Life History and Current Distribution of the Fairy Shrimp Chirocephalus Carnuntanus (Brauer, 1877) (Crustacea: Anostraca)
NORTH-WESTERN JOURNAL OF ZOOLOGY 11 (1): 102-109 ©NwjZ, Oradea, Romania, 2015 Article No.: 141303 http://biozoojournals.ro/nwjz/index.html
Life history and current distribution of the fairy shrimp Chirocephalus carnuntanus (Brauer, 1877) (Crustacea: Anostraca)
Zsófia HORVÁTH1,* and Csaba F. VAD2
1. WasserCluster Lunz, Dr. Carl Kupelwieser Promenade 5, A-3293 Lunz am See, Austria. 2. Doctoral School of Environmental Sciences, Eötvös Loránd University, Pázmány Péter sétány 1/A, H-1117 Budapest, Hungary. * Corresponding author, Z. Horváth, E-mail: [email protected]
Received: 23. January 2014 / Accepted: 15. September 2014 / Available online: 02. January 2015 / Printed: June 2015
Abstract. The anostracan Chirocephalus carnuntanus is presumably a Pannonian endemic large branchiopod species. In this paper, we provide the first detailed data on its life history characteristics and reviewed its recent distribution. The life cycle study was conducted in an intermittent soda pond (formed in a bomb crater). C. carnuntanus was found to be univoltine, which is in accordance with previous assumptions. The first nauplii occurred after ice-break in mid-February and the population disappeared two months later at the end of April, much earlier than the desiccation of the pond. During population development, we observed a gradual change in the sex ratio after the appearance of pre-adults, from approximately equal ratios to almost exclusive male dominance, probably due to the higher cost of egg production. Based on its recent distribution, this species is rather rare and sporadic, restricted only to the lowlands of the Carpathian Basin. Particular caution should be taken by habitat managers to help maintaining suitable habitats for the long- term conservation of this species.
Key words: Chirocephalus carnuntanus, life history, secondary habitat, sex ratio, soda pond.
Introduction 1). Apart from two exceptions – Moscow and the northern Bohemian Lowland (the latter habitat no Large branchiopods (Crustacea: Anostraca, Noto- longer exists, Merta, Lukáš, pers. comm. 2011), it is straca, Spinicaudata and Laevicaudata) are charac- only present in the Carpathian Basin (Brtek & teristic inhabitants of temporary waters and can be Thiéry 1995) and Löffler (1993) regarded it an en- considered a flagship group of these systems (Belk demic Pannonian species. In the Basin, it inhabits 1998, Gascón et al. 2012). They are threatened intermittent and temporary habitats (Eder et al. worldwide owing to the loss and destruction of 1997, Horváth et al. 2013a), being present in astatic their habitats. Being restricted to temporary wa- soda pans of Hungary and Austria (Eder & Hödl ters, they can be used to assess the quality and 2003, Horváth et al. 2013a), sodic pools of Serbia functioning of these systems. Recent studies re- (Petkovski 1993, Petrov & Cvetković 1997, Lukić et vealed their key ecological roles either by affecting al. 2012), and some other freshwater temporary invertebrate community structure through preda- pools of Hungary (Boven et al. 2008), western Ro- tion and competition (Yee et al. 2005, Jocque et al. mania (Demeter & Stoicescu 2008), northern Serbia 2010, Waterkeyn et al. 2010) or by providing a ma- (Lukić et al. 2012) and western Slovakia (Lukáš jor food source for migratory waterbirds (Horváth 2000) (Fig. 2, Table 1). et al. 2013b). Despite the growing number of stud- On the biology of C. carnuntanus, very little is ies on their ecology, our knowledge on the major- known (Eder et al. 1996). The species is presumed ity of species is still poor, although detailed to be univoltine (Löffler 1993, Eder et al. 1997), as demographic studies, investigations targeting individuals occur generally in early spring and their resting egg banks and knowledge on their disappear when water warms up. Horváth et al. ecological requirements should be essential for (2013a) also found them exclusively in spring. Ac- their long-term conservation. This is even more cording to Eder & Hödl (2003), C. carnuntanus is pronounced in the case of rare species with re- presumably a filter-feeder, similarly to most stricted geographic area. anostracans. Eder (2006) and Boven et al. (2008) The genus Chirocephalus represents the second suggested that C. carnuntanus is a habitat general- largest genus of anostracans with 53 species ist, which was confirmed by Horváth et al. (2013a) (Rogers 2013). Several of these species have re- in a large-scale study conducted in astatic soda stricted geographic distributions. A typical exam- pans. Along with Branchinecta orientalis Sars, 1901 ple is Chirocephalus carnuntanus (Brauer, 1877) (Fig. and Branchinecta ferox (H. Milne Edwards, 1840), it Life history of Chirocephalus carnuntanus 103 has a keystone role in the trophic web of shallow astatic soda pans in the Carpathian Basin, by pro- viding important food source for migratory wa- terbirds (Horváth et al. 2013b). Our aim was to study the life history of the species in an intermittent pond. We also aimed to compare the body lengths of males and females using new and previous data formerly collected from different soda pans in Hungary and Austria (during a large-scale sampling campaign in 2010) and to review the recent distribution of this rare species (based on data from the last 20 years).
