Sperm competition in the damselfly Enallagma hageni Walsh (Odonata: Coenagrionidae): benefits of multiple mating to males and females

Summary Sperm competition was investigated in the non-territorial damselfly, Enallagma hageni. Using irradiated (sterile) male techniques, the last male to mate was found to fertilize up to 95% of the eggs of the first clutch laid after mating. Dissection of females collected before, during, and after copula showed that a male removes a maximum of 87% of the sperm by volume of a previous mate. These data verify an earlier estimate of lifetime reproductive success in this species which was based on mating success, and suggest that indirect dissection methods offer minimum estimates of sperm precedence. Male E. hageni have ample opportunity to benefit from sperm precedence, since at least 10% of the receptive females encountered had already mated once that day, but still contained complete or partial clutches of eggs. Female E. hageni benefit directly from high sperm precedence because it allows them to exchange matings for guarding service by males during oviposition bouts under water.     

Why do male Callosobruchus maculatus beetles inseminate so many sperm?

Male Callosobruchus maculatus F. (Coleoptera: Bruchidae) inseminate more sperm than females can effectively store in their spermathecae. This study examines the adaptive significance of excess sperm transfer by measuring components of male and female reproductive success in response to manipulating the number of sperm inseminated. The number of sperm transferred during copulation was reduced from 56,000 ±4,462 to 8,700±1,194 by sequentially mating males to virgin females. Reducing the number of sperm inseminated by the first male to mate had no effect on the extent of sperm precedence, but reducing the number of sperm inseminated by the second male resulted in a significant reduction in the extent of sperm precedence. When large numbers of sperm are inseminated the remating refractory period of females is increased. These results indicate that males transferring large numbers of sperm during copulation have a two-fold advantage at fertilization; they are more effective at preempting previously stored sperm and they are likely to father more offspring by delaying the time of female remating. The transfer of excess sperm does not appear to serve as nonpromiscuous male mating effort; the number of eggs laid, their fertility and the subsequent survival of zygotes were unaffected by manipulating the number of sperm inseminated. The underlying mechanisms of sperm precedence were also examined. Simple models of sperm displacement failed to accurately predict the patterns of sperm precedence observed in this species. However, the results do not provide conclusive evidence against the models but rather serve to highlight our limited understanding of the movement of sperm within the female's reproductive tract.     

Re-copulation and post-copulatory mate guarding increase immediate female reproductive output in the dragonfly Nannophya pygmaea Rambur

After copulation, male Nannophya pygmaea dragonflies mate guard by hovering over ovipositing females and repelling conspecific males. Copulation is not always a prerequisite for oviposition in the females of this species because females can store the sperm received during previous visits/copulations. An oviposition episode consists of several bouts of oviposition separated by periods of perching. We conducted two types of male-removal experiments to examine the effects of mating and post-copulatory mate guarding on the oviposition behaviour of females. In the first experiment, we removed all males from the habitat to eliminate the effect of re-copulation, mate-guarding and harassment by males. In the second experiment, we removed males immediately after copulation to eliminate the effects of guarding and other post-copulatory male-female interactions. We compared these experimental data with data obtained under natural conditions. The dipping rate in an oviposition bout was not influenced by copulation or guarding. However, guarded females made more dips per episode than did solitary females. The proportion of time actually spent ovipositing (total bout duration/oviposition episode duration) of guarded females was higher than that of solitary females. Solitary females often oviposited in more than one territorial site, while guarded females usually oviposited within a single territorial site during an oviposition episode. Because males tend to hold territories at sites where egg survival is high, guarded females (and the male guardian) benefit from guarding in terms of egg hatchability. The possible benefits for solitary females are discussed.     

Variation in copulation duration in Mnais pruinosa pruinosa Selys (Odonata: Calopterygidae)

Summary The Japanese calopterygid damselfly Mnais pruinosa pruinosa shows three distinct copulation durations (Siva-Jothy and Tsubaki 1989). A variety of factors which might influence copulation duration were investigated. Results indicate that the temperature in the vicinity of copulating pairs, the rate at which males encountered receptive females, territory quality, the rate of agonistic interactions between males, female gravidity and female willingness to oviposit have no effect on copulation duration. The most significant factor was the location of the site at which males captured their mates. We suggest that under natural conditions the location of the capture site provides males with reliable, indirect information about a female's intention to oviposit: it is important for males of this species to acquire such information since sperm precedence, and therefore reproductive success, is in part dependant on the interval experienced by females between copulation and subsequent oviposition. The results are discussed and it is suggested that the three observed mate-securing tactics of M.p. pruinosa are facultative and information-dependant.     

