Density affects mating mode and large male mating advantage in a fiddler crab

Fiddler crabs show two different mating modes: either females search and crabs mate underground in male burrows, or males search and crabs mate on the surface near female burrows. We explored the relationship between crab density, body size, the searching behavior of both sexes, and the occurrence of both mating modes in the fiddler crab Uca uruguayensis. We found that crabs change their mating mode depending on their size and crab density. Crabs mated mostly on the surface at low densities, and underground at high densities. The proportion of wandering receptive females but not courting males accounted for the variation in mating modes. This suggests that whether crabs mate underground (or on the surface) is determined by the presence (or absence) of searching females. We found that the change in the mating mode affected the level of assortative mating; males mating underground were bigger than those mating on the surface, suggesting active female choice. Given that fiddler crabs experience multiple reproductive cycles, they are prone to showing behavioral plasticity in their mating strategy whenever the payoffs of using different mating modes differ between reproductive events. Our results suggest that the incorporation of different levels of environmental variability may be important in theoretical models aimed at improving our understanding of the evolution of alternative mating tactics and strategies.     

Size-dependent mating preference of the male fiddler crab Austruca perplexa

In many animals, females prefer large males to small males, which allow large males to be choosier than small males when selecting a mate. We investigated the courtship intensity of small- and large-sized male fiddler crabs (Austruca perplexa) by examining their claw-waving rates (waves/min) towards small- and large-sized females. We found that large males showed a greater preference for large females by producing more waves/min towards them, whereas small males did not show any apparent preference for either large or small females. Moreover, the waving rate of large males was positively correlated with female size, but there was no correlation between waving rate and female size in small males. These results indicate that large males in a population become choosier and show strong mate choice, which is most likely due to their greater preference among females.     

Melatonin and locomotor activity in the fiddler crab Uca pugilator

The influence of melatonin on locomotor activity levels was measured in the fiddler crab Uca pugilator. First, activity in untreated, laboratory-acclimated crabs was measured over 48 hours in a 12L:12D photoperiod; this study showed a nocturnal increase in activity. In eyestalk-ablated crabs, overall activity was significantly reduced, and no significant activity pattern occurred. Next, crabs were injected with melatonin or saline (controls) at various times during the 12L:12D photoperiod (0900h, 1200h, and twice at 2100h; each trial was separated by 3-4 days) and monitored for 3 hr post-injection. Control crabs had low activity during early photophase, high at mid-photophase, increasing activity during the first scotophase trial, and decreasing activity during the second scotophase trial. Melatonin had no significant influence on activity when injected during the early-photophase activity trough or early-scotophase activity decline, but significantly increased activity when injected during the mid-photophase activity peak and early-scotophase activity incline. Next, crabs were injected during an early scotophase activity trough and monitored throughout the twelve-hour scotophase. Melatonin did not increase activity until the mid-scotophase activity increase, approximately 6 hours later, showing that the pharmacological dosage persisted in the crabs' systems and had later effects during the incline and peak of activity but not the trough. Eyestalk-ablated crabs were injected with melatonin or saline during early photo- and scotophase. Melatonin significantly increased activity in the photophase but not the scotophase trial, indicating that the responsiveness to melatonin continues following eyestalk removal, but the timing may not match that of intact crabs. Melatonin may be involved in the transmission of environmental timing information from the eyestalks to locomotor centers in U. pugilator.     