Materials and Methods
Study area The study area is located 30 km south of the Hungarian capital, Budapest, near to the village of Apaj in Kiskunság on the Danube-Tisza Interfluve (Fig. 2). Within an area of approximately 1 km2, more than 100 bomb craters can be found, which were created in bombings during or shortly after World War II. As this part of Kiskunság has sodic soil, the craters now act as secondary ponds and pools Figure 1. Adult female of C. carnuntanus (on the top; holding sodic water. They are astatic, usually drying out photo: Imre Potyó) from the sodic bomb crater in summer and filling up during late autumn or winter, pond (on the bottom; photo: Zsófia Horváth). while their diameter varies between approximately 3 and 12 m (Fig. 1). Many of them are surrounded by vegetation of common reed (Phragmites australis) and sedges (mainly mesh size: 45 μm) was pushed along in five 2 m long Bolboschoenus maritimus) and are entirely or partly filled transects in the ponds, of which three were close to the up (with a max. depth less than 40 cm), but there are still surface and two to the bottom, to sort out bias due to pos- some that are deeper (>70 cm), with a bottom free of sible vertical distribution patterns. For larger animals, a marsh plants. These deeper ponds have higher salinity hand net (opening surface: 346 cm2) was pushed along in and water evaporates later than from the former ones. Al- similar transects with a larger mesh size (5 mm), which together, these latter ponds can be regarded similar to enabled faster movements without pushing away the larger natural astatic soda pans of the region (Boros et al. animals in front of the opening of the net. Later on, as wa- 2014, Horváth et al. 2014). ter evaporated and the ponds became shallower, sam- pling was modified and water was collected initially only Sampling and data analysis from the same depth (in three 2 m long transects), and Sampling was conducted in two periods: first a short sur- later with a one-litre plastic beaker (with a total of 5 litres vey was carried out in March 2010, when C. carnuntanus of water collected), when water depth became lower than was discovered in two of the deeper, more saline craters the diameter of the nets (it happened well after the disap- (with conductivity values of 2.8 and 2.1 mS cm-1 on 11th pearance of C. carnuntanus). Water volume sampled was March 2010), and then these two craters (C1 and C2) were calculated based on the surface of the opening of the ac- chosen for a life history study, which was carried out in tual net and the length of transects. 2011 (see Table 1 for their exact location). The life history Anostracan samples were preserved in 70% ethanol study began on 17th Feb 2011, immediately after the first solution and identification was based on the keys of Eder ice break (there was a second ice break afterwards, as the & Hödl (1996). ponds froze again later in February). Sampling was con- The length of each collected animal was measured ducted in 1–2 week intervals (more frequently when C. with 0.5 mm precision. Nauplii and smaller specimens carnuntanus was present, apart from late February, when were measured after preservation, while larger-sized thick ice layers prohibited sampling), until the ponds specimens were measured on the field and afterwards re- dried out at the end of August. We investigated the tim- leased back to the pond. To estimate sex ratio in the ing of the hatching, the duration of population presence, population, if less than 10 animals were caught with the the temporal change in the ratio of males and females hand net, we searched for additional individuals for an- along with the body length and the ratio of animals reach- other 15 minutes. These additional specimens were not ing the adult stage. included in the density calculation, and were only used Two types of anostracan samples were collected. To for sex ratio and length measurements and were also re- collect nauplii, a plankton net (opening diameter: 15 cm, leased back to the pond afterwards. 104 Z. Horváth & C.F. Vad
Along with the changes in sex ratio and average ductive organs) appeared in the second half of length of adults, juvenile mortality and maximum lifetime March, while adults (with eggs in the egg sacks of were also calculated. females and well-developed reproductive organs Water depth at the deepest point of the crater was and second pair of antennae in the case of males) measured at each sampling occasion, along with water temperature, pH and conductivity, which were deter- were first recorded in early April and disappeared mined by using a WTW Multiline P4 field equipment before May. (with TetraCon 325 and SenTix 41 electrodes). Three litres Among the measured environmental factors, of water were additionally collected for the measurement temperature, conductivity, turbidity