Role of chlorophyll and carbohydrate contents in survival of Posidonia oceanica cuttings transplanted to different depths

Cuttings of the marine phanerogam Posidonia oceanica (L.) Delile were collected and transplanted in January 1991 at four stations of different depths between 5 and 20 m in the underwater reserve at Monaco (1 ha in area). From January to July, samples of the transplants were collected to study the effect of depth on survival and on chlorophyll and carbohydrate concentrations. In the leaves, chlorophyll concentrations were significantly lower (1.00 mg g^-1 dry wt) in cuttings taken from 5 m depth than in cuttings taken from 20 m (1.99 mg g^-1). Transplantation of cuttings from 5 to 20 m or from 20 to 8 m did not significantly change the chlorophyll concentration in leaves for a period of 6 mo after transplantation. Cutting and transplantation result in stress, as indicated by a decrease in pigment concentration without change of depth. However, the carbohydrate concentration in rhizomes from 5 m (32 mg g^-1 dry wt), is significantly higher than in material from 20 m (17 mg g^-1). Cuttings transplanted from 5 to 20 m lost most of their carbohydrate reserves (reaching a minimum of 2.5 mg g^-1 in May); cuttings transplanted from 20 to 8 m maintained higher reserves (with a minimum of 20 mg g^-1 in June). This may explain why material transplanted from 5 to 20 m exhibited only 52% survival, while material transplanted from 20 to 8 m displayed 80 to 92% survival. Material taken from populations adapted to low light at 20 m is probably more resistant to transplantation stress, particularly when replanted at a higher light intensity in shallow water.     

Patterns of seagrass (Posidonia oceanica) flowering in the Western Mediterranean

The intensive reconstructive sampling (1957–2004, 39 localities), a systematic direct observation (1992–2004, 1 locality) and particular direct observations (66 localities) of Posidonia oceanica meadows were analysed together with temporal series of flowering available in the literature (19 localities). This allowed the examination of temporal and spatial variability in annual flowering prevalence (FP, the fraction of meadows flowering in a given year) and of annual meadow flowering intensity (FI, number of inflorescences per shoot) for the period 1979–2004 across the Western Mediterranean, as well as spatial variability of flowering frequency (FF, the fraction of years that a given meadow has flowered) and shoot flowering probability (Pf, fraction of flowering stalks appeared per annual segment). Each year, on an average 17% of the investigated meadows flowered, ranging from 3 to 86% of meadows among the years. The highest annual FP and FI values were obtained in 2003 (FP=0.86 and mean FI=0.23±0.03 inflorescences shoot⁻¹). A secondary peak of FP and mean FI occurred 9 years earlier, in 1994 (FP=0.44 and mean FI=0.08±0.02). Both peaks of flowering occurred after hot summers. Flowering synchrony in particular years across the Western Mediterranean and clines of increased meadow flowering frequency towards the North and East, suggests the existence of large-scale environmental mechanisms controlling the floral induction. On the other hand, meadow FF and Pf were highly heterogeneous among and within the meadows, indicating that local factors also may play a significant role in flowering induction. When flowering, the Western Mediterranean meadows showed an average 0.11±0.02 inflorescences shoot⁻¹, but FI greatly varied among and along the series (from 0.002 to 0.54 inflorescences shoot⁻¹) and decreased significantly with depth but was independent of meadow shoot density and meadow latitude or longitude. The shoot flowering probability was quite low (0.007±0.002 inflorescences shoot⁻¹ year⁻¹) and exponentially increased with shoot age.     

