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TheancientBalkanlakesharboranewendemicspeciesofDiaphanosomaFischer,1850(Crustacea:Branchiopoda:Cladocera)

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ZOOTAXA

ISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)Copyright © 2014 Magnolia Press

Zootaxa 3784 (5): 539–549

www.mapress.com/zootaxa/Article

http://dx.doi.org/10.11646/zootaxa.3784.5.3

http://zoobank.org/urn:lsid:zoobank.org:pub:63CBEC43-F367-4088-A1AD-1F85D23A8C12

The ancient Balkan lakes harbor a new endemic species of Diaphanosoma

Fischer, 1850 (Crustacea: Branchiopoda: Cladocera)

NIKOLAI M. KOROVCHINSKY1 & TRAJAN K. PETKOVSKI2 1A.N. Severtsov Institute of Ecology and Evolution of the Russian Academy of Sciences, Leninsky prospect 33, 119071 Moscow, Rus-

sian Federation. E-mail: nmkor@yandex.ru2Briselska 12, Taftalidze 2, 1000 Skopje, Republic of Macedonia. E-mail: dr.petkovski@yahoo.com

Abstract

Diaphanosoma macedonicum sp. nov. is described from material collected from the ancient Lakes Dojran and Prespa, lo-

cated in the central part of the Balkan Peninsula. It can be regarded as a member of the “D. mongolianum” species group.

It is characterized by specific, but not readily visible features, such as the absence of a thorn near the posterior valve mar-

gins, as well as setules between setae of the ventral valve inflection, and the presence of more chitinized integument. The

discovery of this new species previously identified as "Diaphanosoma brachyurum (Liévin)" highlights the necessity of

more detailed investigations of the zooplankton of Balkan lakes potentially populated by greater numbers of endemic cla-

doceran species. A short overview of the ancient lakes in the Central Balkans is provided.

Key words: Diaphanosoma macedonicum sp. nov., Crustacea, Cladocera, endemic species, ancient Balkan lakes, Lake

Dojran, Lake Prespa

Introduction

Investigations of the freshwater plankton in the Balkan lakes started at the end of the XIXth and the beginning of the

XXth century. In particular, Richard (1892) was the first investigator who listed six zooplankton species from

Macedonian Lake Dojran, including Daphnella brachyura (Livéin, 1848) var. minor Sars (= Diaphanosoma

brachyurum [Liévin, 1848]). Later, Gjorgjevic (Georgevitch) (1906, 1907) recorded Diaphanosoma brandtianum

Fischer, 1850 (= D. brachyurum, see Lilljeborg, 1901 and Korovchinsky 1992, 2004) from Lake Dojran and other

lakes, and noted some morphological differences between his specimens and other known forms of the genus

Diaphanosoma, proposing assignment of the former to a separate unnamed variety. Most of the subsequent authors

who studied Lake Dojran and Lake Prespa recorded only D. brachyurum (see Parenzan 1931; Popovska-Stankovič

1958, 1990; Guseska et al. 2012). More recently, in the 1980s, other species of the genus were found in some other

Balkan lakes: D. lacustris Kořinek, 1987 (syn. D. birgei lacustris Kořinek, 1987); D. mongolianum Ueno, 1938,

and D. orghidani Negrea, 1982 (Kořinek 1981, 1987; Korovchinsky 1987; Zarfdjian et al. 1990; Michaloudi et al.

1997).

In the majority of the aforementioned publications, taxa of the genus under consideration from the Balkan

region were either not described or were characterized with only a limited set of morphological detail, with the

exception of the description of diaphanosomas from one Bulgarian reservoir by Korovchinsky (1987). Such a

superficial study of the material might have led to incorrect identifications, which makes necessary a more detailed

reinvestigation of the species composition of Diaphanosoma from Balkan lakes.

An examination of samples collected in the Macedonian parts of Lake Prespa and Lake Dojran has revealed

representatives of the new species of Diaphanosoma, which is described in the present paper.

