

KMWA.1179

tirigie

ESKI08

1086

615

WAMC55611

KMWA.171

koendersae

KIES1A

1110

587

1210

621

WAMC55612

KMWA.174

koendersae

KIES 1A

1353

767

1340

845

WAMC55613

KMWA.172

koendersae

KIES 1A

1190

629

1189

672

WAMC55614

KMWA.175

koendersae

KIES 1A

1240

689

1340

729

OC3376

KMWA.176

koendersae

KIES 1A

1260

714

1380

749

WAMC55617

KMWA.1260

koendersae

KIES 1A

1321

804

WAMC55618

KMWA.1261

koendersae

KIES 1A

1309

725

WAMC55620

KMWA.664

pinderi

SIKE03

1395

810

1507

840

WAMC55621

KMWA.665

pinderi

SIKE03

1508

898

WAMC55622

KMWA.666

pinderi

SIKE03

1450

810

OC3378

KMWA.667

pinderi

S1KE03

1433

865

WAMC55624

KMWA.671

pinderi

SIKE05

1410

813

1533

875

WAMC55625

KMWA.672

pinderi

S1KE05

1528

927

WAMC55626

KMWA.181

shieli

PSW036

1380

807

1467

888

WAMC55627

KMWA.290

shieli

PSW036

1311

745

1475

852

OC3379

KMWA.293

shieli

PSW036

1480

888

WAMC55628

KMWA.183

shieli

PSW036

1500

933

WAMC55629

KMWA.184

shieli

PSW036

1470

984

WAMC55630

KMWA.294

shieli

PSW036

1370

813

1480

907

WAMC55632

KMWA.1090

muggon

SIKE20

1015

603

1078

638

WAMC55633

KMWA.1084

muggon

S1KE20

1018

606

WAMC55634

KMWA.1085

muggon

SIKE20

1107

654

WAMC55635

KMWA.1086

muggon

S1KE20

1117

668

WAMC55636

KMWA.1087

muggon

SIKE20

1119

685

WAMC55637

KMWA.690

muggon

S1KE20

1040

629

1108

655

WAMC55638

KMWA.691

muggon

S1KE20

1111

699

WAMC55639

KMWA.692

muggon

SIKE20

1101

681

WAMC55640

KMWA.693

muggon

S1KE20

1078

660

WAMC55641

KMWA.1345

muggon

SIKE20

1082

636

WAMC55642

KMWA.1346

muggon

S1KE20

1021

603

WAMC55643

KMWA.1347

muggon

SIKE20

1043

621

650

OC3382

KMWA.1348

muggon

SIKE20

1058

629

1124

667

OC3383

KMWA.1349

muggon

SIKE20

1049

621

1121

672

WAMC55644

KMWA.1350

muggon

SIKE20

1004

600

1056

631

European Journal of Taxonomy 111 : 1-36 ( 2015 )

bootstrap replicates and the parameters of jModeltest for all 88 models), respectively. Genetic diversities

and relationships within and between populations were illustrated with parsimonious networks at the

95% probability limit with TCS 1.21 (Clement et al. 2000). Selected sequences of all species have been

submitted to Genbank (accession numbers KP006531-KP006599; see Table 1).

Testing for cryptic diversity

In the COI phylogenies of Bennelongia , well-supported phylogenetic clades (with bootstraps above 75%

or posterior probabilities above 0.85) were identified, which could represent different species following

the evolutionary genetic species concept (Birky & Barraclough 2009). We then used MEGA version

6.0 (Tamura et al. 2013) to estimate sequence diversities within and between these phylogenetic clades,

either using the number of differences (p) or the Tamura-3 parameter model with gamma distribution

and 1000 bootstrap replicates. Following Birky et al. (2010), we then corrected the obtained estimates

of sequence diversities for sample size. According to the 4 theta rule, sequence diversities between two

sister clades must be no less than 4 to 4.3 times larger than within the two clades, depending on the

number of sequences per clade (Birky et al. 2010). The 4 theta rule has been used previously on bdelloid

rotifers (Fontaneto et al. 2007, 2009; Birky & Barraclough 2009; Birky et al. 2011), sexual and asexual

ostracods (Bode et al. 2010; Schon et al. 2012), including other Bennelongia species (Martens et al.

2012, 2013; Shearn et al. 2012), asexual prokaryotes (Birky et al. 2010) and sexual vertebrates and

invertebrates (Birky 2013).

Abbreviations used in text and figures

CpD/CpV

CpRF

DPaW

K25

KMWA

Fpp

FVe

FVi

QFD

Rpp

RVe

RVi

carapace

carapace in dorsal/ ventral view

carapace in right lateral view

Department of Parks and Wildlife

female

height of valves

inn er list

electrical conductivity standardised to a water temperature of 25°C

original working numbers given to specimens dissected and illustrated by the first author

(KM)

length of valves

left prehensile palp

lateral shield of hemipenis

left valve

left valve, external view

left valve, internal view

male

medial shield of hemipenis

Northern Territory

Ostracod Collection in the Royal Belgian Institute of Natural Sciences (Brussels, Belgium)

Ostracod Slide dissected by Stuart Halse (SIT), retrieved from the voucher collection of

DPaW (Perth)

Queensland

right prehensile palp

right valve

right valve, external view

right valve, internal view

South Australia

MARTENS K., HALSE S. & SCHON L, Bennelongia lineages in Western Australia

Temp = temperature in °C

W = width of carapace

WA = Western Australia

WAMC = Western Australian Museum, Crustacean Collection (Perth, Australia)

Specimens in bold in Table 1 represent the holotypes of the new species.

Chaetotaxy of the limbs follows the model proposed by Broodbakker & Danielopol (1982), revised for

A2 by Martens (1987). Higher taxonomy of the Ostracoda follows the synopsis by Horne et al. (2002).

Results

Results of molecular screening

Both phylogenetic methods generated COI trees with similar topologies consisting of seven well-

supported phylogenetic clades (see the consensus tree in Fig. 2). Of these seven clades, five, namely B.

triangulata sp. nov., B. muggon sp. nov., B. koendersae sp. nov., B. pinderi sp. nov. and B. tirigie sp.

nov., match with the novel species described here from morphological data (see below). A sixth clade

100

1.00

B. triangulata

B. muggon

B. nimala

0.99

Too* 15

B. koendersae

Ttcri B. pinderi

Macroscapha walterae

0.05

Fig. 2. Phylogenetic tree constructed with Bayesian Inference (BI) and Maximum Likelihood (ML)

methods from COI sequences belonging to a total of 54 ostracods from the Bennelongia nimala and

B. triangulata lineages, respectively, with the marine ostracod Macroscapha walterae from Genbanlc

(accession number GU566887) as outgroup. Numbers above and below nodes illustrate statistical

support for this particular node. Numbers above nodes are % bootstrap values of ML analyses with 1000

replicates, numbers below nodes in italics represent Bayesian posterior probabilities (ranging from 0

to 1). Both methods, BI and ML, resulted in the same tree topology. Different phylogenetic clades are

indicated by different colours.

Fig. 3. Parsimonius networks, based on COI sequences of the Bennelongia nimala and B. triangulate/ lineages. Squares represent ancestral sequences

(or haplotypes), small circles missing haplotypes. The size of squares and large ovals is proportional to the number of individuals with the same

sequence in the analysed population. The networks were constructed with up to 14 mutations steps connecting different sequences or haplotypes.

Different phylogenetic clades are indicated by different colours (see Fig. 2).