Detritus-Bacteria-Meiofauna interactions in a seagrass bed (Posidonia oceanica) of the NW Mediterranean

The biochemical composition of the sediment organic matter, and bacterial and meiofaunal dynamics, were monitored over an annual cycle in aPosidonia oceanica bed of the NW Mediterranean to test the response of the meiofauna assemblage to fluctuations in food availability. Primary production cycles of the seagrass and its epiphytes were responsible for relatively high (compared to other Mediterranean systems) standing stocks of organic carbon in sediments (from 1.98 to 6.16 mg Cg^–1 sediment dry weight). The biopolymeric fraction of the organic matter (measured as lipids, carbohydrates, and proteins) accounted for only a small fraction (18%) of the total sedimentary organic carbon. About 25% of the biopolymeric fraction was of microphytobenthic origin. Sedimentary organic carbon was mostly refractory (56 to 84%) and probably largely not utilizable for benthic consumers. The biopolymeric fraction of the organic matter was characterized by high carbohydrate concentrations (from 0.27 to 5.31 mg g^–1 sediment dry weight in the top 2 cm) and a very low protein content (from 0.07 to 0.80 mg g^–1 sediment dry weight), which may be a limiting factor for heterotrophic metabolism in seagrass sediments. RNA and DNA concentrations of the Sediments varied significantly during the year. High RNA and DNA values occurred during the microphytobenthic bloom and in correspondence with peaks of bacterial abundance. Bacteria accounted for a small fraction of the total organic carbon (0.65%) and of the biopolymeric organic carbon (4.64%), whilst microphytobenthos accounted for 3.79% of total organic carbon and for 25.08% of the biopolymeric carbon. Bacterial abundance (from 0.8 to 5.8 × 10⁸ g^–1 sediment dry weight) responded significantly to seasonal changes of organic matter content and composition and was significantly correlated with carbohydrate concentrations. Bacteria might be, in the seagrass system, an important N storage for higher trophic levels as il accounted for 25% of the easily soluble protein. pool and contributed significantly to the total DNA pool (on average 12%). Total meiofaunal density ranged from 236 to 1858 ind. 10 cm^–2 and was significantly related, with a time lag, to changes in bacterial standing stocks indicating that microbes might represent an important resource. Bacterial abundance and biomass were also significantly related to nematode abundance. These results indicate that bacteria may play a key role in the benthic trophic     

Trophic diversity of idoteids (Crustacea, Isopoda) inhabiting the Posidonia oceanica litter

The coexistence of three idoteid species in Posidonia oceanica litter raises the question of trophic diversity and their role in the litter degradation process. Hence, diet composition of Idotea balthica, Idotea hectica and Cleantis prismatica was studied using a combination of gut contents and stable isotopes analysis. Gut content observations indicate that P. oceanica dead leaves are an important part of the ingested food for the three species, although their tissues are constituted of only a small to medium fraction of P. oceanica carbon. Our results also underlined the potential role of these species in the degradation of P. oceanica litter by mechanically fragmenting the litter and by assimilating a small to medium fraction of carbon. Moreover, we showed that there were considerable inter- and intra-specific differences in diet composition. Diet differed between juveniles and adults for I. balthica. Crustaceans are an important food source for adults of I. balthica, while I. hectica indicated a major contribution of algal material. C. prismatica showed an intermediate diet. This trophic diversity is probably one of the factors allowing these species to coexist in the same biotope.     

Primary production of Posidonia oceanica in the Mediterranean Basin

Primary production of the marine phanerogam Posidonia oceanica (Linnaeus) Delile was measured by lepidochronological analyses at 22 sites in the Mediterranean Sea (Corsica, France, Italy, Sardinia and Turkey), between 1983 and 1992, to determine spatial and temporal variations. Leaf production (blade and sheath) ranged from 310 to 1 540 mg dry wt shoot^–1 yr^–1, depending on site and depth. Rhizome production ranged from 24 to 120 mg dry wt shoot^–1 yr^–1 (6% of average leaf production). At some sites the results obtained by lepidochronological analysis were consistent with earlier results obtained by classic methods (e.g. leaf-marking). While primary production per shoot (mg dry wt shoot^–1 yr^–1) displayed no significant differences between sites, primary production of the P. oceanica meadow (g dry wt m^–2 yr^–1) decreased with increasing depth at all sites studied. This decrease correlated with reduced density of the meadow (number of shoots per m²) with increasing depth. Past primary production was also extrapolated at three sites at the island of Ischia (Italy) for a period of 5 yr in order to determine interannual variations over a period of several years. While major variations were recorded for the surface stations (5 and 10 m depth), production remained stable at the deepest station (20 m depth). Given the large geographical scale of the study (location, depth range), it would appear that while P. oceanica production remains considerable, the values recorded in the literature on the basis of classical analyses (surface stations) represent maxima, and cannot be generalised for meadows as a whole.     

