Colonization of a new man-made river (Marchfeldcanal, Lower Austria) by benthic copepods

The role of copepods in the colonization and emigration processes of benthos and drift of an irrigation canal were studied. During the 3 first years of existence of the canal, fifteen species of copepods were recorded (1 diaptomid, 11 cyclopoids, 3 harpacticoids). Copepod succession in the canal was initially represented by lentic species, then by predatory species like Macrocyclops albidus that developed with the proliferation of filamentous algae and epibionts during the first two years. Primary consumers appeared from the beginning of the flowing conditions but started to dominate only from the third year. Smaller species associated with the bed sediments like the harpacticoid species Nitocra hibernica, were late colonizers. The latter as well as Paracyclops fimbriatus and Eucyclops serrulatus were the dominant copepod species in the canal. Total copepod density in the benthos ranged from 15,800 to 68,000 ind./m², with a mean of 7.6% of the total zoobenthos. Copepods constituted up to 25% of the total faunal drift density, ranging from 50 to 140 ind./m³. Their highest abundance was observed around midnight. A three-fold increase of the discharge temporarily affected the drift density, but did not change the diel rhythm of the drift. Cyclopoid copepods namely P. fimbriatus (54.77%), copepodid cyclopoids (20.76%) and E. serrulatus (16.11%) formed the main part of the drift samples. The harpacticoid Nitocra hibernica was a minor participant in the drift processes.     

The female antennules of Paracyclops (Cyclopoida: Cyclopidae): their significance for systematics

The setation patterns of female antennules of some Paracyclops species are analysed with reference to the hypothetical 28-segmented antennule of ancestral copepod proposed by Huys and Boxshall in 1991. Comparison between Paracyclops species has revealed a setation pattern common to all species thus far examined. Several modifications of the female antennules have been identified as apomorphic states and can be used in phylogenetic studies. P. waiariki has 12-segmented antennules and exhibits more plesiomorphic characters than the reduced 8-segmented antennules of other species, such as P. fimbriatus. The results confirm that simple segment numbers are unusable as characters in cladistic analysis. Both P. affinis and P. yeatmani, for example, have 11-segmented antennules but analysis of segmental homologies has revealed that the 11-segmented state is convergent in these two species.     

Environmental impact assessment of sediment dumping in the southern Baltic Sea using meiofaunal indicators

An experimental sediment dumping was carried out in the southern part of the Mecklenburg Bight in June 2001. Foraminiferans and ostracods from superficial sandy sediment were studied in a time series from before dumping until March 2004 in order to assess changes in associations and recolonization patterns of both groups. Additionally, an area sampling covering the dumping site and its surroundings from 15.5 to 20.7 m water depth made it possible to compare associations inside and outside the dumping area as well as the water depth dependent distribution of foraminiferans and ostracods. Salinity values vary within the high alpha-mesohaline and low polyhaline range. The dominating species are Ammotium cassis (Foraminifera) and Sarsicytheridea bradii (Ostracoda). The diversity is low (Fisher alpha index from 0.4 to 3.2 for foraminiferans and 1.0 to 2.5 for ostracods), but higher within the dumping site samples. These higher values are explainable by input of allochthonous tests and valves representing additional species. After the sediment dumping it took two and a half years to re-establish the total foraminiferan association and the total foraminifer/ostracod ratio within the dumping site. Total foraminiferan abundance increases remarkably with water depth (mean 83 tests in 100 ml) driven by higher nutrient availability and more suitable salinity and temperature values within the zone of the oscillating halocline. The distribution of shallow water species such as Cribroelphidium excavatum, Eucythere argus and Hirschmannia viridis, within the transient water layer A. cassis, Nodulina dentaliniformis, S. bradii and Palmoconcha laevata and below Eggerella scabra indicate the depth position of the halocline. Water depth and sediment dumping influence are the main driving factors for the distribution of foraminifer and ostracod associations within the study area. However, a significant sedimentological difference between samples inside and outside the dumping area is not recognizable.     

