The burrows of alpheid shrimp associated with gobiid fish in the northern Red Sea

The burrows of 4 sympatric associations of alpheid shrimp (Alpheus djiboutensis 1 and 2, and Alpheus sp. A and B) and non-burrowing gobiid fish were studied in the lagoon of a narrow fringing reef, in the northern Red Sea. Epoxy resin casts were used to determine the subterranean structure of burrows. The burrows were irregular, and in close contact with hard objects within the substrate. The number and structure of openings and their daily displacement were type-specific. On the other hand, no type specificity was observed in the deeper parts of the burrow. The mechanism of the formation of new openings by the combined activities of the fish Cryptocentrus sungami and the shrimp A. djiboutensis 2 is described. A positive correlation in the size of the partners seems to be due to competition of fish for larger burrows.     

Observations on the burrows and burrowing behaviour of two mud-dwelling decapod crustaceans,Nephrops norvegicus and Goneplax rhomboides

The burrows of the Norway lobster Nephrops norvegicus (L.) and of the crab Goneplax rhomboides (L.) were studied in Loch Torridon, Scotland. Polyester resin casts of burrows in the sea were made by divers to reveal their subsurface form. Tunnels made by N. norvegicus were usually simple, with two or more openings on the mud surface, and penetrated to a depth of about 30 cm. G. rhomboides burrows did not descend more than about 15 cm beneath the surface, but were usually more complex than the lobster burrows and had several openings. The methods of burrow construction used by the two crustaceans are described from aquarium observations. Neither N. norvegicus nor G. rhomboides show obvious morphological adaptations for burrowing, and it is suggested that the fossorial habit was adopted very early by decapods. The burrows of N. norvegicus do not seem to have assumed any functions in addition to the original one of providing refuge from predators. There is not sufficient known of the biology of the crab to indicate whether the same is true in its case.     

Functional burrow morphology of Biffarius arenosus (Decapoda: Callianassidae) from southern Australia

Burrows of the thalassinidean shrimp Biffarius arenosus (Poore, 1975) were investigated by in situ resin-casting on an intertidal sandflat in Western Port, southern Australia. Even though burrow casts exhibited interspecific variation, all had at least two openings, a U-shaped top and a series of tunnels and chambers. The space occupied by the burrows was deeper than it was wide; a maximal depth of 58 cm was recorded. Inhalant and exhalant shafts were restricted relative to tunnel diameter, and were often arranged in a multi-layered U-shape. Additional shafts branching from deeper sections of the burrows were occasionally observed. The main section of a burrow, extending from the base of the U, usually consisted of a downward spiral or an irregular spiral combined with some straight tunnel sections, all with a circular cross-section. The central spiral branched into further tunnels and chambers. Nearly all casts possessed peripheral chambers positioned either at the edges of horizontal lattices or in the deepest sections of the burrow. No evidence of plant material was found in peripheral chambers. Major burrow features were consistent with B. arenosus adopting a deposit-feeding trophic mode and collecting its food below the sediment surface. Received: 17 July 1998 / Accepted: 26 November 1998     

Mesocosm and in situ observations of the burrowing shrimp Calocaris templemani (Decapoda: Thalassinidea) and its bioturbation activities in soft sediments of the Laurentian Trough

In vivo observations in laboratory mesocosms and aquaria, accompanied with in situ photographic surveys, have shown that the burrowing shrimp Calocaris templemani has a significant impact on bottom sediment dynamics and geochemistry in the St. Lawrence Estuary. This burrowing shrimp establishes and maintains complex semi-permanent burrows made up of several interconnected, ‘U-shaped’ galleries with generally four or more openings to the sediment surface. In the Estuary, at 345 m depth, Calocaris average density was estimated at 3.4 individuals m^?2. Observed individual burrows reached a maximum volume of 0.54 L. C. templemani displaces this volume of mostly anoxic sediments from the subsurface layers (down to 15 cm) to the sediment surface, thereby obscuring some of the natural stratification patterns. With an estimated turnover rate of about 8 L m^?2 year^?1 of sediment, our calculations suggest that over a period of about 18.75 years, all the sediment to a depth of 15 cm will have been reworked by C. templemani alone.     

