Distribution and abundance of marine wood borers on the west coast of Peninsular Malaysia

Twenty one species of marine wood borers were recorded in this study. There are seventeen species of bivalves of which sixteen are from the family Teredinidae and one from the family Pholadidae. The crustaceans comprised the family Sphaeromatidae with three species and the family Limnoriidae with one species. Seven of the fourteen known genera of the teredinids were recorded. The bivalve wood borers were widely distributed in the local mangroves. Martesia striata was the most abundant wood borer at the Degaussing Range jetty, Lumut. Lyrodus pedicellatus was the most abundant shipworm in wooden panels. Shipworms were abundant on the subtidal panels where the genus Bankia, was most common while M. striata was abundant on the intertidal panels. M. striata numbers were higher on the top surfaces as compared to the under surfaces of wooden panels.     

Seasonal settlement and succession of fouling communities in Kalpakkam, east coast of India

Seasonal distribution and community succession of macrofoulants were studied using concrete panels in the coastal waters of Kalpakkam, east coast of India, for a period of two years. The panels were suspended at 1 m, 4 m and 7 m depths and categorised into short-term and long-term exposures. A high total of 105 fouling taxa were recorded. The major fouling organisms observed were hydroids, barnacles, mussels, anthozoans and ascidians. Considerable faunistic and biomass variations were noticed both with respect to season and depth. The month of panel exposure had a significant influence on the succession of fouling communities. On the short-term panels, the maximum fouling biomass was 64 kg m^–2 in 30 days at 4 m depth, whereas on the long-term panels, it was 250 kg m^–2 after 216 days at 4 m depth. A comparison with the biomass values reported from elsewhere shows that biomass build-up in Kalpakkam coastal waters is one of the highest ever reported. Such a very high biomass accumulation is due to the extremely dense settlement of mussels, especially the green mussel,Perna viridis (L).     

A lifetime at depth: vertical distribution of southern elephant seals in the water column

Although numerous studies have addressed the migration and dive behaviour of southern elephant seals (Mirounga leonina), questions remain about their habitat use in the marine environment. We report on the vertical use of the water column in the species and the potential lifetime implications for southern elephant seals from Marion Island. Long-term mark-resight data were used to complement vertical habitat use for 35 known individuals tagged with satellite-relay data loggers, resulting in cumulative depth use extrapolated for each individual over its estimated lifespan. Seals spent on average 77.59% of their lives diving at sea, 7.06% at the sea surface, and 15.35% hauled out on land. Some segregation was observed in maximum dive depths and depth use between male and female animals—males evidently being physiologically more capable of exploiting increased depths. Females and males spent 86.98 and 80.89% of their lives at sea, respectively. While at sea, all animals spent more time between 300 and 400 m depth, than any other depth category. Males and females spent comparable percentages of their lifetimes below 100 m depth (males: 65.54%; females: 68.92%), though males spent 8.98% of their lives at depths in excess of 700 m, compared to females’ 1.84% at such depths. Adult males often performed benthic dives in excess of 2,000 m, including the deepest known recorded dive of any air-breathing vertebrate (>2,133 m). Our results provide a close approximation of vertical habitat use by southern elephant seals, extrapolated over their lifespans, and we discuss some physiological and developmental implications of their variable depth use.     

Marine depth use of sea trout Salmo trutta in fjord areas of central Norway

The vertical behaviour of 44 veteran sea trout Salmo trutta (275–580 mm) in different marine fjord habitats (estuary, pelagic, near shore with and without steep cliffs) was documented during May–February by acoustic telemetry. The swimming depth of S. trutta was influenced by habitat, time of day (day v. night), season, seawater temperature and the body length at the time of tagging. Mean swimming depth during May–September was 1·7 m (individual means ranged from 0·4 to 6·4 m). Hence, S. trutta were generally surface oriented, but performed dives down to 24 m. Mean swimming depth in May–September was deeper in the near‐shore habitats with or without steep cliffs (2·0 m and 2·5 m, respectively) than in the pelagic areas (1·2 m). May–September mean swimming depth in all habitats was slightly deeper during day (1·9 m) than at night (1·2 m), confirming that S. trutta conducted small‐scale diel vertical movements. During summer, S. trutta residing in near‐shore habitat progressively moved deeper over the period May (mean 1·1 m) to August (mean 4·0 m) and then reoccupied shallower areas (mean 2·3 m) during September. In winter (November and February), individuals residing in the innermost part of the fjords were found at similar average depths as they occupied during the summer (mean 1·3 m). The swimming depths of S. trutta coincide with the previously known surface orientation of salmon lice Lepeophtheirus salmonis. Combined with previous studies on horizontal use of S. trutta, this study illustrates how S. trutta utilize marine water bodies commonly influenced by anthropogenic factors such as aquaculture, harbours and marine constructions, marine renewable energy production or other human activity. This suggests that the marine behaviour of S. trutta and its susceptibility to coastal anthropogenic factors should be considered in marine planning processes.     