Succession of the ecosystems of the Aral Sea during its transition from oligohaline to polyhaline water body

During 22 field trips from 1990 to 2002 (mainly the western basin of the Large Aral) data on salinity, phytoplankton, zooplankton, zoobenthos and fish fauna have been collected. In 2002, the salinity of the western basin reached 75 ppt, while that in the eastern basin, 150 ppt. In 1999–2002, 159 species of planktonic algae have been recorded. This is approximately twice as low as recorded before. The diversity of Cyanophyta, Pyrrhophyta and Chlorophyta in particular has dropped in the past few years. As before, currently Bacillariophyta is the most diverse plankton. However, the composition of dominants has changed. Once previously dominant species, Actinocyclus ehrenbergii, vanished from the plankton of the Aral Sea and was replaced by such diatoms as Amphora coffeaformis, A. coffeaformis var. acutiuscula and Synedra tabulata var. parva.Since 1970s, a gradual decrease in the diversity of zooplankton has been taking place. Since 1997, the formerly dominant Calanipeda aquaedulcis vanished, which apparently was the reason for the emergence of Moina salina and Artemia parthenogenetica. Since 2000, artemia has been dominant in the plankton of the Aral Sea, constituting 99% of the zooplankton biomass.In the 1970–1980s, a rapid decrease in the biodiversity of the zoobenthos was observed. In the 1990s, most aboriginal and introduced species became extinct. Currently, the bivalve mollusk Syndosmya segmentum, the ostracod Cyprideis torosa and larvae of the dipteran Chironomus salinarius can still be recorded in the western basin. In the eastern basin no benthos is observed.By 1998, in the Large Aral, only five fish species survived: baltic herring Clupea harengus membras, flounder Platichthys flesus luscus, atherine Atherina boyeri caspia and bullheads Neogobius fluviatilis and Potamoschistus caucasicus. Since 2002, only flounder and atherina have been recorded in the western basin of the Large Aral. No fish have been recorded in the eastern part of the Aral Sea in 2002.With increasing salinity and transition of the Aral Sea from an oligohaline to a polyhaline water body, its biota is becoming drastically poorer. Almost all local species became extinct in the Aral; however, some still survive (including some endemics) in some lakes around the Aral Sea. In the near future, artemia will be the only animal in the Aral Sea.     

Theme Issue 30(2): "Economics of Marine Protected Areas"

Most Florida manatees depend on localized warm-water refuges in the southern two-thirds of Florida to survive winter; about 60% use outfalls from 10 power plants, whereas 15% use 4 natural warm-water springs. Future availability of these refuges is in doubt; most of these power plants may be retired within the next 20 years and groundwater withdrawals for human use threaten natural springs. This article examines possible effects on manatees from losing major warm-water refuges and alternative management actions. Because of manatee site-fidelity patterns, plant retirements may increase cold-stress-related deaths and significantly decrease manatee abundance. A forward-looking management strategy is urgently needed before decisions are made to retire plants now used by large numbers of manatees. Possible management alternatives include: gradually weaning manatees off plant outfalls, maintaining the flow of springs now used by manatees, enhancing access to suitable warm-water springs now little used or unused by manatees, constructing new non-industry dependent warm-water refuges, and creating new thermal basins to retain warm-water pockets able to support overwintering manatees.     

The egg morphology of 7 Acartiidae species: a preliminary survey of the ootaxonomy of calanoids

This is the first contribution to acartiid taxonomy based on egg morphology. Eggs laid under laboratory conditions by maintaining female specimens of Acartia adriatica, A. clausi, A. italica, A. margalefi, A. tonsa, Paracartia latisetosa and Pteriacartia josephinae were studied by S.E.M. Most species produced eggs with intraspecific variation in their external morphology. In spite of such variation, however, it was possible to separate three species groups.     

Trace element levels in some commercially valuable fish species from coastal waters of Mediterranean Sea, Egypt

Concentrations of 5 trace elements (Cd, Pb, Cu, Cr, Zn) were determined in muscle of fishes collected from two coastal areas of the Egyptian coast of the Mediterranean Sea west of Alexandria (El-Mex Bay and Eastern Harbour). Sardinella aurita, Alepes djedaba, Siganus luridus, Siganus rivulatus, Sphyraena chrysotoenia, and Scomberomorus commerson were collected from El-Mex Bay. While Boops boops, Lithognathus mormyrus, Sparisoma cretense, Serranus cabrilla and Synodus saurus were collected from the E.H. In all fishes zinc was highest (up to 57 mg/kg) followed by Cr, Cu, Pb and Cd. The concentrations of several metals in fishes were significantly different among the species in each area. For all trace element examined, the concentrations decreased significantly with body weight of some fishes. In contrast, a positive correlation with body weight was found especially for Zn and Cu concentrations in 5 fish species and for Pb in 4 fish species. The concentrations of Cd, Pb and Zn were higher in fishes from El-Mex Bay (3.76, 6.49 and 57.21 mg/kg) than those from Eastern Harbour, whereas the opposite trend was observed for Cu and Cr, revealing the direct sources of trace metal pollution present in El-Mex coastal area. Levels of Cr surpassed the Maximum Permissible Concentration in most fish tissues, followed by Pb and Cd in some species. Cu and Zn presented concentrations below the maximum permissible levels in fish tissues.     

