**10. Phytoplankton population**

186 Ecosystems Biodiversity

Table 3. Annual variations (A) in abundance (ind.m3) and (B) in biomass (wet weight, g.m3) of total zooplankton, Copepoda, Cladocera and merozooplankton species in the Southern Caspian Sea (before and after Mnemiopsis invasion). (C) Seasonal variations in abundance and biomass of zooplankton species in 1996 from Hossieni et al. (1996). (D) Full list of

Cladocera species reported by Hossieni et al. (1996).

In the present study, a total of 226 phytoplankton species were identified. While diatoms constituted 45% of the total species number, chlorophytes, cyanophytes, dinoflagellates and euglenophytes formed 20, 17, 11 and 8% of phytoplankton species, respectively (Fig. 7). Number of species in spring (91 species) and summer (101 species) were higher than in autumn (86 species) and winter (77 species).

Fig. 7. Variations in species number of different phytoplankton groups in the southern Caspian Sea during 2001–2006 and 1986–1994

The highest monthly mean phytoplankton abundance and biomass were 396 × 106 ± 299 × 106 cells m-3 in January 2002 and 1,789 ± 1,761mg m-3 in May 2002 (Fig. 8). Minimum abundance and biomass values were observed in August 2003 (1 × 106 ± 1 × 106 cells m-3 and 7 ± 5 mg m-3) (Fig. 8).The overall average cell abundance and biomass of phytoplankton during 2001–2006 were 64 × 106 ± 76 × 106 cells m-3 and 250 ± 360 mg.m-3, respectively. While diatoms were the most abundant phytoplankton group during 1996, after the introduction of *M. leidyi* the abundances of cyanophytes (in autumn) and dinoflagellates (in winter) exceeded diatom abundance in 2001 and 2002 (Fig. 8 and 9). Excluding 2005, diatom abundance was again high during 2003–2006. An unprecedented bloom of the toxic cyanophyte *Nodularia* sp. was observed between the second half of August and the end of

*Mnemiopsis leidyi* Invasion and Biodiversity Changes in the Caspian Sea 189

Fig. 9. Seasonal changes a in abundance and b in biomass of different phytoplankton groups before (Hossieni et al. 1996) and after Mnemiopsis leidyi invasion in the southern Caspian

**11.1 Increased chlorophyll levels in the southern Caspian Sea after ML invasion**  A significant correlation was observed between satellite derived chlorophyll *a* (Chl *a*) concentrations and the biomass of the invasive comb jellyfish *Mnemiopsis leidyi* in the

Sea (values are depth and station averages)

**11. Other factors that have to be considered** 

September in 2005. The bloom area covered ˜20,000 km2 (CEP 2006, Fig. 9). According to the sampling on 20 September 2005, in addition to *Nodularia* sp., another cyanophyte Oscillatoria sp. was also high in abundance. Abundance of Nodularia sp. was 18 ×106 cells m-3 at 7 m depth and 1,006 × 106cells m-3 at 20 m depth. Average cyanophyte abundance and biomass at 7 and 20 m depths were 582 9 106 cells m-3 (of which 512 cells m-3 was Nodularia sp.) and 1,655 mg m-3. The highest seasonal means of phytoplankton abundance and biomass were 179 × 106 cells m-3 and 880 mg m-3 in winter during 2001– 2006.

Fig. 8. Annual variations in the abundance and biomass of phytoplankton, zooplankton and Mnemiopsis leidyi in the southern Caspian Sea during 2001–2006 (values are depth and station averages). 1996 values are from Hossieni et al. (1996), spring 2001 values are from Kideys et al. (2001)

September in 2005. The bloom area covered ˜20,000 km2 (CEP 2006, Fig. 9). According to the sampling on 20 September 2005, in addition to *Nodularia* sp., another cyanophyte Oscillatoria sp. was also high in abundance. Abundance of Nodularia sp. was 18 ×106 cells m-3 at 7 m depth and 1,006 × 106cells m-3 at 20 m depth. Average cyanophyte abundance and biomass at 7 and 20 m depths were 582 9 106 cells m-3 (of which 512 cells m-3 was Nodularia sp.) and 1,655 mg m-3. The highest seasonal means of phytoplankton abundance and

Fig. 8. Annual variations in the abundance and biomass of phytoplankton, zooplankton and Mnemiopsis leidyi in the southern Caspian Sea during 2001–2006 (values are depth and station averages). 1996 values are from Hossieni et al. (1996), spring 2001 values are from

Kideys et al. (2001)

biomass were 179 × 106 cells m-3 and 880 mg m-3 in winter during 2001– 2006.

