European Journal of Taxonomy 8: 1-56

http://dx.doi.Org/lQ.5852/ejt.2012.8

ISSN 2118-9773

WWW, europeanj oumaloftaxonomv. eu

2012 • Koen Martens, Stuart Raise & Isa Sehon

This work is licensed under a Creative Commons Attribution 3,0 License.

Research article

Nine new species of Bennelongia De Deckker «& McKenzie, 1981

(Crustacea, Ostracoda) from Western Australia,

with the description of a new subfamily

Martens K., Raise S. feSchonl. 2012. Nine new species ofDe Deckker & McKenzie, 1981 (Crustacea,

Ostracoda) from Western Australia, with the description of a new subfamily. European Journal of Taxonomy 8:

1-56. http://dx.doi.Org/10.5852/eit.2012.8

Koen MARTENSi’2, Stuart HALSE^ and Isa SCHONi’4

^Royal Belgian Institute of Natural Seienees, Freshwater Biology, Vautierstraat 29,

B-1000 Brussels, Belgium (eorresponding author)

E-mail: [email protected]

^University of Ghent, Department of Biology, K.E. Eedeganekstraat 35, B-9000 Gent, Belgium

^Bennelongia Environmental Consultants, 5 Bishop Street, Jolimont WA 6014, Australia

"^University of Hasselt, Researeh Group Zoology, Agoralaan Building D,

B-3590 Diepenbeek, Belgium

Abstract. The genus Bennelongia De Deekker & MeKenzie, 1981 is most likely endemie to Australia and

New Zealand and, up to now, only two deseribed speeies in this genus had been reported from Western

Australia. Extensive sampling in Western Australia revealed a mueh higher speeifie diversity. Here,

we deseribe nine new speeies in three lineages, within the genus Bennelongia. B. cygnus sp. nov. and

B.frumentasp. nov. in the .6. cygnus linQagQ,B. gwelupensissp. no\.,B. coondinerensis sp. no\.,B. cuensis

sp. nov., B. lata sp. nov. and B. bidgelangensis sp. nov. in the B. australis lineage, and B. strellyensis

sp. nov. and B. kimberleyensis sp. nov. (from the Pilbara and Kimberley regions respeetively) in the

B. /?/>?/?/-lineage. For six of the nine speeies, we were also able to eonstruet moleeular phytogenies and to

test for eryptie diversity with two different methods based on the evolutionary genetie speeies eoneept,

namely Birky’s 4 x rule and the GYMC model These analyses support the speeifie nature of at least four

of the five new speeies in the B. australis lineage and of the two new speeies in the B. pinpi lineage. We

also deseribe Bennelongiinae n.subfam. to aeeommodate the genus. With the nine new speeies deseribed

here, the genus Bennelongia now eomprises 15 speeies, but several more await formal deseription.

Keywords. Evolution, Ostraeoda, eryptie speeies, biodiversity. Western Australia.

Introduction

Non-marine habitats eover only about 0.8% of the total surfaee and only 0.01% of the total water supply

of the planet Earth. Yet freshwater eomprises about 12% of all known animal speeies (c. 126,000 -

Eeveque et al. 2005; Balian et al. 2008), ineluding c. 25% of all known vertebrate speeies, and 40% of all

known fish speeies. Temporary habitats are strongly negleeted eompared to permanent lakes and rivers

when it eomes to researeh on biodiversity, ineluding eeosystem fimetioning, and espeeially regarding

taxie and moleeular diversity. Nevertheless, 60% of all red list speeies from England (lUCN 2012) oeeur

European Journal of Taxonomy 8 : 1-56 ( 2012 )

in temporary ponds and, when eorreeted for surfaee areas, temporary ponds have signifieantly higher

levels of biodiversity than most lakes and rivers. Gnammas (small, usually shallow roek pools) and

freshwater elay pans are often the only available surfaee water in the interior parts of (Western) Australia,

and thus eonstitute important habitats for aquatie biodiversity on this eontinent (Finder et al. 2000).

There are presently about 2000 subjeetive speeies and c. 200 genera of extant non-marine ostraeods

(Martens et al. 2008). But large parts of ostraeod diversity remain undeseribed today, and notable

hotspots for sueh undoeumented diversity are Airiean and Australian temporary habitats (Martens et al.

loc.cit.). Half of the known speeifie (1000 speeies) and generie (100 genera) diversity of non-marine

ostraeods belongs to one of the 13 known families, the Cyprididae. Many representatives of this group

oeeur in temporary water bodies, beeause several of their biologieal attributes (drought resistant eggs,

mixed / parthenogenetie reproduetion and ability to swim) pre-adapt them to form sueeessful radiations

in these habitats (Home & Martens 1998; Whatley 1992).

Australia eomprises less than 10% of all deseribed non-marine ostraeod speeies (176 against a total

of 1936), but the eontinent has one of the highest levels of endemieity (Martens et al. 2008). These

measures of Australian taxie diversity and endemism are undoubtedly underestimates for three main

reasons. Firstly, beeause mueh taxonomie work remains to be done aeross a great variety of animal and

plant groups, espeeially in the lesser known ones sueh as Ostraeoda. Sereening of extant eolleetions

reveals a large number of easily reeognisable, yet thus far undeseribed speeies and genera (e.g. Halse

2002). Seeondly, new eolleetions from thus far less explored areas and eeosystems will doubtlessly

yield undeseribed taxa. A ease in point is the high and previously eompletely unknown biodiversity of

Ostraeoda in ground waters in the Pilbara region (Karanovie 2007). Finally, several traditional ‘speeies’

are aetually turning out to be speeies eomplexes, eomprising sometimes dozens of eryptie speeies, sueh

as in the European (Bode et al. 2010) and the Australian (Koenders et al submitted) populations of the

morphospeeies Eucypris virens (Jurine, 1820).

The genus Bennelongia is interesting for reasons that go beyond a general knowledge of the biodiversity

of Australian inland waters. Home & Martens (1998) foreshadowed that Bennelongia might be the last

tme deseendant genus of the Mesozoie (and now extinet) lineage of Cypridea Bosquet, 1852, whieh

was also eharaeterised by the anterior beak-like expansion of the left valve and whieh was the dominant

ostraeod lineage in Cretaeeous non-marine waters when dinosaurs mled the Earth. This eould mean

that the tme Cyprididae, whieh now eomprise c. 50% of all living non-marine ostraeods (Martens et al.

2008), might be umelated to Cypridea and eould have deseended from a few Jurassie aneestral lineages

that already existed while Cypridea dominated non-marine freshwaters, mueh like the mammals eould

only expand their diversity after the dinosaurs had died out. Home & Colin (2005) meanwhile further

developed this idea and showed that Bennelongia is the one genus that somewhat resembles tme

Cypridea in the internal marginal valve morphology, while other eandidate genera sueh as Cypris O.F.

Muller, 1776 and Chlamydotheca Saussure, 1858 have eompletely different morphologies.

The aim of our revision of Bennelongia is firstly to identify the lineages within Bennelongia and deseribe

the overall diversity of speeies in these lineages and, seeondly, to analyse the phytogeny of these

speeies using morphologieal and moleeular data. We will use this information to test the hypothesis

that Bennelongia is the only tme deseendant genus of the Cypridea fauna with an antero-ventral beak.

The present paper is thus the first in a series of eontributions that sets out to test the ''Bennelongia as

surviving Cyprided" hypothesis.

Here, we deseribe nine new speeies of Bennelongia from Western Australia. Five of these are sibling

speeies within the B. australis lineage and for four of these their speeifie status is eonfirmed by moleeular

methods. In addition, we reveal the existenee of two sibling speeies within the B. pinpi lineage by

moleeular methods and deseribe these as new speeies. We do this beeause eloser morphologieal

examination revealed stable, miero-seale eharaeter states that allow their identifieation. Two further new

speeies are deseribed in a new lineage in the genus, the B. cygnus lineage.

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

Material and methods

Collections

Ostraeods were eolleeted from pans and lakes with a hand net with mesh size of 250 pm during field

trips between 2006 and 2011 (see below). Material for morphologieal analyses originated from both

these ‘new’ eolleetions and earlier eolleetions from all over Western Australia, mostly eolleeted by SH

and preserved in a eolleetion housed at the Department of Environment and Conservation (Woodvale,

Perth). The moleeular analyses worked only with newly eolleeted material, for whieh analyses were of

either living speeimens or speeimens sorted in the field and preserved in 100% ethanol. Consequently,

moleeular analyses were limited to six of the nine new speeies. Fieldtrips for reeent eolleetions were in

Gaseoyne/Murehinson (April 2006 and July 2011), Pilbara (April 2006), Kimberley (July 2010) and the

surroundings of Bunbury and Perth (July 2010). Eoeations of populations used for the present paper are

indieated on the map in Figure 1. Type material of the new speeies is deposited in the Western Australian

Museum, Perth, WA (WAM numbers) and in the Ostraeod Colleetion of the Royal Belgian Institute of

Natural Seienees, Brussels, Belgium (OC numbers).

Morphological analyses

Ostraeods were disseeted with valves stored dry in mieropalaeontologieal slides and soft parts in

glyeerine in sealed slides. Drawings of soft parts were made with a camera lucida with a eompound

Fig. 1. Map of Western Australia with loealities of Bennelongia

- speeies deseribed in the present paper.

European Journal of Taxonomy 8 : 1-56 ( 2012 )

microscope (Leica, DM 2500 at Beimelongia Environmental Consultants, Perth). Valves were illustrated

and measured using seanning eleetron mieroseopy (Philips XL30 SEM at RBINS, Brussels).

Molecular analysis

We used the Qiagen Blood and Tissue extraetion kit to extraet DNA from 53 individual ostraeods,

belonging to four speeies of the Bennelongia australis and two speeies of the B. pinpi, lineages

respeetively, following the manufaeturer’s protoeol. PCR amplifieation of part of the mitoehondrial

COl region was eondueted with universal primers (Folmer et al. 1994) in a T personal Thermobloek

(Biometra) with the following eonditions: 25 pi volumes of the HotStar Master Mix (Qiagen; 1.5 mM

MgC12, 0.1 pM primer, 200pM dNTP, Tris Cl, KCl, (NHQ^SO^, 1.25 U Taq) and 15 min at 95°C, 40

to 42 eyeles of 1 min at 95°C, 1 min at 42° to 44°C, 1 min at 72°C and a final extension step for 10

min at 72°C. Sueeess of PCR amplifieations was verified by agarose gel eleetrophoresis. Gels were

stained with Gelred and photographed. PCR produets were eleaned with the GFX™ PCR DNA and gel

band purifieation kit (GE Healtheare) aeeording to the manufaeturer’s protoeol and sequeneed in both

direetions on an ABl 3130X with the universal COl primers and the Big Dye kit (ABl) following the

manufaeturer’s protoeol.

As no fresh (living) material of the two speeies in the B. cygnus sp. nov. lineage eould be obtained, these

speeies are not represented in the moleeular phylogenetie tree and network.

Analyses of sequence data

Sequenee ehromatograms were visualised with Chromas (Teehnelysium. Pty Etd.). For eaeh individual,

the two strands were aligned with ClustalX (Earkin et al. 2007), subsequently eheeked by eye for

ambiguities, eorreeted and finally trimmed to obtain sequenees of equal lengths. We used BEAST

(Altsehul et al. 1990) to eonfirm the identity of the obtained sequenees in Genbank. The optimal model

of moleeular COl evolution was identified with jModeltest (Posada 2008) using 88 or 24 models and

AlCe analysis. For phylogenetie reeonstruetions, we used a redueed dataset applying Bayesian Inferenee

(Bl) in Mr Bayes 3.1.2 (Huelsenbeek & Ronquist 2001; with 2 million generations, sampling every lOO^*^

generation, a bum-in of 25% and the parameters identified by jModeltest for 24 different models) and the

Maximum-Eikelihood method in PhyME (Guindon & Gaseuel 2003; with 1000 bootstrap replieates and

the parameters of jModeltest for all 88 models). TCS 1.21 (Clement et al. 2000) was used to eonstmet

a parsimonious network at the 95% probability limit for eonneeting different sequenees (or haplotypes)

by using all sequenee data to illustrate genetie diversities and genetie relationships within and between

populations. Seleeted sequenees of all speeies have been submitted to Genbank (aeeession numbers

JQ432553-JQ432558).

Testing for cryptic diversity

We applied two different methods based on the evolutionary genetie speeies eoneept (Birky &

Barraelough 2009), namely the 4 x mle (Birky et al. 2010) and the GYMC model (Pons et al. 2006), to

identify speeies boundaries and umavel eryptie diversities. Similar approaehes have sueeessfiilly been

used in bdelloid rotifers (Fontaneo et al. 2007, 2009; Birky & Barraelough 2009; Birky et al. 2011),

marine ostraeods (Nunes Brandao et al. 2010) and a wide range of asexual prokaryotes (Birky et al.

2010 ).

For the 4 x mle, we used the obtained phytogenies to define as sister elades the elosest two neighbours,

with high statistieal support, whieh eould potentially be different speeies. Sequenee diversity within

and between these elades were estimated with MEGA 5.0 (Tamura et al. 2011) using the number of

differenees (p) or the TN model (allowing for multiple hits) and 1000 bootstrap replieates. Sequenee

diversities were further eorreeted for sample size and sequenee lengths, following the proeedure by

Birky et al. (2010). In order to fulfil the eriteria of the 4 x mle, the sequenee diversity between two

sister elades must be at least 4 to 4.3 times higher than within the two elades, depending on the number

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

of samples per elade (Birky et al. 2010). If the ratio is elose to 4, it is neeessary to also eompare the

sequenee diversities K within and between sister elades allowing for multiple hits. For applying the

GYMC model, we first tested whether the assumption of eloek-wise substitution rates was fulfilled by

eondueting likelihood ratio tests in TREE-PUZZEE 5.0 (Sehmidt et al. 2002). In PAUP 4.0 (Swofford

1998), the Maximum-Eikelihood approaeh was used with eloek assumption to eonstruet an ultrametrie

tree with all Bennelongia sequenees. This tree was imported into Ape (Paradis et al. 2004) and Geiger

(Harmon et al. 2008) in the R working environment, multifiireations were randomly resolved with Ape

and the GYMC model was applied to the latter ultrametrie tree.