Age and growth of the fan mussel Pinna nobilis from south-east Spanish Mediterranean seagrass (Posidonia oceanica) meadows

The calcitic and aragonitic shell of the fan mussel Pinna nobilis L. contains a record of the environmental changes experienced during its growth. Stable-isotope analyses of oxygen (¹⁸O:¹⁶O) in shell carbonate from the calcitic outer shell-layer have been used to validate the periodicity of clearly defined concentric rings on the aragonitic posterior adductor-muscle scar and to estimate the age and growth of fan mussels growing in Posidonia oceanica (L.) meadows at four locations on the south-east Spanish Mediterranean coast. The stable oxygen-isotope records obtained at intervals along a profile across the shell surface enabled seasonal changes in water temperature to be established, and hence seasonal patterns of shell growth to be inferred. Muscle-scar rings were found to be deposited annually in the shell in the spring and early summer (a period of increasing water temperatures), and represent an interruption in the migration of the posterior adductor muscle along the inner surface of the shell. In small pinnids (<25 cm) accretion of the shell is rapid during the first year, but in the second year it is distinctly slower than at the same time the previous year. This slowing down in growth during the second year coincides with the appearance of the “first” distinct muscle-scar ring, indicating that Pinna nobilis does not form a muscle-scar ring during its first year of shell growth. Maximum growth rates were recorded amongst pinnids from Carboneras, where they achieved a length of 59 cm in 8 yr, whilst those from Aguamarga were estimated to be the oldest (attaining a length of 45 cm in 13 yr). Received: 26 January 1998 / Accepted: 8 October 1998     

Influence of hydrodynamic conditions on the production and fate of Posidonia oceanica in a semi-enclosed shallow basin (Stagnone di Marsala, Western Sicily)

An integrated approach using hydrodynamic and transport numerical models, lepidochronology and stable isotope analysis was used to investigate how local hydrodynamic conditions influence the primary production and fate of the seagrass Posidonia oceanica in a Mediterranean semi-enclosed marine system (Stagnone di Marsala). The water mass exchange aptitude of different sectors of the basin was analysed, and data collected were used to select two sectors (colonized by Posidonia oceanica showing the lowest and highest water exchange values) for biological analyses. According to the mean dispersal coefficient differences simulated by the hydrodynamic model, growth rate and primary production of P. oceanica differed between sectors, with average values lower in the central sector where water exchange is lower than in the southern sector. Although P. oceanica coverage and primary production were higher in the southern sector, carbon and nitrogen stable isotope analysis suggests that the transfer of seagrass organic matter to higher trophic levels of the food web was higher in the central sector. The possibility of a link between hydrodynamism, production and fate of organic matter is proposed to explain the observed patterns.     

Influence of nutrients in the feeding ecology of seagrass (Posidonia oceanica L.) consumers: a stable isotopes approach

