jmjmpaleoJournal of Micropalaeontology0262-821X2041-4978The Geological Society of London10.1144/jmpaleo2015-022jmpaleo2015-022Thematic set: Cyprideis torosaNew occurrences of Cyprideis torosa (Crustacea, Ostracoda) in GermanyScharfBurkhard1*HerzogMichael2PintAnna3Ellhornstrasse 21, 28195 Bremen, GermanyBerliner Strasse 24, 39218 Schönebeck, GermanyGeographisches Institut, Universität zu Köln, Albertus Magnus Platz, 50931 Köln, GermanyCorrespondence: burkhard.w.scharf@t-online.de
Living Cyprideis torosa (Jones, 1850) was found in Germany along the North Sea coast and on some islands, within the Baltic Sea and in coastal waters near the Baltic Sea, but also in some inland natural and anthropogenic saline waters. The natural inland waters were probably colonized by birds because this species was found only in permanent saline shallow lakes that provide resting places for migrating birds. Information on the fossil and Recent record and on the biology of C. torosa and its accompanying ostracod fauna is discussed.
Two kinds of saline inland waters are distinguished: thalassic and athalassic waters. Thalassic waters have an ionic composition similar to that of seawater, athalassic waters have a quite different ionic composition from those with dissolved salts derived from seawater and have never been connected to the sea during geologically Recent times (Cole 1983). Brackish waters must have a connection to the sea (De Deckker 1981). Here we provide two examples for athalassic waters in Germany. Some of the inland saline waters in Germany have their origin in the solution of a salt dome underground by groundwater (Fig. 1). Lake Arendsee, northeastern Germany, Sachsen-Anhalt, is an example of such a lake (Fig. 2). During Late Permian time thick layers of salt from a desiccating sea were deposited. The salt became thermoplastic under high pressure and temperature and ascended via fault zones through the cover deposits. Springs with water that had contact with a salt dome are more or less saline. These are natural saline athalassic inland waters. Thus, solution of the top of the salt dome can impact the formation of a lake over the salt dome (Stottmeister 1998; Leineweber et al. 2009; Scharf et al. 2009).
Position of the salt dome under Lake Arendsee (after Stottmeister 1998, modified). Lower Permian to Quaternary sediments, depths (metres) below surface (mbs), the depths of boreholes between Arendsee and Bockleben SW of Arendsee are presented. The saline (evaporite) deposits of the Upper Permian are marked by a bold line.
Location map. B+J denotes villages Barnstorf and Jerxheim with salt marshes; Sal. S denotes Salziger See; 4–9 are the sample sites in the vicinity of Sülldorf village.
Since 1889, a potash factory (Salz- und Kalihersteller K+S in Kassel, Germany) has extracted potash from the salt of a salt dome and discharged its saline waste water into the River Werra and consequently into the River Weser and the North Sea (Fig. 2). This is an anthropogenic saline inland water.
Cyprideis torosa (Jones, 1850) occurs mainly in thalassic brackish waters with fluctuating salinity, from hypersaline to freshwater in range. It is found in brackish coastal waters of Europe, western and central Asia, the Mediterranean region, the Middle East, lakes in central Africa (Meisch 2000), in the Seychelles (Wouters 2002) and South Africa (Wouters 2003).
The aim of this study is to show where Cyprideis torosa can be found in Germany, especially in inland waters of Germany, to compile information on the life conditions of C. torosa and on the accompanying ostracod fauna at sample sites with C. torosa or at these samples sites where we expected to find C. torosa.
Material and methods
The data of the sampled sites are compiled in Table 1. Waters of sites 1–2 are situated on islands in the North Sea, site 3 in the River Peene which connects the Oder lagoon and the Baltic Sea, sites 4–9 are located in the biggest saline spring area of NE Germany, at Sülldorf near Magdeburg (Fig. 2), sites 10–16 are situated in the River Weser's floodplain or in this river itself downstream of the city of Bremen. In Bremen there is a weir in the River Weser which stops the influence of the North Sea, the end of the estuary. Site 17 is a harbour in the River Weser, c. 25 km upstream (south) of Bremen (Fig. 2). Sites 18–21 are ditches or ponds near natural inland salt marshes, north of the Harz mountain range (Janssen 1986; Janssen & Brandes 1989a, b).