Finally, uneonneeted networks from the parsimonious network analyses were eonsidered as possible,

speeies-like entities following the arguments of Hart & Sunday (2007).

Abbreviations used in text and figures

Antennula

Antenna

Carapaee

Height of valves

Eength of valves

Eeft valve

Right valve

Mandibula

Mxl

Maxillula

First thoraeopod

Epp

Eeft prehensile palp

Rpp

Right prehensile palp

Seeond thoraeopod

Third thoraeopod

Caudal Ramus

Eateral shield of hemipenis

—

Medial shield of hemipenis

Northern Territory

South Australia

QED

Queensland

Western Australia

Chaetotaxy of the limbs follows the model proposed by Broodbakker and 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

Results of molecular screening

We obtained COI sequenees for 37 speeimens from the Bennelongia australis lineage and 16 for the

B. pinpi lineage with a length of 522bp eaeh (for taxonomie definition of these lineages, see below). In

BEAST searehes, these sequenees gave low e values and elustered with COI sequenees from other non¬

marine ostraeods, eonfirming their genetie identity and exeluding possible eontamination. Amongst 88

models, jModeltest identified the TIMl+I+G model as the best fitting with the following parameters:

freqA= 0.3101, freqC = 0.1676, freqG = 0.1418, freqT = 0.3806; [AC] = 1.0000, [AG] = 50.4377, [AT]

European Journal of Taxonomy 8 : 1-56 ( 2012 )

= 11.2835, [CG] = 11.2835, [CT] = 81.8876, [GT] = 1.0000; p-inv = 0.5870 and gamma shape = 1.6460.

Among 24 models, the GTR+1 model was ehosen.

In the phylogenetie tree, the outgroups {Heterocypris spee. and Eucypris virens) eluster elearly outside

of the Bennelongia sequenees (Figure 2). We ean furthermore distinguish six well-supported elusters in

the phylogenetie tree (Figure 2) with high posterior probabilities and bootstrap values. The speeimens

belonging to the B. australis lineage form four groups, eonsisting of the speeies B. gwelupensis sp. nov.,

B. coondinerensis sp. nov., B. lata sp. nov. and .8. cuensis sp. nov., while the speeimens belonging to the

B. pinpi lineage form two distinetive groups, here deseribed as B. kimberleyensis sp. nov. from the

Kimberley and B. strellyensis sp. nov. from the Pilbara, respeetively. Phylogenetie relationships among

0.99

949

0.91

759

Heterocypris spec.

— Eucypris virens

1.0

937

\-145 B. gwelupensis

r 133 B. gwelupensis

l-| 316 B. gwelupensis

414 B. gwelupensis

304 B. gwelupensis

1.0

r 89 B. coondinerensis

983

1.0

970

91 B. coondinerensis

— 74 B. lata

73 B. lata

1.0

r 148 B. cuensis

964

^150 B. cuensis

0.95

888

1000

1.0

^ ^ p95 B. strelleyensis

99 B. strelleyensis

— 344 B. kimberleyensis

p 332 B. kimberleyensis

j-347 B. kimberleyensis

— 350 B. kimberleyensis

364 B. kimberleyensis

1000

0.05

Fig. 2. Phylogenetie tree eonstrueted with Baysian Inferenee of 53 COl sequenees of Bennelongia

and with Heterocypris spee. and Eucypris virens as outgroups. Numbers above and below nodes show

statistieal support for this partieular node. Numbers above nodes are Baysian posterior probabilities

(ranging from 0 to 1), below nodes bootstrap values of Maximum Likelihood (ML) analyses of 1000

replieates are provided. Both methods, B1 and ML, resulted in the same tree topology. The tree shows

four supported elusters in the B. australis lineage, eongruent with speeies deseribed in the present paper,

and two supported elusters for the sibling speeies in the B. pinpi lineage.

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

these six new Bennelongia speeies eannot be fully resolved with our COI data, as the deeper nodes are

polytomous and only weakly supported.

Besides phylogenetie trees, sequenees are also visualized in the form of genetie networks (Figure 3),

whieh are more suitable to display elose genetie relationships (Clement et al. 2000). Six distinet and

uneonneeted network groups ean be identified from the analysed Bennelongia individuals. Almost all

of the 11 individuals belonging to B. kimberleyensis sp. nov. eontain different sequenees or haplotypes

(with one exeeption, where one haplotype is shared between two individuals), while B. coondinerensis

6. cuens/^

148, 149, 151

B. cuensis

cuensis^

B. strellyen^

96 B. strellyen

strellyen^

Fig. 3. Parsimonious network, based on COI sequenees of Bennelongia. Squares represent aneestral

sequenees (or haplotypes), small eireles missing haplotypes. The size of squares and large eireles is

proportional to the number of individuals with the same sequenee in the analyses population. The

network was eonstrueted at the 95% probability limit, whieh usually ineludes up to 8 steps for eonneeting

different sequenees (or haplotypes). Thus, the two sequenees of B. lata being 10 mutational steps apart

needed to be eonneeted manually by lowering the eonneetion probability. The speeimens belonging to the

B. australis group form four groups, eonsisting of the speeies B. gwelupensis sp. nov., B. coondinerensis

sp. nov., B. lata sp. nov. and B. cuensis sp. nov., while the speeimens belonging to the B. pinpi group

forms two distinetive lineages, with B. kimberleyensis sp. nov. and B. strelleyensis sp. nov. from the

Kimberly and the Pilbara, respeetively.

European Journal of Taxonomy 8 : 1-56 ( 2012 )

sp. nov., B. cuensis sp. nov. and strelleyensis sp. nov. from Pilbarahave few, elosely relatedhaplotypes.

Bennelongia gwelupensis sp. nov. is eharaeterized by a single, eommon sequenee appearing in 15

different speeimens from different populations. Also three of the four remaining haplotypes from this

speeies are shared between individuals. Finally, B. lata sp. nov. eomprises the most distant sequenees, as

the two haplotypes are separated by 10 mutational steps, whieh is beyond the 95% probability limit for

eonneeting haplotypes with TCS.

For the 4 x rule, we used three different methods for phylogenetie reeonstruetions to identify elosest

neighbours within phylogenetie trees. The following three sister elades were found: Bennelongia

kimberleyensis sp. nov. - B. strelleyensis sp. nov., Bennelongia gwelupensis sp. nov. - B. cuensis sp. nov.

and B. lata sp. nov. - B. coondinerensis sp. nov. As required for the 4 x rule, we then eompared sequenee

diversity 0 within and between the representatives of eaeh of the three sister elades. In all eases (see

Table 1), sequenee diversity was higher than the required 4 x and varied between 4.8 x (5. gwelupensis

sp. nov. - B. cuensis sp. nov.) up to 16.4 x (5. lata sp. nov. - B. coondinerensis sp. nov.). Even when using

K as the measure of sequenee diversity (taking multiple hits into aeeount), the ratio was higher than 4 x

and varied between 4.7 and 55.6 (see Table 1). Thus, the eriteria of the 4 x rule are fulfilled for all three

sister elades and we identify a total of six different Bennelongia speeies. The eondueted likelihood ratio

tests eonfirm that COl substitution rates behave in a eloek-like fashion among all analysed Bennelongia

speeimens. Therefore, the ultrametrie COl tree ean be eonstrueted with a single, global moleeular eloek.

When applying the GYMC algorithm to this ultrametrie tree, the likelihood of the null model is 350.4,

whieh is signifieantly lower than the likelihood of the GYMC model (445.0) and the likelihood ratio

test is highly signifieant (p<0.001). The better fit of the GYMC model to our data implies that speeies-

like entities ean indeed be reeognized. In our ultrametrie tree (available from IS on request), the GYMC

model reeognizes 29 different sequenees (with no standard deviation) and five speeies (again, with no

standard deviation): the outgroup, Heterocypris spee., a seeond speeies eonsisting ofB. cuensis sp. nov.,

B. coondinerensis sp. nov. and B. lata sp. nov., a third speeies with B. gwelupensis sp. nov., a fourth with

B. strelleyensis sp. nov. from the Pilbara and a fifth with B. kimberleyensis sp. nov. fi*om the Kimberley.

In eonelusion, the GYMC model identifies fewer speeies than the 4 x rule and the parsimonious networks.

Table 1. Results of tests

using the 4 x

rule in six sibling speeies

in 2 lineages of Bennelongia.

Sister elades

max. 0/K

within elade

0/K between

elades

Ratio 0/K

nl, n2

B. kimberlyensis -

B. strellyensis

0.012/

0.012

0.140/

0.300

11.4/

25.2

12.3

B. gwelupensis -

0.039/

0.189/

4.9/

21.5

B. cuensis

0.003

0.189

55.6

B. lata -

B. coondinerensis

0.050/

0.020

0.810/

0.093

16.4/

4.7

2.4

0 = sequenee diversity; K = sequenee diversity taking multiple hits into aeeount. Nearest neighbours

or sister elades were defined from the COl tree eonstrueted with Bayesian Inferenee (see Figure 2),

Maximum Likelihood (not shown) and Neighbour-joining (not shown). In order to fulfil the eriteria of

the 4 X rule for speeies status, the ratio of the mean sequenee diversity within as eompared to between

the two nearest neighbours of one sister elade needs to be 4 or more, depending on the number of

speeimens per elade (Birky et al. 2010). In our analyses, these eriteria are elearly fulfilled for all three

sister elades and regardless whether sequenee diversity is estimated as 0 or K.

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

Taxonomic descriptions

Class Ostraeoda Eatreille, 1806

Subelass Podoeopa G. W. Muller, 1894

Order Podoeopida G.O. Sars, 1866

Suborder Cypridoeopina Baird, 1845

Superfamily Cypridoidea Baird, 1845

Family Cyprididae Baird, 1845

Subfamily Bennelongiinae subfam. nov.

Diagnosis

Adults with asymmetrieal valves with EV overlapping RV. EV antero-ventrally with a beak-like expansion

of the valve margin; anterior ealeified inner lamella with two ineomplete inn er lists, forming a suleus

and extending in the lip-like strueture. RV with short antero-ventral inner list, and antero-ventrally either

with a small, transparent Tape!’ (see below) or with the valve forming a bulbous expansion.

A1 and A2 with medium-long natatory setae; expopodite of A2 with two long and one short seta. Md-

palp with alpha-seta long and smooth, beta-seta stout and hirsute in distal half, gamma-seta relatively

slender and hirsute in distal half T2 (walking leg) with penultimate segment divided. T3 a eleaning leg.

Caudal ramus and attaehment slender. Hemipenis eonsisting of at least two asymmetrieal penal sheets.

Genus allocated:

Bennelongia De Deekker & MeKenzie, 1981

Remarks

Bennelongia was previously alloeated to the subfamily Cypridinae Baird, 1845 within the family

Cyprididae Baird, 1845, but sinee the regional revision of this subfamily by Martens (1990, 1992, 2007)

it is elear that Bennelongia laeks the additional post-labyrinthal eoils of the spermiduet in the hemipenis

that are eharaeteristie of Cypridinae s.s. In addition, the remarkable morphology of the anterior part of

the valves sets this genus aside from all other extant genera in the Cyprididae. Whether Bennelongiinae

subfam. nov. will eventually need to be transferred to the Cyprideidae Martin, 1940 will, hopefully, be

resolved by the time our revision of Bennelongia is eompleted. Meanwhile the new subfamily is lodged

in the Cyprididae.

Some general aspeets of the morphology of Alboa De Deekker, 1981 indieate that it might be elose

to Bennelongia and thus might also be a eandidate to be ineluded in Bennelongiinae subfam. nov.,

but initial moleeular sereening shows Alboa is eloser to Eucypris (Vavra, 1891) Daday, 1900 and

Heterocypris Claus, 1832 than to Bennelongia (results not shown).

Genus Bennelongia De Deekker & MeKenzie, 1981

Diagnosis (modified after De Deekker & MeKenzie 1981)

Adults with asymmetrieal valves, espeeially anteriorly. EV overlapping RV anteriorly, ventrally and

posteriorly, sometimes also dorsally.

EV antero-ventrally with a beak-like expansion of the valve margin; anterior ealeified inner lamella with

two ineomplete inner lists, a ventro-proximal one and a dorso-distal one, both extending to about the

European Journal of Taxonomy 8 : 1-56 ( 2012 )

middle of the valve and slightly or signifieantly overlapping eaeh other; ventro-proximal list protruding

and forming a suleus running along the proximal inner list and extending in the lip-like strueture.

RV with short antero-ventral inner list, and antero-ventrally either with a small, transparent flange (here

termed ‘lapel’) or with valve forming a bulbous expansion; RV set with tubereles along most of the

anterior, ventral and posterior valve margin.

Juveniles with eompletely different valve morphology, with symmetrieal valves, without beak-like

expansion on LV or lapel on RV and with external surfaee either pitted or retieulated or set with wart¬

like tubereles in most lineages in the genus.

A1 and A2 with medium-long natatory setae. Md-palp with alpha-seta long and smooth, beta-seta stout

and hirsute in distal half, gamma-seta relatively slender and hirsute in distal half Mxl-palp with distal

segment reetangular. T1 with prehensile palps in males strongly asymmetrieal, right palp with broad

terminal segment, left palp with siekle shaped distal segment. T2 with seta dl > dl, sometimes twiee

as long, penultimate segment of endopod (segment 3) divided. T3 a eleaning leg. Caudal ramus and

attaehment slender. Hemipenis eonsisting of at least two, asymmetrieal penal sheets.

Remarks

The extent of the differenee in morphology between juveniles and adults in most lineages of Bennelongia

is unusual in non-marine Ostraeoda. The differenee of the extent of this dimorphism between the

lineages, and its relevanee, will be diseussed elsewhere in a separate paper.

Differential diagnosis

Bennelongia is immediately distinguishable from all other eypridid genera by the peeuliar morphology

of the anterior parts of the valves, espeeially of the LV (see above).

