Temperature effects on vital rates of different life stages and implications for population growth of Baltic sprat

Baltic sprat (Sprattus sprattus balticus S.) is a key species in the pelagic ecosystem of the Baltic Sea. Most stocks of small pelagic species are characterized by natural, fishery-independent fluctuations, which make it difficult to predict stock development. Baltic sprat recruitment is highly variable, which can partly be related to climate-driven variability in hydrographic conditions. Results from experimental studies and field observations demonstrate that a number of important life history traits of sprat are affected by temperature, especially the survival and growth of early life stages. Projected climate-driven warming may impact important processes affecting various life stages of sprat, from survival and development during the egg and larval phases to the reproductive output of adults. This study presents a stage-based matrix model approach to simulate sprat population dynamics in relation to different climate change scenarios. Data obtained from experimental studies and field observations were used to estimate and incorporate stage-specific growth and survival rates into the model. Model-based estimates of population growth rate were affected most by changes in the transition probability of the feeding larval stage at all temperatures (+0, +2, +4, +6?°C). The maximum increase in population growth rate was expected when ambient temperature was elevated by 4?°C. Coupling our stage-based model and more complex, biophysical individual-based models may reveal the processes driving these expected climate-driven changes in Baltic Sea sprat population dynamics.     

Effect of temperature on viability and axial muscle development in embryos and yolk sac larvae of the Northeast Arctic cod (Gadus morhua)

Cod (Gadus morhua L.) eggs may develop and hatch within temperatures of −1.5 to 12 °C, but little is known about the effects of very low temperatures on larval characteristics. Eggs of the Northeast Arctic cod (Gadus morhua) were incubated at 1, 5 or 8 °C from Day 1 after fertilisation until hatching, and transferred to 5 °C after hatching. Histological samples of the axial musculature were taken at hatching and 5 d after hatching, and the data on muscle cellularity from these samples were related to survival and hatching, size, developmental data and viability of the yolk sac larvae. All larvae hatched at the same developmental stage. Incubation of eggs at 1 °C produced shorter larvae with a larger yolk sac and more, small deep fibres at hatching than larvae from eggs incubated at 5 or 8 °C. The larval size difference was still present 5 d after hatching, a time at which the larvae from 1 °C-incubated eggs were less developed and less resistant to an acute viability stress test (65 ppt salinity). Although there were no differences between temperature groups in number and size of muscle fibres 5 d after hatching, the deep fibres of the 1 °C-group contained less myofibrils than the two other groups. The phenotype of the larvae at hatching was thus affected within these incubation temperatures. Although all groups were transferred to the same temperature after hatching, the lowest egg incubation temperature (1 °C) still had a negative effect 5 d after hatching, as these larvae were both smaller, less resistant to stress and had less functional muscles at the time of first feeding. Our conclusion is therefore that 1 °C is close to, or below, the lower thermal tolerance limit for normal functional development of Northeast Arctic cod. The results are discussed in relation to larval viability and recruitment of this species in the wild. Received: 4 February 1998 / Accepted: 10 July 1998     

Potential impact of climate warming on the recruitment of an economically and ecologically important species, the European lobster (Homarus gammarus) at Helgoland, North Sea

A laboratory-based study was performed to assess the impact of climate warming on the recruitment of the endangered population of the European lobster (Homarus gammarus) at Helgoland (North Sea, German Bight). Egg-bearing females collected in situ just after spawning in late summer were subjected to various seasonal temperature regimes. Regimes with elevated temperatures (mild winters) resulted in a strong seasonal forward shift of larval hatching. Hatching took place at significantly lower temperatures than under regimes with normal winters. Experiments on larval development across a range of constant temperatures showed that no successful larval development occurred at temperatures below 14°C. Larval survival increased from 9% at 14°C to 80% at 22°C, while duration of larval development decreased correspondingly from 26 to 13 days. We hypothesize that an ongoing warming of the North Sea will strongly affect the recruitment success of the Helgoland lobster, mainly resulting from a decoupling of the seasonal peak appearance of larvae from optimal external conditions (temperature, food availability) for larval development.     