Fig. 9. Seasonal changes a in abundance and b in biomass of different phytoplankton groups before (Hossieni et al. 1996) and after Mnemiopsis leidyi invasion in the southern Caspian Sea (values are depth and station averages)

#### **11. Other factors that have to be considered**

#### **11.1 Increased chlorophyll levels in the southern Caspian Sea after ML invasion**

A significant correlation was observed between satellite derived chlorophyll *a* (Chl *a*) concentrations and the biomass of the invasive comb jellyfish *Mnemiopsis leidyi* in the

*Mnemiopsis leidyi* Invasion and Biodiversity Changes in the Caspian Sea 191

CEP. 2006. A Study on the Harmful Algal Bloom in the Southwestern Basin of the Caspian

Davis, B.A.S., Brewerb, S., Stevensona, A.C., Guiotc, J. and Data Contributors. 2003. The

Esmaeili, A., Abtahi, B., Khoda bandeh, S., Talaeizadeh, R., Darvishi, F. and Terershad, H.

*Environmental Sciences and Technology*, *Islamic Azad University*, 3, 63–69. Fazli, H. and Roohi, A. 2003. The impacts of *Mnemiopsis leidyi* on species composition, catch

University Research bulletin, Astrakhan (KaspNIRKh) No. 3: 99-104. GESAMP. 1997. Opportunistic settlers and the problem of the ctenophore *Mnemiopsis leidyi* 

temperature of Europe during the Holocene reconstructed from pollen data,

2000. First report on occurrence of a combjelly in the Caspian Sea. *Journal of* 

and CPUE of Kilka in Iranian commercial catch. UNESCO, Caspian Floating

invasion in the Black Sea. Reports and Studies no. 58, GESAMP (Joint Group of Experts on the Scientific Aspects of Marine Environmental Protection), London. Gordina, A. D.; Tsytsugina, V. G.; Ovsyanyj, E. I.; Romanov, A. S.; Kemp, R. B., 2004:

Condition of pelagic fish spawn in coastal waters of the Black Sea near Sevastopol.

mesozooplankton by the example of the Copepoda community in Sevastopol Bay

feeding in the Southern Caspian Sea, 2sd International applied biological Congress,

Wilson, G. R. Harbison and H. J. Dumont. 2000. Invasion of Caspian Sea by the

*Mnemiopsis* in the Caspian Waters of Iran. A report prepared for the Caspian

Levels in the Southern Caspian Sea Following an Invasion of jellyfish, Hindawi Publishing Corporation, Research Letters in Ecology, Article ID 185642, 4 pages,

Rostamian, M., Rostami, H. and Negarestan, H. 2004. Physiological characteristics of the

Ctenophores on the Fisheries of the Black Sea and Caspian Sea. *Oceanography-Black* 

Prusova IYu, Skryabin VA, Uysal Z, Zagorodnyaya JuA. 1998. Long-term changes in the biomass and composition of fodder zooplankton in coastal regions of the Black Sea during the period 1957–1996. In: Ivanov LI. and Oguz T (eds.) Ecosystem Modelling as a Management Tool for the Black Sea. Kluwer Academic Publishers,

Gubanova AD, Polikarpov IG, Saburova MA, Prusova IYu. 2002. Long-term dynamics of

Harbison, G. R., Madin, L. P. and Swanberg, N. R. 1978. On the natural history and distribution of oceanic ctenophores. Deep-Sea Research 25, 233-256. Hashemian, A. and Roohi, A., 2004. A survey of *Mnemiopsis leidyi* impacts on Sturgeon fish

Ivanov P. I., A. M. Kamakim, V. B. Ushivtzev, T. Shiganova, O. Zhukova, N. Aladin, S. I.

comb jellyfish *Mnemiopsis leidyi* (Ctenophora). *Biological Invasions* 2:255–258. Kideys A. E., Ghasemi S., Ghninejad D., Roohi A., Bagheri S. 2001. Strategy for Combatting

Kideys, A., Finenko, F., Anninsky, B., Shiganova, T., Roohi, A., Tabari, M., Youseffyan, M.,

ctenophore *Beroe ovata* in Caspian Sea water, *Marine Ecology*, Vol. 266: 111–121. Kideys A.E., A. Roohi, S. Bagheri, G. Finenko, L. Kamburska. 2005. Impacts of Invasive

Kovalev AV, Gubanova AD, Kideys AE, Melnikov VV, Niermann U, Ostrovskaya NA,

Environment Programme, Baku, Azerbaijan, Final Report, July 2001. Kideys, A. E. 1994. Recent dramatic changes in the Black Sea ecosystem: The reason for the sharp decline in Turkish anchovy fi sheries. Journal of Marine Systems 5:171-181. Kideys, A. E., Roohi, A., Elif, E. D., Melin, F.and Beare, D. 2008. Increased Chlorophyll

Sea, Caspian Environment Programme, 15 pages

Quaternary Science Reviews 22 (2003) 1701–1716.

*Hydrobiol. J.* 40, 43–55.

(1976–1996) Oceanology 42 (4): 512-520.

Mashhad Free University, Mashhad, Iran.

and doi:10.1155/2008/185642.

*Sea Special Issue* 18(2):76-85.

