Reproductive biology of a filter-feeding amphipod, Leptocheirus pinguis , with extended parental care

Leptocheirus pinguis (Stimpson, 1853) is a widely distributed, abundant, endobenthic amphipod that engages in extended parental care, i.e. females host their juveniles in their burrows for extended time periods. I examined reproduction and population biology of L. pinguis at mean low water (MLW) in muddy sediment in Lowes Cove, Maine, USA. Cores around individual burrows were taken monthly in 1994 and 1995, and four seasonal samples were taken at different tidal heights. During the major reproductive periods in spring/early summer and in the fall, females produced several consecutive broods and hosted growing offspring in their burrows. Juveniles remained in their mothers' burrows until they reached a length of 5 mm (approximately one-third adult size) or more. At the study site, the majority of amphipods in individual burrows were adult females. Following the main reproductive periods, subadult individuals were found in their own burrows, but densities did not increase following the reproductive period in spring/early summer 1994, probably because large numbers of L. pinguis emigrated via the water column between June and December. L. pinguis is an annual species. Many members of the cohort born in spring/early summer start reproducing in the fall, and survive until the following spring when they produce several broods. Members of the cohort born in the fall start reproducing the following spring and also produce several consecutive broods. Both the spring/early summer and fall cohorts die off after the major reproductive period in the following spring/early summer. High standing stocks of microphytobenthos occur in soft-bottoms at MLW, and I conclude that L. pinguis can engage in extended parental care there because its food is abundant year-round. The limited expandability and low stability of burrows in soft-bottoms at MLW do not permit long persistence of parent–offspring groups in L.␣pinguis. Received: 27 March 1997 / Accepted: 30 July 1997     

Reproductive biology of a wood-boring isopod, Sphaeroma terebrans, with extended parental care

The isopod Sphaeroma terebrans Bate, 1866 burrows in aerial roots of the red mangrove Rhizophora mangle L. The burrows serve as shelter and as a reproductive habitat, and females are known to host their offspring in their burrows. I examined the reproductive biology of S. terebrans in the Indian River Lagoon, a shallow lagoon stretching for ∼200 km along the Atlantic coast of Florida, USA. Reproductive isopods were found throughout the year, but reproductive activity was highest in the fall and during late spring/early summer. During the latter periods, large numbers of subadults established their own burrows in aerial roots. The average numbers of S. terebrans per root were high during the fall, but decreased during the winter and reached lowest levels at the end of the summer. Females reached maturity at a larger size than males, but also grew to larger sizes than the males. The average size of females varied between 8 and 10 mm, the average size of males between 6.5 and 8.5 mm. The number of embryos female⁻¹ was strongly correlated with female body length. No indication for embryo mortality during development was found. Parental females (i.e. with juveniles in their burrows) hosted on average between 5 and 20 juveniles in their burrows (range 1 to 59). Most juveniles found in female burrows were in the manca stage and 2 to 3 mm in body length. Juveniles did not increase in size while in the maternal burrow, and juveniles of similar sizes could also be found in their own burrows. Males did not participate in extended parental care, since most of them left the females after copulation. Many females that were born in the summer produced one brood in the fall and a second during winter/early spring. Females that were born in the fall produced one brood during spring/early summer, but then probably died. Extended parental care in S. terebrans is short compared to other peracarid crustaceans. It is concluded that this reproductive strategy in S. terebrans serves primarily to shelter small juveniles immediately after they emerge from the female body, when their exoskeleton is still hardening and their physiological capabilities are still developing. Thus, in S. terebrans, extended parental care probably aids to protect small juveniles from adverse physical conditions in their subtropical intertidal habitat. Received: 9 December 1998 / Accepted: 24 June 1999     

Population biology of Dyopedos monacanthus (Crustacea: Amphipoda) on estuarine soft-bottoms: importance of extended parental care and pelagic movements