Nitrogen inputs to coastal environments can considerably alter the abundance of primary producers. However, how herbivores modify their trophic signatures and adjust to changes in food resource conditions remains controversial. Here, we assess the effect of nutrient availability on the diet shifts of the two main Mediterranean herbivores, the Sparid fish Sarpa salpa L. and the sea urchin Paracentrotus lividus (Lmk.) that feed mostly on the seagrass Posidonia oceanica L. (Delile), epiphytes and benthic macroalgae. To do this, we (1) investigate the patterns of isotopic composition (δ¹³C and δ¹⁵N signatures) of the two herbivores and their potential food sources in three areas of contrasting nutrient conditions and, (2) we assess the diet shift along this nutrient gradient by estimating the isotopic nutrient enrichment (i.e., the contribution of δ¹³C and δ¹⁵N signatures in consumers’ tissues relative to potential food sources). Food web signatures of δ¹³C were similar among the three study sites, and no patterns of δ¹³C shift were observed in their diets. In contrast, there was a consistent increase in N contents and δ¹⁵N along the nutrient gradient for all primary producers and their consumers. The rate of δ¹⁵N enrichment was also clearly distinctive between the two herbivores: in P. lividus it increased by 61% along the nutrient gradient, while in S. salpa it remained constant. Our results suggest that sea urchins behave as facultative omnivores and feed on vegetable or mixed diets depending on the trophic status of the system. It is unclear, however, if this modification is behavioral or the consequence of mere changes in the availability of food items, as animal epiphytes (e.g., hydrozoans, bryozoans and ascidians) can also became more abundant on seagrass leaves under increased nutrient conditions. In contrast, adult fish appear to feed on vegetal material independent of nutrient availability in the ecosystem.     

Flowering of the seagrass Posidonia oceanica in a north-western Mediterranean coastal area: temporal and spatial variations

Sexual reproduction in Posidonia oceanica (L.) Delile is likely to be highly variable both in space and time. However, there has been little effort to identify at what spatial scale variability exists in the distribution and abundance of reproductive shoots or the magnitude of these variations. Such information is important for elucidating several aspects of the reproductive strategy of this species. Here, by using a reconstructing technique, I estimated the rate at which flowering events occurred over the last decades in three P. oceanica meadows along the coastline of the north-western Mediterranean (Italy). Then, variation in inflorescence frequency (i.e. the percentage of shoots that flowered in a particular year of the total number of shoots present in the sample in that year) was examined on a hierarchy of spatial scales, ranging from a few metres to meadows separated by tens of kilometres over a 3-year period. Separate analyses were also conducted to examine the patterns of spatial variability of several plant characteristics (i.e. shoot age and density of vegetative shoots) that may influence the timing of flowering. In total, ten reconstructed flowering events were recorded during the period 1983–1999. Despite the similar climatic conditions experienced in the three meadows surveyed, no inflorescence scars were found in one meadow. In the other two meadows, the flowering rate fluctuated considerably from year to year (0–26%), and simultaneous flowerings were detected only in 3 years (1993, 1994 and 1995). The pattern of spatial variability in the inflorescence frequency varied among years. An investigation of the flowering events recorded along the north-western Mediterranean coast reveals that in the studied period flowering also occurred across a broad range of meadows. Therefore, flowering could be a far more widespread phenomenon in this sector of the Mediterranean basin than previously thought. Analysis of variance showed significant variation in 1993 at the mean inflorescence frequency on very small scales (metres to tens of metres), whereas variation was not found at any of the scales investigated in 1994 and 1995. Overall, variation in the inflorescence frequency within individual meadows (65–93%) was at least an order of magnitude greater than that among separate meadows (0.5–19%). Patterns of spatial variation in the plant characteristics examined did not reflect those of flowering. It may be hypothesised that internal rhythms of populations, in conjunction with changes in environmental conditions, which are consistent over relatively large spatial (kilometres), are involved in the control of flowering induction. Environmental heterogeneity (in terms of nutrient availability) on a very local scale (metres to tens of metres) and intrinsic factors might affect the distribution of reproductive shoots within P. oceanica meadows.Communicated by R. Cattaneo-Vietti, Genova     

Comparative analysis of macrofaunal species richness and composition in Posidonia oceanica, Cymodocea nodosa and leaf litter beds