Sample sites
Site
Water body
Coordinates
Date
Temperature (°C)
Salinity (psu)
Conductivity (µS cm–1)
Remarks
1
Pond
53° 38′ 20″ N06° 52′ 11ʺ E
1976–87
750–2500m = 1257 (n = 13)
New house pond on the North Sea island Memmert
2
Pond
53° 42′ 51″ N08° 08′ 47″ E
19 July 1974
House pond on the North Sea island Mellum
3
Harbour
54° 02′ 48″ N13° 46′ 48″ E
7 September 1995
3.5
Harbour of Wolgast, River Peene
4
Brook
52° 01′ 23″ N11° 33′ 09″ E
4 October 2014
14
4
3190
Sülldorf: effluent of a spring
5
Pond
52° 01′ 24″ N11° 33′ 09″ E
25 August 2014
77 400
Sülldorf: limnocrene with a diameter of c. 3 m with saline spring.
5
4 October 2014
11
50
6
Spring
52° 01′ 23″ N11° 33′ 09″ E
4 October 2014
11
67
Sülldorf: limnocrene with a diameter of 0.3 m
7
Brook
52° 01′ 23″ N11° 33′ 09″ E
4 October 2014
15
5
3240
Sülldorf: fast-flowing brook Sülze with sand substrate and many submerged macrophytes at the shore
8
Pond
52° 01′ 29″ N11° 33′ 26″ E
25 August 2014
44 900
Sülldorf: artificial pond near the brook Sülze, thick layer of biogenically precipitated small calcite plates
8
4 October 2014
19
33
9
Pond
52° 01′ 30″ N11° 33′ 28″ E
25 August 2014
44 900
Sülldorf: same pond as 8, thick layer of biogenically precipitated small calcite plates
10
Ditch
53° 19′ 35″ N08° 31′ 24″ E
21 October 2012
Ditch in the floodplain of the River Weser
10
4 August 2013
26
1560
11
Pit
53° 16′ 00″ N08° 30′ 48″ E
5 May 2012
13
1046
Loam pit in River Weser flood plain
12
Pit
53° 11′ 22″ N08° 30′ 59″ E
13 December 2009
5
1277
Loam pit in River Weser floodplain
12
18 April 2010
18
1453
12
4 September 2010
17
1260
13
Ditch
53° 11′ 22″ N08° 30′ 59″ E
13 December 2009
5
1277
Effluent of pit 12
13
4 September 2010
17
1260
14
Ditch
53° 11′ 25″ N08° 30′ 52″ E
4 September 2010
19
1073
c. 150 m downstream of ditch 13
15
River arm
53° 11′ 03″ N08° 31′ 46″ E
13 December 2009
7
1205
Arm of the River Weser
16
River
53° 04′ 08″ N08° 48′ 52″ E
17 October 2010
12
1284
River Weser at Bremen
17
Harbour
52° 59′ 45″ N09° 03′ 04″ E
7 March 2014
9
1330
Harbour of the River Weser at Achim-Uesen
17
12 July 2014
23
1282
18
Ditch
52° 06′ 02″ N10° 48′ 33″ E
18 July 2015
19
2
1634
Barnstorf: effluent of a salt marsh
19
Spring
52° 06′ 01″ N10° 48′ 34″ E
18 July 2015
12
3
2370
Barnstorf: spring within ditch 18
20
Pond
52° 06′ 01″ N10° 48′ 47″ E
18 July 2015
24
5
6140
Barnstorf: artificial pond for fishing
21
Pond
52° 04′ 15″ N10° 55′ 11″ E
18 July 2015
25
8
9100
Jerxheim: artificial pond below the salt marsh ‘Seckertrift’, dug 10 to 15 years ago (C. Evers, Braunschweig, pers. comm.)
The method of collecting and separating the living ostracods from the surrounding sediment is described by Scharf et al. (2014). The sampled area was mostly c. 0.25 m2. At each site, the temperature was measured with a standard mercury thermometer, the conductivity with a Multi-Parameter PCTestrTM 35 of EUTECH Instruments OAKTON®, the salinity with the refractometer REF 211 (Arcarda® GmbH), the pH-value colourimetry (pH-indicator solution pH 4.0–10.0, MERCK®), and the coordinates were taken by means of Garmin GPSmap76 (WGS 84).
The ‘Index and bibliography of nonmarine/marine Ostracoda’ by E. K. Kempf (see Viehberg et al. 2014) was used intensively and species listed in this work can be found in this publication. The ostracods are deposited in the collection of BS.