The suspension-feeding amphipod Dyopedos monacanthus (Metzger, 1875) is a common epibenthic amphipod that lives on self-constructed “mud whips” (built from filamentous algae, detritus and sediment particles) in estuaries of the northern North Atlantic Ocean. The population biology of D. monacanthus at a shallow subtidal site in the Damariscotta River Estuary (Maine, USA) was examined between July 1995 and July 1997. The resident population at the study site was dominated by adult females during most months of the year. High percentages of subadults were found in late summer/early fall. Often, between 10 and 20% of the adult females were paired with males, and the percentage of ovigerous females varied between 40 and 100%, indicating continuous reproduction. The percentage of parental females varied between 40 and 80% during most months, but dropped to levels below 20% during summer/early fall. The average size of amphipods on their own mud whips was ∼4 mm during the summer/early fall, after which it increased continuously to >7.0 mm in March or April, and then dropped again. In March and April, the average number of eggs and juveniles female⁻¹ was ∼100 eggs and 55 juveniles, while during the summer/early fall the average number of eggs female⁻¹ was <20 and that of juveniles female⁻¹ was <10. Many juveniles grew to large sizes (>1.4 mm) on their mothers' whips in winter/early spring but not in the summer/fall. The average number of amphipods at the study site was low in late summer/early fall (<50 individuals m⁻²), increased steadily during the winter, and reached peak densities of >3000 individuals m⁻² in April 1996 (>1600 individuals m⁻² in May 1997), after which densities decreased again. The decrease of the D.␣monacanthus population at the study site coincided with a strong increase of amphipods found pelagic in the water column. This behavioural shift occurred when temperatures increased and benthic predators became more abundant and active on shallow soft-bottoms, suggesting that D. monacanthus at the study site is strongly affected by predation. The effects are direct (by predation on amphipods) and indirect (by reducing duration of extended parental care and enhancing pelagic movements). Both extended parental care and pelagic movements are important behavioural traits of D.␣monacanthus (and other marine amphipods), and significantly affect its population dynamics. Received: 18 January 1998 / Accepted: 27 May 1998     

Ventilation or nest defense—parental care trade-offs in a fish with male care

Brood guarding animals face many critical trade-offs. Sand goby males (Pomatoschistus minutus) build nests with larger openings during low oxygen conditions, presumably to enhance ventilation. However, this may make the nest easier for egg predators to detect and harder for guarding males to defend. Manipulating oxygen level and predator presence (a small crab) for small and large males, we found support for a parental trade-off between fanning and nest defense. An increased fanning activity resulted in less time for guarding. Small males and males in low oxygen showed a higher fanning expenditure than large males and males in high oxygen, but surprisingly, filial cannibalism did not differ between these groups. Males built larger nest openings in low than high oxygen. However, males in both high and low oxygen treatments reduced their nest opening size in the presence of a predator, again indicating an important trade-off between ventilation and nest defense.     

Selective deposit-feeding by Lumbrineris cf. latreilli (Polychaeta: Lumbrineridae), with a new method for assessing selectivity by deposit-feeding organisms

The burrowing and feeding behaviour of an intertidal population of Lumbrineris cf. latreilli Audouin & Milne-Edwards, 1834 in Westernport Bay, Victoria, Australia was studied in 1982. Gut contents showed they were depositfeeders, ingesting predominantly sediment. Individuals were discretely motile, constructing semi-permanent burrows from which they fed upon subsurface deposits. Individuals in this population of L. cf. latreilli were selectively ingesting smaller particles. This is consistent with the predictions of optimal foraging theory modified for deposit-feeders.     