Geochemistry of burrow waters vented by a bioturbating shrimp in Bermudian sediments

Waters collected at various times from the burrows of Callianassa spp. shrimp are characterized chemically by concentrations of nutrients, sulfide and organic carbon, intermediate between those of pore waters and overlying waters of Coot Pond, Bermuda. Although the shrimp vigorously flush their burrows, irrigation events are insufficently frequent to maintain a burrow water composition completely like that of oxygenated overlying waters; in instances where departure is extensive, models describing the diffusion of interstitial-water solutes into the water column via burrows would have to take this into account. Measured oxidation potentials are substantially reduced with respect to those of the water column, and relatively high levels of reduced arsenic (As III) are encountered. Burrow irrigation provides a mechanism for the rapid, conservative advection of solutes across the sediment-water interface and therefore is at least partially responsible for the concentration and speciation variations typical of coastal seawater. Of the solutes tested, only total dissolved inorganic arsenic occurs at seawater concentration in the burrows, implying that some specialized mechanism intercepts the diffusion of pore water arsenic into the burrows.     

Males are attracted by their own courtship signals

Courting male fiddler crabs Uca terpsichores (1 cm carapace width) sometimes build mounds of sand called hoods at the entrances to their burrows. Males wave their single enlarged claws to attract females to their burrows for mating. It was shown previously that burrows with hoods are more attractive to females and that females visually orient to these structures. In this study, we test whether males also use their hoods to find their burrows. We first determined the maximum distance that males can see and find a burrow opening without a hood. Males were removed from their burrows and placed on the sand at a range of distances from a burrow opening. If they were more than about 8 cm (seven units of eye-height) away, they were unable find the burrow. In contrast, males that were burrow residents used a non-visual path map to return to their burrows from much greater distances. To determine if hoods help males find their burrows when there are errors in their path maps, we moved residents 1–49 cm on sliding platforms producing errors equal to the distances they were moved. Males with self-made hoods or hood models at their burrows relocated their burrows at significantly greater distances than did males with unadorned burrows. Hood builders also relocated their burrows faster. Hence, hoods have two functions: they attract females and they provide a visual cue that males use to find their burrows quickly and reliably when their path maps fail. An erratum to this article can be found at     

The benefit of obligate versus facultative strategies in a shrimp–goby mutualism

Obligate mutualists are predicted to benefit more from mutualism than facultative mutualists and harbor traits that help derive this extra benefit. I tested these predictions with a shrimp–goby mutualism. Individual shrimp (Alpheus floridanus) construct burrows that are shared with individual gobiid fishes that warn shrimp when emergence from burrows is unsafe. The benefit to gobies is refuge from predation. I compared predator avoidance performance of obligate (Nes longus) and facultative (Ctenogobius saepepallens) shrimp-associated gobies with access to a shell, a shrimp burrow, or no shelter. The two gobies performed similarly with shells and no shelter, but N. longus outperformed C. saepepallens with shrimp burrows. Thus, N. longus benefits more from mutualism than C. saepepallens. Also, N. longus has four behavioral traits that likely allow it to benefit from mutualism more than does C. saepepallens: (1) diving into the nearest shrimp burrow when confronted with predators, (2) long flight initiation distance, (3) long time-until-re-emergence after taking refuge, and (4) remaining close to shelter. The latter three likely come with a cost to foraging or mate acquisition as they limit searching range outside the burrow and detract from time spent outside burrows. Thus, N. longus likely harbors traits that allow it to deal with these costs. I discuss a framework for determining whether traits crucial to obligate association with shrimp (both for reaping benefits and dealing with costs) are pre-adaptations or are created through coevolution with shrimp.     

Mating behaviour of Macrophthalmus hirtipes (Brachyura: Ocypodidae)

 We examined the mating behaviour of the New Zealand ocypodid crab Macrophthalmus hirtipes in the laboratory between February and June 1998. This species has a discrete breeding season. Mating and moulting were not linked and only intermoult females with mobile gonopore opercula were attractive to males. Allometry and compatibility of gonopods and gonopores of different-sized crabs was investigated. Under laboratory conditions, the opercula of intermoult females remained mobile on average for 11.4 d, but the duration of receptivity did not appear to be under female control. The operational sex ratio in the laboratory fluctuated greatly, but was always male-dominated. During the period of opercular mobility, females mated many times with several different males. Matings in the absence of burrows were relatively short (mean duration = 23 min, max. = 122 min) and the mating behaviour of M. hirtipes lacked courtship and mate-guarding. Males used a search-intercept method to acquire mates, with very low levels of intrasexual competition. There was no evidence of mate preference in M. hirtipes, and males spent just as long mating with ovigerous females as with non-ovigerous ones. Although M. hirtipes has ventral-type spermathecae, as do several other ocypodid crabs, it is unclear whether this promotes last-male sperm precedence. The role of burrows in modifying the mating behaviour of M. hirtipes in the field remains to be established. Received: 7 January 2000 / Accepted: 5 June 2000     

