J. Limnol., 70(1): 149-155, 2011
J.L. NOTES
DOI: 10.3274/JL11-70-1-18
First record of Hemidiaptomus (Gigantodiaptomus) superbus (Schmeil, 1895) in
Italy, with notes on distribution and conservation status (Copepoda, Calanoida,
Diaptomidae)
Federico MARRONE*, Giuseppe ALFONSO1), Danio MISEROCCHI2) and Sabrina LO BRUTTO
Department of Animal Biology "G. Reverberi", University of Palermo, Via Archirafi 18, 90123 Palermo, Italy
1)
Laboratory of Zoogeography and Fauna, Department of Biological and Environmental Sciences and Technologies (Di.S.Te.B.A.),
University of Salento, 73100 Lecce, Italy
2)
Museo Civico di Storia Naturale di Ferrara, Via De Pisis 24, 44121 Ferrara, Italy
*e-mail corresponding author: federico.marrone@neomedia.it
ABSTRACT
Hemidiaptomus (Gigantodiaptomus) superbus (Schmeil, 1895) was found in a temporary pool in the Pineta di Classe coastal
pine-wood (Ravenna, northern Italy). This is the first official record of a species belonging to the subgenus Gigantodiaptomus of
Italian fauna, and it widens the known distribution of this rare diaptomid to the Mediterranean basin. In order to supply data on this
rare and poorly known species, original drawings for the Italian population have been provided, and some overlooked
morphological features have been highlighted. In light of the growing evidence of the existence of a number of cryptic species in the
Diaptomidae family, and considering the variability observed in the ornamentation of the fifth pair of female legs, the ribosomal
DNA marker 16S was used to compare the specimens collected in Italy with those from the terra typica of the species. Molecular data
confirmed the strict conspecificity of the two populations being studied. The environmental parameters relating to the new Italian site
are described and compared with the few data currently available in the literature. A careful review of all the published data dealing
with H. (G.) superbus has been performed, and the paucity of recent records is stressed, together with the opportunity to support
dedicated conservation measures for the protection of this species.
Key words: temporary ponds, zooplankton, biogeography, biodiversity, 16S rDNA
1. INTRODUCTION
The Palaearctic diaptomid genus Hemidiaptomus
Sars 1903 includes 19 species, occurring in temporary
water bodies from Mongolia to Spain and Morocco
(Dussart & Defaye 2002; Stepanova 2005a). These species are currently divided into three subgenera, i.e.
Hemidiaptomus s. str. Sars 1903, Gigantodiaptomus
Kiefer 1932, and Occidodiaptomus Borutzky 1991, all
of which are mostly parapatric and present in limited
co-occurrence areas (Kiefer 1978; Dussart & Defaye
2002; Marrone et al. 2010). The subgenus Gigantodiaptomus, a taxon considered of genus rank by some
authors (Stepanova 2005b; Walter 2010), includes five
species occurring in Asia and central Europe. Of these,
Hemidiaptomus (Gigantodiaptomus) superbus (Schmeil
1895) has proved to be the rarest and most enigmatic
species of the group. In spite of its widespread distribution, which covers central/eastern Europe and the Caucasus, few records are currently available and most of
these are old reports lacking iconography and in need of
confirmation.
Hemidiaptomus (G.) superbus has been described by
Schmeil (1895) as 'Diaptomus' superbus, based on
specimens collected from temporary ponds in the 'Biederitzer Busch bei Magdeburg' (Germany) and the draw-
ings published one year later (Schmeil 1896). Kiefer
(1932a, 1932b) transposed it to the subgenus Gigantodiaptomus of the genus Hemidiaptomus, whilst stressing
peculiarities of H. (G.) superbus, which are not typical
of the other species of that subgenus. This attribution is
nowadays widely accepted (e.g., Dussart & Defaye
2002) with the only exception of Einsle (1993), who has
assigned the species to the subgenus Hemidiaptomus s.
str., and Stepanova (2005a), who has raised the subgenus Gigantodiaptomus to genus level.
Its description having been documented, the species
was sighted in Denmark (Jensen 1905 in: Tollinger
1911), the Ukraine (Belousov, 1908), France
(Beuchamp 1918), Algeria (Roy 1924), Croatia and
Serbia (Krmpotić 1925; Ternjej & Stankovic 2007; I.