Feeding habits of a pelagic amphipod,Themisto japonica

Vertical distribution, chlorophylla (chla) and phaeopigment concentrations in the gut, and natural nitrogen isotope ratio ( ¹⁵N) were investigated for pelagic amphipodsThemisto japonica (Bovallius) collected from the Sea of Japan in July 1987. Differences in diel vertical migration behavior were clearly observed between small and largeT. japonica. Many small (<5 mm body length) amphipods appeared in the phytoplankton-rich shallow layers. Their gut pigment concentrations were higher (mean 0.52 ± 0.15µg chla g^–1 amphipod) than those of large amphipods (mean 0.33±0.14µg g^–1); this implies that the amphipods fed on a large amount of phytoplankton during the early stage of life. The ¹⁵N values of small amphipods were lower (5.7 to 6.3) than those of large amphipods (6.8 to 11.7), reflecting the lower trophic level of small amphipods compared to large ones. The ¹⁵N values for small amphipods were similar to those of herbivorous zooplankton. The amphipods' feeding behavior thus changes from herbivorous to carnivorous as they grow.     

Population and reproductive biology of two sibling amphipod species from ascidians and sponges

 The population dynamics and reproductive biology of an ascidian- and a sponge-dwelling amphipod were examined. The two undescribed amphipod species, Leucothoe“ascidicola” and L.“spongicola”, are closely related to each other, and occur in ascidians and sponges, respectively, along the Florida Atlantic coast. L. “ascidicola” was abundant in solitary ascidians during fall 1997, disappeared during spring/early summer, and became abundant again in September 1998. During the time when L. “ascidicola” were absent from their hosts, a copepod became a frequent inhabitant of the ascidians but disappeared again when L.“ascidicola” returned to the ascidians in September 1998. The numbers of L.“spongicola” in sponges increased substantially during spring, when high reproductive activity was observed. Following this reproductive peak, both adult and juvenile amphipods apparently left the sponges, and during the summer amphipod numbers in the sponges were very low. Another small amphipod species, which often co-occurred with L.“spongicola”, showed less seasonal variation and was found in sponges throughout the whole study period. The percentage of ovigerous females per host unit was usually lower in the ascidian-dwelling than in the sponge-dwelling amphipods. In solitary ascidians, L.“ascidicola” amphipods usually occurred in groups of several adults, yet there never was more than one ovigerous female per ascidian. In contrast, several ovigerous L.“spongicola” females were found to cohabit in the same spongocoel. This suggests that intrasexual aggression may be stronger among reproductive amphipod females in the ascidians than in sponges. The size distributions of juvenile cohorts indicate that juvenile L.“ascidicola” remain for relatively long time periods in the parental ascidian, where they may reach sexual maturity. In contrast, in L.“spongicola”, only cohorts of very small juveniles could be identified, indicating that juveniles disperse shortly after emerging from the female's brood pouch. It is concluded that extended parental care is of very short duration or does not occur in the sponge-dwelling amphipod L. “spongicola”, possibly because fast-growing sponges with a highly branched spongocoel system do not allow long-lasting coexistence of parent-offspring groups. In contrast, the discrete character of the solitary ascidians may enhance the potential for exclusion of other species, resource monopolization by reproductive females, and furthermore for long-lasting extended parental care in the ascidian-dwelling amphipod. Groups of single parents together with cohorts of large juveniles are reported in the literature for amphipods and isopods from brachiopods, bivalves and ascidians, suggesting that these discrete biotic microhabitats may favor the evolution of extended parental care in peracarid crustaceans. Received: 30 July 1999 / Accepted: 8 May 2000     