We investigated macrofaunal species richness and composition in Posidonia oceanica, Cymodocea nodosa and Leaf litter beds within a coastal area of the Gulf of Oristano in proximity of the Cabras lagoon (western Sardinia, Italy). A total of 124 taxa were found, of which 116 were identified at the species level. They were analyzed based on both taxonomic and substrate affinity classification. Presence/absence analysis revealed that P. oceanica, C. nodosa and Leaf litter were all characterized by a conspicuous number of soft-bottom polychaetes (e.g., Prionospio multibranchiata and Ampharete acutifrons) and crustaceans (e.g., Corophium sextonae and Dynamene bidentatus), also known as detritivores. There were also major differences between the three habitats investigated. Consistent with its structural complexity, P. oceanica showed the highest species richness [E(S ₅₀)] and the most diversified macrofaunal assemblages, both in terms of taxonomic groups and taxa associated with different substrates. The two seagrasses, however, showed a similar species composition and differed from Leaf litter for the exclusive presence of hard-bottom species (e.g., the tunicate Phallusia fumigata) and seagrass-associated species (e.g., the polychaete Syllis garciai and the decapod Paguristes syrtensis). In contrast, Leaf litter showed the most differences between the habitats, and was characterized by the bivalves Abra alba and Cerastoderma glaucum, not found in seagrass beds, and by Loripes lacteus and Ruditapes decussatus. Leaf litter also had the highest content of organic matter (26.7% ± 1.4) and total organic carbon (10.3% ± 0.4). Our results confirmed the facilitative role of living seagrasses, in particular P. oceanica, as related to their structural complexity, for numerous species from different substrates (e.g., hard bottom species). This study also showed that leaf litter beds act as a particular environment where sediment instability, leaf breakdown, and organic matter enrichment and decomposition strongly influence animal distribution. Finally, our results highlighted the ecological and trophic importance of seagrass-derived detritus and the associated macroinvertebrate detritivores within seagrass-dominated systems.     

Rooting response in stem cuttings from five species of mangrove trees: effect of auxins and enzyme activities

An experiment was conducted to optimise rooting in stem cuttings of the mangrove tree species Bruguiera parviflora, Cynometra iripa, Excoecaria agallocha, Heritiera fomes and Thespesia populnea. Cuttings were girdled or non-girdled and treated with the auxins indole-3-acetic acid (IAA), indole-3-butyric acid (IBA) and naphthalene-acetic acid (NAA) and amylase, IAA-oxidase, peroxidase and polyphenol oxidase activities were measured. The best rooting performance (in terms of root number) was in stem cuttings of E. agallocha followed by C. iripa and H. fomes when treated with IBA + NAA. IAA and IBA combinations were more effective in T. populnea and B. parvifolia. Variations in rooting response were markedly reflected in amylase, IAA-oxidase, peroxidase and polyphenol oxidase activities. Sharp increases in amylase and polyphenol-oxidase activities were associated with enhanced rooting in E. agallocha, C. iripa and H. fomes. On the contrary, lowered IAA-oxidase and peroxidase activities favoured rooting in T. populnea and B. parviflora. A higher number of roots occurred in pre-girdled cuttings in comparison to non-girdled cuttings. Received: 30 March 1999 / Accepted: 6 October 1999     

Wanted dead or alive: high diversity of macroinvertebrates associated with living and ‘dead’ Posidonia oceanica matte

The Mediterranean endemic seagrass Posidonia oceanica forms beds characterised by a dense leaf canopy and a thick root-rhizome ‘matte’. Death of P. oceanica shoots leads to exposure of the underlying matte, which can persist for many years, and is termed ‘dead’ matte. Traditionally, dead matte has been regarded as a degraded habitat. To test whether this assumption was true, the motile macroinvertebrates of adjacent living (with shoots) and dead (without shoots) matte of P. oceanica were sampled in four different plots located at the same depth (5–6 m) in Mellieha Bay, Malta (central Mediterranean). The total number of species and abundance were significantly higher (ANOVA; P<0.05 and P<0.01, respectively) in the dead matte than in living P. oceanica matte, despite the presence of the foliar canopy in the latter. Multivariate analysis (MDS) clearly showed two main groups of assemblages, corresponding to the two matte types. The amphipods Leptocheirus guttatus and Maera grossimana, and the polychaete Nereis rava contributed most to the dissimilarity between the two different matte types. Several unique properties of the dead matte contributing to the unexpected higher number of species and abundance of motile macroinvertebrates associated with this habitat are discussed. The findings have important implications for the conservation of bare P. oceanica matte, which has been generally viewed as a habitat of low ecological value.     