Ternjej, pers. comm.), Azerbaijan (Weisig 1931), Germany (Elster 1931, Gillandt & Martens, 1975; Gillandt
et al. 1983), Iran (Brehm 1937), and Poland (Błędzki
2008) (Fig. 1). Based on the absence of setae on the
distal part of the endopodites of the fifth pair of female
legs, the H. (G.) superbus populations found in the Caucasus were considered by Brehm (1937) as a different
variety of the European H.(G.) superbus, and the author
gave it the name of 'Hemidiaptomus superbus var. hyrcanensis'. This taxon, accepted by Löffler (1961) and
Kiefer (1978), was later synonymised with H. (G.)
superbus by Dussart & Defaye (2002).
150
F. Marrone et al.
Fig. 1. Occurrence localities of H. (G.) superbus. Sources: 1: Schmeil, 1895; 2: Jensen, 1905; 3: Belousov, 1908; 4: Beuchamp,
1918; 5: Krmpotić, 1925; 6: Weisig, 1931; 7: Gillandt & Martens, 1975; 8, 9: Elster, 1931; 10: Brehm, 1937; 11: Błędzki, 2008; 12:
I. Ternjej, pers. com.; 13: Present work. The record of the species for Algeria (Roy, 1924) is most likely erroneous (see text) and thus
has not been reported on the map.
Unfortunately, most of the reports of the species
were published without drawings, which precludes
checks as to their reliability e.g., Hemidiaptomus (G.)
superbus was officially recorded in the Reghaïa forest,
near Algiers (Roy, 1924), a site where extensive surveys
carried out in subsequent years led to the finding of the
only Hemidiaptomus (Occidodiaptomus) ingens
(Gauthier 1928). Furthermore, Roy & Gauthier (1927)
did not include H. (G.) superbus in their work on the
copepod fauna of Algeria, thus implying that the authors
themselves deemed that the previous record of the species in Algeria was erroneous. Currently, reports of the
species are considered doubtful (Dussart 1989), or have
not been reported at all (Mouelhi et al. 2000; Hamaidi et
al. 2010) in the Reviews dealing with the copepods of
Algeria and the Maghreb.
Given the unreliability of some of the older records,
all those relating to Hemidiaptomus (G.) superbus,
which lack adequate iconography, should be confirmed.
Thus, the presence of the species can only be considered
certain for Germany (Schmeil 1895; Elster 1931, Gillandt & Martens 1975; Gillandt et al. 1983) and the
southern coast of the Caspian Sea (Azerbaijan and Iran)
(Weisig 1931; Brehm 1937; Löffler 1961). No record of
the species has been published for the last seventy years,
with the exception of a single population from Peverstof, Germany (Gillandt & Martens 1975; Gillandt et al.
1983). H. (G.) superbus is currently unanimously considered as one of the rarest Palaearctic diaptomid species (Dussart 1967, 1989; Kiefer 1978; Einsle 1993). By
means of an extensive faunal survey, the aim of which
was to update the checklist and distribution of Italian
inland water copepods, various specimens of H. (G.)
superbus were collected in a temporary pool near
Ravenna (northern Italy).
2. MATERIALS AND METHODS
The study material was collected on 6 June 2010 in
four temporary pools in the Pineta di Classe near
Ravenna, Italy. The sampled sites are shallow, clearwater temporary pools, which are located in a mature
coastal pinewood. The bioclimatic characterization of
the area is based on the classification formulated by
Rivas-Martinez (1995). Water temperature, pH and specific conductivity were registered with a MM40 Crison
multimeter. Microcrustacean samples were collected by
means of a 200 μm mesh-sized hand net and fixed in
situ with 80% ethanol. Calanoid copepods were isolated, dissected under the stereomicroscope and identified to the species level, according to Kiefer (1978),
Borutzky et al. (1991) and Einsle (1993). Co-occurring
Cladocera, Cyclopoida, and Ostracoda were identified
according to Margaritora (1985), Alonso (1996), Meisch
(2000), Einsle (1993), as updated following the recent
literature. Drawings of selected specimens (total length,
females: 4200 μm; males: 3400 μm) were made with a
Zeiss® Standard 18 compound microscope, which is
equipped with a camera lucida, after tissue cleaning
with in hot lactic acid. Preserved specimens and
mounted slides have been stored in the authors' collection.