Brief description of the Balkan Lakes, Lake Prespa and Lake Dojran. In the central part of the Balkan

Peninsula, there is a belt of ancient lakes, consisting of two different principal lake groups (Cvijic 1911). The

western group belongs to the South Adriatic-Ionian Sea Basin and consists of three waterbodies, the largest and

Accepted by M. Alonso: 28 Feb. 2014; published: 1 Apr. 2014 539

deepest Lake Ohrid (Ohridsko Ezero), a large Lake Prespa (Prespansko Ezero), as well as a small, ever-shallowing

Lake Maliq. These lakes are the remnants of the vast Palaeolake Dessaret (Albrecht & Wilke 2008). One more

ancient lake is situated in Epirus in northwestern Greece, Lake Ioannina/Pamvotis (Janinsko Ezero), which belongs

to the same geographical region as the Dessaret lake group and contains some pre-glacial faunistic remnants.

Further to the east, there are several lakes of the second principal lake group belonging to the Aegean Sea

catchment. The eastern part of the latter includes the Paionian Lake Dojran (Dojransko Ezero) and, in the

Mygdonian Basin extending toward the Aegean Sea, there is the ever-shallowing Lake Koronia (Ajvasilsko Ezero)

and the larger and deeper Lake Volvi (Besicko Ezero). The western part of this lake group comprises the so-called

Eordean lakes, the formerly very large Lake Vegoritida (Ostrovsko Ezero) and some other smaller lakes.

Lake Prespa consists of two water bodies, Great and Small Lake Prespa, which were separated due to

accumulation of the sediment deposits from the River Stara Reka (Cvijic 1911). Great Lake Prespa is situated at the

southwestern border of Macedonia and has a surface area of 254 km2 shared between Macedonia (176.8 km2),

Albania (49.4 km2), and Greece (47.8 km2). It is the highest tectonic lake in the Balkans, located at an elevation of

849 m msl, with a length of 27.3 km, a width of 17.0 km, an average depth of 14.0 m and a maximum depth of 58.0

m (Albrecht & Wilke 2008). The level of the water in Great Lake Prespa fluctuates considerably because the lake

has no major tributaries and a portion of the water volume escapes permanently downward via karst sinkholes and

an underground drainage system into Lake Ohrid, 160 m lower in elevation. Both lakes, Great and Small Lakes

Prespa, are believed to have been formed during the Pliocene/Pleistocene Epoch, about 2–5 million years ago. Lake

Prespa, in contrast to Lake Ohrid, is much shallower and thus has probably been more susceptible to complete

desiccation at various times in the past as a result of its karstic outflows. This lake could be classified as oligo-

mesotrophic and the level of its endemism (30 known endemic species, see Petkovski 2003) is much lower than

that recorded for Lake Ohrid (212 endemic species), likely due to the smaller depth of the former (58 m vs 289 m)

(Frogley & Preece 2004; Albrecht & Wilke 2008).

Lake Dojran, with a surface area of 42.66 km2 (length 8.9 km, width 7.0 km), average depth of 6.5 m and a

maximum depth of 10.0 m, is located at the southeastern border of Macedonia and is shared with Greece. It lies at

an altitude of 148 m msl. The lake was formed in a closed, karstic basin created by a combination of Tertiary

volcanic and tectonic activities. The lake basin itself represents a remnant of the Plio-Pleistocene Paionian Lake,

about three times larger and ten times deeper than the existing Lake Dojran (Cvijic 1911). Evaporative losses by

the lake are substantial, but it receives inflows via submerged springs and pulses of snow-melt waters from the

Belasica and Karadag mountains. Because of karstification of the underground substratum, Lake Dojran is calcite-

and chloride-rich, alkaline, eutrophic and highly productive (Griffiths et al. 2002). In the past, Lake Dojran’s

primary significance was as an inland fishery, having been claimed to have had the highest rate of lacustrine fish

productivity in Europe (Naumovski 1991). This lake hosts 12 known endemic taxa (Petkovski 2003).