Treatment at different depths and vertical mixing within a 1-m deep horizontal subsurface-flow wetland

The aim of this study was to examine the variation in treatment performance at three depths, and the degree of vertical mixing, within a 1.0 m deep horizontal subsurface-flow constructed wetland (HSSF-CW) planted with Schoenoplectus tabernaemontani (Gmel.) Palla, and treating primary settled municipal wastewater in sub-tropical New South Wales, Australia. Water samples were collected from the upper (0.17 m), middle (0.5 m), and lower (0.83 m) depths at five equi-spaced sample points along the longitudinal axis of the 8.8 m² bed during two trials. Analysis of covariance (ANCOVA) indicated that the rate of pollutant concentration reduction between the three depths was not significantly different (p > 0.05) for all of the measured parameters (dissolved oxygen (DO), hydrogen electrode potentials (E_h), 5-day biochemical oxygen demand (BOD₅)) total nitrogen (TN), TKN, and NH₄-N. Thus, it can be concluded that the break-down of contaminants as wastewater moved through the HSSF-CW was approximately uniform across the 1.0 m depth profile. The lack of a significant depth effect can be largely explained by the substantial amount of vertical mixing that was observed when a pulse of lithium tracer was injected into the middle depth of the first intermediate sampling point. The tracer rapidly migrated vertically into the upper and lower depths as water moved through the bed and was almost completely mixed between the three depths by the time it reached the last intermediate sampling point.The majority of BOD₅ removal occurred within the first-third of the bed where vegetation cover was poor. Performance of the bed declined over time from Trial 1 to Trial 2, possibly due to a cumulative build-up of organic matter within the substrate as a result of limited oxygen transfer throughout the 1.0 m depth of substrate via root leakage or diffusion across the air–water interface. Root penetration was limited to the upper 0.4 m of the substrate, with the majority of below-ground biomass forming a dense mat in the upper 0.2 m. A comparison of two-parameter (K–C^*) first-order volumetric rate constant (K_v20) with those obtained from 0.4 to 0.6 m deep HSSF-CW in the same region indicate that a doubling of the wetted depth resulted in no improvement in BOD₅ removal and a decline in TN removal on an areal basis. Further investigations are warranted, comparing the performance of replicated beds spanning a range of depths (e.g. 0.25, 0.5 and 1.0 m) in order to quantitatively determine the optimal depth of HSSF-CWs treating domestic wastewater.     

Geographic differences in vertical connectivity in the Caribbean coral Montastraea cavernosa despite high levels of horizontal connectivity at shallow depths

The deep reef refugia hypothesis proposes that deep reefs can act as local recruitment sources for shallow reefs following disturbance. To test this hypothesis, nine polymorphic DNA microsatellite loci were developed and used to assess vertical connectivity in 583 coral colonies of the Caribbean depth‐generalist coral Montastraea cavernosa. Samples were collected from three depth zones (≤10, 15–20 and ≥25 m) at sites in Florida (within the Upper Keys, Lower Keys and Dry Tortugas), Bermuda, and the U.S. Virgin Islands. Migration rates were estimated to determine the probability of coral larval migration from shallow to deep and from deep to shallow. Finally, algal symbiont (Symbiodinium spp.) diversity and distribution were assessed in a subset of corals to test whether symbiont depth zonation might indicate limited vertical connectivity. Overall, analyses revealed significant genetic differentiation by depth in Florida, but not in Bermuda or the U.S. Virgin Islands, despite high levels of horizontal connectivity between these geographic locations at shallow depths. Within Florida, greater vertical connectivity was observed in the Dry Tortugas compared to the Lower or Upper Keys. However, at all sites, and regardless of the extent of vertical connectivity, migration occurred asymmetrically, with greater likelihood of migration from shallow to intermediate/deep habitats. Finally, most colonies hosted a single Symbiodinium type (C3), ruling out symbiont depth zonation of the dominant symbiont type as a structuring factor. Together, these findings suggest that the potential for shallow reefs to recover from deep‐water refugia in M. cavernosa is location‐specific, varying among and within geographic locations likely as a consequence of local hydrology.     