European Journal of Taxonomy 111 : 1-36 ( 2015 )

MARTENS K., HALSE S. & SCHON L, Bennelongia lineages in Western Australia

Table 2. Results of tests for genetic species boundaries using the 4 theta rule for species of the Bennelongia

nimala and B. triangulata lineages, respectively. 0 = population genetic parameter theta, indicating

genetic variability within populations. D = genetic distance between sister clades. n 1 , n 2 = number of

sequences for each sister clade. 0 and D were either calculated as p distances or with the Tamura-3

parameter model (in italics). With the exception of B. koendersae sp. nov. and Spec. 15 (see Fig. 2),

no sister clades with high statistical support could be identified in the obtained COI trees; therefore, all

phylogenetic clades were compared to each other. In order to fulfil the criteria of the 4 theta rule for

genetic species status, the ratio of the mean sequence diversity within a phylogenetic clade (theta, 0) as

compared to the sequence diversity between this clade and its nearest phylogenetic neighbour (D) needs

to be 4 or more, depending on the number of specimens per clade (Birky et al. 2010). Comparisons, for

which these criteria are fulfilled, are printed in bold. (Table continued on next page.)

Species

max. 0

Ratio D/0

n 1 , n 2

(within clades)

(between clades)

B. koendersae - Spec. 15

0.0091

0.049

5.38

4,2

0.0091

0.053

5.82

B. koendersae - B. tirigie

0.0081

0.114

14.07

4, 14

0.0083

0.135

16.27

B. koendersae - B. pinderi

0.0148

0.106

7.16

4, 10

0.0154

0.124

8.05

0.0200

0.124

6.20

B. koendersae - B. nimala

0.0204

0.148

7.25

4,9

B. koendersae - B. muggon

0.0050

0.137

27.40

4,8

0.0052

0.168

32.31

B. koendersae - B. triangulata

0.0189

0.0196

0.129

0.157

6.83

8.01

4,21

Spec. 15 — B. tirigie

0.0091

0.113

0.133

12.42

14.62

2, 14

Spec. 15 - B. pinderi

0.0148

0.097

6.55

2, 10

0.0154

0.112

7.27

Spec. 15 — B. nimala

0.0200

0.140

7.00

2,9

0.0204

0.171

8.38

Spec. 15 — B. muggon

0.0091

0.143

15.71

2,8

0.0091

0.177

19.45

Spec. 15 — B. triangulata

0.0189

0.137

7.25

2,21

0.0196

0.170

8.67

B. pinderi - B. tirigie

0.0148

0.109

7.36

10, 14

0.0154

0.128

8.31

B. pinderi - B. nimala

0.0200

0.159

7.95

10,9

0.0204

0.204

10.00

European Journal of Taxonomy 111 : 1-36 ( 2015 )

B. pinderi - B. muggon

0.0148

0.148

10.00

10, 8

0.0157

0.186

11.85

B. pinderi - B. triangulata

0.0189

0.144

7.62

10,21

0.0196

0.181

9.23

B. tirigie - B. nimala

0.0200

0.146

7.30

14,9

0.0204

0.184

9.02

B. tirigie - B. muggon

0.0081

0.142

17.53

14, 8

0.0083

0.177

21.33

B. tirigie - B. triangulata

0.0189

0.136

7.20

14,21

0.0196

0.167

8.52

B. nimala - B. muggon

0.0200

0.144

7.20

9,8

0.0204

0.177

8.68

B. nimala - B. triangulata

0.0200

0.142

7.10

9,21

0 .0204

0.175

8.58

B. muggon - B. triangulata

0.0189

0.146

7.72

8,21

0.0196

0.179

9.13

contains the sequences of B. nimala (see below). Two additional sequences in the tree belonging to clade

Spec. 15 could not be investigated morphologically because the entire specimens had been used for the

molecular analyses. The sequences of Spec. 15 cluster together with B. koendersae sp. nov. with good

statistical support while the relationships among the other phylogenetic clades cannot be derived from

the COI tree because of lack of statistical support (Fig. 2). The seven phylogenetic clades are completely

congruent with seven isolated parsimonious networks (Fig. 3). Among the network structures, it appears

that B. triangulata sp. nov. is genetically most diverse, as this species contains 16 different haplotypes

(sequences) being separated by up to 15 mutational steps. With 10 haplotypes, the network of B. tirigie

sp. nov. is genetically the second most diverse species while the other species contain 7 ( B. nimala ), 5

( B. pinderi sp. nov.) and 4 (B. koendersae sp. nov. and B. muggon sp. nov.) haplotypes, respectively,

and their network structures are more simple. When comparing genetic diversities between and within

each phylogenetic clade, the genetic distances between all seven phylogenetic clades clearly exceed the

distances within each clade by more than 4 times (Table 2), thus fulfilling the criterion of the 4 theta

rule (Birky et al. 2010). The seven phylogenetic clades and networks can thus be regarded as different

genetic species according to the phylogenetic species concept. We found no evidence for cryptic genetic

diversity as all genetic species matched the morphological species with the exception of Spec. 15, for

which no morphological data are available.

MARTENS K., HALSE S. & SCHON L, Bennelongia lineages in Western Australia

Taxonomic descriptions

Class Ostracoda Latreille, 1806

Subclass Podocopa G.O. Sars, 1866

Order Podocopida G.O. Sars, 1866

Suborder Cypridocopina Baird, 1845

Superfamily Cypridoidea Baird, 1845

Family Cyprididae Baird, 1845

Subfamily Bennelongiinae Martens et al. , 2012

Genus Bennelongia De Deckker & McKenzie, 1981

Diagnosis

See Martens et al. (2012).

Bennelongia nimala - lineage

Diagnosis of the B. nimala- lineage

All species in this lineage with strongly calcified and heavily ornamented valves, also in adults external

valve surfaces set with large pustules, spines and short but stiff setae. Nearly all species with yellowish-

brownish colour. Most species also with very pronounced anterior LV/RV overlap, most pronounced of

all lineages in this genus. Some species with an inner ‘eyelet’ in the anterior part of the RV, close to the

lapel, just as in the species of the B. barangaroo lineage (see Martens et al. 2013), to which this lineage

is most closely related.

Bennelongia nimala De Deckker, 1981

Fig. 4A-N

Bennelongia nimala n. sp. - De Deckker, 1981: 105-108, figs 10-11.

Abbreviated redescription.

Valves in inner view (Fig. 4A-B) relatively high, with almost straight dorsal margin and greatest height

situated well in front of the middle; ventral margin anteriorly without mandibular curve. FV (Fig. 4A)

with antero-distal il running all the way down into the beak, but not connecting with ventral inner list;

antero-proximal il running slightly over hallway along the anterior margin; posterior il tuberculate and

running halfway up the posterior margin, next to the pointed valve margin. RV (Fig. 4B) with antero-

ventral lapel large, ventrally pointed and heavily serrated (Fig. 4F-N). Both valves with heavy external

ornamentation, consisting of pits, smaller and larger tubercles (Fig. 4C-D, F-G, K).

Cp (Fig. 4G, K, M) with FV overlapping RV on all sides, but moderately so; CpRF with FV forming

an antero-dorsal hump over RV. CpD with greatest width situated in the middle in females (Fig. 4G),

anteriorly with strong and slightly asymmetrical rostrum, dorsally set with parallel rows of tubercles.

From De Deckker (1981): Soft parts as typical of the genus. Hemipenis with lobe Is broad and plump,

antero-ventral extremity broadly rounded; lobe ms with antero-ventral extremity broad and rounded.

Rpp with unusually long and narrow distal segment. Fpp with sickle-shaped terminal segment, relatively

long and slender, also proximal segment long and narrow.

European Journal of Taxonomy 111: 1-36 (2015)

Fig. 4. Bennelongia nimala De Deckker, 1981 (all all from Kakadu National Park, NT). A. LVi

(WAMC55593). B. RVi (WAMC55593). C. RVe (WAMC55589). D. LVe (WAMC55589). E. RVi,

detail of central muscle scars (WAMC55593). F. LVe, detail of surface ornamentation (WAMC55589).

G. CpD (OC3373). H. LVi, detail of caudal side (WAMC55593). I. LVi, detail of anterior side

(WAMC55593). J. CpD (juvenile A-3, specimen lost). K. CpRL (OC3373). L. RVi, detail of anterior

margin (WAMC55593). M. CpRL, detail of anterior margin, showing external view of lapel (OC3373).

