European Journal of Taxonomy 30: 1-10

http://dx.doi.org/10.5852/ejt.2012.30

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2012 • Vanhove M.P.M. et al..

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Research article

um: Isid: zoobank, org: pub:489A6B C2-795C-4945- A642-440D4F 92 A126

A new species of Gyrodactylus (Monogenea, Gyrodactylidae),

an ectoparasite from the endemic Iranocichla hormuzensis

(Teleostei, Cichlidae), the only Iranian cichlid

Maarten P.M. VANHOVE* '-’, Walter A. BOEGER’, Fidel MUTEREZl BUKINGA",

Filip A.M. VOLCKAERT^ Tine HUYSE^ & Antoine PARISELLE"

Laboratory of Biodiversity and Evolutionary Genomies, Department of Biology, KU Leuven,

Charles Deberiotstraat 32, B-3000 Leuven, Belgium.

Emails: [email protected] . [email protected].

[email protected]

^ Laboratorio de Eeologia Moleeular e Parasitologia Evolutiva, Departamento de Zoologia, Setor de

Cieneias Biologieas, Universidade Federal do Parana, Caixa Postal 19073, CEP 81531-980, Curitiba,

PR, Brazil. Email: [email protected]

^ Seetion de Parasitologie, Departement de Biologie, Centre de Reeherehe en Hydrobiologie, B.P 73,

Uvira, Demoeratie Republie Congo, via B.P. 254, Bujumbura, Burundi.

Email: [email protected]

6ISE-M, IJMR5554 CNRS, IJR226 IRD, Universite Montpellier II - CC 063, F-34095 Montpellier

Cedex 5, Franee. Email: [email protected]

^ lehthyology Unit, Afriean Zoology Department, Royal Museum for Central Afriea,

Leuvensesteenweg 13, B-3080 Tervuren, Belgium.

* Corresponding author. Email: [email protected]

^ um:lsid:zoobank.org:author:F40A361D-F45C-4C91-86AF-115962126120

^ um:lsid:zoobank.org:author:318E38BE-6AB8-4D77-BFA9-36F5E8A546E0

^ um:lsid:zoobank.org:author:B521076B-7ED5-4532-8EEA-CB33BF4019F7

^ um:lsid:zoobarLk.org: author: AC726421-5168-4787-A939-0CCBCBEE5114

^ um:lsid:zoobank.org:author:72A8D565-7DCB-4103-A4B9-84248E88FF4B

^ um:lsid:zoobank.org:author:DE920C99-1505-4BlF-A523-E375BlDC6201

Abstract. Iranocichla hormuzensis oeeupies a biogeographieally peculiar position. This endemie of

southern Iran is the only Iranian eiehlid. While it is phylogenetieally related to Afriean oreoehromine

members of the eiehlid family, it remains unclear how it has dispersed into its current range. It is one

of the many lasting enigmas of cichlid biogeography. Monogenean fish parasites may provide useful

additional information in such cases. Therefore, I. hormuzensis was examined for these fiatworms. A

gyrodactylid parasite is reported and compared to congeners from the Palearctic and from cichlids. In

this way, we verily whether it shows affinities to parasites from fishes that are either biogeographieally

or phylogenetieally close to Iranocichla hormuzensis. The species is new to science and is described as

Gyrodactylus jalalii sp. nov. This is the first description of a parasite infecting /. hormuzensis. Because

of the fixation method or age of the material, DNA could not be isolated. Due to the lack of genetic data,

no conclusions can be drawn on its phylogenetic positioning. Indeed, Gyrodactylus phytogeny cannot be

European Journal of Taxonomy 30 : 1-10 ( 2012 )

inferred from morphologieal eharaeteristies alone. Moreover, the eongeners phenotypieally reminiseent

of the new speeies belong to a Gyrodactylus elade whieh is highly diverse in geographie range and host

ehoiee. Henee, there is no evidenee l inkin g the new speeies to an exelusively Afriean or eiehlid-bound

Gyrodactylus lineage.

Keywords. Gyrodactylus jalalii sp. nov., Oreoehromini, Platyhelminthes, Pereiformes, speeies

deseription.