Mate guarding,copulation strategies and paternity in the sex-role reversed,socially polyandrous red-necked phalarope<Emphasis Type="Italic"> Phalaropus lobatus</Emphasis>

In a recent review, Westneat and Stewart (2003) compiled evidence that extra-pair paternity results from a three-player interaction in which sexual conflict is a potent force. Sequentially polyandrous species of birds appear to fit this idea well. Earlier breeding males may attempt to use sperm storage by females to obtain paternity in their mates subsequent clutches. Later-breeding males may consequently attempt to avoid sperm competition by preferring to pair with previously unmated females. Females may bias events one way or the other. We examined the applicability of these hypotheses by studying mating behavior and paternity in red-necked phalaropes (Phalaropus lobatus), a sex-role reversed, socially polyandrous shorebird. Male red-necked phalaropes guarded mates more strongly than other shorebirds. Males increased within-pair copulation attempts during their mates fertile period, and maintained or further increased attempts towards the end of laying, suggesting an attempt to fertilize the females next clutch; these attempts were usually thwarted by the female. Paired males sought extra-pair copulations with females about to re-enter the breeding pool. Multilocus DNA fingerprinting showed that 6% of clutches (4/63) each contained one chick sired by a male other than the incubator, producing a population rate of these events of 1.7% (n=226 chicks). Male mates had full paternity in all first clutches (n=25) and 15 of 16 monogamous replacement clutches. In contrast, 3 of 6 clutches of second males contained extra-pair young likely fathered by the females previous mate. Previously mated female phalaropes may employ counter-strategies that prevent later mating males from discriminating against them. The stability of this polyandrous system, in which males provide all parental care, ultimately may depend on females providing males with eggs containing primarily genes of the incubating male, and not a previous mate.Communicated by M. Webster     

Copulatory plug displacement and prolonged copulation in the Iberian rock lizard (Lacerta monticola)

Male Iberian rock lizards (Lacerta monticola) produce copulatory plugs that adhere firmly inside the female cloaca and occlude both oviducts. These plugs do not prevent rival male insemination, as they neither reduce female attractiveness or receptivity to rival males nor do they function as chastity belts. Prior to copulation, males bite various regions of the female body, including the cloacal region. A previous plug in the female cloaca is expelled only after rival male intromission. We hypothesized that: (1) such male pre-copulatory behaviours function to loosen plug adherence to the female cloaca, thereby facilitating intromission, and that (2) the hemipenis plays a role in displacing a previous plug prior to the delivery of a new plug. Neither of these hypotheses was supported. Instead, our results indicate that rival males can intromit the hemipenis past a previous plug in the female cloaca and deliver their own plug underneath it. Consequently, previous plugs are pushed away from the oviductal openings and even dislodged from the female cloaca. Copulation duration was determined both by the time used by males to deliver a plug and by the fact that males prolonged copulation beyond plug delivery. There seemed to be sexual conflict over prolonged copulation, which was resolved by the male/female head-length ratio. The adaptive value of Iberian rock lizard copulatory plugs and prolonged copulation in Iberian rock lizards is discussed in the context of sperm competition.Communicated by S. Downes     

Circadian rhythmicity of locomotor behavior and oviposition in female Teleogryllus commodus

Summary Locomotor activity of the Teleogryllus commodus is under circadian control, with _LL=25.3 h and _DD=23.4 h. In LD 12:12 h, running occurs either exclusively during darkness (57%), mainly in the dark phase (35%), or with substantial activity peaks in both phases (8%). For oviposition, _LL>24 h and _DD<24 h; in LD 12:12 h, 80% of the studied deposited their eggs mainly during the light phase and 20% primarily during darkness. In the discussion, a temporal correlation between stridulation, spermatophore formation, and locomotion, and oviposition is established, which serves to make an encounter and mating between the sexes as likely as possible.Dedicated to Prof. Dr. M. Gersch on the occasion of his 70th birthday     

Extra-pair paternity and intraspecific brood parasitism in wild zebra finches Taeniopygia guttata,revealed by DNA fingerprinting

