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| Habitat & community ecology | |||||
| The section on HABITAT & COMMUNITY ECOLOGY includes a selection of topics such as interspecific competition, considered here, and COMMUNITY INTERACTIONS, INTRASPECIFIC COMPETITION, GENE FLOW, and HERMIT-CRAB COMPETITION, considered in other sections. | |||||
| Interspecific competition | |||||
Competition in crabs, as in other animals, may be exploitative, where use of a common resource by one organism denies its use to another organism, or interference, where one organism actively prevents or interferes with another organism's use of a common resource. It can occur interspecifically or intraspecifically, and may involve food, mates, space, or shelter. Most attention in the scientific literature has been paid to sympatric, congeneric pairs of species competing for space, and to hermit crabs fighting over shelter in the form of shells. This section starts with studies of west-coast crabs in general and concludes with a section on invasive green crabs Carcinus maenas |
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Research study 1 |
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Laboratory tests show that juvenile Hemigrapsus are also able to tolerate conditions of siltation and low salinity that Pachygrapsus cannot (see graph on Right). The author concludes that the species are able to coexist through a balance of predatory/competitive capabilities of Pachygrapsus and burrowing/physical stress-tolerance capabilities of Hemigrapsus. Photo of Pachygrapsus crassipes courtesy Jackie Soanes, Bodega Marine Laboratory, California NOTE mean burrow size in the upper region is 10 cm2, 3 times that of the middle region, and 5 times that of the lower region |
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Research study 2 |
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Five species of spider crabs Loxorhynchus crispatus, Pugettia producta, P. richii, Mimulus foliatus, and Scyra acutifrons inhabit kelp-forest habitats along the central California coast. Their numerical abundances and apparent similarily of habitats suggest the potential for strong competitive interactions within members of the guild. However, a comprehensive study at Hopkins Marine Station, Pacific Grove, California shows that in the major dimensions of body size, microhabitat occupation, and foods eaten, sufficient differences exist to suggest that interspecific competiton is minimal. First, as illustrated in the drawings above, the body size at maturity of the 5 species spans an order of magnitude in carapace width from small S. acutifrons (1cm) to large L. crispatus (10cm).
Finally, analyses of stomach contents show partitioning of food resources from dietary generalists such as L. crispatus foraging on a broad range of invertebrates and giant kelps, to strict dietary specialists such as P. producta grazing on kelp Macrocystis pyrifera, to M. foliatus and P. richii capturing drift pieces of the same kelp (see histograms on Left). Scyra acutifrons is an intermediate dietary specialist that feeds on detritus, sponges, and pieces of kelp, and Loxorhynchus crispatus also exploits a wide range of foodstuffs. Niche separation in the spider-crab guild is therefore multidimensional, with overlap in one resource generally being compensated by different utilisation of another resourch. |
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Research study 3 |
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NOTE an easy guide to distinguishing the species is that P. cinctipes has burgundy-coloured antennae, while P. eriomerus has straw-coloured antennae |
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Research study 4 |
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Results show that in high-intertidal cages containing P. cinctipes adults there are significantly more P. cinctipes megalopae than in CONTROL cages (see graph above Left). Essentially no P. eriomerus megalopae settle in the high cages. Similarly, in low-intertidal cages, fewer P. eriomerus megalopae settle in cages with P. cinctipes adults or in CONTROL cages, than in cages containing P. eriomerus adults (see histogram on Right). Finally, significantly more P. cinctipes megalopae settle in low-intertidal cages containing P. cinctipes adults than in CONTROL cages or in cages containing P. eriomerus adults (see graph lower Left). In summary, the megalopae preferentially settle with conspecifics regardless of intertidal height. This settlement strategy gives a “kick-start” to the spatial separation of the 2 species, without the newly settled juveniles being subjected to long, hazardous, and possibly fatal migrations up or down the shore. Near the time of settlement the megalopae tend to be found close to the water surface. Thus, on incoming tides the larvae would wash over rocks containing conspecific adults, first, the lower dwelling P. eriomerus, and then, the higher dwelling P. cinctipes. The larvae presumably perceive species-specific chemical cues emanating from the adults, and drop down to join them.