Vanhove M.P.M., Boeger W.A., Muterezi Bukinga F., Volckaert F.A.M., Huyse T. & Pariselle A. 2012. A new

species of Gyrodactylus (Monogenea, Gyrodactylidae), an ectoparasite from the endemic Iranocichla hormuzensis

(Teleostei, Cichlidae), the only Iranian cichlid. European Journal of Taxonomy 29: 1-10. http://dx.doi.org/10.5852/

eit.2012.30

Introduction

The faseination and seientifie interest that eiehlids raise in many seientists and hobbyists is well

reviewed by Barlow (2000). This author aptly shows the importanee of eiehlid fishes in a wide range of

fields in evolutionary biology, from eeology to behavior and from developmental biology to speeiation.

Numbering about 1350 deseribed speeies, with hundreds remaining to be diseovered (Nelson 2006),

these “model” fishes are the most speeies-rieh vertebrate family worldwide (Koeher 2004). Counting

only deseribed speeies, Ciehlidae is the largest non-ostaryophysan freshwater fish family (Kullander

2003) . Its speeies riehness reaehes its peak in the more than 900 Afriean representatives. Some relatives

to Afriean eiehlids oeeur in the Middle East (belonging to Astatotilapia Pellegrin, 1904; Oreochromis

Gunther, 1889; Sarotherodon Riippell, 1852; Tilapia Smith, 1840 and Tristramella Trewavas, 1942)

(Werner & Mokady 2004) and one in Iran. The latter is Iranocichla hormuzensis Goad, 1982, the only

representative of the monotypie Iranocichla Coad, 1982. The speeies was first reported by Behnke

(1975) and Saadati (1977) before being formally deseribed (Coad 1982). This southern Iranian endemie

eiehlid is found in mostly saline riverine systems draining into the Strait of Hormuz (Persian Gulf). The

question rises how it arrived in this biogeographieally isolated range.

Most eiehlid speeies in the Middle East have eongeners among the Afriean “tilapiine” {Oreochromis,

Tilapia and Sarotherodon'. Sehwarzer et al. 2009) or haploehromine {Astatotilapia. Werner & Mokady

2004) eiehlids. Tristramella and Iranocichla, however, are endemie genera. Phylogenetieally, they are

also plaeed in Oreoehromini, one of the tribes into whieh “tilapiine” eiehlids were split by Sehwarzer

et al. (2009). Thus, they are elosely affiliated to Afriean eiehlids. Middle-Eastern eiehlids at present

only oeeur in the Eevant and these Iranian systems. This faet might indieate a reliet-type eonsequenee

of a reeent (Pleistoeene to Reeent) aridifieation event and deerease in temperature (overview in Coad

1982). Furthermore, the Persian Gulf was a river valley between 90,000 and 10,000 years ago. See

also Arndt et al. (2003) for a diseussion on how the low level of the Mediterranean during Pleistoeene

episodes is suggested to have allowed freshwater dispersal from the Nile to the Eevant. Although those

eonditions would have allowed dispersal of eiehlids, it is also possible that the marine environment

would not represent a barrier to the spread of these animals, anyway (see referenees below). Indeed,

oeeupying mostly saline streams shows that I. hormuzensis is salt-tolerant (Coad 1982). Moreover,

several “tilapiines” are known to disperse between rivers through braekish eoastal waters (Nelson 2006).

Conversely, Murray (2001) favors marine dispersal through the Tethys Sea/Indian Oeean to eoastal

dispersal in view of the absenee of eiehlids along the rest of the region’s eoastlines.

Either way, both freshwater and marine dispersal pathways eould explain the presenee of this eiehlid in

waters draining into the Strait of Hormuz. Be it on a loeal or a global seale, the history behind eurrent

eiehlid distribution aeross eontinents has not been unequivoeally umaveled. The roles of intereontinental

VANHOVE M.P..M. et al., Gyrodactylus from the GoMid Iranocichla hormuzensis

dispersal, on the one hand, and of viearianee (following the break-up of Gondwana) and intra-eontinental

freshwater dispersal, on the other hand, are still highly debated. A potential souree of information that

remains largely unexplored is the (monogenean) parasite fauna of eiehlid speeies (reviewed in Pariselle et

a/. 2011). Many parasites have a elose assoeiation with their host and a shorter generation time than their

host, henee providing an alternative perspeetive on host evolution. Eetoparasitie monogenean flatworms

often do not tolerate substantial ehanges in salinity, as indieated by their distribution and phylogenetie

patterns worldwide (although there are exeeptions, e.g., this is less the ease in Gyrodaetylidae, eertainly

when infeeting euryhaline hosts, see below). One might therefore hypothesize that eertain freshwater

monogeneans rarely survive dispersal through barriers represented by marine environments. Henee,

the phylogenetie relationships within a lineage of monogenean eetoparasites refleet both eontinental

patterns and host phytogeny. Freshwater flsh hosts most likely retain their monogenean assemblage when

dispersing through eontinental surfaee waters. Conversely, it was suggested that most freshwater Ashes

might lose their autoehtonous parasites during marine dispersal events. This would allow subsequent

reeolonisation by “loeaf ’ parasite speeies onee arriving in freshwater at the other end of a marine barrier

(Pariselle et al. 2011).