Detection of lipid peroxidation and cytotoxicity induced by aluminium (Al) and cobalt (Co) ions in barbunia root tip cells

The aim of this study was to investigate the cytotoxic effects of different concentrations of Aluminium (Al) and Cobalt (Co) heavy metal ions on Phaseolus vulgaris L. cv Barbunia (Fabaceae) root tips. We used the germination percentage (GP), root length (RL), weight gain (WG) and micronucleus (MN) frequency as indicators of cytotoxicity and correlated these data with statistical parameters. Additionally to the cytogenetic analysis, lipid peroxidation and DNA analyses were performed in root tips of barbunia seeds treated with Al and Co metals. The seeds were divided into five groups as control, Al and Co treatment groups. They were treated with 25 and 50 ppm doses of Al and Co during 7 days. The results indicated that there was an alteration in the GP, RL, WG and MN frequency depending on the treatment dose in the seeds exposed to Al and Co metal ions when compared with the controls. Al and Co metal ions at both the doses significantly reduced the GP, RL and WG in seeds of all the treatment groups. The highest GP was observed in seeds of the control group (in proportion as 96%). 25 and 50 ppm doses of Co and Al caused 30, 50 and 42, 64% decrease of seed germination, respectively. In the control group, the final weights of all the seeds increased about 1.31 g when compared to initial weight. The mean RL of control seeds were measured as 3.71 cm at the end of experimental period. In Co andAl groups, the final weights of seeds increased about 0.34 g and 0.19 g according to initial weight at 50 ppm dose, respectively But, Al and Co ions caused a dose-dependent increase in the frequency of MN. The highest frequency of MN was observed at 50 ppm dose of Al and least frequency of MN was observed at 25 ppm dose of Co. Besides, 25 and 50 ppm concentrations of Al and Co significantly enhanced the lipid peroxidation and caused an increase in malondialdehyde (MDA) levels at both the doses. In roots treated with 25 and 50 ppm doses of Al, the increase of MDA was about 62 and 136% according to control, respectively In Co-treated roots, the increase of MDA was about 31 and 91% according to the control at 25 and 50 ppm doses, respectively The investigated parameters (except MN and MDA) were higher in the seeds exposed to Co than the seeds treated with Al. Moreover, it was observed that the yields of DNA in the seeds treated with Al andCo metals were lower than recorded in the controls. Hence, DNA yields exposed to Al and Co were run ahead on agarose gel according to the control group. The results of the present study indicate that Al and Co metal ions have toxic effects on barbunia root tip cells, and the selected parameters such as the GP, RL, WG, MN and MDA are very sensitive and useful biomarkers for biomonitoring these effects.     

Neutron-activation analysis of the elementary composition of the marine phanerogamPosidonia oceanica from a reference area in Port Cros National Park (French Mediterranean)

The marine phanerogamPosidonia oceanica (L.) Delile was collected from a presumably unpolluted reference area in Port Cros National Park, Mediterranean Sea, in November 1985, and its elemental composition was studied by gamma spectrometry after neutron activation and by atomic-absorption spectrophotometry. Neutron-activation analysis revealed 23 elements and atomic-absorption spectrophotometry 4 others. Including all known records by other investigators, this brings the total of elementary components identified inP. oceanica up to 42: antimony, arsenic, barium, bromine, cadmium, cesium, calcium, carbon, cerium, chromium, cobalt, copper, europium, gold, hafnium, hydrogen, iodine, iron, lanthanum, lead, magnesium, manganese, mercury, molybdenum, nickel, nitrogen, phosphorus, potassium, rubidium, samarium, scandium, selenium, silver, sodium, strontium, tantalum, terbium, thorium, uranium, ytterbium, zinc, zirconium. These elements display significant differences in their distribution in the various parts of the plant (roots, rhizomes, and growing and freshly budded leaves).     