Material and methods

Material was collected in November 1989 in the Macedonian portion of Lake Dojran (5 parth., 57 gam. and 34

males, coll. T. Petkovski) and in November 1958 in Great Lake Prespa (76 gam. and 28 males, coll. O. Popovska-

Stankovič).

Samples were examined under a dissecting microscope Lomo MBS-10 and a compound microscope Olympus

BX-41 with camera lucida. Measurements of bodies and body parts were made according to Korovchinsky (2004).

Measurements of the parts of swimming antennae and their setae were made from a female individual from Lake

Dojran.

Abbreviations. AnL= length of swimming antennae; BL= body length; DE=diameter of eye; gam.=

gamogenetic females; HH= head height; HL= head length; I= large two-segmented setulated seta on distal corner

of gnathobases of tl I – tl V; i= curved setulated seta on gnathobase of tl I; J= modified thorn-like naked, hooked

seta on distal corner of gnathobases of tl III – tl V; n = number of filtering setae of endopodite and gnathobase of

thoracic limbs; p = not-setulated seta on proximal corner of gnathobases of tl II – tl V; parth. = parthenogenetic

females; tl I…tl VI= thoracic limbs of first…sixth pairs, UB – upper antennal branch.

KOROVCHINSKY & PETKOVSKI 540 · Zootaxa 3784 (5) © 2014 Magnolia Press

Results

Taxonomy

Class Branchiopoda Latreille, 1817

Order Ctenopoda G.O.Sars, 1865

Superorder Cladocera Latreille, 1829

Family Sididae Baird, 1850

Genus Diaphanosoma S. Fischer, 1850

Diaphanosoma macedonicum sp. nov.

(Figs. 1–4)

Daphnella brachyura (Liévin, 1848): Richard 1892, p. 151,152. Diaphanosoma brandtianum Fischer, 1850 n. var.: Gjorgjevic 1906, p. 208, 214–217, Fig. 14–17; Georgevitch, 1907: p. 6. Diaphanosoma brachyurum (Liévin, 1848): Parenzan 1931, p. 40; Popovska-Stankovič, 1958, p. 131; 1990, p. 56, 57; Guseska

et al., 2012, p. 940.

Etymology. the species name is derived from the name of the country (Macedonia), in whose the lacustrine zones

the species was originally found.

Type material. Holotype. A female from Lake Dojran with body length 1.31 mm deposited in the collection

of type material of the Zoological Museum of M.V. Lomonosov Moscow State University (Ml 125).

Paratypes. Eight females and two males from the same lake in the same collection (Ml 126), other paratypes

are deposited in the authors’ collections.

Diagnosis. Body conically elongated, slightly narrowing anteriorly, with head of comparatively large or

medium size, having rather strongly developed dorsal part. Swimming antennae long, with the upper branch

reaching or surpassing the posterior valve margin; basipodite massive, with a small, sharp spine on the outer side of

its apical end. Proximal segment of upper two-segmented antennal branch bearing conspicuous apical spine

dorsally. Formula for antennal seta 4–8 / 0–1–4. Shell with conspicuous dorsoposterior angle seeming more

chitinized than usual, having conspicuous dorsal and lateral ribs separating shell valves. Posteroventral valve

margin armed with a row of 12–27 small comparatively sparsely situated denticles having thin setules between

them. Posterior valve margin with rows of marginal and submarginal densely situated spinules and few setules

between them but lacking internal submarginal thorn. Male’s antennal basipodite with large apical outer spine. Tl I

with large terminal hook, having a number of denticles on its inner margin, and a naked, basally widened seta with

a constriction close to it. Copulative appendages tubular.