An unusual abbreviated larval life in the estuarine burrowing prawn Callianassa kraussi (Crustacea: Decapoda: Thalassinidea)

The larval stages of Callianassa kraussi Stebb. were previously unknown. Larvae were bred out in the laboratory and found to be incapable of swimming. There are two larval stages, lasting 3 to 5 days in sea water of 34 to 35 S at 20 °C. Observations under simulated field conditions in the laboratory, and in the field—where burrow casts were made using a polyester resin, indicated that the larvae metamorphose to the first post-larval stage in the parent burrow, and then burrow directly into the wall of the parent burrow.     

Diurnal variations in the behaviour of the hagfish Eptatretus burgeri

The Japanese hagfish Eptatretus burgeri may be found in relatively shallow water, and is thus accessible to field observations by SCUBA diving. A population of hagfish living in a bay at a depth of about 10 m in Koajiro Bay, Kanagawa Prefecture, Japan has been observed periodically throughout a year by day and night diving. During the day, with rare exceptions, the hagfish remains within its burrow in the soft, muddy substrate; its anterior tip is sometimes just visible within the single opening of the burrow. There are no heaps of mud associated with the burrows, which seem to be relatively impermanent structures. During the night, many hagfish emerge from their burrows, either partially or completely. Partially emerged individuals protrude as much as a third of their length beyond the burrow opening, while completely emerged individuals either rest uncovered on the substrate surface or swim actively near the bottom. An emerged hagfish prefers to reenter a preexisting burrow; when one is not available, however, it quickly excavates a new burrow by swimming vigorously down into the mud. No hagfish were found in the investigated locality from the middle of June to the middle of October.     

Mating system of the burrowing crab Neohelice granulata (Brachyura: Varunidae) in two contrasting environments: effect of burrow architecture

Few studies conducted in crustaceans have demonstrated how habitat features could shape the mating systems. Here, the burrow of Neohelice granulata was considered as a resource used for mating, and its architecture was characterized in two contrasting study sites: Mar Chiquita Lagoon (MCL), an estuary composed of muddy sediments, and San Antonio Oeste (SAO), a marine bay composed of gravel sediment. Burrow features differed between study sites and occupant gender. Large males constructed burrows with a chamber in MCL and with a widened entrance in SAO, while small males constructed the same narrow burrows as females at both study sites. Field experiments demonstrated that burrows with chambers or widened entries are places used for copulation, although successful post-copulatory guarding was displayed only in those with chambers. The intensity of the agonistic encounters and the success of males in winning resources (burrows/females) also depend on the habitat characteristics. N. granulata shows a resource defense mating system where males employ different mating strategies according to the burrow architecture to ensure mating success.     

Paternity analysis of two male mating tactics in the fiddler crab, Uca mjoebergi

The fiddler crab Uca mjoebergi mates both underground in male-defended burrows and on the surface near female-defended burrows. The reproductive tract of Uca species facilitates last-male precedence, suggesting that males that do not guard-mated females are likely to gain very little paternity if the female re-mates with another male. Here, we test the reproductive success of burrow and surface matings using paternity analysis. We found that 100?% of the females that mated in burrows extruded a clutch of eggs. Furthermore, we show conclusively, for the first time in a fiddler crab species, that last-male sperm precedence results in the majority of the female’s eggs being fertilised by the burrow-mated male. In contrast, surface matings resulted in significantly fewer females extruding eggs (5.6?%). Paternity analysis also revealed that more than half of the clutches from burrow-mated females showed low levels of extra-pair paternity from previous matings. Although multiple matings appear common in U. mjoebergi, burrow-mated males that guard females are guaranteed a successful mating with extremely high rates of assured paternity. Surface matings therefore appear to be an opportunistic tactic that may increase male reproductive success in a highly competitive environment.     