三突伊氏蛛雌蛛不同交配史对雄蛛交配行为的影响

近年来越来越多研究表明,雄性产生精子(精液)也需付出代价。在多次交配的动物中,雄性为获得最大生殖潜力,必须依据配偶的质量策略性地调整每次交配的生殖投入。雄性策略性的生殖投入主要体现在两个方面,一是精子竞争(sperm competition),二是柯立芝效应(Coolidge effect)。目前精子竞争研究主要集中于昆虫类群,而柯立芝效应研究集中于高等脊椎动物,同时验证结果也时常与假说不一致。以多次交配的三突伊氏蛛为材料,以雄蛛交配行为为指标,在蜘蛛类群中探讨和验证雄性精子竞争强度假说和柯立芝效应。设定3个交配组合:2只雄蛛依次与1只雌蛛各交配1次(A组)、2只雄蛛依次与2只雌蛛各交配1次(B组)和1只雄蛛与1只雌蛛交配2次(C组),分析比较3个交配组合的三突伊氏蛛第1次交配和第2次交配在交配潜伏期、交配持续时间和交配回合数方面的差异,比较三突伊氏蛛雌蛛不同交配史对雄蛛交配行为的影响,以此验证雄性精子竞争强度假说和柯立芝效应。研究结果表明A和B组的三突伊氏蛛第2次交配的交配潜伏期和交配持续时间显著长于第1次交配。同时,C组的三突伊氏蛛第1次交配的交配潜伏期和交配持续时间与第2次交配都没有显著差异。同时,A、B和C组的三突伊氏蛛第1次交配的交配回合数与第2次交配都没有显著差异。研究结果支持精子竞争强度假说,而不支持柯立芝效应。     

Visual shape discrimination in the fiddler crab,Uca pugilator

1. Research into the visual shape discrimination abilities of compound‐eyed animals has almost exclusively been limited to insects, the crustaceans having been virtually ignored. The two groups have many dissimilarities, having primarily adapted in different habitats to different lifestyles. Differences may exist in visual systems and visually mediated behavior.

2. Fiddler crabs (Uca pugilator), without training, differentially approached dissimilar silhouettes presented simultaneously, demonstrating visual discrimination between stationary, geometric shapes of equal‐area. The strength of response was ordered hierarchically: vertical rectangle, horizontal rectangle, triangle, square, circle.

3. Basic geometric shapes were used to facilitate replication and comparison with research findings from other species.

     

Plasticity in male mating behavior modulates female life history in fruit flies

In many species, intense male-male competition for the opportunity to sire offspring has led to the evolution of selfish reproductive traits that are harmful to the females they mate with. In the fruit fly, Drosophila melanogaster, males modulate their reproductive behavior based on the perceived intensity of competition in their premating environment. Specifically, males housed with other males subsequently transfer a larger ejaculate during a longer mating compared to males housed alone. Although the potential fitness benefits to males from such plasticity are clear, its effects on females are mostly unknown. Hence, we tested the long-term consequences to females from mating with males with distinct social experiences. First, we verified that competitive experience influences male mating behavior and found that males housed with rivals subsequently have shorter mating latencies and longer mating durations. Then, we exposed females every other day for 20 days to males that were either housed alone or with rivals, and subsequently measured their fitness. We found that females mated to males housed with rivals produce more offspring early in life but fewer offspring later in life and have shorter lifespans but similar intrinsic population growth rates. These results indicate that plasticity in male mating behavior can influence female life histories by altering females’ relative allocation to early versus late investment in reproduction and survival.     