Type species

Bennelongia harpago De Deekker & MeKenzie, 1981 (Queensland, Australia).

Other species allocated (only area of type locality given)

See Table 2.

Table 2. Speeies presently deseribed in Bennelongia and their distribution (speeies in bold are newly

deseribed here). Only eertain distributions, based on type loealities, are given here. * = type speeies

Bennelongia australis (Brady, 1886): SA

Bennelongia barangaroo De Deekker, 1981: WA

Bennelongia bidgelongensis sp. nov.: WA, Gaseoyne

Bennelongia coon diner ensis sp. nov.: WA, Pilbara

Bennelongia cuensis sp. nov.: WA, Yilgam

Bennelongia cygnus sp. nov.: WA, Swan Valley

Bennelongiafrumenta sp. nov.: WA, Wheatbelt

Bennelongiagwelupensis sp. nov.: WA, Perth, southwest eoast

"^Bennelongia harpago De Deekker & MeKenzie, 1981: QLD

Bennelongia kimberleyensis sp. nov.: WA, Kimberley

Bennelongia lata sp. nov.: WA, Gaseoine-Murehinson region

Bennelongia nimala De Deekker, 1981: NT

Bennelongia pinpiDe Deekker, 1981: QLD

Bennelongia strellyensis sp. nov.: WA, Pilbara

Bennelongia tunta De Deekker, 1982: QLD

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

Distribution

The genus is most likely endemie to Australia and New Zealand and ean be eonsidered as one of the

more typieal ostraeod groups of the Australasian region. De Deekker (1981a) suggested that Strandesia

flavescens Klie, 1932 and Strandesia feuerborni Klie, 1932, both from Indonesia (Sumatra, Java), might

also belong in this genus, but Savatenalinton & Martens (2010) and Martens & Savatenalinton (2011)

retained both speeies in their original genus, Strandesia Stuhlmann, 1888.

General valve morphology (Figure 4)

In order to allow aeeurate deseriptions of speeies in this genus, it is neeessary to unequivoeally establish

the homology of the different marginal struetures in both valves, as these have undergone remarkable

evolutionary ehanges (Figure 4A).

In the EV, the valve margin and two inner lists are of importanee. The valve margin shows an anteroventral

beak-like expansion. The anterior ealeified inn er lamella earries two inner lists, a proximal ventral and a

distal dorsal one. Together, these lists line a suleus, a depression in the ealeified inner lamella whieh further

expands in the beak-like expansion. The proximal inn er list ean be signifieantly elevated (Figure 4B).

In the RV, an outer list, the valve margin, a selvage and two inner lists form a eomplex strueture. The

outer list is invariably modified, eompletely or partially. In B. cygnus sp. nov., the middle part of the

outer list is still a list, the posterior part is modified into a row of tubereles and the anterior part in a

row of tubereles and a lapel (Figure 4C). In most (all?) speeies of the B. australis group (see below),

the entire outer list is modified into a row of tubereles and the antero-ventral lapel (Figure 4D). In the

speeies of the B. pinpi lineage, the entire outer list is modified into a row of tubereles, whieh stops

where in other speeies the lapel is formed and where in these speeies the bulbous expansion oeeurs

(Figure 4E). What looks to be the anterior valve margin in the RV is aetually a selvage, whieh ean be

modified in plaees, with swollen or fiange-like parts near the lapel. The aetual anterior valve margin

is strongly redueed, but ean in most speeies still be deteeted as a row of small tubereles set with setae

(Figure 4C-E). The ealeified inner lamella also has two ineomplete inner lists, here the ventral one is

distal and the dorsal one more proximal. Internal to the ventral inner list, the ealeified inner lamella

forms a shallow suleus, whieh matehes the deeper one in the EV. The posterior selvage is more easily

reeognizable as sueh.

Most Bennelongia speeies have some form of valve ornamentation, either a pitted surfaee or set with

tubereles and/or with long setae. This is speeies-dependent (though seemingly with similar patterns

within a lineage) and ean vary with water-ehemistry. But even speeies with almost smooth valve surfaees

(e.g. the B. cygnus lineage, see below) invariably have a field of external tubereles along the anterior

margin on the RV (see Figure 4F, G - not to be eonfused with the marginal row of tubereles in the RV,

whieh is mostly internally visible). This field ean be narrower or wider, aeeording to speeies, but is

always present. Its fimetion remains unknown.

Bennelongia cygnus - lineage

Remarks

This lineage so far eonsists of two speeies, here deseribed as new: B. cygnus sp. nov. and B.frumenta sp.

nov. They are eharaeterized by a relatively small earapaee (E = 1.6 mm) with triangular shape in lateral

view, a RV with a pointed lapel and by a hemipenis of the B. australis-typQ, i.e. with ms with oblique,

nearly straight distal margin and a boot-shaped ES, a siekle shape Epp and a Rpp with elongated seeond

segment.

European Journal of Taxonomy 8 : 1-56 ( 2012 )

Fig. 4. General morphology of Bennelongia valves. A. B. cygnus sp. nov., CpV (WAM.C49378).

B B. cuensis sp. nov., LV dorso-anterior view, tilted, showing elevated inner list (WAM.C49408).

C. B. cygnus sp. nov., $, RV, antero-ventral view, tilted, showing tear-like lapel and row of tubereles

(OC.3311). D. B. lata sp. nov., $, RV, antero-ventral view, tilted, showing broad lapel and row of

tubereles (speeimen lost). E. B. kimberleyensis sp. nov., $, RV, antero-ventral view, tilted, showing

absenee of lapel and expansion of valve (WAM.C49433 allotype). F. B. frumenta sp. nov. $, CpRL

view (WAM.C49383). G. B. gwelupensis sp. nov. $, CpRL view, detail of antero-ventral part (WAM.

C49395A). Not to seale.

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

Bennelongia cygnus sp. nov.

(Figs 5-9)

Bennelongia sp. - Davies & Christidis, 1997: 82, fig. 8.3.22.

Etymology

The type loeality of this new speeies is situated in the Swan Valley near Perth. We thus name this speeies

after the (blaek) swan, Cygnus in Eatin.

Diagnosis

Valves triangular in lateral view, weakly pitted, relatively narrow in dorsal view and with rostrum weakly

developed. EV with beak weakly developed. RV with lapel long and narrow, ventrally tear-like, almost

pointed. Hemipenis with Is extending beyond ms. Is distally rounded and with bluntly pointed apex, ms

dorsally with additional lobe-like expansion. Epp with distal segment siekle-shaped, but relatively short.

Rpp with sub-trapezoidal distal segment.

Measurements (all measurements in jim)

Male: RV: E = 1330; H = 933-954. EV: E = 1330-1410; H = 968-969.

Female: RV: E = 1450-1600; H = 1000-1070. EV: E = 1550-1750; H = 1030-1130. Cp: E = 1690-1790;

W= 1120-1130; H= 1100.

Type locality

Ellen Brook Nature Reserve, Swan Valley, near Perth (WA); approximate eoordinates: 31° 44’ 00”S

116° 01’ 00”E. Material used for the present deseription was eolleeted on 25 Sep. 1991, 3 Oet. 1997 and

2 Oet. 1998, all by SH.

Type material

Holotype

Male (WAM.C49370), with soft parts disseeted in a sealed slide and valves stored dry in a

mieropalaeontologieal slide.

Allotype

Female (WAM.C49371), with soft parts disseeted in a sealed slide, and valves stored dry in a

mieropalaeontologieal slide.

Paratypes

Five males disseeted and stored as the holotype (WAM.C49372-49375, OC.3310); RV + EV of one

female (OC.3311).

Several in toto speeimens in EtOH (voueher speeimens WAM.C49376).

Other material investigated

Perth Airport unnamed swamp, eolleeted by SH on 12 Sep. 1995, approximate eoordinates: 31° 56’

00”S 115°58’00”E

Goonaping Swamp, eolleeted by Adrian Pinder and Jane MeRae on 3 Oet. 1997, approximate

eoordinates: 32° 27’ 46”S 119° 25’ 1”E. (illustrated speeimens from this loeality: RV + EV of a female

(WAM.C49377); two female earapaees (WAM.C49378) all from OSTllC).

Christopher Brook, eolleeted by Adrian Pinder and Jane MeRae on 28 Oet. 1997, approximate

eoordinates: 32° 10’ 12”S 116° 47’ 39”E. (illustrated speeimens from this loeality: RV + EV of a female

(WAM.C49379) from OSTllA).

European Journal of Taxonomy 8 : 1-56 ( 2012 )

Cobertup Swamp, collected by Andrew Storey and SH on 19 Oct. 1996, approximate coordinates: 34°

27’00”S 116°49’00”E.

One Tree Hill, collected by SH and Adrian Finder on 11 Aug. 1999, approximate coordinates: 29° 35’

19”S 115°26’31”E.

Fig. 5. Bennelongia cygnus sp. nov. A. EVi ($, Ellen Brook, OC.3311). B. RVi ($, Ellen Brook, Idem).

C. EVi ((?, Ellen Brook, OC.3310). D. EVi (f, Ellen Brook, Idem). E. Cp dorsal ($, Goonaping Swamp,

WAM.C49378A). F. Cp ventral ($, Goonaping Swamp, Idem). G. CpRe ($, Goonaping, WAM.

C49378B). H. RVi (?, Goonaping, WAM.C49377). I. RVi (detail anteriorly, $, Ellen Brook, OC.3311).

J. RVi (detail anteriorly, f, Ellen Brook, OC.3310). Scales: A-H = 500 pm; 1-J = 200 pm.

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

Fig. 6. Bennelongia cygnus sp. nov. (all from holotype S, WAM.C49370). A. A2. B. T2. C. A2, detail of

distal ehaetotaxy. D. T3. E. Al. Seales: A, B, E = 232 pm; D = 184 pm; E = 37 pm.

European Journal of Taxonomy 8 : 1-56 ( 2012 )

Fig. 7. Bennelongia cygnus sp. nov. (all from holotype 3, WAM.C49370). A. Md-coxa. B. Md-palp.

C. Md-palp. Detail of P-seta and accompanying setae. D. Md-palp, showing a, p, and y-setae.

E. Mxl-palp, distal chaetotaxy. F. Mxl, partial chaetotaxy. G. Md-palp, chaetotaxy of distal segment.

Scales: A = 232 pm; B, D, F = 92 pm; C, E, G = 37 pm.

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

Fig. 8. Bennelongia cygnus sp. nov. (all from holotype 3, WAM.C49370). A. Hemipenis, showing

internal anatomy of labyrinth. B. Outline of other hemipenis. C. Eeft prehensile palp. D. Right Tl,

showing partial ehaetotaxy and right prehensile palp. E. Right Tl, showing distal ehaetotaxy of

endopodite. Seales: A-D = 92 pm; E = 37 pm.

European Journal of Taxonomy 8: 1-56 (2012)

Differential diagnosis

Bennelongia cygnus sp. nov. defines the B. cygnus lineage by its triangular shape, the simple type

of hemipensis and the pointed lapel. It ean be distinguished from the other speeies in this lineage,

B. frumenta sp. nov. (see below), by the absenee of a eavity in the selvage near the lapel, the presenee

of a dorsal lobe on the ms of the hemipenes, the short seeond segment of the Lpp and the broad base of

the seeond segment of the Rpp.

Fig. 9. Bennelongia cygnus sp. nov. f. A. Outline of hemipenis (WAM.C49375). B. Outline of seeond

hemipenis of same individual (WAM.C49375). C. Right prehensile palp (WAM.C49375). D. Left

prehensile palp (WAM.C49375). E. Caudal ramus (holotype S, WAM.C49370). F. Supporting braneh

of eaudal ramus (holotype S, WAM.C49370). Seales: A-D = 92 pm; E,F = 184 pm.

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

Additional descriptions

Male valves (Figure 5C, D) slightly smaller and more highly arehed than female valves (Figure 5A, B,

H), otherwise very similar in appearanee. Both valves triangular, with greatest height situated in the

middle of the valves, dorsal margins equally sloping to both anterior and posterior margins, ventral

margin almost straight. Valves weakly pitted and set with few, very short setae.

EV (Figure 5A, C) with posterior ealeified lamella narrow, inner list running along valve margin and

ereating a narrow suleus; the latter eontinuing towards the anterior side and widening up in between both

inner lists (see diagnosis of genus); antero-ventral beak weakly developed.

RV (Figure 5B, D, H) of similar shape as FV, smaller and slightly less high; posterior and ventral margin

set with tubereles, anterior ealeified lamella with short inner list; lapel relatively long, rather ventrally

situated and tear-shaped at its ventral edge (Figure 51, J) ; posterior side with long inner list (reaehing

almost up to dorsal margin) and with selvage elearly inwardly displaeed.

Width of earapaee in dorsal (Figure 5E) and ventral (Figure 5F) views more than half the length, greatest

width situated in the middle, FV overlapping RV on all sides, espeeially anteriorly and posteriorly

(Figure 5), anterior rostrum very weakly built, to almost absent.

A1 (Figure 6E) with all segments relatively short and narrow, natatory setae long, ehaetotaxy as typieal

of the family.

A2 (Figure 6A, C) with 5 natatory setae extending beyond tips of end elaws, basie ehaetotaxy and sexual

dimorphism in ehaetotaxy of penultimate segment as typieal of the family: in female with elaws G1-G3

and zl-3 setae; in males with G1 a short elaw, G2 a large elaw and G3 a seta of intermediate length; zl

and z2 large elaws, z3 a long seta.

Md eoxa (Figure 7A) relatively slender, without speeial features. Mandibular palp (Figure 7B-D, G)

with ehaetotaxy as typieal of the family, endelaws and gamma seta unusually slender.

Mxl (Figure 7E, F) with seeond palp segment reetangular, c. twiee as long as basal width, 3 elaws of

this segment relatively slender. Third endite with smooth ‘zahnborsten’. Sideways direeted bristles on

first endite of unequal length.

T1 (Figure 8C-E) with endite bearing 16, mostly hirsute, setae of unequal length. Females with endopod

a palp bearing 3 unequal apieal setae. Males with endepod developed in asymmetrieal prehensile palps.