Here, a Gyrodactylus speeies parasitizing I. hormuzensis is deseribed. We analyze its affinity to

eongeners infeeting other eiehlids and to speeies known from Ashes of the same region. Beeause of the

ability to switeh hosts, gyrodaetylids are expeeted to mainly mirror eontinental affinities rather than

the relationships between its host speeies (Boeger et al. 2003), depending of eourse on the time-seale

and the dispersal routes used. The result might henee shed light on the pathway eiehlids used to reaeh

southern Iran.

Material and Methods

Eight host Ashes (Axed and stored in ethanol or in formaldehyde) were retrieved from eolleetions of the

Royal Museum for Central Afriea (Tervuren, Belgium) (MRAC B2-28-P-1-8). Branehial arehes, body,

flns and reeipient were inspeeted for parasites under a Wild M8 stereomieroseope. Monogeneans were

removed with a disseetion needle. They were treated with 10% sodium dodeeyl sulphate for a eouple

of minutes, before being Axed using ammonium pierate-glyeerine (Malmberg 1957) and mounted on a

slide under a eover-slip. Measurements were adapted from Shinn et al. (2004) using a Zeiss Axio Imager

ZI mieroseope at a magnifleation of 100 x (oil immersion, 10 x oeular) under differential interferenee

eontrast, with anAxioCamMRS eamera andAxioVision v.4.2.8 software. For drawings, some speeimens

were stained with GomorEs triehrome and mounted in Canada balsam for study of their soft anatomy;

other speeimens were eleared and mounted in Hoyer’s or Gray and Wess’ media for study of their

selerotized struetures (all solutions prepared as in Humason, 1979). Illustrations were prepared with

the aid of a digital eamera (5 megapixels) and a projeetor attaehed to an Olympus BX51 mieroseope

equipped with phase eontrast. Taxon and author names in this study follow Esehmeyer (2012) for hosts

and MonoDb I http://www.monodb. org i for Monogenea.

Results

A single (ineomplete and therefore unfortunately unidentifled) aneyroeephaline monogenean

(Daetylogyridae, Aneyroeephalinae) was reeovered. All other monogeneans found on the I. hormuzensis

speeimens belong to Gyrodactylus.

European Journal of Taxonomy 30: 1-10 (2012)

Class Monogenea Van Beneden, 1858

Subclass Polyonchoinea Bychowsky, 1937

Order Gyrodactylidea Bychowsky, 1937

Family Gyrodactylidae Van Beneden & Hesse, 1863

Subfamily Gyrodactylinae Van Beneden & Hesse, 1863

Genus Gyr6)(iac(y/i/^ von Nordmann, 1832

Gyrodactylus jalalii sp. nov.

um:lsid:zoobank.org:act:A89D45DC-7CA2-4807-AF2E-E34F9A9BB060

Figs 1-2

Etymology

The species epithet,ya/a///, honours prof dr. Behiar Jalali Jafari (1953-2010) (obituary in Shamsi 2010).

He was a researcher in aquatic animal health and fish parasitology at the Veterinary Department of the

Islamic Azad University (Iran) and a keen student of monogeneans. The authors express the hope that

this patronym might serve as an indication for the respect and appreciation this kind man enjoyed from

his colleagues.

Type material examined

Thirty-one specimens, twenty of which (ethanol-preserved) were used for measurements. The holotype

(MNHN HEE301) and paratypes (MNHN HEE302-HEE305) are deposited in the Museum National

d’Histoire Naturelle (Paris, France). Paratypes are deposited in the Natural History Museum (Eondon,

United Kingdom) (NHMUK 2012.9.10.1-2012.9.10.2), the Royal Museum for Central Africa (Tervuren,

Belgium) (MRAC MT: 37711-37713), the Harold W. Manter Eaboratory of Parasitology (Eincoln,

Nebraska) (HWME-49758) and the United States National Parasite Collection (Beltsville, Maryland)

(USNPC 106050).

