An energetic evaluation of a predator–prey (leech–amphipod) couple in Montezuma Well, Arizona, USA

1. The highly stable conditions in Montezuma Well, Arizona, have contributed to a very close predator–prey relationship between the endemic leech, Erpobdella montezuma , and its endemic amphipod prey, Hyalella montezuma , which forms nearly 90% of the leech diet.
2. Using a simple bioenergetic model, it was determined why a benthic opportunistic foraging strategy, typical of most other erpobdellids, is inferior to the specialized foraging behaviour utilized by E . montezuma and why reproduction by the leech is restricted during certain times of the year.
3. Our model demonstrated that to attain a biomass in excess of 2000 mg (maximum field size) during periods of low amphipod densities in the winter E . montezuma must feed on alternative prey types and have a foraging activity < 4 h per day.
4. Leech reproduction 4–6 months prior to the onset of winter food shortages of prey increases juvenile and adult survivorship with highest reproductive success for E . montezuma occurring in June–July.
5. Rather than being an obligate forager on H . montezuma, E . montezuma exploits a periodically abundant amphipod during most of the year in addition to alternate prey types during the winter thereby achieving an overall higher rate of energy gain than could be achievable by other closely related species of erpobdellids utilizing an opportunistic foraging approach.     

The affect of temperature on the metabolism and behaviour of an endemic amphipod, Hyalella montezuma, from Montezuma Well, Arizona, U.S.A.

1. Hyalella montezuma is endemic to Montezuma Well, Arizona, and is exposed to minimal diel and seasonal temperature fluctuations in the pelagic zone (21 ± 4 °C). Juvenile H . montezuma feed in the pelagic zone during the day and migrate into the littoral vegetation at night, while adults remain primarily in the littoral vegetation.
2. Oxygen consumption ( V O₂) of adult and juvenile H . montezuma was measured at 20, 25 and 30 °C. The V O₂ of both adult and juvenile H . montezuma increased with temperature. However, the V O₂ of juveniles was significantly greater than that of adults at all temperatures, with greatest divergence at 30 °C where mean juvenile V O₂ (6.31 μl mg^–1 dry weight (DW) h^–1) was almost twice that of adults (3.60 μl mg^–1 DW h^–1).
3. Survivorship of juveniles was significantly lower (54%) at 30 °C than at 27.5 °C (95%) after 4 h, whereas adults showed at least a 93% survivorship at both temperatures.
4. Our data suggest that temperature may have been the proximate cue that elicited the diel horizontal migration of juvenile H . montezuma in Montezuma Well, with the behaviour maintained and enhanced by intensive invertebrate predation in the pelagic and littoral zones.     

The affect of temperature on the metabolism and behaviour of an endemic amphipod, Hyalella montezuma, from Montezuma Well, Arizona, U.S.A.

1. Hyalella montezuma is endemic to Montezuma Well, Arizona, and is exposed to minimal diel and seasonal temperature fluctuations in the pelagic zone (21 ± 4 °C). Juvenile H . montezuma feed in the pelagic zone during the day and migrate into the littoral vegetation at night, while adults remain primarily in the littoral vegetation.
2. Oxygen consumption ( V O₂) of adult and juvenile H . montezuma was measured at 20, 25 and 30 °C. The V O₂ of both adult and juvenile H . montezuma increased with temperature. However, the V O₂ of juveniles was significantly greater than that of adults at all temperatures, with greatest divergence at 30 °C where mean juvenile V O₂ (6.31 μl mg^–1 dry weight (DW) h^–1) was almost twice that of adults (3.60 μl mg^–1 DW h^–1).
3. Survivorship of juveniles was significantly lower (54%) at 30 °C than at 27.5 °C (95%) after 4 h, whereas adults showed at least a 93% survivorship at both temperatures.
4. Our data suggest that temperature may have been the proximate cue that elicited the diel horizontal migration of juvenile H . montezuma in Montezuma Well, with the behaviour maintained and enhanced by intensive invertebrate predation in the pelagic and littoral zones.     

Concomitant diel vertical migration of a predatory leech and its amphipod prey

SUMMARY.