Rpp (Figures 8D, 9C) with basal segment stout and only slightly longer than the largest width, subapieally

with two unequal sensory organs; seeond segment trapezioidal, with blunt dorsal and pointed ventral

edge. Fpp (Figures 8C, 9D) with first segment more slender, almost twiee as long as wide, distal segment

siekle-shaped but relatively short (F = < half F of first segment).

T2 (Figure 6B) a walking leg with seta dl > d2.

T3 (Figure 6D) a eleaning limb.

CR (Figure 9E) and its attaehment (Figure 9F) slender.

Hemipenes (Figures 8A, B; 9A, B) almost symmetrieal, with Is slender, with rounded dorsal margin and

bluntly pointed distal edge; ms eonsisting of two sub-lobes, one reetangular, the seeond one elongated

and ventrally direeted, with rounded edge.

Ecology and distribution

The speeies appears to be eomparatively widespread in freshwater bodies in south-western parts of

Western Australia. The speeies was found in elear or darkly eoloured water with eonduetivity ranging

from 80-3120 pS em'^and pH 6.8-8.5.

European Journal of Taxonomy 8: 1-56 (2012)

Remarks

One male speeimen (WAM.C49375 - Figure 9A, B) had aberrant morphology of hemipenes and Rpp.

The hemipenes were asymmetrieal, with one being typieal of the speeies, the other bearing an additional

thumb-like expansion of the Is. The same speeimen also had an additional distal thumb-like expansion

of the seeond segment of the Rpp. Rather than eonsidering this a different speeies, we deeided that it is

most likely a teratologieal speeimen.

Bennelongia frumenta sp. nov.

(Figs 10-11)

Etymology

Bennelongia frumenta sp. nov. is typieal of the south-western side of the Western Australian wheatbelt.

Wheat = frumenta in Latin.

Diagnosis

Valves triangular in lateral view, weakly pitted, width more than half the length in dorsal / ventral view

and with rostrum well-developed. LV with beak weakly developed. RV with lapel triangular, ventrally

pointed; selvage near lapel with a eavity, visible with SEM, but espeeially with transparent light.

Hemipenis with Is slightly extending beyond ms. Is distally rounded and with bluntly pointed apex; ms

ventrally and dorsally without lobe-like expansion. Rpp with trapezoidal distal segment; with nearly

straight distal margin and very narrow base. Lpp with distal segment siekle-shaped, elongated, almost

as long as first segment.

Measurements (all measurements in jim)

Male: RV: L = 1310; H = 907. LV: L = 1430; H = 933. Cp: L = 1320-1330; H = 841; W = 810-826.

Female: RV: L = 1320-1390; H = 851-905. LB: L = 1460-1500; H = 962. Cp: L = 1400-1520; H = 878;

W = 901-907.

Type locality

Kodjinup Melaleuea Swamp, 6 km N of Lake Muir in the Cranbrook shire (WA), eolleeted by Adrian

Finder and Jane MeRae on 2 Oet. 1998, approximate eoordinates; 34° 23’45”S 116°39’ 1”E(0STR11A,

SPS105).

Type material

Holotype

Female (WAM.C49380), with soft parts disseeted in a sealed slide and valves stored dry in a

mieropalaeontologieal slide.

Allotype

Male (WAM.C49381), with soft parts disseeted in a sealed slide, valves kept in EtOH (deealeified).

Paratypes

LV+RV of a female (OC.3312); one female earapaee (WAM.C49383); three male earapaees (WAM.

C49382A-C); soft parts of one male (WAM.C49384 - valves lost).

Several in toto speeimens in EtOH (voueher speeimens WAM.C49385).

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

Fig. 10. Bennelongiafrumenta sp. nov. A. EVi(holotype KodjinupMelaleueaSwamp, WAM.C49380).

B. RVi ($, Kodjinup Melaleuea Swamp, Idem). C. EVi {S, West Kulunilup Swamp, OC.3313). D. RVi

{S, West Kulunilup Swamp, Idem). E. Cp dorsal ($, Kodjinup Melaleuea Swamp, WAM.C49383).

F. Cp ventral ($, Kodjinup Melaleuea Swamp, WAM.C49388). G. Cp ventral {S, West Kulunilup

Swamp, WAM.C49382). H. Cp dorsal {S, Kodjinup Melaleuea Swamp, WAM.C49382). I. RVi

(detail anteriorly, holotype $, Kodjinup Melaleuea Swamp, WAM.C49380). J. RVi (detail anteriorly,

$, Kodjinup Melaleuea Swamp, Idem). K. CpRe ($, Kodjinup Melaleuea Swamp, WAM.C49383).

Seales: A-H, K = 500 pm; I-J = 200 pm.

European Journal of Taxonomy 8: 1-56 (2012)

Other material investigated

Wetland south east of Kodjinup Swamp, collected by Andrew Storey and SH on 21 Oct. 1997,

approximate coordinates: 34° 23’ 00”S 116° 40’ 00”E. Specimens from this locality: a dissected male

(WAM.C49386); LV+RV of a female (WAM.C49387).

Job^s Sump, collected by Jane McRae and Mick Smith on 10 Oct. 1997, approximate coordinates: 32°

21’ 15”S 117° 39’ 27”E (SPS060).

West Kulunilup Swamp, collected by Andrew Storey and SH on 22 Oct. 1997, approximate coordinates:

34° 20’ 00”S 116° 47’ 00”E. Specimens from this locality: several voucher specimens in EtOH

(OSTR13E), slide #38, a dissected male (OC.3313) and a female CP (WAM.C49388).

Unnamed claypan, Pingrup, collected by Adrian Pinder on 13 Sep. 2007, approximate coordinates: 33°

26’ 49”S 118° 30’ 41”E. Specimens from this locality: several voucher specimens in EtOH (OSTR13G,

ABP051); EV+RV of a female (OC.3314).

Lake Wheatfield, collected by David Calc on 26 Oct. 2005, approximate coordinates: 33° 48’ 46”S 121°

55’ 38”E (SPM005B).

Differential diagnosis

Bennelongia frumenta sp. nov. belongs to the B. cygnus lineage because of its triangular shape, the

simple type of hemipenis and the pointed lapel. It can be distinguished from the other species in this

lineage, B. cygnus sp. nov. (see above), by the presence of a cavity in the selvage near the lapel, the

absence of a dorsal lobe on the ms of the hemipenes, the elongated second segment of the Epp and the

narrow base of the second segment of the Rpp.

Fig. 11. Bennelongia frumenta sp. nov. (all allotype S, WAM.C49381, Kodjinup Melaleuca, type

locality). A. Outline of hemipenis. B. Right prehensile palp. C. Eeft prehensile palp, detail of distal part.

D. Eeft prehensile palp. E. T2 (walking leg). Scales: E = 232 pm; A-C= 92 pm; D = 37 pm.

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

Additional description

Valves triangular (Figure lOA-D), with greatest height situated in the middle; EV overlapping RV on all

sides (Figure lOK), dorsally only in the first half of the earapaee, ventral margin almost straight; width

of earapaee in dorsal and ventral views (Figure lOE-H) more than half the length, greatest width situated

in the middle, anterior rostrum strong; earapaee weakly pitted and set with few, very short setae.

Male valves slightly smaller and more highly arehed than female valves, otherwise very similar in

appearanee.

EV (Figure 10A, C) with posterior ealeified inner lamella narrow, inner list running along valve margin

and ereating a narrow suleus; this suleus eontinuing towards the anterior side and widening up in between

both anterior inner lists; antero-ventral beak weakly developed.

RV (Figure lOB, D) of similar shape as EV, smaller and slightly less high; posterior and ventral margin

set with tubereles, anterior ealeified inner lamella with short inner list, the latter forming a ‘eavity’ in

and on the selvage near the lapel; lapel relatively long, rather ventrally situated and ventrally pointed

(Figure 101, J); inner margin of posterior ealeified inner lamella with long inner list (reaehing almost up

to dorsal margin) and with selvage submarginal.

Most appendages as typieal of the genus and without speeial features.

Rpp (Figure 1 IB) with basal segment elongated, c. 1.5 x the eentral width; subapieally with two unequal

sensory organs; seeond segment trapezoidal, with blunt dorsal and pointed ventral edge, distal margin

nearly straight, base of segment very narrow.

Fpp (Figure IIC, D) with first segment more slender, more than twiee as long as wide, subapieally

with an elongated outgrowth, subapieally with a short sensory organ; distal segment siekle-shaped and

elongated, longer than dorsal margin of first segment.

Walking leg (Figure HE) stout and hirsute.

Hemipenes (Figure 11 A) symmetrieal; FS with rounded dorsal margin and bluntly pointed distal edge;

MS with oblique but straight distal margin, ventrally with a broadly rounded lobe, ventrally without

additional lobe.

Ecology and distribution

The speeies has thus far been found in a variety of seasonal and, less eommonly, permanent wetlands and

streams in the south-western Wheatbelt of Western Australia and in higher rainfall areas. The speeies has

been reeorded in water with eonduetivity 1550-9260 pS em'^ and pH 6.0-9.6.

Bennelongia australis - lineage

Remarks

Brady (1886) deseribed Chlamydotheca australis based on empty valves eolleeted by Prof R. Tate from

Penola (South Australia). The deseription was ineomplete and, as pointed out by De Deekker (1981a),

most likely also erroneous: although the FV (Brady 1886, plate 9 fig. 7) is undoubtedly of a speeies

of Bennelongia, the RV is not. De Deekker (loc.cit.) investigated the type material of Chlamydotheca

australis in the British Musuem (presently the Natural History Museum, Fondon) and found that the

slide eontained the illustrated Bennelongia FV as well as a seeond FV, most likely belonging to a speeies

of Heterocypris. It is likely that this is the valve illustrated by Brady (1886, plate 9 fig. 8). As it is the RV

that shows most speeifie features of Bennelongia speeies, the true identity of Bennelongia australis will

be impossible to establish based on the type material only. New material from Penola will be neeessary

European Journal of Taxonomy 8: 1-56 (2012)

and topotypes must be established to determine what speeies is the true B. australis. Below, we deseribe

5 sibling speeies in this lineage, whieh ean be identified based on shape of the valves, form of the lapel

on the RV, shape of the hemipenis and of the prehensile palps. The identity of four of these speeies was

eonfirmed by moleeular sereening (see above).

De Deekker (1981a) extensively redeseribed what he thought to be B. australis, based on material from

Western Australia (mainly from pools near Leonora and Cunderdin). Based on the illustrated lapels, it

is elear that this redeseription is based on at least two different speeies within the B. australis lineage,

neither of whieh are present in our eolleetions. Given the regional speeifieity of the speeies within this

lineage, it is unlikely that the true B. australis, deseribed from South Australia is amongst the speeies

used by De Deekker (1981a).

Davies & Christidis (1997) also illustrated a speeimen of what they labelled R australis, eolleeted from

lakes in and around Perth. We have eolleeted material from two of the same lakes and deseribe this speeies

below as B. gwelupensis sp. nov. The speeies figured by Davies & Christidis (1997) as Bennelongia sp.

was deseribed above as B. cygnus sp. nov. Karanovie (2008) illustrated speeimens from Pilbara as

B. australis s.l. These speeimens belong to B. strellyensis sp. nov. in the B. pinpi lineage (see below).

The B. australis lineage is eharaeterized by relatively large earapaees, LV with intermediately sized

beaks and RV with lapel. Most speeies deseribed here also have rounded dorsal margins and have

pronouneed anterior (and sometimes posterior) rostrum in dorsal view.

Bennelongia coondinerensis sp. nov.

(Figs 12, 13, 14A, B)

Etymology

The speeies is named after its type loeality, Coondiner Pool, Pilbara.

Diagnosis

Cp with pronouneed anterior rostrum, and weaker posterior rostrum. LV with pronouneed anteroventral

beak and rounded dorsal margin. Lapel on RV tear-shaped, dorsally sloping towards valve margin.

Hemipenis with MS with straight margin, ventrally widely produeed as a rounded lobe; Is in one

hemipene rounded, and distally bluntly pointed, in seeond hemipenis distally with thumb-like proeess;

tips of Is and ms well-separated. Is extending elearly beyond ms. Rpp with distal segment triangular,

rather broad. Lpp with proximal segment bearing reetangular apieal outgrowth; distal segment siekle-

shaped, tapering and rather elongated.

Measurements (all measurements in jim)

Male: RV: L = 1640, H = 971. LV: L = 1690, H = 911. Cp: L = 1690-1710; W = 911-913.

Female: RV: L = 1750, H = 1040. LV: L = 1860, H = 1120. Cp: L = 1810-1890; W = 1020-1080.

Type locality

Coondiner Pool, Pilbara, WA (sample K1ES14); approximate eoordinates: 22° 43’26”S 119° 39’ 23”E.

All material used for the present deseription eolleeted on 24 Apr. 2006 by the authors.

Type material

Holotype

Male (WAM.C49389), with soft parts disseeted in a sealed slide and valves stored dry in a miero-

palaeontologieal slide.

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

Allotype

Female (WAM.C49390), with soft parts disseeted in a sealed slide and valves stored dry in a miero-

palaeontologieal slide.

Paratypes

One male disseeted and stored as the holotype (OC.3315); three male earapaees (WAM.C49391A,

WAM.C49392), two female earapaees (WAM.C49391B-C). 2RV+1EV of a female (OC.3317A-C).

Several in toto speeimens in EtOH (WAM.C49393).

Other material investigated

Ethel Creek Clay pan, Pilbara, approximate eoordinates: 22° 49’ 32”S 120° 15’ 32”E. Colleeted by the

authors on 24 Apr. 2006.

Differential diagnosis

The speeies belongs to the B. australis group beeause of the generally large size (E > 1500 pm), the

presenee of a lapel on the RV and of a strong anterior rostrum in dorsal view. It ean be distinguished

from the other speeies in this lineage by the rounded dorsal margin of the EV, the tear-shaped lapel, the

broad seeond segment of the Rpp, the faet that tips of Is and ms of the hemipenes are well-separated

from eaeh other and that Is extends well beyond the ms.