N. RVi, tilted, detail of anterior margin and of lapel (WAMC55593). Scale = 1 mm for A-D, G, K; 500

pm for J; 200 pm for E-F, H-I, L-N.

MARTENS K., HALSE S. & SCHON L, Bennelongia lineages in Western Australia

New material investigated

Kakadu National Park, Coonjimba Billabong, Gulungul Creek, Ranger, Jabiru region, NT (sample

CNJ-R-109). Approximate coordinates: 12°34’37” S, 132°52 , 30.1” E. Collected by Russell Shiel on 9

May 2009. Several females (WAMC55591-55593).

Kakadu National Park, Corndori Billabong , Gulungul Creek, Ranger, Jabiru region, NT (sample

CORN 9/v/09). Approximate coordinates: 12°37 , 50” S, 120°53 , 06” E. Collected by Russell Shiel on 9

May 2009. Several females (WAMC55594, OC3373).

Unnamed lagoon, Adelaide River Floodplain , NT (sample NT/12/01). Approximate coordinates:

12°53’06.5” S, 131°12’03.0” E. Collected by the authors on 28 Jul. 2012. Several females (OC3371-

3372, WAMC55585-55586, 55589). K25 = 58 pS/cm, Temp = 30,2°C, pH = 8.0, depth = ca. 0.3 m.

Bennett Dam, Adelaide River Floodplain , NT (sample NT/12/03). Approximate coordinates:

12°57 , 08.3” S, 131°09’59.7” E. Collected by the authors on 25 Jul. 2012. Several females

(WAMC55587-55588). K25 = 58 pS/cm, Temp = 28°C, pH = 7.2, depth = ca. 0.5 m.

Unnamed lagoon, Mary River Park, NT (sample NT/12/08). Approximate coordinates: 12°54 , 52.1” S,

131°39’23.7” E. Collected by the authors on 26 Jul. 2012. Several females (WAMC55590).

Type locality

Georgetown Lagoon, Jabiru, NT.

Measurements (all measurements in pm - see Table 1 for measurements of all newly collected

specimens illustrated with SEM)

Measurements of type material from De Deckker (1981):

Holotype c ?: RV: L = 1340, H = 760, LV: L = 1500, H = 840.

Paratype $: RV: L = 1540, H = 860; LV: L = 1640, H = 960.

Measurements of new material (only $

RV: L = 1450-1550, H = 780-900; LV: L = 1480-1750, H = 870-1010, W = ca. 1020.

Differential diagnosis

The species can be separated from all other congeners belonging to the B. nimala lineage by the

pointed caudal section of the LV and by the large, pronounced and heavily serrated lapel on the RV. The

moderate antero-ventral LV/RV overlap distinguishes this species specifically from B. tirigie sp. nov., B.

koendersae sp. nov. and B. maggon sp. nov.

Ecology and distribution

Bennelongia nimala is an NT species. It occurs in vegetated and unvegetated freshwater lagoons, from

which it derives its name.

Bennelongia tirigie sp. nov.

urn:lsid:zoobank.org:act:7323D117-25B3-4059-B42C-079286494218

Figs 5A-N, 6A-D

Abbreviated description

Valves in inner view (Fig. 5A, C-D, F) relatively elongated, with rounded dorsal margin and greatest

height situated well in front of the middle; ventral margin with pronounced mandibular curve anteriorly.

European Journal of Taxonomy 111: 1-36 (2015)

LV (Fig. 5A, D) with antero-distal il running only halfway along the anterior valve margin, antero-

proximal il running almost all the way up along the valve margin; posterior il tuberculate and running

halfway up the posterior margin. RV (Fig. 5C, F) with antero-ventral lapel relatively large, but bent

closely to valve surface and therefore less conspicuous (Fig. 5K-N). Valves with heavy external

ornamentation, mostly consisting of small tubercles (Fig. 5B, E, G-J).

Fig. 5. Bennelongia tirigie sp. nov. (all from type locality: Tirigie Claypan, Murchison/Gascoyne,

WA). A. LVi (allotype ?, WAMC55596). B. CpRL ($, WAMC 55600). C. RVi ($, WAMC55596).

D. LVi (holotype <J, WAMC55595). E. CpRL (& WAMC55597). F. RVi (f, WAMC55595).

G. CpD (?, WAMC55599). H. CpV ($, WAMC55598). I CpD (& WAMC55597). J. CpV (<?,

WAMC55597). K. RVi, detail anterior margin (§, WAMC55596). L. RVi, tilted, detail anterior margin

(5, WAMC55596). M. RVi, detail anterior margin (c?, WAMC55595). N. RVi, tilted, detail anterior

margin (f, WAMC55595). Scale = 1 mm for A-J; 200 pm for K-N.

MARTENS K., HALSE S. & SCHON L, Bennelongia lineages in Western Australia

Cp (Fig. 5B, E, G-J) with largest LV/RV overlap of all Bennelongia species known to date. CpD and

CpV with greatest width situated slightly behind the middle in males (Fig. 5I-J), in the middle in females

(Fig. 5G-H), anteriorly with strong and asymmetrical rostrum.

Soft parts as typical of the genus. Hemipenes (Fig. 6A-B) almost symmetrical, edge of lobe ms almost

straight, lobe Is with extremity ventrally pointed. Fpp (Fig. 6C) with distal segment rather narrow.

Rpp (Fig. 6D) with distal segment rather broad, elongated and distally rounded, sensory organ on first

segment stout.

Etymology

The species is named after its type locality, Tirigie Claypan in Gascoyne, WA.

Type material

Holotype

S (WAMC55595), with soft parts dissected in a sealed slide and valves stored dry in a micropalaeon-

tological slide.

Allotype

9 (WAMC55596), with valves stored dry in a micropalaeontological slide and soft parts used for

molecular screening.

Paratypes

Numerous females and males from the type locality, either dissected or stored as the holotype, as

carapaces used for SEM or in alcohol. See Table 1 for listing of specimens (WAMC5597-55601).

Other material investigated

Crackers Swamp , Dandaragan, WA (sample SIEK1). Approximate coordinates: 30°54’36” S,

115°35’30.2” E. All material collected by the authors on 5 Apr. 2006. Several females.

Minilya Pool, Gascoyne, WA (sample SIKE19). Approximate coordinates: 23°54’25” S, 114 o 0E47.3”

E. All material collected by the authors on 7 Jul. 2011. K25 = 693 pS/cm, Temp = 17.3°C, pH = 7.3.

Unnamed crabhole swamp on Winning Station, Gascoyne, WA (sample ESKI05). Approximate

coordinates: 23°15 , 22” S, 114 0 44’57.8” E. All material collected by the authors on 6 Apr. 2013. Several

females (WAMC55602-55604).

Dam on Winning Station, Gascoyne, WA (sample ESKI06). Approximate coordinates: 22° 14’16.0”

S, 114°42 , 34.0” E. All material collected by the authors on 6 Apr. 2013. Several males and females

(WAMC556055-55609).

Unnamed large claypan on Winning Station, Gascoyne, WA (sample ESKI08). Approximate

coordinates: 23°07’39.6” S, 114°34’41.3” E. All material collected by the authors on 6 Apr. 2013.

Several males and females (WAMC55610, OC3374-3375).

Type locality

AUSTRALIA: Tirigie Claypan, Gascoyne, WA (sample SIEK4), approximate coordinates: 24°38’29” S,

113°59 , 44” E. All material collected by the authors on 7 Apr. 2006.

European Journal of Taxonomy 111: 1-36 (2015)

Differential diagnosis

The large frontal LV/RV overlap and the shape of the lapel on the RV distinguish this species from all

others in the B. nimala lineage. The shape of the Is on the hemipenes and of the distal segment of the Rpp

enables this species to be distinguished from others of the B. nimala lineage for which males are known:

B. nimala has a broader Is and an even narrower distal segment on the Rpp; B. koendersae sp. nov. has an

Is that is longer, more robust and not so pointed, while the distal segment of its Rpp is broadly triangular,

with almost straight margins; in B. regina Shearn et al., 2012 the distal segment of the Rpp is evenly

rounded while the Is of the hemipenes end in small, birdhead-like lobes.