Results
The abundance of Cyprideis torosa and the accompanying ostracod fauna can be found in Table 2. The most specimens of C. torosa were found in locality 12, a loam pit in the floodplain of the River Weser with more than 2000 individuals per sample. In a ditch in the estuary's floodplain of the River Weser 450 individuals were present in a sample (10). In all other sample sites the number of C. torosa was fewer.
Ostracod distribution
Site
Date
Cyprideis torosa
Accompanying ostracods and [reference]
males
females
juveniles
1
19 July 1974
1
5
14
Heterocypris salina III, Plesiocypridopsis newtoni IV, Potamocypris unicaudata II, Sarscypridopsis aculeata IV [Hollwedel & Scharf 1988]
2
17 June 1970
(1)
Candona candida II, Cyclocypris ovum IV, Eucypris virens II, Notodromas monacha I [Hollwedel & Scharf 1988]
3
7 September 1995
5
15
2
Candona neglecta (I), Darwinula stevensoni III, Limnocythere inopinata I
4
4 October 2014
Candona neglecta I, Ilyocypris bradyi II, Prionocypris zenkeri (I), Pseudocandona albicans I
Heterocypris incongruens (I), H. salina (II), Ilyocypris sp. (II)
10
21 October 2012
32
29
74
Candona neglecta IV, Cypria ophtalmica II, Darwinula stevensoni I, Fabaeformiscandona levanderi (I), Ilyocypris sp. I, Prionocypris zenkeri (II)
10
4 August 2013
90
110
250
Candona neglecta II, Cypria ophtalmica I, Darwinula stevensoni II, Ilyocypris bradyi I, I. monstrifica I
11
5 May 2012
1
Candona candida (I), C. neglecta IV, Cypria ophtalmica III, Cypridopsis vidua I, Cypridopsis. sp. II, Eucypris virens (I), Fabaeformiscandona levanderi (I), Ilyocypris bradyi (I), I. monstrifica (I), Plesiocypridopsis newtoni (I), Potamocypris smaragdina II, Sarscypridopsis aculeata I
12
13 December 2009
28
50
68
Candona candida I, C. neglecta IV, Candonocypris novaezelandiae I, Cypria ophtalmica I, Cypridopsis vidua II, Darwinula stevensoni I, Herpetocypris reptans (I), Ilyocypris decipiens II, I. monstrifica II, Physocypria kraepelini IV, Prionocypris zenkeri (I), Stenocypria fischeri (I)
12
18 April 2010
600
650
1050
Candona candida II, C. neglecta V, Cypria ophtalmica (I), Cypridopsis vidua I, Candonopsis sp. I, Darwinula stevensoni II, Herpetocypris chevreuxi (I), Ilyocypris bradyi I, I. sp. (I), Isocypris beauchampi I, Limnocypthere inopinata (I), Physocypria kraepelini V, Prionocypris zenkeri I, Stenocypria fischeri (I)
12
4 September 2010
4
10
2
Candona neglecta II, Candonocypris novaezelandiae (I), Cypria ophtalmica III, Cypridopsis vidua II, Herpetocypris reptans (I), Ilyocypris bradyi I, I. decipiens I, I. monstrifica IV, Notodromas monacha (I), Physocypria kraepelini I
13
13 December 2009
4
9
17
Candona neglecta IV, Cypria ophtalmica IV, Cypridopsis vidua II, Herpetocypris chevreuxi I, Ilyocypris decipiens (I), Isocypris beauchampi I, Limnocythere inopinata (I), Notodromas monacha (I), Physocypria kraepelini II, Prionocypris zenkeri (I)
13
4 September 2010
3
3
2
Candona neglecta III, Cypria ophtalmica II, Cypridopsis vidua (I), Herpetocypris reptans (I), Ilyocypris bradyi I, I. decipiens I, I. monstrifica IV, Limnocythere inopinate (I), Physocypria kraepelini II, Prionocypris zenkeri (I)
14
4 September 2010
1
Candona neglecta II, Cypria ophtalmica I, C. subsalsa I, Cypridopsis vidua II, Herpetocypris reptans I, Ilyocypris bradyi (I), I. monstrifica I, Notodromas monacha (I), Physocypria kraepelini I, Pseudocandona sp. (I), Stenocypria fischeri I
15
13 December 2009
(3)
Candona neglecta I, Cypria ophtalmica I, Cypria subsalsa III, Cytheromorpha fuscata (I), Darwinula stevensoni I, Ilyocypris bradyi I, Leptocythere ilyophila II, Physocypria kraepelini I, Pseudocandona sp. II
16
17 October 2010
(1)
Cypria subsalsa III, Cypridopsis vidua (I), Darwinula stevensoni I, Limnocythere inopinata I, Ilyocypris bradyi II, I. decipiens (II), Pseudocandona sp. (I), Psychrodromus sp. (I)