The shape of the lapel somewhat resembles that of the seeond speeies redeseribed by De Deekker

(1981a) as B. australis (from a pool elose to Cunderdin), yet these latter speeimens have mueh more

highly arehed valves, while also the edge of the beak in the EV is less pointed than in B. coondinerensis

sp. nov.

Additional description

Valves in lateral view with rounded dorsal margin (Figure 12A-D), EV overlapping RV on all sides

(Figure 12K, E), greatest height anterior to the middle; in dorsal and ventral views (Figure 12E-H) with

greatest width in the middle of the earapaee; anterior rostrum well-developed, posterior side weakly

pointed, EV dorsally ridge-like; external surfaee weakly pitted and set with setae of intermediate length.

EV in inner view (Figure 12A, C) with rounded dorsal margin, greatest height situated in the middle;

antero-ventral beak-like expansion rather large; posterior part of ventral margin markedly sloping in

dorsal direetion.

RV (Figure 12D) in inner view with greatest height situated in front of the middle, dorsal margin almost

straight for about the middle third; posterior selvage elearly inwardly displaeed in the posterior half

of the valve, posterior inner list merging with posterior selvage at about halfway the length of the

latter; lapel tear-shaped (Figure 121, J), dorsally sloping towards the valve margin, ventrally abruptly

eurving towards it; antero-ventral inner list running to about halfway the lapel; selvage at height of lapel

expanded and slightly striate.

Most appendages as typieal of the genus and without speeial features.

Rpp (Figure 13 A) with first segment c. 1.5 times as long as wide, subapieally with one long but slender,

and one short sensory organ; seeond palp segment triangular, rather broad, with almost straight distal

margin; apieally with one small sensory organ.

Epp (Figure 13B) with first segment elongated, more than twiee as long as eentral width, subapieally

with one stout sensory organ, apieally with a reetangular outgrowth, bearing one small sensory organ;

seeond palp segment siekle-shaped and relatively elongated (E > V 2 E of first segment).

T2 (Figure 13C) a hirsute walking leg, with seta dl> seta d2.

European Journal of Taxonomy 8: 1-56 (2012)

Fig. 12. Bennelongia coondinerensis sp. nov., all from Coondiner Pool (Type locality). A. LVi (allotype

WAM.C49390). B. RVi (?, Idem). C. LVi (holotype S, WAM.C49389). D. RVi (S, Idem). E. Cp

dorsal (?, WAM.C49391C). F. Cp ventral (?, WAM.C49391B). G. Cp ventral {S, WAM.C49392B).

H. Cp dorsal {S, WAM.C49392A). I. RVi (detail anteriorly, holotype S, WAM.C49389). J. RVi (detail

anteriorly, allotype ?, WAM.C49390). K. CpRe {S, WAM.C49391A). L. CpRe (?, WAM.C49391C).

Scales: A-H, K-L = 1000 pm; J = 200 pm; 1 = 100 pm.

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

Hemipenes (Figure 14A, B) with tips of Is and ms well separated from one another, Is reaehing well

beyond tip of ms, distal part of ms produeed into an elongated lobe; distal part of Is bluntly pointed with

distal margin rounded in one hemipenis. Is almost reetangular with distal thumb-like proeess in the other.

Ecology and distribution

Bennelongia coondinerensis sp. nov. has thus far been found in two loealities in the Pilbara region.

Both elay pans had turbid waters (through suspended elay), with a thin layer of planktonie algae in the

top few eentimetres of turbid water. This ostraeod speeies has been reeorded in water with eonduetivity

104-197 pS em'^ and pH 7.7-10.2. This high pH is almost eertainly owing to photosynthetie aetivity of

the mentioned algae.

Fig. 13. Bennelongia coondinerensis sp. nov. S- A. Tl, showing right prehensile palp (holotype S,

WAM.C49389). B. Eeft prehensile palp (Idem). C. T2 (OC.3315). Seales: C = 232 pm; A,B = 92 pm.

European Journal of Taxonomy 8 : 1-56 ( 2012 )

Fig. 14. A-B. Bennelongia coondinerensis sp. nov. (holotype S, WAM.C49389);

C-F. Bennelongia gwelupensis sp. nov. (cJ, WAM.C49397). A. Hemipenis outline. B. Other hemipenis,

showing inner anatomy. C. Right prehensile palp. D. Hemipenis outline. E. Left prehensile palp.

F. Outline of seeond hemipenis, same individual. Seales: D, F = 232 pm; A, B, C, E = 92 pm.

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

Bennelongia gwelupensis sp. nov.

(Figs 14C-F, 15)

Bennelongia australis (Brady, 1886) - Davies & Christidis 1997: 81, figs 8.3.21a,b.

Etymology

Named after the type loeality. Fake Gwelup, Perth.

Diagnosis

Cp with pronouneed anterior rostrum, and weaker posterior rostrum. FV with pronouneed anteroventral

beak and rounded dorsal margin. Fapel on RV almost evenly rounded, dorsally sloping towards valve

margin. Hemipenis with MS with straight margin, ventrally widely produeed as a bluntly pointed lobe;

Is in both hemipenes evenly rounded, distally with blunt tip; tips of Is and ms well-separated. Is and ms

nearly equally long. Rpp with distal segment triangular, relatively elongated. Fpp with proximal segment

bearing reetangular apieal outgrowth with rounded tip; distal segment siekle-shaped, elongated, as long

as first segment, and with almost parallel margins, exeept for distal part.

Measurements (all measurements in jim)

Female: RV: F = 1710; H = 1040. FV: F = 1830; H = 1110. Cp: F = 1610-1820; H = 1080; W = 860-916.

Type locality

Fake Gwelup, Perth (WA); approximate eoordinates: 31° 52’ 37”S 115° 47’ 18”E. Material used for the

present deseription eolleeted on 23 Dee. 2008 and on 15 July 2010 by the authors.

Type material

Holotype

A female (WAM.C49394), with soft parts disseeted in glyeerine in a sealed slide, valves stored dry in a

mieropalaeontologieal slide.

Allotype

As the population eolleeted at the type loeality is fully parthenogenetie, no males oeeurred (at least at

the time of eolleetion) and no allotype ean be designated.

Paratypes

Four female earapaees (WAM.C49395A-D) and several in toto speeimens in EtOH (WAM.C49396).

Other material investigated

Big Carine Swampy Perth, eolleeted by the authors on 15 July 2010, approximate eoordinates: 31° 51’

08”S 115°47’04”E.

Several pools between Bunbury and Perth, eolleeted by the authors on 04 July 2010 and 31 July 2010.

The only male found thus far was in a dense population in a large diteh next to a parking lot beside

Forrest Highway, just northeast of the Greenlands Road interseetion, 32° 38’ 29”S 115° 48’ 34”E (sample

MR/10). Speeimens from this loeality: a male (WAM.C49397) disseeted and stored as the holotype; 3

female earapaees (WAM.C49398A-C). All other pools, as well as Fake Gwelup and Big Carine Swamp,

held only parthenogenetie populations.

Small unnamed streamlet^ inflowing in Leschenault Estuary, eolleeted by the authors on 04 July 2010

and 31.7.2010 This is the most southern loeality thus far of this speeies; approximate eoordinates: 33°

19’ 10.3”S 115° 41’ 17.3”E (sample MR/09). Speeimens from this loeality: RV+FV of one female and

a Cp of a female (OC.3316A-B);

European Journal of Taxonomy 8 : 1-56 ( 2012 )

Differential diagnosis

This species belongs to the B. australis group because of the generally large size (L > 1500 pm), the

presence of a lapel on the RV and of a strong anterior rostrum in dorsal view. It can be distinguished

from the other species in this lineage by the rounded dorsal margin of the LV, the rounded lapel, the

rather narrow second segment of the Rpp, the fact that tips of Is and ms of the hemipenes are well-

separated from each other and that Is and ms are almost equally long.

Additional description

Valves in lateral view with rounded dorsal margin (Figure 15A-B), LV overlapping RV on all sides

(Figure 15C, H), greatest height anterior to the middle; in dorsal and ventral view (Figure 15E,G) with

greatest width in the middle of the carapace; anterior rostrum well-developed (Figure 15F), posterior

side bluntly pointed, LV dorsally ridge-like; external surface weakly pitted and almost devoid of setae.

LV (Figure 15A) in inner view with rounded dorsal margin, greatest height situated anterior to the

middle; antero-ventral beak-like expansion rather large; ventral margin almost straight over most of its

length.

RV (Figure 15B) in inner view with greatest height situated well in front of the middle, dorsal margin

almost straight for about the middle third; posterior selvage submarginal, posterior inner list merging

with posterior selvage at about halfway the length of the latter; lapel (Figure 15G, 1, J) rounded, dorsally

sloping towards the valve margin, ventrally gently curving towards it; antero-ventral inner list running

to about halfway the lapel; selvage at height of lapel expanded, not striate.

Most appendages as typical of the genus and without special features.

Rpp (Figure 14C) with first segment c. 1.5 times as long as wide, subapically with one long but slender,

and one short sensory organ; second palp segment triangular, rather elongate, with sinuous distal margin;

apically with one small sensory organ.

Lpp (Figure 14E) with first segment elongated, more than twice as long as central width, subapically

with one long but slender sensory organ, slightly swollen in the middle; apically with a short and distally

rounded outgrowth, bearing one very small sensory organ; second palp segment nearly straight, sickle¬

shaped and relatively elongated (E = E of first segment); distal part of this segment straight and with

nearly parallel margins.

Hemipenes (Figure 14D, F) symmetrical, with tips of Is and ms well separated from one another. Is

and ms almost equally long, distal part of ms produced into an elongated lobe; distal part of Is bluntly

pointed with distal margin rounded.

Ecology and distribution

The species is common in Perth wetlands, where it occurs in large populations most of the year, and

along the south-west coast of Western Australia. Its most southern known locality is a small stream

entering Eeschenault Estuary in Bunbury, where it occurred together with Cyprideis australiensis and

Eucypris virens. The species has been recorded in water with conductivity 483-3880 pS cm'^ and pH 6.8.

Remarks

Davies & Christidis (1997) mention the presence of B. australis in 11 Perth wetlands. We have been able

to identify two of these populations (Take Gwelup, Big Carina swamp) as B. gwelupensis sp. nov., but

most likely all of the populations of these 11 wetlands belong to this species.

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

Fig. 15. Bennelongia gwelupsensis sp. nov. (all from Eake Gwelup, Perth - type loeality).

A. EVi (holotype WAM.C49394). B. RVi (Idem). C. Cp RE (WAM.C49395A). D. CpV (detail

anteriorly, WAM.C49395B). E. CpD (WAM.C49395C). F. Cp anteriorly (animal inverted) (WAM.

C49395D). G. CpV (WAM.C49395B), H. CpRE (detail anteriorly, WAM.C49395A). I. RVi (detail

anteriorly, holotype $, WAM.C49394). J. RVi (detail anteriorly, tilted. Idem).

Seales: A-C, E,G = 1000 pm; F = 500 pm; D,H-J = 200 pm.

European Journal of Taxonomy 8 : 1-56 ( 2012 )

Bennelongia lata sp. nov.

(Figs 16, 17A-C)

Etymology

The species has a broad lapel on the RV. Broad = lata in Latin.

Diagnosis

Cp with pronounced anterior rostrum, posteriorly bluntly pointed. LV with relatively weak anteroventral

beak and perfectly rounded dorsal margin. Lapel on RV broad, running parallel to selvage. Hemipenis

with ms with straight margin, ventrally widely produced as a broad lobe; Is in both hemipenes boot

shaped, distally with blunt tip; tips of Is and ms close together. Is slightly longer than ms. Rpp with distal

segment triangular, broad. Lpp with proximal segment without apical outgrowth; distal segment sickle¬

shaped, elongated, as long as first segment, and with almost parallel margins, except for distal part.

Measurements (all measurements in jim)

Male: RV: L = 1660-1700; H = 930-962. LV: L = 1770-1810; H = 995-1030. Cp: L = 1670-1800;

H = 881-995.

Female: Cp: L = 2000-2040; H = 1130; W = 1120-1170.

Type locality

Yandoo Billabong, Boolathana Station, Gascoyne, WA (sample S1EK3); approximate coordinates 24°

38’ 25”S 113° 55’ 20”E. All material was collected on 7 Apr. 2006 by the authors.

Type material

Holotype

Male (WAM.C49399), with soft parts dissected in a sealed slide, valves stored dry in a micropalaeonto-

logical slide.

Allotype

EV and RV of a female (WAM.C49400) stored dry in a micropalaeontological slide.

Paratypes

Two males dissected and stored as the holotype (WAM.C49401, OC.3318); two males and one female

(WAM.C49402A-C) carapaces in a micropalaeontological slide.

Several juvenile specimens: one cp of a male (A-1) (WAM.C49403); 2 Cp and 1 EV female (A-1) and 1

Cp female (A-3) (WAM.C49404A-D).

Several specimens in EtOH (WAM.C49405).

Other material investigated

Tirigie Claypan^ Boolathana Station^ Gascoyne, WA (sample SIEK4), approximate coordinates: 24°

38’ 29”S 113° 59’ 44”E. All material was collected on 7 Apr. 2006 by the authors.

Differential diagnosis

The species belongs to the B. australis group because of the generally large size (E > 1500 pm), the

presence of a lapel on the RV and of a strong anterior rostrum in dorsal view. It can be distinguished

from the other species in this lineage by the very rounded dorsal margin of both valves, the elongated

lapel running parallel to the selvage, the very broad second segment of the Rpp, the fact that tips of Is and

ms of the hemipenes are hardly separated from each other and that the Is is slightly longer than the ms.

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

Additional description

Valves in lateral view (Figure 16A-D) with rounded dorsal margin, EV overlapping RV on all sides

(Figure 16K), greatest height in the middle; in dorsal and ventral views (Figure 16E-H) with greatest

width in the middle of the earapaee; anterior rostrum well-developed, posterior side bluntly pointed,

EV dorsally less ridge-like than in the other speeies of this lineage; external surfaee weakly pitted and

sparsely set with short setae.