and TSS of turbidity (with a Lovibond PC Checkit turbidity meter) gradually increased from February until all ani- and the concentration of chlorophyll-a and total sus- mals disappeared (beginning of May). Water pended solids (TSS). TSS was measured by filtering water depth constantly decreased, while pH and chloro- (100−1000 ml) through pre-dried and pre-weighted cellu- lose acetate filters after oven-drying (at 105 oC). For chlo- phyll-a concentration did not show any clear pat- rophyll-a concentrations, water (100−1000 ml) was filtered tern. Ranges and medians of the recorded vari- through glass microfibre filters, and concentration was ables are presented in Table 2. determined with a Secomam Anthelie UV-vis spectropho- Juvenile mortality was high: the ratio between tometer after hot methanol extraction (Wetzel & Likens, the density of pre-adults (on 25th March) and the 1991, Pápista et al. 2002). No acidic correction for phae- max. density of larvae (on 1st March) was 0.2, opigments was made. while between the adults (on 10th April) and lar- For the intersexual length difference analysis, we used pre-adult and adult individuals of C. carnuntanus vae it was 0.08, which means that 20.4% of animals that were collected on 10th April 2011 from C1 (when hatched after ice break entered the pre-adult stage males and females were present in equal numbers; N=16) and only 7.7% reached the adult stage. or previously in March 2010 from 10 soda pans in Hun- The population comprised of only one cohort gary and another in Austria (see Horváth et al. 2013a for and no nauplii or metanauplii could be observed further details on the exact locations). Of these latter together with pre-adult or adult animals. More samples, we only used those which contained more than than one month was needed for the first appear- 20 animals and at least 5 specimens of each sex. This meant four Hungarian pans (Bogárzó at Solt, Büdös-szék, ance of pre-adults from hatching, and afterwards, Kelemen-szék and Zab-szék at Szabadszállás, total it took more than two weeks until animals reached N=137). For the comparison of the mean body length of adult stage. From the mass-appearance of nauplii, males and females, we first tested for normality and the the population existed for nearly two months, homogeneity of variances in each case (C1 pond and the which should be close to the maximum life time of four soda pans mentioned above). Subsequently, separate C. carnuntanus given its univoltine feature. The independent two-group t-tests (in case of normal distribu- growth of individuals was the fastest in late tion and homogeneous variances; for C1 pond and Bogárzó) or Wilcoxon tests (if the population could not be March, when they reached pre-adult stage (Fig. assumed to be normally distributed; for Büdös-szék, 3b). Kelemen-szék and Zab-szék) were performed. Analyses By the appearance of pre-adults in the second were made in R (R Development Core Team, 2009). half of March, the crater population was domi- To review the recent distribution of C. carnuntanus, nated by females (Fig. 4). The ratio gradually we collected and summarised all literature data reporting changed: both sexes were equally represented in its occurrence from the last 20 years (starting from 1995). early April, but later on, most females disap- We excluded locations reported without the year of oc- currence. Additionally, we used material and data avail- peared. able at the Collection of Crustacea and Other Aquatic In- Adult animals (measured on 10th April) had a vertebrates of the Hungarian Natural History Museum mean length of 11.3 (1 SD: 1.5) mm (without (collected between 1995 and 2014). furca). Adult females had a slightly larger body length compared to adult males in the soda pond (although it was not significant; Table 3), which Results was similar in three of the other investigated soda
During our life history study, animals were found pans. In Zab-szék, the mean length of males was only in crater C1. Hatching began at the time of significantly larger, while their maximum length the first ice break in early February, when water was smaller than that of females. temperature was only 2oC, while most nauplii and Based on available data, C. carnuntanus has metanauplii were found in the beginning of been recorded from 54 locations in the last 20 March, after the second ice break (Fig. 3a). Pre- years (Table 1, Fig. 2), mostly from Hungary (41 records). It occurred in various habitat types, from
adults (large individuals with noticeable repro- Life history of Chirocephalus carnuntanus 105
Table 1. List of occurrences of C. carnuntanus from the last 20 years (Cou- Country, AT- Austria, HU- Hungary, RO- Romania, RS- Serbia, SK- Slovakia, a.l.- approx. location, b.c.- bomb crater, u.d.- unpublished data, p.c.- personal communication).