Egg and early larval stages of Baltic cod, Gadus morhua, are robust to high levels of ocean acidification

The accumulation of carbon dioxide in the atmosphere will lower the pH in ocean waters, a process termed ocean acidification (OA). Despite its potentially detrimental effects on calcifying organisms, experimental studies on the possible impacts on fish remain scarce. While adults will most likely remain relatively unaffected by changes in seawater pH, early life-history stages are potentially more sensitive, due to the lack of gills with specialized ion-regulatory mechanisms. We tested the effects of OA on growth and development of embryos and larvae of eastern Baltic cod, the commercially most important fish stock in the Baltic Sea. Cod were reared from newly fertilized eggs to early non-feeding larvae in 5 different experiments looking at a range of response variables to OA, as well as the combined effect of CO₂ and temperature. No effect on hatching, survival, development, and otolith size was found at any stage in the development of Baltic cod. Field data show that in the Bornholm Basin, the main spawning site of eastern Baltic cod, in situ levels of pCO₂ are already at levels of 1,100 μatm with a pH of 7.2, mainly due to high eutrophication supporting microbial activity and permanent stratification with little water exchange. Our data show that the eggs and early larval stages of Baltic cod seem to be robust to even high levels of OA (3,200 μatm), indicating an adaptational response to CO₂.     

Effects of temperature and delayed feeding on growth and survival of larvae of three species of subtropical marine fishes

In larvae of the bay anchovy Anchoa mitchilli (Valenciennes), the sea bream Archosargus rhomboidalis (Linnaeus), and the lined sole Achirus lineatus (Linnaeus), growth, survival, and starvation times were investigated at temperatures of 22° to 32°C. The rate at which hours after hatching until starvation decreased in relation to temperature for unfed larvae did not differ significantly among the 3 species, ranging from-5.4 to-6.3 h per degree increase in temperature. The total number of hours until starvation did differ for all 3 species: lined soles survived longest, bay anchovies were intermediate, and sea bream survived the least time. At 28°C, unfed sea bream could survive 90.1 h, bay anchovy 102.3 h, and lined sole 119.8 h. The eyes pigmented at nearly the same time after hatching for sea bream and bay anchovy, but took about 20 h longer at all temperatures for lined sole. Quadratic equations best described the relationship between hours after hatching when the eyes became pigmented and temperature. Eye-pigmentation times became nearly constant for all 3 species at temperatures above 28°C. At 28°C, eyes pigmented about 27 h after hatching for bay anchovy and sea bream but not until 47 h for lined sole. Hours after eye pigmentation when unfed larvae starved was a measure of the effective time that larvae had to commence feeding. Bay anchovies and lined soles were nearly alike in this respect, but sea bream starved at tewer hours after eye pigmentation. Slopes of regressions representing decrease in times to staration for increasing temperatures ranged from-3.7 to-4.4 h per degree increase in temperature, and were not significantly different among the 3 species. At 28°C, unfed lined soles starved at 70 a after eye pigmentation, bay anchovies starved at 72.5 h, and sea bream at only 62 h. Yolk absorption was most rapid for all species during the first 20 h after hatching, and was faster at higher temperatures. Amounts of yolk remaining at the time eyes became pigmented were less at higher temperatures for bay anchovy and lined sole, but were greater for sea bream, suggesting that sea bream used yolk more efficiently at higher temperatures. Either no yolk or small traces (>0.20%) remained at 24 h after eye pigmentation in all 3 species. Feeding was delayed for periods of 8, 16, 24, 32, 40 and 48 h after eye pigmentation for all species at a series of experimental temperatures from 24° to 32°C. Growth and survival were affected when food was withheld for more than 24 h at 28°C, but survival did not decrease markedly until food was withheld at least 8 h longer. At lower temperatures food could be withheld longer and at higher temperatures for less time. Feeding can be initiated by most larvae for several hours after all visible yolk reserves have been exhausted. All species tested can survive for 24 to 40 h after eye pigmentation at 24° to 28°C without food and still have relatively good growth and survival when food is offered. If the “critical period” is considered relative to time of hatching, lined soles need not find food for 3 to 3.5 days after hatching, but bay anchovy and sea bream must feed within 2.5 days of hatching.     