EV (Figure 16A, C) in inner view with rounded dorsal margin, greatest height situated in the middle;

antero-ventral beak-like expansion moderate; ventral margin almost straight over most of its length.

RV (Figure 16B, D) in inner view with greatest height situated slightly in front of the middle, dorsal

margin rounded; posterior selvage submarginal, posterior inner list merging with posterior selvage;

remnant of valve margin visible between lapel and valve margin; antero-ventral inner list running to

about halfway the lapel; selvage at height of lapel expanded, not striate, lapel (Figure 161, J) broad and

long, see speeifie name.

Most appendages as typieal of the genus and without speeial features.

Rpp (Figure 17C) with first segment c. 1.5 times as long as wide, subapieally with one long but slender,

and one short sensory organ; seeond palp segment triangular, broad, with straight distal margin; apieally

with one small sensory organ.

Fpp (Figure 17A) with first segment elongated, more than twiee as long as eentral width, subapieally

with one long, but slender sensory organ, slightly striate in the middle; apieally without distinet

outgrowth, but with one small sensory organ; seeond palp segment nearly straight, siekle-shaped and

relatively elongated (F = c. F of first segment); distal part of this segment straight and with nearly

parallel margins.

Hemipenes (Figure 17B) asymmetrieal, with tips of Is and ms situated elosely together. Is slightly longer

than ms, distal part of ms produeed into a blunt lobe; distal part of Is boot-shaped, bluntly pointed with

distal margin rounded in one hemipenis, with thumb-like expansion in the other.

Ecology and distribution

The speeies is known from two semi-permanent turbid, freshwater billabongs in the Gaseoyne area,

namely the type loeality Yandoo Billabong and Tirigie Claypan (see above). No further environmental

data are available.

Bennelongia cuensis sp. nov.

(Figs 17D-H, 18)

Etymology

The speeies is named after a small outbaek town. Cue, near its type loeality.

Diagnosis

Cp with pronouneed anterior rostrum, posteriorly bluntly pointed; in right lateral view, FV overlapping

RV on all sides, but with distinet overlap on the dorsal side. FV with relatively rounded anteroventral

beak and almost straight dorsal margin. Fapel on RV broad, running parallel to selvage. Hemipenis with

ms with straight and broad distal margin, ventrally produeed as a long lobe, dorsally also protruding

main peniferum as a rounded lobe; Is asymmetrieal, in both hemipenes with rounded distal margin and

distally with blunt tip, but in one hemipenis without and in the other hemipenis with additional (and

unusually elongated) thumb-like expansion; tips of Is and ms well-separated. Is signifieantly longer than

European Journal of Taxonomy 8 : 1-56 ( 2012 )

Fig. 16. Bennelongia lata sp. nov. (all from Yandoo Billabong, type locality). A. LVi (allotype WAM.

C49400). B. RVi (?, Idem). C. LVi {S, WAM.C49401). D. RVi ((?, Idem). E. Cp dorsal (?, specimen

lost). F. Cp ventral (?, WAM.C49402A). G. Cp ventral (S, WAM.C49402B). H. Cp dorsal (S, WAM.

C49402C). I. RVi (detail anteriorly, allotype WAM.C49400). J. RVi (detail anteriorly, tilted, allotype

$, WAM.C49400). K. Cp RL view ($, specimen lost). Scales: A-H, K = 1000 pm; 1, J = 200 pm.

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

Fig. 17. A-C. Bennelongia lata sp. nov. (holotype S, WAM.C49399); D-H. Bennelongia cuensis sp. nov.

(D-G = holotype S, WAM.C49406; H = OC.3319). A. Eeft prehensile palp and detail of apieal sensory

organs on first segment. B. outline of hemipenes (note that distal shield is folded in left hemipenis).

C. Right prehensile palp. D. Right prehensile palp. E. Outline of hemipenis. F. Outline of seeond

hemipenis, same individual. G. Eeft prehensile palp. H. Eeft prehensile palp. Seales: B, E, F = 232 pm,;

A, C, D, G, H = 92 pm; A’ = 37 pm.

European Journal of Taxonomy 8: 1-56 (2012)

ventral lobe of ms. Rpp with distal segment triangular, relatively elongated and with narrow base. Lpp

with proximal segment with small, triangular apieal outgrowth; distal segment siekle-shaped, elongated,

as long as first segment, and with almost parallel margins, exeept for distal part.

Measurements (all measurements in jim)

Male: RV: L = 1810; H = 1040. LV: L = 1930; H = 1130. Cp: L = 1820-1960. H = 1090. W = 1040-1080.

Female: Cp: L = 2170-2230; H = 1360; H = 1180-1300.

Type locality

Pool along Great Northern Highway, near Cue, WA, approximate eoordinates: 27° 15’ 20”S 117° 58’

58”E. All speeimens eolleeted by SH on 6 Sep. 2006.

Type material

Holotype

Male (WAM.C49406), with soft parts disseeted in a sealed slide, RV stored dry in a mieropalaeontologieal

slide (LV lost).

Allotype

A earapaee of a female (WAM.C49407) stored dry in a mieropalaeontologieal slide.

Paratypes

Three males (WAM.C49408-49409, OC.3319) disseeted and stored as the holotype; two male earapaees

(WAM.C49410).

Several speeimens in EtOH (WAM.C49411).

Other material investigated

The speeies is thus far known from its type loeality only.

Differential diagnosis

The speeies belongs to the B. australis group beeause of the generally large size (L > 1500 pm - it is

the largest speeies here deseribed), the presenee of a lapel on the RV and of a strong anterior rostrum

in dorsal view. It ean be distinguished from the other speeies in this lineage by the eonsiderable dorsal

overlap of the RV by the LV, the straight dorsal margin of the LV, the elongated lapel running parallel to

the selvage, the rather narrow seeond segment of the Rpp, the faet that tip of Is and ms of the hemipenes

are well-separated from eaeh other and that the Is extends mueh longer than the ventral lobe of the ms.

Bennelongia cuensis sp. nov. is similar to the previously deseribed speeies, B. lata sp. nov., espeeially

in the shape of the lapel on the RV, but ean be easily distinguished by the shape of the earapaee in right

lateral view (rounded with limited dorsal overlap in B. lata sp. nov., sub quadrate with eonsiderable

dorsal and ventral overlap in B. cuensis sp. nov.) and espeeially by the shape of the ms on the hemipenis

(ventrally bluntly pointed and short and without dorsal lobe in B. lata sp. nov., with elongated and

narrow lobes on both dorsal and ventral sides in B. cuensis sp. nov.)

Additional description

Valves (Figure 18A, B) in lateral view with straight dorsal margin, LV overlapping RV on all sides

(Figure 18G, H), but with eonsiderable dorsal overlap, greatest height situated in the middle; in dorsal

and ventral views (Figure 18C-F) with greatest width in the middle of the earapaee; anterior rostrum

well-developed, posterior side bluntly pointed, LV dorsally ridge-like; external surfaee weakly pitted

and set with short setae.

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

EV (Figure 18A) in inner view with rather straight dorsal margin, greatest height situated in the middle;

antero-ventral beak-like expansion moderate and rounded; ventral margin slightly sinuous in the middle.

RV (Figure 18B) in inner view with greatest height situated slightly in front of the middle, dorsal margin

rounded; posterior selvage sub marginal, posterior inner list merging with posterior selvage; remnant of

valve margin visible between lapel and valve margin; antero-ventral inner list running to about halfway

Fig. 18. Bennelongia cuensis sp. nov. (all from pan near Cue, type loeality). A. EVi {S, WAM.C49408).

B. RVi {S, Idem). C. Cp dorsal ($, speeimen lost). D. Cp ventral ($, same speeimen lost). E. Cp ventral

((?, WAM.C49410A). F. Cp dorsal {S, Idem). G. CpRE (allotype ?, WAM.C49407). H. CpRE ((?,

WAM.C49410B). I. RVi (detail anteriorly, S, WAM.C49408). J. RVi, detail anteriorly (tilted S, Idem).

K. RVi (detail anteriorly, tilted S, Idem). Seales: A-H = 1000 pm; I-K = 200 pm.

European Journal of Taxonomy 8: 1-56 (2012)

the lapel; selvage at height of lapel expanded, striate; lapel (Figure 181-K) broad, running parallel to

selvage, slightly striate.

Most appendages as typieal of the genus and without speeial features.

Rpp (Figure 17D) with first segment e 1.5 times as long as wide, subapieally with one long but slender,

and one short sensory organ; seeond palp segment elongate, with narrow base, apieally with one small

sensory organ.

Lpp (Figure 17G, H) with first segment elongated, more than twiee as long as eentral width, subapieally

with one long, stout sensory organ; apieally with small, rounded outgrowth, a small sensory organ at the

base of this lobe; seeond palp segment nearly straight, siekle-shaped and relatively elongated, as long as

first segment; distal part of this segment straight and with nearly parallel margins.

Hemipenes (Figure 17E, F) asymmetrieal, with tips of Is and ms well-separated. Is extending well

beyond ms, ventral part of ms produeed into an elongated, pointed lobe, dorsally also extending beyond

peniferum with a rounded lobe; distal part of Is bluntly pointed with distal margin rounded in one

hemipenis, with elongated thumb-like expansion in the other.

Ecology and distribution

The speeies is known from its type loeality only, a small seasonal elaypan with fresh turbid water. No

further environmental data are available.

Bennelongia bidgelangensis sp. nov.

(Figs 19, 20)

Etymology

The speeies is named after it type loeality, Bidgelang Pool on Callytharra Springs Station in the Gaseoyne,

WA.

Diagnosis

Cp with pronouneed anterior rostrum, posteriorly bluntly pointed; highly arehed in right lateral view,

LV overlapping RV on all sides. LV with relatively rounded anteroventral beak, rounded dorsal margin

and large postero-ventral fiange. RV in lateral view highly arehed, with highest point of dorsal margin at

about 1/3 of the total length. Lapel on RV broad, dorsally slightly sloping towards selvage. Hemipenis

with ms with straight distal margin, ventrally produeed as a pointed lobe, dorsally slightly protruding

main peniferum as a bluntly pointed lobe; Is asymmetrieal, in both hemipenes with straight distal margin

and distally with blunt tip, but in one hemipenis with additional elongated thumb-like expansion; tips

of Is and ms almost touehing. Is very slightly longer than ventral lobe of ms; hallway pemiferum with

additional small internal lobe. Rpp with distal segment elongated and with narrow base. Lpp with

proximal segment with broad, reetangular apieal outgrowth; distal segment siekle-shaped, elongated, as

long as first segment, and with almost parallel margins, exeept for distal part.

Measurements (all measurements in jim)

Male: RV: L = 1700-1780; H = 952-1030. LV: L = 1890; H = 1040-1110. Cp: L = 1920; W: 1030.

Female: RV: L = 1950; H = 1170. LV: L = 2130; H = 1260. Cp: L = 2030-2090; H = 1230; W = 1130-

1210 .

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

Type locality

Bidgelang Pool, Callytharra Springs Station, Gaseoyne (WA), approximate eoordinates: 24° 34’ 32”S

115° 36’ 01”E. All material eolleeted by SH on 25 Aug. 1994 (OSTR14D).

Type material

Holotype

Male (WAM.C49412), with soft parts disseeted in a sealed slide, valves stored dry in a miero-

palaeontologieal slide.

Allotype

EV and RV of a female (WAM.C49413) stored dry in a mieropalaeontologieal slide.

Paratypes

Three males disseeted and stored as the holotype (WAM.C49414, WAM.C49417, OC.3320); one male

(WAM.C49415A) and three females (WAM.C49415B-D) earapaees stored dry in a mieropalaeonto¬

logieal slide, RV+EV of one male (WAM.C49416).

Several speeimens in EtOH (WAM.C49418).

Other material investigated

The speeies is known from its type loeality only.

Differential diagnosis

The speeies belongs to the B. australis group beeause of the generally large size (E > 1500 pm), the

presenee of a lapel on the RV and of a strong anterior rostrum in dorsal view. It ean be distinguished

from the other speeies in this lineage by the highly arehed aspeet of earapaee and valves, the strongly

produeed postero-ventral flange on the EV (unique eharaeter for this speeies in the B. australis lineage),

the broad and elongated lapel on the RV, dorsally sloping towards the selvage, the narrow seeond segment

of the Rpp, and the faet that the tips of Is and ms of the hemipenes are almost touehing eaeh other.

Additional description

Valves in lateral view (Figure 19A-D) with middle part of dorsal margin almost straight, EV overlapping

RV on all sides (Figure 19K), greatest height situated posteriorly from the middle; in dorsal and ventral

views (Figure 19E-H) with greatest width in the middle of the earapaee; anterior rostrum well-developed,

posterior side bluntly pointed, EV dorsally ridge-like; external surfaee weakly pitted and set with short

setae.

EV (Figure 19A, C) in inner view with slightly rounded dorsal margin, greatest height situated in the

middle; antero-ventral beak-like expansion moderate and rounded; ventral margin slightly sinuous in the

middle; postero-ventral flange strongly produeed.

RV (Figure 19B, D) in inner view with greatest height situated at about 1/3 of total length, dorsal margin

rounded; posterior selvage not sub-marginal, well inwardly displaeed, posterior inner list merging with

posterior selvage below half of the height of the valve; remnant of valve margin visible between lapel and

valve margin; antero-ventral inner list running to about 1/4 of length of the lapel; selvage at height of lapel

expanded, striate; lapel broad, dorsally slightly sloping towards selvage, slightly striate (Figure 191, J).

Most appendages as typieal of the genus and without speeial features.

Rpp (Figure 20A, E) with first segment c. 1.5 times as long as wide, subapieally with one long and

basally stout, and one very short sensory organ; seeond palp segment elongate and with narrow base,

with straight distal margin; apieally with one small sensory organ.

