Utility of geometric morphometrics for inferring feeding habit from mouthpart morphology in insects: tests with larval Carabidae (Insecta: Coleoptera)

Feeding habits are important life‐history traits in animals; however, methods for their determination are not well established in many species. The larvae of the beetle family Carabidae are an example. The present study tested the utility of geometric morphometrics of mouthpart morphology to infer the feeding habits of carabid larvae. Using Pterostichus thunbergi as a model system, larval feeding habits were inferred using geometric morphometrics of mouthparts and the results were compared with those obtained from rearing experiments. The rearing experiments indicated that P. thunbergi larvae are carnivores that require snails as an essential part of the diet. Through geometric morphometrics, associations between mouthpart morphology and larval feeding habits were confirmed for species in which these two traits are known. A discriminant analysis using these associations classified P. thunbergi larvae as snail/slug feeders, which is a result compatible with the rearing experiments. Geometric morphometrics also revealed that morphological integration and ontogenetic shape change might play roles in the diversification of mouthpart morphology. Overall, these results demonstrate the utility of the geometric morphometrics of mouthparts to infer feeding habit and to clarify the mechanisms of mouthpart morphological diversification in the study group, and the results also serve as a basis for future studies of other insect groups.     

Zooplankton capture by a coral reef fish: an adaptive response to evasive prey

Synopsis High-speed cinematography and video using modified Schlieren optics and laser illumination helped elicit details of prey capture mechanisms used by Chromis viridis while feeding on calanoid copepods and Artemia. Chromis viridis is capable of a ram-jaw, low-suction feeding, as well as a typical suction feeding behavior described for other species of planktivores. By adjusting the degree of jaw protrusion and amount of suction used during a feeding strike, this fish can modulate its feeding strikes according to the prey type being encountered. The ram-jaw feeding mode enables C. viridis to capture highly evasive calanoid copepods within 6 to 10 msec. The use of specialized feeding behavior for evasive prey and the ability to vary feeding behavior are adaptations for feeding on evasive prey.     

Feeding behavior and related functional morphology of the mayflyEphemerella needhami (Ephemeroptera: Ephemerellidae)

The feeding behavior and functional morphology associated with feeding in Ephemerella needhamiMcDunnough larvae were studied using videomacroscopic techniques, gut content analysis, and scanning electron microscopy. Two stereotypic feeding cycles were employed by the larvae. In the maxillary brushing cycle, the maxillae are the primary food-gathering organs, with the main food being detritus deposited on the filamentous alga Cladophora.In the mandibular biting cycle, the mandibles are the primary food-gathering organs used to bite Cladophora filaments.Epiphytic diatoms on Cladophorawere another important part of the diet. Behavioral similarities are apparent in the choreography and synchronization of mouthpart movements among mayflies from several families. Functional morphological comparisons are drawn with hypognathous E. needhami, Cloeon dipterum, Baetis rhodani,and Siphlonurus aestivalis, aswell as the prognathous Heptageniidae. Differences in mouthpart usage and structure are related to the relative development of setal fields and combs and the feeding microhabitat.     

Comparative population dynamics of the rotifers Brachionus angularis and Keratella cochlearis

The results indicate that the two rotifer species Brachionus angularis and Keratella cochlearis have different life history strategies. Brachionus angularis appeared to be an r-strategist, showing higher reproductive, mortality and population growth rates than K. cochlearis. Additional evidence is offered suggesting that K. cochlearis is a K-strategist. Although K. cochlearis had a lower growth rate than B. angularis, this was compensated by a lower mortality rate, especially in pre-reproductive individuals. In addition, K. cochlearis allocated energy to reproduction more efficiently than B. angularis.     

Mouthpart morphology and feeding behavior of the invasive kudzu bug,Megacopta cribraria (Hemiptera: Plataspidae)

The invasive kudzu bug, Megacopta cribraria, was first reported in North America in 2009 and has subsequently spread through most of the southeastern United States, causing yield loss in soybean. Since detection in the USA, research has focused mainly on managing this newly established pest, but many important characteristics of the pest's mouthpart morphology and feeding behavior are unknown. Qualitative and quantitative comparisons of nymph and adult mouthparts and sensilla were made through scanning electron microscopy and light microscopy, and feeding behavior was examined using electropenetrography (EPG) and paraffin histology. Morphologies observed were similar to what has previously been reported for other piercing–sucking hemipterans. The relationship between rostrum length and body size (pronotum width and dorsal length) exhibited negative allometry. Rostrum length exhibited an isometric relationship with interocular width. Adult females (n=9) probed soybean stems 1.3±0.8 times in 9 h, with an average probe time of 2.3±1.3 h. EPG waveforms were characterized and correlated with behavior. Salivary sheaths were shown to terminate in the vascular tissue; four of five sheaths terminated in the phloem. This is the first time that the feeding behavior of a member of the Plataspidae has been recorded using EPG. Results add to our current limited knowledge of plataspid mouthpart morphology and provide a baseline for further research on the feeding behaviors of M. cribraria and other soybean‐feeding hemipterans.     