Description. Parthenogenetic and gamogenetic female. They differ only in the presence of dark resting eggs

in brood pouches in the latter (Fig. 1I). Body conically elongated, slightly narrowing anteriorly, with head of

comparatively large or medium size (length 37.5–43.2 %, 31.3–37.7 % and height 22.1–25.4 %, 17.0–23.0 % of

body length in specimens from Lake Dojran and Lake Prespa, respectively), having rather strongly developed

dorsal part, shape of which may vary in preserved specimens (Fig. 1A, 2A–C).

Eye small (6.0–7.2 %, 6.5–7.6 % of body length) and situated closer to ventral side or anteroventral corner of

head.

Antennules small (Fig. 1A, C) with nine small aesthetascs and rather long sensory seta.

Swimming antennae long (73.3–84.7 %, 73.2–87.7 % of body length), with the upper branch reaching or

surpassing the posterior valve margin (47.5–53.4 %, 49.1–56.1 % of antennal length) (Fig. 1A). Their basipodite is

massive, with a small, sharp spine on the outer side of its apical end closer to the base of the upper branch (Fig. 1D)

and long naked seta on the dorsal side of this apex (Fig. 1K). A similar seta sits on the posterodorsal side of the

basipodital proximal part (Fig. 1J). Proximal segment of upper two-segmented antennal branch bears conspicuous

apical spine dorsally and outgrowth of slightly varied shape laterally (Fig. 1E). Distal segment of this branch with

Zootaxa 3784 (5) © 2014 Magnolia Press · 541NEW SPECIES OF DIAPHANOSOMA

larger spine provided with outer and inner outgrowths near its base (Fig. 1F). Proximal segment of lower three-

segmented antennal branch fused with the neighboring one internally (Fig. 1H), but has a clear articulation with it

FIGURE 1. Diaphanosoma macedonicum sp. nov., female, Lake Dojran, Macedonia. A, general lateral view. B, general dorsal view. C, antennule. D, basipodital distal outer end. E, distal part of proximal segment of upper antennal branch of two individuals. F, apical end of distal segment of upper antennal branch. G, distal part of lower antennal branch. H, proximal part of lower antennal branch, inner side. I, resting eggs in brood pouch of gamogenetic female. J, dorsal seta of proximal part of antennal basipodite. K, dorsal seta of distal part of antennal basipodite.

KOROVCHINSKY & PETKOVSKI 542 · Zootaxa 3784 (5) © 2014 Magnolia Press

FIGURE 2. Diaphanosoma macedonicum sp. nov., female (A–D, I – Lake Prespa, E–H, J–L – Lake Dojran) A–C, head, lateral view. D, E, posterior part of shell. F–I, armament of ventroposterior valve margin. J, ventral valve inflection. K, postabdomen, lateral view. L, postabdominal claw.

Zootaxa 3784 (5) © 2014 Magnolia Press · 543NEW SPECIES OF DIAPHANOSOMA

FIGURE 3. Diaphanosoma macedonicum sp. nov., female, Lake Dojran, Macedonia. A, distal part of endopodite of thoracic limb of first pair (tl I). B, apical end of exopodite of tl I. C, gnathobases of tl I. D, thoracic limb of second pair (tl II). E, setae on distal corner of gnathobases of thoracic limb of third pair (tl III). F, thoracic limb of sixth pair (tl VI).

externally. Formula for antennal seta 4–8 / 0–1–4. Lateral setae of upper branch are comparatively short (25.3–27.3

% of body length) and armed uniformly with rough setules of the “swimming type”, while the two apical ones are

considerably longer (50 % of body length), having proximally the same setules but distally, thin, sparsely situated

setules of the “sensory type”. Three apical setae of lower branch are slightly shorter (38.7–43.3 % of body length)

with armament of similar structure.