NOTE supportive data for this are provided in later laboratory experiments at the Bamfield Marine Sciences Centre, British Columbia showing that P. cinctipes megalopae swim less in the presence of conspecific adults than in control treatments where adults are absent (see graph on Right). The megalopae are attracted to adults and settle within 2-4d. If no adults are present, then settlement is delayed and the megalopae swim for a further 2-3wk |
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Research study 5 |
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Research study 6 |
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Casual observation of a mixed kelp forest would suggest that different species of kelp-inhabiting crabs, such as Pugettia gracilis and Oregonia gracilis, are distributed more or less randomly, but this is not the case. Monthly SCUBA surveys at 6 habitats in Kachemak Bay, Alaska over a 16mo period by researchers from the University of Alaska Fairbanks show that each species has its own special microhabitat-resource NOTE several other decapods are present at the sites, but the 2 species listed are by far the most common NOTE 3 of the sites contain both high-canopy kelps (Nereocystis luetkeana) and understory kelps (Laminaria spp., Saccharina spp. Agarum clathratum, Costaria costata, and Cymathaere triplicate), while the remaining 3 sites contain only understory kelps
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| Invasive green crabs Carcinus maenas | |||||
Research study 1 |
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First records of the North Atlantic green crab Carcinus maenas on the west coast date from 1989-90 in San Francisco Bay, California. Its mode of introduction is uncertain, but could involve ballast water, fouling growth on ships’ hulls, imported live bait (and/or imported clawed lobsters), or NOTE green crabs invaded eastern NA in early 1800s and South Africa in the 1980s |
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Research study 2 |
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The green crab Carcinus maenas is a notably aggressive species. Fortunately, its distribution here on the west coast is limited to more protected habitats. Potential prey populations in more wave-exposed habitats are unlikely to be affected. In subtidal sand/mud habitats, one would predict that green crabs would be out-competed by the larger Dungeness crab C. magister; however, casual laboratory observations indicate that C. maenas will actually prey upon Cancer magister and Hemigrapsus oregonensis of equal or smaller size, and larger-sized Cancer magister appear not to eat the green crabs. However, the 2 species may compete for common prey items. After enclosing single individuals of Carcinus maenas in wire-net enclosures on soft-bottom habitats in Bodega Harbor, California for 12d, then assessing potential prey abundances in experimental and control enclosures, researchers record significant decreases in abundance of bivalves, cumaceans, amphipods (1 species only studied), but not of tanaidaceans, amphipods (1 species), polychaetes, and phoronids, although some trends are apparent. The bivalves, representing 2 species of Transennella, are numerically the second and third most abundant invertebrates in the cores (to 5cm depth) and represent an important food item of Dungeness crabs. With the exception of the polychaete Exogene sp., most of the other non-eaten species are in low abundances. NOTE Transennella spp. are noted by the authors to reach >10,000 individuals . m-2 in various coastal embayments NOTE tanaidaceans are the most abundant invertebrates, numbering 300/10cm-dia x 5cm-deep core in the control enclosures Transiennella sp. |
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Research study 3 |
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So, what is the predicted impact of invasion of European green crabs Carcinus maenas onto west-coast shores and other areas of the world? This is estimated by comparing invasion characteristics for 3 areas now hosting green crabs (western NA, eastern NA, and South Africa) with the same characteristics in its native range in Europe. The characteristics are: 1) habitat usage, 2) diet preferences, 3) size of individuals, 4) rate of range expansion, and 5) demonstrated and potential impacts.
In comparison, diet preferences
Finally, potential ecological impact seems quite varied and probably indicates the strong The approach is a nice one but falls short of the author’s expectations mainly through lack of good comparative data. It would be interesting to see how the addition of another decade’s worth of data will add to the picture. Overall, this is a lovely study. NOTE inclusion of echinodermata is a bit unusual, in that presumably many other taxa are not present in the diets of green crabs and yet are not shown here NOTE the authors actually conclude that ecological impact is “relatively similar” across the 3 invasions. However, if “impacts on fishes and birds” is removed as being too uncertain, and “impacts on bivalve molluscs” and “impacts on echinoderms” are removed for reason that they are already included in “diet preferences”, then the remaining entry “impacts on other crabs” is quite varied (ranging from “-“ or no impact to “++” or high impact |
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Research study 4 |
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In 1993 Carcinus maenas was recorded in Bodega Bay, about 100km north of San Francisco, and in 1997 in Coos Bay, Oregon, several hundred kilometers further north. Carcinus maenas has aggressive feeding habits and has already become a serious pest for mariculturists – particularly those attempting to rear Manila clams Venerupis phillippinarium, themselves introduced into the San Francisco Bay area in the 1940’s. NOTE the species was also observed at Nootka Sound, British Columbia in 2002 |
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Research study 5 |
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Field experiments in which juvenile green crabs are caged with similar-sized H. oregonensis in 0.3m2 traps or cloth cages for 1-2d periods show that the indigenous species is no match for the invader.