• 1 The concomitant diet vertical migrations of the predaceous leech Erpobdella montezuma and its pelagic amphipod prey (Hyalella montezuma) were monitored for 1 year in Montezuma Well, Arizona, U.S.A.
• 2 High densities of H. montezuma occurred in the subsurface strata of the water column during the day, but a substantial portion of the population migrated into the surface 1 m at sunset. E. montezuma remained in the lower water strata during the day, but migrated vertically after sunset to exploit the high densities of H. montezuma near the surface. Densities of E. montezuma progressively increased in the upper strata of the water column after sunset as light energy fell below 1 μEin m^?2 s^?1.
• 3 It is suggested that the synchronized nocturnal migrations of the predator E. montezuma in relation to its prey H. montezuma, increases the foraging efficiency of the leech in the highly predictable environment of Montezuma Well.

     

Population dynamics and production of the pelagic amphipod Hyalella montezuma in a thermally constant system

SUMMARY. 1. The population dynamics and annual production of the multivoltine. pelagic amphipod Hyalella montezuma were studied over a 3-year period in the thermally constant environment of Montezuma Well. Arizona.
2. H. montezuma showed two maxima which coincided with spring and autumn phytoplankton maxima. Juveniles comprised over 85% of the population in the pelagic zone compared to 37% in the littoral vegetation and there were significantly more females in the littoral vegetation. It appears that juvenile and adult H. montezuma show different habitat preferences.
3. Total annual mean production for H. montezuma calculated by the size frequency method and adjusted for multiple generations, was 357 kg ha⁻¹ yr⁻¹, which is higher than single-species production estimates reported for most zoobenthic amphipods and multivoltine planktonic crustaceans. Average energy production for H. montezuma was 4640 kJ ha⁻¹ yr⁻¹ in the pelagic zone and 1072 kJ ha⁻¹ yr⁻¹ in the littoral vegetation.
4. Average cohort P / ratios for H. montezuma were higher in the pelagic zone (5.5) than in the litttoral vegetation (3.7). Juveniles had higher cohort P ratios than adults in the pelagic zone, while the reverse relationship was true in the littoral vegetation. We propose that different size-selective predators may contribute to the differences in P ratios for juveniles and adults in these two habitats.     

The evolutionary importance of mechanoreception in three erpobdellid leech species

Summary The ability of an erpobdellid leech, Erpobdella montezuma, to discriminate between two congeneric amphipod prey (Hyalella montezuma and Hyalella azteca) through mechanoreception was examined. Both E. montezuma and H. montezuma, a pelagic filter feeder, are endemic to the near thermally constant environment of Montezuma Well, Arizona, and form a highly specialized predator-prey relationship. In contrast, H. azteca, a benthic detritivore, is widely distributed throughout North America. Erpobdella montezuma was highly responsive to the prey signals of the endemic H. montezuma and showed only a limited response to H. azteca. An inverse relationship occurred between the frequency of attack and size of the leech. The abilities of Erpobdella punctata and Nephelopsis obscura, neither found in Montezuma Well, to detect the signals of both amphipod prey were also examined. These species with non-specialized diets showed a low response to the signals of both prey species. This study demonstrates that very close predator-prey relationships can develop over evolutionary time in isolated aquatic systems through the use of mechanoreception.     

The recent evolutionary origin of the phenotypically novel amphipod Hyalella montezuma offers an ecological explanation for morphological stasis in a closely allied species complex

Numerous molecular studies have identified morphologically cryptic, freshwater invertebrate species, but have not suggested possible mechanisms for their phenotypic stasis. The amphipod crustacean genus Hyalella contains numerous morphologically cryptic species in the H. azteca complex, as well as a small number of morphologically very divergent, narrowly endemic taxa. One such taxon, Hyalella montezuma, is the sole planktonic filter-feeder within the North American amphipod fauna, and is known only from Montezuma Well, a fishless travertine spring mound in Arizona, USA. In this study, we conduct a phylogenetic analysis of mtDNA sequence data using likelihood, Bayesian and cladistic approaches to determine both the relationship of H. montezuma to the H. 'azteca' species complex, and to ascertain if its morphological and ecological differentiation have been comparatively recent. The results show that H. montezuma has a very close phylogenetic affiliation with one lineage in the H. azteca complex, indicating that its origin has been recent. We present evidence suggesting that fish predation is an important ecological factor, which constrains morphological and ecological diversification within the genus Hyalella, and that Montezuma Well has provided a relaxation on this constraint.     

Diel vertical migration of a pelagic amphipod in the absence of fish predation

The pelagic amphipod, Hyalella montezuma, migrates vertically into the surface waters at twilight in Montezuma Well, Arizona, USA despite the absence of fish predators or thermal stratification. We suggest that a persistent, dense, neustonic algal assemblage may provide a food resource incentive for the twilight ascent.     