European Journal of Taxonomy 8: 1-56 (2012)

Fig. 19. Bennelongia bidgelangensis sp. nov. (all from Bidgelang Pool, type locality). A. LVi (allotype

WAM.C49413). B. RVi (?, Idem). C. LVi (holotype WAM.C49412). D. RVi ((?, Idem). E. Cp

dorsal (?, WAM.C49415B). F. Cp ventral (?, WAM.C49415C). G. Cp ventral ((?, WAM.C49415A).

H. Cp dorsal {f. Idem). I. RVi (detail anteriorly, holotype WAM.C49412). J. RVi (detail anteriorly,

tilted, (5, Idem). K. Cp RL ($, WAM.C49415D). Scales: A-H, K = 500 pm; 1, J = 200 pm.

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

Epp (Figure 20C, D) with first segment elongated, length c. 1.5 x eentral width as eentral width,

subapieally with one long, stout sensory organ; apieally with broad, reetangular outgrowth, a small

sensory organ at the base of this lobe; seeond palp segment nearly straight, siekle-shaped and relatively

elongated, as long as first segment; distal part of this segment straight and with nearly parallel margins.

Hemipenes (Figure 20B) asymmetrieal, with tips of Is and ms almost touehing. Is and ms almost equally

long; ventral part of ms produeed into a bluntly pointed lobe, dorsally also extending beyond peniferum

with a rounded lobe; Is almost reetangular, distal margin almost straight in both hemipenes, ventrally

bluntly pointed in one hemipenis, with elongated thumb-like expansion in the other.

Fig. 20. Bennelongia bidgelangensis sp. nov. (A-C = holotype S, WAM.C49412; D-E = WAM.C49414).

A. Right prehensile palp. B. Outline of hemipenes. C. Eeft prehensile palp. D. Eeft prehensile palp.

E. Right prehensile palp. Seales: B = 232 pm; A, C-E = 92 pm.

European Journal of Taxonomy 8: 1-56 (2012)

Ecology and distribution

The species is known from its type locality only, a permanent river pool along the mostly dry Wooramel

River. Water conductivity was 106 pS cm'^ and pH 7.7.

Bennelongia pinpi - lineage

Remarks

Bennelongia pinpi De Deckker, 1981 was described from Pine Tree Creek Lagoon in Queensland and is

characterized by a very pronounced antero-ventral beak-like expansion in the LV, more so than in any

other species of Bennelongia, and also by the absence of an antero-ventral lapel on the RV and by the

presence of an antero-ventral expansion of the RV.

The following two species are allocated to this species group based on the latter two features, as the

beak on the LV is not particularly large in either of them. An additional character might be the heavily

sclerotized dorsal margins of the A1 in this species group (Figure 24F).

Bennelongia strellyensis sp. nov.

(Figs 21, 22)

Bennelongia australis s.l. (Brady, 1886)-Karanovic 2008: 282-286, figs 10-12.

Etymology

Named after the type locality, Strelley Station, Pilbara (WA).

Diagnosis

Cp with pronounced anterior rostrum, posteriorly bluntly pointed; rounded in right lateral view, LV

overlapping RV on all sides, external valve surface centrally almost completely smooth and devoid of

setae. LV with pronounced anteroventral beak, and highly arched, rounded dorsal margin. RV in lateral

view rather elongated, with highest point of dorsal margin at about 1/3 of the total length; lapel on RV

absent, but valves building a triangular bulbous expansion in the same location; anterior selvage with

fiange-like expansion and with additional inner submarginal list.

Hemipenis symmetrical, heavily sclerotized. ms forming a three-d im ensional structure, embracing Is

along three sides (proximal, ventral, distal). Is a large, slightly curved lobe, with almost parallel sides,

distally bluntly pointed in ventral direction. About halfway the hemipenal body, one additional lobe

present.

Rpp with distal segment short and rounded, with blunt tip bearing a sensory organ. Lpp with distal

segment sickle-shaped, rounded and short.

Measurements (all measurements in jim)

Male: RV: L = 1710-1900; H = 1090-1130. LV: L = 790-2010; H = 1080-1220. Cp: L = 1860-1880;

H= 1120; W= 1140-1150.

Female: RV: L= 1870; H= 1130. LV: L= 1980; H= 1200. Cp: L= 1880-2000; H= 1170; W: 1160-1230.

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

Type locality

Unnamed roadside pool on Strelley Station, Pilbara, WA (KIES3), approximate eoordinates: 20° 3E

24”S 119° 03’ 40”E. All material eolleeted on 22 Apr. 2006 by the authors.

Type material

Holotype

Male (WAM.C49419), with soft parts disseeted in a sealed slide, valves stored dry in a mieropalaeontolo-

gieal slide.

Allotype

EV and RV of a female (WAM.C49420) stored dry in a mieropalaeontologieal slide.

Paratypes

Three males (WAM.C49421, 49422, OC.3321) disseeted and stored as the holotype; three females

(WAM.C49423A-C) and two males (WAM.C49424A,B) earapaees stored dry in a mieropalaeontolo¬

gieal slide; RV+EV of a male (WAM.C49425).

Several juvenile speeimens: RV+EV of males (A-1) (WAM.C49426); 3 Cp of females (A-1) and 3 Cp of

females (A-3) (WAM.C49427AF).

Several speeimens in EtOH (WAM.C49428).

Other material investigated

Cooliarin Pooly Pilbara (WA), eolleeted by Adrian Finder on 10 Sep. 2004, approximate eoordinates:

20° 30’ 20”S 118° 37’ 20”E (PSW033). Speeimens from this loeality: RV+EV of two females (WAM.

C49429, 49330).

Errawallana Springy Pilbara (WA), eolleeted by Adrian Finder on 10 Sep. 2004, approximate

eoordinates: 21° 37’ 48”S 117° 46’ 14”E (PSW055). Speeimens from this loeality: a male, disseeted and

stored as the holotype (WAM.C49431).

Differential diagnosis

Bennelongia strellyensis sp. nov. belongs to the B. pinpi lineage, and therefore differs from all other

speeies in the genus not belonging to this lineage by a eombination of the following eharaeters: large

speeies, with well-developed anterior rostrum and bluntly pointed posterior end in dorsal view; RV

without antero-ventral lapel, but with bulbous expansion of the valve there; antero-ventral selvage with

an additional submarginal inner list. EV with antero-ventral beak well-developed. Hemipenis stout and

well-selerotized; ms a ventral, 3-dimensional strueture enveloping the Is. Rpp with short and rounded

seeond segment. Epp with seeond segment siekle-shaped and short.

Within this lineage, B. strellyensis sp. nov. differs from B. pinpi by the length of the antero-ventral

beak in the EV (longer in B. pinpi) and by the shape of the valves in lateral view (more highly arehed in

B. pinpi). The differenees between .6. strellyensis sp. nov. and .8. kimberleyensis sp. nov. will be outlined

below.

Additional description

Valves in lateral view (Figures 21A-D) with rounded dorsal margin, EV overlapping RV on all sides

(Figures 2IK, E) , greatest height in the middle; in dorsal and ventral views (Figures 21E-H) with

greatest width in the middle of the earapaee; anterior rostrum well-developed, posterior side bluntly

pointed, EV. Dorsally only anteriorly ridge-like; external valve surfaee eentrally almost eompletely

smooth and devoid of setae.

European Journal of Taxonomy 8: 1-56 (2012)

Fig. 21. Bennelongia strelleyensis sp. nov. (all from Strelley station, Pilbara, type locality).

A. LVi (allotype ?, WAM.C49420). B. RVi (?, Idem). C. LVi {S, WAM.C49421). D. RVi {S, Idem^

E. Cp dorsal (?, WAM.C49423A). F. Cp ventral (?, WAM.C49423B). G. Cp ventral (3, WAM.

C49424B). H. Cp dorsal {3, WAM.C49424A). I. RVi (detail anteriorly, allotype $, WAM.C49420).

J. RVi (detail anteriorly, $, Idem). K. CpRL ($, WAM.C49423C). L. CpRL ( 3 , WAM.C49424A).

Scales: A-H, K = 1000 pm; L = 500 pm; 1-J = 200 pm.

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

EV (Figure 21 A, C) in inner view with rounded dorsal margin, greatest height situated in front of the

middle; antero-ventral beak-like expansion large.

RV (Figure 2 IB, D) in inner view with greatest height situated well in front of the middle, dorsal margin

rounded; posterior selvage submarginal; lapel on RV absent, but valves building a triangular bulbous

expansion in the same loeation (Figure 211, J); anterior selvage with flange-like expansion and with

additional inner submarginal list dorsally merging with selvage.

Most appendages as typieal of the genus and without speeial features.

Rpp (Figures 22 B, 24D) with first segment c. twiee as long as eentral width, subapieally with one longer

(but less so than in speeies of the previous lineage), and one short sensory organ; seeond palp segment

triangular, broad, without elear ventro-apieal eomer and rounded distal margin; apieally with one small

sensory organ.

Fpp (Figure 22C, D) with first segment elongated, more than 2.5 x as long as eentral width, subapieally

with one large and stout sensory organ, apieally with rounded outgrowth, without sensory organ; seeond

palp segment siekle-shaped short, and asymmetrieally eurved over c. 180°, distally with a small sensory

organ.

Hemipenes (Figure 22A) symmetrieal. Is protruding well beyond ms, distally bluntly pointed towards

the ventral side, proximally with nearly parallel sides; ms aetually eonsisting of three lobes: distal part

of ms (msl) produeed into a three-dimensional lobe, embraeing the Is on proximal and distal sides (this

not visible in drawing, whieh is made of a flattened hemipenis in a slide) and with a small additional

dorsal lobe; ms2 with distal margin nearly straight, ms3 straight and ineomplete. (Remark: ms2 in some

populations weakly developed and almost invisible - see below).

Fig. 22. Bennelongia strellyensis sp. nov. (all from Strelley Station, Pilbara, type loeality; (J; A-C =

holotype S, WAM.C49419; D=OC.3321). A. Hemipenis outline. B. Right prehensile palp. C. Feft

prehensile palp. D. Feft prehensile palp. Seales: A = 232 pm; B-D = 92 pm.

European Journal of Taxonomy 8: 1-56 (2012)

Ecology and distribution

The species is common in moderately turbid claypans and roadside pools, and in clear water springs,

in the Pilbara where it may occur in high densities. The species has been recorded in waters with

conductivity 82-1391 pS cm ' and pH 7.5-8.9.

Remarks

The present species closely resembles B. kimberleyensis sp. nov. described below, as was the case of

the 5 sibling species in the previous B. australis lineage. The specific status of both new taxa was

nevertheless confirmed by molecular methods (see results above), where all the Pilbara and all the

Kimberley specimens clustered together in two distinct groups.

Bennelongia kimberleyensis sp. nov.

(Figs 23, 24)

Etymology

The species is named after the area in which it occurs, namely the Kimberley in the northern part of

Western Australia.

Diagnosis

Cp with pronounced anterior rostrum, posteriorly bluntly pointed; rounded in right lateral view, LV

overlapping RV on all sides, external valve surface centrally almost completely smooth and devoid of

setae. LV with pronounced anteroventral beak, and highly arched, rounded dorsal margin. RV in lateral

view rather elongated, with highest point of dorsal margin at about 1/3 of the total length; lapel on RV

absent, but valves building a triangular bulbous expansion in the same location, this expansion very

pronounced; anterior selvage with fiange-like expansion and with additional inner submarginal list.

Hemipenis symmetrical, heavily sclerotized. ms forming a three-dimensional structure, embracing Is along

three sides (proximal, ventral, distal). Is a large, slightly curved lobe, with almost parallel sides, distally bluntly

pointed in ventral direction. About hallway the hemipenal body, one additional lobe present.

Rpp with distal segment short and triangular, with a distinct dorso-apical comer, and with blunt tip

bearing a sensory organ. Lpp with distal segment sickle-shaped, very curved, and short.

Measurements (all measurements in jim)

Male: RV: L = 1998; H = 1159. LV: L = 2093; H = 1230. Cp: L = 2150-2177; H = 1289; W = 1298.

Female: RV: L = 2068; H = 1243. Cp: L = 2139-2230; H = 1300; W = 1223-1268.

Type locality

Taylors Lagoon, Broome, collected by the authors on 21 July 2011, approximate coordinates: 17° 49’

184” S 122°41’936”E (KlMB-01).

Type material

Holotype

Male (WAM.C49432), with soft parts dissected in a sealed slide, valves stored dry in a micro-

palaeontological slide.

Allotype

LV and RV of a female (WAM.C49433) stored dry in a micropalaeontological slide.

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

Paratypes

Two males (WAM.C49434, OC.3322) disseeted and stored as the holotype; three female (WAM.C49435)

and three male (WAM.C49436) earapaees stored dry in mieropalaeontologieal slides; RV+EV of a male

(WAM.C49437) stored dry in mieropalaeontologieal slides.

Several speeimens in EtOH (WAM.C49438).

Other material investigated (all from Kimberley)

Parry Lagoons, eolleeted by Andrew Storey on 21 Sep. 2000, approximate eoordinates: 15° 34’ 00”S

128° 17’ 00”E (sample NS-028). Speeimens from this loeality: two males, disseeted and stored as the

holotype (OC.3323; WAM.C49441); RV+EV of a male (WAM.C49440); soft parts of a male (WAM.

C49442); RV+EV of a female (WAM.C49439).

Small dam beside natural wetland along Great Northern Highway, eolleeted by the authors on 21 July

2011, approximate eoordinates: 17° 45’ 05”S 122° 56’ 14”E (KIMB-04).

Roadside pool E of Fitzroy River^ Willare, eolleeted by the authors on 21 July 2011, approximate

eoordinates: 17° 44’ 19”S 123° 38’30”E (KIMB-05).

Roadside pool containing cane grass, E of Eitzroy River along Great Northern Highway, eolleeted by

the authors on 21 July 2011, approximate eoordinates: 17° 44’ 22”S 123° 37’ 54”E (KIMB-06).

Myall Bore, Epool, eolleeted by the authors on 22 July 2011, approximate eoordinates: 17° 20’ 53”S

123° 39’59”E(KIMB-10).