Seasonal carbon distribution of copepods in the eastern Weddell Sea, Antarctica

Copepods were sampled by a multiple opening-closing net in the eastern Weddell Sea during various seasons (late winter/early spring, summer, autumn). Total copepod biomass integrated over the upper 1000 m varied seasonally between 1.7 mg C m⁻³ in late winter/early spring and 3.7 mg C m⁻³ in autumn. After the dark season the copepods were rather evenly distributed vertically and highest biomass levels were found in the mid-water layers between about 200 m and 500 m. By contrast, especially in summer but also in autumn copepod biomass concentrated in the uppermost water layer. A total of 64 calanoid species were identified in the upper 1000 m with maximum species numbers in the deepest layer. The large calanoids Calanus propinquus, Calanoides acutus, Metridia gerlachei, Euchaeta antarctica and the small calanoid Microcalanus pygmaeus prevailed and accounted for 60–70% of total copepod biomass, while the small poecilostomatoid Oncaea and the cyclopoid Oithona species comprised about 20%. Hence, the distribution pattern of the entire copepod biomass is strongly influenced by the life cycles of a few dominant species.     

Mesozooplankton species distribution in the NW and N Iberian shelf during spring 2004: Relationship with frontal structures

We have analysed the mesozooplankton community structure in the southern Bay of Biscay shelf and its relationship with the hydrographic conditions during spring 2004. According to thermohaline characteristics, we observed two frontal zones of distinct origin along the shelf (around 7° and 3°W), that allowed us to differentiate three different hydrographic domains. The westernmost part of the shelf (WC), defined by the presence of relatively warm and salty water related to the presence of the Iberian Poleward Current (IPC), the easternmost region (EC), characterized by colder and fresher water and subject to the influence of freshwater inputs from the Adour river in the French coast, and a region in the Central Cantabrian Sea (CC), where thermohaline characteristics were intermediate between these two extremes. The mixing layer depth (MLD) regime in these areas was also different: the WC region was characterized by a mixed water column, whereas in the EC region the river discharges produces stratification of the upper meters of the water column (< 10 m); in the CC region, we found a distinct vertical mixing regime that separated coastal (stratification) from shelf (mixed water column) stations, giving rise to a notorious across-shelf front. We found a good match between the aforesaid hydrographic regions and the distribution of mesozooplankton species composition and community assemblages: the Mantel correlation between physical variables and mesozooplankton distribution was highly significant (n = 63, r = 0.70, α < 0.001). In the WC region, the community was dominated by Paracalanus parvus, Oithona helgolandica, Acartia clausi and Clausocalanus pergens, while in the EC region the most dominant species were Noctiluca scintillans, Oncaea media and Temora longicornis. The CC region showed similar composition of copepods than the WC region, but larvaceans (Oikopleura spp. and Fritillaria spp.) were more abundant in the CC region than in the WC region. Within each zone, the relative abundances of the dominant species differed between coastal and shelf locations.     

