Zoosyst. Evol. 87 (2) 2011, 291 295 / DOI 10.1002/zoos.

201100007

The first representative of the fully aquatic leaf beetle genus

Macroplea Samouelle, 1819 (Coleoptera, Chrysomelidae)
in Turkey: Macroplea mutica (Fabricius, 1792), with notes
on its biology, habitat, host plant and distribution
Ibrahim Trkgl1, Ali Nafiz Ekiz*, 2, Ali Gk3 and Blent Sen4
1
2
3
4

Fisheries Research Institute, Elazg, Turkey

Usak University, Faculty of Art and Science, Department of Biology, 64200 Usak, Turkey
Sleyman Demirel University, Faculty of Art and Science, Department of Biology, 32260 Isparta, Turkey
Frat University, Faculty of Fisheries, 23119, Elazg, Turkey

Abstract
Received 2 October 2010
Accepted 15 February 2011
Published 23 September 2011

Key Words
New record
Palearctic
morphology
phylogeography

Macroplea Samouelle, 1819 is a small genus of fully aquatic leaf beetles distributed in
the Palearctic. Up to now, none is recorded from Turkey, and in this paper, we report
Macroplea mutica (Fabricius, 1792), from Turkey for the first time. We present important morphological characters of larvae, pupae and adults collected from Eastern Anatolia. We also noted the biology, habitat features, host plant associations and the current
distribution of the species. The relatively large but fragmented distribution of M. mutica
is discussed on the bases of restricted underwater locomotion and reduced ability of
flight of the species. The importance of its existence in Anatolian refugium is evaluated with regard to the phylogeography of the species.

Introduction
Macroplea Samouelle, 1819 is a species poor group of
the Donaciinae, the only aquatic group within Chrysomelidae. The genus consists of only five species, and
all are distributed in the Palearctic region: M. appendiculata (Panzer, 1794), M. japana (Jacoby, 1885),
M. mutica (Fabricius, 1792), M. pubipennis (Reuter,
1875) and M. skomorokhovi Medvedev, 2006 (Askevold
1990; Lays 2002; Silfverberg 2010).
As the other members of the genus Macroplea,
M. mutica is fully aquatic and obligatorily associated
with aquatic plants, with larvae and pupae submerged
and adults often found under water (Konstantinov 2003;
Jch & Balke 2008). In other words, it lives underwater
throughout its life cycle. For this reason, the studies
about this species are very restricted and the details of
its biology, especially in larval stages, are poorly
known. M. mutica is primarily halophile living in
brackish water at the coasts of North Sea, Baltic Sea,
Mediterranean and Caspian Sea but also known from

* Corresponding author, e-mail: alinafiz@yahoo.com

# 2011 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim

freshwater habitats (e.g. from lakes in Germany, Hungary, Italy and Sardinia) (Nilsson 1996; Mende et al.
2010; Silfverberg 2010).
During the macrobenthic faunal surveys conducted at
Lake Hazar (Elazg) between 20072008, a large number of larvae, pupae (cocoons) and a few adult specimens belonging to the Donaciinae were collected by
the first author. After careful examination of these samples, we identified the larvae, pupae and adult specimens as M. mutica. As far as we know, none of the species of Macroplea were recorded from Turkey so far.
Existence of the species at Lake Hazar (Elazg province, Turkey) is quite interesting with regard to its current distribution (see discussion).
Consequently, the objectives of this paper are: i) to report Macroplea (namely M. mutica) in Turkey, for the
first time, ii) to re-evaluate the important taxonomic characters of larvae, pupae (cocoons) and adults on the base
of the collected specimens, iii) to illustrate larvae, pupae
and adults (the spermatheca is illustrated for the first
time) iv) to contribute the habitat and host plant associa-

292

Tu rkgu lu , I. et al.: The first representative of genus Macroplea (Coleoptera, Chrysomelidae) in Turkey

Figure 1. Map showing the study area (Collecting localities are shown as black circles).

tions of the species, v) to interpret the existence of M. mutica in Turkey with regard to phylogeographical aspects.