Reproductive biology of the planktonic copepod Temora stylifera

Laboratory studies on the reproductive cycle of the planktonic copepod Temora stylifera, collected from the Gulf of Naples between October 1987 and March 1988, showed that females oscillated between a light and dark gonadal condition. Histological preparations indicated that the dark condition corresponded to a ripe stage of oogenesis as compared to the light or unripe maturation state of primary oocytes. The number of eggs released within 24 h was strongly dependent on the reproductive status of the female at the time of incubation. Light unfed females rarely reverted to a dark or ripe condition and egg deposition in such cases was almost always zero. Dark females produced eggs within 24 h under all experimental food conditions, reverting from dark to light soon after egg deposition. Successively, the length of the interclutch period was strongly dependent on the type of food present. A high percentage of infertile eggs was almost always present. We suggest that short-term studies, not taking into consideration the condition of gonads at the time of incubation, will underestimate potential egg production rates. Females maintained for longer than three or four days produced mainly infertile eggs. After remating, females reinitiated the production of fertile eggs but egg viabilities never reached 100% hatching success, indicating poor efficiency in the fertilization of mature oocytes. We hypothesize a possible cause and effect relationship between egg viabilities and seasonal fluctuations in population densities.Died tragically at sea during a cruise     

Reproductive biology in two species of deep-sea squids

Deep-sea squids, Moroteuthis ingens and Gonatus antarcticus, were collected in the slope waters off the Falkland Islands and their reproductive systems preserved and investigated onshore. Changes in oocyte length-frequencies at maturation and spawning, and their fecundity were studied. These squids, as well as many other species, are characterised by a synchronous oocyte growth and ovulation. Oviducts are not used for ripe egg accumulation and consequently the universal scale of Lipinski (1979) cannot be applied to assign female maturity. M. ingens spawns near the bottom; its fecundity is 168–297 thousand eggs. Maximum egg size is 1.8–2.7 mm. G. antarcticus spawns midwater; its fecundity is 10–25 thousand eggs. Egg size is 3.2–3.3 mm. In M. ingens spawning takes place in the austral autumn and winter, in G. antarcticus—in austral winter. Our data and the literature data show that the so-called “synchronous ovulation” probably occurs in all deepwater squids. This pattern is very rare among fish, but is quite common among benthic octopods that brood their egg masses.     

Reproductive biology and ecology of a deep-demersal fish, alfonsino Beryx splendens, over the seamounts off New Caledonia

The reproductive biology of the alfonsino Beryx splendens was studied by histological examinations, gonadosomatic index and macroscopic scales of maturation of a large sample of gonads. Alfonsino is a gonochoric species. The size-frequency distribution of the sex ratio was bimodal and considered to be due to size dimorphism. In New Caledonia, the breeding period of this species occurs during the southern summer, with a peak in December to January. The spawning stage is attained at a minimum fork length of 28 cm for females and 33 cm for males. The size at which 50% of the population attain sexual maturity (FL₅₀) is 33.2 cm for females and 34.5 cm for males. Maximum potential fecundity is estimated to lie between 270 000 to 675 000 eggs for fish between 34 and 40 cm in fork length. It was possible to differentiate vegetative zones, in which juvenile alfonsino grow until they reach maturity, from reproductive zones (fishing grounds) which are inhabited by mature individuals. The larvae and juveniles could be carried from the reproductive zone to the vegetative zone by currents in an oceanic eddy system. Received: 26 April 1996 / Accepted: 20 September 1996     

Asymmetric sexual conflict over parental care in a biparental cichlid

Theoretical models predict that parents should adjust the amount of care both to their own and their partner’s body condition. In most biparental species, parental duties are switched repeatedly allowing for repeated mutual adjustment of the amount of care. In the mouthbrooding cichlid Eretmodus cyanostictus, terms are switched only once with females taking the first share. The timing of the shift of the clutch between mates strongly determines both partners’ brooding period and thereby their parental investment. Females signal their readiness to transfer the young several days before the male finally takes them, suggesting sexual conflict over the timing of the shift. In a lab experiment, we reduced the body condition of either the female or the male of a pair to test whether energy reserves affect the timing of the shift and whether female signalling behaviour depends on energetic state. Males with a lowered condition took the young later and incubated for a shorter period, which prolonged the incubation time of their female partners. When female condition was lowered, female and male incubation durations remained unchanged, although females signalled their readiness to shift more intensely. Our results suggest that males adjust their parental investment to own energy reserves but are unresponsive to their mate’s condition. Females appear to carry the entire costs for the male’s adjustment of care. We propose that intrinsic asymmetries in the scope for mutual adjustment of parental investment and the costs of negotiation crucially influence solutions of the conflict between sexes over care.     