Life cycle strategies of epipelagic copepods in the Southern Ocean

Twelve epipelagic copepod species were reviewed to compare their adaptations to the short primary production season and low temperatures which characterise the Southern Ocean. The species show a spectrum of adaptations, but three broad life cycle strategies were defined: (1) herbivorous in summer, a short reproductive period and winter diapause at depth (Calanoides acutus and possibly Ctenocalanus citer); (2) predominantly omnivorous/detritivorous diet, an extended period of feeding, growth and reproduction and less reliance on diapause at depth (Metridia gerlachei, Calanus propinquus, Calanus simillimus, Oithona similis, Microcalanus pygmaeus, and possibly Oncaea curvata and Oithona frigida); (3) overwintering and feeding within sea ice as early nauplii or copepodids (Stephos longipes and Paralabidocera antarctica). The large species Rhincalanus gigas appears to be intermediate between strategies (1) and (2). Contrasting species from groups (1) and (2), namely C. acutus and O. similis, were selected for more detailed comparison. For C. acutus, maximum (probably food saturated) feeding and egg production rates are well below equivalent values for Calanus spp. at lower latitudes. Likewise, summer growth and moulting rates are slower, and the growth season of this herbivore is only 2–4 months. Therefore, both the low summer temperatures and short primary production season seem to dictate a long (1 year) life cycle for C. acutus. A collation of data on O. similis revealed that its abundance increases about tenfold from the Antarctic shelf northwards to the Polar Frontal Zone, where abundances are similar to those in temperate and tropical shelf seas. In contrast to C. acutus, O. similis appears to remain in the epipelagic and reproduce there year-round, although the food sources which sustain this are still uncertain.     

Seasonal productivity dynamics in the pelagic central Benguela System inferred from the flux of carbonate and silicate organisms

Flux of bulk components, carbonate- and silicate-bearing skeleton organisms, and the δ¹⁵N-isotopic signal were investigated on a 1-year time-series sediment trap deployed at the pelagic NU mooring site (Namibia Upwelling, ca. 29°S, 13°E) in the central Benguela System. The flux of bulk components mostly shows bimodal seasonality with major peaks in austral summer and winter, and moderate to low export in austral fall and spring. The calcium carbonate fraction dominates the export of particulates throughout the year, followed by lithogenic and biogenic opal. Planktonic foraminifera and coccolithophorids are major components of the carbonate fraction, while diatoms clearly dominate the biogenic opal fraction. Bulk δ¹⁵N isotopic composition of particulate matter is positively correlated with the total mass flux during summer and fall, while negatively correlated during winter and spring. Seasonal changes in the intensity of the main oceanographic processes affecting the NU site are inferred from variations in bulk component flux, and in the flux and diversity patterns of individual species or group of species. Influence from the Namaqua (Hondeklip) upwelling cell through offshore migration of chlorophyll filaments is stronger in summer, while the winter flux maximum seems to reflect mainly in situ production, with less influence from the coastal and shelf upwelling areas. On a yearly basis, dominant microorganisms correspond well with the flora and fauna of tropical/subtropical waters, with minor contribution of near-shore organisms. The simultaneous occurrence of species with different ecological affinities mirrors the fact that the mooring site was located in a transitional region with large hydrographic variability over short-time intervals.     

The role of pelagic fish as forage for the demersal fish community in the southern Bay of Biscay

The small pelagic fish represents the link between nekton and demersal communities, and they are an important food source for some demersal fish species. In the autumns between 1993 and 2002, 74,869 stomach contents of 25 demersal fish species were analysed during the scientific surveys organized by the Instituto Español de Oceanografía. These species represent the demersal fish community of the southern Bay of Biscay. An important part of their diet (39% by volume) was composed of pelagic fish species (e.g. Engraulis encrasicolus, Gadiculus argenteus, Micromesistius poutassou, Trachurus trachurus, Sardina pilchardus, Scomber scombrus). Among all these prey species, the relevance of M. poutassou and G. argenteus stood out because of their high abundance both in the diets and during the scientific surveys. The relevance of pelagic fish as prey increased with predator size, reaching more than 60% of the diet by volume in the length range 25–29 cm. However, fish predators larger than 50 cm depended less on pelagic fish (33%), since they were also able to feed on other sources such as megafaunal invertebrates, mainly cephalopods. The pelagic fish resource was primarily exploited by 12 demersal fish species, with Merluccius merluccius and Zeus faber being the main ones feeding on small pelagics, which was related to the vertical movement of predators and prey through the water column. Survey abundance indices were used as indicators of prey abundance in the ecosystem, both for biomass and number. There was evidence for density-dependant feeding by predators on E. encrasicolus, G. argenteus and T. trachurus, while the main discrepancies between abundance in the stomachs and in the surveys were due to differential availability of prey length classes in the environment. Small prey species and individuals were exploited by most demersal fish while large prey species were less accessible to predators.     