Measurements (all measurements in pm - see Table 1 for measurements of all specimens illustrated

with SEM)

Holotype $ (WAMC55595): RV: L = 1080, H = 603; LV: L = 1190, H = 650.

Allotype $ (WAMC55596): RV: L = 1180, H = 692; LV: L = 1300, H = 737.

Ecology and distribution

Bennelongia tirigie sp. nov. is a common species in turbid seasonal water bodies of the Gascoyne region

of WA. Its range extends south to the northern Swan Coastal Plain, where it has been collected from a

seasonal freshwater swamp of low turbidity.

Bennelongia koendersae sp. nov.

um:lsid:zoobank.org: act: 6CA91421-02FF -4A2E-A0E6-89F417057C81

Figs 6E-H, 7A-N

Abbreviated description

Valves in inner view (Fig. 7A, C-D, F) relatively elongated, with rounded dorsal margin and greatest

height situated well in front of the middle; ventral margin anteriorly without pronounced mandibular

curve. LV (Fig. 7A, D) with antero-distal il running only halfway along the anterior valve margin,

antero-proximal il ru nn ing slightly higher along the anterior valve margin; posterior il tuberculate and

running halfway up the posterior margin. RV (Fig. 7C, F) with antero-ventral lapel large, rounded and

strongly serrated (Fig. 7K-N). Valves with heavy external ornamentation, mostly consisting of small

tubercles (Fig. 7B, E, G-J).

Cp (Fig. 7B, E, G-J) with large LV/RV overlap, but less so than in B. tirigie sp. nov. CpD and CpV with

greatest width situated in the middle in males (Fig. 7G-H), slightly behind the middle in females (Fig.

7I-J), anteriorly with strong and asymmetrical rostrum.

Soft parts as typical of the genus. Hemipenes (Fig. 6E-F) almost symmetrical, edge of ms slightly

sinuous. Is with extremity rounded. Lpp (Fig. 6G) with distal segment rather broad, distal half of second

segment with parallel margins. Rpp (Fig. 6H) with distal segment large, subtriangular and with almost

straight margins, sensory organ on first segment stout.

Etymology

The species is named after Dr Annette Koenders (Edith Cowan University, Joondalup, WA), in recognition

of her contribution to our knowledge of the genetic diversity of various Australian invertebrate groups.

Type material

Holotype

S (WAMC55611), with soft parts dissected in a sealed slide and valves stored dry in a micropalaeonto-

logical slide.

MARTENS K., HALSE S. & SCHON L, Bennelongia lineages in Western Australia

Fig. 6. Bennelongia tirigie sp. nov. (A-D, WAMC55595, holotype) and B. koendersae sp. nov. (E-

H, WAMC55615, paratype), all <$. A-B. Outlines of hemipenes. C. Lpp. D. Rpp. E-F. Outlines of

hemipenes. G. Lpp. H. Rpp. Scale = 73 pm for A, D-E; 31 pm for B-C, F-H.

European Journal of Taxonomy 111: 1-36 (2015)

Allotype

9 (WAMC55612) carapace stored dry in a micropalaeontological slide.

Paratypes

Numerous males and females from the type locality, either dissected and stored as the holotype, as carapaces

used for SEM or in alcohol (WAMC55613-55619, OC3376-3377). See Table 1 for listing of specimens.

Fig. 7. Bennelongia koendersae sp. nov. (all from type locality: claypan at Strelley Station, Pilbara,

WA). A. LVi (?, OC3376). B. CpRL ($, WAMC55618). C. RVi (?, OC3376). D. LVi (holotype

WAMC55611). E. CpRL (f, WAMC55613). F. RVi ($, WAMC55611). G. CpD WAMC55613).

H. CpV (cJ, specimen lost). F CpD (allotype ?, WAMC55612). J. CpV (?, WAMC55617). K. RVi,

detail anterior margin (§, OC3376). L. RVi, tilted, detail anterior margin (§, OC3376. M. RVi, detail

anterior margin (c?, WAMC55611). N. RVi, tilted, detail anterior margin (c?, WAMC55611). Scale =

1 mm for A-J; 200 pm for K-N.

MARTENS K., HALSE S. & SCHON L, Bennelongia lineages in Western Australia

Type locality

AUSTRALIA: Strelley Station, shallow unnamed claypan, Pilbara, WA (sample KIES 1 A), approximate

coordinates: 20°24 , 58” S, 118°59’4” E. All material collected by the authors on 21 Apr. 2006.

Measurements (all measurements in pm - see Table 1 for measurements of all specimens illustrated

with SEM)

Holotype $ (WAMC55611): RV: L = 1110, H = 587; LV: L = 1210, H = 621.

Paratype ? (WAMC55612): RV: L = 1240, H = 689; LV: L = 1340, H = 729.

Differential diagnosis

Bennelongia koendersae sp. nov. can be distinguished from all other species in the B. nimala lineage

by the size and shape of the antero-ventral lapel on the RV, and from B. tirigie sp. nov. by the less

pronounced anterior LV/RV overlap. Bennelongia koendersae sp. nov. can be distinguished from those

species in the lineage where males are known (B. nimala , B. regina , B. tirigie sp. nov.) by the large and

triangular second segment of the Rpp.

Ecology and distribution

The species is only known from its type locality, which is a small, ephemeral, turbid claypan.

Remarks

In a second sample from the 2006 KIES collecting expedition in Pilbara, a species close to B. koendersae

sp. nov. was recognised with molecular methods only. This species (as Spec. 15 in Fig. 2) looked very

much like B. koendersae sp. nov. and the only two specimens available were both screened in toto by

molecular methods to establish conspecificity. Although these specimens cluster close to B. koendersae

sp. nov. in the phylogenetic COI tree, they are considered to represent a second species (Fig. 2). As no

further specimens are available, it could not be checked whether the species is cryptic or morphologically

distinguishable from B. koendersae sp. nov. We thus leave this clade in open nomenclature.

Locality of Spec. 15

Ethel Creek claypan , east of Roy Hill Station, Pilbara, WA (sample KIES 15). Approximate coordinates:

22 0 4E28” S, 119°58’45” E. Specimens collected by the authors on 24 Apr. 2006.

Bennelongiapinderi sp. nov.

urn:lsid:zoobank.org:act:B2630A2B-D21E-4A5E-8B15-D5E0D9C13FAF

Figs 8A-G

Abbreviated description

Valves in inner view (Fig. 8A, C) relatively elongated, with rounded dorsal margin and greatest height

situated slightly in front of the middle; ventral margin anteriorly without pronounced mandibular curve.

LV (Fig. 8A) with antero-distal il running over halfway along the anterior valve margin, antero-proximal

il running about 4/5 way up along the anterior valve margin; posterior il tuberculate, but more delicately

so than in the preceding three species, and running halfway up the posterior margin. RV (Fig. 8C) with

antero-ventral lapel pronounced and rounded but with a smooth edge (Fig. 8F-G). Valves with external

ornamentation mostly consisting of small tubercles, less pronounced than in the three preceding species

(Fig. 8C-E).

Cp (Fig. 8C-E) with large LV/RV overlap, but less so than in the two preceding species. CpD and CpV

with greatest width situated in the middle in females, anteriorly with strong and asymmetrical rostrum.

European Journal of Taxonomy 111: 1-36 (2015)

Soft parts as typical of the genus.

Male unknown.

Etymology

The species is named after Adrian M. Pinder (DPaW, Science and Conservation Division, Kensington,

WA) in recognition of his substantial contribution to our knowledge about the taxonomy and ecology of

freshwater invertebrates of Australia, especially freshwater Oligochaeta.

Type material

Holotype

5 (WAMC55620), valves stored dry in a micropalaeontological slide and soft parts used for molecular

screening.

Paratypes

Numerous females from the type locality, either stored as the holotype, as carapaces used for SEM or in

alcohol (WAMC55621-55623, OC3378). See Table 1 for listing of specimens.