Burrow morphology of the ghost shrimp<Emphasis Type="Italic"> Nihonotrypaea petalura</Emphasis> (Decapoda: Thalassinidea: Callianassidae) from western Kyushu,Japan

The callianassid shrimp Nihonotrypaea petalura (Stimpson, 1860) is a common member on boulder beaches in Japanese waters. Its burrow morphology was investigated, based on 28 resin casts collected from a steeply sloping beach with dense boulders and 30 from a more gently sloping beach with less dense boulders in Ariake Sound, southern Japan. The structure and dimensions of the burrows from the two beaches were basically the same, and thus a combined mean value could be given to most of their constituent elements. In its entire dimensions, the burrow is greater in lateral extent than depth, with a mean maximum horizontal extension of 145 mm and a mean maximum depth of 119 mm for the shrimp with a mean total length of 31.8 mm. The burrow winds along boulders or cobbles and consists of, from top to bottom, a single surface opening with an ejected mound, a top shaft leading to the uppermost chamber at a mean depth of 48–56 mm, a passage with a regular cross section that is wider than that of the top shaft, and bulbous chambers (mean no.=4.7) with an irregular cross section associated with branches (mean no. per burrow=1.2). Bulbous chambers are much larger than the uppermost chamber and are usually connected by passages, with some directly attached to each other. The combined architecture of these features is unique and relatively simple among the burrows of all callianassid species, the majority of which inhabit bare soft sediments. The structure and function of the N. petalura burrow are discussed in relation to lifestyle.Communicated by T. Ikeda, Hakodate     

Some direct observations on the ecology and behaviour of the Norway lobster Nephrops norvegicus

A detailed investigation of a small area of sea bed occupied by the Norway lobster Nephrops norvegicus (L.) was carried out by diving and television observations at depths of 30 m in Loch Torridon, Scotland. The density of burrows was 1/2 m², but only a proportion of these were occupied by N. norvegicus. Although about 70% of the larger burrows were occupied by N. norvegicus, giving a density of 1 lobster/8 m², very few juveniles (carapace length less than 30 mm) were found in the area. Many of the small burrows were occupied by the gobiid fish Lesueurigobius friesii (Collett). There was evidence that N. norvegicus frequently change their burrows, and fighting for burrows was observed. N. norvegicus remain within their burrows during the day, emerge around sunset to forage for food during the night, and then return to their burrows at dawn. This, and other aspects of their burrowing behaviour, have a marked effect on the commercial trawl catches of N. norvegicus which show large seasonal and diurnal variations in size and sex composition.     

Underlying reasons of the controversy over adverse effects of Bt toxins on lady beetle and lacewing larvae

In their recent study, Hilbeck et al. (2012) report that Cry1Ab causes lethal effects on larvae of the ladybird beetle Adalia bipunctata when fed directly to the predator. Such toxic effects were not previously observed in a direct feeding study conducted by us (álvarez-Alfageme et al. 2011). Because Hilbeck et al. (2012) claim that our study design did not allow us to detect any adverse effects we provide arguments for the value and relevance of our study in this commentary. Furthermore we discuss two additional published studies that have not revealed any direct effects of Cry1Ab on larvae of A. bipunctata and are not mentioned by Hilbeck et al. (2012). One of the studies was conducted in our laboratory under more realistic exposure conditions (álvarez-Alfageme et al. 2011). Feeding A. bipunctata larvae with spider mites reared on Bt maize did not reveal any adverse effects on lethal and sublethal parameters of the predator. This was despite the fact that the larvae had ingested high amounts of biologically-active Cry1Ab protein. Thus, we do not see verified evidence that A. bipunctata larvae are sensitive to Cry1Ab at realistic worst-case exposure concentrations. This, together with the fact that A. bipunctata will be little exposed to Cry1Ab under field conditions, allows us to conclude that the risk of Bt maize to this predator is negligible. Support for this comes from the results of many Bt maize field studies that have not revealed evidence for direct Cry1Ab-effects on non-Lepidoptera species.     

Contrast in the importance of arrow squid as prey of male New Zealand sea lions and New Zealand fur seals at The Snares,subantarctic New Zealand

New Zealand sea lions (Phocarctos hookeri) are threatened by incidental bycatch in the trawl fishery for southern arrow squid (Nototodarus sloanii). An overlap between the fishery and foraging sea lions has previously been interpreted as one piece of evidence supporting resource competition for squid. However, there is currently no consensus about the importance of squid in the diet of New Zealand sea lions. Therefore, we investigated this importance independently of spatial and temporal differences in squid availability through a simultaneous study with sympatric New Zealand fur seals (Arctocephalus forsteri), a species known to target arrow squid. Diet sampling at The Snares (48°01′S 166°32′E), subantarctic New Zealand, in February 2012 coincided with peak annual catch in the nearby squid fishery. Diets were deduced by analyses of diagnostic prey remains from scats (faeces) and casts (regurgitations). The contribution of each prey species to the diet was quantified using the per cent index of relative importance (% IRI) that combined frequency of occurrence, mass and number of prey items. Arrow squid was a minor component in sea lion scats (2 % IRI), and none was found in their casts. In contrast, arrow squid was the major component in fur seal scats and casts (93 and 99 % IRI, respectively). This study found that New Zealand sea lions ate minimal squid at a time when squid was clearly available as evidenced by the diet of New Zealand fur seals; hence, there was no indication of resource competition between sea lions and the squid fishery.     