Human Impacts on Cable Beach,Broome (Western Australia)

There are concerns that Cable Beach, which is of outstanding natural beauty and a major tourist attraction for Broome and the Kimberley region of northwest Australia, is suffering damage from recreational use, particularly that caused by off-road vehicles. To assess current levels of human impact on the shore and its fauna, the southern-most kilometer of the Beach, covering an area to which vehicles had access and an adjacent area closed to vehicles, was surveyed for human usage and shore crab abundance. Human usage over the recording period was high in the area with high vehicular access. The amount of litter, although considerable, was well below that recorded in some other tourist destinations elsewhere in the world. Burrows of both the ghost crabs Ocypode spp, and the sand bubbler, Scopimera inflata, were less numerous in areas of high vehicle use than those of low use. Both species may be valuable in monitoring human impacts on Cable Beach.     

Morphological and biochemical differentiation in Antarctic krill

During the February 1981 cruise FIBEX MD-25 between 30–50°E and 61–64°S, hydrography showed the presence of two gyres, confirmed by the geostrophic circulation relative to 1000 m from Levitus climatology, at the borders of these gyres concentrations of highly morphologically differentiated krill were found. Gaussian component analysis of krill samples, pooled by sectors, showed three cohorts of Euphausia superba in the western sector and one in the eastern sector. Across the sampling area, Thysanoessa macrura and E. superba occurred at separate stations. Analysis of cohorts in T. macrura separated two size groups in both the western and the eastern sectors. The use of a Differentiation Index (D.I.) [Färber-Lorda, J., 1990. Somatic length relationships and ontogenetic morphometric differentiation of Euphausia superba and Thysanoessa macrura of the southwest Indian Ocean during summer (February 1981). Deep-Sea Res. 37, 1135–1143.], based on somatic lengths, allows studying certain morphological differences within the populations sampled. Morphologically different and bigger males II (D.I. from 2.8 to 3.5) were present only in the southern transect while smaller males I (D.I. from 3.5 to 5.0) were present over the entire area. Biochemical composition of both species showed significant differences among stations for protein, lipids, and carbohydrates. A significant difference in lipid content was found between males I, and males II. For T. macrura, percentage of lipid content in mature animals was much higher than that in E. superba. The D.I. size distribution showed that when populations of E. superba were highly differentiated (corresponding to mature animals) in morphology, lipid content was high, and they were located near a gyre. Differences in morphometry can influence distribution of the species, because different developing stages have different swimming capacities. It is shown that, together with hydrography and trophic conditions, lipid content and morphometry of krill populations, are different but complementary aspects that help to understand krill ecology and distribution.     

Ecophysiological growth characteristics and modeling of the onset of the spring bloom in the Baltic Sea

The onset of spring bloom in temperate areas is a transition period where the low productive, winter phytoplankton community is transformed into a high productive spring community. Downwelling irradiance, mixing depth and the ability of the phytoplankton community to utilize the light, are key parameters determining the timing of the onset of the spring bloom. Knowing these parameters would thus provide tools for modeling the spring bloom and enhance our knowledge of ecophysiological processes during this period.Our main objective with this study was to provide data for the growth characteristics of some key species forming the spring bloom in the Gulf of Finland, and to apply those results in a simple dynamic model for the onset of the spring bloom, in order to test if the timing of the spring bloom predicted by the models corresponds to field observations. We investigated the photosynthetic characteristics of three diatoms and two dinoflagellates (Chaetoceros wighamii, Melosira arctica, Thalassiosira baltica, Scrippsiella hangoei and Woloszynskia halophila), at low temperatures (4–5 °C). All of these species are common during spring bloom in the Baltic Sea.Cultures of these species were acclimated to different irradiance regimes prior to measurements of photosynthesis, respiration, pigment concentration and light absorption. We did not find a positive relationship between respiration and growth rate, and we hypothesize that this relationship, which is well established at higher temperatures, is negligible or absent at low temperatures (< 10 °C). Photosynthetic maximum (P_m), and maximum light utilization coefficient (α) was lowest and respiration (R) highest in the dinoflagellates.We made a model of the onset of the spring bloom in the western part of Gulf of Finland, using the obtained data together with monitoring data of mixing depth and water transparency from this area. Model results were compared to field observations of chlorophyll-a (Chl-a) concentration. There was a good agreement between the model predictions and the observed onset of the spring bloom for the diatoms. S. hangoei, however, was not able to reach positive production in the model, and W. halophila had the similar growth characteristics as S. hangoei. Consequently, these species must have other competition strategies enabling them to exist and grow during spring bloom.     