Summary The frequency of extra-pair parentage in a wild population of zebra finches Taeniopygia guttata was examined by DNA fingerprinting. A total of 25 families, comprising 16 pairs of parents and 92 offspring (in broods of 1 to 6) were examined. Ten cases of extra-pair parentage, presumed to constitute intraspecific brood parasitism, were detected (10.9% of offspring or 36% of broods), including one possible instance of quasi-parasitism (parasitism by a female fertilized by the male nest owner). The average number of parasitic eggs per clutch detected by fingerprinting was 1.10±0.32 SD, very similar to the one egg difference in average clutch size between parasitised (6.0±0.82) and unparasitised nests (5.0±0.95). Two cases of extra-pair paternity (EPP) were detected among 82 offspring whose maternity was confirmed: 2.4% of offspring, or 8% of broods. In both cases EPP accounted for only a single offspring within a brood. Behavioural observations show that EPP occurs through extra-pair copulation rather than rapid mate switching. The results are discussed in the light of what is known about the fertile period and sperm precedence patterns in this species. Offprint requests to: T.R. Birkhead     

Parental care of nestlings by male red-winged blackbirds

Summary Nestling feeding by males is less common among birds with polygynous mating systems than in monogamous species, because of the pronounced trade-off between parental behavior and the attraction of additional mates. In this study, however, we found that male red-winged blackbirds (Agelaius phoeniceus) commonly assisted females in feeding nestlings in several Ontario marshes. Male parental care was additional to that provided by females, and it significantly enhanced the fledging success of nests (Table 2). Male redwings did not help to feed all nests on their territories: primary and secondary nests were much more likely to receive male parental care than tertiary and later nests. Contrary to expectation, male parental care was not restricted to the nests of primary females: a greater proportion of secondary than primary nests were assisted (Tables 4a and b). The presence of new females settling on the territory at the same time that a resident female had nestlings, resulted in males deferring parental care until a later brood. This suggests that males trade off the recruitment of females against parental care to an existing brood. Although the number of nestlings fledging from a male's territory was strongly influenced by the number of females in the harem, males could additionally increase their reproductive success by feeding nestlings in one or more nests on their territories (Fig. 2). The reproductive success of females was significantly enhanced by male assistance in feeding nestlings (Table 3). However, those females not receiving male assistance on territories of feeding males did not suffer a significantly reduced reproductive success in comparison to females on territories of non-feeding males (Table 2). Males varied considerably in the quality of brood care given. We therefore suggest that the quality of male parental care may be a factor considered by females in choosing a breeding situation.     

Selective males and ardent females in pipefishes

Summary In the pipefishes Syngnathus typhle and Nerophis ophidion, males have been shown to limit female reproductive rate, and females to compete for access to males. Hence, these species fit the criteria for sex-role reversal. Males brood the eggs and provide the offspring with nutrients, oxygen and an osmoregulated environment. Moreover, in S. typhle both sexes prefer a larger mate when given a choice. Sexual selection theory predicts that males should be more choosy than females, and that was experimentally demonstrated in this study. We predicted that S. typhle males should be less eager to copulate than S. typhle females with an unattractive (i.e. small) mate. We measured eagerness as the time from the start of the experiment until copulation occurred. Males with unattractive partners took significantly longer to copulate than females with unattractive partners. Moreover, females invariably initiated the courtship dance, and resumed it quicker after copulation than did the males, again suggesting reproductive hesitation in males. Neither male nor female size per se was correlated with time until copulation. In N. ophidion, where we have previously shown that males prefer larger to smaller females, we found that females did not select males with regard to size. Our results are consistent both with earlier findings (males limit female reproduction and females compete for males) and with operational sex ratios in nature: in seven annual field samples in June, the numbers of S. typhle females with ripe eggs always significantly exceeded numbers of receptive males. Hence, the potential cost of being choosy in terms of lost matings is much higher in females than in males. In conclusion, S. typhle females were somewhat choosy, but less so than males, whereas N. ophidion females were not choosy at all. Correspondence to: A. Berglund     

Cloacal erection promotes vent apposition during forced copulation in the New Zealand stitchbird (hihi): implications for copulation efficiency in other species