An evaluation of lekking behavior in the fiddler crab Uca spp

The reproductive behavior of Uca spp. has been extensively studied, especially the relationships between the dimorphically enlarged male claw and reproductive success. In contrast, little is known about the apparent congregations of adult males in marsh areas lacking vegetative cover where they engage in behaviors thought to attract mates. Similar congregations and displays in avian and mammalian species are termed 'leks.' In order to test the hypothesis that open-area assemblages of Uca spp. are functioning as leks, we examined the sex ratios, juvenile/adult ratios, and the percentage of time that adult males spend eliciting reproductive behaviors within open and vegetated areas of marsh habitats. Moreover, to evaluate whether differences in sediment-based food resources could explain open-area aggregations, we compared substratum organic content of open and vegetated areas of marsh habitats. Substrate was also examined to determine if grain size composition varied between open and vegetated areas and might preclude the construction of breeding burrows in vegetated areas of the marsh. Three species of Uca from four marsh habitats in biogeographically distinct regions of North America were sampled including Dauphin Island, Alabama, Hunting Island, South Carolina, Saxis, Virginia, and Wallops Island, Virginia. Comparisons of male/female and juvenile/adult ratio means indicated that greater numbers of adult males occurred in open areas of all marshes. In addition, adult males allocated significantly greater time to reproductive behaviors in open rather than in vegetatively covered areas across-all biogeographic regions and among all species. Food levels (sediment organic content) in open areas were equal to or less than sediment organic contents in vegetated areas in marsh habitats at Dauphin Island and Hunting Island, the two marshes where the variable was examined. Similarly, substratum granulometry analysis revealed no significant differences between open and vegetatively covered areas of the marshes at Dauphin Island or Hunting Island that might influence choice of burrow location. Collectively, these observations support the hypothesis that lek behavior is an integral component of the reproductive repertoire in Uca spp.     

RXR isoforms and endogenous retinoids in the fiddler crab, Uca pugilator

The pleiotropic effects of circulating ecdysteroids in the adult fiddler crab, Uca pugilator, during molting, regeneration, and reproduction are mediated by a limited number of receptor proteins. We hypothesize that hormonal effects in vivo may be the result of complex interactions between at least two receptor heterodimer conformations that differentially respond to multiple ecdysteroid/retinoid signals. Two splicing variants of the fiddler crab retinoid-X-receptor (UpRXR) differ from one another by the addition of a 33 amino acid insert in the ligand-binding domain. We show here that the ecdysteroid receptor in the fiddler crab (UpEcR) behaves differently depending upon the UpRXR isoform with which it is partnered. The two UpRXR variant partners for UpEcR confer slightly different responses in the binding of Ponasterone A (PA)—a naturally occurring ecdysteroid in the blood of Uca. UpRXR can bind 9-cis retinoic acid (9cRA) as well as terpenoids. 9cRA and the naturally occurring terpenoid, methyl farnesoate, influence the binding of PA to UpEcR/UpRXR dimers. Endogenous retinoids are found in the blastema of regenerating limbs of Uca and they (plus blood-borne terpenoids) may add additional levels of differential response by target tissues. Thus, the two sets of heterodimers tested here may represent different dynamic complexes whose properties are defined by the specific heterodimeric subunits involved and the specific ligands available.     

Neurosecretion in the sinus gland of the fiddler crab,Uca pugnax

The ultrastructure of the sinus gland of the fiddler crab, Uca pugnax, was investigated and found to be similar to that in other crustaceans. Five types of neurosecretory axon terminals were tentatively identified on the basis of the size, shape, and electron density of granules within the axons. Release of neuro-secretory material appears to be by exocytosis.     

Residency and size affect fight duration and outcome in the fiddler crab Uca annulipes

We performed a field experiment to investigate the effect of carapace width, major cheliped length and burrow ownership on the righting success of male fiddler crabs (Uca annulipes) . We removed males from their burrows and released them back into the colony ( n = 82). Released males tended to initiate encounters with burrow owners slightly smaller than themselves. Several general predictions of Sequential Assessment Game models of contest behaviour were supported: (1) residents won more encounters; (2) intruders were more likely to win when larger than residents. When body size (carapace width) was controlled for, intruders with relatively large claws for their body size were more likely to win contests; (3) the duration of encounters was related to the size difference between males; (4) encounters won by the larger male were of shorter duration than those won by the smaller male; (5) encounters won by the resident tended to be of shorter duration than those won by intruders ( P = 0.07); (6) on average, encounter duration was longer when the intruder was larger than the resident. However, the encounters we documented began with seemingly costly behaviour such as pushing and the inter-locking of claws and did not unambiguously escalate from initial low cost behaviours. Sequential assessment of relative fighting ability may therefore not have been occurring. Prior visual assessment of opponents' fighting ability, followed by 'all-out fights' during physical encounters may also provide a plausible explanation for our results.     