17
7 March 2014
12
Candona candida (I), C. neglecta IV, Cypria ophtalmica II; C. subsalsa II, Dawinula stevensoni II, Fabaeformiscandona levanderi II, Ilyocypris bradyi (I), I. monstrifica (I), Limnocythere inopinata (I), Physocypria kraepelini (II), Psychrodromus olivaceus (I)
17
12 July 2014
12
Candona neglecta III, Cypria ophtalmica III, C. subsalsa II, Cypridopsis vidua (1), Darwinula stevensoni II, Limnocythere inopinata II, Physocypria kraepelini (II), Prionocypris zenkeri I
18
18 July 2015
Candona candia II, Cypria opthalmica I, Ilyocypris inermis I, Prionocypris zenkeri (III)
19
18 July 2015
Candona candiada I, Cypria ophtalmica I, Ilyocypris inermis I, Potamocypris sp. (I), Prionocypris zenkeri (IV), Pseudocandona sp. (II)
20
18 July 2015
Cyclocypris ovum IV, Cypria ophtalmica I, Cypridopsis vidua (I), Herpetocypris chevreuxi III, Notocromas monacha (I), Sarscypridopsis aculeata IV
21
18 July 2015
Heterocypris salina IV, Ilyocypris gibba II, Sarscypridopsis aculeata IV
Same sampling sites as Table 1. Abundance of the accompanying fauna: I = 1–3 individuals per sample; II = 4–10 ind.; III = 11–25 ind.; IV = 26–100 ind.; V = >100 ind.; () denotes only subrecent records.
Near Magdeburg is the biggest saline spring of northern Germany (sites 4–9 in Tables 1 and 2). Cyprideis torosa was found at this locality only at sites 8 and 9 and there not alive. Here the effluent of the saline spring is dammed (Fig. 3a, d). The pond was hypereutrophic (many submerged macrophytes and slight smell of H2S) and this is probably the reason why only empty carapaces and valves of C. torosa were present in August and October 2014. The number of dead animals shows that there was an important population of this species. Also, none of the accompanying species was found living (Table 2). At sample sites 15–17 only juveniles were found.
Some studied localities: (a) aerial view of the saltmarsh of Sülldorf, 4–9 are the sample sites (see Fig. 2); (b) sample sites 4–7, in the background the red prostrate glasswort Salicornia ramosissima; (c) sample site 4; (d) pond with sample sites 8 and 9; (e) aerial picture of the saltmarsh with sample sites 18–20; (f). aerial picture of the salt marsh with sample site 21. (a)–(d) from Google Earth 2015.
Among the accompanying species only the following show an abundance of more than 26 individuals per sample (abundance: IV and V): Candona neglecta (localities 10–13, 17), Cyclocypris ovum (locality 2), Cypria ophtalmica (13), Heterocypris salina (21), Ilyocypris monstrifica (12), Physocypria kraepelini (12), Plesiocypridopsis newtoni (1), Prionocypris zenkeri (5, 7, 19), Sarscypridopsis aculeata (1, 20, 21).
DiscussionQuaternary findings of Cyprideis torosa in Germany
Fossil Cyprideis torosa have been recorded from 45 inland localities in Germany, 32 of them at a distance of more than 200 km away from the coasts of the Baltic and the North seas. The species occurred especially during interglacial periods, including the Holocene, and can be used as an indicator for palaeosalinity and palaeoclimate (Gramann 2000; Frenzel et al. 2012, Pint et al. 2012, 2015).
Historical and new findings of living Cyprideis torosa in Germany
In the literature there are some records on the occurrence of Cyprideis torosa in Germany. Klie (1938) mentions the following findings of C. torosa:
Coastal occurrence: Borkum (island within the North Sea), Kolberger Heide (salt marsh at the Baltic Sea near Kiel), Schlei (estuary at the Baltic Sea, 25 km NW of Kiel), Waterneverstorfer See (lake, c. 30 km east of Kiel near the Baltic Sea), Bay of Wismar (Baltic Sea), Saaler Bodden (lake connected with the Baltic Sea, near the island Rügen), Greifswald (Baltic Sea), Frisches Haff (lagoon near Baltic Sea, 50 km east of Gdansk, since 1945 in Poland).