Cloacal protuberances (CP) in male birds result from spermatic engorgement of storage tubules around the cloaca during the breeding season. We examined seasonal changes in the volume and orientation of the CP in the New Zealand stitchbird Notiomystis cincta. The male stitchbird has one of the largest recorded CPs for any species (max = 1,570 mm³), with CP volume increasing by almost 400% between the non-breeding and breeding seasons. While sperm competition has been positively correlated with the magnitude of CP storage in other species, no evidence previously existed for the CP improving copulation efficiency. By measuring the relative orientation of the CP throughout the year, we show that not only does the CP increase in size as males become sexually active, it also changes its orientation by approximately 60°. This results in it shifting from facing posteriorly to becoming almost perpendicular to the abdomen. This cloacal erection improves the apposition of the male and female cloacal openings during face-to-face forced copulation in this species. This provides the first reported evidence supporting the copulation efficiency hypothesis of the avian CP. While the magnitude of seasonal changes in female cloacal volume was similar to males, female cloacal orientation remained virtually unchanged across seasons. This difference between the sexes is likely to reflect differing selection pressures for optimizing sperm transfer. In females, a posterior-facing cloaca is ideal for both waste evacuation and sperm reception, whereas, for the male, a posterior-facing cloaca is well suited for waste evacuation, but possibly hinders sperm delivery. Changes in male cloacal orientation from the non-breeding to the breeding season are a likely reflection of conflict in this dual function. Evidence of changes in CP orientation in another passerine species suggests this phenomenon is widespread and also important for understanding related fields such as sperm competition, forced copulation and constraints on the evolution of the avian intromittent organ.     

Variation in copulation duration in Mnais pruinosa pruinosa Selys (Odonata: Calopterygidae)

Summary Males of the damselfly Mnais pruinosa pruinosa were observed to use three different tactics to secure mates. The mean duration of copulation differed between the three observed tatics and resulted in varying degrees of sperm removal and insemination. It is shown that the last male to mate had almost 100% sperm precedence immediately after copulation regardless of the duration of copulation and therefore the quantity of sperm removed. In situations where less than 100% of rivals' sperm was removed the sperm from different males mixed within the female sperm storage organs over a period of about 6 days: sperm mixing produced variation in last male sperm precedence. The significance of sperm mixing in M. p. pruinosa is discussed in the context of the observed matesecuring tactics and the frequent female habit (37% of observations) of ovipositing without remating during an oviposition bout.     

Paternity assurance by “helpful” males: adaptations to sperm competition in burying beetles

Summary Male burying beetles invest parentally by participating in the burial of a carcass and in provisioning and guarding the larvae that come to the carcass. Since most of the females arriving at a carcass have stored fertile sperm within their spermathecae, sperm transferred by such helpful males must compete with other males' ejaculates for the fertilization of the female's eggs. We showed that these males are able to achieve a high level of paternity (mean=92%). The mechanism they employ is a repeat-mating tactic, i.e., the female is mated very frequently shortly before and during oviposition. Repeated matings are essential for a high reliability of paternity, since single copulations result in the fertilization of only a very small proportion of the female's eggs.     

Spatial mesoscale patterns of West Pacific picophytoplankton as analyzed by flow cytometry: their contribution to subsurface chlorophyll maxima

Vertical distributions of picophytoplankton (ppp) (<2 m) were studied by ship-board flow cytometry during two cruises in Western Pacific waters to Palau and to Australia in 1990. Weak red-fluorescing small ppp, supposed to be free-living prochlorophytes (Chisholm et al. 1988), were abundant in the area surveyed. These ppp, designated the prochlorophytes, were abundant in the surface waters (>10⁴ cells ml^-1) at the northern region (27°03N; 7°11N) in November, whereas in December at the southern tropical stations (0°23.54S; 9°20.30S; 13°50.6S), they formed subsurface maximum layers (>10⁵ cells ml^-1) on a nitracline at a depth of 3.5 to 5.4% surface irradiation. Their fluorescence intensity increased with depth below 10% surface irradiation. The prochlorophytes at a depth of 1% surface irradiation had ten times higher fluorescence than those at the surface layer. The total fluorescence intensity of the prochlorophytes accounted for 32 to 63% of the sum of the total fluorescence intensity of all fluorescing phytoplankton detected at subsurface chlorophyll maxima in the tropical area. These results suggest that distribution of the prochlorophytes is greatly affected by nitracline and by light intensity and that their chlorophyll is a major contributor to the subsurface chlorophyll maximum in the pelagic West Pacific Ocean.     