Herbivory on a chemically defended plant as a predation deterrent in Hyalella azteca

SUMMARY 1. We investigated whether a population of the freshwater amphipod, Hyalella azteca , which consumed plants with defensive secondary compounds, reduced predation as reported for terrestrial and marine systems.
2. Field observations in Montezuma Well, U.S.A., indicated a strong association between the emergent macrophyte, Berula erecta and H . azteca . We hypothesised that this geographically isolated population of sedentary amphipods was able to consume roots of the chemically defended B . erecta as a deterrent against predation. Berula erecta is in the family Apiaceae, which commonly produce coumarins that deter herbivory in terrestrial systems.
3.  Hyalella azteca consumed roots of B . erecta at a significantly greater rate than alternative macrophyte substrata in Montezuma Well. Additionally, H . azteca moulted at a significantly higher rate when consuming B . erecta compared with a diet of periphyton.
4. Two insect predators ( Telebasis salva and Belostoma bakeri ) with different feeding strategies were used to assess the effects of a B . erecta diet on predation rates in the laboratory and in Montezuma Well. Hyalella azteca was preyed on at a significantly lower rate by both predators when given a strict diet of B . erecta roots compared with a diet of periphyton.
5. This is the first experimental evidence that predation on a freshwater herbivore, H . azteca , was reduced when it consumed a chemically defended plant.     

Piscivore diet response to a collapse in pelagic prey populations

Pelagic fish populations in the upper San Francisco Estuary have experienced significant declines since the turn of the century; a pattern known as the pelagic organism decline (POD). This study investigated food habits of piscivorous fishes over two consecutive fall seasons following the decline of pelagic fish prey. Specifically, this study addressed the contribution of pelagic versus benthic prey to piscivorous fish diets, including the frequency of predation on special-status pelagic species, and the spatial variability in prey consumption. The piscivore community was dominated by Striped Bass and also included small numbers of Sacramento Pikeminnow and Largemouth Bass. Overall, pelagic prey items contributed less than 10% of the diet by weight in both years, whereas pre-POD studies gleaned from the literature found contributions of 39–100%, suggesting a major switch from pelagic to benthic prey resources. Between-year variation in piscivore diets reflected differences in environmental conditions associated with variation in freshwater outflow. No special status fish species were detected in any of the piscivore stomachs examined. The consequences of this pelagic to benthic diet shift warrants further investigation to understand its ecological relevance.     

Combined gut‐content and stable isotope trophic analysis of the pelagic stingray Pteroplaytrygon violacea (Bonaparte, 1832) diet from the western North Atlantic Ocean

The understanding of trophic relationships is vital for correctly modeling ecosystems and ecosystem effects of fisheries removals. The pelagic stingray is found in epipelagic sub‐tropical and tropical waters worldwide and is a common bycatch in pelagic longline fisheries. Between August 2008 and November 2011, 156 specimens (81 males; 75 females) were collected during pelagic longline fishing operations in the US South Atlantic Bight and Gulf of Mexico. Stomach content analyses found that the major prey items were cephalopod molluscs (59.18%), followed by actinopterygiian fishes (37.75%), and decapod crustaceans (35.71%). These concentrations of prey items found in the stomachs coincide with previous studies done in the Pacific Ocean. In contrast to previous studies that found high percentages of empty stomachs (63%), the current percentage of empty stomachs was much lower (25.6%), likely due to shorter times between collection and inspection. Stable isotope analysis (δ¹³C and δ¹⁵N) was performed on white muscle in order to correlate the trophic position with gut‐content analysis. The δ¹³C values ranged from ‐18.81‰ to ‐16.70‰, while the δ¹⁵N ranged from 6.11‰ to 11.88‰. Modeling of stable isotope data suggest that while squid are occasionally an important part of the pelagic stingray diet, prey usually consist of shrimp and other pelagic crustaceans. Pelagic stingrays fed within two trophic levels, but their prey appeared to feed on different carbon sources than those found in other pelagic elasmobranchs. A deeper understanding of the pelagic stingray diet sources can help fisheries management as it begins to transition into ecosystem‐based management.     

Diet comparison between pelagic and demersal whiting in the North Sea

Pelagic North Sea whiting Merlangius merlangus preyed upon (pelagic) sprat and sandeel, while demersal whiting preyed upon (demersal) Norway pout and herring. Values of diet breadth were low for both feeding groups using Levin's index. Diet overlaps within layers were low ( D <0·25), while the between layer food overlap was moderate ( D =0·25–0·74) to high ( D >0·74) using Schoener's index. Selection of prey was density dependent. However, prey size also played an important role. The diet of whiting shifted from amphipods and mysids to fish with increasing predator length, and the length of prey consumed increased significantly with length of whiting. The fact that the stomach contents differed between the feeding groups demonstrates the need for a sampling design that includes both pelagic and demersal habitats when trying to quantify the diet of whiting.     