Other material investigated

Unnamed claypan near Murchison River , Murchison, WA (sample SIKE01). Approximate coordinates:

27°50 , 03” S, 114°43 , 37” E. All material was collected by the authors on 5 Jul. 2011. K25 = 28 pS/cm,

Temp = 8.3°C, pH = 5.8.

Fig. 8. Bennelongia pinderi sp. nov. (all $, all from type locality: unnamed claypan in Murchison/

Gascoyne, WA). A. LVi (holotype, WAMC55620). B. CpRL (WAMC55622). C. RVi (WAMC55620).

D. CpD (WAMC55621). E. CpV (OC3378). F. RVi, detail anterior margin (WAMC55620). G. RVi,

tilted, detail anterior margin (WAMC55620). Scale = 1 mm for A-E; 200 pm for F-G.

MARTENS K., HALSE S. & SCHON L, Bennelongia lineages in Western Australia

Roadside ditch on Euardy Station , Murchison, WA (sample SIKE05). Approximate coordinates:

27°35 , 31” S, 114°41 , 43” E. All material was collected by the authors on 5 Jul. 2011. Several females

(WAMC55624-55625). K25 = 31 pS/cm, Temp = 14°C, pH = 6.7.

Type locality

AUSTRALIA: Unnamed swamp, Coolcalalaya Station, Murchison, WA (samples CB06a, SIKE 3).

Approximate coordinates: 27°3r22” S, 115°04’23” E. All material collected by the authors on 5 Jul.

2011. K25 = 33 pS/cm, Temp = 13.6°C, pH = 7.8.

Measurements (all measurements in pm - see Table 1 for measurements of all specimens illustrated

with SEM)

Holotype ? (WAMC55620): RV: L = 1395, H = 810; LV: L = 1507, H = 840.

Differential diagnosis

Bennelongiapinderi sp. nov. is most closely related to B. koendersae sp. nov., but is ca. 15-20% larger,

has a less pronounced valve ornamentation and has a less-developed antero-ventral lapel with a smooth

edge on the RV rather than serrated as in B. koendersae sp. nov.

Ecology and distribution

Bennelongia pinderi sp. nov. has thus far been found in three seasonal or ephemeral water bodies in the

Murchison region of WA.

Bennelongia muggon sp. nov.

urn:lsid:zoobank.org:act:38D3B6B7-30B5-4F70-AE39-A46A3213719C

Fig. 9A-N

Bennelongia cf. nimala nov. sp. - De Deckker & Martens 2013: 6-7, figs 2-10.

Abbreviated description

Valves in inner view (Fig. 9A ,C) relatively high, with rounded dorsal margin and greatest height situated

well in front of the middle; ventral margin anteriorly with slight mandibular curve. LV (Fig. 9A) with

antero-distal il running over halfway along the anterior valve margin, antero-proximal il running less

than halfway up along the anterior valve margin; posterior il running more than halfway up the posterior

margin, but mostly smooth, not tuberculate. RV (Fig. 9C) with antero-ventral lapel pronounced and

droplet-shaped, with delicately serrated edge (Fig. 9C, H-I, K-N). Valves with external ornamentation

mostly consisting of small tubercles (Fig. 9B, D-H).

Cp (Fig. 9C-G) with strong LV/RV overlap, almost as large as in B. tirigie sp. nov. CpD and CpVwith

greatest width situated in the middle in females, anteriorly with less pronounced, asymmetrical rostrum.

Soft parts as typical of the genus.

Male unknown.

Etymology

The species is named after its type locality, a large lake on Muggon Station, Murchison, WA.

European Journal of Taxonomy 111: 1-36 (2015)

Fig. 9. Bennelongia muggon sp. nov. (all from type locality: Lake Muggon, Murchison/Gascoyne,

WA). A. LVi (holotype, WAMC55632). B. CpRL (WAMC55635). C. RVi (WAMC55632). D. CpD,

detail anterior margin (WAMC55634). E. CpD (WAMC55634). F. CpV (WAMC55636). G. CpV, detail

anterior margin (WAMC55636). H. CpRL, detail anterior margin (WAMC55635). I. RVi (WAMC55633).

J. RVi, detail posterior margin (WAMC55633). K. RVi, detail anterior margin (WAMC55632). L. RVi,

tilted, detail anterior margin (WAMC55632). M. RVi, detail anterior margin (WAMC55633). N. RVi,

tilted, detail anterior margin (WAMC55633). Scales = 1 mm for A-C, E-F, I; 200 pm for D, G-H, J-N.

MARTENS K., HALSE S. & SCHON L, Bennelongia lineages in Western Australia

Type material

Holotype

$ (WAMC55632), valves stored dry in a micropalaeontological slide, with soft parts used for molecular

screening.

Paratypes

Numerous females from the type locality, either stored as the holotype, as carapaces used for SEM or in

alcohol (WAMC55633-55645, OC3381-3383). See Table 1 for listing of specimens.

Other material investigated

Unnamed canegrass pan on Boolathana Station , Gascoyne, WA (sample SIKE 12). Approximate

coordinates: 24 0 44 , 4E’ S, 113°43’22” E. All material collected by the authors on 6 Jul. 2011. K25 =

1020 pS/cm, pH = 8.7, Temp = 16.9°C.

Dam at Solomon’s Well , Victoria Plains, WA (sample DJC/04). Approximate coordinates: 31° 11 ’59” S,

116°21 ’47.7” E. All material collected by David J. Cale (DPaW, Kensington) on 9 Sep. 2011. K25 = 120

pS/cm, pH = 6.8, Temp = 14.4°C.

Petrudor Dam, Wheatbelt, WA (sample DJC/15). Approximate coordinates: 30°25 , 19” S, 116°57’40”

E. All material collected by David J. Cale (DPaW, Kensington) on 11 Sep. 2011. K25 = 162 pS/cm, pH

= 7.85, Temp = 22.0°c.

Type locality

AUSTRALIA: Unnamed large lake at Muggon Station, Murchison, WA (sample SIKE 20). Approximate

coordinates: 26°44’ 15” S, 115°29’59” E. All material collected by the authors on 8 Jul. 2011. K25 = 475

pS/cm, pH = 8.9, Temp = 13.4°C.

Measurements (all measurements in pm - see Table 1 for measurements of all specimens illustrated

with SEM)

Holotype $ (WAMC55632): RV: L = 1015, H = 603; LV: L = 1078, H = 638.

Differential diagnosis

This is the smallest of all species described in the present paper. Bennelongia muggon sp. nov. can

further be distinguished from other members of the B. nimala lineage by the size and shape of the antero-

ventral lapel on the RV, the smooth posterior il and the short anterior inner il in the LV.

Ecology and distribution

This is one of the more common species in this lineage, as it has been found in several freshwater dams

and pans from the Gascoyne region south to the Wheatbelt, WA. It occurs in fresh turbid water.

Bennelongia shieli sp. nov.

urn: lsid: zoobank, org: act: 3739D A5 6-41CC-4 7FA-93 A6-44BBFBFFB 5 2E

Fig. 10A-M

Abbreviated description

Valves in inner view (Fig. 10A-B, G, I) relatively high, with almost straight dorsal margin, parallel

to ventral margin in RV, sloping caudally in LV; greatest height situated well in front of the middle;

ventral margin anteriorly with slight mandibular curve. LV (Fig. 10A, G) with antero-distal il running

almost along the entire anterior valve margin, almost connecting with ventral inner list; antero-proximal

European Journal of Taxonomy 111: 1-36 (2015)

il reaching halfway up along the anterior valve margin; posterior il running more than halfway up the

posterior margin, tuberculate for most of its length. LV with 2-3 spines halfway up the posterior margin

(Fig. 10A, D, G). RV (Fig. 10B, I) with antero-ventral lapel elongated and slightly serrate, with a large

tooth (Fig. 10B, I, K-M). Valves with external ornamentation mostly consisting of small tubercles (Fig.