Prey selection by Asplanchna girodi (Rotifera): the importance of prey defence mechanisms

1. Predator (laboratory-grown Asplanchna girodi and prey (several field-collected non-carnivorous rotifer species) interactions were studied in the laboratory as a function of both predator and prey densities. The clearance rates of A, girodi decreased with increasing prey density. Predator density had no effect on the feeding of A. girodi. 2. Asplanchna girodi selectively fed on Keratella cochlearis. Its clearance rates for K. cochlearis were much higher than those for Polyarthra and Brachionus. Short-spined forms (spine length less than 25 μm) showed a significantly higher susceptibility to predation than either the non-spined or the long-spined forms. 3. Large Asplanchna individuals fed selectively on reproductive females of K. cochlearis, thereby reducing the fecundity of their prey.     

A centrifugation method for measuring the relative density (specific gravity) of planktonic rotifers (Rotifera), with values for the relative density of Polyarthra major Burckhardt and Keratella cochlearis (Gosse)

The relative density of Polyarthra major and Keratella cochlearis was measured by centrifugation in solutions of different densities. Ficoll was used as the solute. The mean values for the relative density of living specimens of P. major was found to be 1.0277 ± 0.0062. The value for preserved animals was lower at 1.0160 ± 0.0029. The relative density of fixed specimens of K. cochlearis was measured and found to be approximately 1.0450 ± 0.0175.     

Keratella distribution in North Patagonian lakes (Argentina)

The distribution of Keratella species from 15 different lakes in North Patagonia (Argentina) was analysed. The genus was not present at altitudes above 1000 m. K. tropica was restricted to Patagonian Plateau lakes with a comparatively high conductivity. A morphometric analysis of the widely distributed K. cochlearis was performed. Results showed three groups of K. cochlearis corresponding to Andean lakes, Patagonian Plateau lakes and a Patagonian Reservoir.     

Food limitation and seasonal population declines of rotifers

Summary Keratella cochlearis exhibited consistent seasonal abundance patterns during a four-year study in Little Rock Lake, Wisconsin, U.S.A. In each year, spring population peaks were followed by strong summer reductions. Concomitant with population declines, there were reductions in rotifer egg ratios. Keratella taurocephala abundance patterns were similar to those of K. cochlearis during 1984 and 1985, but not in 1986 and 1987, when spring peaks and summer declines were not apparent. However, summer declines in the egg ratio of K. taurocephala were observed during each year. The reduction in rotifer populations simultaneously with decreased egg ratios suggested that population declines were caused by food limitation. Food-addition experiments conducted in situ in small enclosures indicated that food was limiting for K. cochlearis when its populations were declining, but not during other periods of the year. Keratella taurocephala did not show a consistent response to food addition.     

Observations on the susceptibility of some protists and rotifers to predation by asplanchna girodi

Direct observations of behavioral interactions show that the predator A. girodi: 1) easily ingested Synchaeta pectinata, two forms of Keratella cochlearis cochlearis, and individuals of Conochilus unicornis and C. dossuarius enzymatically dissociated from their matrix; 2) rarely, if ever, captured Kellicottia bostoniensis and intact Conochilus; and 3) generally rejected the peritrich ciliate Rhabdostyla sp. and the dinoflagellate Peridinium sp. Coloniality and secretion of a gelatinous matrix in Conochilus can be viewed as adaptations to limit mortality from invertebrate predation. Intraspecific variability in the feeding responses of A. girodi is considered.     

The mouth apparatus of Lithodes maja (Crustacea: Decapoda) – form,function and biological role