Age- and size-based morphological comparison between the brown alga Sargassum macrocarpum (Heterokonta; Fucales) from different depths at an exposed coast in northern Kyoto, Japan

Although large perennial brown algae are known to show great morphological plasticity responding to specific environmental factors such as depth and wave exposure, there are few reports showing this morphological variability taking into account age or size composition. Here, we conducted age- and size-based morphological comparison between the perennial brown alga Sargassum macrocarpum C. Agardh from their upper depth limit, middle depth, and lower depth limit at an exposed coast in northern Kyoto, Japan. Model II regression was fitted for the relationships between age or stipe diameter (SD) and morphological variables including SD, holdfast weight (HW), number of main branches (MBN), total weight of main branches (MBW), thallus weight (TW), and thallus length (TL) of 30 specimens collected seasonally from each depth zone. The differences between depths in these regression equations were analyzed using SMATR. Although SD, HW, and MBN of the specimens did not differ between depths under both age- and SD-basis, there were significant differences between depths in MBW, TW, and TL, suggesting that the differences in TW between depths are resulted from differences in MBW. Whereas TW at the lower depth limit was lower than that at the upper depth limit or middle depth in many months, TL at the upper depth limit was shorter than that at the middle depth or lower depth limit in some months. These results suggest that S. macrocarpum at shallower depths tend to have short and bushy morphology, whereas those at deeper depths have long and less bushy morphology.     

The fundamental niche of blood oxygen binding in the pelagic ocean

Marine species ranging in size from microscopic zooplankton to large predatory fish move vertically in the ocean water column to forage for food and avoid predators. Oxygen and temperature decrease, often rapidly, from shallow to deeper depths, restricting the ability of species to use the vertical habitat. One physiological trait that determines the tolerance of organisms to low oxygen is the oxygen affinity of oxygen carrier proteins, hemoglobin and hemocyanin, in the blood. To quantify the range of oxygen affinities for marine organisms, we surveyed the literature for measurements of oxygen binding to blood at multiple temperatures to account for its temperature sensitivity. Oxygen affinity is mapped within the ocean environment using the depth at which oxygen pressure decreases to the point at which the blood is 50% oxygenated (P₅₀ depth) as organisms move from the surface to depth in the ocean water column. We find that vertical gradients in both temperature and oxygen impact the vertical position and areal extent of P₅₀ depths. Shifts in P₅₀ due to temperature cause physiological types with the same P₅₀ in the surface ocean to have different P₅₀ depths and physiological types with different P₅₀'s in the surface ocean to have the same P₅₀ depth. The vertical distances between P₅₀ depths are spatially variable, which may determine the frequency of ecological interactions, such as competition and predation. In summary, P₅₀ depth, which represents a key physiological transition point between dexoxygenated and oxygenated blood, provides mechanistic insight into organism function within the water column of the global ocean.     

Vertical zonation and functional diversity of fish assemblages revealed by ROV videos at oil platforms in The Gulf

An assessment of vertical distribution, diel migration, taxonomic and functional diversity of fishes was carried out at offshore platforms in The (Arabian–Iranian–Persian) Gulf. Video footage was recorded at the Al Shaheen oil field between 2007 and 2014 using a remotely operated vehicle (ROV). A total of 12 822 individual fishes, from 83 taxonomic groups were recorded around the platforms. All the species identified are considered native to The Gulf, although Cyclichthys orbicularis and Lutjanus indicus were recorded for the first time in Qatari waters. Several trends were uncovered in the vertical distribution of the fish community; most species were observed between 20 and 50 m depth and fish abundance decreased towards the bottom, with the highest abundances recorded in the upper layers, i.e. down to 40 m depth. Vertical variation in fish diversity, however, was generally not accompanied by differences in vertical movements. Carnivores and invertivores were the dominant trophic groups, being found at each depth range from surface to seabed. The functional indices showed no significant differences between water depths or diel cycles. The study demonstrates that oil platforms represent a hotspot of fish diversity and interesting sites for studying fish communities, abundance and behaviour.     