Dordrecht/Boston/London, Vol. 1, pp.209–219

southern Caspian Sea. By consuming the herbivorous zooplankton, the predatory ctenophore *M. leidyi* may have caused levels of Chl *a* to rise to very high values (∼9mg m*−*3) in the southern Caspian Sea. There might also be several other factors concurrent with predation effects of *M. leidyi* influencing Chl *a* levels in this region, such as eutrophication and climatic changes which play major roles in nutrient, phytoplankton, and zooplankton variations (kideys et al., 2008). The decrease in pelagic fishes due to overfishing, natural, and anthropogenic impacts might have provided a suitable environment for *M. leidyi* to spread throughout this enclosed basin (Fig. 10).

Fig. 10. Spatiotemporal distribution of Chl *a* concentration (mgm*−*3, note the broken scale here), zooplankton abundance (ind *·* m*−*2), *Mnemiopsis leidyi* biomass (g m*−*2 values for June and August 2001 are from Shiganova et al. 2004), and sea surface temperature (*◦*C) obtained from NOAA in the Caspian Sea. Note the strong difference in Chl *a* distributions (as seen from satellite during a warm period, September) before (1998 and 1999) and after *Mnemiopsis leidyi* impact (2001 and 2006) in the lower section of the figure.

#### **12. References**


southern Caspian Sea. By consuming the herbivorous zooplankton, the predatory ctenophore *M. leidyi* may have caused levels of Chl *a* to rise to very high values (∼9mg m*−*3) in the southern Caspian Sea. There might also be several other factors concurrent with predation effects of *M. leidyi* influencing Chl *a* levels in this region, such as eutrophication and climatic changes which play major roles in nutrient, phytoplankton, and zooplankton variations (kideys et al., 2008). The decrease in pelagic fishes due to overfishing, natural, and anthropogenic impacts might have provided a suitable environment for *M. leidyi* to spread

Fig. 10. Spatiotemporal distribution of Chl *a* concentration (mgm*−*3, note the broken scale here), zooplankton abundance (ind *·* m*−*2), *Mnemiopsis leidyi* biomass (g m*−*2 values for June and August 2001 are from Shiganova et al. 2004), and sea surface temperature (*◦*C) obtained from NOAA in the Caspian Sea. Note the strong difference in Chl *a* distributions (as seen

Agassiz L. 1860. Contribution of the natural history of the United States of America. Boston:

Aladin, N. B. and Plotnikov, I. S. 2004. The Caspian Sea, Lake Basin Management Initiative,

Birstein, Y. A., Vinogradov, L.K., Kandakova, N., Kon, M. S., Stakhovaya, T. V. and

Romanova, N. N. 1968. Atlas of the Caspian Sea invertebrates. Food industrial Co.

from satellite during a warm period, September) before (1998 and 1999) and after *Mnemiopsis leidyi* impact (2001 and 2006) in the lower section of the figure.

throughout this enclosed basin (Fig. 10).

**12. References** 

Truber and Co. V.3.p.1-321.

Moscow, 415 pp.

the Caspian Bulletin 4: 112–126.


**9** 

*Sweden* 

**Main Ecosystem Characteristics and** 

 **Alpine Landscapes in Northern** 

**Sweden Under Climate Change** 

A. Allard, L. Nilsson and J. Svensson *Swedish University of Agricultural Sciences* 

J. Jeglum, S. Sandring, P. Christensen, A. Glimskär,

**Distribution of Wetlands in Boreal and** 

Wetlands and peatlands are integral parts of many of the world's biomes, forming important transition zones between upland and aquatic systems. These habitats have a high degree of complexity of hydrology, edaphic conditions, and vegetation composition, contributing to the biodiversity of landscapes and species richness. They act to influence and modify the movement of runoff and groundwater from uplands into streams and lakes, by laying down organic remains (peats), and absorbing and releasing elements, compounds, gases, and particulate and dissolved organic matter. They therefore act as hydrological

Many kinds of wetlands and peatlands can be found, each with a particular hydrology and surface form, moisture and chemical regime, and range of vegetation types and associated biota. Owing to their hydrological characteristics, predominantly peat soils and hydrophytic plants, wetlands and peatlands are key habitats to indicate climate change, particularly changes towards drying (e.g., decreased precipitation, increased runoff from melting glaciers and snow pack). Changes in moisture regime will effect changes in the processes of peat accumulation and decomposition, release of nutrients and dissolved organic matter, and vegetation and species. Drainage for agriculture and forestry, peat harvesting, and development have already caused considerable areas of peatlands to decrease in depth and area. Owing to drying, some peatlands adjacent to uplands have decreased in depth to less than 30 cm, the defined depth for peatlands in Sweden, and thus the total area of peatland

Drying also has caused changes in vegetation, for examples, advances of trees and shrubs from the margins into the centres of peatlands (e.g., Fig. 1; cf. Hebda et al., 2000; Linderholm & Leine, 2004), and the dying of Sphagnum by lowered water levels and being covered over by leaf litter. Hebda *et al.* estimated the zone of influence of water lowering in Burns Bog, a bog on the Fraser River Delta in southern British Columbia, Canada, to extend over 100 m from a peripheral ditch. The Swedish Wetland Inventory, VMI, (Gunnarsson & Löfroth, 2009) was conducted during 25 years and generated results that indicate that about 15% of

**1. Introduction** 

has decreased.

water retainers and biological filters in the landscape.