Underwater oviposition in a damselfly (Odonata: Coenagrionidae) favors male vigilance,and multiple mating by females

Summary Female Enallagma hageni oviposit underwater where they are inaccessible to males. I demonstrate that males guard submerged females rather than perch sites, and are behaviorally distinct from lone males at the water. In contrast to lone males, which always attempt to copulate with females presented to them, guarding males exhibit a conditional latency to remating which corresponds closely to the time required by females to oviposit a complete clutch of eggs. By ovipositing underwater, females decrease the risk that their eggs become exposed. Risks associated with submerged oviposition favor both mate guarding, and multiple, within-clutch matings by females. Both guarding mates and lone males rescue females that float on the water surface as the result of improper resurfacing. Such behavior reduces the mortality risk to females from 0.06 to 0.02 per oviposition bout. By remating between bouts, females benefit from the additional vigilance of lone males, who rescue floating females 1.4 times as often as original mates. A second consequence of multiple mating is an increase in the selective advantage of vigilance by mates. Because receptive females become scarce by early afternoon, whereas male density remains high, a male has little (3%) chance of encountering a second receptive female that day. However, he incurs a 42% risk of losing fertilizations if he abandons a mate. For male E. hageni mate guarding functions in the context of both natural and sexual selection. It insures that a mate lives to lay a complete egg clutch in addition to protecting a male's sperm investment.     

Protandry and mate assessment in the wartbiter Decticus verrucivorus (Orthoptera : Tettigoniidae)

Summary Protandry, or early male emergence, is prevalent in the majority of insects. There are several explanations, both adaptive and incidental for this widespread phenomenon. Here I report the occurrence of protandry in the wartbiter, Decticus verrucivorus, and argue that the explanation for the evolution of protandry in this species is the result of selection acting on males in relation to sperm competition. It has previously been shown that sperm mixing occurs in this species, leading to a reduction in confidence of paternity with number of matings. Emerging early and thereby having a higher probability of mating with virgin females benefits males in two ways: both through a 100% assurance of paternity and because female egglaying rate decreases over time. A singly mated female can lay up to 30% of her lifetime egg production during her first refractory period, during which time the eggs are exclusively fertilized by the first male. Wartbiter males seem also to be able to assess female mating status and transfer larger spermatophores when mating with virgin females than when mating with already mated females. This holds true both for previously mated and virgin males. Thus, it seems that male wartbiters allocate their resources differentially depending on female quality.     

Effects of nitrogen and phosphorus enrichement on in vivo symbiotic zooxanthellae of Pocillopora damicornis

The reef coral Pocillopora damicornis (Linnaeus) was grown for 8 wk in four nutrient treatments: control, consisting of ambient, unfiltered Kaneohe Bay seawater [dissolved inorganic nitrogen (DIN, 1.0 M) and dissolved inorganic phosphate (DIP, 0.3 M)]; nitrogen enrichment (15 M DIN as ammonium); phosphorus enrichment (1.2 M DIP as inorganic phosphate); and 15 M DIN+1.2 M DIP. Analyses of zooxanthellae for C, N, P and chlorophyll a after the 8 wk experiment indicated that DIN enrichment increased the cellular chlorophyll a and excess nitrogen fraction of the algae, but did not affect C cell^-1. DIP enrichment decreased both C and P cell^-1, but the decrease was proportionally less for C cell^-1. the response of cellular P to both DIN and DIP enrichment appeared to be in the same direction and could not be explained as a primary effect of external nutrient enrichment. The observed response of cellular P might be a consequence of in situ CO₂ limitation. DIN enrichment could increase the CO₂ (aq) demand by increasing the net production per unit area. DIP enrichment could slow down calcification, thus decreasing the availability of CO₂ (aq) in the coral tissue.Hawaii Institute of Marine Biology Contribution No. 920     

Male capacity as related to sperm production,pregnancy initiation,and sperm competition in deer mice (Peromyscus maniculatus)

Summary Limitations on male capacity may be important to the evolution of mating strategies and behavior. Sperm counts in successive ejaculates of deer mice (Peromyscus maniculatus) decreased progressively through six ejaculations and remained depressed the following day. Males were as effective in inseminating females in cycling estrus with their second two ejaculates as with their first two and as effective in inseminating postpartum females with their fifth as with their first. In sperm competition situations males competed with rested males as effectively with their fourth, fifth, and sixth ejaculates as with their first. Thus, although sperm counts decrease in successive ejaculates, males suffer no apparent functional deficiency as long as they continue ejaculating. It is not clear whether or not deer mice are honest salesmen but they appear to be honest ejaculators and usually are honest intromitters.     