Rhodolith beds at the easternmost extreme of South America: Community structure of an endangered environment

Rhodolith beds are built by the aggregation of free living marine benthic coralline algae. Herein, we described phytobenthic communities associated with subtidal rhodolith beds in northeastern Brazil and tested the hypothesis that depth affects their structure. We compared macroalgal assemblages from depths of 10, 15 and 20 m. The genus Lithothamnion was dominant in these beds. Rhodolith density was similar at different depths, but volume decreases as depth increases. Sixty-seven species of fleshy algae were collected. The red algal order Ceramiales was dominant. A distinct community corresponds to each sampled depth. The shallower depth presented higher values for biomass, number of species, Shannon-Wiener diversity, and Pielou's evenness. When depth and water transparency increased, the number of species and the abundance of macroalgae decreased.     

Variables associated with the occurrence of Ips beetles,red turpentine beetle and wood borers in live and dead ponderosa pines with post‐fire injury

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Migration depths of adult steelhead Oncorhynchus mykiss in relation to dissolved gas supersaturation in a regulated river system

Adult steelhead Oncorhynchus mykiss tagged with archival transmitters primarily migrated through a large river corridor at depths >2 m interspersed with frequent but short (<5 min) periods closer to the surface. The recorded swimming depths and behaviours probably provided adequate hydrostatic compensation for the supersaturated dissolved gas conditions encountered and probably limited development of gas bubble disease (GBD). Results parallel those from a concurrent adult Chinook salmon Oncorhynchus tshawytscha study, except O. mykiss experienced greater seasonal variability and were more likely to have depth uncompensated supersaturation exposure in some dam tailraces, perhaps explaining the higher incidence of GBD in this species.     

Damaging UV radiation and invertebrate predation: conflicting selective pressures for zooplankton vertical distribution in the water column of low DOC lakes

In nature most organisms have to manage conflicting demands of food gathering, predator avoidance, and finding a favorable abiotic environment (oxygen, temperature, etc.) in order to maximize their fitness. In the vertical water column of lakes with high solar ultraviolet radiation (UV) and invertebrate predators, zooplankton face two particularly strong and conflicting selective pressures. During daylight hours invertebrate predators often induce an upward vertical migration of zooplankton prey while potentially damaging UV forces a downward migration. We used 2.2 m long columns suspended vertically in a lake to conduct 2×2 factorial experiments to examine patterns of depth selection behavior by zooplankton in the presence and absence of both the invertebrate predator Chaoborus and UV. We hypothesized that Chaoborus and UV both affect the distribution of zooplankton and a combination of both factors would lead to a narrowing of depth distribution. We found that when Chaoborus were present zooplankton tended to be distributed at shallower depths in the columns, while in the presence of UV they exhibited a deeper distribution. Chaoborus themselves were always found near the bottom of the columns regardless of the UV treatment. Simultaneous exposure to predators and UV resulted in a peak of zooplankton (especially Daphnia catawba) distribution at intermediate depths. In a significant number of cases, depth range was narrowed in response to Chaoborus, UV, or both.     

Development of survival skills in captive-raised Siberian polecats (Mustela eversmanni) II: Predator avoidance

We exposed naive Siberain polecats (Mustela eversmanni) (aged 2, 3, and 4 months) to a swooping stuffed great horned owl (Buho virginianus) and a stuffed badger (Taxidae taxus) mounted on a remote control toy automobile frame. The first introduction to each was harmless, the second was accompanied by a mild aversive stimulus, the third (1 day after attack) was harmless, and the fourth (30 days after attack) was harmless. Alert behavior increased after a single attack by either predator model. Escape responses of naive polecats did not differ between ages when exposed to the badger, but 4 month old polecats reduced their escape times after a single badger attack. When exposed to the swooping owl, naive 4 month old polecats redponded more quickly than the other two age groups, and 3 and 4 month old polecats reduced escape times after a single owl attack. This indicates an innate escape response to the owl model at 4 months of age, and a short-tert ability to remember a single mild aversive encounter with the badger and owl models at 3 or 4 months of age.     