Shell with slightly arched dorsal side and conspicuous dorsoposterior angle (Fig. 1A). It seems more chitinized

than usual, having conspicuous dorsal and lateral ribs separating shell valves (Fig. 1A, B). Valves with

comparatively short posterior margin smoothly connected with ventral margin and forming a comparatively narrow

inflection bearing 12–14 long, finely setulated setae, the proximal of which are diminished and sit submarginally

(Fig. 2D, E, J). One-two similar setae sit on the ventralmost part of posteroventral margin (Fig. 2F–H, J), which is

armed with a row of 12–27 small comparatively sparsely situated denticles having thin setules between them (Fig.

2D–I). Posterior valve margin with rows of marginal and submarginal densely situated spinules and few setules

between them.

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Postabdomen comparatively narrow with prominent ventrodistal side and long postabdominal setae (about 65

% of body length) (Fig. 2K). Terminal claws with three basal spines, the distal of which is longest and rather

straight (Fig. 2L). Rows of spinules are situated above the basal spines and distally along the outer lateral side of

claws.

Six pairs of thoracic limbs, all with epipodites. Structure and armament of the limbs are schematically

summarized in Table 2. Exopodite of tl I comparatively narrow at its end (Fig. 3B), while the exopodite widens

terminally in limbs tl II–tl VI (Fig. 3D, F). Those of tl I–tl V bear 5–6 terminal and 3–5 lateral, unsegmented, thick

and long, densely setulated setae. Endopodites of tl I–tl V are inconspicuously subdivided into four segments and

bear 29–48 long, two-segmented setulated filtering setae and one (tl I) or two (tl II–tl V) outer setae, similar to those

of the exopodite (Fig. 3A, D: arrows). Among the latter, the subterminal one is conspicuously shorter than the other

(Fig. 3D). Small, thorn-like, naked seta on the end of proximal segment above the row of filtering setae of tl I (Fig.

3: ns). Gnathobase of tl I with a row of eight–nine two-segmented, distally finely setulated setae, one large two-

segmented seta on its distal corner (I) and a shorter, curved, basally widened seta (i) near it (Fig. 3C). Gnathobases of

tl II–tl V are larger, bearing 22–28 filtering setae and one naked seta (p) proximally and one long, two-segmented

seta (I) distally with rough setules; an additional modified naked, hooked seta (J) with few lateral denticles near the

previous one is also present (tl III– tl V) (Fig. 3D, E). Tl VI small and strongly modified (Fig. 3F). Its exopodite

reduces up to the terminal plate and is armed with five terminal and one lateral setae. Endopodite with seven similar

setae and one thorn. Gnathobase with two long setae and three thorns of different sizes and shapes.

TABLE 1. Measurements of body and body parts of adult females of Diaphanosoma macedonicum sp. nov. from two

populations (in each column from top to bottom: range, M, SD, CV).

TABLE 2. Structure and armament of thoracic limbs of Diaphanosoma macedonicum sp. nov. from Lake Dojran,

Macedonia, November 1989 (for explanations of abbreviations, see above and Figure 3).

Male. Most of features are similar to those of females.

Antennules long (44.3–60.7 % of body length), attached to large outgrowths of ventral side of head (Fig. 4A,

B). Their proximal part is thickened and the distal one bears numerous densely situated setules (Fig. 4C).

BL, mm HL : BL, % HH : BL, % DE : BL, % AnL : BL, % UB : AnL, % Number of denticles onpostero-ventral valve margins

1. Lake Dojran (n = 20)

1.09 – 1.33 37.5 – 43.2 22.1 – 25.4 6.0 – 7.2 73.3 – 84.7 47.5 – 53.4 13 – 24

1.20 38.9 24.2 6.6 78.9 49.3 18.1

1.43 0.77 0.33 3.45 1.77

3.69 3.19 4.93 4.37 3.58

2. Lake Prespa (n = 15)

0.90 – 1.12 31.3 – 37.7 17.0 – 23.0 6.5 – 7.6 73.2 – 87.7 49.1 – 56.1 12 – 27

1.02 34.1 19.0 7.2 81.0 52.5 20.9

1.62 1.41 0.44 4.18 2.04

4.74 7.43 6.13 5.16 3.89

Limb pairs Exopodite (apical + lateral setae)