Cou. Settlement Name of habitat Habitat type Lat (N) Lon (E) Reference AT Apetlon Lange Lacke soda pan 47.762° 16.870° Horváth & Vad u.d. 2013 AT Apetlon Ochsenbrunnlacke soda pan 47.810° 16.845° Horváth et al. 2013a AT Apetlon Östliche Hut- soda pan 47.769° 16.875° Eder & Hödl 2003 weidenlacke AT Illmitz Zicksee soda pan 47.768° 16.786° Eder & Hödl 2003 HU Apaj - pool on arable field, 47.104° 19.085° Forró, László, u.d. 1996, wheel-track (a.l.) 1999 HU Apaj - sodic pond (b.c.) 47.124° 19.131° Forró, L., u.d. 2006 HU Apaj C1 sodic pond (b.c.) 47.121° 19.133° present paper HU Apaj C2 sodic pond (b.c.) 47.122° 19.136° present paper HU Dunatetétlen Böddi-szék (N Pool) soda pan 46.766° 19.157° Horváth et al. 2013a HU Dunatetétlen Böddi-szék (SE Pool) soda pan 46.757° 19.157° Horváth et al. 2013a HU Dunatetétlen Böddi-szék (SW Pool) soda pan 46.757° 19.154° Horváth et al. 2013a HU Dunatetétlen Böddi-szék (W Pool) soda pan 46.765° 19.147° Horváth et al. 2013a HU Dunatetétlen Fűzfa-szék soda pan 46.792° 19.158° Horváth et al. 2013a HU Fülöpszállás - pool (a.l.) 46.800° 19.217° Boven 2009 HU Fülöpszállás - pool (a.l.) 46.817° 19.283° Boven 2009 HU Fülöpszállás Kelemen-szék soda pan 46.797° 19.184° Forró, L. u.d. 1996, 1997 Horváth et al. 2013a HU Fülöpszállás Partos-szék sodic meadow pool 46.818° 19.228° Forró, L. u.d. 2001 HU Izsák - wheel-track (a.l.) 46.776° 19.393° Forró, L. u.d. 2005 HU Jászárokszállás - pool on arable field (a.l.) 47.633° 19.964° Forró, L. u.d. 2001 HU Kardoskút - pool on arable field 46.472° 20.590° Forró, L. & Kovács, K. u.d. 2006 HU Kardoskút - meadow pool 46.471° 20.591° Forró, L. & Kovács, K. u.d. 2006 HU Kardoskút - pool on arable field 46.469° 20.593° Forró, L. u.d. 2006 HU Kardoskút - pool on arable field 46.471° 20.591° Forró, L. u.d. 2006 HU Kenézlő - pool on arable field (a.l.) 48.202° 21.540° Forró, L. u.d. 1999 HU Kömlő - pool on arable field 47.591° 20.464° Forró, L. u.d. 1997, 1999, 2006 HU Kunszentmiklós - pool (a.l.) 47.033° 19.083° Boven 2009 HU Kunszentmiklós - pool (a.l.) 47.033° 19.117° Boven 2009 HU Orgovány - pool on arable field 46.770° 19.401° Forró, L. u.d. 2005, 2006 HU Orgovány - pool on arable field 46.764° 19.428° Forró, L. u.d. 2006 HU Páhi - unknown (a.l.) 46.706° 19.382° Forró, L. u.d. 2005 HU Poroszló - pool on arable field 47.662° 20.588° Forró, L. u.d. 2006 HU Pusztaszer Büdös-szék soda pan 46.546° 20.033° Horváth et al. 2013a HU Solt Bogárzó soda pan 46.808° 19.142° Horváth et al. 2013a HU Solt - pool on arable field (a.l.) 46.803° 19.139° Forró, L. & Boros, E. u.d. 2001 HU Soltszentimre - pool on arable field 46.762° 19.236° Forró, L. u.d. 2006 HU Szabadszállás Büdös-szék soda pan 46.866° 19.169° Horváth et al. 2013a HU Szabadszállás Pipás-rét sodic meadow pool 46.878° 19.185° Forró, L. u.d. 1998 HU Szabadszállás Pipás-szék soda pan 46.876° 19.177° Horváth et al. 2013a HU Szabadszállás Zab-szék soda pan 46.837° 19.170° Horváth et al. 2013a HU Soltszentimre - pool (a.l.) 46.767° 19.333° Boven 2009 HU Tiszacsege - pool on arable field (a.l.) 47.697° 20.990° Forró, L. u.d. 1996 HU Tiszafüred - unknown (a.l.) 