Materials and methods

This study is based on the 554 larvae, 30 pupal cocoons and only
eight adult specimens (four male and four female) collected by the
first author from Lake Hazar between 20072008. Lake Hazar is a
tectonic lake located at the west of South-eastern Taurus Mountains,
near Elazg province in East Anatolia (38 310 N, 39 250 E). The mean
altitude of the lake is about 1238 m asl. Localities where specimens
were collected are shown as black circles in Figure 1.
The specimens of larvae, pupae and adults were collected by pulling the submerged host plants and the bottom substrate by using a
Van Veen grab. Specimens were placed in 70 percent ethanol and taken to laboratory for identification. 13 larvae, 9 pupal cocoons and 6
adult specimens (3 males, 3 females) were deposited at the Department of Biology, Faculty of Arts and Sciences, Sleyman Demirel
University, Isparta, Turkey and 541 larvae, 21 pupal cocoons and
2 adult specimens (one male, one female) were deposited in the
Faculty of Fisheries, Frat University, Elazg, Turkey.

Results
Macroplea mutica (Fabricius, 1792)
Figure 2
Taxonomical remarks
Larva. (Figure 2ab) Body creamy-white, covered with
short setae arranged in patches; head and legs pale yellow, mouth parts, claws, abdominal hooks, setae and
ocelli brown; head prognathous, very small (about 18

museum-zoosyst.evol.wiley-vch.de

19 times shorter than whole body length), cutting edge

of mandible with two teeth, antennae three segmented,
antennal ratio: 2 : 1 : 1. Legs short, tarsal claw single,
hook-shaped. Abdomen with ten segments, all segments
with short setae, certain setae on segments 6 and 7 distinctly longer, spiracular atria of eighth segment modified to canaliculated hooks, ninth and tenth segments
reduced.
Pupae. (Figure 2ce) Ovoid, colour changes from
creamy-white to brown with respect to its developmental stage, always attached to the roots of host plant.
Adults. (Figure 2fj) Males and females resemble
each other but males are distinctly smaller than females
(average length 4.7 mm, width 1.7 mm for males and
average length 5.5 mm, width 2.3 mm for females). Body
narrow and slender. Head generally black (brownish in
females), surface micro-granulate, hairless; behind antennal base with very distinct and prominent transverse
tubercles, which are separated by deep longitudinal
groove; antennae long, slender, base and apex of all antennal segments slightly yellowish brown, ratio of antennal segments: 6 : 3 : 4 : 8 : 9 : 9 : 10 : 11 : 11 : 12 : 12. Pronotum 1.2 times wider than its length anteriorly,
generally straw-coloured with two black spots, pronotal
disc with a shallow longitudinal groove (this groove is
deeper in females); anterior margin slightly angled in the
median, lateral margins distinctly projected sidewards in
anterior half and almost parallel posteriorly, each corner
of the pronotum projected bearing a seta, surface of the
pronotum covered with small punctures and tiny setae.
Scutellum cuneiform, black, micro-granulated. Elytra
elongate, almost two times as long as wide in the median
and narrowed backwards, apex emarginated, with a sharp

# 2011 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim

Zoosyst. Evol. 87 (2) 2011, 291 295

293

Figure 2. Macroplea mutica (Fabricius, 1792). ab. Larva, a. Lateral view, b. Ventral view of head and thorax; ce. Pupal cocoons; fg. Adults, f. Male, g. Female; hi. Aedeagus, h. Dorsal view, i. Lateral view, j. Spermatheca.