Endopodite Gnathobase Epipodite

I 5 + 5 (n7 + 1) + (n4) + (n4) + (n30–32) (n8–9) + I + i +

II 6 + 5 (n7–8 + 1) + (n4 + 1) + (n4) + (n28) (n28 + p) + I +

III 6 + 5 (n7–8 + 1) + (n4 + 1) + (n4) + (n32) (n24 + p) + I + J +

IV 6 + 5 (n8 + 1) + (n4 + 1) + (n4) + (n24) (n24 + p) + I + J +

V 6 + 3 (n7 + 1) + (n4 + 1) + (n18) (n22 + p) + I + J +

VI 5 + 1 7 + one thorn 2 + three thorns +

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Antennal basipodite with large apical outer spine (Fig. 4D). Tl I with large terminal hook, having a number of

denticles on its inner margin, and a naked, basally widened seta with a constriction near it (Fig 4F, G). Above it,

comparatively short naked setae located on the ends of each of the three previous segments.

Copulative appendages tubular (about 30 % of body length).

Size. Body measurements of species are presented in Table 1. Female body length 0.75–1.33 mm. Resting eggs

ovally-elongated 0.24–0.29 x 0.16–0.18 mm. Male body length 0.67–1.15 mm.

Intra-and interpopulational variability. On the whole, judging from Table 1, the intrapopulational

variability of specimens from both lakes seems to be low. It is slightly higher in specimens from Lake Prespa,

which are smaller than those from Lake Dojran and have a smaller head and a less developed basipodite of their

swimming antennae – body parts, the development of which seems directly connected. The qualitative variability

of some structures, head shape, armament of swimming antennae, and shell margin, is shown in Figures 1 and 2.

FIGURE 4. Diaphanosoma macedonicum sp. nov., male, Lake Dojran, Macedonia. A, general lateral view. B, head, ventral view. C, setules of distal part of antennule. D, apical end of antennal basipodite, outer side. E, copulatory appendages. F, distal part of thoracic limb of first pair (tl I). G, hook on apical end of tl I.

KOROVCHINSKY & PETKOVSKI 546 · Zootaxa 3784 (5) © 2014 Magnolia Press

Differential diagnosis. The new species is especially close to D. mongolianum differing from it, first of all, in

absence of a thorn near the posterior valve margin. Furthermore, the former possess more strongly chitinized shell

integument and lacks setules between setae of the ventral valve inflection. The difference in integument structure is

very obvious if specimens of the new species from Lake Dojran are compared with co-occurring representatives of

D. mongolianum, which are smaller in size and have a thin, transparent integument. The same features distinguish

D. macedonicum sp. nov. from D. lacustris, which also has more numerous denticles (usually 30–50, sometimes up

to 60) on posteroventral valve margins. D. orghidani Negrea, 1982 widely distributed in Eurasia, including Balkan

lakes, lacks thorn near posterior valve margins but it has quadrangular head, low posterior valve margins, row of

marginal valve denticles shifted far ventrally and copulatory appendages of males sole-like, enlarging distally.

Remarks. Taking into consideration that D. macedonicum sp. nov. can co-occur with D. mongolianum or

possibly with other congeners, it should be noted that the synonymy of the new species described above is

provisional since it might not reflect other different species currently grouped under the names “D. brachyurum”

and “D. brandtianum”. Only a re-investigation of material of previous researchers who studied diaphanosomas

from Balkan lakes can unambiguously elucidate this question.