Type host

Iranocichla hormuzensis Coad, 1982 (Teleostei, Perciformes, Cichlidae).

Type locality

Mehran River, Persian Gulf Basin (2009).

Infection site

Gill filaments, fins, eye.

Description

(measurements in micrometres (pm) and angles in degrees (°); average ± standard deviation, followed

by range and number of measurements in parentheses).

Body (Fig. lA) fusiform, 361.7 ± 64.4 (294.7-528.9; n = 13) long; greatest width 128.1 ± 21.3 (98.8-

164.0; n = 13). Two head organs provided with single spicule each. Cephalic glands lateral to pharynx.

Pharynx composed of two tandem, muscular bulbs. Oesophagus short. Digestive glands lateral to

oesophagus. Caeca two, non-confiuent, reaching level of germarium. Male copulatory organ (MCO)

(Fig. IB) armed with a broad-based, robust, recurved, apical spine 5.7 ±1.1 (4.5-7.6; n = 8) long, 5-7

smaller fianking spines in a single row, becoming more slender from the terminal over the subterminal

towards the median ones (terminology of Garcia-Vasquez et al. 2007). Testis dorsal to germarium.

VANHOVE M.P..M. et al., Gyrodactylus from the GoMid Iranocichla hormuzensis

Germarium immediately posterior to uterus. Uterus with up to 2 embryos. Unieellular glands lateral

to terminations of eaeea, posterior to germarium. Raptor elongate (Fig. lA, D). Anehor (hamulus)

(Fig. IE) 79.9 ± 4.4 (70.4-86.3; n = 20) long; point 33.9 ± 2.3 (28.3-38.4; n = 19) long; base (superfieial

root) 28.2 ± 3.9 (19.7-33.1; n = 20) long; deep root knob-like; groove proximally at the base of the

anehor, serving as artieulation to superfieial (ventral) bar; shaft 49.5 ± 2.2 (45.6-55.0; n = 19) long,

proximally 10.2 ± 0.8 (8.7-11.9; n = 20) wide, distally 5.7 ± 0.8 (4.5-7.4; n = 19) wide; point sharply

eurved, with aperture 30.5 ± 2.5 (26.2-35.4; n = 20), aperture angle 42.7 ± 3.2 (36.9-47.3; n = 19), inner

aperture angle 46.9 ± 5.3 (30.1-53.3; n = 19); inner eurve length 1.9 ± 0.8 (0.7-3.3; n = 18) with point

eurve angle 9.5 ± 4.5 (4.2-22.9; n = 18). Ventral bar (Fig. ID) 33.5 ± 2.5 (29.8-37.7; n = 20) wide, 44.1

± 3.9 (36.4-49.8; n = 20) long; anterior bilateral proeesses slender, pronouneed and 6.0 ±1.1 (4.2-7.8;

n = 20) long with proeess to mid-length 10.4 ± 1.7 (7.5-14.8; n = 20). Median portion of ventral bar

Fig. 1. Gyrodactylus jalalii sp. nov. A. whole mount. B. male eopulatory organ. C. marginal hook.

D. anehor-bar eomplex. E. anehor. Seale bars represent 50 pm (whole mount), 10 pm (marginal hook,

MCO) or 30 pm (anehor, anehor-bar eomplex).

European Journal of Taxonomy 30: 1-10 (2012)

8.3 ± 1.2 (5.7-9.8; n = 20) long; shield (ventral bar membrane) 25.4 ± 3.3 (19.8-30.1; n = 20) long,

subreetangular, elearly striated in mueh the same way as bar proper. Deep (dorsal) bar 23.0 ± 2.8 (18.9-

29.9; n = 20) wide, medially eonstrieted and at eonneetion to deep root of anehors. Marginal hook

(Figs 1C, 2A, B) 31.8 ±4.1 (25.8-43.5; 77 = 20) long, shank with small distal bulb, 26.9 ±4.1 (22.1-40.0;

n = 20) long; siekle (booklet) 5.4 ± 0.3 (4.5-6.1; n = 20) long, 4.3 ± 0.4 (3.6-5.1; n = 20) wide proximally,

4.5 ± 0.5 (3.8-5.3; n = 20) distally; toe depressed, 2.0 ± 0.4 (1.4-3.0; n = 20) long; eonvex platform;

eoneave base; round keel; point of siekle proper as long as shaft, forming an angle of about 90° from

eaeh other; aperture 5.2 ± 0.5 (4.5-6.4; n = 20); instep/areh height 0.6 ±0.1 (0.4-0.8; n = 20).