Being able to utilize many different food resources is probably an important aspect of the success of decapod crustaceans which fill a wide range of various ecological niches worldwide. The phenomenon is facilitated by the complex mouth apparatus found in this group, whose representatives possess six pairs of mouthparts– mandibles, maxillae 1, and maxillae 2 and three maxillipeds, the first three pairs of thoracic appendages which are also specialized to food manipulation. These six pairs are able to perform a number of movements for transporting, aligning, crushing and cutting. Studies into the functional morphology of mouthparts have already been carried out in some decapod species. This study focuses on Lithodes maja, a species of the hitherto understudied king crabs (Lithodidae), chosen on the grounds of their remarkable evolutionary history as ‘derived hermit crabs’. Individuals were filmed while being presented with different kinds of food. To obtain structural information on the individual mouthpart elements as naturally arranged in relation to one another, the shape of the mouthparts was 3D‐reconstructed from micro‐CT scans. These data were complemented by scanning electron microscopy, to analyse the surface structures in detail. There is evidence that the various elements of the mouthparts of L. maja can be sorted into six functional groups: (i) transporting mouthpart elements, (ii) aligning/sorting mouthpart elements, (iii) clutching/holding mouthpart elements, (iv) tearing/cutting/crushing mouthpart elements, (v) current‐generating mouthpart elements and (vi) grooming mouthpart elements. According to our 3D reconstruction, there only seem to be minor differences in morphology and relative position between the mouthparts of L. maja and those of the closely related species, Pagurus bernhardus.     

Morphological and trophic specialization in a subterranean amphipod assemblage

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Food and feeding ecology of postlarval and juvenile Pleuragramma antarcticum (Pisces; Notothenioidei) in the seasonal pack ice zone off the Antarctic peninsula

Summary Stomach contents of 275 postlarval and 269 one year old juvenile Pleuragramma antarcticum caught in February 1982, March 1981 and November 1977 in the Bransfield Strait and adjacent waters were investigated. Juveniles in November 1977 fed mainly on calanoid copepods and Oithona spp. The principal food of postlarvae in February 1982 were Oncaea spp., eggs of calanoid copepods and tintinnids whereas the staple food of juveniles in February 1982 consisted of calanoid copepods of which copepodites of Calanus propinquus were most abundant. Eggs of Euphausia superba were frequently ingested by postlarvae and were the main component of the juvenile diet in the Antarctic Sound. Postlarvae in March 1981 preyed on Oncaea spp. and calanoid eggs. There was a shift from feeding on cyclopoid copepods by postlarvae to feeding on calanoids by juveniles. The food particle size increased as a function of fish length. Maximum prey size was controlled by mouth width up to at least 45 mm standard length. Prey selection was apparently food density dependent with an inverse relationship between food abundance and selective feeding. Density dependent size selection may have resulted in resource partitioning among postlarval and juvenile fishes in February. This specific feeding behaviour is likely to be an adaptation to the low zooplankton stocks of the high Antarctic shelf waters to minimize food competition among the youngest age classes. Predation on krill eggs indicated that the frequent association of one year old juvenile Pleuragramma antarcticum with the Antarctic krill is related to local spawning events.     

Fine structure and functional morphology of the mouthparts of Bittacus planus and Terrobittacus implicatus (Insecta: Mecoptera: Bittacidae)

The Bittacidae are unique in Mecoptera for their adults being predaceous. However, their mouthparts have not been well documented for functional morphology to date. Here, we investigated the mouthpart morphology of the hangingflies Bittacus planus Cheng and Terrobittacus implicatus (Huang & Hua) using scanning electron microscopy. The mouthparts are of the mandibulate type and situated at the tip of an elongated rostrum. The labrum is greatly elongated, roughly twice as long as the subquadrate clypeus. The epipharynx is furnished with a row of basiconic sensilla arranged evenly as a median band extending from the apex to the base. The mandibles are slender and elongated, bearing a sharp lateral and a small mesal tooth. The maxillae are well developed, each consisting of a partially sclerotized cardo and a stipes, a hirsute galea and a lacinia, and a five-segmented maxillary palp. The sensillar pattern on the distal segment of the maxillary palp differs slightly between the two bittacid species. The labium is composed of a postmentum, a prementum, and a pair of two-segmented labial palps. The feeding mechanism of bittacids is briefly discussed in combination with the mouthpart morphology and their feeding habits.     

The first record of two demersal calanoid copepods,Pseudodiaptomus poplesia and P. nihonkaiensis in Korea,with remarks on morphology of the genital area

The demersal calanoid copepods Pseudodiaptomus nihonkaiensis Hirakawa, 1983 and P. poplesia (Shen, 1955) are redescribed from Korean waters. Using scanning electron microscopy, we examine the morphology of the female genital systems of four species of Pseudodiaptomus (P. inopinus, P. marinus, P. nihonkaiensis and P. poplesia), revealing interspecific differences. The zoogeography of these four species is discussed.     