Discussion

In his schematically illustrated description of diaphanosomas from Lake Dojran, Gjorgjevitch (Georgevich) (1906,

1907) noted their differences from known forms and assigned them to a new variety. Some characteristic features

of the latter—a large head with a well-developed dorsal region, long swimming antennae, and the presence of 16–

17 denticles on the posterior valve margin led us to suspect that this author dealt not with D. brachyurum but

probably with the new species under consideration or/and with another close species.

The new species is especially close to D. mongolianum as well as to D. lacustris in most respects of its

morphology, differing from them only in inconspicuous morphological details. For this reason it may be regarded

as a new member of the “D. mongolianum” species group, currently comprising three sibling species. Two of

them, mentioned above, living both in fresh and brackish waters, are widely distributed primarily within Northern

Eurasia (Korovchinsky 1987, 2004), while D. macedonicum sp. nov. seems to have a restricted occurrence in

ancient freshwater Balkan lakes. At least it was not encountered by the first author of the present paper

(Korovchinsky, unpubl. data) in several samples collected from other Balkan waterbodies in Macedonia, Albania

and Greece.

D. macedonicum sp. nov. is actually a new endemic species (and first species of Cladocera) for ancient lakes

Prespa and Dojran. Before, in this region, only a single endemic benthic cladoceran species, Phreatalona smirnovi

(Petkovski et Flössner) (syn. Alona smirnovi Petkovski et Flössner), was known from Lake Ohrid and the

neighboring springs (Petkovski & Flössner 1972; Albrecht & Wilke 2008). One more candidate for endemic status

is Leptodora kindtii (Focke) from Lake Dojran and possibly from other similar Balkan lakes, which locally creates

a highly recognizable separate molecular-genetic clade, unstudied morphologically yet (see Millette et al. 2011).

D. macedonicum sp. nov. is a new species for the European fauna of Cladocera, which is considered

comparatively well known among others (Korovchinsky 1996). Despite this, the new species of the group appears

to have been recorded rather regularly for this region (e.g. Hudeč 1997; Juračka et al. 2010; Kotov & Alonso

2010). This recognition opens the way for further taxonomic discoveries, especially in the Southern Europe

unglaciated in the past (see also Hamrová et al., 2012).

In total, there are few known Сladocera occurring exclusively in ancient lakes. Most of them (seven species of

the genera Kozhowia Vasiljeva et Smirnov, Parakozhowia Kotov, Phreatalona Van Damme et Dumont and

Chydorus Leach) inhabit Lake Baikal (see Smirnov 1984, 1996; Kotov 2000; Sinev & Kotov 2001), all being

bottom dwellers. Another planktonic species, Diaphanosoma modigliani Richard, occurs in the ancient Indonesian

Lakes Toba (Sumatra) and Tempe (Sulawesi) (Korovchinsky 1998). Other cladoceran endemics recorded there

have not yet been described properly and their status remains questionable (Korovchinsky 2013a). This situation is

also the case for other ancient lakes (e.g., Biwa, Great African lakes), the cladocerans of which either are poorly

known or belong to widely distributed species (Korovchinsky 2000, 2013b).

Thus, D. macedonicum sp. nov. appears to be the second known endemic species of Cladocera in the Balkans.

It is so far recorded in two ancient lakes of the Balkan Peninsula and its distribution in other water bodies of the

Zootaxa 3784 (5) © 2014 Magnolia Press · 547NEW SPECIES OF DIAPHANOSOMA

region populated by some, sometimes co-occurring congeners (see Michaloudi et al. 1997; this paper), should be

studied further.

Acknowledgments

NMK thanks Dr. A.A. Kotov (Institute of Ecology and Evolution, Moscow, Russia) for help with literature and

reviewer’s corrections and comments and Prof. N.N. Smirnov (the same Institute) for the improvement of the first

draft of the manuscript. The corrections and comments of second anonymous reviewer helped to improve the

manuscript further as well. Thanks also to Dr. R. Darrell Smith for his review of the English text. The research

work by NMK was supported by a grant from the Russian Foundation for Basic Research (no. 12-04-00207a).

References

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