Reproductive biology of three species of midwater fishes associated with the continental slope of eastern Tasmania,Australia

The reproductive biology of Lampanyctodes hectoris (Günther, 1876), Maurolicus muelleri (Gmelin, 1789) and Diaphus danae Tåning, 1932, from continental-slope waters of eastern Tasmania, was examined between April 1984 and June 1985. L. hectoris spawned in winter and M. muelleri spawned from late winter to early summer. Apart from one ripe male, no reproductive activity was detected in D. danae; this species may be an expatriate in these waters. Fecundity was positively correlated with standard length in L. hectoris, but not in M. muelleri. The ratio of females to males increased with length in all three species. The spermatozoa of L. hectoris and D. danae are atypical of vertebrates and have no tail.     

Manipulation of sex differences in parental care

Summary In a species with biparental care two parents cooperate to provide the appropriate amount of care for the young. Recent theoretical treatments consider the evolutionarily stable investment strategy. Under most conditions, the parental investment of the two partners should be negatively correlated, with the shortfall of one partner being partially compensated for by the other. Previous experimental manipulations of biparental care have involved removal of one partner, yet the response of a widowed bird may differ from that of a mated bird whose partner is doing less than its fair share of parental care. We present the first data involving subtle manipulations of sex differences in parental care where both partners continue to care for the young. This study involves pairs in a nestbox colony of european starlings (Sturnus vulgaris L.) with all brood sizes manipulated to five chicks. Pairs were randomly assigned to three groups: (i) male parental care reduced; (ii) female parental care reduce; and (iii) control pairs. Parental care was manipulated by attaching small weights to the base of a bird's tail feathers. Regardless of sex, nest visitation rate was reduced in the weighted birds with an incomplete compensatory increase by their unweighted partner. Additional parental duties were also considered, including shifts in prey type delivered to the nest, in both weighted birds and their partners. The shift in diet and the overall lower total visitation rate in experimental nests contributed to slower chick growth and lower chick weights than in control nests. The data accord with models suggesting that equality of invesment in biparental species is evolutionarily stable, but reveal new dimensions of parental response that need to be taken into account in theoretical treatments.     

Endocrine influences on parental care during a short breeding season: testosterone and male parental care in Lapland longspurs (Calcarius lapponicus)

In males of socially monogamous birds, plasma testosterone (T) typically declines to low levels during the parental phase. Studies on multiple-brooded species indicate that high T may be incompatible with high-quality paternal care. The length of the breeding season may affect the costs and benefits of high T and its effect on paternal care. We studied the effect of experimentally elevated T on paternal care in a single-brooded species with a short breeding season, the Lapland longspur (Calcarius lapponicus). We monitored T levels and parental behavior in 16 males with subcutaneous T implants, 14 males with empty implants, and 14 unimplanted males. We videotaped nests when nestlings were 2–3 days old and again at 4–5 days. T males with 2- to 3-day-old young visited nests and fed young less often than control males, and the mates of the T males compensated with elevated visits and feedings. However, when nestlings were 4–5 days old, T males visited their nests at normal rates – though feeding movements remained below normal – and T females visited and fed at normal rates. Nestling mass and nest success were similar in both groups. Overall, high T suppresses paternal care in Lapland longspur males. The partial improvement of paternal care when nestlings are older, despite high T, may be related to the short 6-week breeding season of this arctic species, and the consequently reduced benefits of sexual behavior late in the breeding season. Received: 2 February 1998 / Accepted after revision: 2 November 1998     

Reproductive biology of a deep-sea brittle star Amphiura carchara (Echinodermata: Ophiuroidea)