Predation risk and foraging behavior of the hoary marmot in Alaska

Summary I observed hoary marmots for three field seasons to determine how the distribution of food and the risk of predation influenced marmots' foraging behavior. I quantified the amount of time Marmota caligata foraged in different patches of alpine meadows and assessed the distribution and abundance of vegetation eaten by marmots in these meadows. Because marmots dig burrows and run to them when attacked by predators, marmot-toburrow distance provided an index of predation risk that could be specified for different meadow patches.Patch use correlated positively with food abundance and negatively with predation risk. However, these significant relationships disappeared when partial correlations were calculated because food abundance and risk were intercorrelated. Using multiple regression, 77.0% of the variance in patch use was explained by a combination of food abundance, refuge burrow density, and a patch's distance from the talus where sleeping burrows were located. Variations in vigilance behavior (look-ups to search for predators while feeding) according to marmots' ages, the presence of other conspecifics, and animals' proximity to their sleeping burrows all indicated that predation risk influenced foraging.In a forage-manipulation experiment, the use of forage-enhanced patches increased six-fold, verifying directly the role of food availability on patch used. Concomitant with increased feeding, however, was the intense construction of refuge burrows in experimental patches that presumably reduced the risk of feeding. Thus, I suggest that food and predation risk jointly influence patch use by hoary marmots and that both factors must be considered when modeling the foraging behavior of species that can be predator and prey simultaneously.     

Length polymorphism at the avpr1a locus is correlated with male reproductive behavior in a natural population of prairie voles (Microtus ochrogaster)

Laboratory studies have shown that vasopressin can influence sociosexual behavior through its action on the vasopressin 1a receptor (V1aR). There is substantial evidence that the length of a microsatellite in the gene (avpr1a) encoding for the V1aR can affect social attachment to females and paternal behavior in male prairie voles under laboratory conditions. However, previous field studies of prairie voles have failed to detect a strong effect of the length of a male’s avpr1a allele on their sociosexual behavior, but these studies are typically much shorter than the average prairie vole breeding lifespan. We examined the relationship between male avpr1a microsatellite allele length and sociosexual behavior in a natural population of prairie voles for 15 weeks, closer to the lifespan of prairie voles in nature. Contrary to predictions, we found that males with the longest avpr1a microsatellite alleles were significantly more likely to sire offspring with more than one female and to sire offspring that survived until trappable age than males with the shortest avpr1a microsatellite allele lengths. This relationship was the strongest for males with the longest tenure on the study site. As in previous field studies, we did not find evidence of a relationship between a male’s avpr1a genotype and any index of social behavior including male residency status or the number of females with which males associate. This is the first study to support the hypothesis that a male’s avpr1a genotype is a factor underlying variation in the genetic mating system of prairie voles under natural conditions.     

Looking for the clock mechanism responsible for circatidal behavior in the oyster Crassostrea gigas

Valve activity rhythm of the oyster Crassostrea gigas is mainly driven by tides in the field, but in the laboratory, only a circadian clock mechanism has been demonstrated. In an attempt to reconcile these results, the mechanisms underlying the circatidal rhythm were studied in the laboratory under different entrainment or free-running regimes and in the field at Arcachon (44°39′N/1°09′W) in February–April 2011). Results confirm the existence of a circadian clock in C. gigas. Under entrainment regimes (12-h dark/12-h light photoperiod and tidal cycles simulated by a reversing current flow), oysters exhibited both circadian and circatidal cycles. Under free-running conditions (e.g., continuous darkness), the endogenous rhythm appeared to be circadian. There was no experimental evidence for an endogenous circatidal rhythm, even in oysters just transferred from the field, where a clear tidal cycle was expressed. There are two possible mechanisms to explain tidal behavior in C. gigas: an exogenous tidal cue that drives tidal activity and masks the circadian rhythm and an endogenous circatidal clock that is sensitive to tidal zeitgebers and runs at tidal frequency.     

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