triangular spine at outer angle; humeral calli well developed; elytra with regular puncture rows, sutural puncture
row only reaches the anterior third, other rows paired into
six rows posteriorly before reaching the apex of the elytra, size of punctures quite variable such that posterior
punctures are bigger; elytra generally straw-coloured but
elytral suture and inside of puncture rows black forming
stripes, black stripes never reach the apex; surface of elytra pubescent. Underside densely covered with velvet
hairs, impunctate; propleura shiny yellow; prosternum
and metathorax black; sides of abdominal sternites black
with yellowish brown median area. Legs long and slender, femora and tibiae of all legs brownish (hind tibia of

# 2011 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim

females evidently longer), first three tarsal segments

black with yellowish apices, fourth tarsal segments
straw-colored with black apices; length of first and second tarsomeres almost equal about 0.2 mm, third small
about 0.1 mm, fourth relatively longer (about 0.7 mm for
fore and middle legs and 1.0 mm for hind legs). Aedeagus and spermatheca as in Figure 2hi and Figure 2j respectively.
Habitat and host plant. Macroplea mutica was previously recorded from some aquatic plants: Brasenia
schreberi Gmel., Myriophyllum spicatum L., Potamogeton filiformis Pers., P. maritimus L., P. pectinatus L.,

museum-zoosyst.evol.wiley-vch.de

294

Tu rkgu lu , I. et al.: The first representative of genus Macroplea (Coleoptera, Chrysomelidae) in Turkey

Ruppia maritime L., R. rostellata L., Zostera marina L.,

Z. maritime L., Zanichellia palustris L. (Mohr 1966;
Lays 2002; Bienkowski & Bienkowskaja 2004, BioInfo
UK). Most of the specimens including larvae, pupal cocoons and adults were collected from the upper littoral
zone (05 m depth), however; a few larvae were also
collected from 515 m depth. Larvae and pupae were
found attached to the roots of its host plant, Potamogeton pectinatus L., a common submerged macrophyte at
Lake Hazar. Larvae were also found on the soft, sandy,
sedimentary bottom of the lake. Rare adult specimens
were collected from the submerged parts of the same
host plant as larvae and pupae. The number of larval
specimens collected was particularly higher at the sedimentary shores where small streams (Zkkm, Behrimaz
and Krk streams; see Figure 1) flow into the lake.
Biological notes. It was observed that the numbers of larvae were highest in August and September. Seasonal
numbers of the collected larvae indicate that M. mutica
has one generation per year at Lake Hazar. Mating seems
to be substantially occurring in late spring and early summer from May to July. Although not observed in this
study, it is well known for the members of the genus
Macroplea that females lay eggs under water, in gelatinous envelopes. One or two weeks after oviposition, the
eggs begin to hatch. The hatchling larvae attach to the
roots of the host plant by the spiracle hooks on their
eighth abdominal segment. As we collected some of the
larvae from the sandy substrate at the bottom of the lake,
it is quite probable that larvae move to attach another
plant when the host quality deteriorates. Pupation occurs
in waterproof yellowish or reddish brown cocoons. Several respiration holes were observed on the cocoons as
stated in Bienkowski (1996). Both mature larvae and
adults were found overwintering in cocoons.
Distribution. Macroplea mutica is a transpalearctic species distributed in Belgium, Belarus, Denmark, Estonia,
Finland, France, Great Britain, Germany, Hungary, Italy,
Latvia, The Netherlands, Norway, North and South European parts of Russia, Poland, Romania, Sweden, Algeria, Kyrgyzstan, Mongolia, Uzbekistan (Silfverberg
2010) and China (Mende et al. 2010). The species is
recorded from Turkey for the first time.