47.600° 20.810° Forró, L. u.d. 2001 HU Tiszanána - meadow pool (a.l.) 47.569° 20.515° Forró, L. u.d. 1999 HU Ürbőpuszta (Bugyi) - pool on arable field (a.l.) 47.159° 19.170° Forró, L. u.d. 2005 RO Baile Calacea - pool 45.906° 21.174° Demeter & Stoicescu 2008 RO Sânnicolau Mare - pool 46.073° 20.633° Demeter & Stoicescu 2008 RO Timişoara - pool 45.775° 21.195° Demeter & Stoicescu 2008 RS Bačko Gradište - pasture pond 45.538° 20.023° Lukić et al. 2012, Šćiban, M. p.c. 2013 RS Melenci - pool 45.525° 20.325° Petrov & Cvetković 1997 RS Novi Bečej - sodic meadow pool 45.633° 20.185° Lukić et al. 2012, Šćiban, M. p.c. 2013 SK Vysoká pri Morave - pool 48.330° 16.909° Lukáš 2000 106 Z. Horváth & C.F. Vad
Sl ova ki a ● ● ke La rtő l/Fe Hungary Ti sza sied Neu ●● Danube ● ● Aust ria ● ●● ●
n ●● lato ● Ba ●● Figure 2. Distribution of C. carnunta- ke La ●●●● Romani a ●●●●●● nus in the Carpathian Basin ac- ● ● cording to data from the last 20 years. Location of the bomb craters ● is signed with an open circle while ● Se r bi a ● 05025 km other occurrences are indicated ● with filled circles. Exact locations ● ● and habitat types are listed in Ta- ble 1.
a) 6 nauplii + metanauplii 5 preadults 30 -1 adults temperature 4 20 3
2 10 Temperature (C°) Temperature
Density (ind ) l 100 1
0 0 b) 14
12
10
8 Figure 3. Density of larval, 6 preadult and adult forms of C. carnuntanus (a) and their 4 Length (mm) average length (± SD) without 2 furca (b). Sampling dates are
0 presented on real-time axis 3 3 and water temperature is also .0 .0 .04 0.04 17.02 01.03 10.03 18 25 31.03 1 20.04 28 08.05 shown.
large astatic soda pans (which were the most fre- late August. Similarly, populations always con- quent habitat type with 15 records) to small sisted of only one cohort per habitat in our previ- meadow pools and wheel-tracks. ous investigations in 12 soda pans (Horváth et al. 2013a, Horváth & Vad unpubl. data), which con- firms its univoltine feature. Discussion Hatching began when conductivity was 3.07 mS cm-1, and animals were still present with 3.85 Hatching of C. carnuntanus started with the ice mS cm-1. The latter value is somewhat higher than break, which is in accordance with its cold water the upper limit for the species Horváth et al. preference (Eder & Hödl 1996). Previous authors (2013a) found in soda pans. In C2, no hatching oc- (e.g. Löffler 1993, Eder et al. 1997, Horváth et al. curred in 2011, although we also monitored this 2013a) also found this species only in spring, and pond during the whole study period. This may be it also disappeared in our case before May. After parallel with different environmental parameters its disappearance, no newly hatched animals were of the crater in this year (lower salinity, denser found in the crater until it completely dried out in vegetation, lower turbidity). Compared to C1 at
Life history of Chirocephalus carnuntanus 107
Table 2. Local environmental variables (minimum, maximum, median) in soda pond C1 (based on values recorded during the presence of animals) and values measured in C1 and C2 at the start of recruitment (17.02.2011).