Feeding convergence in South American and African zooplanktivorous cichlids Geophagus brasiliensis and Tilapia rendalli

Synopsis Acará, Geophagus brasiliensis, and red-breasted bream, Tilapia rendalli, are important planktivorous cichlids in southern Brazilian lakes and reservoirs. In laboratory experiments, I quantified behavior and selectivity of different sizes of these two fish feeding on lake zooplankton. Feeding behavior depended on fish size. Fish < 30 mm were visual feeders. Fish 30–50 mm either visually fed or pump-filter fed depending on zooplankton size. Fish > 70 mm were pump-filter feeders. Replicate 1 h feeding trials revealed that, as the relative proportions of prey changed during an experiment, acará (30–42 mm, standard length) and tilapia (29–42 mm) shifted from visual feeding on large evasive copepods to filter feeding on small cladocerans and rotifers. Electivity and feeding rate increased with prey length, but were distinct for similar-sized cladocerans and copepods. Visual/filter-feeding fish had lowest electivities for small and poorly evasive rotifers and cyclopoid nauplii. They fed non-selectively on cyclopoid copepodites, had intermediate electivities for calanoid nauplii and small cladocerans, and had highest electivities for large cladocerans, cyclopoid adults, and calanoid copepodites and adults. Although belonging to different cichlid genera and native to South America and Africa, respectively, acará and red-breasted bream (= congo tilapia) exhibited similar selectivity for zooplankton. Apparently, few stereotyped feeding behaviors have evolved during the acquisition of microphagy in fish. Shift in feeding modes allows these two species to optimally exploit the variable and dynamic patchy distribution of planktonic resources.     

Daytime grazing and assimilation rates of planktonic copepods Acanthodiaptomus denticornis and Cyclops vicinus vicinus. Comparison of spatial and resource utilisation by rotifers and cladoceran communities in a eutrophic lake

The clearance rates of ¹⁴ carbon-labelled Chlamydomonas sp. by the copepodite and adult stages of the 2 dominant species of Copepoda were studied in eutrophic Lake Aydat. They varied from 30 to 3,007 µl ind^–1 h^–1 for Acanthodiaptomus denticornis (mean = 600 µl ind^–1 h^–1) and from 20 to 1,133 µl ind^–1 h^–1 for Cyclops vicinus (mean = 340 µl ind^–1 h^–1). Their maximum assimilation efficiencies were 58% and 50%, respectively. These populations collectively could consume the available food in 4 days during September (maximum daily grazing rate = 24%). Mean individual clearance rates could be ranked Acanthodiaptomus denticornis > Ceriodaphnia quadrangula > Chydorus sphaericus > Daphnia longispina > Cyclops vicinus vicinus > Bosmina longirostris > K. cochlearis > K. quadrata and Kellicottia longispina. Like cladocerans and rotifers, the copepods living in this eutrophic lake can feed at low oxygen concentrations.     

Influence of temperature on morphological variation in populations of Keratella cochlearis (Gosse) in Rybnik Reservoir

Morphological changes in a population of Keratella cochlearis were investigated in a reservoir of changed temperature and high trophy. Four morphological forms were distinguished: Keratella cochlearis f. cochlearis, K. cochlearis var. tecta f. micracantha, K. cochlearis var. tecta f. micracantha, K. cochlearis var. tecta f. typica and K. cochlearis f. macracantha. A correlation between the length of the lorica and the posterior spine, and the temperature of water was observed. These four forms of Keratella cochlearis occurred during the entire period of investigations.     

Short term experiments on calanoid-cyclopoid-phytoplankton interactions

To investigate their potential effects on each other in nature, calanoid (Diaptomus clavipes and D. siciloides) and cyclopoid (Acanthocyclops vernalis and Mesocyclops edax) copepod populations were manipulated in 5 liter aquaria in laboratory experiments of 20–60 days duration. Diaptomus generally had a strongly negative effect on both cyclopoid species. The cyclopoids established populations more successfully when introduced to aquaria before calanoids than they did when calanoids were already present. On the other hand, whether introduced earlier or later than the cyclopoids, Diaptomus populations were unaffected by Acanthocyclops and were strongly depressed by Mesocyclops. Diaptomus effects on the phytoplankton were often strong but varied markedly among experiments. They included reduction of populations of edible algae, such as Chlamydomonas, which are essential for both calanoid and cyclopoid nauplii, and large increases in inedible algae, such as Kirchneriella. Feeding experiments revealed that under conditions of food scarcity Acanthocyclops nauplii survived less well than did Diaptomus nauplii. Competition for edible phytoplankton seemed to be a key factor in the calanoid-cyclopoid interactions, since the survival of herbivorous cyclopoid larvae determined the abundance of the predaceous adults. This indicates that the competitive effects of calanoids on cyclopoids often may exceed the predative effects of cyclopoids on calanoids.