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Research study 6 |
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Similarly, in field tests in Bodega Bay Harbor where experimentally placed oyster-shell plots are allowed to be colonised by H. oregonensis2 and C. maenas, the former species also dominates (see graph on Right). In the laboratory, as well, Hemigrapsus consistently dominates equal-sized Carcinus in contests for shelter. However, in arena-type competitions for food, using a screw- and wire-anchored damaged bivalve3 as prey, C. maenas dominates over H. oregonensis. This suggests that the competitive interactions between the 2 species may actually favour H. oregonensis for interference-type space competition, and C. maenas for exploitative-type food competition. NOTE1 notably Hemigrapsus oregonensis, H. nudus and, in southern Oregon and California, Pachygrapsus crassipes NOTE2 density of H. oregonensis in these experimental plots reach in excess of 600 individuals . m-2 NOTE3 either Mytilus spp. or Venerupis philippinarum |
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Research study 7 |
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Finally, lab and field enclosure experiments show that Dungeness crabs tend to move out of oyster-shell refuges also inhabited by green crabs and to go into open-sand habitat (see schematic lower Left). Conversely, in both situations C. magister is more likely to remain in the shell refuge in the absence of C. maenas. The authors conclude that invasive green crabs could have an overall negative impact on the indigenous Dungeness crab fishery through predation on young stages and exclusion from protective oyster-shell habitats. NOTE the shelters are single clam shells, competed for by 2 crabs, one of each species NOTE a recent assessment of economic impact of C. maenas on commercial shell-fisheries on the U.S. west coast using a combination of ecological and economic models concludes that past and present impacts are minor. Future impacts, such as into Alaska, will depend upon degrees of density increases and range expansion. |
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Research study 8 |
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| Laboratory studies in Oregon show that green crabs Carcinus maenas eat a variety of bivalves. Preference for the species tested is in the order: Ostrea (lurida) conchaphila = Cryptomya californica > Macoma nasuta = Venerupis philippinarum. | |||||
Research study 9 |
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![]() After its establishment in San Francisco Bay in 1989, green crabs Carcinus maenas have moved steadily northwards. In 1998, coincidental with an unusually strong El Niño-generated current flowing northwards from September 1997 to April 1998, a strong settlement occurred along the coast as far north as Vancouver Island, British Columbia. Studies on this cohort by a consortium of researchers shows, interestingly, that growth is significantly faster than in east-coast and European counterparts (see graph). Note in the graph that west-coast specimens reach 46mm carapace width by the end of the first summer, and 66mm by the end of the second. Comparable sizes for European specimens are 18 and 43mm, respectively. Warmer water temperatures on the west coast certainly lead to more frequent moulting but, as discussed by the authors, there are likely to be other contributing factors, most notably food limitation and higher incidence of parasitism in the more crowded European populations. Green crabs live for only 4-6yr, so the original colonising year class of 1998 has probably completely died out. Northward-flowing currents of the one just described occur only sporadically, but local reproduction and recruitment appear high enough to maintain permanent populations in Oregon and Washington estuaries. Photograph courtesy Museum Victoria Australia. |
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Research study 10 |
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A novel idea with respect to habitat use by green crabs Carcinus maenas in Willapa Bay, Washington is to model their bioenergetics budgets in 4 habitat scenarios to determine which suits their metabolic expenditures most favourably. The researchers input a variety of ecological and physiological parameters published for C. maenas and, in the single case of excretion, for related brachyuran into the 4 models to determine energy expenditures in the different habitats over a 214d simulation period. Results show, interestingly, that predicted metabolic costs are about 6% higher in intertidal habitats than in subtidal ones. The intertidal-simulation crabs, moreover, are less efficient than subtidal ones in converting consumed food energy into growth. So, where do the real green crabs live in Willapa Bay? Monthly trapping reveals that they live predominantly in mid-intertidal habitats and not at all in subtidal ones. The authors credit the (fairly major) discrepancy from model prediction to their failure to include interspecific interactions, such as agonistic ones that may occur with subtidal competitors such as Dungeness crabs. Thus, although subtidal life is predicted to suit C. maenas better, the species may be excluded from it by unfavourable interactions with C. magister. Although laboratory tests of competitiveness of the 2 species (see Research Study 7 above) actually show that Dungeness crabs are inferior to green crabs, such interactions regardless of the outcome would increase metabolic costs for both participants and, for energetic considerations alone, are best avoided. For readers that may have doubts about the overall NOTE these are the high intertidal, mid-intertidal, and subtidal areas, with the fourth being the subtidal area with migrations into the intertidal region NOTE these include estimates for energy equivalents of consumption, defecation, excretion, and oxygen-uptake rates; air and water temperatures; energy loss to specific-dynamic action; daily tidal cycles; activity “multipliers” during submergence during day and night high tides; energy contents of various bivalve prey; and so on Green crab Carcinus maenas on |
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Research study 11 |
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Research study 12 |
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How suitable is the newly invaded west-coast habitat for growth of green crabs Carcinus maenas? The answer, based upon a comparison of crab sizes between a “home” location in Anglesey, Wales (53oN latitude) and an invaded habitat in British Columbia, Canada (49oN) is, quite suitable indeed. The British Columbia population (both sexes) is significantly larger than the Anglesey population (by about 40%), suggesting a faster growth rate and thus a potentially greater fecundity. Interestingly, while the home population is comprised of green- and red-coloured morphs (about 45 and 55%, respectively, for both sexes), the invasion population is predominantly green in colour (84% of males and 47% of NOTE the area selected for study in British Columbia is Barkley Sound, first colonised in 1999 by just a few individuals, but then more intensively settled in 2005. Water temperatures in this area are similar to those in Anglesey
Green (on Left), orange, and red colour |
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Research study 13 |
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NOTE see Research Study 2 above for a report on an early part of the study done by members of the same research team |
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