Inland occurrences: Mansfeld (with the lake Salziger See (Fig. 2) and lake Süßer See; lake Salziger See was drained in 1892 and C. torosa disappeared at this locality), Oldesloe (saline spring, c. 20 km west of Lübeck (Klie 1925)), and recorded from the Glockensee, near Bad Laer, c. 120 km west of Hannover, Lower Saxony (Gramann 2000). Freshwater lakes mentioned by Klie (1938): Gruber See (lake with a connection to the Baltic Sea, subsequently drained, c. 40 km north of Lübeck), Trammer See (periphery of Plön; Trammer See is situated over a salt dome). Ponds and lakes of central Germany have no evidence for living individuals of C. torosa, only empty valves of C. torosa are present in waters with higher salinity. Five localities are mentioned by Pint et al. (2015): Gründelsloch near Bilzingsleben, the artificial lake of Kelbra, the pond of Stotternheim, Bindersee, and Süßer See.
Cyprideis torosa does not occur in the North Sea (Vesper 1972a), but in ditches along the North Sea coast and on islands within the North Sea (Vesper 1972a; Hollwedel & Scharf 1988; Scharf & Hollwedel 2010). This species is very frequent in the Sehlendorfer Binnensee near the Baltic Sea ([54° 18′ 25″ N, 10° 40′ 59″ E]; Rosenfeld & Vesper 1977; Rosenfeld 1979) and in the shallow brackish waters (called ‘Bodden’) around the island of Rügen within the Baltic Sea (Schäfer 1953; Frenzel 1991, 1996; Frenzel & Oertel 2002; Frenzel & Viehberg 2004; Frenzel & Boomer 2005; Frenzel et al. 2005, 2010; Borck & Frenzel 2006). Many studies on C. torosa, especially concerning nodding, were performed in the Schlei estuary, connected to the Baltic Sea (Jaeckel 1962; Vesper 1972b; Keyser & Aladin 2004).
Cyprideis torosa is probably transported by ship from the estuary of the River Weser to the harbour on the river at Achim-Uesen (site 17) with its artificially increased salinity from the waste water of the potash factory. The finding of C. torosa in lake Gemündener Maar (Scharf 1980) is probably due to a collecting net not thoroughly cleaned (B.S. opinion; the previously collected sample was from a locality in Greece with many specimens of C. torosa). The report of C. torosa in lake Gemündener Maar due to passive transport by migratory birds and unsuccessful colonization (Scharf 1980) is probably an error.
Biology of Cyprideis torosa
There are two forms of Cyprideis torosa. (e.g. Frenzel & Oertel 2002). The valves of the first form show nodes that are absent in the second one. The formation of nodes depends on the salinity of the water (e.g. Vesper 1972b; Frenzel & Boomer 2005; Frenzel et al. 2012; Pint et al. 2012, 2015) and the calcium-ion content of the water (Frenzel et al. 2012) results in a failure of osmoregulation (Keyser & Aladin 2004).
Experimental studies by Frenzel et al. (2012) have shown that noding occurs below a salinity of 14 psu and, therefore, often occurs in freshwater and brackish habitats. Cyprideis torosa occurs above a salinity of 0.5 psu, but also in hypersaline lagoons and lakes. In these environments it often appears monospecifically. The noded form reaches 10% in the salinity range from 2–7 psu and is dominant below 2 psu (Frenzel et al. 2012). It is holeuryhaline and polythermophilic (0–26, rarely up to 32°C). The larval development starts above 12°C. One can find the species in very shallow to shallow waters. It is oligorheophilic. It occurs in estuaries, ponds and lagoons and salt marshes. It prefers mud as a sediment substrate and lives endobenthically and epibenthically. It tolerates low oxygen (data from Frenzel et al. 2010). Cyprideis torosa was found at a pH range between 7.5 and 8.5 on the island Terschelling in The Netherlands (Scharf & Hollwedel 2010). Cyprideis torosa is a brackish-water species (Meisch 2000), it does not occur in freshwater or in strictly marine environments, e.g. the North Sea (Vesper 1972a; Pint et al. 2015).
The species shows a high tolerance to low oxygen and high hydrogen sulphide values in the surrounding water (Gamenick et al. 1996; Jahn et al. 1996). Cyprideis torosa oxidizes sulphide to non-toxic thiosulphate and sulphite and eliminates the oxidation products rather quickly (Jahn et al. 1996). Additionally, the species is able to switch over at anaerobiosis (Jahn et al. 1996). These characteristics allow C. torosa to survive in an environment with often low oxygen content and high hydrogen sulphide concentration, e.g. if the sediment is covered by the macrophyte Fucus vesiculosus. Cyprideis torosa appears as the first immigrant in areas that were earlier deoxygenated and with high content of hydrogen sulphide (Gamenick et al. 1996).