Discussion
Due to lack of sufficient studies, the knowledge about
the aquatic leaf beetles of Turkey is limited to only a
few local faunistic records. Recently, some papers providing both taxonomical and ecological data have been
performed (e.g. Ekiz et al. 2009; Ekiz & Gk 2010).
However, there is still a certain need of new surveys,
because many regions of Turkey are surely hosting a
lot of undiscovered species. Morphologically, the Anatolian population of M. mutica is similar to those described in literature, as well as the recorded host plant
and described habitat in this paper.

museum-zoosyst.evol.wiley-vch.de

The currently known distribution of M. mutica is relatively larger in comparison with its congeners (Silfverberg 2010), but rather fragmented most likely because of
lack of data, and constitutes a case of vicariance (Klsch
et al. 2006; Klsch & Pedersen 2010). Discovery of
M. mutica from East Anatolia enlarges the known distribution area to south and also introduces a new perspective for dispersal of the species. As is known, the locomotion of adults under water is rather restricted to
walking on the substrate and vegetation. Out of water,
they are hardly able to walk on their thin and weak legs
(Mende et al. 2010). According to Mann & Crowson
(1983), the hindwing venation of Macroplea is mostly
reduced, which supports a reduced ability to fly. Mende
et al. (2010) stated that M. mutica was never observed
flying nor can flight be induced out of the water under
various conditions such as light, darkness and warmth.
However, according to Mohr (1985), this species is fully
winged and can fly. Similarly, for a related species M. japana, Zhang et al. (2010) point out that they occasionally
emerge from the water and fly, when disturbed. Nevertheless, M. mutica was able to colonise the Palearctic Region throughout. It is obvious that the dispersal of this
species is passive either by eggs and cocoons attached to
drifting host plants in large brackish water bodies or by
migratory waterfowls (endozoochory) as mentioned by
Mende et al. (2010). Since the host plants of M. mutica
are also preferred food of many waterfowls (Figuerola &
Green 2002). Location of Lake Hazar supports the dispersal by zoochory, because it is on the migration route
of many waterfowl species.
Recently, Mende et al. (2010) discussed the phylogeography of M. mutica to complement the currently
known scenarios of postglacial re-colonization of Europe. They have evaluated 25 European populations of
the species and five specimens from China. They concluded that there are three genetically different populations in Europe (Italy, the Baltic and North Sea populations). They discussed the contribution of a Far Eastern
refugium to postglacial re-colonization of Europe, however; the discovery of M. mutica in Anatolia makes it
more plausible that an Anatolian population was the
origin of the postglacial European populations. As stated in plak (2008), Anatolia was not covered by an
ice sheet, and during each glacial period a very active
faunal communication has occurred, both between Anatolia and Europe through the Balkans and Thrace, and
between Anatolia and Asia through the Caucasus. On
the other hand, a molecular comparison could possibly
confirm this hypothesis.
Our knowledge about Turkish distribution of M. mutica is quite new, but it is probable that this species and
possibly other species of this genus may inhabit different regions of Turkey. Discovering further populations
of M. mutica, especially from western parts of Anatolia,
will surely enlarge our knowledge. Additional surveys
are needed to reveal the actual aquatic leaf beetle diversity of Turkey.