Empirical evaluation of phenotype–environment correlation and trait utility with allopatric and sympatric whitefish, Coregonus lavaretus (L.), populations in subarctic lakes

Adaptive phenotypic divergence of sympatric morphs in a single species may have significant evolutionary consequences. In the present study, phenotypic impacts of predator on zooplankton prey populations were compared in two northern Finnish lakes; one with an allopatric whitefish, Coregonus lavaretus (L.), population and the other with three sympatric whitefish populations. First, we examined whether there were phenotypic associations with specific niches in allopatric and sympatric whitefish. Second, trait utility (i.e. number of gillrakers) of allopatric and sympatric whitefish in utilizing a pelagic resource was explored by comparing predator avoidance of prey, prey size in environment, and prey size in predator diet. The allopatric living large sparsely rakered (LSR) whitefish morph, was a generalist using both pelagic and benthic niches. In contrast, sympatric living whitefish morphs were specialized: LSR whitefish was a littoral benthivore, small sparsely rakered whitefish was a profundal benthivore and densely rakered (DR) whitefish was a pelagic planktivore. In the lake with allopatric whitefish, zooplankton prey did not migrate vertically to avoid predation whereas, in the lake with sympatric whitefish, all important prey taxa migrated significantly. Trait utility was observed as significantly smaller size of prey in environment and predator diet in the lake with DR whitefish than in the lake with only LSR whitefish.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 561–572.     

The diet of the Weddell seal <Emphasis Type="Italic">Leptonychotes weddellii</Emphasis> at Laurie Island,South Orkney Islands

The diet of the Weddell seal Leptonychotes weddellii at the South Orkney Islands was investigated by the analysis of 44 and 26 faecal samples collected from the beaches of Laurie Island from May 1999 to January 2000 and from October to December 2001, respectively. The diet was diverse and both pelagic and benthic-demersal organisms were represented in the samples. Fish were the most frequent prey of the samples in both seasons, and fish and krill were the most numerous prey items. Octopods predominated by mass in 1999, whereas krill constituted the bulk of the diet in 2001, a fact that was not observed previously. Among fish, Gobionotothen gibberifrons was largely the species that contributed mostly to the diet, whereas pelagic fish could not be identified in the samples. The results are compared with information from other study areas and discussed in relation to the consumption of krill.     

The diets of polar cod (Boreogadus saida) from August 2008 in the US Beaufort Sea

Polar cod (Boreogadus saida) play an integral part in the Arctic ecosystems linking the upper and lower trophic levels. Though their estimated biomass is considerable, recent knowledge of their diets in the US Beaufort Sea is sparse. Collections of polar cod from the US Beaufort Sea were made during August 2008 using demersal and pelagic trawls. Polar cod diet composition was quantified as percent prey weight, percent prey count, and frequency of occurrence of prey. The diet composition between the demersal- and pelagic-captured cod showed differences in all these categories. Polar cod captured in the demersal nets primarily fed on fish (by weight), and pelagic cod primarily fed on copepods (frequency of occurrence) and euphausiids (by weight). In general, these dominant preys are different than what has been reported in other studies describing polar cod diets.     

Feeding habits of <Emphasis Type="Italic">Bathydraco marri</Emphasis> (Pisces,Notothenioidei, Bathydraconidae) from the Ross Sea,Antarctica

In comparison with other bathydraconids, all species of the genus Bathydraco are poorly known from an ecological perspective. The diet of juvenile Bathydraco marri Norman, 1938 was studied for the first time in specimens collected in the southwestern Ross Sea during summer 1998. Fish were collected in a single otter trawl catch at 330–340 m depth. The stomach content analysis showed that this species fed exclusively on crustaceans. Overall, 20 prey taxa were identified to genus or species level. Mysids, amphipods and copepods were the most important prey in decreasing order of importance. Other prey, such as Euphausia superba, isopods and tanaids were eaten occasionally and in very small amounts. A multivariate analysis was applied to feeding data to assess ontogenetic or sex-related changes in diet. No difference was detected between sexes, whereas diet of small and large fish differed in some degree. An ontogenetic shift from small and pelagic crustaceans such as copepods to benthic–benthopelagic prey such as amphipods and mysids was observed. Relating present results with published data on physiological characteristics of B. marri, it was possible to infer their feeding behaviour and mode of life. Like other bathydraconids, this species appeared to be an inactive and sluggish fish, which relied on more or less motile benthic or epibenthic prey adopting a “sit and wait” feeding strategy. On the other hand, smaller fish seem to be more active, feeding also on pelagic prey such as copepods that can be seasonally abundant, thus reducing the intraspecific competition for food.     