Marine fouling on floating installations west of Dongsha Islands,the northern South China Sea

Marine fouling on floating installations west of Dongsha Islands, the northern South China Sea, was investigated from August 1987 to October 1990. Two buoy stations were deployed at approximately 110 km and 115 km west of Dongshadao, Dongsha Islands, anchored in water at 325 and 345 m depths, and exposed for 8 and 12 months. The experimental panels were fixed on iron structure frames which were connected to the buoy and its mooring system immersed in water at different depths, and retrieved after 4, 8 and 12 months of exposure. Fouling organisms colonizing the buoys and their mooring systems were sampled after 8 and 12 months of deployment. A total of 86 species was collected and identified at these two stations and the fouling community showed a typical oceanic characteristics. Most of the species were mainly found in the top 50 m depth and a striking vertical zonation of species was also observed with depth. The distance from offshore is a major factor in determining the fouling community composition. In terms of biomass, the fouling community was dominated by pedunculate barnacles, hydroids, and algae, followed by common oysters, pearl oysters and acorn barnacles. Biomass of hard fouling organisms increased over time. Under the influence of hydrological currents propagules of communities around the Dongsha Islands may also affect the development of the fouling communities in the offshore waters.     

ZONATION OF DEEP WATER BENTHIC ALGAE IN THE GULF OF MAINE1

Algal community structure is described for a deep-water rock pinnacle in the Gulf of Maine. Three depth zones of algal dominance were apparent consisting of 1) leathery macrophytes (to 40 m), 2) foliose red algae (to 50 m) and 3) crustose algae (fleshy crusts to 55 m and coralline crusts to 63 m). Microscopic filamentous and erect calcareous algae were also present but inconspicuous. Upright macroscopic filamentous and thin sheet-like forms were not observed on the pinnacle. Sea anemones (Metridium senile) dominated some vertical faces and abrupt prominences in the shallowest regions of the pinnacle (to 24 m) and locally appeared to set the upper vertical limits of kelp and possibly foliose reds. Laminaria sp. formed an open park-like canopy from 24 to 30 m whereas Agarum cribrosum, the deepest kelp, grew as isolated individuals to 40 m. Peyssonnelia sp. and Leptophytum laeve were the deepest occurring fleshy (to 55 m) and calcareous crusts (to 63 m), respectively. The occurrence of these algae at record depths for the Gulf of Maine and for cold water marine environments may be the result of an absence of large herbivores and the high productivity potential of the benthos in these relatively clear waters. By compiling data on depth distribution patterns world-wide, it is evident that the three zone structure of algal morphologies observed in the Gulf of Maine is a global phenomenon.     

Variation in Petroleum Hydrocarbon Chain Lengths with Depth at a Former Crude Oil and Natural Gas ProductionFacility

An investigation was performed at a former crude oil and natural gas production facility to evaluate whether releases from the product flowlines, gathering lines or water injection lines had impacted soil beneath the site. Thirty-six trenches were initially excavated and sampled beneath the former piping runs to a maximum depth of 6?m. After the trenching investigation, nine soil boreholes were advanced and sampled to a depth of approximately 18?m to further delineate the lateral and vertical extent of impacted soil. Soil samples collected from the trenches and boreholes were analyzed for total petroleum hydrocarbons (TPH) in accordance with ASTM Method 2887. The results of the investigation indicated that TPH impacted soil was present within several areas of the 40-ha site. The petroleum hydrocarbons generally had chain lengths ranging from C₆ to C₃₅, characteristic of light crude oil. The impacted soil also contained condensate, the volatile portion of crude oil. Condensate consists of short-chain hydrocarbons (C₁ to C₁₂) and is characterized by low levels of aromatic volatile organic compounds (VOCs). The condensate typically was more prevalent at depths below 4.5?m than the less volatile, longer chain length hydrocarbons. Statistical analysis of TPH data collected during subsequent excavation activities showed that the mean percentage of condensate was significantly greater at depths below 4.5?m than in shallower samples. In contrast, the mean percentage of TPH compounds in the diesel range (C₁₄ to C₂₃) was significantly greater in samples collected at depths above 4.5?m. The difference in the mean percentage of heavier hydrocarbons (C₂₄ to C₄₄+) with depth was not statistically significant.     