Remarks

In eomparison with eongeners parasitizing eiehlids, the striated ventral bar proper and shield, as well

as the eonspieuous ventral bar proeesses, seem most reminiseent to G. zimbae Vanhove, Snoeks, Huyse

& Volekaert, 2011. However, the anterolateral proeesses of the ventral bar of G. zimbae are more ear¬

shaped. In G. zimbae, the ventral bar shield is slender and rounded and the booklet laeks an arehed base

{versus subreetangular shield and eoneave booklet base in G.jalalii sp. nov.). Other eiehlid Gyrodactylus

with relatively large ventral bar proeesses inelude G. shariffi Cone, Arthur & Bondad-Reantaso,

1995 and G. yacatli Gareia-Vasquez, Hansen, Christison, Bron & Shinn, 2011. Just like in G. jalalii

sp. nov., point and shaft of their booklet siekle are at a right angle. These speeies are easily distinguished

from G. jalalii sp. nov. by the smaller size of their haptoral selerites and the ventral bar in partieular

(e.g., anehor 47.5 and 48.4 long, ventral bar shield 14.4 and 8.5 long, in G. shariffi and G. yacatli

respeetively) (Gareia-Vasquez et al. 2011). It should be noted, however, that these two speeies were

deseribed from eultured Oreochromis niloticus (Linnaeus, 1758) in the Philippines, resp. Mexieo. The

authors deseribing G. yacatli eonsider aeeidental infeetion or host switeh a more likely seenario than an

Afriean origin (Gareia-Vasquez et al. 2011). Henee, G. zimbae seems to be the most eomparable eiehlid

parasite whose natural distribution is eertainly Afriean.

Comparison to Palearetie eongeners followed Pugaehev et al. (2009). The rather large ventral bar

proeesses, in eombination with the length of the marginal hooks, and MCO armed with one large apieal

spine and one row of smaller spines of similar size, resemble the morphology of G. ophiocephali Gussev,

1955 from Channa argus (Cantor, 1842) (Pereifonnes: Channidae) and Cyprinus carpio Linnaeus, 1758

(Cypriniformes, Cyprinidae), and to G. tokobaevi Ergens & Karabekova, 1980 from Gymnodiptychus

Fig. 2. A. mierograph detailing the marginal hooks of Gyrodactylus jalalii sp. nov. (photographed in

utero, with an Olympus BX50 mieroseope, using phase eontrast, and Olympus DP-soft v.3.2 software).

B. idem, detail. Seale bars represent 8 pm.

VANHOVE M.P..M. et al., Gyrodactylus from the GoMid Iranocichla hormuzensis

dybowskii (Kessler, 1874) (Cypriniformes, Cyprinidae). However, in G. ophiocephali and G tokobaevi,

the proeesses are longer than the ventral bar proper (median length, i.e., without the shield), whieh is

not the ease in G jalalii sp. nov. Elongate antero-lateral proeesses, albeit not longer than the ventral

bar proper, are also found in G hrabei Ergens, 1957 and G. mariannae Winger, Hansen, Baehmann

& Bakke, 2008, parasites of Cottus Einnaeus, 1758 spp. (Seorpaeniformes, Cottidae). These speeies,

however, have a eomparatively shorter anehor root than G jalalii sp. nov. The longitudinal striae on the

ventral bar shield as well as an overlap in size of anehor and marginal hook are reminiseent of G. lotae

Gussev, 1953 from Lota lota (Einnaeus, 1758) (Gadiformes, Eotidae). This speeies eanbe distinguished

from G. jalalii sp. nov. beeause the new speeies has blunter and larger ventral bar proeesses, and a

marginal hook siekle toe whieh joins smoothly into the platform, whereas this transition leaves a sharp

“bump” in the platform in G. lotae.

Discussion

Just as Barlow (2000) praises eiehlids, early students of Gyrodactylus expressed their high expeetations

for the seientifie interest of these flatworms, as they reeognized the speeies riehness of this genus and

the amount of work and skill it would take to adequately study it (von Nordmann 1832; Van Beneden &

Hesse 1863). Given this seientifieally ehallenging position of both eiehlids and gyrodaetylids, G. jalalii

sp. nov. was deseribed, a monogenean eetoparasite of the only Iranian eiehlid/. hormuzensis. To the best

of our knowledge, it is the first parasite to be formally deseribed from this fish speeies. It brings the total

number of Gyrodactylus speeies deseribed from eiehlid hosts to 15 (Papema 1979; Boeger & Popazoglo

1995; Christison et al. 2005; Pfikrylova et al. 2009, 2012; Gareia-Vasquez et al. 2011; Vanhove et al.