17.02.2011–28.04.2011 17.02.2011 C1 min. C1 max. C1 median C1 C2 pH 9.2 9.4 9.4 9.2 9.5 Water temperature (°C) 2.2 21.5 11.4 2.2 3 Conductivity (mS cm-1) 3.1 3.9 3.2 3.1 1.3 Water depth (cm) 58.5 82 70.5 82 81 Dissolved oxygen (%) 70 302 126.3 144.5 173 Dissolved oxygen (mg l-1) 6.2 35.4 12.0 19.5 24.5 Chlorophyll-a (µg l-1) 1 9.5 5 5.0 17.7 Turbidity (NTU) 19 69 36 28 8.6 Total suspended solids (mg l-1) 8.5 53.4 16 13 7.9
100 female
80 male
60
40 Percentage (%) 20 Figure 4. Temporal changes in sex ratio after the appearance of pre-adults. 0 Dashed line indicates 50%. Sampling 3 3 4 4 4 .0 .0 .0 .0 .0 5 1 0 0 8 dates are presented on real-time axis. 2 3 1 2 2
Table 3. Difference in body length between males and females of C. carnuntanus (without furca). Abbreviations: C1 – our investigated pond, Bog – Bogárzó, Bud – Büdös-szék at Szabadszállás, Kel – Kelemen-szék, Zab – Zab- szék (for exact locations, see Table 1)
C1 Bog Bud Kel Zab Female mean length (mm) ± 1 SD 11.88 ± 0.83 14.64 ± 1.34 13.92 ± 1.02 13.39 ± 1.34 13.69 ± 3.17 Male mean length (mm) ± 1 SD 10.63 ± 1.77 13.56 ± 1.33 12.43 ± 0.96 11.80 ± 1.74 16.47 ± 2.60 Female max. length (mm) 13 17 16 17 22 Male max. length (mm) 13 16 14 17 20 t-value 1.80 1.91 - - - W-value - - 154.5 440.5 45.5 p 0.092 0.070 0.004 <0.001 0.006
the beginning of hatching in 2011 (Table 2), C2 had find even a single nauplius in the crater C2 implies lower salinity and turbidity, moreover, Potamoge- that even hatching from the egg bank was inhib- ton pectinatus already dominated C2 (with an ap- ited. Additionally, when we re-visited the bomb proximately 90% coverage), while C1 was largely craters in 2014 (12th March), we found animals unvegetated (less than 5% coverage), with bare both in C1 (1.96 mS cm-1) and C2 ponds (2.72 mS muddy sediment. In March 2010, when animals cm-1). were observed in this pond, conductivity was 2.04 Its absence in 2011 can also be in accordance mS cm-1, while at the same time in 2011, its con- with the risk-spreading strategy of anostracans, ductivity was only 1.32 mS cm-1. As C. carnuntanus which is described in other similar species, e.g. inhabits freshwater pools with conductivities well Chirocephalus ruffoi Cottarelli & Mura, 1984 (Mura below 1 mS cm-1 (Horváth et al. 2013a), such a 2001). Similarly, C. carnuntanus was not present in drop in this variable alone cannot explain its dis- the Austrian Lange Lacke in 2010 (Horváth et al. appearance. It is more likely that it acted indi- 2013a), while it was abundant in this soda pan in rectly, e.g. it may have contributed to a higher 2013 (Horváth & Vad, unpubl. data). number of predators. But the fact that we did not We observed a gradual change in the sex ratio 108 Z. Horváth & C.F. Vad after the appearance of pre-adults. The earlier dis- univoltine feature. The time period of its occur- appearance of females may be attributable to the rence is therefore very limited, taking place in the higher cost of egg production. Beladjal et al. (2003) first part of spring (in accordance with Horváth et similarly found that the males of Streptocephalus al. 2013a). This also stands for other anostracan torvicornis (Waga, 1842) live much longer than fe- species in the region as many of them are similarly males (they outlive females by more than a fe- spring species (Eder et al. 1997), being presumably male’s lifetime). On the other hand, S. torvicornis univoltine. Specific discovery campaigns and had a maximum lifetime of 325 days, which is raised public awareness could help to better ex- much longer than that of C. carnuntanus (pre- plore the distribution of such large branchiopods sumably between 59 and 71 days in the sodic and help habitat management. bomb crater). In the case of the cold-water species Similarly to the majority of anostracans (Bren- C. carnuntanus, this is due to its disappearance donck et al. 2008), C. carnuntanus is a specialist of with the rapidly increasing water temperature, temporary