The effect of temperature on the development and hatching of resting eggs of non-indigenous predatory cladoceran Cercopagis pengoi in the Gulf of Finland, Baltic Sea

The effect of temperature on the development and hatching of resting eggs of the Ponto-Caspian Cercopagis pengoi was studied experimentally in the Gulf of Finland, Baltic Sea. Morphological changes were monitored as the development of the resting eggs proceeded. Sexual reproduction of the C. pengoi population in the Gulf of Finland was evaluated by combining the data from hatching experiments and resting egg abundances in the sediment. Development time of resting eggs was dependent on temperature: increase in the temperature shortened the time needed until hatching. Hatching success was also dependent on incubation temperature. Almost sixfold increase in hatching success was detected when temperature increased two degrees above the storage temperature. Average resting egg abundances varied between 0.16 and 0.49 eggs cm⁻³ in the 0–6 cm sediment layer.     

Biological effects of oil on early development of the Baltic herring Clupea harengus membras

The effects of petroleum hydrocarbons from two crude oils and one fuel oil (No. 1) were studied on the ontogenic development of the Baltic herring Clupea harengus membras L. Herring eggs exposed to water-soluble fractions of the oils at the time of fertilization showed no decrease in numbers of fertilized eggs compared to eggs exposed 6 or 72 h after fertilization. During embryongenesis, treatment with 3.1 to 8.9 ppm or 3.3 to 11.9 ppm total oil hydrocarbons from light fuel oil and the two crude oils respectively, gave rise to alterations in embryonic activity, decreased heart rate, and premature or delayed hatching. Although many larvae hatched from eggs exposed to contaminated water (3.1 to 11.9 ppm total oil hydrocarbons), the majority of the (70 to 100%) were malformed or dead 1 day after hatching. Exposure of eggs to 5.4–5.8 ppm total oil hydrocarbons resulted in significantly (P<0.001) decreased lengths of the larvae. Increased temperature (from 9° to 14°C) aggravated the effects of the oils. The results are discussed in relation to the potential effects of oil spills and chronic oil pollution on fish eggs and larvae in the Baltic Sea.     

The effects of temperature and salinity on egg production and hatching success of Baltic Acartia tonsa (Copepoda: Calanoida): a laboratory investigation

The functional response of the aspects of reproductive success of a southwestern Baltic population of Acartia tonsa (Copepoda: Calanoida) was quantified in the laboratory using wide ranges in temperatures and salinities. Specifically, daily egg production (EP, # female⁻¹ day⁻¹) was determined for 4 or 5 days at 18 different temperatures between 5 and 34°C and the time course and success of hatching were evaluated at 10 different temperatures between 5 and 23°C. The effect of salinity (0 to 34 psu) on egg hatching success was also examined. The highest mean rates of EP were observed between 22 and 23°C (46.8–50.9 eggs female⁻¹ day⁻¹). When studied at 18 psu, hatching success of eggs increased with increasing temperature and was highest (92.2%) at 23°C. No hatching was observed for eggs incubated at low temperatures (≤12°C) that were produced by females acclimated to temperatures ≤10°C indicating a possible thermal threshold between 10.0 and 13.0°C below which only the production of diapause (or low quality) eggs exists in this population. When tested at 18°C, the hatching success of eggs incubated at 15 different salinities increased asymptotically with increasing salinity and was maximal (81.4–84.5%) between 17 and 25 psu. The high reproductive success observed over wide ranges in temperatures and salinities in this Baltic population demonstrates one of the mechanisms responsible for the cosmopolitan distribution of this species within productive, estuarine and marine habitats.     