Various specimens of Hemidiaptomus (G.) superbus
and environmental data from a temporary pond near the
town of Breitenhagen (Germany) have been included in
the study as comparative material. This site (coordinates
UTM-WGS84 32U: 5756674 N; 703109 E; Altitude: 53
m a.s.l.) was sampled on 4 April 2008, its location being
the same river basin of the type locality of the species
(i.e., the Elbe River basin), and approximately 30 km
from its locus typicus. The site was rich in water
First record of Hemidiaptomus (Gigantodiaptomus) superbus in Italy
macrophytes, water temperature was 12.1 °C, and specific conductivity was 1430 μS cm-1 on the sampling
date. Adult specimens of Hemidiaptomus (G.) superbus
and Hemidiaptomus (G.) amblyodon (Marenzeller 1837)
specimens were collected at the same time.
In light of the growing evidence of the presence of a
cryptic species among diaptomid calanoid copepods
(e.g., Thum & Harrison 2009) and within the genus
Hemidiaptomus in particular (Marrone et al. 2010) and
in order to check for the molecular divergence between
the populations being studied, a fragment of the 16S
ribosomal mtDNA was amplified from a single H. (G.)
superbus specimen per population. Prior to DNA
extraction, specimens were carefully cleaned and
soaked in double-distilled water for 3 hours. DNA was
then extracted using whole specimens and the "DNEasy
Animal Tissue Kit" (QIAGEN). A partial sequence of
the 16S ribosomal DNA subunit was then amplified
using the "Skisto-1" (5' –TGG TAA GGT AGC ATA
ATA AT- 3') e Skisto-2 (5'- CCG GTT TGA ACT CAG
ATC ATG T -3') primers, as described by Thum & Harrison (2009).
The PCR mix consisted of 3.5 μl of Buffer 10×
(THERMOSCIENTIFIC), 3 μL of 25 mM MgCl2, 0.35
μL of dNTP 10 mM, 1 μL of each 10 μM primer, 0.4 μL
of Red Hot Taq Polymerase 5U/μL (THERMOSCIENTIFIC), 1 μL of DNA template, and 23.7 μL of
double-distilled water, producing a total reaction volume of 35 μL. The amplification consisted of an initial
denaturation step of 94 °C for 4 minutes, followed by 35
cycles of 94 °C for 50 seconds, 42 °C for 50 seconds,
and 72 °C for 1 minute, followed by a final extension at
72 °C for 8 minutes. The PCR products were separated
by electrophoresis on 2% agarose gel at 70 V for 1 h
and visualized with a UV Transilluminator. Amplified
bands were cut, purified from agarose gel (using the
Qiaquick Gel Extraction Kit) (QIAGEN) and sequenced
in forward and reverse directions with a ROCHE
GENOME SEQUENCER FLX. Chromatograms were
imported and edited with Chromas Lite 2.01
(TECHNELYSIUM PTY LTD) and exported to be
aligned with BioEdit (Hall 1999); sequence divergence
was calculated with Mega ver. 4 (Tamura et al. 2007).
The sequences have been deposited in GenBank
(Accession numbers: HQ687215 and HQ687216).
3. RESULTS AND DISCUSSION
A population of Hemidiaptomus (G.) superbus has
been located in Italy approximately three decades after
the last published record of the species (Gillandt et al.
1983). The morphology of the Italian specimens correlates well with the description of the species currently
available in the literature (Kiefer 1978; Borutzky et al.
1991; Einsle 1993). However, the presence of a small
chitinous hump on the dorsal side of the second segment
of the exopodite of fifth, right, male leg (Fig. 2H),
which was present in all the studied male specimens,
151
has not to date been reported in the published drawings.
Within the Gigantodiaptomus subgenus, this chitinous
outgrowth is reported to occur in all the species with the
only exception of H. (G.) hungaricus (Kiefer 1978; Stepanova 2005a).
Furthermore, the length of the apical setae on the
endopodite of fifth female pair of legs (Fig. 2B) is
slightly shorter than that reported in the available
drawings of German specimens (Gillandt & Martens
1975; Kiefer 1978; Einsle 1993). This characteristic is
noteworthy as the 'hyrcanensis' variety of H. (G.) superbus was described as lacking these setae in the specimens originating from the Caucasus (Brehm 1937). It
thus seems that a certain variability exists in the ornamentation of the fifth pair of female legs, which also
concerns the presence of a trasversal row of spinules on
the apical part of the second exopodal segment of the
fifth leg: these spinules are present in the German and
Italian specimens (e.g., Fig. 2B) but they are absent in
the specimens from the Balkans (Krmpotić 1925).