Does diet in Celtic Sea fishes reflect prey availability?

Feeding preferences of Celtic Sea fishes were investigated using a database of stomach content records, collected between 1977 and 1994. The diet of cod Gadus morhua , hake Merluccius merluccius , megrim Lepidorhombus whiffiagonis , whiting Merlangius merlangus and saithe Pollachius virens changed markedly as the animals grew larger, and although large predators generally chose larger bodied prey, the variability of prey sizes consumed also increased. Large predators continued to select small, low value, benthic prey ( e.g . Callionymus spp. and Trisopterus spp.) which were easier to catch, rather than larger, more energy lucrative pelagic prey ( e.g . mackerel Scomber scombrus ), even though these pelagic prey‐fishes were nearly always available and were often very abundant. Stock estimates of the International Council for the Exploration of the Sea and U.K. groundfish survey catches were used as indices of prey abundance. Blue‐whiting Micromesistius poutassou and other small pelagic fishes ( Argentina spp. and clupeoids) were identified as being particularly important, and were consumed by some predators more often than would be expected given the abundance of these prey in the environment. There was no evidence for density‐dependent feeding by predators on mackerel and only hake exhibited density‐dependent feeding on horse‐mackerel. Hake, cod and megrim consumed more blue‐whiting when this prey was at higher abundance in the environment. In choosing what prey to consume, predators must balance costs and benefits, considering the quality of prey and the energy expended during search, capture and handling.     

How important are pelagic preys for the kelp gull during chick-rearing at the South Shetland Islands?

An analysis of pellets regurgitated indicated adult kelp gulls (Larus dominicanus) on the South Shetland Islands consumed predominantly intertidal prey, whereas previous studies at Antarctic Peninsula sites have reported kelp gulls consuming predominantly pelagic species. The pellets collected at Nelson Island during the chick-rearing period indicated that the limpet Nacella concinna was their most frequent prey, followed by carrion, gammariids, snails and krill. Fish were scarcely represented. Also, regurgitated stomach contents of chicks showed that limpets and carrion were the most frequent food items, accounting for 70% of the mass. However, gammariids were particularly important by number. Significant differences were observed in the overall comparison of the diet as reflected by the two sampling methods. In general, the importance of pelagic prey was negligible when compared to intertidal or scavenged prey. Our results differ greatly from those reported for the Antarctic Peninsula, where chicks were almost exclusively fed with the pelagic fish Pleuragramma antarcticum. These differences could be related to the abundance of pelagic resources in southernmost latitudes, and/or to the presence of more extensive intertidal foraging areas at the South Shetland Islands. Received: 7 October 1996 / Accepted: 14 July 1997     

Phytoplankton ecology in an unusually stable environment (Montezuma Well,Arizona, U.S.A.)

Relatively minor annual amplitudes of change in certain major nutrients, and especially pH and water temperature were measured in the spring-fed system of Montezuma Well, Arizona during a four year study. phytoplankton diversity was low but for the most part, composition was spatially and temporally constant; total seasonal phytoplankton density was significantly correlated with regional incident light. Phytoplankton species composition changed briefly during and for a short period following the summer monsoon. Ultraplankton (<5 µm diam.) numerically comprised nearly 80% of the phytoplankton community throughout most of the year. The limited residence time of water in the Well may have provided a competitive advantage for cells with high surface area:volume ratios and correspondingly rapid division rates. Nannochloris bacillaris Naum. and Coccomyxa minor Skuja were perennial dominants. Diatom populations did not increase with annual increases in vernal solar radiation. Low pH, high dissolved CO₂, and limited residence time for metabolic inhibitors are considered to be largely responsible for the reduced blue-green populations in the Well. The only flagellated photosynthetic group present in Montezuma Well was the Cryptophyta. Desmid populations were minimal, even though pH was consistently below circumneutral (6.5) and free CO₂ concentrations high. The role of grazing by an amphipod, Hyalella montezuma, on annual phytoplankton abundance is examined.     

Timing of the diet shift from zooplankton to macroinvertebrates and size at maturity determine whether normally piscivorous fish can persist in otherwise fishless lakes

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