Description of the early development of the halibut Hippoglossus hippoglossus and attempts to rear the larvae past first feeding

Rearing experiments on the halibut Hippoglossus hippoglossus (L.) were carried out using gametes from parents caught at a depth of 600 to 700 m off the Norwegian coast in February 1980. After fertilization, the average egg diameter was 3.08 mm, average dry weight 1 038 μg and neutral buoyancy was at 36.5‰ S. The eggs hatched after 20 d at 4.7°C, 18 d at 5°C and 13 d at 7°C. Survival to hatching was better when antibiotics were used. At hatching the larvae were 6.4 mm long, there were no functional eyes or mouth, but prominent neuromast organs were present. Resorption of yolk lasted 50 d at 5.3°C; the eyes and mouth were then functioning and the larva was about 11.5 mm long. The larvae were offered zooplankton as food, but with little success in initial feeding. A few larvae fed and grew in 2 500-litre plastic bags, one reaching a length of 24 mm after 90 d.     

Temporal and spatial variation in the growth rates of Baltic herring (<Emphasis Type="Italic">Clupea harengus membras</Emphasis> L.) larvae during summer

It has been suggested that larval survival determines the year-class strength in most marine fish species. During their growth and development, the ability of the larvae to catch prey and avoid predation will increase. However, the factors affecting short-term changes in the growth of Baltic Sea herring have been little studied in the field. We collected Baltic herring (Clupea harengus membras L.) larvae from five different towing areas in the Archipelago Sea (SW Finland) during May and June 1989, right after the main spawning season. Twenty thousand two hundred and ten larvae were analysed and the area-specific growth rate (i.e. increase in standard length) was estimated by tracing the larval cohorts from the length-frequency data. This represents the first Baltic herring study with daily sampling during a long study period. The growth rate was related to environmental factors, such as temperature, number of zooplankters, and wind speed and direction. Large variation in larval growth rate occurred between areas: lowest and highest growth rates were 0.18 and 0.52 mm·day^-1. Temperature was an important variable controlling larval-fish growth rate. An increase of one 1°C in average water temperature corresponded to an increase in growth rate of 0.043 mm·day^-1. This may have been caused either by a direct temperature effect (changes in metabolic rate) or by the indirect effect of changes in food availability. We also found the densest herring populations in the areas with highest average water temperature. However, temperature and larval growth rate both increased towards the inner archipelago.     

Eggs and early larvae of Thrissocles species (Engraulidae: Pisces)

Eggs of Thrissocles species are found in surface plankton in the Ernakulam Channel (Cochin Harbour)during February to May 1967. The eggs hatch within 24 h. Empty egg shells have characteristic apertures, through which the embryos have emerged; yolk is resorbed 36 h after hatching.Larvae (36 to 72 h groups) assemble at the lighted region of the aquaria during day-time and scatter to different levels at night. Larvae older than 72 h show no inclination to assemble as before. All larvae died between 96 and 110 h after hatching. Many batches of eggs were reared in the same medium, and all of them behaved as described. The results indicate that the right type of food was available in the aquaria for larvae up to a period of 72 h. The volume of water also appears to have a bearing upon the survival rate and longevity of the larvae since, in small aquaria, more larvae died at an early stage.     