10C-F, H). (Remark: one specimen showed a larger tooth on the antero-ventral lapel on the RV; this may

be an aberrant individual).

Cp (Fig. 10E-F, H) with strong LV/RV overlap, but less so than in B. tirigie sp. nov., B. koendersae sp.

nov. and B. muggon sp. nov. CpRL with LV forming an antero-dorsal hump over RV. CpD and CpV

with greatest width situated slightly behind the middle in females, anteriorly with strongly pronounced,

asymmetrical rostrum, dorsally set with parallel rows of tubercles, as in B. nimala.

Soft parts as typical of the genus.

Male unknown.

Etymology

The species is named after Dr Russell Shiel (University of Adelaide, Adelaide) in recognition of his

substantial contribution to the taxonomy and ecology of freshwater invertebrates, especially rotifers and

cladocerans, of Australia.

Type material

Holotype

5 (WAMC55626), with soft parts dissected in a sealed slide and valves stored dry in a micropalaeonto-

logical slide.

Paratypes

Numerous females from the type locality, either stored as the holotype, as carapaces used for SEM or in

alcohol (WAMC55627-55631, OC3379-3380). See Table 1 for listing of specimens.

Type locality

AUSTRALIA: Munreemya Billabong, Pilbara, WA (sample PSW036 = OSTR133). Approximate

coordinates: 20°40U2” S, 120°13’33.6” E. Material collected by Adrian Pinder and Harley Barron on

19 May 2004. K25 = 194 pS/cm, pH = 8.54, Temp = 23°C.

Measurements (all measurements in pm - see Table 1 for measurements of all specimens illustrated

with SEM)

Holotype ? (WAMC55626): RV: L = 1380, H = 807; LV: L = 1467, H = 888.

Differential diagnosis

In shape and valve ornamentation, the species differs from all others described here except B. nimala ,

which it resembles. However, B. shieli sp. nov. has a much wider anterior LV/RV overlap than B. nimala

and is also about 10% smaller. In B. nimala, the anterodorsal inner list almost connects with the ventral

list and clearly overlaps with the anteroventral inner list. In addition, the antero-ventral lapel on the RV

is also different in both species.

Ecology and distribution

The species is known from its type locality only, a freshwater semi-permanent billabong in the northern

Pilbara.

MARTENS K., HALSE S. & SCHON I., Bennelongia lineages in Western Australia

Fig. 10. Bennelongia shieli sp. nov. (all §, all from type locality: Munreemya Billabong, Pilbara,

WA). A. LVi (holotype, WAMC55626). B. RVi (WAMC55626). C. RVe (WAMC55627). D. LVe

WAMC55627). E. CpD (WAMC55628). F. CpV (WAMC55629). G. LVi (WAMC55630). H. CpRL

(OC3379). I RVi (WAMC55630). J. RVi, detail central muscle scars (WAMC55627). K. RVi, detail

anterior margin (WAMC55626). L. RVi, tilted, detail anterior margin (WAMC55626). M. RVi, detail

anterior margin (WAMC55630, aberrant specimen). Scales = 1 mm for A—I; 200 pm for J-M.

European Journal of Taxonomy 111: 1-36 (2015)

Bennelongia triangulata lineage

Diagnosis of the B. triangulata lineage

Large (L > 2 mm) and triangular species, with ventral margins nearly smooth, without an antero-ventral

beak on the LV. Anterior LV/RV overlap moderate. CpD and CpV without pronounced anterior rostrum.

Last 3 juvenile stages with fully flat ventral side, not so in adults.

Remarks

De Deckker & Martens (2013) described the morphology of the last 3 instar juveniles of B. triangulata

sp. nov. (as B. sp. 414) and found that they have, unlike juveniles of any other Benelongia species, a fully

flat ventral side. This feature is an important part of the diagnosis of the lineage.

This lineage and species are atypical for Bennelongia , in that some of the obvious features, such as the

long beak in the B. australis lineage or the large LV/RV overlap in the B. nimala lineage, are missing.

Nevertheless, the structure of the valve margins of both valves is most similar to the other species in the

genus, and also the soft parts, apart from being slightly more elongated, show no structural differences.

We thus maintain this species in Bennelongia.

Bennelongia triangulata sp. nov.

urn:lsid:zoobank.org:act:2EE84F9D-40BD-4866-9E42-5B6815D95ElA

Figs 11A-M, 12A-M, 13A-G

Bennelongia sp. nov. 414 - Halse et al. 2000: table 4, appendix 4.

Bennelongia n. sp. 414 - Pinder et al. 2010: appendix 2.

Bennelongia sp. 414 nov. sp. - De Deckker & Martens 2013: 7-8, figs 11-12.

Abbreviated description

Valves in inner view (Figs 11 A, C, H, J; 12A, C, H, J) subtriangular, with greatest height situated either

in the middle or just before the middle; ventral margin almost straight, with weak mandibular curve.

LV (Figs 11 A, H; 12A, H) with antero-distal il running only halfway along the anterior valve margin,

antero-proximal il running slightly beyond halfway up along the valve margin, this list dorsally strongly

S-shaped; posterior il smooth, not tuberculate, and running only along ventral margin. RV (Figs 11C,

J; 12C, J) with antero-ventral lapel a very narrow ridge, but valve margin protruding beyond selvage as

in the Bennelongia pinpi lineage (Figs 11C, I-M; 12C-D, G, I-J, L-M). Valves with delicate external

ornamentation, consisting of small tubercles (Figs 11D, G, L; 12G, I, M).

Cp (Fig. 11B, D-G, L) with moderate LV/RV overlap. CpD and CpV with greatest width situated in the

middle in both males and females (Figs 11E-F; 12E-F), without anterior rostrum.

Soft parts as typical of the genus, but generally more elongated than in the other species of Bennelongia.

Hemipenes (Fig. 13A, E) symmetrical, edge of ms almost straight, Is with ventral extremity strongly

pointed. Lpp (Fig. 13B, F) with distal segment narrow, tapering towards the point and rather long (the

specimen in Fig. 13C is an exception). Rpp (Fig. 13D, G) with distal segment narrowly subtriangular,

with apical margin straight, sensory organs on first segment unequal, one long, one short.

Etymology

The species is named after its most striking character, the triangular shape of the valves in lateral view.

MARTENS K., HALSE S. & SCHON I., Bennelongia lineages in Western Australia

Fig. 11. Bennelongia triangulata sp. nov. (all A-G & K-M from type locality: canegrass pan on

Wooramel Station, Murchison/Gascoyne, WA; H-J from CB54, Boolathana Station, Carnarvon Basin,

WA). A. LVi (allotype, WAMC55565). B. CpRL (OC3368). C. RVi (WAMC55565). D. CpD, detail

anterior margin (WAMC55569). E. CpD (WAMC55569). F. CpV (WAMC55570). G. CpV, detail

anterior margin (WAMC55570). H. LVi (OC3370). I. LVi, detail posterior margin (OC3370). J. RVi

(OC3370). K. RVi, detail anterior margin (WAMC55565). L. CpRL, detail anterior margin (OC3368).

M. RVi, tilted, detail anterior margin (WAMC55565). Scales = 1 mm for A-C, E-F, H, J; 200 pm for

D, G, I, K-M.

European Journal of Taxonomy 111: 1-36 (2015)

Fig. 12. Bennelongia triangulata sp. nov. (all $, A-G from type locality: canegrass pan on Wooramel

Station, Murchison/Gascoyne, WA; H-M from CB54, Boolathana Station, Carnarvon Basin, WA). A. LVi

(holotype, WAMC55564). B. CpRL (OC3369). C. RVi (WAMC55564). D. RVi, detail anterior margin

(WAMC55564). E. CpD (WAMC55571). F. CpV (WAMC55572). G. RVi, tilted, detail anterior margin

(WAMC55564). H. LVi (WAMC55574). I. RVi, tilted, detail anterior margin (WAMC55564). J. RVi

(WAMC55574). K. LVi, detail central muscle scars (WAMC55574). L. RVi, detail anterior margin

(WAMC55574). M. RVi, tilted, detail anterior margin (WAMC55574). Scale = 1 mm for A-C, E-F, H,

J; 200 pm for D, G, I, K-M.