Environmental Geochemistry of Soils and Waters of Susaki Area, Korinthos, Greece

Soil and water samples were collected from the Susaki area of Korinthos and analysed for heavy metals in order to evaluate their environmental impact. The geology of the studied area includes ultrabasic rocks and Neogene and Quaternary deposits whereas magnesite veins are found within the ultrabasic rocks. In the north part of the studied area post volcanic emissions of H₂S, CO₂ and H₂O vapor continue to the present day. All the samples were analysed for heavy metals by the ICP method. The element ranges (in g g^–1) for soil samples are: Cu 11–63, Pb 5–256, Zn 21–604, Ni 183–2665, Co 12–124, Mn 456–1434, As 5–104, Sr 44–730, V 21–84, Cr 163–2346, Ba 48–218, Zr 3–41, Y 3–13. The metals Pb, Zn, Ni, Co, Cr, Fe, Cu, Mn, As and Sr are enriched in the Susaki soils. The element ranges for water samples are: Cu 65–103ppb, Pb<10ppb, Zn<5ppb, Ni 21–163ppb, Co 2–12ppb, As<30ppb, Cr<20ppb, Ba 36–785ppb, Sb<10ppb, W<10ppb, Bi<30ppb, Mn 0.0–0.9 g g^–1, Fe 0.01–0.22 g g^–1, Na 843–3076 g g^–1, K 98–278 g g^–1, Si 39–65 g g^–1, P 0.1–0.2 g g^–1. There is a natural pollution of soils with elevated concentrations of Ni, Co, Mn, Fe and Cr due to the presence of ultrabasic rocks. Another natural case of As pollution of soils is due to the volcanic activity and the geothermal field in the area. The geochemical data of ground waters and also the ¹⁸, D data showed a mixing in different proportions between sea water and meteoric water.     

Effects of salinity on respiration and nitrogen excretion in two species of echinoderms

The 30-d survival limit of Eupentacta quinquesemita and Strongylocentrotus droebachiensis is 12–13 S. The activity coefficient (1 000/righting time in seconds) of stepwise acclimated sea urchins declined from 16.3 at 30 S to 3.5 at 15 S. Oxygen consumption rates (QO₂) of both species held at 30 S and 13°C were highest in June and lowest in December. During the summer, when environmental salinity is most variable in southeastern Alaska, the QO₂ of both species held at 30, 20 and 15 S varied directly with salinity. Perivisceral fluid PO₂ varied directly with acclimation salinity in sea urchins, but not in sea cucumbers. Perivisceral fluid oxygen content of acclimated sea urchins was significantly lower at 15 and 20 S than at 30 S due to reduced PO₂ and extracellular fluid volume at the lower salinities. The QO₂ of both species varied directly with ambient salinity during a 30-10-30. semidiurnal pattern of fluctuating salinity. No change occurred in the average QO₂ of either species over a 15-30-15. semidiurnal pattern of fluctuating salinity. Sea urchin perivisceral fluid PO₂ declined as ambient salinity fluctuated away from the acclimation salinity in both cycles and increased as ambient salinity returned to the acclimation salinity. Total nitrogen excretion of stepwise acclimated sea cucumbers declined significantly from 30 to 15 S, but there was no salinity effect on total nitrogen excretion in sea urchins. Ammonia excretion varied directly with salinity in stepwise acclimated sea cucumbers (67–96% of total nitrogen excreted), but there was no salinity effect on ammonia excretion (89–95% of total nitrogen excreted) of sea urchins. Urea excretion did not vary with salinity in sea cucumbers (2–4% of total nitrogen excreted) or sea urchins (2–9% of total nitrogen excreted). Primary amines varied inversely with salinity in sea cucumbers (2–30% of total nitrogen excreted), but did not vary with salinity in sea urchins (2–4% of total nitrogen excreted). The oxygen: nitrogen ratio of both species indicated that carbohydrate and/or lipid form the primary catabolic substrate. The O:N ratio did not vary as a function of salinity. Both species are more tolerant to reduced salinity than previously reported, however, rates of oxygen consumption and/or nitrogen excretion are modified by salinity as well as season.