A 350 base-pairs long fragment of the ribosomal
DNA subunit 16S was successfully amplified and
sequenced. The uncorrected p-distance between the
amplified sequences is just 0.9% (Fig. 3), while it often
accounts for 14.9-23% among congeneric diaptomid
species and even among distinct presumed intraspecific
clades (Thum & Harrison 2009). Thus, molecular data
confirm the conspecificity of the two populations under
study and the existence of a certain intraspecific variability in the ornamentation of the fifth pair of female legs.
The co-occurring microcrustacean fauna, collected
in the Pineta di Classe pools (Tab. 1), consists of species
which are widespread in Italy and often related to
slightly brackish water bodies (Marrone et al. 2006a;
Stoch 2006; pers. obs. of the authors). Hemidiaptomus
(G.) superbus was collected in only one out of the four
sampled sites (i.e. 'pool A') on the sampling date but
large diaptomids, probably H. (G.) superbus, had also
been observed in the other pools in May 2010 (D. Miserocchi, personal observation). Coordinates, water temperature, pH and the specific conductivity of the sampled pools are reported in table 2. The pools were a few
square metres in width on the sampling date and the
only observed macrophytes were Alisma lanceolata
With. and Galium palustre L.
According to Rivas-Martinez (1995), both the Italian
and the German areas, where H. (G.) superbus occur, lie
in the temperate-oceanic bioclimatic area. However, all
the other Hemidiaptomus species occurring in Italy, i.e.
Hemidiaptomus (Hemidiaptomus) gurneyi, H. (Occidodiaptomus) ingens s.l., and H. (O.) roubaui (Ruffo &
Stoch 2006; Marrone et al. 2006b; Marrone et al. 2010;
Alfonso & Belmonte 2011) occur in areas characterised
by a Mediterranean pluviseasonal, oceanic bioclimate.
Unfortunately, the scarce and anecdotal data currently
available for Hemidiaptomus (G.) superbus do not clarify the ecological preferences of the species.
152
F. Marrone et al.
Fig. 2. A-C: Hemidiaptomus superbus, female. A: somites 4-5 and urosome; B: fifth pair of legs; C: antennula, segments 1-7. D-H:
H. superbus, male. D: left antennula, segments 1-6; E: somite 5 and genital segment; F: right antennula, segments 1-17; G: right
antennula, last three segments; H: fifth pair of legs. Scale bars: 500 μm.
Tab. 1. List of crustaceans occurring in the sampled pools on 6 June 2010.
Taxa
Pool A
ER003
Pool B
ER004
Pool C
ER005
Pool D
ER006
Copepoda
Calanoida
Diaptomidae
Hemidiaptomus (Gigantodiaptomus) superbus (Schmeil, 1895)
Mixodiaptomus kupelwieseri (Brehm, 1907)
X
X
X
X
X
Copepoda
Cyclopoida
Cyclops divergens (Lindberg, 1936)
Megacyclops viridis (Jurine, 1820)
Diacyclops bisetosus (Rehberg, 1880)
Diacyclops lubbocki (Brady, 1869)
X
X
X
X
X
X
X
X
X
X
X
X
X
Branchiopoda
Anomopoda
Daphniidae
Scapholeberis rammneri (Dumont & Pensaert, 1983)
Ceriodaphnia laticaudata (P.E. Müller, 1867)
Daphnia (Daphnia) curvirostris (Eylmann, 1887)
X
X
X
Ostracoda
Podocopa
Candonidae
Unidentified larvae
Cyprididae
Eucypris virens 'complex' (Jurine, 1820)
Heterocypris salina (Brady, 1868)
X
X
X
X
X
X
X
X
First record of Hemidiaptomus (Gigantodiaptomus) superbus in Italy
153
Fig. 3. Compared 16S rDNA nucleotide sequences for the Pineta di Classe (Ravenna, Italy) and Breitenhagen (Salzlandkreis,
Germany) populations.
Tab. 2. Coordinates, water temperature, pH and water conductivity of the sampled pools on 6 June 2010.
Coordinates
(UTM-WGS84)
Pool A – ER003
Pool B – ER004
Pool C – ER005
Pool D – ER006
4913408 N; 33 283404 E
4913413 N; 33 283411 E
4913411 N; 33 283426 E
4913418 N; 33 283442 E
However, a rather high degree of specific conductivity was recorded for both the Italian and German
sites, which is quite unusual for most Hemidiaptomus
species (e.g., Gauthier 1928; Alonso 1998; Marrone et
al. 2006a).