MARTENS K., HALSE S. & SCHON L, Bennelongia lineages in Western Australia

Fig. 13. Bennelongia triangulata sp. nov. (all S, A-D from CB75a, Cattle Camp Pan, Gascoyne, WA;

E-G holotype (WAMC55564): cane grass pan on Wooramel Station, Murchison/Gascoyne, WA).

A. Outline of hemipenis (WAMC55584). B. Lpp (WAMC55584). C. Lpp (WAMC55575). D. Rpp

(WAMC55584). E. Outline of hemipenis. F. Lpp. F’. Lpp, detail of distal part of first segment. G. Rpp.

Scale = 156 pm for A, E; 73 pm for B-D, F-G.

European Journal of Taxonomy 111: 1-36 (2015)

Type material

Holotype

S (WAMC55564), with soft parts dissected in a sealed slide and valves stored dry in a micropalaeon-

tological slide.

Allotype

5 (WAMC55565), with valves stored dry in a micropalaeontological slide and soft parts used for

molecular screening.

Paratypes

Numerous SS and $ $ from the type locality, either dissected and stored as the holotype, as carapaces

used for SEM or in alcohol (WAMC55566-55573, OC3367-3369). See Table 1 for listing of specimens.

Other material investigated

Unnamed claypan , Gascoyne, WA (sample SIKE11). Approximate coordinates: 24°47’49” S,

114°15’42” E. All material collected by the authors on 6 Jul. 2011. Several males and females (specimens

WAMC55576-55577). K25 = 434 pS/cm, Temp = 15.4°C, pH = 7.6.

Unnamed claypan , Muggon Station, Murchison, WA (sample SIKE21). Approximate coordinates:

26°46’54” S, 115°40’53” E. All material collected by the authors on 8 Jul. 2011. Several males and

females (specimens WAMC55578-55583). K25 = 154 pS/cm, Temp = 13.3°C, pH = 8.0.

Homestead dam, Boolathana Station , Gascoyne, WA (sample ESKI/01). Approximate coordinates:

24 0 39 , 15 ,, S, 113 0 4E37” E. All material collected by the authors on 5 Apr. 2013. Several females

(specimens WAMC55584).

Unnamed canegrass pan, Wooramel Stn , Gascoyne, WA (sample CB35a). Approximate coordinates:

25°40’52” S, 114°13’14” E. Material collected by Stuart Halse on 24 Aug. 1994 (specimen OS256).

Near Cardabia Swamp , Gascoyne, WA (sample CB54). Approximate coordinates: 24°33’10” S,

113 0 45’35” E. Material collected by Stuart Halse on 18 Mar. 1995 (specimen OC3370).

Unnamed claypan, Doorawarrah Stn , Gascoyne, WA (sample CB58b). Approximate coordinates:

24°48’8” S, 114°16’ 15” E. Material collected by Stuart Halse on 23 Aug. 1994 (specimen WAMC55574).

Boolan Pool , Gascoyne, WA (sample CB73). Approximate coordinates: 24°28’38” S, 113°40’36” E. All

material collected by Stuart Halse on 18 Aug. 1994 (specimens OS 121, OS391).

Cattle Camp Pan, Gascoyne, WA (sample CB75a). Approximate coordinates: 24°28’25” S, 114°13’27”

E. Material collected by Stuart Halse on 19 Aug. 1994 (specimen WAMC55575).

Nicabay Flats , Gascoyne, WA. Approximate coordinates: 24°52’7” S, 113 0 43’3” E. Material collected

by Stuart Halse on 15 Aug. 1995 (specimen OS258).

Type locality

AUSTRALIA: Unnamed canegrass pan on Wooramel Station, Gascoyne, WA (samples CB35a and

SIKE07). Approximate coordinates: 25°40’53” S, 114° 13’17” E. All material collected by the authors

on 5 Jul. 201E K25 = 307 pS/cm, Temp = 15.5°C, pH = 7.4.

MARTENS K., HALSE S. & SCHON L, Bennelongia lineages in Western Australia

Measurements (all measurements in pm - see Table 1 for measurements of all specimens illustrated

with SEM)

Holotype $ (WAMC55564): RV: L = 1850, H = 1158; LV: L = 1923, H = 1188.

Allotype $ (WAMC55565): RV: L = 2192, H = 1308; LV: L = 2286, H = 1350.

Differential diagnosis

This species is unlike any other species in the genus, because of the features cited in the diagnosis to the B.

triangulata lineage, especially the large size, triangular shape and lack of an antero-ventral beak on the LV.

Remark

The species comes in two forms, one with valves high (Figs 11A-C; 12A-C) and one with valves more

elongated (Figs 11H, J; 12H, J). In the soft parts, both forms are almost identical, with the exception of

the shape of the second segment of the Rpp. In the high form, this segment is subtriangular, with both

dorsal and distal margins straight (Fig. 13D). In the more elongated form, this segment has a straight

distal, but more sinuous dorsal margin (Fig. 13G).

Morphology, ecology and distribution

Bennelongia triangulata sp. nov. is common in the Murchison region north to the southern Pilbara

region of WA and occurs in turbid claypans. As outlined above, the valve shape can be quite different

(either rather high, or quite elongated), and originally it was thought that these two forms were either

seasonally determined or constituted two species.

New sampling and screening of older collections showed that both forms occur in both summer and

winter. Laboratory cultures reared at different temperatures, day length etc. may show which, if any,

aspects of the environment may affect valve shape in this species. Males with different valve shape have

slightly different morphologies in the Rpp, but as this could be part of a normal range of variability, its

occurrence should be checked on longer series of dissections. Bennelongia timmsi Martens et al., 2013

also showed significant variability in the shape of the Rpp (Martens et al. 2013).

Sadly, all specimens with elongated valves originated from old collections, on which no molecular

work could be done. In spite of several attempts, we have not been able to collect fresh material of the

elongated form. Future work will have to examine whether genetic data suggest that the two forms are

different species. For the moment we retain both forms as being conspecific.

Discussion

B. nimala lineage

The Bennelongia nimala lineage presently consists of seven species, which are united by the presence of

heavily calcified and heavily ornamented adult valves. Valve ornamentation consists of large pustules,

spines and short and stiff setae. It is in the B. nimala lineage that the heavily ornamented valves of the

juveniles of the last instars of most Bennelongia species, as described by De Deckker & Martens (2013),

are strongly retained in the adults. With regard to these features, the species in this lineage show the most

retarded heterochronic development. However, in other features, for example the strongly developed

anterior LV/RV overlap, the species show the most derived valve characters.

Two species of the B. nimala lineage ( B. pinderi sp. nov. and B. muggon sp. nov.) have an eyelet in the

anteroventral part of the RV, much as in the B. barangaroo lineage (see Martens et al. 2013), indicating

that these lineages are more closely related to one another than the valve morphology might indicate at

first glance. Species of the B. barangaroo lineage have valves that are much smoother and only weakly

pitted, although they can in some species be quite hirsute. As is usual in ostracods (Tsukagoshi 1988;

European Journal of Taxonomy 111: 1-36 (2015)

Martens et al. 2004), morphology of the different lineages of Bennelongia shows a mixture of different

heterochronic processes.

As in the other lineages of Bennelongia (Martens et al 2012, 2013; Shearn et al 2012), the antero-

ventral lapel on the RV proved to be a species-specific feature, allowing distinction between species

in the lineage and genus. In the B. nimala lineage, the lapel is most strongly developed in B. nimala ,

where it has the appearance of a toothed comb, and weakest in B. pinderi sp. nov., where it is a small

smooth ridge. Bennelongia muggon sp. nov. has a pronounced triangular lapel, reminiscent of some of

the species in the B. barangaroo lineage ( B. timmsi , B. scanloni).