Pool N side of Gibb Rd, eolleeted by the authors on 22 July 2011, approximate eoordinates: 17° 21 ’ 09”S

123° 44’ 11”E(KIMB-11).

(Identifieation of KIMB04,06,10 & 11 was eonfirmed using bareoding).

Differential diagnosis

The present speeies is slightly larger than B. strellyensis sp. nov. (up to 2.2 mm in Kimberley, up to 2.0

mm in Pilbara), and has a slightly larger antero-ventral beak in the EV, while also the antero-ventral

bulbous expansion in the RV is slightly larger.

The seeond segment of the Rpp in B. kimberleyensis sp. nov. is more triangular, i.e. with a elear dorso-

apieal eomer. The seeond segment of the Epp is less eurved, but this also depends partially on the

position of this segment relative to the first segment (the seeond segment ean hinge). The hemipenis has

a broader Is, while in some Kimberley populations, lobe ms2 is hardly visible, but still present (visible

at least on the ventral side at high magnifieation).

Additional description

Valves in lateral view (Figure 23A-D) with rounded dorsal margin, EV overlapping RV on all sides

(Figure 23K, E), greatest height in the middle; in dorsal and ventral views (Figure 21E-H) with greatest

width in the middle of the earapaee; anterior rostrum well-developed, posterior side bluntly pointed,

EV dorsally only anteriorly ridge-like; external valve surfaee eentrally almost eompletely smooth and

devoid of setae.

EV (Figure 23 A, C) in inner view with rounded dorsal margin, greatest height situated in front of the

middle; antero-ventral beak-like expansion large.

RV (Figure 23B, D) in inner view with greatest height situated well in front of the middle, dorsal margin

rounded; posterior selvage submarginal; lapel on RV absent, but valves building a triangular bulbous

expansion in the same loeation (Figure 231, J); anterior selvage with fiange-like expansion and with

additional inner submarginal list dorsally merging with selvage.

Most appendages as typieal of the genus and without speeial features.

European Journal of Taxonomy 8: 1-56 (2012)

Fig. 23. Bennelongia kimberleyensis sp. nov. (all from Taylors Lagoon, Kimberley, type loeality).

A. LVi (allotype ?, WAM.C49433). B. RVi (?, Idem). C. LVi WAM.C49437). D. RVi ((?, Idem^

E. Cp dorsal (?, WAM.C49435A). F. Cp ventral (?, WAM.C49435C). G. Cp ventral (S, WAM.

C49436A). H. Cp dorsal {S, WAM.C49436B). I. RVi (detail anteriorly, allotype $, WAM.C49433).

J. RVi (detail anteriorly, $, Idem). K. CpRe (?, WAM.C49435B). L. CpRe (S, WAM.C49436Cy

Seales: A-H, K-L = 500 pm; 1-J = 200 pm.

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

Fig. 24. A-C, E, F. Bennelongia kimberleyensis sp. nov. OC.3323, Parry’s Eagoon, Kimberley);

D. B. strellyensis sp. nov. {S, WAM.C49421, Strelley Station, Pilbara). A. Right prehensile palp. B. Eeft

prehensile palp. C. Hemipenis outline. E. T2. F. Al, without ehaetotaxy, showing strongly selerified

dorsal margins of segments. D. Right prehensile palp. Seales: C, E, F = 232 pm; A, B, D = 92 pm.

European Journal of Taxonomy 8: 1-56 (2012)

Rpp (Figure 24A) with first segment c. twiee as long as eentral width, subapieally with one longer

(but less so than in speeies of the previous lineage), and one short sensory organ; seeond palp segment

triangular, broad, with elear ventro-apieal eomer and almost straight distal margin; apieally with one

small sensory organ.

Lpp (Figure 24B) with first segment elongated, more than 2.5 x as long as eentral width, subapieally with

one large and stout sensory organ, apieally with rounded outgrowth, without sensory organ; seeond palp

segment siekle-shaped short, and asymmetrieally eurved over c. 180°, distally with a small sensory organ.

T2 (Figure 24E) a hirsute walking leg.

Hemipenes (Figure 24C) symmetrieal. Is protruding well beyond ms, distally bluntly pointed towards

the ventral side, proximally with nearly parallel sides; ms aetually eonsisting of three lobes: distal part of

ms (msl) produeed into a three-dimensional lobe, embraeing the Is on proximal and distal sides (this not

visible in drawing, whieh is made of a fiattened hemipenis in a slide) and with a small additional dorsal

lobe; ms2 with distal margin nearly straight, ms3 straight and ineomplete.

Ecology and distribution

This speeies oeeurred in a third of all our loealities (pools, elay pans) sampled during 2010 in the south¬

western Kimberley and was also found in the north-east near the mouth of the Ord River. The speeies

appears to be eommon through the Kimberley and oeeurs in fresh, elear or slightly turbid water. No

further environmental data are available.

Discussion

Morphological characters

Presence/absence and morphology of the antero-ventral lapel on the RV

The lapel on the RV is a newly identified eharaeter typieal of most speeies of Bennelongia (with

exeeption of speeies in the B. pinpi lineage). Its homology ean easily be dedueed from the SEM pietures

of the tilted RV: the row of marginal tubereles eontinues into the lapel, so both tubereles and lapel are the

remnants of an outer list (Figure 4C-E). The lapel forms part of the system of filter feeding, whieh also

ineludes the anterior suleus in the EV, and whieh is formed by both anterior inner lists on the ealeified

inner lamella and whieh runs into the antero-ventral beak-like expansion in the EV. This suleus is used

to direet the stream of partieles sueked in during filter feeding and the lapel is neeessary to elose the

entranee of the suleus when valves need to be elosed tightly (e.g. during a predatory attaek or during

periods of desieeation). In speeies without a lapel (see above the speeies of the B. pinpi lineage), the

antero-ventral bulbous expansion of the RV has the same fimetion.

Asymmetrical valves

The extreme anterior asymmetry of the valves is a result of the development of the filter feeding apparatus,

mainly in the EV. The extreme EV/RV overlap means that natatory setae of A1 and A2 extend out of the

valves during swimming in an angle of almost 90 degree to the direetion (plane) of swimming. In most

other swimming ostraeods these setae extend out of the valves in dorsal (Al) and ventral (A2) direetion,

whieh means that their movement is wit hin the same plane as the swimming direetion. Nevertheless,

speeies of Bennelongia are able to swim rapidly, and no speeial morphologieal adaptations to Al andA2

to the different orientation are visible.

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

Diversity and radiation of Bennelongia in (Western) Australia

The type speeies of Bennelongia, B. harpago De Deekker & MeKenzie, 1981, was deseribed from

several loealities in Queensland (De Deekker & MeKenzie 1981). Eater, De Deekker (1981a) also

deseribed B. barangaroo (from several loealities in Western Australia, South Australia, Queensland,

New South Wales and even New Zealand), B. nimala (Northern Territory) and B. pinpi (Queensland)

and redeseribed what he thought to be B. australis (Brady, 1886) using new material from Western

Australia (see above). De Deekker (1982) finally added R tunta from Queensland (Table 2).

Based on the present interpretation of new eharaeters, sueh as presenee/absenee and shape of the lapel

on the RV, the exeellent SEM illustrations of De Deekker (1981a) allow us to determine that at least for

his deseription of B. barangaroo and the redeseription of B. australis, material from several speeies has

been used. This inelusion of multiple speeies was also suggested by the presumed wide distribution of

these speeies as indieated by De Deekker (1981). Analysis of Western Australian material has indieated

that it is unlikely that any of the five speeies in this lineage deseribed here are the true B. australis and

that topotype material near Penola (SA) must be used for a redeseription of the true B. australis. Type

material from B. barangaroo De Deekker, 1981 from the type loeality itself (Take Buehanan, QED)

must be used to establish the identity of that speeies.

There are at present 15 nominal and deseribed speeies in the genus (ineluding the nine speeies

deseribed in the present paper), and if one ignores the presumed wide oeeurrenees of B. barangaroo and

B. australis based on the deseription of multiple speeies, all have elearly delimited regional distributions

(Table 2). Furthermore, it is elear that there are also many more speeies in this genus. The illustrations of

B. australis in De Deekker (1981a) eomprise at least two as yet undeseribed speeies in the B. australis

lineage and there is at least a further undeseribed speeies in the B. nimala lineage (De Deekker 1981a,

figure 9), possibly several more. Our own eolleetions from Western Australia eomprise at least another

ten new speeies in three different lineages within the genus Bennelongia, these will be deseribed

elsewhere. After this a key to the speeies will be provided. Whether or not the largest radiation has taken

plaee in Western Australia remains to be determined. Four very different speeies are already known from

Queensland and the Northern Territory remains virtually unsampled.

Congruence of genetic and morphological species definitions

We have used four different methodologieal approaehes to identify speeies entities from the obtained

mitoehondrial COI DNA sequenee data in speeies from the B. australis and the B. pinpi lineages:

phylogenetie tree eonstruetions, genetie networks, the 4 x rule and the GYMC algorithm. The first three

methods all identify six different speeies, whieh are all deseribed here as nominal speeies: B. gwelupensis

sp. nov., B. coordinerensis sp. nov., B. lata sp. nov., B. cuensis sp. nov., B. strelleyensis sp. nov. and

B. kimberleyensis sp. nov.

Only the GYMC algorithm deviates from other moleeular identifieations, with fewer speeies being

reeognized. Using the GYMC algorithm, three speeies, B. cuensis sp. nov., B. coordinerensis sp. nov. and

B. lata sp. nov., were identified as a phylogenetie speeies, while B. gweulpensis sp. nov., B. strelleyensis

sp. nov. and B. kimberleyensis sp. nov. were reeognized as different. Sinee all newly deseribed

Bennelongia speeies have distinetive morphologieal eharaeter states, it appears that the GYMC algorithm

underestimated the real biologieal diversity. This appears to be a general eharaeteristie of the method,

beeause similar diserepaneies have been deseribed in other faunal groups when results are, for example,

eompared to results of the 4 x rule (Birky et al. 2011; Birky & Barraelough 2009). The failure of the

GYMC algorithm to identify all speeies may be explained by rather reeent speeiation events that eannot

yet be diseriminated from the eoaleseent proeesses used in the GYMC algorithm.

European Journal of Taxonomy 8 : 1-56 ( 2012 )

Cryptic Diversity

The present results show that there is a large potential for eryptie diversity in Bennelongia. Within the

radiations of the B. australis and the B. pinpi lineages, new speeies initially identified with moleeular

teehniques eould also be reeognized morphologieally using new eharaeter and eharaeter states, sueh as

position and shape of the lapel on the RV. Thus, these speeies are not really eryptie and their diseovery just

aeeentuated the need for detailed morphologieal analyses to reveal small, but signifieant, morphologieal

differenees between speeies. However, without the genetie information there may have been insulfieient

use of detailed morphologieal eharaeters.

Martens et al. (2008) foreshadowed that eryptie diversity in non-marine Ostraeoda may be important

and might inerease the number of extant ostraeod speeies by one or even two orders of magnitude. Bode

et al. (2010) demonstrated this eonvineingly by identifying elose to 40 eryptie speeies with the Eucypris

Virens speeies eomplex in Europe. Some of these eryptie speeies have meanwhile also been found in

southern (De Deekker 1981b) and Western Australia (Koenders et al, in press).

As Martens & Savatenalinton (2011) have pointed out, from a speeies eonservation perspeetive, it is

important to undertake further researeh into the extent of eryptie diversity. Sometimes, as with the

speeies of Bennelongia deseribed, this diversity may not be truly eryptie but requires more detailed

morphologieal eharaeters than usually used to distinguish all speeies. Whether eryptie or only near-

eryptie, until these speeies are deseribed they will remain unproteeted by the proeesses of eonservation

planning beeause “what we do not know, we eannot proteef’.

Conclusions

The present eontribution is the first in a series of papers on Australian Bennelongia, with the ultimate aim

to test if Bennelongia is indeed a surviving remnant of the large Mesozoie Cypr/iiea-radiation. Here we

have demonstrated that the taxie diversity of Bennelongia in Western Australia is mueh higher than was

previously assumed. We have deseribed 9 new speeies in three different lineages within the genus and

foreshadow the presenee of at least another 10 new speeies in Bennelongia. We have also demonstrated

the usefulness of moleeular and population-genetie methods to strengthen the reeognition of potentially

eryptie speeies that might otherwise have remained undeteeted, but that ean indeed be reeognised using

new and miero-seale eharaeters and eharaeter states.

Acknowledgements

The authors gratefully aeknowledge the finaneial support by an ABRS-grant (m RF211-33 : ‘Biodiversity

and taxonomy of Ostraeoda (Crustaeea) from temporary water bodies of inland Western Australia’)

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

Australian Department of Environment and Conservation (2006) and Bennelongia Pty Etd (2008, 2009,

2010) for finaneial support during several seientifie stays in Perth, as well as the finaneial eontribution

of the FWO Vlaanderen (Fund for Seientifie Researeh, Flanders) in the form of travel grants in 2010

(V4.172.ION & V4.173.ION) and the projeets 1.5.172.09 (krediet aan navorsers) and G.0118.03N

(projeetonderzoek). KM and IS also thank the people of Bennelongia Pty Etd (Perth, WA) for eontinuous

logistie support (lab spaee, use of mieroseopes ete.) and for unfailing eompanionship in the lab and their

son Emrys for invaluable help with sorting living speeimens in the field as well as in the lab. Julien

Cilis and Claudine Behen (RBINSe, Brussels, Belgium) provided teehnieal assistanee with the SEM

mierographs and with the line drawings, respeetively. Jane MeRae (Bennelongia Pty Etd, Perth, WA)

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

eompiled the SEM plates and Mike Seanlon (Bennelongia Pty Etd, Perth, WA) produeed the map. IS

thanks Annette Koenders (Edith Cowan University, Perth) and Mike Johnson (University of Western

Australia, Perth) for providing laboratory spaee for the moleeular part of this researeh. Prof Patriek

De Deekker (ANU, Canberra, Australia) and Prof W. Birky (Univ of Arizona, USA) are thanked for

their valuable eomments as referees.

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