Combined effects of dissolved oxygen concentration and water temperature on embryonic development and larval shell secretion in the marine snail <Emphasis Type="Italic">Chorus</Emphasis><Emphasis Type="Italic">giganteus</Emphasis> (Gastropoda: Muricidae)

The present study was undertaken to determine the effects of both extracapsular oxygen concentration and temperature on embryonic development in Chorus giganteus. In normoxia increasing water temperature from 12°C to 18°C reduced by 15 days the median time required for the capsules to hatch. Hypoxia (oxygen content at 50% of air saturation) generated a low development rate and totally prevented both shell secretion and larval hatching from the egg capsule. Experimental transfer at weekly intervals, from normoxia to hypoxia and vice versa, induced a decrease and increase in the embryonic ash content, respectively, but did not affect the number of hatched larvae. Such an effect was more pronounced at 12°C than at 15°C or 18°C. The embryonic inability to produce a shell under hypoxia is likely to be a result of the low intracapsular oxygen concentration (IPO₂) generated as the combined effect of a low extracapsular oxygen concentration (environmental) added to the intracapsular embryonic oxygen demands, which lowers the IPO₂ still further. Under such conditions, a decrease in intracapsular pH is likely to take place, and, if so, embryos might divert carbonates away from shell calcification to balance such changes in pH.     

Egg survival,embryonic development,and larval characteristics of northern shrimp (<Emphasis Type="Italic">Pandalus borealis</Emphasis>) females subject to different temperature and feeding conditions

Laboratory experiments on ovigerous females of northern shrimp (Pandalus borealis) were used to assess the effects of temperature and food ration on female condition during incubation and examine how combined effects of temperature and female condition influenced egg survival, embryonic development, and larval characteristics. Ovigerous females were maintained at 2°C, 5°C, and 8°C and fed on a low (three times/week; 2–2.7% W/W) or high ration (five times/week at satiation). The increase in temperature accelerated the developmental time of the eggs but their survival at 8°C was reduced. Conversion efficiency of yolk reserves in developing embryos was significantly reduced at elevated temperatures and larvae hatching at 2°C and 5°C were significantly larger and heavier than those hatching at 8°C. The experimental design did not result in any effect of food ration on the energetic condition of females or on egg characteristics and their biochemical composition. However, lower energy reserves were observed for females held at 8°C.     

Elevated temperature elicits greater effects than decreased pH on the development, feeding and metabolism of northern shrimp (Pandalus borealis) larvae

Climate models predict that the average temperature in the North Sea could increase 3–5 °C and surface-waters pH could decrease 0.3–0.5 pH units by the end of this century. Consequently, we investigated the combined effect of decreased pH (control pH 8.1; decreased pH 7.6) and temperature (control 6.7 °C; elevated 9.5 °C) on the hatching timing and success, and the zoeal development, survival, feeding, respiration and growth (up to stage IV zoea) of the northern shrimp, Pandalus borealis. At elevated temperature, embryos hatched 3 days earlier, but experienced 2–4 % reduced survival. Larvae developed 9 days faster until stage IV zoea under elevated temperature and exhibited an increase in metabolic rates (ca 20 %) and an increase in feeding rates (ca 15–20 %). Decreased pH increased the development time, but only at the low temperature. We conclude that warming will likely exert a greater effect on shrimp larval development than ocean acidification manifesting itself as accelerated developmental rates with greater maintenance costs and decreased recruitment in terms of number and size.     

Effects of temperature on oxygen consumption,growth, and development of embryos and yolk-sac larvae of Siganus randalli (Pisces: Siganidae)

Eggs from laboratory spawnings of the coralreef fish Siganus randalli Woodland were incubated at two temperatures (27 and 30 °C). Eggs and larvae were sampled until larval starvation, while changes in oxygen consumption, growth, yolk utilization, and development were monitored. Oxygen consumption, which peaked at hatching, was higher for embryos incubated at 30 °C than at 27 °C. Rates of oxygen consumption (nl h^-1 individual^-1) at hatching were similar to those for other temperate and tropical species. Rates of oxygen consumption by yolk-sac larvae were highly variable, and these data suggest that larval oxygen consumption prior to yolk-sac absorption may not be significantly influenced by temperature. Rates of yolk depletion were higher for larvae at the higher temperature. After an initial rapid increase in length, length of larvae at 30 °C decreased with age. Egg size, egg weight, and maximum notochord length of larvae differed significantly between spawns. Age-specific oxygen consumption rates by the embryos varied between spawns, but regressions describing oxygen consumption as a function of age did not differ significantly. The initiation and completion of eye pigmentation were used as developmental markers to calculate the amount of yolk remaining for larvae at the different temperatures. Larvae maintained at 30 °C completed eye pigmentation approximately 3 h sooner than those maintained at 27 °C, but had less endogenous reserves. This finding indicates a trade-off between rapid development and efficient utilization of the endogenous reserves. The completion of eye pigmentation in larvae incubated at the higher temperature occurred at midnight and, depending on the amount of time that the larvae have to initiate feeding prior to the point-of-no-return, the timing of completion of eye pigmentation could influence larval survival.     