Foraging costs and accessibility as determinants of giving-up densities in a swan-pondweed system

We measured the patch use behaviour of Bewick's swans (Cygnus columbianus bewickii) feeding on below ground tubers of fennel pondweed (Potamogeton pectinatus). We compared the swans’ attack rates, foraging costs and giving‐up densities (GUDs) in natural and experimental food patches that differed in water depth. Unlike most studies that attribute habitat‐specific differences in GUDs to predation risk, food quality or foraging substrate, we quantified the relative importance of energetic costs and accessibility. Accessibility is defined as the extent to which the animal's morphology restricts its harvest of all food items within a food patch. Patch use behaviours were measured at shallow (ca 0.4 m) and deep (ca 0.6 m) water depths on sandy sediments. In a laboratory foraging experiment, when harvesting food patches, the swan's attack rate (m³ s^?1) did not differ between depths. In deep water the energetic costs of surfacing, feeding and trampling were 1.13 to 1.21 times higher than in shallow water with a tendency to spend relatively more time trampling, the most expensive activity. Taking time allocation as measured in the field into account, foraging in deep water was 1.26 times as expensive as in shallow water. In the lake the GUD in shallow water was on average 12.9 g m^?2. If differences in energetic costs were the only factor determining differences in GUDs, then the deep water GUD should be 14.2 g m^?2. Instead, the mean GUD in deep water was 20.2 g m^?2, and therefore energetic costs explain just 18% of the difference in GUDs. At deep sites, 24% of tuber biomass was estimated to be out of reach, and we calculated a maximum accessible foraging depth of 0.86 m. This is close to the published 0.84 m based on body measurements. A laboratory experiment with food offered at a depth of 0.89 m confirmed that it was just out of reach. The agreement between calculated and observed maximum accessible foraging depths suggests that accessibility largely explains the remaining difference in GUDs with depth, and it confirms the existence of partial prey refuges in this system.     

Vertical distributions and relations of euphausiid populations off Elephant Island,March 1984

Summary Distributional relationships are described for post-larval and larval Euphausia superba and Thysanoessa sp. (probably macrura) and post-larval Euphausia frigida collected in 0–70/80 m and 0–175/200 m depth ranges with a MOCNESS sampler north of Elephant Island (61°S, 55°W) during 17–23 March 1984. Larval E. superba (predominantly calyptopes stage 2 and 3) were rare shallower than 80 m at night. Day catches of post-larval E. suberba were small and night catches were primarily near the top of the thermocline above 50 m depth. Thysanoessa sp. occurred throughout the 0–200 m depth range and was abundant in the upper 80 m both night and day. E. frigida migrated to the upper 80 m at night from deeper day depths. Larval stages of E. superba and bost-larval stages of all three species demonstrated independent and variable vertical distribution patterns both night and day. Changes in E. superba abundance and distributional patterns could to a certain extent be associated with observed environmental changes. An increase in larval and decrease in post-larval E. superba abundances between 0–80 m was associated with an intrusion of cold water at depth. At night, vertically restricted concentrations of post-larval E. superba were associated with shallow mixed layer depths, and a significant vertical separation of developmental stages and size categories was observed only during periods of stratification in the upper 80 m. Fluctuations in the distribution and abundance of Thysanoessa sp. and distribution of E. frigida did not appear to be influenced by physical parameters within the upper 80 m. Within the 0–80 m depth range, the distributions of these two species differed from each other and from E. superba and showed large tow to tow variability that could not be related to physical parameters in the upper water column.     

The development and vertical distribution of populations of gas-vacuolate bacteria in a eutrophic,monomictic lake

The role of gas vacuoles in the vertical stratification of planktonic bacteria is analysed. Measurements made with certain gas-vacuolate bacteria in laboratory culture suggest that only colonial forms could sink or float fast enough to form population maxima in lakes by vertical migration from other depths. It is suggested that in the case of individual cells the importance of the buoyancy provided by gas vacuoles is to minimise sinking rates and thereby to increase residence times of the organisms at depths where conditions support their growth.Changes in the vertical distribution of a number of gas-vacuolate bacteria were followed throughout the year in a monomictic, eutrophic lake (Crose Mere, Shropshire). All were restricted to the anaerobic hypolimnion which developed in summer. The various species formed maxima at different depths and times. With some of them (e.g. species of Thiopedia, Pelonema and Brachyarcus) growth was necessary to explain their development. In others (e. g. species of Pelodictyon and two colourless bacteria) vertical migrations might also have contributed to their development.