The B. nimala lineage occurs in the NT (the nominal B. nimala ) as well as in the western part of WA,

between Pilbara and Perth. Additional information about the frequency of occurrence of the lineage

(though not individual species) is available from Pinder et al. (2010). No representatives of the lineage

have been found to date in Kimberley or south of Perth. One would expect the lineage to occur in

Kimberley to bridge the gap between the occurrence of B. nimala in the NT and B. koendersae sp. nov.

and B. shieli sp. nov. in the Pilbara, but thus far the only Bennelongia species known from Kimberley is

B. kimberleyensis within the B. australis lineage (Martens et al 2012).

In line with earlier findings in other Bennelongia lineages, most species of the B. nimala lineage have

restricted distributions, with B. nimala being known only from the NT, B. koendersae sp. nov. and B.

shieli sp. nov. being restricted to Pilbara and B. pinderi sp. nov. occurring only in the Murchison region.

In contrast, Bennelongia muggon sp. nov. is widespread, being recorded from Carnarvon to south of

Perth, a distance of 800 km. Bennelongia tirigie sp. nov. appears to be even more widespread, with most

records being from north of Carnarvon in the Gascoyne region and an outlying population occurring at

Crackers’ Swamp about 1000 km to the south. In addition to this surprising disjunct distribution, there

are also ecological differences with northern localities all being turbid water claypans, while Crackers’

Swamp is a clear-water body. Nevertheless, the specimens of all localities, including from Crackers’

Swamp, cluster tightly together in the COI-tree and have a very similar morphology, so conspecificity

cannot be doubted at this stage.

The results for genetic species boundaries of the B. nimala lineage match the morphological species

descriptions. With the possible exception of the enigmatic “species 15” from Pilbara, the morphology

of which remains unknown, no cryptic species were discovered with molecular methods in this lineage.

B. triangulata sp. nov.

Bennelongia triangulata sp. nov. is an aberrant species in the genus, as its triangular shape is different

from all other congeneric species. The morphology of the latest instar juveniles is even more different,

with a ventral side that is completely flattened (De Deckker & Martens 2013). Nevertheless, marginal

valve structures are typical of the genus and the soft parts are very similar to those of other Bennelongia

species, albeit slightly more elongated. Accordingly, we have decided to maintain this species within the

genus Bennelongia , despite its different habitus.

This species is common in the turbid clay pans of Murchison, Gascoyne and coastal southern Pilbara

with a latitudinal range of approximately 550 km. More information on the species range and habitat

preferences is available from the collecting records in Halse et al (2000) and Pinder et al (2010).

Bennelongia triangulata sp. nov. can occur in highly arched and more elongated forms (see Figs 11-

12) and both forms have been found in both warmer and colder seasons. In addition, male soft parts

(hemipenes, prehensile palps) are similar in both forms. Rpp (Fig. 13D, G) appear to have a different

morphology according to valve shape, but intermediate shapes of the distal segment have been found in

MARTENS K., HALSE S. & SCHON L, Bennelongia lineages in Western Australia

Table 3. Species presently described in Bennelongia , their lineage and their distribution (species in bold

are newly described here). Only certain distributions, based on type localities and documented range

extensions, are given here. * indicates the type species.

1 . * Bennelongia harpago De Deckker & McKenzie, 1981: QLD

2. Bennelongia tunta De Deckker, 1982: QLD

B. australis lineage

3. Bennelongia australis (Brady, 1886): S A (uncertain species)

4. Bennelongia bidgelangensis Martens et al., 2012: WA, Gascoyne

5. Bennelongia coondinerensis Martens et al., 2012: WA, Pilbara

6. Bennelongia cuensis Martens et al., 2012: WA, Yilgarn

7. Bennelongia gwelupensis Martens et al, 2012: WA, Perth, southwest coast

8. Bennelongia lata Martens et al, 2012: WA, Gascoyne-Murchinson region

B. barangaroo lineage

9. Bennelongia barangaroo De Deckker, 1981: WA

10. Bennelongia calei Martens et al, 2013: WA

11 . Bennelongia dedeckkeri Shearn et al, 2012: QLD, WA

12 . Bennelongia gnamma Martens et al, 2013: WA

13 . Bennelongia hirsuta Martens et al, 2013: WA

14. Bennelongia ivanae Martens et al, 2013: WA

15. Bennelongia mckenziei Shearn et al, 2012: QLD

16 . Bennelongia mcraeae Martens et al, 2013: WA

17 . Bennelongia scanloni Martens et al, 2013: WA

18 . Bennelongia timmsi Martens et al, 2013: WA

B. cygnus lineage

19. Bennelongia cygnus Martens et al, 2012: WA, Swan Valley

20. Bennelongia frumenta Martens et al, 2012: WA, Wheatbelt

B. nimala lineage

21. Bennelongia koendersae sp. nov.: WA

22. Bennelongia muggon sp. nov.: WA

23. Bennelongia nimala De Deckker, 1981: NT

24. Bennelongiapinderi sp. nov.: WA

25. Bennelongia regina Shearn et al, 2012: QLD

26. Bennelongia shieli sp. nov.: WA

27. Bennelongia tirigie sp. nov.: WA

B. pinpi lineage

28. Bennelongia kimberleyensis Martens et al, 2012: WA, Kimberley

29. Bennelongia pinpi De Deckker, 1981: QLD

30. Bennelongia strellyensis Martens et al, 2012: WA, Pilbara

B. triangnlata lineage

31. Bennelongia triangnlata sp. nov.: WA

some specimens, so that the observed differences may be artefacts of the dissected limb’s position on

a slide. Unfortunately, only older material of the elongated forms was available so that molecular data

ca nn ot be compared for the two morphological forms. Prom the network in Pig. 3 it can be seen that B.

triangidata sp. nov. is genetically highly variable, but there are no indications of the existence of cryptic

species, as all haplotypes remain connected.

With the six new species described here, the genus Bennelongia now comprises 31 nominal species,

divided over at least 7 lineages (Table 3), but several new species still await description.

European Journal of Taxonomy 111: 1-36 (2015)

Acknowledgments

The authors gratefully acknowledge the financial support by anABRS-grant (nr RF211-33: ‘Biodiversity

and Taxonomy of Ostracoda (Crustacea) from temporary water bodies of inland Western Australia’)

and of the Edith Cowan University Industry Collaboration grant. KM & IS acknowledge the Western

Australian Department of Parks and Wildlife (2006) and Bennelongia Pty Ltd (2008, 2009, 2010) for

financial support during several scientific stays in Perth, as well as the financial contribution of the FWO

Vlaanderen (Fund for Scientific Research, Flanders) in the form of travel grants in 2010 (V4.172.10N &

V4.173.10N), and the projects 1.5.172.09 (Krediet aan Navorsers) and G.0118.03N (projectonderzoek).

KM and IS also thank the people of Bennelongia Pty Ltd for continuous logistic support (lab space,

use of microscopes, etc.) and for unfailing companionship in the lab, and they thank their son Emrys

for invaluable help with sorting living specimens in the field as well as in the lab. David Cale, Adrian

Pinder (DPaW, Kensington), Andrew Storey (Wetland Research & Management, Perth), Brian Timms

(Newcastle) and Jane McRae (Bennelongia Pty Ltd) are acknowledged for collecting much of the

material described in the present paper.

Julien Cilis and Claudine Behen (RBINS, Brussels) provided technical assistance with the SEM

micrographs and with the line drawings, respectively. Kristiaan Hoedemakers (RBINS) produced the

SEM plates and supervised the entire production of the manuscript. Mike Scanlon (Bennelongia Pty

Ltd) produced the map. IS thanks Annette Koenders (Edith Cowan University, Perth) and Mike Johnson

(University of Western Australia, Perth) for providing laboratory space for the molecular part of this

research. Two anonymous referees are thanked for their valuable comments.

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Manuscript received: 21 July 2014

Manuscript accepted: 27 October 2014

Published on: 3 February 2015

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