waters. These habitats seriously de- while S. torvicornis is rather a summer species clined in the last centuries in Europe (together (Eder et al. 1997). The approximately two-month with permanent ponds, these habitat were re- maximum lifetime we found in the case of the duced by more than 50% for most countries and bomb crater population is similar to other Euro- may even reach 90% in some regions; Hull 1997, pean species e.g. Branchipus schaefferi Fischer, 1834 Wood et al. 2003, Curado et al. 2011). Despite the (77 days; Beladjal et al. 2003) or Tanymastix stag- growing interest in their ecology and conserva- nalis (Linnaeus, 1758) (30–60 days; Mura & Zarat- tion, temporary waters are still neglected and en- tini 2000). dangered by habitat destruction (Blaustein & Sexual dimorphism in the body length of Schwartz 2001, Céréghino et al. 2008). Considering Anostraca varies among the different species. For its habitat preference, together with its limited example, adult females of S. torvicornis have much geographic range, C. carnuntanus can be consid- larger body length (females: 27.68 mm, males: ered vulnerable. 22.23 mm; Beladjal et al. 2003), while Miličić et al. The presence of this rare and (presumably) (2013) found no general difference in the case of B. endemic species in the sodic bomb crater ponds schaefferi. In our case, adult females of C. carnunta- also draws the attention to the conservation im- nus were slightly larger than males, but the in- portance of originally man-created habitats in trasexual difference in the mean body size be- maintaining intermittent and temporary water tween pans (presumably due to local factors, e.g. biodiversity. This could be especially important food quality and quantity) was generally much for the group of large branchiopods, given their larger than the intersexual difference within the presence in wheel-tracks (Boven et al. 2008, Ar- same pan. mitage et al. 2012) and even in military areas According to existing data from the last 20 (Maier et al. 1998, Gołdyn et al. 2012). Such species years, C. carnuntanus has a patchy distribution in would need special attention in respect of local the Carpathian Basin. Most records are from the habitat management for their long-term conserva- Great Hungarian Plain (49 out of 54 total, which tion. means more than 90% of all data) and within this region, most habitats (30) are restricted to the Kiskunság region in Central Hungary (on the Da- Acknowledgements. We thank László Forró for nube–Tisza Interfluve). This is a similar situation providing information regarding material available at the Hungarian Natural History Museum, Balázs Németh and to the distribution map reported by Forró (2000), Lajos Vörös for TSS measurements, Imre Potyó for the who reviewed the distribution of this species in photo of the live animal, Adrienn Tóth for helping with Hungary based on all available historic and new the preparation of the distribution map and the staff of data before 2000 (dating back to the 19th century). the Kiskunság National Park Directorate (Emil Boros, The patchy distribution of the species within István Nagy) for showing us the study area and allowing the region can also partly originate from sampling further access to it. We are grateful to Bartłomiej Gołdyn bias. Most freshwater studies target permanent and an anonymous reviewer for their helpful comments on the manuscript. This research was supported by the systems, and researchers mainly conduct spring European Union and the State of Hungary, co-financed sampling in late April or May. This considerably by the European Social Fund in the framework of reduces the possibility of finding C. carnuntanus in TÁMOP 4.2.4.A/2-11-1-2012-0001 ‘National Excellence the field given its relatively short lifespan and Program’. The authors contributed equally to this work. Life history of Chirocephalus carnuntanus 109
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