Remarks on accompanying ostracods in inland saline waters
In the area of Sülldorf, near Magdeburg, there are many small and big saline springs (Table 1, Figs 2 and 3). They have different salinities: in October 2014 the salinity at sample site 4 was 4 psu; at 5 it was 50 psu; at 6, 67 psu; at 7, 5 psu; at 8 and 9, 33 psu. At site 6, no ostracods were present; site 4 is the effluent of a rheocrene with low discharge. Some ostracod species (Candona neglecta, Ilyocypris bradyi, Prionocypris zenkeri, Pseudocandona albicans) could be found in the short distance (c. 5 m) between the spring and pond 5 (Table 2). These species are known to tolerate waters flowing from springs and to tolerate a slight increase in salinity (Meisch 2000). Site 5 is a pond with a strong underwater spring. One can see the sand moving over this spring. It was surprising to find 20 carapaces and 68 valves of Prionocypris zenkeri, all dead, but some with soft parts. This species lives in freshwater and prefers slowly flowing waters (Meisch 2000). Ostracods of flowing waters must move upstream, because the drift transports them downstream. We assume that P. zenkeri moves up from the brook Sülze (site 7) into the discharge of pond 5, and that they die eventually when they arrive in the pond, caused by the high salinity in the pond.
The famous saline springs north of the Harz Mountains were also studied: springs near the village of Barnstorf (springs and discharge of a salt marsh; sites 18–20) and pond 21 in the salt marsh ‘Seckertrift’ near the village of Jerxheim (Janssen 1986; Janssen & Brandes 1989a, b). There were no living animals nor empty carapaces nor valves of Cyprideis torosa in samples 18–21. The salt marshes near Sülldorf (sites 4–9) as well as those of Barnstorf and Jerxheim have existed for a long time. The difference between these salt marshes and those of Sülldorf and the ancient Salziger See is the extent of the stagnant waters. In Barnstorf and Jerxheim only small discharges exist which are overgrown by reeds so that there is no visible water, while in Sülldorf and in the ancient Salziger See large areas of stagnant waters are visible and are visited by migrating birds. This may be an explanation for the lack of C. torosa in Barnstorf and Jerxheim; however, the samples contained some interesting co-occurring species with two forms of Sarscypridopsis aculeata. In the pond at Jerxheim (21), S. aculeata was in the typical form, but in the separate fishery pond at Barnstorf (20), S. aculeata was found with a distinct postero-dorsal angle in lateral view (compare Meisch 2000, fig. 163 D) which Hollwedel & Scharf (1988) have found also on some islands in the North Sea. We also have the first records of Ilyocypris inermis (sites 18 and 19) and Prionocypris zenkeri (sites 4–8, 10, 12, 13, 17–19) in a saline environment.
Conclusion
In Germany Cyprideis torosa colonizes mainly the brackish shallow habitats in the Baltic Sea itself or in estuaries near the Baltic Sea, and brackish ditches and ponds near the coast of the Baltic and the North Sea. But this species was also found far from the coast, such as in saline springs at Bad Oldesloe (Klie 1925), lakes Salziger and Süßer See (Klie 1925) (Fig. 2) and Glockensee (Gramann 2000), many sites in central Germany (Pint et al. 2015), and a saline pond at Sülldorf (this paper, Fig. 2). It is absent in the small ditches at Sülldorf (this paper), in the salt marshes, small ponds and their drainage ditches at Barnstorf and Jerxheim (this paper) and in the saline ditch at Salzkotten where Candona species and Heterocypris salina were present [51° 39′ 60″ N, 8° 35′ 56″ E] (Schmidt 1913). It seems that C. torosa occurs inland only in saline waters of significant extent which provide resting places for migrating birds. This hypothesis should be examined in future research.
Acknowledgements and Funding
BS thanks Ursula Beddig, Bremen, for assistance during fieldwork and Hans Theede, Universität Kiel, Ragnar Kinzelbach, Universität Rostock, Claude Meisch, Luxembourg, and Finn Viehberg, Universität Köln, for providing literature. We thank Karel Wouters (Brussels), Claude Meisch (Luxembourg) and Patrick De Deckker (Canberra) for their critical reviews of the manuscript, and Alan Lord (Frankfurt) for improving the English text.
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