The scattered geographical and temporal distribution
of the records of occurrence of H. (G.) superbus might
be ascribed to three different reasons, which are not
mutually exclusive: (i) the species could be naturally
rare in nature; (ii) the distribution barycentre of H. (G.)
superbus might indeed occur in relatively unknown
regions of south-eastern Europe and the Caucasus, thus
being present in western Europe only with few, shortliving populations which episodically colonise sites
along the westernmost fringe of the species' distribution
area; and (iii) the species could have experienced a total
collapse in recent decades, having been replaced by
other, more generalist, copepod species. The latter
hypothesis is supported by the fact that, compared to
nine records published between 1895 and 1937
(Schmeil 1895; Jensen 1905; Belousov 1908;
Beauchamp 1918; Roy 1924; Krmpotić 1925; Weisig
1931; Elster 1931; Brehm 1937), only two records of
Hemidiaptomus (G.) superbus were published after
1895 and both of them regarded the same area near the
town of Pevestorf, Germany (Gillandt & Martens 1975;
Water Temperature Specific conductivity
(°C)
(μS cm-1)
24
25
25.2
24.1
1433
864
1329
768
pH
8.04
7.89
7.94
7.95
Gillandt et al. 1983). The occurrence of the species near
Breitenhagen (Germany), which was reported by Elster
(1931), has been confirmed in the ambit of this study;
however, it was not possible to find H. (G.) superbus in
the locus typicus of the species (Schmeil, 1895) in the
Magdeburg area, which is the other site reported by the
same author (Elster 1931). Furthermore, the European
distribution area of H. (G.) superbus overlaps with that
of the more common and widespread congeneric species
Hemidiaptomus (G.) amblyodon (Dussart & Defaye
2002), which could have replaced the H. (G.) superbus
in various sites. This process might be ongoing, e.g.
these two Hemidiaptomus species currently co-occur in
the Breitenhagen pond, both syntopically and synchronically, while Elster (1931) reported for the same
site the occurrence of H. (G.) superbus only.
It is necessary to confirm the current presence of the
species for most of the countries where it has been
recorded, and it would be desirable to check the historical and current occurrence of H. (G.) superbus throughout its known distribution area. Until more data regarding the distribution and consistency of the species in
Europe are available, dedicated, protective measures of
the few, confirmed known occurrence sites of the species (i.e., the pools in the Pineta di Classe in Italy, and
the ponds near Breitenhagen and Pevestorf in Germany)
154
should be implemented, and H. (G.) superbus should be
included on the national 'red lists' for invertebrates (e.g.,
Herbst 1982).
The presence of Hemidiaptomus (Gigantodiaptomus) superbus in Italy, which constitutes the first
Hemidiaptomus record from the Padanian faunal province (Minelli et al. 2006), has increased to 26 the number of diaptomid species currently known to occur in
Italy (Stoch 2006), and it confirms Italy as one of the
European countries with the richest diaptomid copepod
fauna (Dussart & Defaye 2002). This finding is rather
unexpected since no other species belonging to the
Gigantodiaptomus subgenus have been reported south
of the Alps and Pyrenées (see Marrone et al. 2010 for a
review). The Italian finding further stresses the considerable cognitive gaps which affect our knowledge
relating to particular taxa, even in the assumed betterknown countries in the west-Palaearctic area.
ACKNOWLEDGEMENTS
The authors of this paper wish to particularly
acknowledge the valuable support of M. Korn (Senckenberg Natural History Collection, Dresden, Germany),
who kindly provided Hemidiaptomus (G.) superbus
specimens from Breitenhagen. We are also grateful to I.
Ternjej (University of Zagreb, Croatia) and L. Samchyshina (Center of Megalopolis Ecomonitoring and Biodiversity Research NASU, Ukraine) for their stimulating
discussions and for having provided important articles
from the relevant literature. G. Rossetti (University of
Parma, Italy) and F. Stoch (University of L'Aquila,
Italy) kindly identified the ostracods and the cyclopoid
copepods respectively. Finally, we recognise the contribution of Dr. A. Vistoli (Ufficio Ambiente e Sostenibilità, Comune di Ravenna) for having provided
Permission to sample in the SIC ITA4070010 Pineta di
Classe to DM. This study was supported by the University of Palermo, Italy (Fondi d'Ateneo, ex 60%)
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