Behavioral patterns of two fiddler crab species Uca rapax and Uca tangeri in a seminatural mangrove system

Zoos increasingly transform their exhibitions from traditional one‐species enclosures to more natural exhibits, that is, environments that capture part of an ecosystem including a selection of animals and plants that occur there. Thus, enhancing the experience of its human visitors while also allowing its residents to possibly show more natural behavior. In 2017 Royal Burger's Zoo (Arnhem, The Netherlands) created and opened a mangrove‐like environment containing fiddler crabs. Fiddler crabs display a broad range of behaviors, and this research examines which wild‐type behavior and behavioral patterns can be observed on a seminatural mudflat. The behavior shown by Uca rapax and Uca tangeri on the mudflat was counted each hour between 07:00 and 17:00. An asymmetric tidal regime was present in the enclosure including two high water periods. Various known fiddler crab behaviors, including waving and combat, were observed but no copulation. A clear pattern in exposed crabs on the mudflat was found, with low numbers visible in the early morning and the highest numbers present in the early afternoon, while number of visitors did not have a significant effect on this pattern. Interestingly, the highest abundances were not observed around the ebbing tide (07:00–09:00), as observed in the wild, but somewhat later, possibly due to the asymmetric tidal scheme or the interaction of tidal and daily rhythms. This study shows that in captivity, fiddler crabs indeed show a range of natural behaviors which is linked to the tidal and possibly daily rhythm as well.     

Colouration and colour changes of the fiddler crab, Uca capricornis: a descriptive study

Colour changes in animals may be triggered by a variety of social and environmental factors and may occur over a matter of seconds or months. Crustaceans, like fiddler crabs (genus Uca), are particularly adept at changing their colour and have been the focus of numerous studies. However, few of these studies have attempted to quantitatively describe the individual variation in colour and pattern or their adaptive significance. This paper quantitatively describes the colour patterns of the fiddler crab Uca capricornis and their ability to change on a socially significant timescale. The most dramatic changes in colour pattern are associated with moulting. These ontogenetic changes result in a general reduction of the colour pattern with increasing size, although females are more colourful and variable than similarly-sized males. Uca capricornis are also capable of rapid colour changes in response to stress, but show no endogenous rhythms associated with the semilunar and tidal cycles commonly reported in other fiddler crabs. The extreme colour polymorphism and the relative stability of the colour patterns in Uca capricornis are consistent with their use in visually mediated mate recognition.     

Eye and clasper damage influence male mating tactics in the horseshoe crab, Limulus polyphemus

In the horseshoe crab mating system, mated pairs are frequently accompanied by unattached satellite males as they spawn on intertidal beaches. Previous studies have shown that males locate females visually using their lateral (compound) eyes, and that attached (mated) males generally have less heavily worn or damaged carapaces than unattached males. The purpose of this study was to investigate the influences of lateral eye condition and clasper abnormalities on male mating tactics. Sexually mature males had two kinds of eye damage: deterioration caused by disease, and overgrowth by sessile invertebrates, such as bryozoans, mussels, and tube-building polychaetes. The lateral eyes of attached males had significantly less decay than unattached males. On the other hand, coverage of the lateral eyes by encrusting invertebrates was more extensive among attached than unattached males. Although overgrowth did not appear to impair a males ability to pair with a female as severely as eye decay, it is conceivable that amplexus may have occurred before epibiont coverage was sufficient to obscure vision. Male crabs that were experimentally blindfolded by painting their lateral eyes with black nail polish were less likely to reattach to a female than controls. Appendage injuries were more frequent among unattached males than among attached males; in particular, 6.4% of unattached males but 0.0% of attached males had damaged claspers (the modified first legs required for amplexus). Unattached males in the population were older, as judged by the degree of carapace wear, than attached males. Severe visual impairment and/or clasper damage may explain the reduced pairing success of older male horseshoe crabs, and underlie their choice of the alternative satellite male mating tactics.