 New information on the brooding reproduction of Amphiura carchara, and previous studies of several deep-sea congeners, contradict the prevailing notion that direct development is exceptional among deep-sea echinoderms. Over 500 specimens of A. carchara from 2,850 to 4,100-m depths off the coast of California, USA, were studied. The species was found to be gonochoric, although most brooding species of ophiuroids are hermaphroditic. Females each brooded up to 72 embryos at a time, with up to 10 in a single bursa. The embryos of individual adults were often at different stages of development, but those brooded in a single bursa tended to be at the same stage. Thus, the species has characteristics that breach the distinctions between sequential and simultaneous brooding exhibited by shallow-water ophiuroids. The embryos developed from yolky eggs that are large for an ophiuroid, with the mean largest oocytes 0.45 mm, and ranging to 1.28 mm in diameter. Almost all late-stage embryos were positioned with their mouth and arms pressed against the wall of the bursa, possibly to facilitate the uptake of nutrients from the parent. Thus, A. carchara may be matrotrophic. The largest embryos examined had a rudimentary disk skeleton, and arms with four joints and a terminal plate. Emerging juveniles probably differ in disk diameter. The species appears to brood year round, although differences in gonad size, the incidence of brooding, and the relative numbers of early developmental stages in summer and winter samples indicated that there are seasonal trends in reproduction. Received: 14 February 2000 / Accepted: 3 July 2000     

Reproductive biology of deep-water calanoid copepods from the Arctic Ocean

Reproductive modes and egg production were studied in 15 species of meso- and bathypelagic copepods from nine Calanoida families in the Arctic Ocean. During shipboard incubation, females of seven species released eggs freely into the water and females of three species produced membrane-bound egg sacs. One species, Aetideopsis rostrata, produced a mass of eggs looking like an egg sac, but the “sac” lacked a membrane and disintegrated within 2 h. Females of four additional species were encountered with membrane-bound egg sacs in the preserved samples. In most families, only one reproductive mode, either egg-carrying (the Euchaetidae, Augaptilidae), or broadcast spawning (the Heterorhabdidae, Spinocalanidae, Scolecitrichidae, Tharybidae, Bathypontiidae) was observed. In contrast, different genera of the Aetideidae family demonstrated different reproductive modes, with broadcast spawning predominant in the benthopelagic species, and both broadcast spawning and egg-brooding in the planktonic species. Clutch size and egg diameter varied widely between species in both broadcast spawners and egg-brooders. In broadcast spawners, the clutch size varied from 1 to 95 eggs female⁻¹, while the average egg diameter ranged from 152 to 440 μm. The clutch size for egg brooders varied between 3 and 82, while average egg diameter varied from 258 to 732 μm. Deep-water broadcast spawners produced much larger eggs compared to surface-dwelling broadcast spawning species. This larger egg size may result in a reduction, or elimination, of feeding during naupliar stages, thereby improving the survival potential of deep-water species.     

Reproductive biology of the antipatharian black coral Antipathes fiordensis in Doubtful Sound, Fiordland, New Zealand

The reproductive biology of Antipathes fiordensis Grange, a species endemic to south-western New Zealand, was followed from April 1994 to May 1995. Ten colonies were individually tagged in Doubtful Sound and sampled on a monthly basis in order to determine their reproductive activity. The fecundity of each of the five tagged female colonies was determined by estimating the total number of polyps per colony from photographs of each colony and by planimetry, the proportion of gravid polyps per colony, and the mean number of oocytes per gravid polyp. In addition, 56 colonies were sampled in March 1995 to estimate the sex ratio, height at sexual maturity, and mean sizes of females and males. A. fiordensis was found to be a dioecious species which, in the absence of gonads in the polyps, has no obvious external morphological differences between the sexes. Broadcast spawning of gametes is the likely mode of reproduction. Gametogenesis began in November 1994 and was highly synchronous within and between colonies, with spawning occurring in March 1995. The sex ratio in adults was 1:1. Colonies reached sexual maturity between the heights of 70 and 105 cm which, based on existing estimates of growth rate, corresponds to a minimum age for sexual maturity of about 31 yr. The largest oocytes measured ranged from 100 to 140 μm in size. Female colonies produced between 1.3 and 16.9 million oocytes, with the larger colonies dominating the reproductive output of the population. Received: 23 June 1997 / Accepted: 1 August 1997     