Ri质粒诱导喜树毛状根培养基中喜树碱的研究

喜树(Camptotheca acuminate)可产生具有抗癌作用的喜树碱,有许多用细胞培养方法制备喜树碱的尝试,但都因极低的产率而无从深入。植物毛状根培养的应用将成为一种生物碱生产的替代方法。以发根农杆菌(Agrobacterium rhizogenes)A41.2556感染喜树诱导产生毛状根,建立喜树碱毛状根产生体系。甘露碱检测及PCR结果证实发根农杆菌Ri质粒的rolB基因已在喜树毛状根基因组中整合并得到表达。确定了用RP-HPLC测定发根培养基中喜树碱的的方法,比较了不同的吸附树脂对培养基中喜树碱的富集能力,从培养30d的喜树培养基中获得喜树碱干质量分数为0.14mg·g-1。     

萘乙酸和Zn对大豆Cd胁迫伤害的缓解效应

为探讨植物生长调节剂对大豆Cd胁迫伤害的缓解效应,采用营养液培养试验方法,研究了喷施萘乙酸(NAA)和加Zn处理对3个Cd胁迫大豆幼苗的影响.结果表明,加Zn和喷施NAA均可降低Cd胁迫大豆幼苗叶片丙二醛(MDA)和脯氨酸(PRO)含量,可减轻膜脂的过氧化作用及蛋白质的水解;喷施NAA还可降低Cd胁迫大豆幼苗POD活性,提高硝酸还原酶(NR)活性;加Zn对Cd胁迫大豆幼苗NR活性的降低缺乏抑制作用,但降低沔1101叶片中POD活性,提高湘04-6和特早熟毛豆的POD活性,品种之间表现明显的差异性.     

Dispersal, pair formation and social structure in gibbons (Hylobates lar)

We report observations on reproduction, natal dispersal, pair formation, and group structure based on longitudinal observations of several white-handed gibbon (Hylobates lar) groups spanning 18 years. Our observations are at odds with the traditional view that gibbons live in nuclear family groups consisting of a pair of adults and their offspring, and that parents exclude young from the family territory when they reach adult size. In the relatively dense Khao Yai study population, dispersing young usually obtain mates by replacing adults in existing territories, which creates non-nuclear families. Six subadults, five males and one female, matured and dispersed at an average age of 10 years, or about 2 years after reaching adult size. Average natal dispersal distance was 710 m, or between one and two territories away. At least two dispersing males replaced adults in neighboring groups. In one case, forcible displacement of the resident male resulted in a group which included a young juvenile presumably fathered by the previous male, two younger juveniles (probably brothers) from the new male's original group, and (later) offspring of the new pair. Social relations within this heterogeneous group remained harmonious: the adults groomed all the young and play occurred between all preadult members. In only two out of a total of seven cases of dispersal seen did two subadults pair and disperse into new territorial space. Nonreproducing subadults which delay dispersal may be tolerated by the adults provided that they contribute benefits to the adults or their offspring. Possible benefits include behaviors such as grooming, social play with juveniles, and support of the adult male in defending the territory. Delayed dispersal is probably advantageous in a saturated environment where there is no room for floaters, but subadults may also gain indirect fitness benefits by aiding siblings and other relatives. Received: 24 January 1997 / Accepted after revision: 12 January 1998     

Lactate formation in Callianassa californiensis and Upogebia pugettensis (Crustacea: Thalassinidea)

The ability to accumulate lactate as a result of laboratory exposure to anoxia was examined in 2 species of mud-dwelling shrimp, the ghost shrimp Callianassa californiensis and the mud shrimp Upogebia pugettensis. Hemolymph lactate accumulated to a much greater degree in the mud shrimp, even though the overall levels of lactate dehydrogenase activity were similar in tissues of the two species. Carbohydrate reserves, estimated as glycogen, were not significantly depleted in muscle or midgut gland of either species as a result of anoxic stress. The results are discussed in relation to possible metabolic strategies employed to cope with the hypoxic environment of these crustaceans.