The effects of temperature and salinity on reproductive success of <Emphasis Type="Italic">Temora longicornis</Emphasis> in the Baltic Sea: a copepod coping with a tough situation

At specific locations within the Baltic Sea, thermoclines and haloclines can create rapid spatial and temporal changes in temperature (T) and salinity (S) exceeding 10°C and 9 psu with seasonal ranges in temperature exceeding 20°C. These wide ranges in abiotic factors affect the distribution and abundance of Baltic Sea copepods via species-specific, physiological-based impacts on vital rates. In this laboratory study, we characterized the influence of T and S on aspects of reproductive success and naupliar survival of a southwestern Baltic population of Temora longicornis (Copepoda: Calanoida). First, using ad libitum feeding conditions, we measured egg production (EP, no. of eggs female⁻¹ day⁻¹) at 12 different temperatures between 2.5 and 24°C, observing the highest mean EP at 16.9°C (12 eggs female⁻¹ day⁻¹). Next, the effect of S on EP and hatching success (HS, %) was quantified at 12°C for cohorts that had been acclimated to either 8, 14, 20 or 26 psu and tested at each of five salinities (8, 14, 20, 26 and 32 psu). The mean EP was highest for (and maximum EP similar among) 14, 20 and 26 psu cohorts when tested at their acclimation salinity whereas EP was lower at other salinities. For adults reared at 8 psu, a commonly encountered salinity in Baltic surface waters, EP was relatively low at all test salinities—a pattern indicative of osmotic stress. When incubated at 12°C and 15 different salinities between 0 and 34 psu, HS increased asymptotically with increasing S and was maximal (82.6–84.3%) between 24 and 26 psu. However, HS did depend upon the adult acclimation salinity. Finally, the 48-h survival of nauplii hatched and reared at 14 psu at one of six different temperatures (10, 12, 14, 16, 18 and 20°C) was measured after exposure to a novel salinity (either 7 or 20 psu). Upon exposure to 7 psu, 48-h naupliar mortality increased with increasing temperature, ranging from 26.7% at 10°C to 63.2% at 20°C. In contrast, after exposure to 20 psu, mortality was relatively low at all temperatures (1.7% at 10°C and ≤26.7% for all other temperatures). An intra-specific comparison of EP for three different T. longicornis populations revealed markedly different temperature optima and clearly demonstrated the negative impact of brackish (Baltic) salinities. Our results provide estimates of reproductive success and early survival of T. longicornis to the wide ranges of temperatures and salinities that will aid ongoing biophysical modeling examining climate impacts on this species within the Baltic Sea.     

Yolk resorption,onset of feeding and survival potential of larvae of three tropical marine fish species reared in the hatchery