# 2011 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim

Zoosyst. Evol. 87 (2) 2011, 291 295

References
Askevold, I. S. 1990. Reconstructed phylogeny and reclassification of
the genera of Donaciinae (Coleoptera: Chrysomelidae). Quaestiones Entomologicae 26: 601664.
Bienkowski, A. O. 1996. Life cycles of Donaciinae (Coleoptera: Chrysomelidae). In Jolivet, P. H. & Cox, M. L. (eds). Chrysomelidae
Biology. SBP Academic Publishing, Netherlands: pp. 155171.
Bienkowski, A. O. & Bienkowskaja, M. J. O. 2004. Morphology, systematics and host plants of Palearctic Donaciinae larvae. In Jolivet, P., Santiago-Blay, J. A. & Schmitt, M. (eds). New Developments in the Biology of the Chrysomelidae. SBP Academic
Publishing, Netherlands: pp. 481502.
BioInfo (UK): http://www.bioinfo.org.uk/html/t34145.htm (last visit
15.02.2011).
plak, B. 2008. The analogy between interglacial and global warming for the glacial relicts in a refugium: A biogeographic perspective for conservation of Anatolian Orthoptera. In Fattorini, S.
(ed.). Insect Ecology and Conservation. Research Signpost, India:
pp. 135164.
Ekiz, A. N. & Gk, A. 2010. Taxonomic studies on Donacia Fabricius, 1775 (Coleoptera, Chrysomelidae, Donaciinae) of Southwestern Turkey with notes on their geographic distributions, habitats
and host plant associations. Zoosystematics and Evolution 86
(2): 213219.
Ekiz, A. N., Gk, A., Borowiec, L. & Bilginturan, S. 2009. Redescription of Donacia kraatzi Weise, 1881, a poorly known aquatic leaf
beetle, (Coleoptera, Chrysomelidae, Donaciinae), with notes on
its taxonomic status and geographical distribution. Deutsche
Entomologische Zeitschrift 56: 299303.
Figuerola, J. & Green A. J. 2002. Dispersal of aquatic organisms by
waterbirds: A review of past research and priorities for future studies. Freshwater Biology 47: 483494.
Jch, M. A. & Balke, M. 2008. Global diversity of water beetles (Coleoptera) in freshwater. Hydrobiologia 595: 419442.
Konstantinov, A. S. 2003. Chrysomelidae: Aquatic leaf beetles of China. In Jch, M. A. & Ji, L. (eds). Water Beetles of China (Vol. 2).

# 2011 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim

295

Zoologisch-Botanische Gesellschaft in sterreich and Wiener Coleopterologenverein, Wien: pp. 563572.

Klsch, G., Bistrm, O. & Pedersen, B. V. 2006. Species delimitation
in the leaf beetle genus Macroplea (Coleoptera, Chrysomelidae)
based on mitochondrial DNA, and phylogeographic considerations. Insect Systematics and Evolution 37: 467479.
Klsch, G. & Pedersen, B. V. 2010. Molecular phylogeny of reed beetles (Coleoptera, Chrysomelidae, Donaciinae): The signature of
ecological specialization and geographical isolation. Molecular
Phylogenetics and Evolution 48: 936952.
Lays, P. 2002. Notes on the Donaciines (Coleoptera: Chrysomelidae:
Donaciinae). Bulletin de la Societe Royale Belge dEntomologie 138: 7784.
Mann, J. S. & Crowson R. A. 1983. Observations on the internal anatomy and classification of Donaciinae (Coleoptera, Chrysomelidae). Entomologists Monthly Magazine 119: 1727.
Mende, M., Bistrm, O., Meichssner, E. & Klsch, G. 2010. The
aquatic leaf beetle Macroplea mutica (Coleoptera: Chrysomelidae)
in Europe: population structure, postglacial colonization and the
signature of passive dispersal. European Journal of Entomology
107: 101113.
Mohr, K. H. 1966. Familie: Chrysomelidae. In Freude, H., Harde,
K. W. & Lohse, G. A. (eds). Die Kfer Mitteleuropas Bd. 9.
Goecke & Evers, Krefeld: pp. 95299.
Mohr, K. H. 1985. Beitrge zur Insektenfauna der DDR: Coleoptera
Chrysomelidae: Donaciinae, Orsodacninae, Criocerinae, Clythrinae. Beitrge zur Entomologie 35: 219262.
Nilsson, A. N. 1996. Aquatic insects of North Europe, a taxonomic
handbook (Vol. 1). Apollo Books, Stenstrup.
Silfverberg, H. 2010. Subfamily Donaciinae. In Lbl, I. & Smetana,
A. (eds). Catalogue of Palearctic Coleoptera (Vol. 6). Apollo
Books, Stenstrup: pp. 354358.
Zhang, J., Wheeler, G. S., Purcell, M. & Ding, J. 2010. Biology, distribution, and field host plants of Macroplea japana in China: An
unsuitable candidate for biological control of Hydrilla verticillata.
Florida Entomologist 93: 116119.

museum-zoosyst.evol.wiley-vch.de