Marine molluscs as biomonitors for heavy metal levels in the Gulf of Suez, Red Sea

Levels of the heavy metals Copper (Cu), Zinc (Zn), Lead (Pb), Cadmium (Cd), Chromium (Cr), Nickel (Ni), Iron (Fe) and Manganese (Mn) were determined in coastal water, sediments and soft tissues of the gastropod limpet, Patella caerulea, and the bivalve, Barbatus barbatus, from seven different stations in the western coast of the Gulf of Suez. The concentrations of heavy metals in water ranged between 3.37–4.78, 18.83–21.46, 2.75–3.17, 0.22–0.27, 0.99–1.21, 2.69–3.65, 3.75–4.56 μg L^− 1 and 23.82–32.78 mg g^− 1 for Cu, Zn, Pb, Cd, Cr, Ni, Mn and Fe, respectively. The corresponding concentration values in the sediments were 8.65–12.16, 51.78–58.06, 36.52–42.15, 3.23–3.98, 9.03–12.75, 34.31–49.63, 3.28–4.56 and 64.20–70.22 μg g^− 1 for Cu, Zn, Pb, Cd, Cr, Ni, Mn and Fe, respectively. The highest accumulated metals were Fe, Zn and Mn in both P. caerulea and B. barbatus, while the lowest one was Cd. The accumulation of metals was more pronounced in P. caerulea than B. barbatus. The highest concentrations of all metals in water, sediments and mollusca were recorded at Adabiya harbour north of the Gulf, while the lowest concentrations were recorded at Gabal El-Zeit and Hurghada. Land based activities and ships awaiting berth are the main source of metal pollution in the northern part of the Gulf.     

Effects of closed trawl fishery areas on some elasmobranch species in the Cantabrian Sea

This study examines the effects of certain closed areas to the trawl fishery on some Rajidae species and small-spotted catshark in the southern area of the Bay of Biscay. The results are based on abundance index (number/haul) and catch length composition obtained from a series of bottom trawl surveys carried out in the area (1993–2005). Abundance trends are similar considering or not closed areas, but with much lower values excluding closed areas. The length distributions revealed that in some of the species like Scyliorhinus canicula the larger specimens were the most affected by this decrease in number. Finally, the effects and consequences of one of these marine protected areas (MPAs), an artificial reef were examined in more detail for all the fishes and invertebrates. The analysis of the yield series showed an increase in total biomass after the placement of the reefs not only for S. canicula and rays but also teleosts fishes like Pagellus acarne, Trachurus trachurus, Mullus surmuletus and Trachinus draco and particularly invertebrates (sea urchins). On the contrary, the main target species of the trawl fishery (hake, monkfish and megrim among others) did not increase.     

Particle flux, and composition of sedimenting matter, in the Greenland Sea

Vertical flux of particulate material was recorded with moored sediment traps during 1988/1989 in the Greenland Sea at 72°N, 10°W. This region exhibits pronounced seasonal variability in ice cover. Annual fluxes at 500 m water depth were 22. 79, 8.55, 2.39, 3.81 and 0.51 g m⁻² for total flux (dry weight), carbonate particulate biogenic silicate, particulate organic carbon and nitrogen, respectively. Fluxes increased in April, maximum rates of all compounds occurred in May–June, and consistently high total flux rates of around 100 mg m⁻²d⁻¹ prevailed the summer. The increasing flux of biogenic particles measured in April is indicative of an early onset of algal growth in spring. Small pennate diatoms dominated in the trap collections during April, and were still numerous during the high flux period when Thalassiosira species were the most abundant diatoms. During May–June, up to 22% of the Thalassiosira cells collected were viable-looking cells. The faecal pellet flux increased after the May–June event. Therefore we conclude that the diatoms settled as phytodetritus, most likely in rapidly sinking aggregates. From seasonal nutrient profiles it is concluded that diatoms contribute 25% to new production during spring and 50% on an annual basis. More than 50% of newly produced silicate particles are dissolved above the 500 m horizon. High new production during spring does not lead to a pronounced sedimentation pulse of organic matter during spring but elevated vertical export is observed during the entire growth period.