This paper provides basic early life-history information on milkfish (Chanos chanos), seabass (Lates calcarifer) and rabbitfish (Siganus guttatus) which may explain in part the observed differences in their survival performance in the hatchery. Egg size, larval size, amount of yolk and oil reserves and mouth size are all greater in milkfish than in seabass, and greater in the latter than in rabbitfish. During the first 24 h after hatching, rabbitfish larvae grow much faster than milkfish and seabass larvae at similar ambient temperatures (range 26°–30°C, mean about 28°C). The eyes become fully pigmented and the mouths open earlier in seabass and rabbitfish (32–36 h from hatching) than in milkfish (54 h). Seabass larvae learn to feed the earliest. Yolk is completely resorbed at 120 h from hatching in milkfish, and yolk plus oil at 120 h in seabass and 72 h in rabbitfish at 26° to 30°C. Milkfish and seabass larvae have more time than rabbitfish to initiate external feeding before the endogenous reserves are completely resorbed. Delayed feeding experiments showed that 50% of unfed milkfish larvae die at 78 h and all die at 150 h from hatching. Milkfish larvae fed within 54 to 78 h after hatching had improved survival times: 50% mortality occurred at 96 to 120 h, and 10 to 13% survived beyond 150 h. Unfed seabass larvae all died at 144 h, while 6 to 13% of those fed within 32 to 56 h after hatching survived beyond 144 h and well into the subsequent weeks. Unfed rabbitfish larvae all died at 88 h, while 7 to 12% of those fed within 32 to 56 h after hatching survived beyond 88 h. A delay in initial feeding of more than 24 h after eye pigmentation and opening of the mouth may be fatal for all three species.Contribution No. 167 from the SEAFDEC Aquaculture Department     

Fertilization and early developmental stages of eggs of Clupea harengus pallasi n. maris-albi under fresh-water conditions

Eggs of the White Sea herring Clupea harengus pallasi Val. are laid in the intertidal zone, where they are liable to come under the influence of fresh water. The present paper deals with the effect of fresh-water influence on the egg of the White Sea herring, under experimental conditions. In fresh water, egg and sperm are in an unactivated state and, after transfer into sea water, are capable of re-activation, even after 212/ h exposure. The treated eggs develop until the hatching of normal and viable larvae. The number of eggs which are able to complete development depends upon the period of freshwater exposure, although the correlation between the length of that period and survival rates is not very close. Apparently, this is due both to the ability of C. harengus pallasi eggs to develop parthenogenetically and different qualities of eggs (which were obtained from different females). The morphological features of White Sea herring eggs, induced by experimental fresh-water influence, are very similar to those typical for herring eggs of the genus Ilisha, obtained from natural brackish waters. However, such changes are not characteristic of the eggs of forms from natural fresh and brackish-waters.     

Reproductive isolation between intertidal and subtidal<Emphasis Type="Italic"> Scoloplos armiger</Emphasis> (Polychaeta,Orbiniidae) indicates sibling species in the North Sea

Two distinct modes of development in the common polychaete Scoloplos armiger (O. F. Müller, 1776) occur in the North Sea region: holobenthic development in egg cocoons and pelagic larvae hatching from suspended eggs. In the northern Wadden Sea near the island of Sylt, we observed that egg cocoons are produced intertidally while pelagic larvae originate from the adjacent subtidal zone. A previous genetic comparison between these subtidal and intertidal populations revealed distinct gene pools, suggesting that reproductive differences are not phenotypic but heritable. In this study, crossbreeding experiments show that intertidal and subtidal populations are reproductively isolated. Couples with males and females from different habitats had no offspring. Production of egg cocoons is determined by female origin from the intertidal zone. Pelagic larvae occurred only in couples with subtidal females and subtidal males. Intertidal males have spermatozoa with heads twice as long as those from subtidal males and a significantly shorter flagellum. We suspect that deviating sperm morphology may cause the reproductive breakdown at the fertilization stage. Juveniles hatching from cocoons have shorter anal cirri compared to juveniles that metamorphosed from pelagic larvae. We conclude there to be two sympatric sibling species in S. armiger: 'type I' in intertidal areas, which have egg cocoons, no pelagic larvae, elongated sperm heads, shortened sperm flagella and anal cirri; and a subtidal 'type S', lacking egg cocoons but with pelagic larvae, short sperm heads, long sperm flagella and anal cirri.Communicated by O. Kinne, Oldendorf/Luhe