Divergence between populations of a monogamous polychaete with male parental care: Premating isolation and chromosome variation

Low dispersal and sexual selection are characteristic of the coastal polychaeteNereis acuminata Ehlers 1868 [also known asNereis arenaceodentata Moore 1903 andNereis (Neanthes) caudata Delle Chiaje 1841]. We assessed levels of premating isolation between populations of this polychaete. Four North American populations were used, two from the Atlantic and two from the Pacific. Worms from all sites (1) were collected in 1987 and 1988 from the same habitat type, (2) were morphologically similar and keyed out asN. acuminata, and (3) reproduced monogamously and exhibited male parental care, an extremely rare reproductive mode in marine invertebrates. There was no evidence from 10-min or 36-h trials of premating isolation between the two Pacific populations. Incomplete premating isolation was found between the two Atlantic populations. High aggression and non-pairing occurred in some 10-min trials between males and females. However, in 36-h trials males and females from the two Atlantic populations always paired to mate. Complete premating isolation was found between Atlantic and Pacific populations. During 10-min trials, males and females from different oceans often attacked and then avoided each other, and they never paired to mate. Nor did they pair to mate in longer, 36-h trials. One Pacific and one Atlantic population were compared for tolerance to cold temperature. Pacific individuals were less able to tolerate cold water than Atlantic individuals. Two Atlantic populations studied had karyotypes with 11 pairs of small acrocentric chromosomes (2n=22), while the two Pacific populations had nine pairs of large metacentric or submetacentric chromosomes (2n=18). Such extreme dissimilarity in karyotype was not expected considering the similarity in morphology, habitat, and reproductive mode. Results suggest strongly that the Atlantic and Pacific populations have been allopatric for a long time, and are different species.     

The costs of male parental care and its evolution in a neotropical frog

Summary Parental care is practiced exclusively by males of the Puerto Rican frog, Eleutherodactylus coqui. Males brood clutches of direct-developing eggs in non-aquatic nest sites and defend eggs against cannibalistic nest intruders. Here, I report on energetic and mating costs incurred by males that provide parental care, and suggest how these proximate costs affect male fitness and the evolution of male parental care in this species. Energetic costs are small for brooding males in comparison to non-brooding, calling males. Brooding males had a higher frequency of empty stomachs and lost small, but significant, fractions of their initial body mass during parental care. Abdominal fat bodies of brooding males during the middle third of parental care were significantly smaller than those of calling males; those of males brooding eggs in earlier or later stages were not different. The mating cost of parental care is greater. Most brooding males cease calling during parental care. However, gravid females are available (i.e., known to mate) on most nights during the principal breeding season; hence non-calling males miss potential opportunities to mate. A mating cost was estimated by calculating nightly mating probabilities for calling males in a plot where nightly calling male densities and daily oviposition schedules were known. On average, a male exhibiting normal calling behavior would be expected to obtain a new mate once every 35.7 days. Hence a brooding male that ceased calling for a 20-day parental care period would miss, on average, 0.56 additional mates. Males that were more successful than average in attracting mates could miss up to 1.63 matings. A marginal value model (Fig. 1) is used to analyze the net effect on male fitness of parental care benefits and costs in E. coqui (Fig. 3). The model indicates that males garner the highest reproductive success by providing care from oviposition through hatching. There is no stage during the pre-hatching period at which a desertion strategy would yield higher reproductive success. In fact, the model suggests that males should provide full parental care even in the face of much higher mating costs than currently obtain in the system.