2011 ).

Jalali et al. (2001) observed that the highly distinet zoogeographieal regions of Iran and the high degree

of endemieity of its freshwater fishes gave rise to a diverse monogenean fauna. These authors did

not mention Gyrodactylus in their overview. Indeed, Jalali et al. (2005) noted that representatives of

this genus from the endemie freshwater fishes of the eountry are basieally unknown and undeseribed.

They did not report gyrodaetylids from Iranocichla. This eiehlid seems distributed mostly in the

Mesopotamian faunal region of Iran, from whieh Jalali et al. (2001) reported a rather unexplored,

endemie and speeifie monogenean fauna, eontaining both Palearetie and Afriean elements. For this

reason, G. jalalii sp. nov. was eomparedto Palearetie and Afriean eongeners. Morphologieal similarities

are unsurprisingly apparent (see above). Haptoral morphology alone does not allow reliable phylogenetie

inferenees for speeies of Gyrodactylus (Zi^tara & Eumme 2004). Unfortunately, we were unsueeessful

in amplifying DNA from G. jalalii sp. nov. Henee, genetie data are at present unavailable for the new

speeies, as is the ease for several eongeners used in this eomparison. However, it is noteworthy that

some of the aforementioned speeies show affinities in moleeular phylogenetie reeonstruetions. Indeed,

G. mariannae is elose to G. hrabei, whieh is genetieally similar to G.flesi Malmberg, 1957 (Winger et

al. 2008). This speeies elusters with G. lotae and G. zimbae, among other speeies, in a Gyrodactylus

elade eontaining representatives from various subgenera as defined by Malmberg (1970) and from many

eontinents and host taxa (Zi^tara & Eumme 2004; Vanhove et al. 2011). Despite the need for genetie

data, assuming, that G. jalalii sp. nov. is indeed related to this lineage, this faet alone does not make

it a very suitable eandidate to provide information on biogeographieal dispersal pathways of eiehlid

fishes. Indeed, several speeies also belonging to this diverse elade have been observed to show a broad

salinity toleranee (G. branchialis Huyse, Malmberg & Volekaert, 2004 and G. ostendicus Huyse &

Malmberg, 2004: Huyse et al. 2006) or the ability to switeh infeetion sites (G. arcuatus Byehowsky,

1933: Raeymaekers et al. 2008) or hosts (G. arcuatus'. Huyse et al. 2003, 2006). However, while there

is henee no proof that G. jalalii sp. nov. belongs to an exelusively Afriean or eiehlid-infeeting lineage,

this broad eeologieal speetrum enhanees the potential of these parasites as biogeographieal markers,

eonditional to the availability of moleeular data.

European Journal of Taxonomy 30: 1-10 (2012)

Acknowledgements

M.P.M.V. is a PhD fellow, andX.H. a post-doctoral fellow, of the Research Foundation-Flanders (FWO-

Vlaanderen). W. A.B. is a research fellow of CNPq - Brazil. F.M.B. received financial support from IRD

(BEST) for a training session in Belgium, during which part of the morphological work for this study

was performed. Prof L. Schoofs, Prof L. Arckens and Dr. E. Meelkop (KU Eeuven) are thanked for

the use of the ZEISS microscope, and Dr. Sh. Shamsi (Charles Sturt University, Wagga Wagga) for

help in obtaining literature. S. Asadollah (Isfahan University of Technology) and the late Prof B. Jalali

Jafari (Islamic Azad University, Teheran) are acknowledged for kindly providing us with Iranocichla

specimens and with information on this fish. Two anonymous reviewers provided useful suggestions.

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49: 5-28.

Manuscript received: 17 May 2012

Manuscript accepted: 7 November 2012

Published on: 20 November 2012

Topic editor: Rudy Jocque

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

the EJT eonsortium: Museum National d’Histoire Naturelle, Paris, Franee; National Botanie Garden

of Belgium, Meise, Belgium; Royal Museum for Central Afriea, Tervuren, Belgium; Natural History

Museum, Fondon, United Kingdom; Royal Belgian Institute of Natural Seienees, Brussels, Belgium;

Natural History Museum of Denmark, Copenhagen, Denmark.